|
|
Line 5,519: |
Line 5,519: |
|
| |
|
|
| |
|
| ==Part III Organogeny==
| | {{Footer}} |
| | |
| Chapter VIII Endodermal Derivatives
| |
| | |
| The tissues derived directly from the endoderm are for the
| |
| most part of the epithelial type and form the inner lining of the
| |
| gastrocoel and the organs that arise therefrom. These organs
| |
| are grouped into two closely connected organ systems, the digestive system and the respiratory system. The digestive (enteric)
| |
| tube, however, becomes ensheathed in splanchnic mesoderm which
| |
| contributes largely to the ultimate structure of the organ systems
| |
| just mentioned. Furthermore, this tube opens to the exterior
| |
| at both the anterior and posterior ends by means of two ectodermal pits, the stomodeum and proctodeum, respectively. All
| |
| three germ layers, therefore, contribute to the organogeny of these
| |
| systems.
| |
| | |
| The stomodeum. — There is an ectodermal invagination on
| |
| the ventral side of the head to form the stomodeum (Fig. 118),
| |
| | |
| Notochord Neural tube
| |
| | |
|
| |
| | |
| canal
| |
| | |
| Fig. 118. — Diagram of an early vertebrate embryo, to show endodermal derivatives.
| |
| | |
| which is bounded on the sides by the maxillary ridges and on the
| |
| rear by the mandibular ridges. The rupture of the oral plate,
| |
| which separates the stomodeum from the fore-gut, results in the
| |
| formation of the oral cavity, or mouth. From the stomodeum
| |
| another invagination, the hypophysis, grows upward in front of:
| |
| the fore-gut, and eventually fuses with an evagination from the
| |
| | |
| floor of the neural tube, the infundibulum, to form the pituitary
| |
| 181
| |
| 182 ENDODERMAL DERIVATIVES
| |
| | |
| gland, an organ of internal secretion. As the stomodeum joins
| |
| the fore-gut a little posterior to the anterior end of the latter
| |
| cavity, there is a blind pocket of endoderm, anterior to the
| |
| mouth, called the preoral gut.
| |
| | |
| The oral cavity. — The cavity of the mouth is a compound
| |
| structure, derived in part from the ectodermal stomodeum and
| |
| | |
|
| |
| | |
|
| |
| | |
| Epithelium —=y4r3
| |
| of mouth Snosie
| |
| | |
| Enamel
| |
| | |
| Dentine
| |
| | |
| Fig. 119. — Diagram to show origin of vertebrate tooth (lower jaw).
| |
| | |
| in part from the endodermal fore-gut. The boundary line between these is soon lost after the rupture of the oral plate owing
| |
| to unequal local growth of the different regions of the mouth.
| |
| The boundaries of the mouth are the upper jaws, formed from the
| |
| maxillary ridges, and the lower jaws, derived from the mandibular
| |
| | |
| Visceral pouches
| |
| | |
|
| |
|
| |
| | |
| Dorsal epithelial bodies stomach
| |
| Esophagus
| |
| | |
| Dorsal pancreas
| |
| | |
|
| |
| | |
| Mid-gut
| |
| | |
| Ventral pancreas
| |
| | |
| epithelial bodies Hepatic diverticulum
| |
| | |
| Tongue
| |
| | |
| Fia. 120. — Diagram showing derivatives of vertebrate fore-gut.
| |
| | |
| ridges. On these ridges the teeth arise in exactly the same way
| |
| as the placoid scales of the elasmobranchs (page 230). Two elements are concerned: an ectodermal enamel organ, shaped like an
| |
| inverted cup; and a mesodermal dental papilla, which fills the
| |
| cavity of the enamel organ. The enamel organ gives rise to the
| |
| outer enamel layer of the tooth, while the papilla forms the dentine
| |
| (Fig. 119). The dentine is in the general form of a hollow cone,
| |
| THE INTERNAL GILLS 183
| |
| | |
| the cavity of which is filled with connective tissue, nerves, and
| |
| blood vessels. The tongue (Fig. 120) is also a compound organ,
| |
| arising from an endodermal primary tongue which is formed from
| |
| the floor of the pharynx in the region of the hyoid arches, and
| |
| from an ectodermal secondary tongue which arises from the
| |
| floor of the oral cavity in front of the thyroid gland (page 184).
| |
| Into the tongue a migration of mesoderm takes place, by means
| |
| of which the musculature is formed. The glands of the mouth
| |
| (salivary glands, etc.) arise from the ectodermal lining of the
| |
| mouth. The taste buds, however, are endodermal (Holtfreter,
| |
| 1933). The connection between the oral cavity and the nasal
| |
| cavity will be discussed in Chapter X.
| |
| | |
| The pharynx. — The region of the fore-gut which follows the
| |
| oral cavity is the pharynx, particularly important on account of
| |
| the respiratory organs and other structures which arise from it.
| |
| | |
| Respiratory organs. — Respiratory exchange may take place
| |
| in any thin epithelium in which the blood corpuscles are brought
| |
| into contact with the oxygen-carrying medium. ‘These epithelia
| |
| may be either ectodermal or endodermal in origin. Thus, we
| |
| find that among the amphibia, respiration may take place in the
| |
| skin as a whole (lungless salamandeys) ; ; An specialized outgrowths
| |
| on the visceral arches, external gills (N. ecturus); or in the so-called
| |
| “hairs ” of the African frog, Astylosternus. In this group are to
| |
| be found also examples of endodermal respiratory organs, the internal gills and lungs. Internal gills are otherwise found only
| |
| among the fish, while the lungs are characteristic respiratory
| |
| organs also of the amniotes.
| |
| | |
| The internal gills. -- The internal gills (branchiae) arise in the
| |
| visceral clefts (Fig. 120) common to all chordates. Among the
| |
| aquatic vertebrates these are typically six in number (see Table 8,
| |
| page 131). In the cartilage fish the first cleft (the spiracle) opens
| |
| on the dorsal side of the head and is otherwise modified. The
| |
| clefts are separated by the visceral arches, of which the first is
| |
| known as the mandibular arch and the second is called the hyoid
| |
| arch. The visceral clefts are formed by the coming together of
| |
| paired evaginations of the endoderm (visceral pouches) and complementary invaginations of the ectoderm (visceral grooves).
| |
| The ectoderm and endoderm come into direct connection to form
| |
| closing plates. Later, these plates rupture and a series of finger184 ENDODERMAL DERIVATIVES
| |
| | |
| like projections grow out into the cleft from the anterior and
| |
| posterior sides of each arch. These filamentous processes usually
| |
| fuse to form a demibranch (Fig. 197). The demibranchs in some
| |
| fish are apparently of endodermal origin, while in the amphibia
| |
| they are derived from the ectoderm. It is interesting to note that
| |
| in the spiracle of the cartilage fish a gill-like structure, the
| |
| pseudobranch, develops. In amphibians and the amniotes generally the first visceral pouch does not open to the exterior but
| |
| gives rise to the tympanic cavity and auditory tube (see Chapter
| |
| X). In all fish except the elasmobranchs, a projection grows
| |
| back from the hyoid arch to cover the remaining visceral clefts.
| |
| This is the operculum. Internal gills do not appear in the development of the amniotes; but the visceral clefts, or at least the
| |
| visceral pouches and grooves, are of invariable occurrence.
| |
| | |
| The lungs. — In all the vertebrates except the cyclostomes and
| |
| cartilage fish, there develops from the pharynx a sac (or a pair
| |
| of sacs) which becomes the air bladder in pisces and the lungs
| |
| in tetrapoda. We shall confine our attention here to the development of the lungs (Fig. 120). The first indication of lung formation is the appearance of a longitudinal groove in the floor of the
| |
| pharynx posterior to the last pair of visceral pouches. This is
| |
| the tracheal groove. This groove separates from the pharynx,
| |
| the process commencing at the posterior end, so that the dorsal
| |
| portion of the tube, or esophagus, is separated from the ventral
| |
| portion, or trachea, except for a narrow opening, the glottis.
| |
| The trachea grows backward rapidly and divides into two lobes,
| |
| the primordia of the lungs. There is some evidence that the
| |
| trachea is bifurcated from its first appearance, suggesting that
| |
| the lungs arise from paired primordia. In the birds and mammals
| |
| the lung primordia subdivide many times to form the bronchi, or
| |
| branches of the respiratory tree.
| |
| | |
| The thyroid gland. — This structure arises as a median ventral
| |
| evagination of the pharyngeal floor between the primary and the
| |
| secondary tongue primordia or at the level of the hyoid arches.
| |
| The diverticulum grows downward and expands at its distal
| |
| end (Fig. 120). Eventually, its connection with the pharyngeal
| |
| floor, the thyroglossal duct, becomes occluded and disappears,
| |
| and the gland itself subdivides into a mass of vesicles which
| |
| migrate backward and assume somewhat different positions in
| |
| THE STOMACH 185
| |
| | |
| various vertebrates, often ending as a paired organ on either side
| |
| of the trachea.
| |
| | |
| The epithelial bodies.— In all the vertebrates there arise,
| |
| from the upper or lower angles of the visceral pouches, small
| |
| buds of epithelium which often give rise to endocrine glands of
| |
| varying — and mostly unknown — function (Figs. 120, 121).
| |
| The dorsal buds (except among the mammals, where conditions
| |
| are reversed) contribute in varying number to the formation of a
| |
| large gland, the thymus, which loses connection with the pharynx
| |
| and moves backward to its definitive position, which differs
| |
| according to the form studied. The remainder of the dorsal
| |
| | |
|
| |
|
| |
|
| |
| | |
|
| |
|
| |
|
| |
| | |
| Thymus
| |
| I Thyroid 1 Thyroid 1
| |
| aD ap
| |
| Thyroid Tr L H
| |
| Thymus pare
| |
| thyroi
| |
| Para- Para- mw ¥
| |
| thyroid thyroid Th
| |
| <4 ymus w
| |
| Y
| |
| B
| |
| Ultimo- Ultimobranchial branchial
| |
| | |
| Ultimobranchial
| |
| | |
| Fig. 121. — Diagrams showing origin of epithelial bodies in A, frog; B, chick;
| |
| and C, man.
| |
| | |
| bodies become lymphoid and degenerate. The ventral buds
| |
| (absent in fish) detach themselves from the pharyngeal wall
| |
| and take up varying positions. Among the mammals it is
| |
| the ventral buds which form the thymus, while the dorsal buds
| |
| of the third and fourth pouches move to the sides of the thyroid
| |
| gland where they are known as the parathyroids.
| |
| | |
| The esophagus. — The digestive canal behind the pharynx
| |
| becomes specialized into four regions: (1) the esophagus; (2)
| |
| the stomach; (3) the intestine and its derivatives; and (4) the
| |
| cloaca. Of these, the esophagus (Fig. 120) remains comparatively
| |
| unspecialized; it is a narrow tube, short in the anamniotes, elongate in the amniotes. No digestive glands are found in this region.
| |
| | |
| The stomach. — This portion of the digestive tract is distinguished by its dilation (Fig. 120) into a large sac or series of
| |
| 186 ENDODERMAL DERIVATIVES
| |
| | |
| sacs, and by the development of a thick wall of muscle from the
| |
| splanchnic mesoderm in which it is enveloped. The stomach is
| |
| rich in glands which aid in digesting the passing food.
| |
| | |
| The intestine. — All the regions of the digestive tract mentioned
| |
| so far are derived from the fore-gut. The intestine is derived in
| |
| part from the fore-gut, in part from the mid-gut, and in part from
| |
| the hind-gut. It is impossible to indicate exactly which regions
| |
| arise from these divisions of the gut, as both the fore-gut and the
| |
| hind-gut expand at the expense of the mid-gut during the consumption of the yolk. As was said in the discussion of the development of body form, the division of the alimentary canal
| |
| into these regions is the result of the method by which the head
| |
| and tail are formed. The intestine becomes subdivided in various
| |
| ways in the different groups, but we need notice only the most
| |
| anterior of these, the duodenum, which is that portion of the
| |
| intestine immediately succeeding the stomach and generally held
| |
| to be derived from the fore-gut. The intestine is richly glandular
| |
| throughout its length, but from the duodenum, in particular, we
| |
| find developed two most important glands, the liver and the
| |
| pancreas (Fig. 120).
| |
| | |
| The liver. — This gland arises from the ventral side of the duodenum as an evagination which grows forward, expanding into
| |
| a vesicle at the distal end and retaining its connection with the
| |
| duodenum by a narrow hollow stalk, the common bile duct,
| |
| (Fig. 120). The sac-like distal end becomes subdivided, by the
| |
| ingrowth of mesenchyme, into many tubules which often anastomose. In this process of growth and subdivision the liver grows
| |
| about the vitelline veins (Chapter [X) and breaks these up into
| |
| a system of hepatic capillaries. The cavity of the sac becomes
| |
| the gall-bladder, to which the bile, formed in the glandular
| |
| portion of the liver, is carried by means of the hepatic ducts. It
| |
| releases these secretions into the duodenum via the common
| |
| bile duct (ductus choledochus).
| |
| | |
| The pancreas. — This gland arises usually from three diverticula of the duodenum (Fig. 120), but the number of primordia
| |
| is variable. One appears on the dorsal side of the duodenum
| |
| just posterior to the stomach; the others arise on the ventral
| |
| side, usually in connection with the hepatic diverticulum. The
| |
| primordia increase in size, and break up into masses of secretory
| |
| THE FROG 187
| |
| | |
| tubules at the distal end of each. The primordia unite and their
| |
| proximal ends become the pancreatic ducts, one or more of which
| |
| may be suppressed in later organogeny. The pancreas, as well
| |
| as elaborating a digestive pancreatic juice discharged through the
| |
| pancreatic duct, forms a hormone (insulin), which is carried away
| |
| by the blood stream. It functions therefore as an endocrine
| |
| gland in addition to its digestive function. Insulin, as is well
| |
| known, is important in the treatment of diabetes.
| |
| | |
| The cloaca. — The intestine behind the duodenum is variously
| |
| subdivided in the different vertebrate classes, but all are alike in
| |
| the possession of a terminal region which receives in addition the
| |
| ends of the nephric ducts and of the genital ducts (see Chapter IX).
| |
| From the cloaca also arises the urinary bladder and the allantois
| |
| of the amniotes.
| |
| | |
| The cloaca, like the pharynx, communicates with the exterior
| |
| by means of an aperture lined with ectoderm, which arises as
| |
| a median ventral pit, the proctodeum (Fig. 118), just in front
| |
| of the tail region. The proctodeum is formed at the point where
| |
| the blastopore was obliterated and is separated from the hind-gut
| |
| temporarily by means of the cloacal plate, which is comparable
| |
| with the oral plate. For a time there is a blind pocket of endoderm posterior to the cloaca, which is known as the postcloacal
| |
| gut. The region of the cloaca anterior to the entrance of the
| |
| nephric ducts is known as the rectum; its aperture is called the
| |
| vent. In mammals the rectum becomes separated from the remainder of the cloaca, which is then known as the urogenital
| |
| sinus. Each of these cavities has a separate exit, the two openings
| |
| being the anus and the urogenital aperture, respectively.
| |
| | |
| THE FROG (SEE ALSO CHAPTER XI).— The mouth of the tadpole does not open until a few days after hatching. It remains
| |
| round during larval life and is enclosed by the mandibular ridges.
| |
| Outside these, folds of ectoderm project as the larval lips, on which
| |
| horny larval teeth develop. These larval structures are lost at
| |
| metamorphosis, when the definitive jaws and teeth are formed
| |
| in the usual way. The tongue is compound, arising from a primary tongue and a gland field, relatively late in larval life. The
| |
| hypophysis is solid (Fig. 181).
| |
| | |
| Six visceral pouches appear, of which the first never ‘becomes
| |
| perforated, its closing plate becoming the tympanum of the ear,
| |
| 188 ENDODERMAL DERIVATIVES
| |
| | |
| and its cavity persisting as the tubo-tympanic cavity. Of the
| |
| five remaining pouches, the second and third open to the exterior before the first and fourth, and the fifth remains vestigial.
| |
| External gills appear on the third, fourth, and fifth arches (that
| |
| on the fifth arch being rudimentary), but are resorbed later when
| |
| covered by the operculum. This structure fuses with the body
| |
| surface on the right side, but on the left it opens to the exterior
| |
| by an opercular aperture. The internal gills appear as demibranchs commencing on the anterior side of the third arch. The
| |
| first three gills, therefore, have two demibranchs, while the fourth
| |
| has but one, formed from the anterior side of the sixth arch. The
| |
| visceral clefts, gills, and opercular cavity are lost as separate
| |
| structures by cell proliferation and reorganization just before
| |
| metamorphosis. The lungs appear early in larval life as solid
| |
| primordia of the pharynx. These acquire cavities prior to the
| |
| formation of the tracheal groove which is relatively late in formation. The thyroid arises, just before hatching, as a solid diverticulum of the pharynx; it soon detaches itself and divides into
| |
| two bodies which later become vesicular. The two thymus
| |
| glands are formed from epithelial bodies on the dorsal side of the
| |
| first and second visceral pouches. Epithelial bodies arise from
| |
| the ventral sides of the second visceral pouches. It has been
| |
| claimed that those of the third and fourth pouches become the
| |
| carotid glands. The sixth pharyngeal pouches give rise to the
| |
| ultimobranchial (suprapericardial) bodies. (Fig. 121A.)
| |
| | |
| The esophagus is short, and the stomach a simple dilation.
| |
| The liver arises as a backward ventral diverticulum of the duodenum (Fig. 181). All three pancreatic primordia appear and
| |
| fuse; the dorsal duct disappears, while the two ventral ducts
| |
| fuse to become the adult pancreatic duct. The intestine of the
| |
| tadpole, which is long and coiled (about nine times the body
| |
| length), becomes resorbed during metamorphosis until it is about
| |
| one-third of its larval length (Fig. 122).
| |
| | |
| The postcloacal gut loses its connection with the neural tube
| |
| (neurenteric canal) during the backward growth of the tail. The
| |
| urinary bladder does not appear until after metamorphosis.
| |
| | |
| THE CHICK (SEE ALSO CHAPTER XII). — The mouth opens on the
| |
| third day of incubation. The teeth are represented only by the
| |
| tooth ridges which are the first stage in the appearance of the
| |
| THE CHICK 189
| |
| | |
| enamel organs. These appear on the sixth day of incubation and
| |
| disappear shortly after the cornification of the jaws. This results in the formation of the beak and the egg tooth, the latter
| |
| a horny projection on the upper jaw which is used in breaking
| |
| through the shell at the time of hatching, and soon after disappears. The primordia of the tongue appear on the fourth day.
| |
| | |
| Five visceral pouches appear, of which the first three open to
| |
| the exterior during the third day of incubation (Fig. 218). The
| |
| | |
|
| |
| | |
| Bladder
| |
| | |
|
| |
| | |
| Fig. 122. — Digestive tube in A, tadpole, and B, frog, to show actual shortening of
| |
| intestine. (After Leuckart wall-charts.)
| |
| | |
| first cleft closes during the fourth day, and the dorsal part of the
| |
| pouch becomes the tubo-tympanic cavity. With the extension
| |
| of the cervical flexure, the remaining pouches are crowded together and disappear. The thyroid appears on the second day,
| |
| separates from the pharynx on the fourth, and on the seventh
| |
| divides inte two bodies which migrate backward to the junction
| |
| of the common carotid and subclavian arteries. The thymus
| |
| arises from the dorsal epithelial bodies of the third and fourth
| |
| visceral pouches, while the parathyroid rudiments arise from
| |
| 190 ENDODERMAL DERIVATIVES
| |
| | |
| the ventral epithelial bodies. The fifth pouch gives rise to the
| |
| ultimobranchial bodies. The lung primordia (Fig. 123) appear on
| |
| the third day and grow back, becoming surrounded by mesenchyme. The primary bronchi subdivide to form a respiratory
| |
| tree, some branches of which extend among the viscera and even
| |
| into the hollow bones, as the accessory air sacs.
| |
| | |
| The esophagus is relatively long; and a dilation, the crop,
| |
| forms at its posterior end. The stomach is divided into an anterior proventriculus, which contains the gastric glands, and a
| |
| | |
| Visceral arches
| |
| | |
|
| |
|
| |
| | |
| Dorsal
| |
| pancreas
| |
| | |
| Yolk stalk
| |
| | |
| Fig. 123. — Endodermal derivatives in a 72-hour chick.
| |
| | |
| muscular gizzard at the posterior end. The liver primordium
| |
| arises at the edge of the anterior intestinal portal on the second
| |
| day and, therefore, presents the aspect of an anterior ventral
| |
| and two posterior lateral diverticula for a short time. These
| |
| fuse, however, by the end of that day, as the backward extension
| |
| of the fore-gut continues. Three pancreatic diverticula are
| |
| formed, the dorsal one on the third day, the ventral ones on the
| |
| fourth. They fuse in later development, and either two or three
| |
| of the ducts persist. The anterior portion of the mid-gut becomes
| |
| the small intestine, the large intestine arising from the posterior
| |
| | |
| region..
| |
| MAN 191
| |
| | |
| The cloaca is first distinguishable on the fourth day, when the
| |
| proctodeum also is first apparent. The cloaca is ultimately
| |
| divided into three regions: an anterior portion, the coprodeum,
| |
| into which the rectum enters; an intermediate part, the urodeum,
| |
| into which the nephric ducts and gonoducts enter; and the terminal proctodeum.
| |
| | |
| MAN (SEE ALSO CHAPTER XIII). — The mouth opens in the second or third week, and, like that of all vertebrates, develops lips
| |
| (fifth week). Ten teeth papillae and enamel caps, the primordia
| |
| of the milk teeth, appear in each jaw. This is a long-drawn-out
| |
| process, the germs of the third molar not appearing until the
| |
| fifth year of infancy. The tongue arises from swellings on the
| |
| first three arches, the secondary tongue, or gland field, appearing
| |
| as the tuberculum impar, which does not, however, appear to
| |
| contribute to the ultimate structure of the tongue.
| |
| | |
| Five pairs of visceral pouches appear, none of which becomes
| |
| perforated. The first gives rise to the tubo-tympanic cavity.
| |
| The ventral portion of the second persists as the fossa in which
| |
| the tonsil develops. The dorsal epithelial bodies from the third
| |
| and fourth pair of pouches become the parathyroids. The
| |
| ventral epithelial bodies of the third pair of pouches unite to form
| |
| the thymus gland. Similar bodies from the fourth pair may give
| |
| rise to vestigial thymus-like bodies which remain attached to
| |
| the parathyroids from the same pouch. The fifth pair become
| |
| the ultimobranchial bodies. The thyroid gland undergoes an
| |
| incomplete division into two lobes which remain connected by a
| |
| narrow isthmus. The lungs (Fig. 124) arise toward the end of the
| |
| fourth week, from a laryngo-tracheal groove. The cartilages and
| |
| musculature of the larynx arise from the branchial arches.
| |
| | |
| The esophagus, at first relatively short, lengthens as the backward movement of the heart and lungs displaces the stomach.
| |
| The latter organ arises as a dilation of the fore-gut posterior
| |
| to the esophagus. Continued growth, mainly on the dorsal
| |
| surface, produces the greater curvature, and a displacement of
| |
| the whole organ so that the cephalic end is moved to the left
| |
| and the caudal end to the right. This is followed by a rotation
| |
| of the stomach on its long axis through 90° to the left. The liver
| |
| ari ring the third week as a ventral groove in the duodenum.
| |
| The-pancreas appears slightly later, with either two or three
| |
| 192 ENDODERMAL DERIVATIVES
| |
| | |
| primordia according to whether or not one of the ventral primordia
| |
| is suppressed. The ventral pancreatic duct persists and opens
| |
| into the common bile duct. The point of division between small
| |
| and large intestines is marked by the formation of a blind pouch,
| |
| | |
| Visceral arches
| |
| | |
|
| |
|
| |
|
| |
|
| |
| | |
| ( Hypophysis )
| |
| | |
| Stomach
| |
| | |
| Ventral
| |
| | |
| pancreas Intestinal loop
| |
| | |
| Dorsal
| |
| pancreas,
| |
| | |
|
| |
|
| |
|
| |
|
| |
| | |
| Allantoic stalk
| |
| | |
| bladder
| |
| | |
| Metanephric
| |
| | |
| ( Mesonephric duct ) liverticulum
| |
| | |
| a7
| |
| | |
| Fig. 124. — Endodermal derivatives in 10-mm. pig. (From a wax reconstruction
| |
| by G. W. Hunter and L. T. Brown.)
| |
| | |
| the cecum. The distal end of the cecum does not grow as rapidly
| |
| as the proximal region and so remains a finger-like projection
| |
| known as the vermiform appendix. The small intestine, growing
| |
| more rapidly than the large, is thrown into a set of six primary
| |
| coils, each of which develops secondary coils.
| |
| | |
| The cloaca becomes divided, by a frontal partition, into a
| |
| SUMMARY 193
| |
| | |
| dorsal rectum and a ventral urogenital sinus. The cloacal
| |
| membrane is correspondingly divided into a rectal and a urogenital plate, and the final openings are the anus and the urogenital aperture. The urogenital sinus later is divided into a
| |
| phallic portion (see page 211) and a vesico-urethral portion. The
| |
| latter gives rise to the urinary bladder at its distal end, and to.
| |
| the urethra at its proximal end.
| |
| | |
| SUMMARY
| |
| | |
| The endoderm gives rise to the epithelial lining of the following
| |
| structures:
| |
| | |
| A. Fore-gut
| |
| I. Oral cavity (also partly from ectoderm of stomodeum)
| |
| | |
| Teeth (also partly from ectoderm)
| |
| Tongue
| |
| II. Pharynx
| |
| Trachea and lungs
| |
| Thyroid
| |
| Visceral pouches
| |
| Auditory tube and chamber
| |
| Fossa of palatine tonsil
| |
| Thymus
| |
| Parathyroids
| |
| Ultimobranchial bodies
| |
| III. Esophagus
| |
| | |
| IV. Stomach
| |
| V. Duodenum
| |
| | |
| Liver
| |
| Pancreas
| |
| B. Mid-gut
| |
| I. Intestine
| |
| C. Hind-gut
| |
| I. Cloaca (also partly from ectoderm of proctodeum)
| |
| Rectum
| |
| | |
| Urogenital sinus
| |
| | |
| Urinary bladder
| |
| | |
| Urethra (also partly from mesoderm, page
| |
| 204)
| |
| 194 ENDODERMAL DERIVATIVES
| |
| | |
| REFERENCES
| |
| | |
| Keey, L. B. 1934. Developmental Anatomy, 3rd Ed., Chaps. 7 and 8.
| |
| | |
| Brachet, A. 1921. Traité d’embryologie des vertébrés, Part 2, Bk. 1, Chap. 5;
| |
| Bk. 2, Chap. 4.
| |
| | |
| Hertwig, O. 1906. Handbuch, Vol. 2, Chaps. 1, 2, and 4.
| |
| | |
| Keibel and Mall. 1910-1912. Human Embryology, Chap. 17.
| |
| | |
| Kellicott, W. E. 1913. Chordate Development.
| |
| | |
| Kerr, J.G. 1919. Textbook of Embryology, Vol. II, Chap. 3.
| |
| | |
| Kingsley, J.S. 1926. Comparative Anatomy of Vertebrates, 3rd Ed.
| |
| | |
| Lillie, F. R. 1919. The Development of the Chick, 2nd Ed.
| |
| | |
| MeMurrich, J. P. 1923. The Development of the Human Body, 7th Ed.
| |
| CHAPTER IX
| |
| MESODERMAL DERIVATIVES
| |
| | |
| The middle germ layer arises as three different aggregates of
| |
| cells between the ectoderm and endoderm: the notochord; the
| |
| mesoderm; and the mesenchyme. The origin of the notochord
| |
| has already been described, and its later history will be discussed
| |
| in connection with the skeleton. Organs of mesenchymatous
| |
| origin will be taken up in connectiqn with the history of the
| |
| region from which their mesenchyme originates. Of the structures derived from the mesoderm, we shall consider first those
| |
| arising from the lateral mesoderm, then those whose origin is from
| |
| the intermediate mesoderm, and finally those derived from the
| |
| axial mesoderm.
| |
| | |
| A. THE COELOM AND ITS MESENTERIES
| |
| | |
| Cavities may appear in all three divisions of the mesoderm;
| |
| if in the myotomes, they are known as myocoels; if in the
| |
| nephrotomes, they are called nephrocoels; the cavity of the
| |
| lateral mesoderm is the coelom (Fig. 76). In some forms the
| |
| three cavities are confluent. The connection, however, is a temporary one, and the myocoels soon disappear. In other forms
| |
| they make a transitory appearance and are entirely disconnected
| |
| with the other cavities, and in many vertebrates myocoels are
| |
| never formed. The nephrocoels will be considered with the
| |
| nephric organs. The coelom_ in amphioxus has a metameric
| |
| origin from the ventral portions of the enterocoels, which become
| |
| confluent at this point by the disappearance of the intervening
| |
| anterior and posterior partitions. In vertebrates the coelomic
| |
| cavity arises from the splitting of the lateral mesoderm into a
| |
| dorsal somatic and a ventral splanchnic layer. In the amniotes
| |
| this: split continues out into the extra-embryonic mesoderm, thus
| |
| giving rise to the exocoel, or cavity of the chorion. The coelom
| |
| does not extend anterior to the visceral arches. Transitory
| |
| | |
| cavities have been found in the arches and, indeed, in the head
| |
| 195
| |
| 196 MESODERMAL DERIVATIVES
| |
| | |
| itself, and these have been interpreted as the remains of a cephalic
| |
| coelom. It will appear later that these are more probably the
| |
| rudiments of cephalic myotomes. The coelom does not extend
| |
| into the tail.
| |
| | |
| Somatopleure and splanchnopleure. — The somatopleure has
| |
| already been defined as the outer layer of the lateral mesoderm
| |
| together with the ectoderm with which it becomes associated.
| |
| Between these two there is an invasion of mesenchymatous cells
| |
| from the dermatomes and myotomes which give rise to the
| |
| corium of the skin (see Chapter X) and to its dermal musculature (see page 239). The somatic mesoderm lining the outer
| |
| wall of the coelom becomes the outer peritoneal lining. The
| |
| splanchnopleure is the inner layer of the lateral mesoderm plus
| |
| the endoderm with which it is associated. Between these two
| |
| occurs a migration of mesenchyme cells which give rise to the
| |
| splanchnic musculature and blood vessels, while the splanchnic
| |
| mesoderm itself forms the inner peritoneal lining of the coelom.
| |
| | |
| The mesenteries (Fig. 125).—JIn all the vertebrates, the
| |
| coelom is divided for a time into right and left halves by sagittal
| |
| partitions above and below the alimentary canal, known as the
| |
| dorsal mesentery and the ventral mesentery, respectively. These
| |
| are formed by the inward growth of the splanchnic mesoderm
| |
| above and below the digestive tube and the subsequent fusion
| |
| of these sheets in the median line. The ventral mesentery disappears posterior to the liver, probably in connection with the
| |
| coiling of the intestine. The dorsal mesentery (Fig. 125) persists
| |
| as the support of the alimentary canal, and frequently becomes
| |
| subdivided into regions which are named from the supported
| |
| organ, such as the mesogastrium which supports the stomach, the
| |
| mesoduodenum, etc. In the formation of the ventral mesentery,
| |
| two organs, the heart and the liver, owing to their ventral position,
| |
| are caught in between the two advancing sheets of splanchnic
| |
| mesoderm. In these regions, therefore, the ventral mesentery is
| |
| divided into an upper and a lower half. The ventral mesentery
| |
| dorsal to the heart becomes the dorsal mesocardium; that part
| |
| which is ventral to the heart is the ventral mesocardium (Fig.
| |
| 126A). Both eventually disappear as the heart increases in size
| |
| and complexity. In the region of the liver, the dorsal half of the
| |
| mesentery becomes the dorsal mesohepar, while the ventral porTHE MESENTERIES 197
| |
| | |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
| | |
| Pericardial cavity—__f 4 Dorsal mesocar, a dium
| |
| | |
| Ventricle of heart
| |
| | |
| A
| |
| | |
| Ventral mesocardium +- Septum transversum
| |
| | |
|
| |
| | |
| Liver Stomach
| |
| Ventral mesentery st omac
| |
| (faeiform Hgament) Ventral mesentery
| |
| | |
| (lesser omentum)
| |
| Dorsal mesogastrium
| |
| | |
| Dorsal pancreas
| |
| | |
| Fia. 125. — Diagram of mesenteries in early human embryo from left side. A, B,
| |
| and C indicate planes of sections shown in Fig. 126. (From Arey after Prentiss.)
| |
| | |
|
| |
|
| |
|
| |
| | |
| Neural tube
| |
| Notochord Neural tub
| |
| Aorta NotochordPostcardinal vein Aorta
| |
| | |
| Dorsal mesentery,
| |
| | |
| — Fore-gut
| |
| Lesser
| |
| ‘omentum
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
| Liver
| |
| | |
|
| |
| | |
| Peritoneal
| |
| cavity
| |
| Foalciform
| |
| ligament
| |
| | |
| B Cc
| |
| | |
| Fra. 126. — Diagrams of mesenteries in early human embryo as seen in transverse
| |
| sections. Compare lig. 125. (From Arey after Prentiss.)
| |
| 198 MESODERMAL DERIVATIVES
| |
| | |
| tion is the ventral mesohepar (Fig. 126B). The primordia of the
| |
| pancreas lie originally in the dorsal and ventral mesenteries,
| |
| respectively, but with the rotation of the stomach all are included
| |
| in the dorsal mesentery. The peritoneal supports of the nephric
| |
| and genital organs will be considered in the following section.
| |
| The spleen (see page 224) arises in the mesogastrium, close to the
| |
| wall of the alimentary canal, and is probably mesodermal in
| |
| origin.
| |
| | |
| Later divisions of the coelom. — The coelom becomes divided
| |
| into an anterior pericardial cavity surrounding the heart, and
| |
| a posterior abdominal cavity surrounding the viscera, by the
| |
| septum transversum, a transverse partition which grows out
| |
| from the bridge of mesoderm surrounding the vitelline veins
| |
| | |
| ( Coe nos
| |
| | |
| ‘cavity
| |
| | |
| Pericardial
| |
| cavity
| |
| | |
| Liver
| |
| | |
|
| |
|
| |
|
| |
|
| |
| | |
| fx-+- Peritoneal
| |
| cavity
| |
| | |
| Abdominal
| |
| “cavity |
| |
| | |
| Fie. 127. — Diagrams of coelom and its divisions in A, fish, B, amphibia, reptiles
| |
| and birds, and C, mammals. (After Kingsley.)
| |
| | |
| where they cross the coelom en route from the body wall to the
| |
| heart (Fig. 127A). These cavities are connected during a large
| |
| part of the embryonic period by pericardio-peritoneal canals
| |
| where the septum has failed to unite with the ventral body wall.
| |
| In the amniotes, additional septa develop behind the lungs and
| |
| separate the pleural cavities, which contain the lungs, from the
| |
| remainder of the abdominal cavity, which is now known as the
| |
| peritoneal cavity (Fig. 127B). The pleural cavities are separated
| |
| from each other in the median line by the mediastinum. In the
| |
| mammals (Fig. 127C) the partition separating the lungs from the
| |
| viscera receives musculature from the myotomes and becomes the
| |
| diaphragm.
| |
| | |
| THE FROG (SEE ALSO CHAPTER XI.) — In the frog, the ventral
| |
| mesentery disappears as soon as it has been formed, except in the
| |
| region of the heart and liver. The ventral mesocardium appears
| |
| THE NEPHRIC ORGANS 199
| |
| | |
| before the dorsal mesocardium is formed, and disappears soon
| |
| after, to be followed by the disappearance of the dorsal mesocardium. The ventral mesohepar also has but a short period of
| |
| existence. The septum transversum receives much of its substance from the mesodermal sheath of the liver. No pleural
| |
| cavities are formed.
| |
| | |
| THE CHICK (SEE ALSO CHAPTER xII.) — In the chick, both dorsal and ventral mesenteries are formed. The latter, however, persists only in the region of the fore-gut, and gives rise to the
| |
| mesocardia, which soon disappear; the dorsal mesohepar, which
| |
| becomes the gastro-hepatic omentum, and the ventral mesohepar, which becomes the falciform ligament. The septum transversum is not completed until the eighth day of incubation.
| |
| The pleural cavities are cut off from the pericardial cavities by
| |
| a pleuro-pericardial septum, and from the peritoneal cavity by
| |
| the pleuro-peritoneal septum.
| |
| | |
| MAN (SEE ALSO CHAPTER XIII).— From the first, the pericardial cavity is distinguishable from the abdominal cavity,
| |
| inasmuch as it never communieates directly with the extraembryonic coelom as does the abdominal cavity. As in the chick,
| |
| its posterior boundary is coterminous with that of the fore-gut,
| |
| but it is in communication with the abdominal cavity by means
| |
| of the parietal recesses, passages which correspond to the peritoneo-pericardial canals of the anamniotes. The recesses are
| |
| divided frontally by the vitelline veins into dorsal and ventral
| |
| parietal recesses. With the formation of the septum transversum,
| |
| the ventral recesses are incorporated into the pericardial cavity.
| |
| The dorsal recesses become the pleural cavities; and the pleuroperitoneal septum, which divides them from the peritoneal
| |
| cavity, is formed by the upward growth of the diaphragm. The
| |
| musculature of this organ arises from the fourth cervical myotome
| |
| during the backward growth of the diaphragm. The rotation of
| |
| the stomach results in a rearrangement of the mesenteries, for
| |
| an account of which the reader is referred to Hertwig or Keibel
| |
| and Mall.
| |
| | |
| . B. THE NEPHRIC ORGANS
| |
| | |
| The nephric or excretory system of vertebrates is essentially
| |
| a paired series of tubes (nephridia), developed in the intermediate
| |
| mesoderm, which collect nitrogenous wastes from the blood and
| |
| 200 MESODERMAL DERIVATIVES
| |
| | |
| discharge them to the exterior by two longitudinal ducts emptying
| |
| into the cloaca. The intermediate mesoderm in the anterior
| |
| part of the body is divided into
| |
| nephrotomes corresponding to the
| |
| somites. There are three different
| |
| types of kidneys among the vertebrates (Fig. 128). The first is the
| |
| pronephros, which arises from the
| |
| anterior nephrotomes and is the
| |
| functional kidney in the larval
| |
| stages of the fish and the amphibians. The second is the mesonephros, which arises from nephro
| |
| 9 } Pronephros
| |
| | |
|
| |
| | |
| Mesonephric duct tomes posterior to the pronephros
| |
| and is the functional kidney of
| |
| Metanephros adult anamniotes and embryonic or
| |
| Mefanephric duct f4¢4] amniotes. The third is the
| |
| Cloaca metanephros which is the functional
| |
| | |
| rae, kidney of adult amniotes.
| |
| | |
| Fig. 128. — Diagram to show rela- The pronephros. — This organ is
| |
| tionships of vertebrate excretory formed during development by all
| |
| systems. :
| |
| | |
| vertebrates, but is best developed
| |
| | |
| in larval types like the frog, where it arises from nephrocoels
| |
| (Fig. 129) in the anterior nephrotomes (III, IV, V), the dorsal
| |
| ends of which grow caudally and unite with each other to form
| |
| the pronephric duct which grows backward toward the cloaca.
| |
| The ventral ends of the nephrocoels open into the coelom, and
| |
| these openings, the nephrostomes, become lined with long cilia.
| |
| The tubules meantime elongate and become contorted as they project into the surrounding posterior cardinal vein. Median to each
| |
| nephrostome, the splanchnic mesoderm bulges out and in this
| |
| projection develops a net of capillaries, or glomerulus, which becomes connected with the dorsal aorta. The pronephros is functional, at most, for a short time; and it disappears as the mesonephros develops to replace it.
| |
| | |
| The mesonephros. — The mesonephros, like the pronephros,
| |
| is developed by all vertebrates. It becomes the adult kidney
| |
| of the anamniotes, but is functional during the embryonic (and
| |
| fetal) period only of the amniotes. Portions of the mesonephros
| |
| THE MESONEPHROS 201
| |
| | |
| become associated with the genital organs of the adult (see next
| |
| section).
| |
| | |
| The mesonephros also develops as a series of segmental nephrocoels, but in the nephrotomes posterior to those containing the
| |
| | |
|
| |
|
| |
|
| |
|
| |
| | |
| Primary
| |
| tubule
| |
| | |
| Nephrostomal
| |
| _
| |
| | |
| IP \nc_Nephrostome
| |
| ECS
| |
| | |
| i
| |
| A
| |
| | |
|
| |
| | |
|
| |
| | |
| Fig. 129. — Diagrams showing three stages in the development of the pronephric
| |
| tubule. (After Felix.)
| |
| | |
| pronephric ducts with which they unite (Fig. 130). After the
| |
| degeneration of the pronephros, the tube is known as the mesonephric or Wolffian duct. The ventral ends of the nephrocoels
| |
| acquire coelomic nephrostomes in the anamniotes. In amniote
| |
| development, nephrostomes are seldom formed. A glomerulus
| |
| connected with the dorsal aorta and the cardinal veins arises in
| |
| connection with each tubule, as in the pronephros. An important
| |
| difference between the pronephros and the mesonephros lies in
| |
| the fact that the number of nephric tubules in each nephrotome
| |
| is greater in the mesonephros (Fig. 131). These arise by the
| |
| constriction of the posterior median part of each nephrocoel into
| |
| a small vesicle which gives rise to a secondary tubule; each of
| |
| these tubules acquires a glomerulus and nephrostome at the
| |
| 202 MESODERMAL DERIVATIVES
| |
| | |
| proximal end. The connection of these secondary
| |
| tubules with the Wolffian
| |
| duct, however, is attained
| |
| by an evagination from the
| |
| duct itself which grows out
| |
| as the collecting duct to
| |
| meet the developing
| |
| secondary tubule. From
| |
| these secondary tubules,
| |
| tertiary ones bud off and
| |
| develop in like manner,
| |
| acquiring connections with
| |
| the collecting duct through
| |
| an evagination of this
| |
| canal. As many as eight
| |
| tubules may be formed in
| |
| a single segment by this
| |
| process of budding. This
| |
| complexity is greatest at
| |
| the posterior end of the
| |
| mesonephros. In the amBowman’s capsule niotes, the mesonephros
| |
| | |
| Fig. 130. — Diagrams showing four stages in (except for that portion
| |
| development of mesonephric tubule. (From associated with the genital
| |
| Arey after Felix.) organs) disappears after
| |
| | |
| Mi esonephric duct
| |
| | |
|
| |
|
| |
|
| |
|
| |
| | |
| Anlage of
| |
| mesonephric tubule!
| |
| | |
|
| |
| | |
| the metanephros has
| |
| been formed. v OT Aorta
| |
| | |
| The metanephros. “(WS NN
| |
| — The metanephros,
| |
| which is found as a
| |
| separate kidney only
| |
| | |
|
| |
| | |
|
| |
| | |
| Mesonephric
| |
| | |
|
| |
| | |
| in adult amniotes, iry 3ry 2ry Iry 2ry duct. ry ty
| |
| . . —_—— ened
| |
| | |
| probably is equiva- Collecting Nephrostomes Secretory
| |
| tubules P tubules
| |
| | |
| lent to the posterior
| |
| portion of the meso- F1¢. 181. — Diagram to show origin of secondary and
| |
| | |
| terti . :
| |
| 7 eph ros of the anam- ( ar nary meson yPhne tubules from primary tubules
| |
| | |
| niotes, which it resembles greatly in its organogeny.
| |
| THE METANEPHROS 203
| |
| | |
| The region in which the metanephros arises is, like that in
| |
| which the earlier kidneys are found, the intermediate mesoderm.
| |
| But in the posterior region of the body this mass is never segmented into separate nephrotomes. The first indication of metarephros formation is the appearance of an evagination from the
| |
| dorsal surface of the mesonephric duct near the point at which
| |
| the latter enters the cloaca. This evagination grows dorsally and
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
| 4 5
| |
| Fig. 182. — Diagrams to show origin and development of metanephric tubules.
| |
| Collecting tubule in center, secretory tubules to right and left, the one on the right
| |
| relatively more advanced. (From Arey after Huber.)
| |
| | |
| then turns forward to become the metanephric duct, or ureter, in
| |
| much the same manner as the collecting ducts of the mesonephros
| |
| arose. The metanephric duct then sends out into the nephrogenous tissue evaginations which increase in length and branch
| |
| repeatedly to form the collecting tubules of the metanephros.
| |
| Around the distal end of each tubule, a small mass of the nephrogenous tissue condenses and acquires a lumen like the nephrocoels
| |
| of the pronephros and mesonephros (Fig. 132; 1, 2). From these
| |
| vesicles the secretory nephric tubules arise by a process of elonga204 MESODERMAL DERIVATIVES
| |
| | |
| tion and later fuse with the collecting tubules just described
| |
| (Fig. 132; 3,4). In each of the tubules a capsule develops for the
| |
| reception of a glomerulus which later acquires a connection with a
| |
| branch of the renal artery (Fig. 1382; 4, 5). Development proceeds from the posterior end toward the anterior, instead of in the
| |
| opposite direction as in the earlier types of kidneys. The portion
| |
| of the Wolffian duct nearest to the cloaca is absorbed by it so that
| |
| the ureter has an opening into the cloaca separate from that of the
| |
| mesonephric duct. From the region of the cloaca into which the
| |
| ureters open is formed the urinary bladder and urethra (page
| |
| 193). In mammals, at least, the enlarging bladder includes part
| |
| of the ureter.
| |
| | |
| The later history of the kidneys and their ducts is considered
| |
| in the next section.
| |
| | |
| THE FROG (SEE ALSO CHAPTER XI). — Threc pronephric tubules
| |
| are formed (somites II, III, IV), each with a nephrostome. The
| |
| region of the coelom into which these open is cut off ventrally
| |
| by the development of the lungs and becomes the pronephric
| |
| chamber. The glomeruli soon unite to form a glomus. Before
| |
| metamorphosis the pronephric tubes, and that portion of the
| |
| duct into which they open, degenerate.
| |
| | |
| Mesonephric tubules appear in the nephrotomes (somites VIIXII). These have nephrostomes in early larval life; but at the
| |
| time the pronephroi degenerate the portion of each mesonephric
| |
| tubule next to the nephrostome (peritoneal canal) breaks away
| |
| from the remainder of the tubule and fuses with the posterior
| |
| cardinal vein. The mesonephros is the functional kidney of the
| |
| adult, and the Wolffian duct, therefore, functions as the ureter.
| |
| | |
| THE CHICK (SEE ALSO CHAPTER xi). — About twelve pronephric tubules arise (somites V-XVI) beginning on the second day of
| |
| incubation. Nephrostomes are formed, but glomeruli do not
| |
| appear until the third and fourth days of incubation, at which
| |
| time the pronephros is degenerating. The pronephric duct arises
| |
| at the ninth somite.
| |
| | |
| Mesonephric tubules arise from the intermediate mesoderm
| |
| between somites XII and XXX, the more anterior of which
| |
| develop nephrostomes. The main part of the mesonephros, however, arises between somites XX and XXX, where the continued growth of the tubules causes the projection of this region
| |
| THE GENITAL ORGANS 205
| |
| | |
| into the coelom as the Wolffian body. It is extremely doubtful
| |
| whether the mesonephros ever functions as a kidney, as it begins
| |
| to degenerate on the eleventh day.
| |
| | |
| The metanephros arises on the fourth day of incubation, from
| |
| two primordia as usual, the intermediate mesoderm in somites
| |
| XXXI-XXXIII, and an evagination of the mesonephric duct,
| |
| comparable to the collecting ducts of the mesonephric tubules,
| |
| which becomes the ureter, pelvis, and collecting ducts.
| |
| | |
| MAN (SEE ALSO CHAPTER XIII). — Pronephric tubules arise in
| |
| somites VII—XIII, develop nephrostomes and glomeruli, but
| |
| degenerate rapidly.
| |
| | |
| Mesonephric tubules appear in the intermediate mesoderm
| |
| between the sixth cervical and fourth lumbar segments, but
| |
| those of the cervical and thoracic segments soon degenerate.
| |
| Nephrostomes are formed by the more anterior tubules but have
| |
| only a transitory existence. The mesonephros does not function
| |
| as a kidney.
| |
| | |
| The metanephros has a double origin as in the chick.
| |
| | |
| C. THE GENITAL ORGANS
| |
| | |
| The genital organs may be grouped into two classes: (1) the
| |
| primary genital organs, or gonads, in which the germ cells develop; and (2) the accessory genital organs, whose original function is the discharge of the germ cells from the body.
| |
| | |
| The gonads consist of the germ cells and the subordinate tissues,
| |
| blood vessels, nerves, connective tissue, ete., which make up a
| |
| large part of these glands. In an earlier chapter it has been
| |
| shown that the primordial germ cells may first appear in the
| |
| endoderm of the gut wall and thence migrate by way of the
| |
| splanchnic mesoderm, dorsal mesentery, and peritoneum to their
| |
| definitive position in a thickening of the peritoneum on the
| |
| mesial side of the nephrotomes. This thickening is called the
| |
| genital ridge (Fig. 133B). A considerable body of evidence is
| |
| accumulating to indicate that germ cells may also arise from the
| |
| cells of the genital ridge itself.
| |
| | |
| The genital ridge is now invaded by mesenchymal cells, and
| |
| projects into the coelomic cavity. In some amphibians, a metameric arrangement corresponding to the myotomes has been
| |
| recorded, but following this the segments unite. The anterior
| |
| 206 MESODERMAL DERIVATIVES
| |
| | |
| and posterior ends of the ridge degenerate, and the middle portion enlarges and is separated by a longitudinal groove from the
| |
| mesonephros so that it hangs in the coelom suspended by a fold
| |
| of the peritoneum, known as the mesorchium in the male or the
| |
| mesovarium in the female. The germ cells have by this time
| |
| become transformed into gonia (Chapter III) and the germ glands
| |
| are known as gonads.
| |
| | |
| Within the gonads, the gonia come to lie in nests, close to
| |
| the germinal epithelium. Tubular outgrowths from the nephric
| |
| | |
| Glomerulus
| |
| Wolffian
| |
| di
| |
| | |
|
| |
| | |
| Fig. 133. — Diagrams to show early development of the gonads in transverse sections. A, testis. B, genital ridge. C, ovary. (After Corning.)
| |
| | |
| tubules of the mesonephros approach these nests. The later
| |
| history of the gonads differs in the two sexes.
| |
| | |
| Testis. — In the male, the nests of spermatogonia become
| |
| tubules which connect with the tubules growing in from the
| |
| mesonephros (Fig. 133A). The testicular parts of these canals
| |
| are known as the seminiferous tubules, the nephric portions as
| |
| the efferent ductules. The walls of the seminiferous tubules are
| |
| composed of spermatogonia and sustentacular cells which act
| |
| as nurse cells to the developing sperm. Between the tubules lie
| |
| partitions of mesenchyme which make up the stroma of the testis
| |
| and contain the interstitial cells, which are supposed to be concerned in the formation of the male hormone. _ It is because of the
| |
| presence of these cells that the testis is sometimes spoken of as
| |
| the “ interstitial gland.” It is now well established that the
| |
| testis secretes a “male”? hormone whose presence in the blood
| |
| has much to do with the male secondary characters. Eventually,
| |
| the tubules become separated from. the surrounding germinal
| |
| OVARY 207
| |
| | |
| epithelium by the development of a mesenchymatous layer called
| |
| the tunica albuginea.
| |
| | |
| Ovary. — In the female, the nests of odgonia become separate
| |
| follicles (Fig. 133C) which never attain connection with the mesonephric tubules. These tubules consequently degenerate. A
| |
| follicle consists of a single o6gonium surrounded by follicle cells
| |
| which may be compared to the sustentacular cells of the male.
| |
| In the mammalian ovary the primary follicle is greatly enlarged
| |
| to form a vesicular (Graafian) follicle (Fig. 184), which protrudes
| |
| | |
| Tunica externa
| |
| | |
| Tunica interna
| |
| | |
| Stratum granulosum
| |
| | |
| Cumulus odphorus Ovum
| |
| | |
| Nucleus
| |
| | |
|
| |
| | |
| Fig. 134. —Section of human vesicular (Graafian) follicle. (From Arey after
| |
| Prentiss.)
| |
| | |
| from the surface of the ovary. The follicle cells multiply and
| |
| secrete a follicular fluid which presses the outer wall (stratum
| |
| granulosum) away from the egg and a layer of follicle cells
| |
| immediately surrounding it. These form a projection (cumulus
| |
| odphorus) into the cavity of the follicle. When ovulation takes
| |
| place the wall of the follicle is ruptured, and the egg,. ‘still surrounded by its investment of follicle cells, now known as the
| |
| corona r | radiata (page 41), is washed out with the_ follicular fluid.
| |
| After ovulation the follicle cells enlarge, ‘multiply, and secrete a
| |
| yellow “substance, the whole forming a corpus luteum. Hisaw
| |
| has identified hormones from corpus luteum which produce
| |
| 208 MESODERMAL DERIVATIVES
| |
| | |
| definite effects on the uterus and other parts of the female body
| |
| associated with pregnancy and parturition. The existence of
| |
| female hormones formed in the ovary is now definitely proved.
| |
| These hormones appear to be formed in the follicles and to be
| |
| quite distinct from the hormones derived from the corpus luteum
| |
| (Hisaw). The tunica albuginea of the ovary develops much later
| |
| than that of the testis but is also of mesenchymal origin.
| |
| | |
| The genital ducts. — The sperms formed in the seminiferous
| |
| tubules of the testis are discharged into the mesonephric tubules
| |
| and thence make their way into the mesonephric duct, which
| |
| accordingly becomes the male genital duct. The ova, on the
| |
| other hand, are discharged directly into the cavity of the coelom
| |
| whence they are received into a new tube, the oviduct, by means
| |
| of an opening, the ostium tubae (abdominale). The mesonephric
| |
| duct is often called the Wolffian duct; the oviduct is frequently
| |
| called the Miillerian duct. Both ducts appear in every embryo
| |
| (Fig. 135A), but the later histories of the two differ according to
| |
| the sex.
| |
| | |
| The Wolffian duct.— In the male (Fig. 135B), the efferent
| |
| ductules toward the posterior end of the series become occluded,
| |
| leaving only a few at the anterior end functional. These lose
| |
| their renal corpuscles and shorten greatly. In the amniotes,
| |
| where the metanephros acts as the functional kidney, this anterior
| |
| group becomes the epididymis, while the more posterior, nonfunctional vestige becomes the paradidymis. The mesonephrie
| |
| duct persists as the deferent duct. At the point where the deferent duct enters the cloaca, there develops a dilation, the seminal
| |
| vesicle. In the female (Fig. 185C), the anterior portion of the
| |
| mesonephros persists as the vestigial epodphoron, and the posterior portion becomes the paroédphoron. Traces of the Wolffian
| |
| duct sometimes persist, as in mammals, where this structure is
| |
| known as Gartner’s canal.
| |
| | |
| The Millerian duct. — This canal arises in the elasmobranchs
| |
| by the constriction of the pronephric duct into two tubes, of
| |
| which the ventral becomes the Miillerian duct, while the dorsal
| |
| tube becomes the Wolffian duct. The opening of the Miillerian
| |
| duct into the coelom, the ostium tubae abdominale, is a persistent
| |
| nephrostome. In all other vertebrates, this duct arises independently of and after the formation of the Wolffian duct, a
| |
| ESTROUS CYCLE 209
| |
| | |
| fact possibly correlated with the delayed functioning of the oviduct as compared with the primary renal function of the Wolffian
| |
| duct. In these vertebrates the duct arises in the mesoderm
| |
| lateral to the Wolffian duct and grows both forward and backward until the abdominal and cloacal openings are formed. It
| |
| is not formed until late in embryogenesis. In the female (Fig.
| |
| 135C), the posterior ends of the ducts are usually dilated as
| |
| | |
|
| |
|
| |
|
| |
| | |
| Epididymus
| |
| | |
| Epodphoron
| |
| | |
|
| |
|
| |
| | |
| Parodphoron
| |
| | |
|
| |
|
| |
|
| |
|
| |
| | |
|
| |
| | |
| B
| |
| | |
|
| |
| | |
| deferens
| |
| Seminal
| |
| vesicle
| |
| Utriculus
| |
| prostaticus
| |
| | |
| Fia. 135. — Diagrams showing origin and early development of genital ducts. A,
| |
| early stage showing mesonephros, gonads, (male on left, female on right) and
| |
| ducts. B, later stage in male, showing in broken lines the structures which degenerate. C, later stage in female. (After Felix.)
| |
| | |
| storage chambers, and not infrequently fuse to form a uterus.
| |
| In the male (Fig. 135B), the Miillerian duct degenerates, but
| |
| vestiges are to be found even in the adult, such as the appendix
| |
| testis and prostatic utricle of man, which represent the anterior
| |
| and posterior ends of the female duct, respectively.
| |
| | |
| Estrous cycle. — Most vertebrates have an annual breeding
| |
| season. Among the mammals, however, the fact that the young
| |
| develop for a longer or shorter period (of gestation) in the uterus
| |
| of the mother is associated with a periodical set of changes in the
| |
| 210 MESODERMAL DERIVATIVES
| |
| | |
| activity of the uterus which are known as the estrous cycle.
| |
| There are three main stages: proestrum, estrus, and anestrum.
| |
| | |
| During the proestrum the blood vessels of the uterine wall are
| |
| congested, and in some animals (dog) there is destruction of the
| |
| uterine wall accompanied by the discharge of blood into the
| |
| cavity of the uterus.
| |
| | |
| In estrus the destructive changes of the proestrum are repaired
| |
| while the cavity itself often contains the secretions of the uterine
| |
| glands and the materials discharged in the preceding period
| |
| (“ uterine milk’’). It is in this period that ovulation usually
| |
| takes place and the wall of the uterus is in the condition most
| |
| favorable for the implantation of the blastocyst. The estrus receives its name from the fact that this is the time in which the
| |
| sexual drive is strongest. If implantation (page 140) and pregnancy do not take place, a condition known as pseudopregnancy
| |
| occurs in some animals (rat, rabbit, etc.). In the closing stages
| |
| of the estrus, the wall of the uterus returns to its normal condition, accompanied in some animals (dog) by slight hemorrhages. This period of repair is distinguished (Marshall) as the
| |
| metestrum.
| |
| | |
| The estrus is succeeded by the anestrum, a name given to the
| |
| interval lasting until the next proestrum commences. In many
| |
| mammals estrus occurs but once during the breeding season, but
| |
| in others it may take place more frequently. The period between
| |
| each estrus and the next proestrum is sometimes known as a
| |
| diestrum in these polyestrous species.
| |
| | |
| There is a considerable difference of opinion among the authorities as to the exact relation between ovulation and menstruation,
| |
| a term applied to the periodic hemorrhages characteristic of the
| |
| female primate. It is assumed that the period of ovulation corresponds to the estrus, but the clinical evidence is not clear as to
| |
| whether the menstrual discharge is comparable to that of the
| |
| proestrum or that of the closing stages of the estrus itself.
| |
| | |
| The external genitalia. — The genital organs so far considered
| |
| are common to all vertebrates and are sometimes spoken of as the
| |
| internal genitalia. External genitals are found only in those
| |
| animals in which fertilization is internal. These organs serve the
| |
| function of transmitting or receiving the sperm at the time of
| |
| copulation. Internal fertilization is a phenomenon which has
| |
| THE EXTERNAL GENITALIA 211
| |
| | |
| been observed in all classes of the vertebrates, but it is characteristic of all amniotes.
| |
| | |
| Although the external genitalia differ in the sexes, they are
| |
| embryologically homologous. Two types are recognized, duplex
| |
| and simplex. In the duplex type, characteristic of the sauropsids, sac-like extensions arise on each side of the cloaca, which
| |
| in the male become the hemipenes or intromittent organ, while
| |
| in the female they remain vestigial.
| |
| | |
| In the simplex type, characteristic
| |
| of mammals, a single median ectodermal prominence arises anterior Genital tubercle
| |
| to the cloacal aperture, to become Phallus
| |
| the phallus (Fig. 136). In the
| |
| male, the phallus enlarges and encloses the greater part of the urogenital sinus. In this way it becomes the penis, while the enclosed Fie. 136.— Diagram to show the
| |
| sinus becomes the penile urethra, °"i#in_of the mammalian external
| |
| genitalia. (After Ielix.)
| |
| In the female mammal, the phallus
| |
| becomes the vestigial clitoris, while the sides of the urogenital
| |
| sinus remain open as the labia minora which guard the opening
| |
| of the urogenital vestibule. At the base of the phallus is a swelling, the genital tubercle, from which labio-scrotal folds arise on
| |
| either side of the urogenital opening. In the male they fuse to
| |
| form the scrotum, an external sac into which the testes descend;
| |
| in the female they remain separate as the labia majora.
| |
| | |
| TABLE 9
| |
| Homo.ocies oF THE MAMMALIAN GentTraL System
| |
| | |
| Anus.
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
| Male Indifferent Female
| |
| Testis Gonad Ovary
| |
| Epididymis Mesonephros Epoéphoron
| |
| Paradidymis Paroéphoron
| |
| Ductus deferens Mesonephric Gartner’s canal
| |
| | |
| (Wolffian) duct
| |
| | |
| Appendix testis Miillerian duct Uterus
| |
| Prostatic utricle Vagina
| |
| Penis Phallus Clitoris
| |
| | |
| Labia minora
| |
| | |
|
| |
| | |
|
| |
| | |
| Scrotum Labio-scrotal swellings | Labia majora
| |
| 212 MESODERMAL DERIVATIVES
| |
| | |
| THE FROG (SEE ALSO CHAPTER XI). — The genital ridges arise
| |
| soon after hatching. Sex can be distinguished at the time when
| |
| the embryo is about 30 mm. in body length. The anterior portion of each genital ridge degenerates and becomes a fat body.
| |
| | |
| The Wolffian duct in the male acquires connection with the
| |
| testis by means of some of the mesonephric tubules (vasa efferentia), and serves as the deferent duct as well as the ureter. A
| |
| seminal vesicle is formed. A rudimentary Miillerian duct appears. In the female the Wolffian duct functions solely as a
| |
| ureter while the Miillerian duct becomes the oviduct.
| |
| | |
| No external genitalia are developed.
| |
| | |
| THE CHICK (SEE ALSO CHAPTER XII).— The genital ridge arises
| |
| with the mesonephros as the urogenital ridge. Of this the
| |
| anterior region gives rise to the gonad on the mesial side. Sex
| |
| is not distinguishable until the seventh day of incubation. In
| |
| the female, the right ovary develops only partially and finally
| |
| disappears.
| |
| | |
| The Wolffian duct becomes the deferent duct, connected with
| |
| the testis by vasa efferentia forming the epididymis. The persisting mesonephric tubules of the posterior region of the mesonephros form a paradidymis. In the female a vestigial epodphoron and parodphoron represent these bodies respectively. The
| |
| Miillerian ducts degenerate in the male without ever acquiring
| |
| a cloacal exit. In the female the right Miillerian duct disappears while the left becomes the oviduct. The shell gland
| |
| appears on the twelfth day of incubation, but the cloacal opening
| |
| is not formed until the hen is six months old.
| |
| | |
| No external genitalia are formed, although hemipenes are
| |
| formed in some other birds.
| |
| | |
| MAN (SEE ALSO CHAPTER XIII).— The genital ridge arises on
| |
| the mesial side of the mesonephros. Sex is not distinguishable
| |
| until after the fifth week.
| |
| | |
| Each Wolffian duct functions as a deferent duct, and both epididymus and paradidymis are formed, as is a seminal vesicle
| |
| at the distal end. In the female, epodphoron and paroédphoron
| |
| are formed, while some portion of the duct itself may persist
| |
| as Gartner’s canal. The Miillerian ducts become the uterine
| |
| tubes, which unite at their posterior ends to form the uterus
| |
| and vagina. The latter is partially closed by a semicircular
| |
| THE ADRENAL ORGANS 213
| |
| | |
| fold, the hymen, where it enters the urogenital sinus. In the
| |
| male, vestiges of the anterior end of each Miillerian duct persist
| |
| as the appendix testis, while the posterior end is represented by
| |
| the rudimentary prostatic utricle. The dilation of the bladder
| |
| results in the inclusion of the ureters (metanephric ducts) in
| |
| its walls. The genital ducts (Wolffian or Miillerian ducts) empty
| |
| into the urogenital sinus posterior to the bladder, in a region
| |
| which constricts to form the urethra. About this develop a
| |
| number of outgrowths which acquire cavities and form the prostate gland in the male, and the para-urethral glands of the female.
| |
| The external genitalia are of the mammalian type.
| |
| | |
| D. THE ADRENAL ORGANS
| |
| | |
| Closely associated with the nephric organs are the mesodermal
| |
| interrenal glands, which frequently become associated with the
| |
| suprarenal glands, of ectodermal origin, to form the so-called
| |
| | |
| Uy
| |
| | |
| Sympathetic @~
| |
| | |
| ganglion
| |
| | |
| Suprarenal @. 2D QO} \
| |
| Inter - ——ep
| |
| renal
| |
| Genital
| |
| ridge
| |
| A
| |
| | |
| Fig. 137. — Diagrams to show the origin of the suprarenal and interrenal components
| |
| of the adrenal gland. A, origin as shown in cross section (after Corning). B,
| |
| condition in amphibia. C, in birds. D, in Tammals. .
| |
| | |
|
| |
| | |
|
| |
| | |
| \
| |
| | |
| Suprarenal
| |
| | |
| Interrenal
| |
| | |
|
| |
| | |
|
| |
| | |
| Interrenal
| |
| | |
| Suprarena}
| |
| | |
| adrenal glands. All are endocrine (or ductless) glands. The
| |
| suprarenal portion of the adrenal forms the powerful hormone
| |
| epinephrin (adrenalin); the interrenal portion secretes a hormone
| |
| known as cortin (Swingle), which is employed in the treatment
| |
| of Addison’s disease.
| |
| 214 MESODERMAL DERIVATIVES
| |
| | |
| The interrenals. — These arise as paired thickenings of the
| |
| splanchnic mesoderm mesial to the nephrocoels. In some of
| |
| the amphibians there are traces of a segmentation which is soon
| |
| lost by fusion. There is no direct connection between the interrenal and the mesonephros. These glands may fuse to form an
| |
| elongate median organ or become associated with the suprarenals.
| |
| | |
| The suprarenals. — Although these glands are found in the
| |
| vicinity of the mesonephros, they originate from the sympathetic
| |
| ganglia (ectodermal) as described in the following chapter. They
| |
| are separate structures in the fish, but unite with the interrenals
| |
| in the tetrapods.
| |
| | |
| The adrenals (Fig. 137). — These compound glands are not
| |
| found in the fish. In the amphibians the suprarenal portion of
| |
| the gland is external to the interrenal portion. In the chick they
| |
| are intermingled. In the amniotes, however, the interrenal substance (cortex) surrounds the suprarenal (medulla).
| |
| | |
| E. THE VASCULAR SYSTEM
| |
| | |
| The vascular system is mesenchymatous in origin. It consists
| |
| of separate cells, the blood corpuscles, floating in a fluid matrix,
| |
| | |
| Blood island Ectoderm Somatic mesoderm Splanchnic mesoderm Blood vessel Blood cells
| |
| | |
|
| |
| | |
| : Enloderm fused to yolh
| |
| | |
| Fia. “138. — Diagrams showing three stages in the development of seplary from
| |
| blood island based on transverse sections of the area vasculosa in a seven somite
| |
| chick. (From Arey.)
| |
| | |
| the blood plasma, in a closed system of interconnected tubes, the
| |
| blood vessels. Some vessels become lined eternally with muscle
| |
| fibers, and in one locality this muscular development gives rise
| |
| THE BLOOD CORPUSCLES 215
| |
| to a pulsating heart by means of which the blood is kept in
| |
| circulation.
| |
| | |
| Origin of the blood-vascular system. — The first indications of
| |
| the vascular system are found in the splanchnopleure as blood
| |
| islands (Fig. 138). In the telolecithal vertebrates this is always
| |
| in the extra-embryonic splanchnopleure. These blood islands
| |
| originate as local aggregates of mesenchyme. Later, the inner
| |
| cells separate as corpuscles, while the outer ones form the endothelial lining of a vesicle. These vesicles anastomose with each
| |
| other to form the extra-embryonic vitelline circulation.
| |
| | |
| The blood corpuscles. — The first corpuscles formed are the
| |
| inner cells of the blood islands. Later corpuscles are budded off
| |
| | |
| . (" 2
| |
| @ “SoC
| |
| | |
| °e@ ef
| |
| | |
| Fie. 139. — Stages in the development of human red blood corpuscles. A, hemoblasts. B, megaloblasts (anamniote type). C, D, normoblasts (sauropsid type).
| |
| E, normoblasts in process of becoming F, erythrocytes. (From Arey after Prentiss.)
| |
| | |
|
| |
| | |
|
| |
| | |
| from the walls of the capillaries into their cavities. Mesenchymal cells in regions where the capillary network is forming
| |
| may develop into blood corpuscles and enter the blood stream.
| |
| These first corpuscles are the hemoblasts (Fig. 139).
| |
| | |
| Hemoblasts become differentiated into the different types of
| |
| blood corpuscles in the following blood-forming centers: (1)
| |
| the yolk sac; (2) the embryonic capillaries; (3) the liver, the
| |
| spleen, and the lymph glands; (4) the bone marrow. In the
| |
| adult the lymph glands give rise to lymphocytes, and the bone
| |
| marrow to all types of corpuscles.
| |
| | |
| The erythrocytes, or red corpuscles, are distinguished by the
| |
| presence of hemoglobin which gives them their color. In the
| |
| 216 MESODERMAL DERIVATIVES
| |
| | |
| anamniotes the erythrocytes have a large vesicular nucleus with
| |
| granular chromatin and a distinct cell membrane. In the sauropsida, the erythrocytes have a small compact nucleus. The
| |
| mammalian erythrocyte is distinguished by the absence of the
| |
| nucleus in the adult. In the development of mammals there is a
| |
| succession of erythrocytes: first the anamniote type; then the
| |
| sauropsid type; and finally the mammalian erythrocyte, which
| |
| is produced by the extrusion of the nuclei from the blood cells
| |
| of the sauropsid type (Fig. 139).
| |
| | |
| The leucocytes, or white corpuscles, are of many types, for
| |
| a discussion of which the reader is referred to the textbooks on
| |
| histology. The preponderance of evidence indicates that these,
| |
| like the erythrocytes, are derived from the hemoblasts.
| |
| | |
| Origin of the intra-embryonic vessels. — The first embryonic
| |
| blood vessels (Fig. 140) are the vitelline veins which appear at the
| |
| ventro-lateral margins of the fore-gut. These vessels unite in
| |
| the region of the anterior intestinal portal to form the heart, then
| |
| separate as the ventral aortae, which bend up around the pharynx
| |
| in the mandibular arch as the first aortic arches, and continue
| |
| backward as the dorsal aortae. These fuse at a very early stage as
| |
| the dorsal aorta, from which branches are sent to each myotome
| |
| and to the vitelline circulation. The posterior ends of the vitelline veins fuse in small-yolked forms, such as the frog, to form a
| |
| subintestinal vein which continues back to the tail. In largeyolked forms like the chick, the vitelline veins are widely separated
| |
| and brought into connection only by the sinus terminalis which
| |
| makes a circuit of the area vasculosa. The vitelline veins are
| |
| the ventral venous channels of the splanchnic circulation. <A
| |
| dorsal set of vessels soon originates independently to form the
| |
| somatic venous circulation. The first of these to appear are the
| |
| anterior cardinal (precardinal) veins of the head. A similar pair,
| |
| the posterior cardinal (postcardinal) veins, arise in connection
| |
| with the nephric region. These, however, do not discharge their
| |
| contents directly into the heart but into the anterior cardinals.
| |
| The portions of the original anterior cardinals proximal to this
| |
| juncture with the posterior cardinals are now called the common
| |
| cardinal veins.
| |
| | |
| The heart. — Although the heart is primitively a paired organ,
| |
| we have seen that the two primordia are soon fused into a single
| |
| 217
| |
| | |
| THE HEART
| |
| | |
|
| |
|
| |
|
| |
| | |
| Dorsal aorta
| |
| | |
|
| |
| | |
| Aortic arch
| |
| Ventral aorta
| |
| | |
|
| |
|
| |
| | |
| Vitelline Vitelline
| |
| vein A area vasculosa
| |
| | |
| Caudal
| |
| artery
| |
| | |
|
| |
|
| |
|
| |
|
| |
| | |
|
| |
|
| |
| | |
| osterior cardinal vein
| |
| | |
| Aortic arches
| |
| Anterior cardinal vein
| |
| | |
|
| |
|
| |
| | |
| Internal
| |
| Common cardinal vein
| |
| | |
| carotid
| |
| artery
| |
| | |
|
| |
|
| |
| | |
|
| |
|
| |
|
| |
|
| |
| | |
|
| |
| | |
| Ventral
| |
| aorta
| |
| | |
| External Vitelline
| |
| carotid vein
| |
| artery
| |
| | |
| Internal carotid
| |
| External carotid
| |
| | |
| I y_S
| |
| c—~\
| |
| —
| |
| (EAI ESXJ
| |
| ¢ IAN
| |
| | |
| CS Anterior cardina)
| |
| Dorsal aorta
| |
| Common cardinal
| |
| . | ==— Vitelline vein
| |
| 4 Posterior cardinal
| |
| Vitelline artery —= _
| |
| C
| |
| | |
| Fig. 140. — Diagrams to show fundamental plan of embryonic circulation. A,
| |
| early stage in side view. _B, later stage in side view. C, same from above, aortic
| |
| | |
| Ventral aorta
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
| roots pulled apart.
| |
| 218 MESODERMAL DERIVATIVES
| |
| | |
| median tube connected with the ventral aortae in front, and the
| |
| vitelline veins (and later the common cardinals) behind. Around
| |
| the endocardial lining there develops a coat of muscle fiber which
| |
| later becomes striated to form the myocardium. Outside this
| |
| is a lining of splanchnic mesoderm which forms the epicardium,
| |
| continuous with the lining of a part of the coelom surrounding
| |
| the heart, which will later be cut off by the septum transversum
| |
| to form the pericardium. In this the heart is suspended by a
| |
| dorsal and a veéritral mesentery known respectively as the dorsal
| |
| and ventral mesocardia.
| |
| | |
| The later history of the heart is one of growth and subdivision
| |
| into special chambers. Because the local growth of the heart
| |
| is limited by the anterior and posterior walls of the pericardium
| |
| | |
|
| |
| | |
| D E
| |
| | |
| Fie. 141. — Diagrams to show early stages in development of vertebrate heart.
| |
| A, paired heart tubes. B, same fused. C, primary flexure. D, later ‘‘S”’ stage.
| |
| E, after antero-dorsal displacement of atrium.
| |
| | |
| and by the mesocardia in which it is suspended, any extension
| |
| in length must be accompanied by coiling. The primary flexure
| |
| of the heart is toward the right, thus changing the shape of the
| |
| organ from a straight tube to a C-shaped one. Further growth
| |
| results in the twisting of the heart into the shape of an S. Still
| |
| later, the original posterior loop of the S is pushed forward and
| |
| dorsad so that it comes to lie above the morphologically anterior
| |
| end (Fig. 141).
| |
| | |
| The original chambers of the heart are produced by local
| |
| dilations, of which the most posterior is the sinus venosus; next to
| |
| this is the atrium; in front of this, the ventricle; and finally,
| |
| the bulbus arteriosus. The sinus is the chamber into which the
| |
| primitive veins enter; the atrium is a thin-walled distensile
| |
| THE ARTERIES 219
| |
| | |
| chamber; the ventricle is a thick-walled, muscular, pulsating
| |
| pump; and the bulbus is the chamber from which the blood
| |
| efitéts the primitive arteries.
| |
| | |
| These chambers undergo different changes in the various types
| |
| of vertebrates. Of these, the most important is a progressive
| |
| differentiation, completed in the mammals and birds, of the
| |
| atrium and ventricle into separate right and left halves, of which
| |
| the right side receives venous blood from all parts of the body
| |
| and transmits it to the lungs for respiratory exchange. From
| |
| the lungs the blood is returned to the left side of the heart and
| |
| thence conveyed to all parts of the body.
| |
| | |
| The arteries (Fig. 142). — The ventral aortae fuse into a single
| |
| median tube sending branches into each of the visceral arches.
| |
| | |
|
| |
|
| |
| | |
|
| |
|
| |
| | |
| Anterior
| |
| mesenteric
| |
| | |
| Fig. 142. — Diagrams to show principal arteries; A, in side view, B, cross section
| |
| through mesenteric.
| |
| | |
| These branches, which unite with the dorsal aortae, are usually
| |
| six in number and are known as the aortic arches. Anterior to
| |
| these the ventral aortae continue forward as the external carotid
| |
| arteries. Similar forward extensions of the dorsal aortae are
| |
| known as the internal carotid arteries. In the region of the
| |
| aortic arches the dorsal arteries remain separate as the aortic
| |
| roots (radices aortae). Behind them, as has been mentioned, the
| |
| paired vessels fuse as the median dorsal aorta.
| |
| 220 MESODERMAL DERIVATIVES
| |
| | |
| The aortic arches. — In larvae breathing by means of external
| |
| gills, a loop from each aortic arch grows out into the gill developing on the visceral arch with which it is associated. These loops
| |
| are short-circuited when the external gills disappear.
| |
| | |
| In forms with internal gills, each aortic arch breaks up into
| |
| capillaries in the demibranch and becomes divided into a ventral
| |
| afferent branchial artery and a dorsal efferent branchial artery.
| |
| | |
| In vertebrates with a pulmonary respiration, aortic arches I
| |
| and II, in the mandibular and hyoid arches, respectively, disappear. Arch III, in the first branchial arch, persists as the
| |
| connection between the internal and external carotid arteries,
| |
| | |
| Internal External
| |
| carotid carotid
| |
| | |
|
| |
|
| |
|
| |
|
| |
| | |
| I
| |
| clgy TT
| |
| Il
| |
| | |
| IV
| |
| | |
| Vv Subclavian
| |
| VI
| |
| | |
|
| |
| | |
|
| |
| | |
| Pulmonary
| |
| arteries
| |
| | |
| LEFT
| |
| | |
|
| |
| | |
| B Cc D
| |
| | |
| Fig. 143. — Diagrams of aortic arches. A, hypothetical primitive type. B, in
| |
| frog. C, in chick. D, in man. (After Kingsley.)
| |
| | |
| while the dorsal aorta between arches III and IV disappears.
| |
| Arch IV becomes the systemic arch connecting the dorsal and
| |
| ventral aortae (Fig. 143B). In birds (Fig. 143C), the arch on the
| |
| left side disappears; in mammals (Fig. 143D), that on the right
| |
| degenerates. Arch V is greatly reduced and frequently disappears or has at most a vestigial and transitory existence. From
| |
| arch VI there grow back to the lungs the pulmonary arteries.
| |
| The portion of the sixth arch distal to the pulmonary arteries
| |
| is reduced in caliber and is known as the ductus arteriosus. It
| |
| becomes occluded and degenerates in all the amniotes except
| |
| some few reptiles.
| |
| | |
| Intersegmental arteries. — From the dorsal aortae are given
| |
| off small branches between the myotomes (Fig. 142B). Some of
| |
| these intersegmental arteries persist as the intervertebral arteries. The more anterior ones becomes united on either side by
| |
| THE VITELLINE VEINS 221
| |
| | |
| a dorsal longitudinal vertebral artery. These vertebrals subsequently fuse to form an anterior basilar artery which divides behind the pituitary, the two halves uniting with the internal
| |
| carotid on either side. The posterior halves of the vertebral
| |
| arteries fuse to form the spinal artery which runs back beneath
| |
| the spinal cord. In the region where the anterior limb buds are
| |
| developing, intersegmental arteries grow out, to give rise to the
| |
| subclavian arteries. Similarly, in the region of the pelvic limb
| |
| buds, intersegmental arteries give rise to the iliac arteries. In
| |
| the amniota, the allantoic arteries grow out from the iliac arteries
| |
| into the walls of the allantois. These become so important
| |
| that for some time it appears as though the iliac arteries were
| |
| derived from the allantois instead of the reverse. These allantoic arteries, which degenerate at the time of birth, are known as
| |
| the umbilical arteries in mammals as they traverse the umbilical
| |
| cord and supply the placenta.
| |
| | |
| Other important intersegmental arteries become the renal
| |
| arteries of the kidneys and the genital arteries of the gonads.
| |
| | |
| Mesenteric arteries. — From the dorsal aorta, a number of
| |
| ventral branches, originally paired, but soon fused to become
| |
| median vessels, pass down the dorsal mesentery. They unite
| |
| with the capillaries of the yolk sac which they supply with blood.
| |
| Later, some of them develop branches over the alimentary canal
| |
| which persist after the loss of the yolk sac as the coeliac and
| |
| mesenteric artcries.
| |
| | |
| The veins. — There are two primitive venous systems: the
| |
| somatic system, comprising the cardinal veins; and the splanchnic,
| |
| including the vitelline (omphalomesenteric) and, in amniotes, the
| |
| allantoic (umbilical) veins. The cardinal veins are replaced by
| |
| caval veins; the vitelline veins become transformed into a
| |
| hepatic-portal system. The allantoics disappear at hatching
| |
| (or birth). Finally, there are the pulmonary veins. In general,
| |
| the history of these transformations may be summed up in the
| |
| statement that the primitive independent venous systems become
| |
| transformed into a system wherein an accompanying vein is
| |
| developed for every artery.
| |
| | |
| The vitelline veins (Fig. 144). — These vessels, and their continuation, the subintestinal vein (in small-yolked forms), are the
| |
| first vessels formed in the embryo. In the amniotes, two veins
| |
| 222 MESODERMAL DERIVATIVES
| |
| | |
| grow out from these into the wall of the allantois to become the
| |
| allantoic veins of the sauropsida (umbilicals of mammals). In
| |
| man, however, the umbilical veins actually appear before the
| |
| vitelline veins.
| |
| | |
| It has been noted previously that the vitelline veins pass
| |
| around the liver on their way to the heart. As the liver enlarges,
| |
| it surrounds the vitelline veins, and these become broken up in
| |
| the liver tissue to form a great capillary network. In the amniota, the allantoic (umbilical) veins are similarly absorbed. The
| |
| proximal portions of the vitelline veins, from the liver to the sinus
| |
| venosus, are now known as the hepatic veins; the distal portions
| |
| | |
| 64 Common
| |
| ardinal
| |
| p
| |
| | |
|
| |
| | |
|
| |
| | |
| Ductus
| |
| { venosus
| |
| ‘
| |
| | |
| Fig. 144. — Diagrams to show three stages in the development of the hepatic-portal
| |
| venous system, based on conditions in man. (After Hochstetter.)
| |
| | |
| are called the portal veins. Of the umbilical veins, the right degenerates; the left for a time maintains a direct connection
| |
| through the liver to the hepatic veins, known as the ductus
| |
| venosus. This connection disappears at the time of birth. After
| |
| the disappearance of the yolk, the portal vein and its tributaries,
| |
| of which the most important is the mesenteric vein, carry blood
| |
| from the digestive canal to the liver.
| |
| | |
| The anterior cardinal veins. — The original plan of the cardinal
| |
| system is that of an H in which the upper limbs represent the
| |
| anterior cardinals; the cross-bar the common cardinals, with
| |
| the heart in the middle of the cross-bar; and the lower limbs
| |
| represent the posterior cardinals (Fig. 145). The anterior cardinals arise as a drainage system for the blood passing into the head
| |
| from the carotid arteries.
| |
| | |
| The anterior cardinals are often called the internal jugular
| |
| THE POSTERIOR CARDINALS 223
| |
| | |
| veins. From these, parallel veins, known as the external jugular
| |
| veins, branch off in the ventral region of the head. Veins from
| |
| the vertebral region (vertebral veins) and from the pectoral appendages (subclavian veins) soon develop. In most vertebrates
| |
| the common cardinals and the proximal portion of the anterior
| |
| cardinals, i.e., up to the point where these tributary veins diverge, persist as the precaval veins. In some mammals, a crossconnection is formed between the anterior cardinals, after which
| |
| the portion of the left anterior cardinal, proximal to the anasto
| |
|
| |
| | |
| Anterior.
| |
| cardinal Anterior
| |
| | |
| anastomosis
| |
| | |
| Pre cava ot} Coronary
| |
| | |
| sinus
| |
| Common
| |
| cardinal
| |
| Post
| |
| | |
| Sub - cava
| |
| | |
| cardinal
| |
| | |
| Post
| |
| cardinal]
| |
| | |
| Kidney
| |
| | |
|
| |
| | |
| A
| |
| | |
| Fia. 145. — Diagrams to show three stages in the development of the caval venous
| |
| system. Generalized (supra-cardinals omitted).
| |
| | |
| mosis, and the left common cardinal become the coronary vein
| |
| draining the wall of the heart. The corresponding vessels on
| |
| the right side persist as the precaval (anterior caval) vein.
| |
| | |
| The posterior cardinals. — Each posterior cardinal lies dorsal
| |
| to the mesonephros which it drains. Beneath each mesonephros
| |
| is developed a subcardinal vein. In the anamniotes these veins
| |
| arise as tributaries of the posterior cardinals, returning blood from
| |
| the tail where they are united to form the caudal vein. Later,
| |
| they lose direct connection with the parent vessels and return
| |
| blood from the tail region to the mesonephros as the renal-portal
| |
| veins. The posterior portions of the subcardinals fuse as the
| |
| interrenal vein, which acquires a secondary connection with the
| |
| 224 MESODERMAL DERIVATIVES
| |
| | |
| hepatic vein, and persists as the postcaval vein. In the amniotes the postcaval vein is a complex which arises partly from
| |
| the hepatic veins, partly from appropriated portions of the
| |
| posterior cardinals and subcardinals, and partly from the supracardinals, a pair of vessels dorso-mesial to the posterior cardinals.
| |
| It eventually replaces the posterior cardinals, so that the only
| |
| blood vessels entering the right side of the heart are (1) the precaval vein returning blood from the head, pectoral region, and appendages; and (2) the postcaval vein returning blood from the
| |
| trunk and pelvic appendages as well as all blood from the digestive
| |
| canal conveyed by way of the hepatic-portal system.
| |
| | |
| The pulmonary veins. — These enter the left atrium and are
| |
| new vessels which grow backward from the heart to the developing
| |
| lungs.
| |
| | |
| The lymphatic system. — This system serves to return to the
| |
| veins the blood plasma which has escaped from the capillaries
| |
| (Fig. 146). It contains white blood corpuscles of the ameboid
| |
| type (lymphocytes) which have the power of making their way
| |
| through the capillary walls. The lymphatics apparently originate as intercellular spaces in mesenchyme which later become
| |
| confluent and acquire a limiting endothelium. Like the blood
| |
| vessels, the lymphatic capillaries anastomose and form larger vessels which drain into the veins. The walls of these central vessels
| |
| are often muscular, and localized areas known as lymph hearts
| |
| are found. So, too, localized distensible sacs, the lymph sacs, are
| |
| notunknown. Some of these become lymph glands. The spleen,
| |
| already alluded to in the section on mesenteries (page 198), is a
| |
| hemolymph gland in which both lymphocytes and erythrocytes
| |
| are proliferated.
| |
| | |
| THE FROG (SEE ALSO CHAPTER XI). — In the frog (Fig. 147),
| |
| the primordia of the vitelline veins first appear and grow together as a loose aggregate of cells in front of the liver. Around
| |
| this the coelom grows in from right and left to form the pericardium. Meantime the primordium of the heart endocardium
| |
| develops from the loose aggregate of cells referred to above.
| |
| The inner wall of the coelom (splanchnic mesoderm) becomes the
| |
| myocardium. The atrium is divided by an interatrial septum
| |
| into two auricles, right and left. The ventricle remains a single
| |
| chamber.
| |
| THE FROG, : 225
| |
| | |
| Superficial lym phatics
| |
| | |
| Jugular lymph sac
| |
| | |
| Subclavian lymph sac
| |
| oly
| |
| as ao] Lymph gland
| |
| | |
| yi) Deep lymphatics
| |
| 4 /// Thoracic duct
| |
| ‘ i Retroperitoneal lymph sac
| |
| YA Cisterna chyli
| |
| | |
| Posterior lymph sac
| |
| Superficial lymphatics Lymph gland
| |
| | |
|
| |
| | |
|
| |
| | |
| Fig. 146. — Reconstruction of primitive lymphatic vessels in human fetus of two
| |
| months. (From Arey after Sabin.)
| |
| | |
| Arteries External Aortic
| |
| | |
|
| |
| | |
| : Ventral
| |
| carotid __ arches
| |
| : fiver 2orta Dorsal
| |
| Internal; | WIWIVV VI - aorta
| |
| | |
| carotid ; ;_ Vitelline + Caudal
| |
| | |
|
| |
| | |
|
| |
|
| |
|
| |
|
| |
| | |
|
| |
| | |
|
| |
| | |
| ‘ Caudal
| |
| | |
|
| |
| | |
|
| |
| | |
| Heart :,. ’
| |
| | |
| Anterior > ey :
| |
| cardia Posterior. *Vitelline
| |
| Veins cardinal cardinal"
| |
| | |
|
| |
| | |
| Subintestinal
| |
| | |
| Fie. 147. — Diagram of embryonic vascular system of early tadpole. (After
| |
| Kingsley.)
| |
| 226 MESODERMAL DERIVATIVES
| |
| | |
| Aortic loops develop in the external gills, corresponding to
| |
| aortic arches III, IV, and V. After the appearance of the internal gills, the ventral limb of the loop becomes the afferent
| |
| branchial artery, while the dorsal limb becomes the efferent
| |
| branchial artery. A similar differentiation takes place in arch VI.
| |
| With the loss of branchial respiration, arch III becomes the proximal portion of the carotid arteries, arch IV the systemic arch
| |
| which persists on both sides, and arch V disappears, while from
| |
| arch VI arise vessels which carry blood to both the lungs (pulmonary arteries, Fig. 143B) and skin (cutaneous arteries).
| |
| | |
| The vitelline veins anterior to the liver fuse to become the
| |
| hepatic vein: posterior to the liver, the right vitelline vein disappears, the left becomes the hepatic-portal vein. The anterior
| |
| cardinal veins become the internal jugular veins; the common
| |
| cardinals become the precaval veins. The posterior cardinal
| |
| veins fuse between the mesonephroi, and a new vein grows back
| |
| from the hepatic vein to the right posterior cardinal, to form the
| |
| postcaval vein. The posterior cardinals, anterior to their junction with the postcaval, degenerate. Posterior to this junction
| |
| they persist as the renal-portal veins carrying blood from the
| |
| iliac veins to the kidneys.
| |
| | |
| THE CHICK (SEE ALSO CHAPTER XII). — In the chick (Fig. 148),
| |
| the endocardium of the heart arises as the forward extension of
| |
| the vitelline veins, which soon fuse as the pericardial | primordia
| |
| are brought together beneath the head. The myocardium is
| |
| formed as in the-frog. The right and left halves of the heart
| |
| are completely separated by three septa: the septum aorticopulmonale, which divides the bulbus into a chamber on the right
| |
| ‘leading to the pulmonary arteries and one to the left leading to
| |
| the dorsal aorta; the interventricular septum, which divides the
| |
| ventricle; and the interatrial septum, which divides the atrium
| |
| into two auricles. This separation is completed at the end of
| |
| the first week of incubation. The sinus venosus is incorporated
| |
| in the right auricle.
| |
| | |
| Six aortic arches are formed: I and II disappear on the third
| |
| and fourth days of incubation; IIT forms the proximal portion
| |
| of the internal carotid artery; IV disappears on the left side
| |
| but persists as the systemic arch on the right; V disappears; the
| |
| pulmonary arteries arise from VI, but the distal portion of the
| |
| MAN 227
| |
| | |
| right arch remains as the ductus arteriosus until the chick hatches
| |
| (Fig. 143C).
| |
| | |
| The vitelline veins unite behind the sinus venosus to form the
| |
| meatus venosus which later becomes the hepatic vein. The
| |
| mesenteric vein becomes the portal vein, and the vitelline veins
| |
| disappear at hatching. The allantoic veins grow backward from
| |
| the common cardinals to join the capillaries of the allantois;
| |
| the right allantoic degenerates on the fourth day, and the left
| |
| acquires a new connection with the meatus venosus, by way of the
| |
| | |
| 3
| |
| | |
| $432 Aortic _ _ g 2
| |
| | |
| EZES arches Eg gs s 3 a
| |
| <Timmivvvr § 2 a
| |
| | |
| arene EERIUNNVT SS 88 5 2 3
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
| cardinal
| |
| | |
| Veins
| |
| | |
|
| |
| | |
|
| |
| | |
| Common ‘ Allantoie oornne
| |
| | |
| cardinal
| |
| | |
| Posterior
| |
| ardinal “PON ON
| |
| Vitelline | “NS
| |
| | |
| Fie. 148. — Diagram of embryonic vascular system of chick. (After Kingsley.)
| |
| | |
| left hepatic vein. The allantoic vein degenerates at hatching.
| |
| Two precaval veins are formed from the proximal portions of
| |
| the anterior cardinals and common cardinals. The posterior
| |
| caval vein arises from (1) a branch of the meatus venosus which
| |
| grows back to meet the right subcardinal vein, (2) the fused subcardinals which carry blood from the mesonephros, and (38) the
| |
| renal veins which develop in connection with the metanephros.
| |
| The anterior ends of the posterior cardinals disappear, while the
| |
| posterior ends supply the mesonephros and, after its degeneration, the common iliac veins, which pass directly to the postcaval vein.
| |
| | |
| MAN (SEE ALSO CHAPTER XIII). The heart arises in man
| |
| (Fig. 149) much as in the chick; but the subsequent partition228 MESODERMAL DERIVATIVES
| |
| | |
| ing of this organ into right and left halves is more complicated,
| |
| for two atrial septa are formed. The ventricle is separated by an
| |
| interventricular septum, and the bulbus is divided by two septa
| |
| which unite to form the septum aortico-pulmonale. The sinus
| |
| venosus is incorporated in the right atrium.
| |
| | |
| The aortic arches are formed and have the same history as
| |
| those of the chick, with the exception that it is the left fourth
| |
| aortic arch which becomes the systemic arch (Fig. 143D).
| |
| | |
| The anterior portion of the right vitelline vein becomes the
| |
| hepatic vein; the hepatic-portal arises from the posterior portion
| |
| of the vitelline veins anterior to their junction with the mesenteric
| |
| | |
| Postcardinal veins Precardinal veins
| |
| Descending aorte
| |
| | |
|
| |
| | |
| Sinus venosus
| |
| Vitelline veins
| |
| Fia. 149. — Diagram of embryonic vascular system in man: (From Arey after Felix.)
| |
| | |
| vein. The anterior cardinals are united by an anastomosis (left
| |
| innominate vein), and the left common cardinal disappears with
| |
| the exception of the coronary vein. The right common cardinal,
| |
| together with that portion of the anterior cardinal as far as the
| |
| branching of the left innominate, becomes the precaval vein.
| |
| The postcaval vein is a complex formed from (1) a branch of
| |
| the hepatic vein, (2) the anterior portion of the fused subcardinals, (3) part of the fused supracardinals, and (4) the fused
| |
| posterior portion of the posterior cardinals. The anterior portions of the posterior cardinals separate from these veins, unite by
| |
| means of an anastomosis, and drain into the right precaval vein.
| |
| They are then known as the azygos (right) and hemiazygos (left)
| |
| veins. Of the umbilical veins, the left only persists, with a
| |
| SKELETOGENOUS REGIONS 229
| |
| | |
| direct connection through the liver by means of the ductus
| |
| venosus. At birth this duct closes and the umbilical vein dis
| |
| appears.
| |
| F. THE SKELETON
| |
| | |
| The skeleton of vertebrates consists of a system of supporting
| |
| and protecting elements developed from mesenchyme. These
| |
| elements pass through several conditions in later development.
| |
| The primordia of the skeletal elements are preformed in connective tissue. These become transformed into cartilage, a
| |
| process known as chondrification, through the activities of specialized cells, the chondrioblasts. Cartilage in turn is transformed into bone, through the action of osteoblasts, the process
| |
| being known as ossification. Bones that pass through these three
| |
| stages are known as cartilage bones. In the formation of some
| |
| bones, the cartilaginous stage is omitted; these are known as
| |
| membrane bones.| Both cartilage and bone are typically surrounded by a membrane of mesenchyme which is called the
| |
| perichondrium or periosteum, as the case may be. The separate
| |
| elements of the skeleton are connected with each other by ligaments, by cartilage, or in a bony union.
| |
| | |
|
| |
| | |
| Transverse
| |
| septum
| |
| | |
|
| |
| | |
| Sagittal
| |
| septum
| |
| | |
| Fia. 150. — Diagram to show the skeleton-forming regions as seen in the tail region
| |
| of a vertebrate. (After Kingsley.)
| |
| | |
| Skeletogenous regions. — The principal regions where skeleton
| |
| may be formed in the vertebrate body (Fig. 150) are (1) the
| |
| 230 MESODERMAL DERIVATIVES
| |
| | |
| dermis of the skin, (2) the median sagittal planes between the
| |
| myotomes on the dorsal and ventral sides of the body, (3) the
| |
| right and left frontal planes between the dorsal and ventral
| |
| muscle masses, (4) the transverse planes between the myotomes,
| |
| (5) around the notochord, neural tube, and axial blood vessels,
| |
| (6) in the visceral arches, and (7) in the paired appendages.
| |
| Skeletal elements formed in (1) are called the dermal skeleton;
| |
| those formed in (2) to (5), the axial skeleton; those formed in (6),
| |
| the visceral skeleton; and those formed in (7), the appendicular
| |
| skeleton. The skull contains elements from all but the appendicular skeleton.
| |
| | |
| The dermal skeleton. — Among living vertebrates the most
| |
| primitive example of derm bones are the placoid scales (Tig. 151)
| |
| of the eartilage fish which are
| |
| formed in exactly the same
| |
| way as teeth (Chapter VIIT).
| |
| In the dermal skeleton two
| |
| types of bones are distinguished. The investing bones
| |
| (dermal plates) serve to envelop regions of the head
| |
| | |
| scale (Squalus acanthias) to show originof and trunk. The substituting
| |
| | |
| primitive dermal bone. Compare Tig. bones become so closely allied
| |
| | |
| 119. (After Kingsley.) . . .
| |
| | |
| with the cartilaginous bones as
| |
| to become fused with them or even to replace them in ontogeny.
| |
| Many of the cranial bones are of this type. They may be distinguished by the fact that they
| |
| | |
| x * : Seles ee ORS IS SS Ectoderm
| |
| do not pass through a cartilagi- SaaS EE Dermatomé
| |
| | |
| Myotome
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
| nous stage in development.
| |
| The axial skeleton. — The
| |
| primitive axial skeleton is the
| |
| notochord, whose origin has
| |
| been discussed in Chapter V. 1 NTA
| |
| Around this a connective tissue Fig. 152. — Section through sclerotome
| |
| heath is f db h of lizard (Scleporus) to show arcualia.
| |
| snea 1s forme y mesenchy- (After Kingsley.)
| |
| mal cells. The mesenchyme
| |
| from each sclerotome now forms four little blocks, the arcualia
| |
| (Fig. 152), two dorsal to the notochord and two ventral, from
| |
| which the arches and centra of the vertebrae are formed, as well
| |
| THE STERNUM 231
| |
| | |
| as the primordia of the ribs. The posterior arcualia of each somite unite with the anterior arcualia of the succeeding myotome
| |
| to form the definitive vertebra, which thus comes to lie at the
| |
| point of separation between two myotomes. Eight elements are
| |
| thus concerned with a single vertebra: right and left dorsal
| |
| arcualia from the anterior half sclerotome, and from the posterior
| |
| half sclerotome, and the corresponding ventral elements.
| |
| | |
| The vertebrae. — In the prevertebral masses so formed appear
| |
| centers of chondrification, one on each side of the spinal cord
| |
| and one or more below the cord. These form, respectively, the
| |
| neural arch and the centrum of the vertebrae (Fig. 153). In the
| |
| tail region, two centers of chondrification arise below the centrum,
| |
| | |
|
| |
| | |
| FZ NSY
| |
| CZ AIGMSITANY
| |
| USSD
| |
| | |
| Fia. 153. — Section to show ossification centers in human vertebra and ribs. (After
| |
| JXollman.)
| |
| | |
| enclosing the caudal prolongation of the dorsal aorta, and form a
| |
| hemal arch. With the chondrification of the vertebrae the notochord disappears in all but the most primitive vertebrates, persisting only between the vertebrae as nuclei pulposi of the intervertebral discs. Finally the vertebrae become ossified, and the
| |
| spines, zygapophyses, and other differentiations are developed.
| |
| | |
| The ribs. — Except in the caudal region, lateral processes
| |
| arise from the vertebral primordia and grow out into the myosepta. They later become cartilaginous, and finally true bone.
| |
| These are the ribs, of which there are two types, dorsal and ventral, distinguished according to the part of the vertebra from which
| |
| they originate.
| |
| | |
| The sternum. — The sternum, or breast bone, arises in the
| |
| amphibians from the coalescence of two longitudinal bars of
| |
| cartilage, which later articulate with the coracoids of the pectoral
| |
| girdle, but do not come in contact with the ribs. In the amniota,
| |
| 232 MESODERMAL DERIVATIVES
| |
| | |
| the sternum arises from the fusion of the ventral ends of the anterior rib rudiments. In this way there arise two longitudinal
| |
| bars, from which the unpaired sternum
| |
| , <—_Chviele is formed by fusion along the mesial
| |
| line (Fig. 154).
| |
| Ends The skull. — The skull is a complex
| |
| or of skeletal elements, arising from the
| |
| chondrocranium, or primitive cranium
| |
| of cartilage bones, which is derived in
| |
| part from the protective covering of
| |
| the brain and sense organs (neurocranium), and in part from the supporting elements of the visceral arches
| |
| Fia. 154. — Diagram to show ori- (gnlanchnocranium). This is supplegin of mammalian sternum. . .
| |
| (After Kingsley.) mented by numerous investing and
| |
| substituting bones from the original
| |
| dermal skeleton (dermocranium).
| |
| | |
| Neurocranium.— The neurocranium arises from the head
| |
| mesenchyme which, as has been said, cannot be traced to any
| |
| definite somites. In this mass, which completely invests the
| |
| brain and sense organs, definite centers of chondrification appear.
| |
| These masses unite to form the chondrocranium of the cartilage
| |
| fish (Fig. 155). If the notochord be used as a point of orientation,
| |
| | |
|
| |
| | |
| Presternum
| |
| | |
|
| |
| | |
| Mesosterna
| |
| | |
| Sphenolateral
| |
| | |
| Otic capsule a
| |
| 4. Occipital
| |
| | |
| vertebrae
| |
| | |
| Nasal
| |
| capsule
| |
| | |
|
| |
| | |
|
| |
|
| |
| | |
| Visceral arches
| |
| | |
| Fig. 155. — Diagram showing components of chondrocranium (Squalus acanthias).
| |
| (After Kingsley.)
| |
| | |
| on either side of it is found a parachordal bar. In front of each
| |
| of these is a separate rod; these are the trabeculae. Between
| |
| the two parachordals and around the notochord, the basilar plate
| |
| arises as the support of the epichordal brain. The trabeculae
| |
| also fuse in front, to form the ethmoid plate which supports
| |
| OSSIFICATION OF THE CHONDROCRANIUM 233
| |
| | |
| the prechordal brain, but remain separate at their posterior
| |
| ends to form an opening through which the pituitary projects
| |
| downward. In front of the ethmoid plate the trabeculae grow
| |
| forward as the cornua. Dorsal to each trabecula, another longitudinal bar, the sphenolateral, arises. Between these two bars
| |
| the cranial nerves make their way to the exterior.
| |
| | |
| Around each of the major sense organs a cartilaginous capsule
| |
| develops. The olfactory capsules unite with the cornua, ethmoid, and sphenolaterals. The optic capsule rarely develops
| |
| fully, usually persisting in the conncctive tissue stage as the
| |
| sclera of the eyeball. The otic capsule, however, becomes completely chondrified and unites with the parachordals and the
| |
| latero-sphenoids. Between the two otic capsules and sphenolaterals arises a dorsal plate which forms a roof for the brain.
| |
| In the amniotes, one or more neck vertebrae are consolidated with
| |
| the occipital region.
| |
| | |
| The splanchnocranium.— The digestive canal in the head
| |
| region consists of the mouth, oral cavity, and pharynx, the walls
| |
| of the pharynx being penetrated by the visceral clefts. As there
| |
| is no coelom in this region, the lateral mesoderm is not divided but
| |
| gives rise to mesenchyme which foreshadows the cartilaginous
| |
| bars supporting the wall of this part of the body. These visceral
| |
| arches are the mandibular, hyoid, and four (or more) branchial
| |
| arches. The mandibular arch divides into dorsal and ventral
| |
| portions, of which the dorsal portion becomes the pterygoquadrate
| |
| cartilage (upper jaw of cartilage fish) while the ventral portion
| |
| becomes the meckelian cartilage (lower jaw). The hyoid arch
| |
| divides into a dorsal hyomandibular cartilage, and a ventral hyoid
| |
| cartilage which is usually divided into several centers of chondrification. The hyomandibular acts as a suspensory element for
| |
| the jaws in the fish. It is homologized with a bone of the middle
| |
| ear, the columella, in amphibians, and the stapes of mammals
| |
| (see page 269). The hyoid gives rise to the support of the
| |
| tongue. The branchial arches are usually divided into four parts
| |
| and act as gill supports in the anamniota and disappear or become
| |
| laryngeal cartilages in amniota.
| |
| | |
| Ossification of the chondrocranium. — The limits of this text
| |
| will not permit of an enumeration of all the bones formed from
| |
| the chondrocranium (Figs. 156, 157, 158). They may be grouped,
| |
| 234 MESODERMAL DERIVATIVES
| |
| | |
| however, as follows: (1) the occipitals, formed from the occipital
| |
| vertebrae; (2) the sphenoids, arising from the parachordals,
| |
| basilar plate, trabeculae, and latero-sphenoids; (3) the ethmoids,
| |
| | |
| Premaxilla
| |
| | |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
| | |
| Vomer
| |
| Maxilla
| |
| | |
| Ethmoid
| |
| Palatine Parasphenoid
| |
| Orbito sphenoid
| |
| Pterygoid Jugal
| |
| Alisphenoid
| |
| Squamosal
| |
| | |
| J
| |
| | |
| Quadrate +
| |
| Prootic ~ A 5 “
| |
| Opisthotic
| |
| | |
| Supratemporal oo
| |
| Basioccipital
| |
| | |
| Fia. 156. —— Diagram showing components of vertebrate skull, generalized. Ventral
| |
| view. Chondrocranium stippled, dermal elements in outline. (After Kingsley.)
| |
| | |
| from the ethmoid plate and nasal capsule; (4) the otics, from the
| |
| otic capsule. The pterygoquadrate bar gives rise to the pterygoid bones and the quadrate (which in mammals becomes the
| |
| incus of the middle ear). The mcckelian cartilage gives rise to the
| |
| | |
| Premaxilla
| |
| | |
| Interparietal
| |
| | |
| Quadratojugal
| |
| | |
| Squamosal
| |
| | |
|
| |
| | |
| Fig. 157. — Dorsal view of skull diagrammed in Fig. 156.
| |
| | |
| articular bone at its distal extremity. This becomes the malleus,
| |
| another ear-bone, of the mammals. The remainder of the
| |
| meckelian persists as cartilage. In the hyoids and the branchials,
| |
| bones are formed which retain the names of their cartilaginous
| |
| predecessors.
| |
| THE GIRDLES 235
| |
| | |
| The dermocranium (Figs. 156, 157, 158). — The derm bones
| |
| which invest and, to some extent, supplant the elements of the
| |
| | |
| chondrocranium are too numerous to be more than mentioned Premaxilla
| |
| here. The dorsal derm bones
| |
| | |
| are, from front to rear, the nasals, Maxilla
| |
| | |
| frontals, and parietals, together
| |
| | |
| with a number of smaller bones Jugal
| |
| | |
| which appear in variable quantity
| |
| in the different classes. The
| |
| | |
|
| |
|
| |
|
| |
| | |
| Postorbital —}
| |
| | |
| Nasal
| |
| Lachrymal
| |
| | |
| Sclerotics
| |
| | |
| Frontal
| |
| | |
| Postfrontal
| |
| | |
| principal lateral elements, from Squamosal
| |
| front to rear, are the premaxillae, #4708"!
| |
| | |
|
| |
| | |
| Fibula
| |
| | |
| Tibia Ulna
| |
| | |
| Radius
| |
| | |
| Tarsals 09520 Carpals
| |
| ‘0
| |
| | |
| oO oO
| |
| Metatarsals Ui it Metacarpals
| |
| | |
| f ¥V Phalanges
| |
| WY
| |
| 0
| |
| | |
| Fig. 159. — Diagram of appendicular skeleton, tetrapod type, showing homologies of pectoral elements
| |
| above and to left; pelvic
| |
| elements below and _ to
| |
| right. (After Kingsley.)
| |
| | |
| Parietal
| |
| Supratemporal
| |
| Interoccipital
| |
| | |
|
| |
| | |
| Dermoccipital
| |
| | |
| Via. 158. — Lateral view of skull diagrammed in Figs. 156, 157.
| |
| | |
| maxillae, jugals, quadratojugals, and
| |
| squamosals. The floor of the chondrocranium is invested by the parasphenoids,
| |
| palatines, and vomer. The lower jaw is
| |
| invested by a series of bones of which the
| |
| most important is the dentary.
| |
| | |
| The appendicular skeleton. — The simplest forms of appendages, the unpaired
| |
| and paired fins of fish, contain a skeleton
| |
| consisting of parallel cartilaginous rods,
| |
| which are divided into proximal portions,
| |
| basalia, embedded in the body, the distal
| |
| portions, radialia, extending into the free
| |
| appendages. The paired appendages of
| |
| fish are paddle-like fins; in tetrapods they
| |
| are jointed legs. In both, the skeleton is
| |
| divided into a basal girdle and a free appendicular skeleton (Fig. 159).
| |
| | |
| The girdles. — The girdles are in the
| |
| form of inverted arches, of which the
| |
| pectoral girdle is united to the axial skeleton in fish and free in the tetrapods,
| |
| while the pelvic girdle, usually free in
| |
| | |
| fish, is united to the axial skeleton in the tetrapods. Each arch
| |
| 236 MESODERMAL DERIVATIVES
| |
| | |
| typically consists of three portions. The dorsal one in the pectoral girdle is the scapula; in the pelvic girdle it is called the
| |
| ilium. The two ventral elements of the pectoral girdle are the
| |
| precoracoid (anterior) and the coracoid (posterior), while the
| |
| corresponding elements of the pelvic girdle are the pubis and
| |
| ischium. In the shoulder region, the clavicle, a derm bone,
| |
| becomes associated with the pectoral girdle.
| |
| | |
| The free appendages. — The pectoral and pelvic appendages
| |
| are very similar. Each has three segments: proximal, intermediate, and distal. The proximal segment of the pectoral appendage contains one bone, the humerus, while the corresponding
| |
| bone of the pelvic limb is called the femur. The intermediate
| |
| portion of the pectoral limb possesses two bones, the radius and
| |
| ulna; while the corresponding bones of the pelvic appendage are
| |
| the tibia and fibula. The distal segment is divided into three
| |
| regions of which the proximal portion contains nine or ten bones,
| |
| the carpalia of the pectoral appendage, tarsalia of the pelvic.
| |
| The intermediate portion contains five metacarpalia or metatarsalia, respectively. The distal portion contains the free
| |
| phalanges of the fingers or toes.
| |
| | |
| TABLE 10
| |
| | |
| Homo.oaies oF APPENDICULAR SKELETON
| |
| | |
|
| |
| | |
|
| |
| | |
| Pectoral Gencral Pelvic
| |
| ; Girdle
| |
| Seapula | lium
| |
| Procoracoid Pubis
| |
| Coracoid Ischium
| |
| | |
|
| |
| | |
| Free appendage
| |
| | |
| Humerus Femur
| |
| | |
| Radius Tibia
| |
| | |
| Ulna Fibula
| |
| Carpalia Tarsalia
| |
| Metacarpalia Metatarsalia
| |
| Phalanges (I-V) Phalanges (I-V)
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
| Origin of the appendicular skeleton. — All the bones of the
| |
| appendicular skeleton, with the exception of the clavicle, are
| |
| formed from a mesenchymal blastema in the limb buds by the
| |
| appearance of centers of chondrification. The origin of this
| |
| ORIGIN OF THE APPENDICULAR SKELETON 237
| |
| | |
| mesenchyme is probably from the somites, but the details of the
| |
| process are still imperfectly understood.
| |
| | |
| THE FROG.! — Nine vertebrae are formed, of which the first is known as the
| |
| cervical vertebra, or atlas, the succeeding seven are the abdominal vertcbrac, and
| |
| the last is called the sacral vertebra as it is to this that the pelvic girdle is attached.
| |
| No caudal vertebrac are formed, but thrce strips of cartilage enclose the notochord
| |
| and form the primordium of the adult urostyle. Dorsal ribs are differentiated, but
| |
| these remain rudimentary and fuse with the transverse processes of the vertebrae.
| |
| The sternum arises from the fusion of two longitudinal bars of cartilage which never
| |
| attain connection with the ribs. It persists anterior and posterior to the pectoral
| |
| girdle.
| |
| | |
| The cartilage bones of the skull are the exoccipitals, prodtics, stapes, ethmoids,
| |
| and the pterygoquadrate (in part), articulare, mentomeckelian, hyoid, and branchials.
| |
| The derm bones are the fronto-parictal, nasals, premaxillac, maxillac, quadratojugals,
| |
| squamosals, parasphenoid, palatines, vomers, and dentaries.
| |
| | |
| In the pectoral girdle develop the scapula, coracoid, and precoracoid, the last of
| |
| which is replaced by the clavicle. In the pelvic girdle only the ilium and ischium
| |
| ossify. Only four digits are present in the hand, the thumb (pollex) being absent.
| |
| | |
| THE CHICK. — There are sixteen cervical vertebrae, of which the first is the atlas,
| |
| and the second, which has appropriated the centrum of the first, is the axis; five
| |
| thoracic vertebrae; about six lumbar vertebrac; two sacrals; and about fifteen
| |
| caudals. The last thoracic, all lumbars, and sacrals and five caudals are fused to
| |
| the pelvic girdle. The last four caudals are fused into a pygostyle. Dorsal ribs are
| |
| formed by the cervical and the thoracic vertebrae. The sternum arises from two
| |
| longitudinal bars of cartilage which unite in the median line. It is distinguished by
| |
| the development of a large keel (carina) for the attachment of the pectoral muscles.
| |
| | |
| The cartilage bones of the skull are the basioccipital, exoccipitals and supraoccipitals; prodtics, epiotics, and opisthotics; basisphenoid, orbitosphenoids, and
| |
| alisphenoids; the ethmoid; quadrate, articular, meckelian cartilage; stapes, hyoid,
| |
| and branchials. The derm bones are the frontals, parictals, nasals, lachrymals, premaxillac, maxillae, jugals, quadratojugals, squamosals, pterygoids, palatines, parasphenoids, vomer, angular, supra-angular, opercular, and dentary.
| |
| | |
| The pectoral girdle devclops a scapula and coracoid, together with a dermal
| |
| clavicle. Ilium, ischium, and pubis ossify separatcly in the pelvic girdle. Five
| |
| digits are performed in the pectoral appendage; of these the first and fifth fail to
| |
| develop further. Five also appear in the embryonic skeleton of the pelvic appendage;
| |
| the fifth soon disappears, and the first is extremely short and develops no phalanges.
| |
| | |
| MAN. — Seven cervical vertebrae, including the axis and atlas, twelve thoracic,
| |
| five lumbar, five sacral, and four caudal vertebrae are formed. Of these, the sacral
| |
| vertebrae are united to the pelvis, and the caudal vertebrae are frequently fused to
| |
| form the coccyx. Primordia of ribs are formed by all vertebrae except those following the first caudal. Only the thoracic segments, however, develop complete ribs.
| |
| The sternum arises from two longitudinal primordia with which the first eight or
| |
| nine ribs acquire cartilaginous connections.
| |
| | |
| The cartilage bones of the skull are the occipital (in part), the sphenoid, the
| |
| ethmoid, the turbinates, temporals (in part), the stapes, malleus, incus, and hyoid.
| |
| The malleus and incus are the representatives of the articular and quadrate. The
| |
| | |
| 1 The details of the skeleton in this and succeeding paragraphs arc for reference
| |
| only.
| |
| 238 MESODERMAL DERIVATIVES
| |
| | |
| derm bones are the occipital (in part), temporals (in part), frontal, parictals, lachrymals, nasals, vomer, maxillae, zygomatics, palatines, and mandible, the last-named
| |
| bone representing the fused dentaries. It is apparent that many of the bones of the
| |
| human skull are the result of the fusion of separate centers of ossification which
| |
| represent skull elements of the lower vertebrates. The second and third visceral
| |
| arches contribute to the formation of the hyoid, the others to the laryngeal cartilage.
| |
| | |
| The pectoral girdle consists of the scapula, with which is fused the coracoid.
| |
| There is no precoracoid, but a dermal clavicle is present. The centers of ossification
| |
| that represent the pubis, ischium, and ilium fuse to form an innominate bone. The
| |
| free appendages terminate in five digits. In conclusion, it should be mentioned
| |
| that the adult condition of the human skeleton is not attained until the age of
| |
| twenty-five.
| |
| | |
| G. THE MUSCLES
| |
| | |
| The musculature of the vertebrate is derived from mesenchyme (Fig. 160), of which the greater part originates from the
| |
| myotomes and gives rise to striated muscle cells, controlled by the
| |
| central nervous system, the skeletal musculature. A portion,
| |
| | |
|
| |
|
| |
|
| |
|
| |
| | |
| Sclerotome
| |
| | |
| Neural tube
| |
| | |
| Notochord
| |
| Aorta
| |
| | |
| Dorsal
| |
| appendicular
| |
| muscle mass
| |
| | |
| Ventral
| |
| appendicular
| |
| Gut muscle mass
| |
| Splanchnie
| |
| mesoderm
| |
| | |
| Somatic
| |
| mesoderm
| |
| | |
| Fig. 160. — Diagram of transverse section through vertebrate embryo in region of
| |
| limb bud, to show origin of appendicular muscles. (After Corning.)
| |
| | |
| however, originates from splanchnic mesoderm and gives rise to
| |
| non-striated (smooth) muscle cells(found in the skin, surrounding
| |
| the alimentary canal, blood vessels, and the urogenital organs.
| |
| They are/controlled by the autonomic nervous system (page 254),
| |
| and make up the visceral musculature. Several exceptions to
| |
| these general statements should be noted. The muscle cells of
| |
| the heart are striated; the muscles derived from the visceral
| |
| arches are both)striated and controlled by the central nervous
| |
| CRANIAL MUSCLES 239
| |
| | |
| system, although derived from lateral mesoderm. It will be
| |
| noted later that the muscles of the iris of the eye (page 266) and
| |
| of the sweat glands (page 246) are apparently ectodermal in
| |
| origin.
| |
| | |
| Dermal musculature. — In the skin are found striped muscles
| |
| which are derived from skeletal musculature (see below) but
| |
| which have lost their attachment to the skeleton. The dermal
| |
| musculature is best developed in the amniotes. The muscles of
| |
| expression in man are dermal muscles supplied by the seventh
| |
| cranial nerve (see Chapter X).
| |
| | |
| Axial musculature. In this section are included all the
| |
| muscles arising from the myotomes and attached to parts of the
| |
| axial skeleton, which they move. They are originally metameric, but their later history is obscured by subsequent migration, fusion, splitting, budding, and degencration. The intercostals, between the ribs, however, preserve their original metamerism, which in the others may be traced to some extent by the
| |
| innervation, since the connection between a spinal nerve and
| |
| the muscle mass it supplies is established early in organogeny
| |
| and remains constant. Thus it can be shown that the musculature of the diaphragm,
| |
| supplied by the phrenic
| |
| nerve, arises from a cervical myotome.
| |
| | |
| Cranial muscles. —
| |
| Like the cranium, the associated muscles are derived from different
| |
| sources and consist of
| |
| skeletal and _ visceral
| |
| muscles. The muscles
| |
| | |
| of the eyeball arise from Fia. 161.— Head of embryo dogfish (Squalus
| |
| | |
| : _ _ acanthias) showing preotic somites (A, B, C)
| |
| the three preotic myo and cranial nerves (V, VII, [X, X). (After
| |
| tomes (Fig. 161), of which — jingsley.) 7
| |
| | |
| the first supplies all the
| |
| | |
| muscles of the eyeball except the superior oblique, derived from
| |
| the second myotome, and the lateral rectus, supplied by the third
| |
| head myotome. These are innervated by the third, fourth, and
| |
| sixth cranial nerves, respectively. The tongue musculature is
| |
| 240 MESODERMAL DERIVATIVES
| |
| | |
| derived from the myotomes associated with the occipital vertebrae
| |
| and supplied by the twelfth cranial nerve. The muscles of mastication, the facial muscle, and the laryngeal muscles, together with
| |
| those of the ear bones, arise from the visceral arches (Fig. 162),
| |
| | |
| Glossopharyngeal _, Facial
| |
| | |
| ig
| |
| | |
|
| |
|
| |
|
| |
| | |
| Trigeminal
| |
| | |
|
| |
| | |
| ma
| |
| \i Me
| |
| | |
| KS
| |
| | |
| . id Mandibular
| |
| Branchial arches Fiyel arch
| |
| | |
| Fia. 162. — Diagram to show primitive visceral muscles in relation to visceral skeleton and cranial nerves. (Hypothetical after Wilder.)
| |
| | |
| ‘
| |
| | |
|
| |
| | |
| Vagus
| |
| \\\
| |
| i
| |
| | |
|
| |
| | |
|
| |
|
| |
| | |
|
| |
|
| |
| | |
|
| |
|
| |
| | |
|
| |
| | |
| and are supplied by cranial nerves V, VII, [X, X, and XI (see
| |
| Chapter X).
| |
| | |
| Appendicular muscles. — In the anamniotes, these muscles
| |
| arise from the myotomes; among the amniotes, their origin is
| |
| doubtful, as the limb bud develops as an undifferentiated mass of
| |
| mesenchyme surrounded by ectoderm. In this blastemal mass,
| |
| | |
|
| |
|
| |
| | |
| Dorso - medial
| |
| muscle primordia
| |
| | |
| Procorocoid
| |
| | |
| Humerus
| |
| Ventro - lateral
| |
| muscle primordia
| |
| | |
| Fig. 163. — Reconstruction of the pectoral muscle masses in a 17-mm. Necturus:
| |
| (Prepared by H. F. DeBruine.)
| |
| | |
| muscles and bones are laid down, the differentiation proceeding
| |
| from the proximal toward the distal end. The pectoral muscles
| |
| differentiate before those of the pelvic appendage. The appenSUMMARY 241
| |
| | |
| dicular muscles are found in antagonistic groups: protractors,
| |
| which move the limb forward; and retractors, which have the
| |
| opposite effect; levators, which raise the limb; and depressors,
| |
| which contract in the opposite direction. Like the axial muscles,
| |
| these have become highly modified and specialized among the
| |
| tetrapods (Fig. 163).
| |
| | |
| Visceral muscles. — Under this head are included the muscles
| |
| lining the alimentary tract, lungs, vascular organs, and urogenital system. All arise in the mesoderm which surrounds the
| |
| endothelial lining of the organs concerned. The muscle cells of
| |
| the heart arise as smooth muscle cells which become striated in
| |
| later development. It is interesting in this connection that the
| |
| smooth muscle cells of the bladder of the dog have been transformed into what are apparently striate muscles when this organ
| |
| is made to pulsate rhythmically by continued irrigation.
| |
| | |
| SUMMARY
| |
| | |
| The following structures are derived from the middle germ
| |
| layer:
| |
| | |
| A. The notochord
| |
| | |
| B. The mesoderm
| |
| | |
| I. The lateral mesoderm
| |
| Epithelium of the coelom
| |
| Pericardial cavity
| |
| Pleural cavity
| |
| Peritoneal cavity
| |
| | |
| Mesenteries
| |
| Dorsal mesentery
| |
| Ventral mesentery
| |
| Mesocardia
| |
| Mesohepares
| |
| | |
| II. The intermediate mesoderm
| |
| | |
| Kidneys
| |
| Pronephros
| |
| Mesonephros
| |
| Metanephros
| |
| 242 MESODERMAL DERIVATIVES
| |
| | |
| Genitalia
| |
| Gonads
| |
| Genital ducts
| |
| Wolffian (mesonephric) duct
| |
| Miillerian (oviducal) duct
| |
| External genitalia (also ectodermal)
| |
| | |
| Adrenal glands
| |
| | |
| Interrenals
| |
| (Suprarenals from ectoderm)
| |
| | |
| C. The mesenchyme
| |
| III. (Principally from splanchnic mesoderm)
| |
| | |
| The blood corpuscles
| |
| Blood plasma
| |
| Blood vessels
| |
| Heart
| |
| Arteries
| |
| Veins
| |
| The lymphatics
| |
| | |
| IV. (Principally from the axial mesoderm)
| |
| | |
| Connective tissue
| |
| Skeleton
| |
| Dermal
| |
| Axial
| |
| _ Cranial
| |
| Chondrocranium
| |
| Neurocranium
| |
| Splanchnocranium (or visceral
| |
| skeleton)
| |
| Dermocranium
| |
| Appendicular
| |
| Musculature
| |
| Dermal
| |
| Axial
| |
| Cranial
| |
| Appendicular
| |
| Visceral (from splanchnic mesoderm)
| |
| REFERENCES 243
| |
| | |
| REFERENCES
| |
| | |
| Allen, E. 1932. Sex and Internal Seerction.
| |
| | |
| Arey, L. B. 1934. Developmental Anatomy, 3rd Ed., Chaps. 9-13.
| |
| | |
| Brachet, A. 1921. Traité d’embryologie des vertébrés, Part II, Bk. 1, Chap. 2;
| |
| Bk. 2, Chaps. 1+4.
| |
| | |
| Hertwig, O. 1906. Handbuch, Vol. 1, Chap. 5; Vol. 3, Chaps. 1-7.
| |
| | |
| Jenkinson, J. W. 1913. Vertebrate Embryology, Chap. 7.
| |
| | |
| Keibel and Mall. 1910-1912. Human Embryology, Chaps. 11-13, 15, 18, and 19.
| |
| | |
| Kellicott, W. E. 1913. Chordate Development.
| |
| | |
| Kerr, J.G. 1919. Textbook of Embryology, Chaps. 4-6.
| |
| | |
| Kingsley, J.S. 1926. Comparative Anatomy of Vertebrates, 3rd Ed.
| |
| | |
| —— 1925. The Vertebrate Skeleton.
| |
| | |
| Lillie, F. R. 1919. The Development of the Chick, 2nd Ed.
| |
| | |
| MeMurrich, J. P. 1923. The Development of the Human Body, 7th Ed.
| |
| | |
| Vialleton, L. 1924. Membres et ceintures des vertébrés tétrapodes.
| |
| CHAPTER X
| |
| ECTODERMAL DERIVATIVES
| |
| | |
| The ectoderm, being the external germ layer, gives rise to the
| |
| outer layer of the skin, the epidermis, which continues into all
| |
| the openings of the body} Of these, the development of the
| |
| mouth, the cloaca and its derivatives, and the visceral clefts
| |
| has been discussed. There remain for consideration the openings of the nostrils, the chamber of the eye, and the external
| |
| auditory meatus. These will be taken up in connection with the
| |
| sense organs, which, together with the nervous system, form in
| |
| development a sensory-nervous complex.
| |
| | |
| A. THE INTEGUMENT
| |
| | |
| The integument consists of two parts, the ectodermal epidermis, and the mesodermal dermis. The epidermis soon de
| |
| Periderm
| |
| | |
|
| |
| | |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
| | |
| Periderm
| |
| | |
| Stratum intermeditum
| |
| | |
| Stratum germinativum
| |
| | |
| Corium
| |
| | |
| after Prentiss.)
| |
| | |
| laminates into two layers, the deeper germinativum, from which
| |
| new strata are proliferated towards the exterior, and an outer
| |
| | |
| periderm or embryonic skin (Fig. 164). Beneath the periderm,
| |
| 244
| |
| DERIVATIVES OF THE CORNEUM 245
| |
| | |
| the outer cells of the germinativum are transformed into a horny
| |
| layer, the corneum. The underlying dermis is essentially a
| |
| supporting layer of mesenchyme cells derived largely from the
| |
| outer side of the myotome, a region which is sometimes known
| |
| as the dermatome. In the dermis are formed blood vessels, connective tissue, bone, and muscle. The bony scales of fish are
| |
| dermal in origin.
| |
| | |
| Derivatives of the corneum. — In the amniotes the horny layer
| |
| of the epidermis is frequently fragmented to form horny scales
| |
| | |
| Epidermis
| |
| —_—
| |
| | |
| i
| |
| i
| |
| | |
| Germinativum
| |
| h~ Dermis
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
|
| |
| | |
| Germinativum
| |
| Corneum
| |
| | |
|
| |
| | |
| " Blood vessel = i!
| |
| Fiq. 165.— Diagrams showing similar development in A, scale; B, feather; and C,
| |
| | |
| hair. (After Kingsley.)
| |
| | |
| (Fig. 165A), such as those of reptiles, or those found on the legs
| |
| of birds, or the tails of rats, ete. Among the birds, scales are
| |
| largely replaced by feathers which originate in much the same
| |
| | |
| Unguis
| |
| | |
|
| |
| | |
| Al Cy
| |
| | |
| Fia. 166. — Diagrams to show ectodermal primordia of A, nail; B, claw; and C, hoof.
| |
| Above in sagittal sections; below ventral view. (After Kingsley.)
| |
| | |
| manner as scales. The epidermal plate, however, grows down
| |
| like a cup to enclose a core of dermal origin (Fig. 165B). The
| |
| epidermal sheath gives rise to the quill and barbs, while the core
| |
| gives rise to the pulp, by means of which nutriment is supplied
| |
| to the developing feather. Among the mammals, hair arises in a
| |
| 246 ECTODERMAL DERIVATIVES
| |
| | |
| very similar fashion. An epidermal plate grows down into the
| |
| dermis to form the hair bulb, the proximal end of which invaginates to receive a mesodermal core, the hair papilla, while
| |
| around the whole is a mesodermal hair sheath (Fig. 165C). The
| |
| hair papilla, however, does not grow out into the center of the
| |
| hair as does the pulp of the feather. Claws, nails, and hoofs
| |
| arise from the union of two epidermal primordia like those of
| |
| scales, a dorsal unguis and a ventral subunguis (Fig. 166).
| |
| Derivatives of the germinativum. — The germinativum, in addition to producing the more superficial layers of the epidermis,
| |
| gives rise to the glands of the skin
| |
| Unicellular Multicellular (Fig. 167). Among the anamnigland gland .
| |
| — — otes, these glands are usually unicellular and produce the mucus
| |
| which serves to diminish the friction of the skin against the water
| |
| while swimming. Unicellular
| |
| glands frequently aggregate to
| |
| | |
|
| |
|
| |
| | |
| 3 Epi 2 | dermis
| |
| eo:
| |
| | |
|
| |
|
| |
| | |
| Se esa £ produce multicellular glands, such
| |
| RS Sars
| |
| Chromatophore 7 as the flask glands and cement
| |
| | |
| glands of the anamniotes, or the
| |
| sebaceous (oil) and sudoriparous
| |
| (sweat) glands of the mammals.
| |
| The mammary glands of mammals are modified sudoriparous
| |
| glands secreting the milk by which the new born are nourished
| |
| through infancy.
| |
| | |
| Derivatives of the dermis. — Two types of pigmentation are
| |
| to be distinguished in the integument. The first is produced by
| |
| pigment secreted in the ectodermal epidermis, i.e., the melanin,
| |
| of the frog tadpole. The second is produced by chromatophores,
| |
| which are mesenchyme cells of the dermis. These secrete pigment granules and move toward the light to form a layer immediately below the epidermis, some even wandering into the epidermis
| |
| itself.
| |
| | |
| THE FROG. — The ectoderm of the frog embryo is ciliated at
| |
| 6-mm. body length and remains so until the length of 20 mm. is
| |
| attained, when the cilia disappear except on the tail which remains ciliated until metamorphosis. The jaws and oral combs
| |
| of the tadpole are derivatives of the corneum and consist of rows
| |
| | |
| Fig. 167. — Section of Protopterus skin
| |
| to show glands. (After Kingsley.)
| |
| THE NERVOUS. SYSTEM 247
| |
| | |
| of horny denticles forming replacement series. The oral gland,
| |
| or sucker, is a multicellular mucous gland derived from the
| |
| germinativum and elevated by the elongation of its gland cells.
| |
| It arises as a crescentic groove posterior and ventral to the point
| |
| where the stomodeum will appear, then becomes V-shaped, and
| |
| finally divides by the degeneration of the middle portion. The
| |
| cement gland atrophies soon after the opening of the mouth. The
| |
| pigmentation of the skin is derived from two sources, the melanin
| |
| of the egg which is distributed to the epidermis, and the mesenchymal chromatophores (Fig. 199) which develop in the dermis.
| |
| | |
| THE CHICK. — The scales on the legs are typical reptilian scales
| |
| and are derived from the corncum; they sometimes bear feathers
| |
| in the young’bird and so form a transition between scales and the
| |
| characteristic avian feathers. The claws arise in the corneum
| |
| from two primordia, a dorsal “ claw-plate ” and a softer “ clawsole.” To prevent the sharp claws tearing the embryonic membranes, the concavity of the claw is filled with a pad known as the
| |
| neonychium, derived from the corneum, which is lost after hatching. The beak arises from the corneum around the upper and
| |
| lower margins of the jaws. The egg tooth is a horny prominence
| |
| on the dorsal side of the upper jaw, appearing on the sixth day of
| |
| incubation but not taking on its ultimate shape until the fourteenth. It serves to aid in breaking the shell and is lost after
| |
| hatching.
| |
| | |
| MAN. — The nails arise from nail-plates and sole-plates, of
| |
| which the latter are rudimentary structures. They are covered
| |
| during fetal life by the eponychium, consisting of the periderm
| |
| and outer layers of the corneum. The hairs are arranged in
| |
| patterns which have been’ interpreted as reminiscences of the
| |
| ancestral scalés. The first growth of hair is called the lanugo;
| |
| it is cast off, except over the face, soon after birth. The mammary glands arise from two longitudinal thickenings of the
| |
| epidermis, known as the milk ridge. In later development the
| |
| gland resembles an aggregation of sudoriparous glands.
| |
| | |
| B. THE NERVOUS SYSTEM
| |
| | |
| Although the nervous system and sense organs arise together
| |
| and remain in functional continuity, it has become customary to
| |
| distinguish the sense organs (receptors) from the nerves (trans248 ECTODERMAL DERIVATIVES
| |
| | |
| mittors) by which stimuli are passed on to the muscles or glands
| |
| (effectors). | Both the nervous system and the sense organs arise
| |
| from specialized regions of the dorsal ectoderm, knowy respectively as the neural plate and the sense plates (placodes)4 These
| |
| represent an inward growth from the germinativum as opposed
| |
| to the outward growth which produces the epidermis. In the
| |
| frog this division is clearly indicated by a line of cleavage between the outer epidermal ectoderm and the inner nervous ectoderm. Both the neural plate and the sensory placodes withdraw from the surface and become subepidermal by a process of
| |
| invagination. In this connection it is interesting to note that
| |
| the optic placode is incorporated and invaginates with the neural
| |
| plate so that when the retina of the eye develops, it does so from
| |
| the brain. |
| |
| | |
| The neural tube. —'The neural plate is an elongate structure,
| |
| extending from the blastopore to the head region.! Local growth
| |
| results in the incurving of this plate to produce a neural groove
| |
| with conspicuous lips, the neural folds. As this growth continues the groove sinks inward and the lips meet above it, thus
| |
| converting the groove into a neural tube, which breaks away from
| |
| the overlying epidermis and sinks into the interior. The cells
| |
| at the margin of the neural plate form, at each dorso-lateral
| |
| angle of the neural tube, a bar known as the neural crest, which
| |
| subsequently segments into the ganglia.
| |
| | |
| The neurons. — The inner lining of the neural tube, corresponding to the outer layer of the neural plate, is called the
| |
| ependyma. This is the center of cell proliferation (Fig. 170).
| |
| Two types of cells are formed: the supporting cells, or spongioblasts; and the embryonic nerve cells, or neuroblasts. The
| |
| neuroblasts migrate out of the ependyma and form an intermediate mantle layer in which they become transformed into
| |
| neurons. These nerve elements have a prolongation at one
| |
| end known as the axon or nerve fiber, while at the other are
| |
| branched projections called dendrites. The axons grow out
| |
| from the mantle layer into the outer layer of the cord, known as
| |
| the marginal layer, where they secrete the medullary sheaths
| |
| which act as insulating coats. Not all axons become medullated.
| |
| Similar changes take place in the ganglia, whereby neurons and
| |
| supporting cells are differentiated.
| |
| TYPES OF NEURONS 249
| |
| | |
| Types of neurons. — We may distinguish four types of neurons
| |
| (Fig. 168), as follows: (1) Afferent neurons arising in the ganglia
| |
| and sending theiraxons
| |
| to the dorsal region of
| |
| theneuraltube. These
| |
| convey excitations
| |
| from the sensory receptors to the neural tube.
| |
| Two sub-types are distinguished: (a) the somatic sensory neurons,
| |
| conveying excitations
| |
| from the exterior; and
| |
| (b) splanchnic sensory
| |
| neurons, conveying excitations from the viscera. (2) Efferent
| |
| neurons, with their
| |
| bodies in the ventral
| |
| region of the neural
| |
| tube, sending their
| |
| axons to effectors
| |
| (muscles or glands).
| |
| Two sub-types are recognized: (a) somatic
| |
| motor and (b) splanchnic motor. These af- Fig. 168.— Diagram to show cross-sections of the
| |
| ferent and efferent neu- spinal cord at three levels, the posterior level above.
| |
| rons form the periph- The dotted lines indicate the paths of neurons whose
| |
| | |
| bodies lie wholly within the cord, suprasegmental
| |
| to the left.
| |
| | |
|
| |
| | |
| Effector
| |
| | |
| eral nervous system.
| |
| (3) The intersegmental
| |
| neurons have their bodies in the ventral portion of the neural tube
| |
| and their axons are usually directed towards its posterior end.
| |
| They serve to connect efferent neurons in the different segments
| |
| of the body. (4) The suprasegmental neurons have their bodies
| |
| usually in the dorsal portion of the neural tube and their axons are
| |
| directed toward the anterior end of the tube, i.e., the brain.
| |
| They serve to convey afferent excitations toward the brain and
| |
| in that organ give rise to the great brain centers. The axons of
| |
| 250 ECTODERMAL DERIVATIVES
| |
| | |
| these last two types of neurons form the descending and ascending bundles of the brain and cord.
| |
| | |
| The spinal cord. —4 The spinal cord, or neural tube exclusive
| |
| of the brain, retains its primitive characteristics, - The cavity,
| |
| or neurocoel, persists as the central canal. (Between each pair
| |
| of vertebrae/ the afferent and efferent neurons’ form a pair of
| |
| spinal nerves which run out into the myotomes and hence have
| |
| a metamerism equivalent to that of the myotomes, an important
| |
| point in considering the homologies of the muscles. { In the region
| |
| of the pectoral and pelvic appendages, several of the segmental
| |
| nerves combine to form the brachial and the sacral plexus, respectively. The cord becomes surrounded by an envelope of mesenchyme known as the meninx, which in the higher vertebrates
| |
| becomes divided into an inner pia mater and an outer dura mater.
| |
| The development of the nerves will be taken up in a later section.
| |
| | |
| The brain. — Whereas the cord is largely composed of afferent,
| |
| efferent, and intersegmental neurons, by which certain reflex
| |
| actions are directed, the anterior end of the neural tube enlarges
| |
| and differentiates into the complex brain (Fig. 169). Here arise
| |
| several centers in which the impulses received mainly from the
| |
| major sense organs, nose, eye, and ear, are correlated. The brain
| |
| may be divided into two major regions: the archencephalon, or
| |
| prechordal brain; and the deutencephalon, or epichordal brain.
| |
| With continued local growth, the archencephalon grows down in
| |
| front of the notochord, thus forming the first or cranial flexure.
| |
| At the same time, three dilations appear: the prosencephalon
| |
| from the archencephalon; the mesencephalon at the point of the
| |
| flexure; and the rhombencephalon from the deutencephalon.
| |
| It is convenient to associate the future history of the prosencephalon with that of the nose, the mesencephalon with that of
| |
| the eye, and the rhombencephalon with that of the ear.
| |
| | |
| The prosencephalon. — The later history of the prosencephalon
| |
| is complicated by the fact that\the optic placode is included in
| |
| the neural tube at this point. Accordingly, we find he prosencephalon dividing into an anterior telencephalon and a posterior
| |
| diencephalon.
| |
| | |
| The telencephalon. — The anterior part of the telencephalon
| |
| becomes the olfactory lobe, which receives the afferent neurons
| |
| from the nose. From the roof develops the cerebrum, ‘which beTHE DIENCEPHALON 251
| |
| | |
| comes the most complex and important center of association’.
| |
| From the floor arises the optic part of the hypothalamus.* There
| |
| are two cavities, or telocoels (also known as the lateral ventricles).
| |
| | |
| - Deutencephalon
| |
| | |
|
| |
|
| |
| | |
| gE
| |
| | |
| Neyrenterie EX
| |
| | |
| cana.
| |
| | |
| Diencephalon
| |
| | |
|
| |
| | |
| Telencephalon
| |
| | |
| Fig. 169. — Diagrams to show early development of the vertebrate brain in sagittal
| |
| sections. A, prechordal and epichordal divisions. B, primary brain vesicles.
| |
| C, definitive vesicles. The longitudinal broken line indicates division between roof
| |
| and floor regions. (After von Kuppfer.)
| |
| | |
| The diencephalon. — The roof of the diencephalon gives rise
| |
| to the thalamus in front, and the metathalamus behind; from
| |
| the latter springs a dorsal diverticulum, the epithalamus. This
| |
| structure, often known as the epiphysis, gives rise to something
| |
| very much resembling an unpaired eye in early embryonic life;
| |
| this later becomes the pineal gland of the adult, one of the so252 ECTODERMAL DERIVATIVES
| |
| | |
| called endocrine glands. / The eyes take their origin from the side
| |
| of the diencephalon. The floor of the diencephalon gives rise to a
| |
| ventral diverticulum — the infundibulum, which grows downward
| |
| to meet the advancing hypophysis from the stomodeum (see page
| |
| 181). The two later fuse to form the pituitary glandJanother of
| |
| the endocrine series. Behind the infundibulum, the floor of the
| |
| diencephalon forms the mammillary part of the hypothalamus.
| |
| It is evident that the thalamencephalon, often used as a synonym
| |
| of the diencephalon, differs from it by the inclusion of the optic
| |
| part of the hypothalamus, which is derived from the telencephalon
| |
| although indistinguishable from the mammillary part of the
| |
| hypothalamus in the adult. The thalami contain nuclei (masses
| |
| of neurons) which receive afferent impulses from the optic,
| |
| general sensory, and acoustic organs, and transmit impulses to
| |
| and from the other centers of the brain. The cavity of the
| |
| diencephalon persists as the diacoel (third ventricle).
| |
| | |
| The mesencephalon. — The roof of the mesencephalon gives
| |
| rise to the corpora bigemina (quadrigemina in mammals), or optic
| |
| lobes, the centers which receive afferent impulses from the eyes
| |
| transmitted through the diencephalon.| The floor of the mesencephalon is the anterior portion of the brain stem, from which the
| |
| motor neurons of the cranial nerves depart. The third and fourth
| |
| cranial nerves originate from the mesencephalon. Its cavity is
| |
| the mesocoel (or aqueduct).
| |
| | |
| The rhombencephalon. —‘The hind-brain, like the fore-brain,
| |
| is divided into two regions, metencephalon and myelencephalon,
| |
| respectively. ~
| |
| | |
| The metencephalon. — The roof of the metencephalon gives
| |
| rise to the cerebellum, the center associated with hearing except
| |
| in mammals), the lateral line organs of anamnidtes, and the sense
| |
| of equilibrium. The floor of the metencephalon is part of the
| |
| brain stem, and from it arises the pons, a bundle of axons connecting the two sides of the cerebellum. The cavity is the
| |
| metacoel.
| |
| | |
| The myelencephalon. — The roof of the myelencephalon is
| |
| covered by a thin roof plate, the choroid plexus. Its floor forms
| |
| the posterior portion of the brain stem. (The cranial nerves,
| |
| from V to XII inclusive, depart from this portion of the stem,
| |
| which merges imperceptibly into the spinal cord. Its cavity,
| |
| THE SPINAL NERVES 253
| |
| | |
| hardly distinguishable from that of the metencephalon, is called
| |
| the myelocoel (fourth- ventricle).
| |
| | |
| The spinal nerves. af The nerves are segmentally arranged
| |
| bundles of afferent and efferent neurons originally associated with
| |
| the myotomes. The afferent neurons arise in the ganglia, the
| |
| efferent in the floor of the spinal cord. Accordingly, a typical
| |
| spinal nerve has two roots in the cord: a dorsal afferent root
| |
| uniting with the ganglion; and a ventral efferent root which
| |
| unites with the dorsal root after the other has attached itself to
| |
| the ganglion ¥Fig. 170). The nerve trunk then divides into
| |
| branches, each containing afferent and efferent neurons, which
| |
| are called rami and supply the body wall, although one (the com
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
| | |
|
| |
| | |
| Dorsal root Marginal layer
| |
| Somatic sensory neuron foi rs Ependymal layer
| |
| Visceral sensory neuron fd i \\\ Manile layer
| |
| | |
| Spinal ganglion
| |
| Visceral motor neuror
| |
| | |
| Somatic motor neuro
| |
| | |
| Dorsal ramus
| |
| | |
| Q
| |
| | |
| Lat. terminal . O
| |
| division CV\’\A (
| |
| Ventral terminal division of , Aorta teen A
| |
| Spinal nerve .
| |
| Ramus communicans Sympathetic ganglion
| |
| | |
| Fia. 170. — Diagram to show the neuron components of a spinal nerve. Transverse section of 10 mm. human embryo. (From Arey after Prentiss.)
| |
| | |
| municating ramus) connects with a sympathetic ganglion, derived
| |
| from a spinal ganglion, through which the splanchnic afferent and
| |
| efferent neurons serve the viscera.
| |
| | |
| It has been shown by Coghill that the development of behavior is closely paralleled by the development of the connections (synapses) between the neurons. Thus in the urodele,
| |
| Ambystoma, the first reflex of the embryo, a bending away from a
| |
| light touch on the skin, does not take place until an intermediate
| |
| 254 ECTODERMAL DERIVATIVES
| |
| | |
| neuron in the spinal cord has established synaptic relations with
| |
| the sensory tract on one hand and a floor plate cell which already
| |
| has established a synaptic relation to the motor tract on the
| |
| opposite side of the spinal cord (Fig. 171).
| |
| | |
|
| |
|
| |
|
| |
| | |
| Floor Piole Ceir
| |
| | |
|
| |
|
| |
| | |
| Motor Fibre
| |
| | |
| Notochord
| |
| | |
|
| |
| | |
| Fig. 171. — Diagram to show in transverse section of Ambystoma larva, neurons
| |
| concerned in earliest reflex. (From Coghill, ‘““Anatomy and the Problem of
| |
| Behavior.’’)
| |
| | |
| The autonomic nerves. — The brain, spinal cord, and cranial
| |
| and spinal nerves are grouped by anatomists as the central
| |
| nervous system. Associated with this is the autonomic nervous
| |
| system, consisting of nerves and ganglia and controlling the smooth
| |
| muscles of the viscera and blood vessels, and some glands. This
| |
| system arises from the neural plate, like the central nervous system, but from the lateral margins which become the neural crests.
| |
| At the time when the neural crests are dividing into the cerebrospinal ganglia, some of the cells migrate inward toward the dorsal
| |
| aorta, where they aggregate and multiply to form the chain
| |
| ganglia. The chain ganglia on each side become connected by
| |
| fore and aft extensions which form the sympathetic trunks. They
| |
| retain a connection with the cranial and spinal ganglia by means of
| |
| the communicating rami, and send out nerves along the principal
| |
| blood vessels. From the chain ganglia, by secondary and tertiary
| |
| THE AUTONOMIC NERVES 255
| |
| | |
| migrations, arise the prevertebral and visceral ganglia. In the
| |
| head the four sympathetic ganglia (ciliary, sphenopalatine, otic,
| |
| | |
| Glossopharyngeal
| |
| | |
|
| |
|
| |
| | |
| Semilunar ganglion Vagus
| |
| ganglion | ganglion
| |
| Nile
| |
| Aorta
| |
| Ciliary ZO 4
| |
| ganglion @-— / /
| |
| @ tic
| |
| Sphenopalatine / ganglion Z )
| |
| ganglion
| |
| | |
| Spinal
| |
| OZ) ganglion
| |
| Submaxillary (a
| |
| ganglion
| |
| | |
|
| |
| | |
| cartine Za § —<_)
| |
| | |
| plexus @
| |
| Chai n PSS
| |
| | |
| $
| |
| BS
| |
| cit Ly SS
| |
| | |
| <i SS
| |
| Soma eS
| |
| | |
| Prats geeg™ Se
| |
| © “
| |
| oe a
| |
| Te ALD
| |
| aft PA
| |
| dy j | \y sO
| |
| Fig. 172. — Diagram to show migrations of autonomic ganglia in human develop
| |
| ment. (After Strecter.)
| |
| | |
| and submaxillary) arise from the semilunar ganglion of the fifth
| |
| cranial nerve, and later acquire connections with the chain
| |
| ganglia (Fig. 172).
| |
| 256 ECTODERMAL DERIVATIVES
| |
| | |
| It has already been noted (page 214) that some of the cells from
| |
| the autonomic ganglia (chromaffin cells) migrate to the vicinity
| |
| of the mesonephros to form the suprarenal gland.
| |
| | |
| The cranial nerves. — The cranial nerves, or nerves of the
| |
| head regions, contain not only splanchnic and somatic afferent
| |
| and efferent neurons comparable to those of the spinal
| |
| nerves, but also special afferent neurons from the nose,
| |
| eye, ear and lateral line system. There are ten cranial
| |
| nerves in the anamniotes,
| |
| twelve in the amniotes (Figs.
| |
| 173, 174). To these should
| |
| be added in all cases the terminal nerve, unknown when
| |
| the cranial nerves were first
| |
| classified.
| |
| | |
| O. Terminal, a ganglionated nerve from the organ of
| |
| Jacobson entering the cerebral lobe with functions unknown, probably sensory.
| |
| Fig. 173. — Diagram to show origin of cranial I. Olfactory, a non-gangli
| |
| nerves in man. (After His.) onated sensory nerve from
| |
| the olfactory sensory region
| |
| | |
|
| |
| | |
| of the nose to the olfactory lobe.
| |
| | |
| II. Optic (ophthalmic), a non-ganglionated sensory nerve from
| |
| the retina of the eye to the floor of the diencephalon where the
| |
| fibers from the two eyes cross (optic chiasma). Each set of
| |
| fibers then enters the brain and runs to the optic lobe on the opposite side of the brain to that on which the eye is located.
| |
| | |
| Il. Oculomotor (motor oculi), a motor nerve, somatic with
| |
| some sensory elements, from the floor of the mesencephalon to all
| |
| muscles of the eyeball except the superior oblique and the lateral
| |
| rectus.
| |
| | |
| IV. Trochlear, a motor nerve, somatic with some sensory elements, from the roof of the mid-brain to the superior oblique
| |
| muscle of the eyeball.
| |
| THE CRANIAL NERVES 257
| |
| | |
| V. Trigeminal, a mixed nerve. Its somatic sensory neurons
| |
| arise in the semilunar ganglion, the motor elements in the floor of
| |
| the myelencephalon. The sensory neurons are somatic (general
| |
| cutaneous). The motor neurons supply the jaws (mandibular
| |
| arch).
| |
| | |
| VI. Abducens (pathetic), a somatic motor nerve with some
| |
| sensory elements, arising from the myelencephalon and supplying the external rectus muscle of the eyeball.
| |
| | |
| VII. Facial, a mixed nerve. The afferent neurons arise in the
| |
| geniculate ganglion and are splanchnic in nature, supplying the
| |
| | |
|
| |
|
| |
|
| |
|
| |
| | |
| Visceral
| |
| arches
| |
| | |
| t
| |
| | |
|
| |
| | |
| Pay
| |
| | |
| Somatic sensory «.....- .
| |
| Visceral sensory — — — —
| |
| Somatic motor
| |
| Visceral motor —-—-—
| |
| | |
|
| |
| | |
| Fig. 174. — Diagram showing relationships between cranial nerves and parts supplied. A, B, C, head somites. Arabic numerals, visceral arches. Roman
| |
| numerals, nerves.
| |
| | |
| hyoid arch, and also the tongue of mammals. In the anamniotes, an associated ganglion gives rise to a lateral branch with
| |
| afferent components from the lateral line organs. The efferent
| |
| neurons supply the hyoid arch in the lower vertebrates and the
| |
| facial region in the amniotes. The rami of the fifth and seventh
| |
| nerves are closely associated.
| |
| | |
| VIII. Acoustic (auditory), a ganglionated sensory nerve arising
| |
| from the acoustic ganglion and bearing afferent neurons from the
| |
| ear. In higher vertebrates it becomes differentiated into the
| |
| 258 ECTODERMAL DERIVATIVES
| |
| | |
| vestibular and cochlear nerves, each with its own ganglion
| |
| produced by the division of the acoustic ganglion.
| |
| | |
| IX. Glossopharyngeal, a mixed nerve. The afferent neurons
| |
| arise in the petrosal and the superior ganglion and are principally
| |
| splanchnic. They divide into a prebranchial branch running into
| |
| the hyoid arch and a postbranchial branch into the first branchial
| |
| arch. The efferent components are principally found in the
| |
| postbranchial branch.
| |
| | |
| X. Vagus, a mixed nerve arising by the fusion of several primitive cranial nerves, which supplied the arches with afferent
| |
| (from the jugular ganglion) and efferent neurons. In addition,
| |
| the vagus gives off a visceral branch to the stomach, lungs, etc.,
| |
| and in the anamniotes a lateral branch to the lateral line organs
| |
| of the trunk (from the nodosum ganglion).
| |
| | |
| XI. Accessory, a motor nerve which innervates the muscles of
| |
| the shoulder girdle and is found only in the amniotes. A ganglion
| |
| (of Froriep) disappears before the embryo becomes adult.
| |
| | |
| XII. Hypoglossal, also a motor nerve, which innervates the
| |
| tongue in the amniotes. In the anamniotes the tongue is innervated by so-called “ occipital’? nerves which possibly are the
| |
| fore-runners of the hypoglossal, prior to the appropriation of the
| |
| occipital region by the head.
| |
| | |
| Metamerism of the nervous system.— The metameric arrangement of the nerves, like that of the segmental arteries, is
| |
| purely secondary and dependent upon the primary metamerism
| |
| of the mesoderm. The nerves, however, are more conservative
| |
| than the vascular organs or myotomic derivatives. For example, the diaphragm of mammals is supplied by muscles from
| |
| one of the cervical myotomes, and the innervation of the diaphragm (phrenic nerve) still arises from the cervical region.
| |
| Many attempts have been made to reconstruct the metamerism
| |
| of the head, by a study of the cranial nerves, following Bell’s law:
| |
| that every original cranial nerve has, like a spinal nerve, a dorsal
| |
| sensory and ventral motor root.
| |
| | |
| This problem has been complicated by the fact that in the
| |
| head there are two types of metamerism, (1) primary as indicated
| |
| by the head myotomes in the elasmobranch embryo, and (2)
| |
| secondary (branchiomeric) as indicated by the visceral arches
| |
| (Fig. 174). Accordingly, there are two types of musculature,
| |
| METAMERISM OF THE NERVOUS SYSTEM 259
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
| TABLE 11
| |
| NEURONE COMPONENTS OF CRANIAL NERVES AND FUNCTIONS
| |
| Nerve Afferent Afferent Efferent Efferent
| |
| erv Somatic Splanchnic Somatic Splanchnie
| |
| I Smell
| |
| IT Vision
| |
| III Movement
| |
| of eyeball
| |
| IV Movement
| |
| of eyeball
| |
| Vv General Movement
| |
| cutaneous of jaw
| |
| VI Movement
| |
| of eyeball
| |
| VII Taste Hyoid and
| |
| facial movement
| |
| and salivation
| |
| Vill Hearing and
| |
| equilibration
| |
| Ix Taste and Salivation,
| |
| pharyngeal pharyngeal
| |
| sensation movement
| |
| x Visceral Movement of
| |
| sensation viscera and
| |
| pharynx
| |
| XI Movement of
| |
| pharynx and
| |
| shoulder
| |
| XII Movement
| |
| of tongue
| |
| 260 ECTODERMAL DERIVATIVES
| |
| | |
| (1) somatic as represented by the muscles of the eyeball, and (2)
| |
| splanchnic as represented by the muscles of the jaws and visceral
| |
| arches. Two types of efferent neurons, therefore, are present,
| |
| (1) somatic and (2) splanchnic. The splanchnic motor neurons
| |
| of the cranial nerves differ from those of the trunk, however, in
| |
| that no sympathetic neurons intervene between them and the
| |
| muscles which they supply. There are altogether three sets of
| |
| afferent neurons: (1) the general sensory or cutaneous, which
| |
| correspond to the somatic sensory neurons of the trunk; (2)
| |
| splanchnic sensory, which correspond to those of the trunk; and
| |
| (3) lateral, belonging to the lateral line system. The cranial
| |
| nerves are evidently not serially homologous, as can be seen from
| |
| Table 11.
| |
| | |
| Finally, we must mention the neuromeres which have been
| |
| reported in various vertebrate embryos. These are formed by
| |
| constrictions in the cranial portion of the neural tube and interpreted by some authors as the remains of a neural metamerism.
| |
| They seem in many forms to correspond with the cranial nerves
| |
| and more probably represent areas of local growth prior to the
| |
| outgrowth of the nerves themselves.
| |
| | |
| The general problem of the metamerism of the head still awaits
| |
| solution. The latest summary, that of Brachet, indicates the
| |
| probable number of segments in the primitive head as six. Three
| |
| of these are ephemeral, and their somites give rise to mesenchyme.
| |
| The three posterior segments are associated with the first three
| |
| visceral clefts bounded by the first four arches, each of which has
| |
| its own cranial nerve: the trigeminal of the mandibular arch; the
| |
| facial of the hyoid; the glossopharyngeal of the first branchial;
| |
| the vagus of the second branchial arch. According to this interpretation, the posterior clefts and arches are reduplications supplied by new branches of the vagus, while the accessory and
| |
| hypoglossal are secondarily acquired spinal nerves.
| |
| | |
| THE FROG (SEE ALSO CHAPTER XI). — The prechordal and epichordal divisions of the brain are demarcated by the notochord,
| |
| and the division into the three primary vesicles is but slightly
| |
| indicated. The brain of the frog never develops neuromeres.
| |
| The optic lobes are corpora bigemina. The division into myelencephalon and metencephalon is incomplete, and no pons is formed.
| |
| There are forty pairs of spinal nerves in the tadpole, reduced to
| |
| THE SENSE ORGANS 261
| |
| | |
| ten in the adult. There are but ten of the cranial nerves (XI and
| |
| XII not included). The sympathetic ganglia originate from the
| |
| cranial and spinal ganglia by the emigration of ganglion cells.
| |
| | |
| THE CHICK (SEE ALSO CHAPTER XII). — The divisions of the
| |
| brain into the three primary and five secondary vesicles is well
| |
| marked. Eleven neuromeres are formed, of which three are found
| |
| ‘in the prosencephalon, two in the mesencephalon, the remainder in
| |
| the rhombencephalon. “Three flexures are formed: (1) cranial
| |
| in the floor of the mesencephalon; (2) cervical at the junction
| |
| of the myclencephalon and the spinal cord; and (3) pontine
| |
| in the floor of the myclencephalon. A pons is formed. There
| |
| are fifty pairs of nerves developed in the chick of eight days
| |
| (Lillie), of which thirty-cight are spinal and twelve cranial, including the eleventh and twelfth which are not incorporated in
| |
| the head of the frog.
| |
| | |
| MAN (SEE ALSO CHAPTER XIII). — The particular feature of importance in the development of the human brain is the great
| |
| increase in size and complexity of the cerebral hemispheres of
| |
| the telencephalon. The optic lobes are quadripartite (corpora
| |
| quadrigemina), of which the two anterior lobes are especially
| |
| associated with vision, the two posterior ones with hearing.
| |
| | |
| C. THE SENSE ORGANS
| |
| | |
| The nervous system receives stimuli not only from outside the
| |
| body but also from within, such as those concerning the tension of
| |
| the muscles. For the reception of stimuli, special organs — the
| |
| sense organs — are developed. Of these the most conspicuous
| |
| are the eyes, the ears, and the nose. In addition, it must be
| |
| remembered that the entire skin functions as a sense organ by
| |
| means of special receptors, and that stimuli are received from the
| |
| viscera and other internal structures by means of free nerve
| |
| terminations.
| |
| | |
| Of the special sense organs, the eye is most specialized in its
| |
| mode of development. It is responsive to photic stimuli. ‘The
| |
| nose represents a concentration of chemical sense receptors, more
| |
| highly developed than the scattered taste buds of the head,
| |
| which are confined in adult mammals to the cavity of the mouth.
| |
| The ear, responsive to slower vibrations (pressure, sound) in the
| |
| surrounding medium, originates in a manner similar to that of
| |
| A Nasal pit
| |
| | |
| Frontal
| |
| | |
| process Oronasal
| |
| | |
| groove
| |
| | |
| Mandible
| |
| Mandible
| |
| | |
| Mandible
| |
| | |
|
| |
| | |
|
| |
| | |
| Oy
| |
| 2K
| |
| xX
| |
| | |
|
| |
| | |
|
| |
| | |
| Primitive
| |
| palate
| |
| | |
| c. QS b
| |
| | |
| Fig. 175. — Diagrams showing early stages in development of nose. A, nasal placodes (in black). B, same now on ventral
| |
| surface of head. C, nasal pits. D, nasal grooves, anterior-ventral view. E, nasal tubes, ventral view, lower jaw removed.
| |
| | |
|
| |
| | |
| ~ "
| |
| 2,
| |
| x
| |
| | |
|
| |
| | |
| x
| |
| | |
| CALS.
| |
| Xx
| |
| XH
| |
| ieee
| |
| Y
| |
| x)
| |
| | |
|
| |
| | |
| x
| |
| | |
|
| |
| | |
|
| |
| | |
| AXA
| |
| | |
|
| |
| | |
| ©
| |
| | |
|
| |
|
| |
| | |
|
| |
| | |
| 262
| |
| | |
| ECTODERMAL DERIVATIVES
| |
| THE NOSE 263
| |
| | |
| the lateral line system. This system is highly developed in
| |
| the aquatic anamniotes, vestigial or absent in the amniotes. The
| |
| ear, on the other hand, is more highly developed in the amniotes.
| |
| | |
| The nose. -+ The nose arises as a pair of local thickenings of the
| |
| ectoderm at the anterior end of the head (Fig. 175). These
| |
| thickenings are hereafter known as the nasal (olfactory) placodes.
| |
| Later they invaginate to form the nasal (olfactory) pits,)which
| |
| persist as the nose of all fish except the air-breathing dipnoi.
| |
| Here also should be noted the fact that the cyclostomes are
| |
| peculiar in the possession of a single median nasal pit. Among
| |
| the tetrapods(the nasal pits elongate to become oro-nasal grooves,
| |
| the anterior ends of which become connected with the developing
| |
| mouth into which they are carricd.\ The original anterior ends
| |
| | |
|
| |
|
| |
|
| |
|
| |
| | |
| Nasolachyrmal
| |
| duct
| |
| | |
|
| |
| | |
| Auditory
| |
| : : tube
| |
| External Dee Pai NE Pharynx
| |
| “nares. Gil y Hard st
| |
| / ; Palate oft
| |
| | |
| Palate
| |
| Fig. 176. — Sagittal hemi-section through human nose. (After Howden.)
| |
| | |
| of the nasal pits, therefore, come to lie at the posterior end of the
| |
| mouth and open into the pharynx as the internal nares, while the
| |
| original posterior ends become the external nares (Fig. 175E).
| |
| The nasal cavity is later separated from the oral cavity by the
| |
| ingrowth of the maxillary, palatine, and pterygoid bones, which
| |
| form the hard palate YFig. 176). Jacobson’s organ arises as a
| |
| pocket of the olfactory epithelium. Its function is unknown.
| |
| The olfactory epithelium contains ciliated cells connected to the
| |
| olfactory lobe by means of the first cranial nerve) which is
| |
| 264 ECTODERMAL DERIVATIVES
| |
| | |
| peculiar in that its ncurons run directly to the brain. without the
| |
| interposition of ganglion cells. Jacobson’s organ receives a
| |
| branch of the trigeminal nerve.
| |
| | |
| The eye. — The optic placodes are incorporated into the neural
| |
| plate, where they can be distinguished as lateral thickenings of
| |
| the margin at points which will later be included in the diencephalon. (Fig. 177). When the tube is formed, the relation of the
| |
| sensory epithelial cells to the exterior is, of course, reversed.
| |
| The optic placodes “ invaginate,” but, owing to their relation
| |
| to the neural tube, the result is an apparent “ evagination ” from
| |
| the tube towards the exterior. This produces the outgrowths
| |
| which later, by constriction, give rise to the proximal optic stalks
| |
| and distal optic vesicles. At the point where the optic vesicle
| |
| touches the ectoderm, two reactions take place: (1) a local thickening of the ectoderm, called the lens placode, from which the lens
| |
| of the eye develops; and (2) an invagination of the optic vesicle
| |
| whereby this vesicle is transformed into a two-layered optie cup
| |
| This invagination continues into the optic stalk to produce a
| |
| groove called the choroid fissure.
| |
| | |
| The lens. —< The lens placode invaginates to form the lens pit,
| |
| which then withdraws still further from the surface and becomes
| |
| closed in by the union of its external lip to form the lens vesicle.
| |
| The lens‘vesicle becomes solid by the elongation of the cells on the
| |
| internal side which assume a clear transparent appearance.)
| |
| | |
| The optic cup. — The inner layer of the cup becomes the sensory portion of the retina, the outer layer the pigmented portion.
| |
| Jt will be recalled that the sensory epithelium of the eye is inverted, and as a result the rods and cones, or sensory elements,
| |
| of the retina are pointed away from the light.’ In the pigmented
| |
| layer of the retina, melanin is secreted. Meantime the cavity of
| |
| the optic cup becomes filled with a clear fluid secreted by the
| |
| surrounding cells, which later becomes viscous and forms the
| |
| vitreous humor.
| |
| | |
| The envelopes of the eyeball (Fig. 178). — Around the optic
| |
| cup and stalk, a layer of mesenchyme accumulates, which later
| |
| differentiates into an inner delicate layer called the choroid which
| |
| contains pigment and capillaries and: may be compared with the
| |
| pia mater of the brain, and an outer dense layer known as the
| |
| sclera, which may be compared with the dura mater of the brain.
| |
| THE ENVELOPES OF THE EYEBALL 265
| |
| | |
| The external portion of the sclera over the lens makes contact
| |
| with the epidermis-and becomes transparent to form the cornea.
| |
| | |
| ‘
| |
| | |
| eS SS
| |
| | |
| E
| |
| | |
| Fig. 177. — Diagrams showing early stages in development of vertebrate eye. A,
| |
| optic placodes (in black). B, same after formation of neural tube. C, optic
| |
| vesicles and lens placodes. D, optic cups and lens pits. E, optic cups and lens
| |
| vesicles.
| |
| | |
| The epidermis over the eye forms the conjunctiva. In some
| |
| vertebrates, sclerotic cartilage, or even bone, is formed, the vestige of an optic capsule. ) The edge of the choroid, together with
| |
| 266 ECTODERMAL DERIVATIVES
| |
| | |
| the marginal retina, gives rise to the iris, a circular curtain surrounding the opening of the cup which is called the pupil of the
| |
| eye. The muscles of the.iris are apparently of ectodermal origin.
| |
| The iris divides the space between the lens and the cornea into
| |
| two chambers, an anterior and a posterior chamber, which are
| |
| filled with a fluid, the aqueous humor. The muscles of the
| |
| | |
|
| |
|
| |
| | |
| Posterior chamber:
| |
| SL Ciliary
| |
| | |
| Optic nerve
| |
| Fig. 178. — Horizontal section of human eye. (After Howden.)
| |
| | |
| eyeball are six in number, arising from the three head myotomes.
| |
| They are innervated by the oculomotor, trochlear, and abducens
| |
| nerves.
| |
| | |
| The optic nerve. — The afferent neurons pass from the retina
| |
| into the optic cup and form a bundle which passes out through
| |
| the choroid fissure and into the optic stalk, and so to the optic
| |
| chiasma on the floor of the diencephalon, where they cross and
| |
| make their way to the optic lobes on the opposite side.
| |
| | |
| The lateral line system. — This is a diffuse sensory organ consisting of sense buds arranged in rows over the head and body of
| |
| aquatic anamniotes. Its function apparently is to detect slow
| |
| vibrations in the water. The origin of the lateral line system is a
| |
| lateral thickening of the sensory ectoderm which later breaks up
| |
| into separate suprabranchial placodes. These are found in the
| |
| THE INNER EAR 267
| |
| | |
| embryos of the amniotes but soon degenerate. The lateral line
| |
| system is of particular interest inasmuch as the lateral thickening referred to is in some cases continuous with the otic placode
| |
| which gives rise to the ear. The principal nerve supplying the
| |
| lateral system is the facial, although trigeminal, glossopharyngeal, and vagus often contain lateral line components.
| |
| | |
| The ear. — The ear becomes differentiated into the vestibule or
| |
| equilibratory organ and the cochlea or organ of hearing. Three
| |
| parts of the ear are distinguished (Fig. 180). The inner ear,
| |
| giving rise to the vestibule and the cochlea, arises from an ectodermal otic (auditory) placode. The middle ear appears in the
| |
| amphibians, and it is derived from the endodermal first visceral
| |
| pouch. The outer ear, found only in the amniotes, is an ectodermal derivative of the first visceral groove and an outgrowth
| |
| from the mandibular and hyoid arches
| |
| | |
| The inner ear. — This originates from the otic placode, which
| |
| invaginates to form an otic (auditory) pit (Fig. 179) and later
| |
| closes over to withdraw from the epidermis as the otic (auditory)
| |
| vesicle or otocyst., In some vertebrates (elasmobranchs) the
| |
| vesicle retains its connection with the exterior by means of a
| |
| hollow stalk, the endolymphatic duct. Usually this connection
| |
| is lost and the endolymph duct of the adult is a new formation.
| |
| The vesicle divides into a ventral saccule and a dorsal vestibule
| |
| or utricle. The saccule gives rise to the éndolymph duct and the
| |
| lagena, which in mammals becomes th@ coiled cochlea or organ of
| |
| hearing, while the utricle gives rise by constriction to three semicircular canals, each with a dilation at one endf the ampulla.
| |
| The sensory epithelium is restricted to the lagena and ampullae}
| |
| The cavity of these structures is known as the membranous
| |
| labyrinth, and contains a fluid, the endolymph. Concretions, the
| |
| otoliths, may appear in the endolymph of the vestibular portion.
| |
| Around this labyrinth ‘the otic capsule, Jor skeletal labyrinth} is
| |
| formed. ‘This later ossifies to give rise to the otic bones. (The
| |
| skeletal labyrinth contains a fluid known as the perilymph. In
| |
| vertebrates with a middle ear) two openings are formed in the
| |
| skeletal labyrinth, the fenestra rotunda, closed by a membrane,
| |
| and the fenestra ovale, into which the stapes projects.g The
| |
| acoustic nerve, whieh is ganglionated, divides into a vestibular
| |
| and a cochlear nerve, each with its separate ganglion. }
| |
| 268 ’ ECTODERMAL DERIVATIVES
| |
| | |
| TN
| |
| | |
| A
| |
| | |
| B
| |
| fo \0\
| |
| Fie. 179. — Diagrams showing early stages in development of inner ear, A, otic
| |
| | |
| placodes (in black). B, otic pits. C, otic vesicles (otocysts).
| |
| | |
| Semicircular
| |
| canal
| |
| | |
| Endolymph
| |
| duct
| |
| | |
|
| |
| | |
| Saccule
| |
| | |
| Cochlea
| |
| | |
| Fig. 180. — Frontal section of human ear. Semi-diagrammatic. (After Howden.)
| |
| THE FROG 269
| |
| | |
| The middle ear. — The middle ear arises from the first visceral
| |
| pouch, which constricts into a proximal auditory (Eustachian)
| |
| tube and a distal tympanic cavity which is separated from the
| |
| exterior by the tympanic membrane,(a persistent closing plate
| |
| formed from ectoderm and endoderm. Through the tympanic
| |
| cavity there is a chain of bones (auditory ossicles) connecting the
| |
| tympanum with the fenestra ovalis. In anurans and sauropsids,
| |
| this chain of auditory ossicles consists of the columella and stapes
| |
| (hyomandibular). In the mammals, the columella is replaced by
| |
| the incus and malleus, equivalent to two other jaw bones, the
| |
| quadrate and articulare, respectively. The muscles of the middle
| |
| ear, tensor tympani and stapedial muscles, arise from the mesoderm of the mandibular and hyoid arches, respectively, and are
| |
| innervated by the facial and glossopharyngeal nerves.
| |
| | |
| The outer ear.—— The external ear consists of the external
| |
| auditory meatus, derived from the first visceral groove, and the
| |
| pinna, which arises from tubercles on the mandibular and hyoid
| |
| arches. It is composed of mesoderm and ectoderm, contains
| |
| muscles, and is strengthened by cartilage. The innervation is
| |
| from the facial nerve.
| |
| | |
| THE FROG (SEE ALSO CHAPTER XI).— In the development of
| |
| the nose, the nasal groove stage is suppressed. Instead, a thickening develops from the olfactory pit into the mouth as far as the
| |
| pharynx. This acquires a lumen which connects the olfactory
| |
| pit to the pharynx. The development of the eye presents no
| |
| especial peculiarities. The endolymph duct is a dorsal evagination from the otocyst. The semicircular canals are each formed
| |
| by the appearance of a pair of ridges in the cavity of the utricle
| |
| which fuse to enclose the cavity of the canal. The saccule gives
| |
| rise to two ventral diverticula, the cochlea and basilar chamber.
| |
| The function of the latterisunknown. The tubo-tympanic cavity
| |
| arises from the first visceral pouch, which in the frog is vestigial
| |
| and has no cavity. From this rudiment a strand of cells grows
| |
| dorsad and later acquires a lumen. It loses its connection with
| |
| the pharynx and moves backward to the ear region where it
| |
| acquires a secondary connection with the pharynx (Fig. 181).
| |
| The tympanic membrane is apparently entirely ectodermal. The
| |
| columella, which connects the tympanum with the inner ear, arises
| |
| from two primordia: the inner stapedial plate, which is a part
| |
| 270 ECTODERMAL DERIVATIVES
| |
| | |
| of the otic capsule; and a cartilage derived from the palatoquadrate bar. This cartilage is thought to be homologous with
| |
| the hyomandibular bone of fishes. The lateral line organs arise
| |
| from the fragmentation of a placode known as the placode of the
| |
| tenth cranial nerve, which innervates this series. Similar epibranchial placodes appear on the head and are innervated by the
| |
| | |
| Brain . Utriculus and
| |
| Auditory semi-circular canals
| |
| nerve
| |
| | |
|
| |
|
| |
| | |
| Sacculus
| |
| | |
|
| |
| | |
| CORA ALLL
| |
| XS
| |
| | |
| POOLE LL >
| |
| SER RL,
| |
| SSSR Gina
| |
| ROKR RK ING
| |
| SEO LL
| |
| YY =) ERLE
| |
| ) OX
| |
| | |
|
| |
|
| |
|
| |
| | |
|
| |
| | |
|
| |
|
| |
|
| |
| | |
|
| |
| | |
|
| |
|
| |
| | |
|
| |
| | |
|
| |
|
| |
| | |
| y
| |
| 2 o
| |
| Ree 5x Stape al
| |
| OR
| |
| SIO SK HOS P
| |
| | |
|
| |
| | |
| SSG
| |
| SSK OS
| |
| | |
|
| |
|
| |
| | |
|
| |
| | |
| Fig. 181. — Rana pipiens, diagram to show the parts of the ear. Schematic crosssection through head.
| |
| | |
| seventh and ninth nerves. They are larval sense organs and
| |
| disappear at metamorphosis.
| |
| | |
| THE CHICK (SEE ALSO CHAPTER XII). — The chick has a cleft
| |
| palate due to the incomplete fusion of the palatine processes of
| |
| the maxillae. Jacobson’s organ makes a short appearance as a
| |
| vestigial organ but disappears before hatching. The eye possesses three eyelids, the third (nictitating membrane) arising
| |
| from a separate fold inside that which forms the upper and
| |
| lower lids. The pecten is a vascular plate in the vitreous
| |
| humor, from mesenchyme which enters the choroid fissure. Its
| |
| function is unknown. \The endolymphatic duct arises from the
| |
| dorsal wall of the otocyst. The semicircular canals arise as outpocketings of the otocyst prior to its separation into utricle and
| |
| saccule. The cochlea is more highly developed than in the frog.
| |
| The tubo-tympanic cavity arises from the first pharyngeal pouch.
| |
| The tympanum is formed from ectoderm and endoderm and includes a middle layer of mesenchyme. The columella arises from
| |
| SUMMARY 271
| |
| | |
| a stapedial plate and hyomandibular cartilage. The external
| |
| auditory meatus is short, and no pinna is developed. ©:
| |
| | |
| MAN (SEE ALSO CHAPTER XIII). — The organ of Jacobson is
| |
| rudimentary and may completely disappear in the adult. A
| |
| small fold (plica semilunaris) is the representative of the nictitating membrane. The cochlea is highly differentiated. The tube
| |
| and tympanic cavity form from the first visceral pouch. The
| |
| tympanum apparently is composed of all three germ layers.
| |
| There are three auditory ossicles. The stapes is derived from the
| |
| second visceral arch, while the malleus and incus arise from the
| |
| first visceral arch. They are thought to represent the quadrate
| |
| and articular bones of reptiles, respectively. The pinna arises
| |
| from elevations on the mandibular and hyoid arches.
| |
| | |
| SUMMARY
| |
| | |
| The ectoderm gives rise to the epithelial linings of the following
| |
| structures:
| |
| | |
| A. The epidermis, with the
| |
| apertures of
| |
| Oral cavity
| |
| Visceral clefts
| |
| Cloaca
| |
| | |
| B. The neural plate
| |
| 1. Neural tube
| |
| Brain and cranial nerves
| |
| Prosencephalon
| |
| Telencephalon
| |
| Diencephalon
| |
| Mesencephalon
| |
| Rhombencephalon
| |
| Metencephalon
| |
| Myelencephalon
| |
| Cord and spinal nerves
| |
| | |
| 2. Neural crest
| |
| Ganglia
| |
| Cerebrospinal
| |
| Autonomic
| |
| Suprarenal gland
| |
| 272 ECTODERMAL DERIVATIVES
| |
| | |
| C. Sensory placodes
| |
| | |
| 1. Nose
| |
| | |
| 2. Eye (choroid and sclera from mesoderm)
| |
| | |
| 3. Ear (middle ear from endoderm, ossicles from mesoderm)
| |
| | |
| 4, Lateral line organs
| |
| | |
| REFERENCES
| |
| | |
| Arey, L. B. 1934. Developmental Anatomy, 3rd Ed., Chaps. 14-17.
| |
| | |
| Brachet, A. 1921. Traité d’embryologie des vertébrés, Part H, Bk. 1, Chap. 4.
| |
| | |
| ACoghill, G. E. 1929. Anatomy and the Problem of Behavior.
| |
| | |
| Hertwig, O. 1906. Handbuch, Book II, Chaps. 5-10.
| |
| | |
| Jenkinson, J. W. 1913. Vertebrate Embryology, Chap. 7.
| |
| | |
| Keibel and Mall. 1910-1912. Human Embryology, Chaps. 14 and 16.
| |
| | |
| Kerr, J. G. 1919. Textbook of Embryology, Chap. 2.
| |
| | |
| Kingsley, J.S. 1926. Comparative Anatomy of Vertebrates.
| |
| | |
| Lillie, F. R. 1919. The Development of the Chick, 2nd Ed.
| |
| | |
| MeMurrich, J. P. 1923. The Devclopment of the Human Body.
| |
| | |
| Strong, O. S. 1921. The Nervous System, being Chap. 17 of Bailey and Miller,
| |
| Textbook of Embryology, 4th Ed.
| |
| PART IV
| |
| ANATOMY OF VERTEBRATE EMBRYOS
| |
| | |
| CHAPTER XI
| |
| THE ANATOMY OF FROG EMBRYOS
| |
| | |
| In earlier chapters we have discussed the fertilization of the
| |
| frog’s egg (page 57), its cleavage (pages 97, 103), and germ-layer
| |
| formation (pages 109, 118), and have observed that while the germ
| |
| layers are being laid down the process is complicated by the early
| |
| localization of some of the organ systems, notably the sensorynervous complex (page 129). In this account of later organogeny, three stages of development seem especially significant:
| |
| first, an early embryo of about 3 mm. body length in which the
| |
| visceral grooves are apparent, a stage attained in Rana pipiens
| |
| about the second day after the eggs are laid; second, the newly
| |
| hatched larva of about 6 mm. with external gills developing,
| |
| about two weeks old; third, a young “ tadpole ” stage of about
| |
| 11 mm. with the opercula covering the internal gills, about the
| |
| age of one month.
| |
| | |
| These stages are easily identified even though the lengths and
| |
| ages can be given only approximately, for the rate of development is greatly influenced by the prevailing temperature, and
| |
| the size of the tadpole is determined largely by external factors,
| |
| such as the amount of food available.
| |
| | |
| The student must bear in mind that the sections illustrated in
| |
| this and the two chapters following are for the sole purpose of giving
| |
| him starting points from which he 1s expected to study all the sections
| |
| in the series furnished him. He will probably never encounter
| |
| sections exactly like those selected for these illustrations, but he will
| |
| discover sections very like them from which he can commence his
| |
| own observations.
| |
| | |
| A. THE EARLY EMBRYO (3 MM.)
| |
| | |
| External form. — This stage corresponds approximately to the
| |
| embryo of 33 mm. described by Marshall. The head region,
| |
| through its more rapid growth, has become easily distinguishable
| |
| from the trunk, which bulges ventrally on account of the large
| |
| | |
| amount of contained yolk, and a well-marked tail bud is present.
| |
| 275
| |
| 276 THE ANATOMY OF FROG EMBRYOS
| |
| | |
| The neural folds have fused throughout their length, and enclosed the blastopore. In the head the stomodeum appears
| |
| as an antero-posterior slit on the anterior ventral surface, and is
| |
| enclosed by ridges identifiable as the maxillary processes and
| |
| mandibular arches. On either side and slightly ventral to the
| |
| stomodeum, are the primordia of the sucker or oral gland. At the
| |
| dorso-lateral margins the olfactory placodes have begun to evaginate. Lateral bulges on either side of the head are due to the
| |
| developing optic vesicles. The ear is now in the otic vesicle stage.
| |
| The gill region shows five visceral grooves. Immediately behind
| |
| the last arch, a swelling is caused by the developing pronephros.
| |
| Dorsally, slight furrows indicate the boundaries of thirteen soEpiphysie mites. Beneath the tail
| |
| Optic vesicle bud, the proctodeum
| |
| Prosencephalon has united with the
| |
| | |
| Oral gland hind-gut to form the
| |
| Visceral pouch eloacal aperture.
| |
| | |
| Fore gut Endodermal derivaay Liver tives. - The anterior
| |
| portion of the gastrocoel
| |
| is now a large fore-gut
| |
| with a thin-walled
| |
| lining. From this, on
| |
| a . either side, the begin
| |
| eurenteric . :
| |
| canal nings of three visceral
| |
| Fig. 182. —3 mm. frog embryo, viewed from right pouches can be seen.
| |
| | |
| side as a transparent object. X15.
| |
| | |
| From the fore-gut a narrow evagination grows backward into the floor of the mid-gut as
| |
| the primordium of the liver. The mid-gut is distinguishable by
| |
| its relatively narrow lumen and thick yolk-laden floor. The
| |
| small but thin-walled hind-gut opens above into the neurenteric
| |
| canal by which it is connected with the neurocoel, and opens
| |
| ventrally to the exterior by way of the proctodeum. An axial
| |
| rod, the hypochord, is found beneath the notochord. It originates from the roof of the gastrocoel and disappears soon after
| |
| hatching.
| |
| | |
| Mesodermal derivatives. — The notochord is large and vacuolated and enclosed by two sheaths. The somites have now
| |
| attained their maximum number (13) in the trunk, but are not
| |
| | |
|
| |
|
| |
|
| |
| | |
| Mesencephalon
| |
| | |
| Otic vesicle
| |
| Rhombencephalon
| |
| | |
| Somite I
| |
| Notochord +
| |
| THE EARLY EMBRYO 277
| |
| | |
| yet distinguishable in the tail region. The intermediate mesoderm, after a temporary division into nephrotomes, is now reunited into a nephrotomal band in which spaces have appeared
| |
| opposite the second, third, and fourth somites, indicative of the
| |
| pronephric tubules which are to develop. A thickening along the
| |
| | |
|
| |
|
| |
| | |
| Mesencephalon PD, Prosencephalon
| |
| | |
|
| |
| | |
| Neurenteric
| |
| canal
| |
| | |
| Fia. 1838. —3 mm. frog embryo. Sagittal section: 50.
| |
| | |
| nephrotomal band immediately below the ventro-lateral margins
| |
| of the somites is the primordium of the pronephric duct. Immediately below the floor of the fore-gut, the lateral mesoderm has
| |
| separated into dorsal splanchnic and ventral somatic layers, while
| |
| the contained space is the beginning of the pericardial cavity, the
| |
| only region of the coelom yet apparent.
| |
| | |
| Ectodermal derivatives. — The epidermis at this stage is
| |
| ciliated. The neurocoel, as has been remarked above, is con278 , THE ANATOMY OF FROG EMBRYOS
| |
| | |
| nected with the hind-gut by the neurenteric canal. At the
| |
| anterior end, the brain is distinguishable by its relatively larger
| |
| lumen and by the cranial flexure over the anterior end of the
| |
| notochord. The divisions between the three primary vesicles
| |
| are not marked by the constrictions characteristic of
| |
| many vertebrates, but are
| |
| distinguished by the following points of reference: the
| |
| prosencephalon extends to a
| |
| Optic line projected from a thickvesicle ening on the floor known as
| |
| the tuberculum posterius to
| |
| a point just in front of a
| |
| similar thickening on the
| |
| Hypophysis — dorsal wall; the mesencephOral gland = aon, from the boundary of
| |
| the prosencephalon to a line
| |
| Fia. 184. —3 mm. frog embryo. Transverse connecting the tuberculum
| |
| section through optic vesicle. 50. anda point just behind the
| |
| dorsal thickening; the rhombencephalon merges imperceptibly
| |
| into the spinal cord. From the prosencephalon, the optic vesicles
| |
| extend on either side and cause the external bulges already noted.
| |
| From the ventral side of the prosencephalon, a depression, the
| |
| infundibulum, extends towards the hypophysis, which in the frog
| |
| grows inward as a solid wedge of ectodermal cells anterior to
| |
| the stomodeum. Dorsally, the epiphysis appears as a median
| |
| evagination.
| |
| | |
|
| |
|
| |
| | |
| Fore-gut
| |
| | |
| B. THE LARVA AT HATCHING (6 MM.)
| |
| | |
| External form. — Although the larva, if it may be so called,
| |
| has emerged from the protecting membranes of egg jelly, the
| |
| mouth has not yet opened and for several days the yolk is still
| |
| the sole source of food. The head region is still easily distinguishable from the trunk, while the tail has increased greatly in
| |
| length and has become bilaterally compressed. In the head, the
| |
| stomodeal pit has deepened at the anterior end, and the maxillary processes and mandibular arches are more sharply sculptured. The invagination of the nasal (olfactory) placodes has
| |
| THE LARVA AT HATCHING 279
| |
| | |
|
| |
| | |
| Fig. 185.—3 mm. frog embryo. Transverse section through otic (auditory)
| |
| vesicle. 50.
| |
| | |
|
| |
| | |
| Fig. 186. —8 mm. frog embryo. Transverse section through mid-gut and liver.
| |
| X50.
| |
| 280 THE ANATOMY OF FROG EMBRYOS
| |
| | |
|
| |
| | |
| Fig. 187. — 3 mm. frog embryo. Frontal section through optic stalks, liver, and
| |
| hind-gut. 50.
| |
| THE LARVA AT HATCHING 281
| |
| | |
| continued to the point where they may be called pits, connected
| |
| to the anterior margins of the stomodeal pit by oro-nasal grooves.
| |
| The bulge of the eye is still prominent. The primordia of the oral
| |
| glands have fused to form a U-shaped sucker ventral and posterior to the stomodeum. The visceral grooves are still separated from the visceral pouches by closing membranes, while on
| |
| the third and fourth arches external gills have appeared. Behind
| |
| them the pronephric elevation is well marked, and continues
| |
| backward as a slight ridge marking the pronephric duct.
| |
| Intersomitic grooves are still apparent. On the ventral side
| |
| at the base of the tail is the cloacal aperture.
| |
| | |
| Prosencephalon
| |
| | |
|
| |
|
| |
|
| |
|
| |
| | |
| Optic cup
| |
| | |
| Mesencephalon
| |
| é
| |
| | |
| Otic vesicle a
| |
| | |
| Heart
| |
| Rhombencephalon S External gills
| |
| Pronephros i a— Liver
| |
| | |
|
| |
| | |
| Yolk
| |
| | |
| Myotomes
| |
| | |
| A
| |
| Fig. 188. — 6 mm. frog larva (just hatched). Transparent preparation, viewed from
| |
| right side. X15.
| |
| | |
| Endodermal derivatives. — On either side of the fore-gut are
| |
| to be seen five visceral pouches, although they would hardly be
| |
| recognized as such since they are so compressed. A groove on
| |
| the ventral side of the pharyngeal cavity is the primordium of
| |
| the thyroid gland. At this stage, also, the dorsal epithelial
| |
| 282 THE ANATOMY OF FROG EMBRYOS
| |
| | |
| bodies of the first two visceral pouches (hyomandibular and first
| |
| branchial) may be distinguished. The liver diverticulum has
| |
| increased in length. The hind-gut has lost its connection with
| |
| the neurocoel through the occlusion of the neurenteric canal, but
| |
| now receives the posterior ends of the pronephric ducts.
| |
| | |
| Mesodermal derivatives. — The notochord has grown back
| |
| into the tail. The somites have now become differentiated into
| |
| the myotomes, dermatomes, and sclerotomes, while from the
| |
| myotomes muscle cells have been formed. The pronephros is
| |
| now established. There are three pronephric tubules, each
| |
| opening into the coelom by means of a ciliated nephrostome.
| |
| Opposite these, a mass of capillaries, connected with the dorsal
| |
| aorta, forms the so-called glomus, equivalent to the separate
| |
| glomeruli of other vertebrates. The pronephric tubules grow
| |
| backward into the pronephric ducts, which have acquired lumina.
| |
| At the time of hatching, the primordia of the heart have fused
| |
| to form a tube, twisted slightly and almost S-shaped, suspended
| |
| in the pericardial cavity by a dorsal mesocardium. ‘Two regions
| |
| may be distinguished, the posterior atrium and anterior ventricle. From the ventricle leads the bulbus, arising from the
| |
| fusion of paired primordia. This connects with the dorsal aorta,
| |
| also the result of fusion, by means of aortic arches in the third
| |
| and fourth visceral arches (vestiges of the first and second aortic
| |
| arches have already appeared and disappeared). At a slightly
| |
| later stage, loops from these arches will grow out into the external gills to form a branchial circulation. The anterior ends of
| |
| the dorsal aortae are prolonged to form the internal carotids,
| |
| while the posterior ends unite directly above the heart, and just
| |
| after uniting give off the glomi on either side. Both the somatic
| |
| and splanchnic venous systems are represented at this stage.
| |
| Two vitelline veins unite to enter the heart at the sinus venosus.
| |
| The cardinal veins at this time are represented by irregular
| |
| lacunar spaces in the head and near the pronephros.
| |
| | |
| Ectodermal derivatives.— The epidermis is still ciliated.
| |
| From the prosencephalon the thin-walled cerebral vesicle has
| |
| appeared. The epiphysis is well marked, and the infundibulum
| |
| is in contact with the hypophysis. At this time the primordia
| |
| of cerebrospinal nerves may be distinguished. In the spinal
| |
| nerves, dorsal roots arise from the ganglia produced by the segTHE LARVA AT HATCHING
| |
| | |
|
| |
| | |
| 283
| |
| Infundibulum
| |
| Epiphysi
| |
| Mesencephalan. piphysis
| |
| Prosencephalon
| |
| Rhombencephalon
| |
| Fore gut
| |
| Oral gland
| |
| | |
| Heart
| |
| | |
| Notochord
| |
| | |
| Liver
| |
| | |
| Sittegretee SEN
| |
| Bengawere ee
| |
| | |
| ee
| |
| | |
| pear
| |
| =
| |
| a
| |
| 2
| |
| | |
| os
| |
| | |
| a ‘5-3 eo
| |
| Re Od
| |
| SA eGo
| |
| | |
| a Hoe Raa pide
| |
| Pina SRL
| |
| | |
| eC
| |
| PRES ee
| |
| | |
| BJ
| |
| | |
| eS
| |
| Fs
| |
| “
| |
| | |
| i)
| |
| ce ‘
| |
| ret
| |
| OKs Ve
| |
| on
| |
| | |
| a
| |
| | |
| A)
| |
| | |
| a
| |
| | |
| Yolk
| |
| | |
| Ps
| |
| ae
| |
| oe
| |
| | |
| Fig. 189. — 6 mm. frog larva.
| |
| | |
| Sagittal section, anterior portion. 50.
| |
| 284 THE ANATOMY OF FROG EMBRYOS
| |
| | |
|
| |
|
| |
|
| |
| | |
| Prosencephalon
| |
| | |
| Optic
| |
| cup
| |
| Lens
| |
| Optic . ZH
| |
| stalk FF 5 —_ Notochord
| |
| | |
| Fore-gut
| |
| | |
| Stomodeum
| |
| | |
| Fig. 190. — 6 mm. frog larva. Transverse section through optic cup. 50.
| |
| | |
| Otic vesicle
| |
| | |
| cavity
| |
| | |
| Fig. 191. — 6 mm. frog larva. Transverse section through otic vesicle. 50.
| |
| THE LARVA AT HATCHING 285
| |
| | |
| mentation of the neural crest while the ventral roots arise from
| |
| neuroblasts in the spinal cord. In the head, four ganglia arise
| |
| and with each is associated a placode of nervous ectoderm. From
| |
| the first ganglion and placode, the trigeminal (V) nerve arises.
| |
| The second combination gives rise to the facial (VII) and acoustic
| |
| (VIII) cranial nerves, while the remainder of this placode invaginates to form the otic vesicle. The third ganglion and placode produce the glossopharyngeal (IX) cranial nerve, and the
| |
| | |
| Pronephric
| |
| tubules
| |
| | |
|
| |
| | |
| Fig. 192. — 6 mm. frog larva. Transverse section through pronephros. 50.
| |
| | |
| fourth gives rise to the vagus (X). The fourth placode grows
| |
| back as far as the tail, giving off as it goes small groups of cells
| |
| which later become the lateral line organs of the trunk. Those
| |
| of the head arise from the second and third placodes. At this
| |
| time, also, ganglion cells are migrating toward the dorsal aorta to
| |
| aggregate as the ganglia of the autonomic nervous system. The
| |
| eye is well advanced in development, as the optic vesicles have
| |
| invaginated to form the optic cup and the lens placode has separated from the epidermis and acquired a cavity. The ear is in
| |
| the otic vesicle stage with an endolymphatic duct. The nose is
| |
| still represented by the nasal pits. From the prolongation of
| |
| the fourth placode referred to above, the lateral line system is
| |
| in process of formation.
| |
| 286 THE ANATOMY OF FROG EMBRYOS
| |
| | |
|
| |
|
| |
|
| |
|
| |
| | |
| Visceral 1
| |
| pouch
| |
| | |
| I Visceral
| |
| arch
| |
| | |
| Pronephric
| |
| tubules
| |
| | |
| Segmental
| |
| muscles
| |
| | |
| Fig. 193. —6 mm. frog larva. Frontal section through nasal pit and visceral
| |
| | |
| pouches. 450.
| |
| THE YOUNG TADPOLE 287
| |
| | |
| C. THE YOUNG TADPOLE (11 MM.)
| |
| | |
| External form. — The head and trunk are now fused into a
| |
| common ovoid mass, sharply distinguished from the long bilaterally compressed tail. The mouth is open and equipped with
| |
| horny raspers, while the oral gland is reduced to two vestiges
| |
| on the ventral side of the head. On the dorsal surface, the large
| |
| eyes, now functional, protrude slightly. Anterior to these are
| |
| the external openings of the nasal tubes (external nares). The
| |
| external gills, which were developing at hatching, have now degenerated and been replaced by internal gills concealed from view
| |
| by the opercula. On the left side, the opercular aperture serves
| |
| as a means of egress for the water from which the gills obtain
| |
| their oxygen. The tail, now two-thirds the length of the tadpole, has a dorsal and a ventral fin. Close to the margin of the
| |
| latter, at the base of the tail, is the cloacal opening.
| |
| | |
| Endodermal derivatives. — The mouth has been formed by
| |
| the breaking through of the oral membrane. From the pharynx,
| |
| all the visceral pouches except the hyomandibular and the
| |
| vestigial sixth pouch open to the exterior as visceral clefts; and
| |
| demibranchs have arisen on the anterior and posterior margins of
| |
| the third, fourth, and fifth visceral arches and on the anterior
| |
| margin of the sixth. These are the internal gills which hang down
| |
| into the opercular cavity. The epithelial bodies from the hyomandibular pouch have disappeared. Those from the second pouch
| |
| form the thymus gland, while similar buds arise from the third
| |
| and fourth but presently disappear. The ventral epithelial
| |
| bodies of the second pouch are said to give rise to the carotid
| |
| gland, and those of the third and fourth to “ parathyroids.”
| |
| The fifth pouch never gains communication with the exterior
| |
| but gives rise to the ultimobranchial bodies. The thyroid is
| |
| now separated from the pharynx. In the tadpole the pulmonary
| |
| organs consist of a pair of thin-walled sacs, the lungs, arising
| |
| from a laryngeal cavity connected with the pharynx by a narrow
| |
| opening, the glottis. Posterior to the pharynx comes the esophagus, which was occluded just before the opening of the mouth but
| |
| now possesses a narrow lumen opening into the stomach, which is
| |
| not greatly dilated. The vesicle, which formerly represented the
| |
| liver, persists as the gall bladder and common bile duct, rela288 THE ANATOMY OF FROG EMBRYOS
| |
| | |
|
| |
| | |
| Internal
| |
| gills
| |
| | |
| Intestine
| |
| | |
| Fig. 194. — 11 mm. frog larva.1_ Transparent preparation viewed from right side.
| |
| X15.
| |
| | |
| 1 Figs. 194-198 inclusive are from preparations loaned me by Dr. A. R. Cahn.
| |
| In earlier editions they were labelled 9 mm., as measured after preservation.
| |
| THE YOUNG TADPOLE 289
| |
| | |
|
| |
| | |
| Infundibulum
| |
| | |
| Stomach
| |
| Notochord
| |
| Intestine
| |
| Dorsal aorta Yolk
| |
| Muscles
| |
| of tail
| |
| | |
| ie
| |
| Fig. 195. — 11 mm. frog larva. Sagittal section, anterior part. 40.
| |
| 290 THE ANATOMY OF FROG EMBRYOS
| |
| | |
| tively small in comparison with the great glandular mass of the
| |
| liver. Although the pancreas arose from paired primordia of the
| |
| duodenum, these have now shifted their position so that their
| |
| ducts open into the common bile duct. The intestine is extremely long and coiled into a double spiral. It terminates in a
| |
| slightly dilated rectum, opening into the cloacal cavity which also
| |
| receives the pronephric ducts and opens to the exterior by the
| |
| cloacal aperture.
| |
| | |
| Mesodermal derivatives. — The notochord has elongated toward the posterior end, accompanying the growth of the tail.
| |
| The two most anterior somites have disappeared, leaving eleven
| |
| in the trunk region and a much larger and variable number in
| |
| | |
|
| |
| | |
| Fig. 196. — 11 mm. frog larva. Transverse section, through eye. X40.
| |
| | |
| the tail. In the tail the myotomes have given rise to the dorsal
| |
| and ventral musculature. The pronephros has become larger
| |
| and more complicated through the branching of the pronephric
| |
| tubules. The coelom consists of a pericardial cavity containing
| |
| the heart, whose myocardia have disappeared, and an abdominal
| |
| cavity in which the gut is suspended by the dorsal mesentery.
| |
| These cavities are still continuous up to the time of metamorphosis. In the heart the sinus venosus is now a large transverse sac;
| |
| the atrium is partially divided by the interatrial septum; the
| |
| ventricle has thick muscular walls; and the short bulbus opens
| |
| into the ventral aorta which is divided into proximal and distal
| |
| portions by a pair of valves. The ventral aorta is divided into
| |
| THE YOUNG TADPOLE 291
| |
| | |
| four afferent branchial arteries, the ventral portions of aortic
| |
| arches III-VI. From these the blood passes through the internal
| |
| gills by means of capillaries and is conveyed to four efferent
| |
| branchial arteries, the dorsal portions of the aortic arches referred
| |
| to above, which in turn lead to the dorsal aortae. The carotid
| |
| arteries are connected in front of and behind the infundibulum by
| |
| commissural vessels, and continue forward as the anterior cerebral arteries. From the anterior commissure the basilars run
| |
| backward and the anterior palatines forward. The pharyngeal
| |
| | |
| Myelencephalon
| |
| | |
|
| |
| | |
| Otic vesicle
| |
| | |
|
| |
| | |
| Horizontal
| |
| | |
| Fig. 197. — 11 mm. frog larva. Transverse section through ear. X40.
| |
| | |
| artery, running forward from the dorsal aorta, at a point just
| |
| posterior to the anterior commissure, represents the dorsal portion
| |
| of the mandibular arch; the lingual artery arises independently
| |
| and unites with the first efferent branchial. From the efferent
| |
| branchial arteries of the sixth arch, the pulmonary arteries grow
| |
| backward to the lungs. The vitelline veins have been broken
| |
| up, by their inclusion in the developing liver, into hepatic veins,
| |
| opening into the sinus venosus, and hepatic-portal veins from the
| |
| intestine. The anterior cardinal veins are formed by the union
| |
| of the superior jugular and facial veins and empty into the common cardinals. From the ventral side of the head the inferior
| |
| jugulars drain into the common cardinals. The posterior somatic
| |
| veins are the posterior cartlinals, which return the blood from the
| |
| 292 THE ANATOMY OF FROG EMBRYOS
| |
| | |
| region of the pronephros into the common cardinals. The lymphatic vessels of the tadpole have arisen from the confluence of
| |
| numerous, small intercellular spaces in the mesenchyme.
| |
| Ectodermal derivatives. — The epidermis is no longer ciliated.
| |
| The cerebral vesicle is now subdivided into right and left portions, while immediately behind this is the choroid plexus of
| |
| the diencephalon. The pineal gland is connected with the
| |
| diencephalon by:a small stalk; the pituitary gland has lost all
| |
| connection with the exterior. In the mesencephalon the optic
| |
| | |
| Neural tube
| |
| | |
|
| |
|
| |
| | |
| Notochord
| |
| | |
|
| |
|
| |
| | |
| Intestine
| |
| | |
| Fig. 198. — 11 mm. frog larva. Transverse section through pronephros. X40.
| |
| | |
| lobes are just apparent. The metencephalon is distinguishable
| |
| by the thickness of its walls as compared with the choroid plexus
| |
| of the myelencephalon. All cranial nerves and spinal nerves
| |
| are now established. The eye now contains all elements necessary for functioning; rods and cones of the sensory layer connect
| |
| with the neurons of the optic nerve; pigment is deposited in
| |
| the pigment layer; the choroid and sclerotic layers have been
| |
| formed from mesenchyme; the lens is transparent, as is the
| |
| cornea formed from the ectoderm. The otocyst is partially
| |
| divided by a dorsal partition into an outer saccule and inner
| |
| utricle. The nasal pits have grown backward as solid rods
| |
| which by now have acquired lumina and will soon open into the
| |
| | |
| pharynx.
| |
| REFERENCES 293
| |
| | |
| Diencephalon
| |
| | |
| Ear
| |
| | |
| Notochord
| |
| | |
| Pronephros
| |
| | |
|
| |
| | |
| Wall of
| |
| Intestine
| |
| | |
| Fig. 199. — 11 mm. frog larva. Trontal section through nose, eye, and ear. 40.
| |
| | |
| REFERENCES
| |
| | |
| Huxley, J. S., and de Beer, G. R. 1934. The Elements of Experimental Embryology, Chap. 2.
| |
| | |
| Jenkinson, J. W. 1913. Vertebrate Embryology, Chap. 7.
| |
| | |
| Kellicott, W. E. 1913. Chordate Development.
| |
| | |
| Marshall, A. M. 1898. Vertebrate Embryology, Chap. 3.
| |
| | |
| McEwen, R.S. 1931. Vertebrate Embryology, 2nd Ed., Part 2.
| |
| | |
| Morgan, T. H. 1897. The Development of the Frog’s Egg.
| |
| | |
| Zeigler, H. E. 1902. Lehrbuch der vergleichenden Entwickelungsgeschichte der
| |
| niederen Wirbeltiere. .
| |
| CHAPTER XII
| |
| THE ANATOMY OF CHICK EMBRYOS
| |
| | |
| The traditional stages in the development of the chick Gallus
| |
| domesticus) for laboratory practice are those at the end of each
| |
| of the first three days of incubation. So many important changes
| |
| take place during the second day, however, that it is advisable
| |
| to study an additional stage intermediate between twenty-four
| |
| and forty-eight hours in age. The chick of thirty-three hours
| |
| is selected because the form of the embryo is not yet affected by
| |
| torsion or flexure, and the headfold of the amnion has not yet
| |
| slipped over the head of the chick.
| |
| | |
| As it is a well-known fact that, in these first few days of incubation, embryos of the same age have attained varying degrees of
| |
| development, the length of the embryo has been proposed as a
| |
| mark of identification. The flexures of the body, however,
| |
| make this standard impracticable, and the remaining alternative
| |
| is to select the specific development of some particular structure
| |
| as a basis of arrangement. For this purpose the number of
| |
| somites, suggested by Lillie, is admirable. Still, it must be remembered that on account of the effect of temperature upon the
| |
| rate of development, the number of somites is not correlated
| |
| exactly with the number of hours of incubation, as may be seen
| |
| from the following table.
| |
| | |
|
| |
| | |
|
| |
| | |
| TABLE 12
| |
| Duval Keibel Lillie Patten
| |
| About 24 hours Fig. 76 Vig. 9, 9A Vig. 59 Fig. 36
| |
| | |
| (24 hrs. 6S) (24 hrs. 7-88) (25 hrs. 7S) (27 hrs. 8S)
| |
| | |
|
| |
| | |
| About 33 hours Fig. 93 Fig. 10, 10A Fig. 68 Fig. 39
| |
| (33 hrs. 168) (32 hrs. 9 S) (33 hrs. 128) (33 hrs. 128)
| |
| | |
|
| |
| | |
| About 48 hours Fig. 109 Fig. 16, 16A Fig. 93 Fig.
| |
| (48 hrs. 27-28S)| (52 hrs. 278) a8 hrs. 278) (55 hrs. 2 S)
| |
| | |
|
| |
| | |
| About 72 hours Fig. 115 Fig. 18, 18A g. 117 g. 63
| |
| (68 hrs. 37S) {(67 hrs. 35-37 S) (ak 18s, 35S) ah ee 368)
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
| ° 294
| |
| TWENTY-FOUR HOURS 295
| |
| | |
| A. THE TWENTY-FOUR HOUR STAGE
| |
| | |
| At the end of the first day of incubation, the chick embryo has
| |
| completed the period of cleavage (pages 98, 105) and germ-layer
| |
| formation (pages 111, 121), and is in the early stages of organogeny.
| |
| | |
| Anterior neuropore
| |
| | |
| Head fold Proamnionrn t hy
| |
| | |
|
| |
| | |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
| | |
| Amnio te Anteriorcardiac ntestinal portal
| |
| vesicle Neural fold
| |
| | |
| Neural
| |
| groove
| |
| | |
| 4th
| |
| Somite
| |
| | |
| Area &
| |
| pellucida
| |
| Primitive
| |
| knot
| |
| | |
| Primitive
| |
| streak
| |
| | |
| Area
| |
| vasculosa
| |
| | |
| Fie. 200. — 24 hour chick embryo. Cleared preparation from dorsal side. X25.
| |
| | |
| External form. — The embryo, 3.3 mm. in length, lies along
| |
| the axial line of the slipper-shaped area pellucida which in turn is
| |
| surrounded by the crescent-shaped area vasculosa, whose anterior
| |
| horns, separated by the proamnion, reach about to the level of
| |
| 296 THE ANATOMY OF CHICK EMBRYOS
| |
| | |
| tip of the head. At the anterior end, the head fold of the embryo
| |
| is lifted above the proamnion from which it is separated by the
| |
| subcephalic pocket. In the head fold is contained the fore-gut,
| |
| | |
| 0.59 mm. in length, which opens at its
| |
| | |
| posterior end into the yolk cavity by
| |
| | |
| means of the anterior intestinal portal.
| |
| | |
| On either margin of the portal the pri
| |
| mordia of the vitelline veins are to be
| |
| | |
| recognized in thick bands of splanchnic
| |
| mesoderm. The neural plate has already given rise to the neural folds
| |
| which extend back as far as the first
| |
| somite. They have united just posterior
| |
| to the region where the optic vesicles are
| |
| _ to appear and thus have given rise to a
| |
| neural tube 0.3 mm. in length, which is
| |
| widely open in front and behind as the
| |
| anterior and posterior neuropores, respectively. Behind the head fold the
| |
| axial mesoderm is segmented into six
| |
| somites. Between the neural folds the
| |
| notochord can be recognized as a faint
| |
| line which joins, at its posterior end, the
| |
| | |
| primitive streak, now reduced to 0.83
| |
| | |
| mm. in length.
| |
| | |
| Endodermal derivatives. — The only
| |
| differentiation which has taken place in
| |
| the endoderm consists of the establishment of the fore-gut by means of the
| |
| folding off of the head from the proamnion. As this process continues the fore
| |
| gut will be lengthened at the expense of
| |
| | |
| Fia. 201. — 24 hour chick em- the widely open mid-gut, and the an
| |
| ean Sagittal section. terior intestinal portal: will progress
| |
| steadily backward.
| |
| | |
| Mesodermal derivatives. — The mesoderm proper does not
| |
| extend into the head, but a loose aggregate of mesenchyme
| |
| derived from it is present. Posterior to the head the axial mesoderm is divided into six somites. Transverse sections show that
| |
| TWENTY-FOUR HOURS 297
| |
| | |
| Epidermis , , Brain
| |
| | |
|
| |
|
| |
|
| |
|
| |
| | |
| Mesenchyme
| |
| | |
| Splanchnic
| |
| mesoderm
| |
| | |
| Proamnion
| |
| | |
|
| |
| | |
| Somatic mesoderm
| |
| Fore-gut
| |
| | |
|
| |
| | |
| Ectoderm Endoderm.
| |
| | |
| Fig. 202. — 24 hour chick embryo. Transverse section through brain region. The
| |
| neural folds have met but are not yet fused together. X50.
| |
| | |
|
| |
| | |
| Axial mesoderm Notochord
| |
| ge ie
| |
| See
| |
| SO ete Ry
| |
| oe ayer
| |
| Vitelline vein Amnio-cardiac Splanchnopleure
| |
| vesicle
| |
| | |
| Fig. 203. — 24 hour chick embryo. Transverse section through region of intestinal
| |
| portal. X50.
| |
| | |
| Neural groove
| |
| | |
| | Somite IY
| |
| Ectoderm
| |
| | |
| Exocoel
| |
| | |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
| | |
| PO Blood island
| |
| nom i ram) Canes.
| |
| | |
| Notochord So
| |
| Endoderm ~
| |
| | |
| Fig. 204. — 24 hour chick embryo. Transverse section through fourth somite.
| |
| X50.
| |
| | |
| Intermediate mesoderm
| |
| | |
| Primitive groove
| |
| | |
| Endoderm
| |
| | |
| HH, BY
| |
| Fig. 205. — 24 hour chick embryo. Transverse section through primitive streak.
| |
| x50.
| |
| 298 THE ANATOMY OF CHICK EMBRYOS
| |
| | |
| each has a minute cavity, or myocoel. The intermediate mesoderm does not divide into nephrotomes as in the frog. The
| |
| lateral mesoderm is divided into the somatic and splanchnic
| |
| layers. In the latter, numerous blood islands appear and give
| |
| the characteristic mottled appearance to the area vasculosa. The
| |
| coelom of the embryo is continuous with that of the extra-embryonic regions, or exocoel. In the region on either side of the head,
| |
| between the proamnion and the intestinal portal, the coelom is
| |
| distended into an amniocardiac vesicle, so called because the somatopleure will contribute to the head fold of the amnion, while
| |
| the splanchnic mesoderm will give rise to the primordia of the
| |
| heart, and the cavities of the vesicles will unite to form the
| |
| pericardial cavity. The notochord, from its point of origin, the
| |
| primitive streak, extends forward into the head.
| |
| | |
| Ectodermal derivatives. — The ectoderm at this stage consists of the elongate neural plate, with its groove and folds which
| |
| are already in process of fusion, and the epidermis or non-nervous
| |
| ectoderm.
| |
| | |
| B. THE THIRTY-THREE HOUR STAGE
| |
| | |
| External form. — In the chick embryo, after thirty-three hours’
| |
| incubation, the length has increased to 4.3 mm. There is a
| |
| slight bending of the head downward over the end of the notochord, foreshadowing the cranial flexure. The area vasculosa,
| |
| in which the blood islands are being converted into capillaries,
| |
| now has grown in toward the embryo, so that the area pellucida
| |
| persists only around the head and tail regions. The anterior
| |
| horns of the area vasculosa have met in front, completely inclosing the proamnion. The head has increased in length not
| |
| only by actual forward growth but also by the backward extension
| |
| of the lateral margins of the head fold, so that the enclosed foregut is now 1 mm. long. The vitelline veins are prominent at
| |
| the margins of the intestinal portal and continue on the ventral
| |
| side of the fore-gut to meet at the posterior end of the heart,
| |
| which is now a single tube, slightly bent toward the right. The
| |
| neural folds are fused as far back as the eleventh somite, where
| |
| the posterior neuropore is now known as the rhomboidal sinus.
| |
| The anterior neuropore is about to close, and in the head the
| |
| neural tube shows three regions of dilation which represent the
| |
| THIRTY-THREE HOURS 299
| |
| | |
| Head fold |.
| |
| of amnion
| |
| | |
|
| |
|
| |
| | |
| 4 neuropore
| |
| Prosencephalon
| |
| : Optic
| |
| 7 vesicle
| |
| Mesencephalon
| |
| Foregut
| |
| Rhombencephalon
| |
| & Heart
| |
| | |
| - Vitelline
| |
| vein
| |
| | |
| Somite 6
| |
| | |
| Sinus
| |
| rhomboidialig
| |
| | |
| Primitive
| |
| streak
| |
| | |
| Fig. 206. — 33 hour chick embryo. Cleared preparation from dorsal view. X25.
| |
| 300 THE ANATOMY OF CHICK EMBRYOS
| |
| | |
| fore-brain, mid-brain, and hind-brain, respectively. The sides
| |
| of the fore-brain are evaginating to produce the optic vesicles.
| |
| | |
| Head fold .
| |
| of amnion
| |
| | |
| Prosencephalon—
| |
| Subcephalic F
| |
| pocket
| |
| Mesencephalon
| |
| Fore-gut
| |
| Pericardial
| |
| cavity
| |
| | |
|
| |
| | |
| Rhombencephalon# 6.4. /"
| |
| | |
| Notochord
| |
| | |
| Fig. 207. — 33 hour chick embryo.
| |
| | |
| Sagittal section.
| |
| | |
| are ~ Anterior
| |
| | |
| intestinal
| |
| portal
| |
| | |
| i Primitive
| |
| cm, streak
| |
| | |
| aN
| |
| | |
| X25.
| |
| | |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
| | |
| In the hind-brain, five neuromeres
| |
| can be identified. Twelve somites
| |
| may be counted. The notochord
| |
| extends forward to the fore-brain
| |
| from the primitive streak which is
| |
| now reduced to 0.3 mm.
| |
| | |
| Endodermal derivatives. — The
| |
| anterior end of the fore-gut is in
| |
| contact ventrally with the stomodeum separated only by the oral
| |
| plate, composed of ectoderm and
| |
| endoderm. At the sides, the walls
| |
| of the fore-gut are fused to the ectoderm at points where the first visceral pouches (hyomandibular) will
| |
| be located.
| |
| | |
| Mesodermal derivatives. — The
| |
| somites now number twelve, and
| |
| myocoels are still apparent. The
| |
| mesomere is still unsegmented, but
| |
| pronephric tubules have appeared
| |
| in the region corresponding to somites 5-12. The four posterior
| |
| tubules are growing back to form
| |
| the pronephric duct. In the
| |
| splanchnic mesoderm the blood
| |
| islands are being converted into
| |
| capillaries. The vitelline veins are
| |
| prominent and continue forward
| |
| into the heart, of which the endo
| |
| -cardium and myocardium are dis
| |
| tinct. The heart is supported by
| |
| the dorsal mesocardium, the ventral
| |
| mesocardium having disappeared.
| |
| The primordial tubes, from the
| |
| | |
| fusion of which the heart arose, continue forward as the ventral
| |
| aortae which bend around the pharynx (first aortic arches) and
| |
| THIRTY-THREE HOURS 301
| |
| | |
| continue backward along the dorsal surface of the pharynx as the
| |
| dorsal aortae. At the level of the primitive streak they are lost
| |
| in a capillary nexus which foreshadows the vitelline arteries.
| |
| From a point immediately in front of the optic vesicle, the anterior
| |
| cardinals course backward on either side of the neural tube, bending down ventrally to enter the heart with the vitelline veins.
| |
| The notochord is slightly bent at the anterior end.
| |
| | |
| Ectodermal derivatives. The ncural folds now extend to
| |
| the eleventh somite and have fused throughout the length of the
| |
| head. The anterior neuropore is almost closed. The three
| |
| | |
| Prosencephalon
| |
| | |
| Epidermis Mesenchyme
| |
| | |
| Optic vesicle
| |
| | |
| =» _
| |
| | |
| ts wa
| |
| P Exocoel é OOF
| |
| | |
|
| |
| | |
| Splanchnopleure Sub-cephalic
| |
| pocket
| |
| | |
| Fia. 208. — 33 hour chick embryo. Transverse section through optic vesicles.
| |
| X50.
| |
| | |
| dilations which represent the prosencephalon, mesencephalon,
| |
| and rhombencephalon are distinct. From the prosencephalon
| |
| the two optic vesicles extend to the ectoderm of the sides of the
| |
| head. Five neuromeres may be identified in the rhombencephalon. It has been asserted that in earlier stages three neuromeres may be identified in the prosencephalon and two in the
| |
| mesencephalon, while the first of the five noted above has resulted from the fusion of two original neuromeres destined to
| |
| give rise to the metencephalon. At about this time a shallow
| |
| depression in the floor of the prosencephalon, just in front of the
| |
| tip of the notochord, marks the appearance of the infundibulum.
| |
| The auditory placodes may sometimes be seen in sections as
| |
| thickenings at the level of the constriction separating the last two
| |
| neuromeres on either side.
| |
| 302 THE ANATOMY OF CHICK EMBRYOS
| |
| | |
| Rhombencephalon
| |
| | |
|
| |
|
| |
|
| |
|
| |
|
| |
| | |
|
| |
|
| |
|
| |
| | |
| Notochord
| |
| Fore-gut Otic ( auditory) placode
| |
| Somatopleure TET Dorsal aorta
| |
| Dorsal [Bi soy 7S uateral sulcus
| |
| mesocardium B Feria
| |
| : 5 Way, Oe
| |
| ee ae
| |
| ox ed fhe & wou
| |
| ee se
| |
| Os a lamest ~
| |
| Endocardium Splanchnopleure
| |
| Fig. 209. — 33 hour chick embryo. Transverse section through otic placodes.
| |
| | |
| x50.
| |
| | |
| Spinal cord
| |
| | |
|
| |
|
| |
|
| |
|
| |
| | |
| Dorsal aorta Somite
| |
| | |
| Intermediate mesoderm
| |
| Exocoel
| |
| | |
| Se
| |
| | |
| Vitelline vein
| |
| | |
| Fie. 210. — 33 hour chick embryo. Transverse section through vitelline veins.
| |
| x50.
| |
| | |
| Spinal cord
| |
| | |
| Neural crest Somite
| |
| | |
|
| |
|
| |
|
| |
| | |
| Intermediate mesoderm
| |
| Somatic layer
| |
| | |
| NAPRY
| |
| | |
| Fig. 211. — 33 hour chick embryo. Transverse section through sixth somite. 50.
| |
| FORTY-EIGHT HOURS . 803
| |
| | |
| Cc. THE FORTY-EIGHT HOUR STAGE —
| |
| External form. — The chick at the end of the second day of
| |
| incubation has usually attained a length of 7 mm., but the form
| |
| | |
| of the body has been altered profoundly. As the head has been
| |
| lifted away from the blastoderm, it has increased greatly in size,
| |
| | |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
| | |
|
| |
| | |
| Ww ir ot
| |
| y
| |
| h h, Mesencephalon
| |
| ‘Rhombencephalon
| |
| | |
| Otic vesicle Optic cup
| |
| | |
| Lens vesicle
| |
| Visceral cleft I Prosencephalon
| |
| | |
| 0
| |
| i
| |
| | |
| Sinus venosus—
| |
| Vitelline vein—
| |
| | |
| Atrium
| |
| Bulbus arteriosus
| |
| Ventricle
| |
| | |
| Amniotic fold
| |
| Somite XIV
| |
| | |
| Vitelline artery
| |
| | |
| Neural tube
| |
| | |
| Tail fold
| |
| | |
| aed cent
| |
| Fia. 212.—48 hour chick embryo. Transparent preparation from dorsal view
| |
| (head from right side). X15.
| |
| | |
| and the cranial flexure, which was just appearing in the thirtythree hour chick, has become so pronounced that the anterior
| |
| end of the head is directed backwards. With this growth and
| |
| flexure the head is twisted normally to the right, until it lies on
| |
| one side, a phenomenon known as torsion. At forty-eight hours,
| |
| this torsion involves the chick as far back as the seventeenth
| |
| somite. The posterior end of the chick lies in its original position,
| |
| and at the extreme caudal end a tail fold is being formed. In the
| |
| Fig. 213. — 48 hour chick embryo.
| |
| (304)
| |
| | |
| frontal section due to torsion.
| |
| | |
| x50.
| |
| | |
| Head in sagittal section, somite region in
| |
| | |
| Stomodaeal
| |
| plate
| |
| Telencephalon.
| |
| | |
|
| |
| | |
| Diencephalon
| |
| | |
| Myelencephalon
| |
| | |
|
| |
| | |
| Metencephalon
| |
| | |
|
| |
| | |
|
| |
| | |
|
| |
| | |
| Mesencephalon
| |
| FORTY-EIGHT HOURS 305
| |
| | |
| area vasculosa the capillaries have formed attachments with the
| |
| vitelline arteries and veins, and at the border of this area is a circular vessel, the sinus terminalis. The fore-gut is now 1.4 mm.
| |
| in length, and the first of the three visceral pouches now communicates to the exterior following the rupture of the closing
| |
| plate which separated it from the corresponding visceral groove.
| |
| The second and third visceral grooves are apparent, but their closing plates are still unperforated. In the visceral arches the first
| |
| three aortic arches are apparent, arising from the ventral aorta.
| |
| The heart is now twisted so that the ventricular loop is upper
| |
| Anterior cardinal vein Dorsal aorta
| |
| | |
| Otic pit
| |
| | |
|
| |
|
| |
|
| |
|
| |
| | |
|
| |
|
| |
| | |
| eB
| |
| 5
| |
| | |
| Chorion
| |
| | |
|
| |
|
| |
| | |
| Yolk sac tent
| |
| Notochord eet Blood island
| |
| Visceral groove I a Pigment layer
| |
| | |
| Visceral pouch I Hypophysis Sensory layer
| |
| | |
|
| |
| | |
| Fia. 214. — 48 hour chick embryo. Transverse section through otic pit and optic
| |
| cup. 650.
| |
| | |
| most. The vitelline veins are large and conspicuous, as are the
| |
| vitelline arteries which leave the body at the level of the twentysecond somites. The neural tube is completely closed. In the
| |
| head the five definitive regions of the brain are outlined, the
| |
| prosencephalon having given rise to the telencephalon and diencephalon, and the rhombencephalon to the metencephalon and
| |
| myelencephalon. The eye is now in the optic cup stage, and the
| |
| invagination of the optic vesicle continues down the stalk to form
| |
| the choroid fissure. The lens is in the form of a pit which has
| |
| almost attained the vesicle stage. The ear is represented by an
| |
| otic pit which, owing to the cervical flexure, is about on a level
| |
| with the eye. There are twenty-seven somites at this stage.
| |
| The primitive streak is found only in the tail fold. At this time
| |
| 306 THE ANATOMY OF CHICK EMBRYOS
| |
| | |
| the head fold of the amnion has grown back over the chick as
| |
| far as the sixteenth somite.
| |
| | |
| Endodermal derivatives. — The stomodeum, an ectodermal
| |
| invagination from the ventral surface of the head fold, has formed
| |
| the oral membrane by contact with the fore-gut a little back of its
| |
| most anterior point. Hence there is a blind pocket in front of
| |
| the oral plate, known as the preoral gut. Three visceral pouches
| |
| are present, the first of which opens into the corresponding visceral
| |
| furrow following the rupture of its closing membrane. The
| |
| primordium of the thyroid is represented by a ventral depression
| |
| in the floor of the pharynx at the level of the second visceral
| |
| pouches. The primordia of the lungs (sometimes difficult to
| |
| distinguish) extend to the level of the sinus venosus. The liver
| |
| arises at the level of the anterior intestinal portal from two
| |
| evaginations of the endoderm, one below and one above the
| |
| meatus venosus. ‘The mid-gut now has two shifting boundaries,
| |
| the anterior intestinal portal and the posterior intestinal portal.
| |
| The latter is barely apparent as the opening of a shallow endodermal pocket or hind-gut in the tail fold.
| |
| | |
| Mesodermal derivatives.—-The somites, twenty-seven in
| |
| number, show a varying degree of specialization, with the most
| |
| advanced at the anterior end. In these two regions can be distinguished: a loose aggregate of cells at the median ventral angle
| |
| (the sclerotome); and a cap of epithelial cells at the lateral dorsal
| |
| angle. The cells of this cap nearest the epidermis will form the
| |
| dermatome, while those nearest the neural tube will form the
| |
| myotome.
| |
| | |
| The pronephric tubules in the more anterior somites have disappeared and mesonephric tubules are appearing in the mesomere posterior to the thirteenth somite. The pronephric (now
| |
| the mesonephric) duct has acquired a lumen but has not yet
| |
| attained its complete backward growth.
| |
| | |
| The heart is still tubular, but the ventricular limb of the cardiac
| |
| loop has grown back and over the atrial limb so that the ventricular region is now caudal and dorsal with relation to the
| |
| atrial region. Three aortic arches are present as a rule, but infrequently the third has not developed. From the first aortic arch
| |
| a network of capillaries extends into the head. From these the
| |
| carotid arteries will be formed. The dorsal aortae have fused
| |
| FORTY-FIGHT HOURS 307
| |
| | |
| from a point back of the sixth somite as far as the level of the
| |
| fifteenth somite. The vitelline arteries leave the dorsal aortae at
| |
| the level of the twenty-second somite but the aortae continue
| |
| | |
| Common cardinal Bulbus
| |
| vein arteriosus Chorion
| |
| : I.- poem 2s Cra, /
| |
| Dorsal aorta Fore-gut er ,
| |
| bAY
| |
| | |
| Notochord
| |
| | |
|
| |
|
| |
|
| |
|
| |
|
| |
| | |
| Spinal cord
| |
| | |
| ,
| |
| | |
| as
| |
| | |
| Epidermis 5
| |
| J
| |
| | |
| Amnion
| |
| | |
| “ Somiter
| |
| Coelom
| |
| | |
| ‘Dorsal mesocardium
| |
| Fia. 215. — 48 hour chick embryo. ‘Transverse section through heart. X50.
| |
| | |
| backward as the caudal arteries to the last somite. The vitelline
| |
| veins are fused at their point of entrance into the heart as the sinus
| |
| venosus. The anterior cardinals are prominent and extend from
| |
| a capillary plexus in the head back toward the heart, where they
| |
| | |
| Somite
| |
| | |
| Notochord | Dorsal aorta
| |
| | |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
| | |
| Vitelline vein,
| |
| | |
| Choria
| |
| | |
| - Armnion— Gi
| |
| N
| |
| Coelom —
| |
| | |
| Posterior cardinal
| |
| vein
| |
| | |
| Ventricle
| |
| | |
|
| |
| | |
| Meatus Mid-gut: *
| |
| | |
| venosus
| |
| | |
| Fig. 216. — 48 hour chick embryo. Transverse section through liver. 50.
| |
| | |
| are joined by the posterior cardinals and proceed as the common
| |
| cardinals to: enter the heart in the angles between the sinus
| |
| venosus and the vitelline veins. The posterior cardinals may be
| |
| traced back to the last somite. The heart of the chick commenced
| |
| 308 THE ANATOMY OF CHICK EMBRYOS
| |
| | |
| beating at the forty-fourth hour of incubation, so that the course
| |
| of the blood is through the ventral aorta to the aortic arches and
| |
| thence to the dorsal aorta. From the first aortic arch a network
| |
| of capillaries supplies the head with blood (which is returned by
| |
| way of the anterior cardinals). The main current of the stream
| |
| passes down the dorsal aortae to the point where these fuse to
| |
| form the median dorsal aorta. From the dorsal aorta, the somites
| |
| are supplied by capillaries, which will later become the intersegmental arteries. This blood is returned through the posterior
| |
| cardinals. Leaving the dorsal aorta by way of the vitelline
| |
| arteries, the blood passes through the capillaries of the area
| |
| vasculosa to the sinus terminalis, and thence to the capillary
| |
| drainage of the vitclline veins which return it to the heart.
| |
| | |
| The notochord is bent, not only at its tip (cranial flexure) but
| |
| also at the point where the myelencephalon merges with the spinal
| |
| cord (cervical flexure).
| |
| | |
| Ectodermal derivatives. — The brain now has acquired its
| |
| five definitive vesicles. The telencephalon is enlarged but shows
| |
| | |
| Amniotic raphe
| |
| | |
|
| |
|
| |
|
| |
| | |
|
| |
| | |
| — Posterior cardinal vein
| |
| | |
| Mesonephric tubules
| |
| | |
| Beast
| |
| | |
| Lateral sulcus
| |
| Fig. 217. —48 hour chick embryo. Transverse section through mesonephros.
| |
| 50.
| |
| | |
| no particular differentiation. From the diencephalon project the
| |
| constricted optic stalks which bear the optic cups with their inner
| |
| sensory layer and outer pigmented layer. (The pigment will
| |
| not arise until later.) The invagination by which the cups were
| |
| formed continues down the stalk as the choroid groove. On the
| |
| ventral surface of the diencephalon the infundibulum has deepened. Growing in toward it from the stomodeum is an ectodermal invagination, the hypophysis, which will fuse with the
| |
| infundibulum to form the pituitary gland. The lens of the eye
| |
| SEVENTY-TWO HOURS 309
| |
| | |
| is in the pit stage, resulting from the invagination of a sensory
| |
| placode. When the process is complete, the lens will be a vesicle
| |
| completely withdrawn beneath the surface of the ectoderm, as
| |
| will the otic vesicle, the primordium of the inner ear. Along the
| |
| rhombencephalon and cord, the neural crest is to be seen as a
| |
| narrow band of cells on each dorso-lateral angle.
| |
| | |
| Myelencephalon
| |
| | |
|
| |
| | |
| Otic vesicle %, Metencephalon
| |
| | |
| he 4 te
| |
| | |
| Visceral
| |
| cleft I *
| |
| q Mesencephalon
| |
| wy Choroid fissure
| |
| Optic cup
| |
| Atrium and lens
| |
| : Diencephalon
| |
| Nasal pit - Epiphysis
| |
| Telencephalon
| |
| Ventricle
| |
| Anterior
| |
| limb bud
| |
| 3 Somite 26
| |
| Vitelline 2
| |
| artery
| |
| Vitelline ; .
| |
| . : Posterior
| |
| vem limb bud
| |
| | |
| Fig. 218. — 72 hour chick embryo. Transparent preparation from dorsal view,
| |
| head seen from right side. X15.
| |
| | |
| D. THE SEVENTY-TWO HOUR STAGE
| |
| External form. — At the end of the third day of incubation,
| |
| the total length of the embryo is 9.5 mm., but the curvature of
| |
| the body is so great, on account of the cranial and cervical flexures in addition to the newly developed caudal flexure, that the
| |
| greatest length, from neck to tail, is 7 mm. Torsion involves the
| |
| 310 THE ANATOMY OF CHICK EMBRYOS
| |
| | |
| body as far back as the vitelline arteries and will become complete during the fourth day. Anterior and posterior limb buds
| |
| are now apparent at the levels of somites 17-19 and 26-32 respectively. The tail is curved forward. The fore-gut is still
| |
| 1.4 mm. in length but has undergone further differentiation,
| |
| indicated externally by the fact that the first three visceral
| |
| clefts are open while the fourth is still interrupted by its closing
| |
| plate. In the branchial arches four aortic arches may be seen.
| |
| The telencephalon has given rise to the primordia of the cerebral
| |
| hemispheres, and from the roof of the diencephalon, a small
| |
| evagination represents the epiphysis or primordium of the pineal
| |
| gland. The eye and ear, which were formerly in the same
| |
| transverse section, are now nearly in an antero-posterior relationship. The olfactory pits have made their appearance in the
| |
| head. The semilunar (fifth cranial nerve), geniculo-acoustic
| |
| (seventh and eighth), and petrosal (ninth) ganglia may be seen.
| |
| There are approximately thirty-five somites. The primitive
| |
| streak has disappeared. The amnion is completed by the fusion
| |
| of head and tail folds. The allantois, a small sac-like evagination,
| |
| protrudes ventrally between the posterior limb buds.
| |
| | |
| Endodermal derivatives. — At the end of the third day the oral
| |
| aperture has been formed by the rupture of the oral membrane
| |
| separating the stomodeum and the fore-gut. Immediately anterior to this opening the preoral gut persists. The fore-gut is
| |
| still the same length as in the chick of forty-cight hours, but is
| |
| more complex in structure. The thyroid gland, which appeared
| |
| during the second day, has now become differentiated into the
| |
| distal dilation which will give rise to the gland proper and the
| |
| thyroglossal duct. The first three visceral pouches are open
| |
| to the exterior, but the epithelial buds destined to give rise to the
| |
| thymus and parathyroids are not yet apparent. The fourth
| |
| visceral pouch is still separated from the corresponding groove by
| |
| the closing plate. The laryngeo-tracheal groove has developed
| |
| in the floor of the pharynx just posterior to the fourth visceral
| |
| pouches. At its posterior end the dorsal margins of this groove
| |
| have closed together to form the primordium of the trachea which
| |
| is thus set free from the esophagus above. The trachea is bifurcated at the posterior end, thus giving rise to the two bronchial
| |
| buds which are the primordia of the lungs.
| |
| SEVENTY-TWO HOURS 311
| |
| | |
| The esophagus, which is relatively narrow, is followed by a
| |
| dilation which is to become the stomach. Posterior to this, the
| |
| primordium of the liver may be seen as an evagination from the
| |
| | |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
| | |
| Aortic arches
| |
| | |
| Dorsal.
| |
| | |
| aorta Myelencephalon
| |
| | |
| Metencephalon
| |
| . Roy fandibatam
| |
| | |
| Sinus
| |
| | |
| venosus Isthmus
| |
| | |
| Atrium
| |
| Spinal
| |
| cord ,
| |
| Notochord ++—%
| |
| | |
| >Amnion
| |
| Mesencephalon
| |
| | |
| Diencephalon
| |
| | |
| Telencephalon Epiphysis
| |
| Mesonephros
| |
| | |
| Notochord
| |
| | |
| Spinal cord—
| |
| | |
| Fiq. 219. — 72 hour chick embryo. Sagittal section. X25.
| |
| | |
| ventral floor of the duodenal region of the gut. The dorsal
| |
| pancreas arises from the duodenal region just dorsal to the liver
| |
| at the end of the third day. The ventral primordia will not
| |
| appear for another day.
| |
| 312 THE ANATOMY OF CHICK EMBRYOS
| |
| | |
|
| |
|
| |
|
| |
| | |
| halon
| |
| | |
| fe
| |
| | |
|
| |
| | |
| Fig. 220. — 72 hour chick embryo. ‘Transverse section through otic vesicle. X25.
| |
| | |
| L esicle
| |
| Dorsal aorta Optic ome
| |
| | |
| Aortic arches
| |
| | |
| d cup Sensory layer
| |
| ii
| |
| | |
|
| |
| | |
| Pharynx yyy
| |
| | |
| Visceral arches
| |
| Fia. 221. — 72 hour chick embryo. Transverse section through optic cup. X25.
| |
| | |
|
| |
|
| |
|
| |
| | |
|
| |
|
| |
| | |
|
| |
|
| |
|
| |
|
| |
|
| |
| | |
|
| |
| | |
| /
| |
| Esophagus Primary ‘Common cardinal Bulbus arteriosus “
| |
| . : i
| |
| Chorion Amnion bronchus yon polite eee! oa
| |
| 3 A ang ots + ~
| |
| Somite AfSead- m,
| |
| | |
| \
| |
| rst; Epidermis
| |
| ip
| |
| | |
| 3
| |
| | |
| ‘Yolk sac
| |
| | |
|
| |
|
| |
| | |
| Pleural groove
| |
| Sinus Atrium Nasal pit
| |
| “venosus Pericardial
| |
| cavity
| |
| Fig. 222. — 72 hour chick embryo. Transverse section through heart and lung.
| |
| X25.
| |
| | |
| ‘
| |
| SEVENTY-TWO HOURS 313
| |
| | |
| The mid-gut region is gradually lessened by the advancing
| |
| sulci which are cutting off the body of the embryo from the yolk.
| |
| This region opens into the yolk stalk which is still quite wide.
| |
| | |
| The hind-gut contained in the tail fold has not yet acquired
| |
| its cloacal aperture nor has the proctodeum appeared. The
| |
| floor of the hind-gut between the tail bud and the posterior
| |
| intestinal portal evaginates to give rise to the allantoic primordium.
| |
| | |
| Mesodermal derivatives. — The somites, typically thirty-five
| |
| in number, still show a varying degree of differentiation which is
| |
| carried to its furthest point in the more anterior somites. The
| |
| dermatome is now a thin sheet of cells along the dorso-lateral
| |
| | |
| Posterior
| |
| Dorsal cardinal} Dorsal Li
| |
| _Amnion aorta vein mesentery iver
| |
| | |
|
| |
|
| |
|
| |
|
| |
|
| |
| | |
|
| |
| | |
|
| |
| | |
| ao
| |
| | |
| Spinal
| |
| cord
| |
| | |
| Notochord
| |
| | |
| Coelom
| |
| | |
| Allantoic vein
| |
| | |
|
| |
| | |
| Ventral mesentery
| |
| | |
| Meatus venosus
| |
| Fig. 223. — 72 hour chick embryo. Transverse section through liver. X25.
| |
| | |
| angle of the embryo, with the myotome parallel and internal;
| |
| the sclerotome in these anterior segments is a large and loose
| |
| aggregate of cells investing the neural tube, notochord, and
| |
| aortae.
| |
| | |
| The pronephric tubules have degenerated to a considerable
| |
| extent, but the nephrostomes opening into the coelom may persist. The mesonephric tubules are now in process of development, with those in the more anterior segments most highly
| |
| differentiated. The tubules between the thirteenth and _ thirtieth somites have progressed from the vesicle stage characteristic of those behind the twentieth somite, and some have acquired a lumen and joined the pronephric duct which henceforward is known as the mesonephric duct. A few of the more
| |
| anterior tubules develop nephrostomes, but these soon disappear.
| |
| 314 THE ANATOMY OF CHICK EMBRYOS
| |
| | |
| Behind the twentieth somite, as far back as the thirtieth, only
| |
| vesicles are formed. The mesonephric ducts have grown back
| |
| and united with the cloaca.
| |
| | |
| The heart now shows a constriction between the atrial and
| |
| ventricular region. Four aortic arches are developed, of which
| |
| | |
| Amniotic
| |
| raphe
| |
| | |
| Dermatome, Sclerotome \ Spinal cord
| |
| | |
|
| |
| | |
| Mesonephric
| |
| Vitelline tubule
| |
| artery
| |
| | |
| Lateral sulcus Dorsal
| |
| aorta
| |
| | |
| Fig. 224. — 72 hour chick embryo. Transverse section through vitelline arteries
| |
| leaving body. X25.
| |
| | |
| the first is becoming smaller, and somctimes has disappeared at
| |
| this stage. The internal carotid arteries are now well developed,
| |
| growing forward into the head from the point of union between
| |
| the first arches and the dorsal aortae. From the ventral end of
| |
| the first aortic arch the external carotid takes its origin. The
| |
| | |
| Chorion Mesonephric .
| |
| Amnion duct Somite Dorsal ao
| |
| | |
|
| |
| | |
| Fig. 225. 72 hour chick embryo. Transverse section through allantois. 25.
| |
| | |
| pulmonary is sometimes apparent as a posterior prolongation of
| |
| the ventral aorta at the point where the fifth arches will appear
| |
| during the next twenty-four hours. The intersegmental arteries
| |
| are now apparent as dorsal diverticula from the aorta between
| |
| each pair of somites. The vitelline veins have fused for a short
| |
| distance behind the sinus, thus giving rise to the meatus venosus.
| |
| REFERENCES 315
| |
| | |
| The anterior cardinal vein now possesses many branches from
| |
| the head, among which are three intersegmental veins. The
| |
| posterior cardinal has continued its backward growth dorsal
| |
| to the mesonephric duct as far as the thirty-third somite. It
| |
| receives the intersegmental veins of this region. Where the
| |
| posterior cardinals unite with the common cardinals, a capillary
| |
| network indicates the beginnings of the allantoic veins.
| |
| | |
| Ectodermal derivatives. —-'The brain at the end of the third
| |
| day has its five definitive vesicles even more sharply demarcated.
| |
| From the telencephalon two lateral vesicles have evaginated to
| |
| form the primordia of the cerebral hemispheres. In the diencephalon the epiphysis has appeared as a dorsal evagination.
| |
| On the floor of this vesicle the infundibulum is almost in contact
| |
| with the hypophysis. The mesencephalon is separated from the
| |
| metencephalon by a deep constriction known as the isthmus.
| |
| Along the sides of the myelencephalon may be distinguished the
| |
| following cerebral ganglia: the semilunar of the fifth cranial
| |
| nerve; the acoustico-facialis which will later separate into the
| |
| geniculate ganglion of the seventh and the acoustic of the eighth;
| |
| and the petrosal ganglion of the ninth. The eye has increased in
| |
| size, and the lens is now free from the epidermal ectoderm. The
| |
| ear, too, is in the vesicle stage and possesses a short endolymphatic duct, which has lost its connection with the epidermis.
| |
| On the third day the primordium of the nose is represented by
| |
| two olfactory pits anterior to the mouth.
| |
| | |
| REFERENCES
| |
| | |
| Arey, L. B. 1934. Developmental Anatomy, 3rd Ed., Chap. 18.
| |
| | |
| Duval, M. 1889. Atlas d’embryologie.
| |
| | |
| Keibel and Abraham. 1900. Normaltafeln II, des Huhnes (Gallus domesticus).
| |
| Lillie, F. R. 1919. The Development of the Chick, 2nd Ed.
| |
| | |
| McEwen, R. 8S. 1931. Vertebrate Embryology, 2nd Ed., Part 4.
| |
| | |
| Patten, B. M. 1929. The Early Embryology of the Chick, 3rd Ed.
| |
| CHAPTER XIII
| |
| THE ANATOMY OF THE 10 MM. PIG EMBRYO
| |
| | |
| Pig embryos of 10 to 12 mm. body length are particularly
| |
| instructive for laboratory work in mammalian embryology as they
| |
| | |
| Myelencephalon Metencephalon
| |
| | |
| Ear,
| |
| | |
|
| |
|
| |
| | |
| Mesencephalon
| |
| | |
| Trachea +
| |
| | |
| Anterior __| a —Body stalk
| |
| | |
| limb bud
| |
| Roots of
| |
| spinal nerves Posterior
| |
| | |
| Fig. 226. — 10 mm. pig embryo. Transparent preparation from right side. X11.
| |
| | |
| are sufficiently large for the study of external structures and yet
| |
| small enough to afford serial sections for a detailed study of the
| |
| anatomy. The primordia of practically all the organ systems,
| |
| excepting the skeleton and musculature, are present. In comparing the accounts given by different authors of this particular
| |
| stage, it should be remembered that a large amount of shrinkage
| |
| | |
| takes place during the preparation of fresh sections, so that, as
| |
| 316
| |
| ENDODERMAL DERIVATIVES 317
| |
| | |
| pointed out by Patten, an embryo of 12 mm. will not measure
| |
| more than 93} mm. when prepared for sectioning. The account
| |
| which follows corresponds in general to the pig (Sus scrofa) of
| |
| 10 mm. described by Keibel, of 12 mm. (Minot), 10 mm. (Prentiss)
| |
| and 9.4 mm. (Patten), but is not so advanced as the 13.5 mm.
| |
| pig (Boyden).
| |
| | |
| External form. — The pig embryo at this stage is relatively
| |
| ‘more advanced than the chick of seventy-two hours. The body
| |
| is sharply flexed, owing to the presence of the cranial, cervical,
| |
| dorsal, and caudal flexures. In the head region the olfactory
| |
| pits are well developed and are connected by the naso-lachrymal
| |
| groove to a depression which surrounds the bulging eyeball.
| |
| The five divisions of the brain are apparent through the relatively thin overlying epidermis. Four visceral grooves can be
| |
| seen, the first of which, or hyomandibular, is the primordium of
| |
| the external auditory meatus. The third and fourth grooves are
| |
| compressed by the cervical flexure into a deeper depression known
| |
| as the cervical sinus. A frontal view of the head shows the oral
| |
| cavity bounded above by the frontal process in the middle, the
| |
| maxillary processes at the side, while the lower jaw is represented
| |
| by the mandibular arch.
| |
| | |
| In the trunk region, the buds of the pectoral and pelvic appendages are large but show no further differentiation. The
| |
| contours of the somites, now forty-four in number, are apparent
| |
| along the back, and ventral to these can be seen the outlines of the
| |
| heart, liver, and mesonephros. In some specimens there appears
| |
| between the limb buds a thickened ridge from which the mammary
| |
| glands develop and which is therefore known as the milk line. |
| |
| The umbilical cord projects from the ventral side of the embryo.
| |
| Between this and the base of the slender tail is a small protuberance, the genital tubercle, or primordium of the external
| |
| genitalia.
| |
| | |
| Endodermal derivatives. — The preoral gut still persists anterior to the oral aperture. Ventral to this, and seen best in
| |
| sagittal section, is the long and slender hypophysis, now in contact with the infundibulum of the diencephalon. Both the
| |
| hypophysis and infundibulum, it should be remembered, are of
| |
| ectodermal origin. The pharynx is dorso-ventrally compressed,
| |
| and from its floor the tongue is arising. Four visceral pouches
| |
| 318 THE ANATOMY OF THE 10 MM. PIG EMBRYO
| |
| | |
| are present, corresponding to the visceral grooves already noted.
| |
| These do not unite to become visceral clefts but remain separated
| |
| by their closing membranes. Between the second and third
| |
| | |
| Metencephalon
| |
| Myelencephalon
| |
| | |
|
| |
|
| |
| | |
| Pharynx
| |
| | |
|
| |
| | |
| Posterior
| |
| vena cava
| |
| | |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
| | |
| //_ . Ductus venosus
| |
| | |
| i Liver
| |
| Duct of ventral pancreas
| |
| | |
| Spinal { nw ;
| |
| artery fDuodenum___Vitelline vein
| |
| on—__ Body
| |
| Dorsal stalk
| |
| pancreas i ses mbilical
| |
| SSS ) rte
| |
| Vitelline 4 7 SN d Af ty
| |
| (ant. mesenteric)’ oN i R
| |
| artery cl
| |
| Notochord oaca
| |
| Metanephros
| |
| Aorta
| |
| | |
| Mesonephros
| |
| | |
| Fia. 227.— 10 mm. pig embryo. Sagittal section. 164.
| |
| | |
| pouches the thyroid gland appears. From the level of the fourth
| |
| pouch a short laryngeal groove is prolonged into the trachea
| |
| which has given rise to the bronchial buds, three in number. Two
| |
| of these, the primary bronchi, have arisen by the bifurcation of
| |
| ENDODERMAL DERIVATIVES 319
| |
| | |
| the trachea; the third or apical bud, which will give rise to the
| |
| eparterial bronchus, develops anterior to the right primary
| |
| bronchus. The esophagus is relatively long and narrow and,
| |
| just posterior to the level of the lung buds, passes into the stomach
| |
| which is dilated and shows a slight dorsal curvature. Posterior
| |
| to the stomach the duodenal glands, liver, and pancreas are well
| |
| developed. The liver, now a large glandular mass traversed by
| |
| | |
|
| |
|
| |
|
| |
|
| |
| | |
| W- Nerve XI
| |
| | |
| Nerve X
| |
| and jugular F Ganglion IX
| |
| ganglion (superior)
| |
| | |
| Otic vesicle
| |
| | |
| ~Ganglion VOI
| |
| Myelencephalon ¥} (acoustic)
| |
| Ganglion Y~}(semilunar) P
| |
| Pog IT a1 Basilar
| |
| Fee eof artery
| |
| Nerve III
| |
| Internal
| |
| | |
| carotid artery
| |
| | |
| Mesencephalon
| |
| | |
|
| |
| | |
| Fig. 228. — 10 mm. pig embryo. Transverse section through otic vesicles. 163}.
| |
| | |
| the capillaries of the hepato-portal veins, retains its original connection with the duodenum as the common bile duct from the
| |
| distal end of which the gall bladder is forming. Both dorsal
| |
| and ventral primordia of the pancreas are present, the duct of
| |
| the latter arising from the common bile duct. The long and
| |
| slender intestine extends into the umbilical cord as the intestinal
| |
| loop, to which the yolk stalk is still attached. Just posterior to
| |
| this, a slight enlargement may sometimes be observed which indicates the boundary between the large and small intestine. The
| |
| hind-gut is dividing into a dorsal rectum and ventral urogenital
| |
| 320 THE ANATOMY OF THE 10 MM. PIG EMBRYO
| |
| | |
| sinus, prolonged into the allantoic stalk. The sinus and rectum
| |
| unite in a common cloaca which has not yet established connection
| |
| with the proctodeum. Immediately posterior to the cloacal plate,
| |
| a small blind pocket represents the postcloacal gut.
| |
| | |
| Spinal cord.
| |
| | |
|
| |
| | |
| it Dorsal root
| |
| co Spinal ganglion
| |
| | |
| re Ventral root
| |
| Dorsal ramus
| |
| | |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
| | |
|
| |
|
| |
|
| |
|
| |
| | |
| Notochord
| |
| | |
| Anterior
| |
| cardinal vein Ganglion X
| |
| Aortic ( nodosum )
| |
| arch OT Radix aortae
| |
| Visceral
| |
| | |
| Visceral arch
| |
| | |
| a Hypophysis
| |
| Anterior
| |
| cardinal vein
| |
| | |
| Sensory layer
| |
| Pigment layer
| |
| | |
| Diencephalon
| |
| | |
| Fig. 229.10 mm. pig embryo. Transverse section through optic cup. 164.
| |
| | |
| Mesodermal derivatives. — The notochord extends from the
| |
| vicinity of the floor of the mesencephalon into the tail, following
| |
| the flexures of the body.
| |
| | |
| The somites have long since become differentiated into the
| |
| myotome, dermatome, and sclerotome. In the tail region, the
| |
| sclerotomes are separated into the cranial and caudal arcualia
| |
| from which the vertebrae will originate.
| |
| | |
| In the pig of 10 mm., the pronephric stage has been passed;
| |
| the mesonephros is at the height of its development, forming a
| |
| great “Wolffian” body with a complicated network of interwoven
| |
| tubules; while the mesonephric duct (originally the pronephric
| |
| duct) may be recognized along the ventral margin. Emerging
| |
| MESODERMAL DERIVATIVES 321
| |
| | |
| from the mesonephros, each duct enters the urogenital sinus at the
| |
| same level as the allantoic stalk. From each duct a narrow stalk
| |
| runs dorsally and forward as the metanephric duct, or ureter,
| |
| which at its distal end is enlarged to form the pelvis of the metanephros. Around the pelvis the posterior portion of the nephrotomal band will produce the secretory tubules of the definitive
| |
| kidney at a later stage. On the median ventral margin of each
| |
| | |
|
| |
|
| |
|
| |
|
| |
|
| |
| | |
| Notochord
| |
| | |
| Dorsal aorta
| |
| | |
| Oesophagus Anterior
| |
| | |
| ardinal vein
| |
| | |
| Left
| |
| atrium
| |
| | |
| Telencephalon
| |
| | |
| Fig. 230. — 10 mm. pig embryo. Transverse section through nasal (olfactory) pit.
| |
| X 163.
| |
| | |
| mesonephros are slight swellings which will later become the
| |
| genital ridges, primordia of the gonads. The coelom is partially
| |
| divided into the pericardial and abdominal cavities by the septum
| |
| transversum. The mesenteries of the principal viscera are in
| |
| evidence. The liver is still suspended in the ventral mesentery.
| |
| A dorsal mesocardium is present.
| |
| | |
| The heart of the 10 mm. pig has the four main chambers established, although not yet completely separated into right and left
| |
| halves. The sinus venosus now enters the right atrium through
| |
| 322 THE ANATOMY OF THE 10 MM. PIG EMBRYO
| |
| | |
| a slit guarded by the valves of the sinus. The right and left
| |
| atria are partially separated by the interatrial septum in which
| |
| can be seen an opening, the foramen ovale. The atrio-ventricular canal leading to the ventricle is partially separated into right
| |
| and left halves by the endocardial cushion. The ventricle is partially divided by the interventricular septum. From the ventral
| |
| aorta three aortic arches curve around the pharynx to unite with
| |
| the dorsal aorta. These are the third, fourth, and sixth aortic
| |
| arches; the first and second have degenerated, while the fifth
| |
| | |
| Spinal cord .
| |
| | |
|
| |
|
| |
|
| |
|
| |
| | |
| Ganglion
| |
| Notochord
| |
| | |
| Anterior
| |
| limb bud
| |
| | |
| Common Dorsal aorta
| |
| cardinal vein
| |
| Eparterial Oesophagus
| |
| bronchus Trachea
| |
| Valves of
| |
| sinus Left atrium
| |
| Right
| |
| atrium
| |
| Left
| |
| Right \ ventricle
| |
| ventricle
| |
| | |
| Fia. 231. — 10 mm. pigembryo. Transverse section through sinus venosus. 16}.
| |
| | |
| seldom appears as a separate structure. The pulmonary arteries
| |
| are growing back from the sixth aortic arches.
| |
| | |
| As prolongations of the original paired ventral and dorsal aortae,
| |
| the external and internal carotid arteries, respectively, run forward into the head. The internal carotid arteries are united at
| |
| the level of the isthmus between the mesencephalon and the
| |
| metencephalon with the basilar artery, which serves to unite
| |
| them with the vertebral arteries, arising from the anastomosis of
| |
| intersegmental arteries in the cervical region. At the 10 mm.
| |
| stage the vertebral arteries have lost their intersegmental connections with the aorta except at the posterior end, where the
| |
| MESODERMAL DERIVATIVES 323
| |
| | |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
| | |
| Anterior
| |
| limb bud:
| |
| | |
| Coelom
| |
| | |
| Posterior
| |
| vena cava
| |
| | |
| Right
| |
| | |
| atrium {
| |
| i Left
| |
| Right i
| |
| ventricle ventricle
| |
| | |
| Notochord
| |
| | |
| Dorsal aorta
| |
| | |
| Glomerulus
| |
| | |
| ketee . g ‘
| |
| Posterior Vea me AN Stomach
| |
| vena cava i
| |
| | |
| by \
| |
| v
| |
| : Septum 4 Pericardial
| |
| ransversum — cavity
| |
| a
| |
| | |
| Fig. 233. — 10 mm. pig embryo. Transverse section through stomach. X16}.
| |
| 324 THE ANATOMY OF THE 10 MM. PIG EMBRYO
| |
| | |
| seventh cervical intersegmental artery persists and grows out into
| |
| the pectoral limb bud to form the subclavian artery. Near the
| |
| point of origin of the subclavian, the dorsal aortae are fused and
| |
| run back as a single median aorta into the tail. Dorsally,
| |
| branches are given off from the aorta as intersegmental arteries
| |
| of the trunk. Laterally, many small branches supply the glomeruli of the mesonephros. Ventrally, the dorsal aorta gives
| |
| off the coeliac artery and anterior mesenteric arteries to the gut.
| |
| | |
|
| |
|
| |
| | |
| Ganglion
| |
| Notochord
| |
| | |
| Left umbilical
| |
| vein
| |
| | |
| Fig. 234. — 10 mm. pig embryo. Transverse section through gall bladder. 163.
| |
| | |
| Two large umbilical (allantoic) arteries run from the dorsal aorta
| |
| into the umbilical cord. The aorta continues into the tail as a
| |
| relatively slender vessel, the caudal artery.
| |
| | |
| The vitelline veins are much smaller than in the chick of
| |
| seventy-two hours, for the yolk sac from which they drew their
| |
| blood is nearly degenerated. In the pig at this stage they drain
| |
| the gut area and cross into the liver where they become the portal
| |
| vein. Within the liver they are broken up into capillaries which
| |
| emerge as the hepatic veins to the sinus venosus. Of the somatic
| |
| MESODERMAL DERIVATIVES 325
| |
| | |
| veins, the anterior cardinals are still prominent and are joined
| |
| by an extensive series of head veins. In the cervical region the
| |
| anterior cardinals receive the dorsal intersegmental veins as well
| |
| as the external jugular from the mandible. As the anterior
| |
| cardinals enter the common cardinal veins, they are joined by the
| |
| posterior cardinals, which have already lost part of their drainage
| |
| | |
| Spinal cord oO .
| |
| | |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
| | |
| Notochord
| |
| Posterior
| |
| cardinal
| |
| vein
| |
| Posterior
| |
| vena
| |
| | |
| Left
| |
| vitelline
| |
| (portal)
| |
| vein
| |
| Left
| |
| mbilical
| |
| vein
| |
| Left
| |
| vitelline
| |
| artery
| |
| | |
| duct
| |
| | |
| umbilical
| |
| artery
| |
| | |
| Fused
| |
| umbilical
| |
| veins
| |
| | |
| Fig. 235. 10 mm. pig embryo. Transverse section through umbilical stalk in
| |
| region of intestinal loop. X16}.
| |
| | |
| area to the subcardinal veins passing through the ventral portions
| |
| of the mesonephroi. Numerous small venous channels serve to
| |
| connect the subcardinals and postcardinals during this period.
| |
| The posterior caval vein has already made its appearance as a
| |
| direct connection from the subcardinals to the liver. The umbilical (allantoic) veins proceeding from the allantois toward the
| |
| heart are fused together in the umbilical cord. In the body they
| |
| 326 THE ANATOMY OF THE 10 MM. PIG EMBRYO
| |
| | |
| pass through the liver, within which they are, like the vitelline
| |
| veins, broken up into capillaries. The left umbilical maintains a
| |
| broad channel through the liver. This vessel, now known as the
| |
| ductus venosus, connects the umbilical with the posterior caval
| |
| vein.
| |
| | |
|
| |
|
| |
|
| |
| | |
| Posterior Mesonephric
| |
| | |
| limb bud duct
| |
| Umbilical
| |
| artery Metanephric
| |
| duct
| |
| | |
| Caudal
| |
| artery
| |
| | |
| Notochord
| |
| Spinal
| |
| cord
| |
| | |
| Fig. 236. — 10 mm. pig embryo. Transverse section through metanephric duct and
| |
| posterior limb buds. X16}.
| |
| | |
| Ectodermal derivatives. — The epidermal derivatives of the
| |
| ectoderm have already been enumerated in the description of
| |
| external form. There remain for consideration the nervous system and sense organs. ‘The five definitive vesicles of the brain
| |
| are well marked. From the telencephalon arise the two lateral
| |
| cerebral vesicles. This division of the brain is separated from
| |
| the diencephalon by two points of reference, the optic recess in
| |
| the floor, and the velum transversum in the roof. From the
| |
| diencephalon spring the optic stalks, leading to the optic cups,
| |
| and the infundibulum, now in contact with the hypophysis as
| |
| mentioned above. The posterior boundary of the diencephalon
| |
| is indicated by the tuberculum posterius arising from the brain
| |
| floor. The epiphysis seldom appears at this stage. The mesencephalon, with the third cranial nerve arising from its floor, is
| |
| ECTODERMAL DERIVATIVES 327
| |
| | |
| demarcated at its posterior end by the deep constriction of the
| |
| isthmus. The metencephalon is distinguished from the myelencephalon by its thicker roof. From the isthmus the fourth
| |
| cranial nerve runs forward laterally over the sides of the brain to
| |
| the mass of mesoderm surrounding the eyeball, from which the
| |
| | |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
|
| |
| | |
| Basilar
| |
| artery
| |
| | |
| Anterior
| |
| cardinal
| |
| | |
| vein Internal
| |
| carotid artery
| |
| Thymus Olfactory pit |
| |
| Visceral 8rd Aortic arch
| |
| | |
| 4th Aortic arch
| |
| 6th Aortic arch.
| |
| Sinus
| |
| | |
| venosus
| |
| | |
| Right
| |
| atrium
| |
| | |
| pouch
| |
| | |
| Ductus
| |
| venosus
| |
| | |
|
| |
| | |
| [J Subcardinal
| |
| #/J anastomosis,
| |
| | |
| Notochord
| |
| | |
| Fig. 237. — 10 mm. pig embryo. Frontal section through aortic arches and ductus
| |
| venosus. X16}.
| |
| | |
| eyeball muscles will be formed. Conspicuous at the anterior
| |
| ventro-lateral margin of the metencephalon is the large semilunar
| |
| ganglion of the fifth cranial nerve. From the floor of the myelencephalon, the sixth cranial nerve emerges to run forward toward
| |
| the eye. Immediately following this, the geniculate ganglion of
| |
| the seventh and the acoustic ganglion of the eighth are in close
| |
| 328 THE ANATOMY OF THE 10 MM. PIG EMBRYO
| |
| | |
| connection. The ninth cranial nerve has two ganglia, the dorsal
| |
| superior ganglion and ventral petrosal, while the tenth similarly
| |
| possesses a dorsal jugular and ventral nodose ganglion. The
| |
| eleventh cranial nerve possesses at this stage a small ganglion
| |
| (of Froriep) which disappears in the adult. The last of the cranial
| |
| nerves, the twelfth, arises from the floor of the myelencephalon
| |
| by a number of small roots and without a ganglion. In the region
| |
| of the spinal cord the segmental nerves arise from the cord by two
| |
| roots, of which the dorsal is associated with a spinal ganglion.
| |
| The trunk is very short and soon divides into three main branches.
| |
| The dorsal and ventral rami run to these respective regions of
| |
| the body wall, while the third, or communicating ramus, unites
| |
| the spinal nerve with a ganglion of the sympathetic chain. The
| |
| sympathetic ganglia may be recognized as small masses of cells
| |
| dorsal to the aorta.
| |
| | |
| The nose is represented by the olfactory pits. The eye is in
| |
| the optic cup stage with a well-marked choroid fissure and
| |
| groove, while the lens is completely separated from the outer ectoderm and is in the vesicle stage. Of the various regions of the
| |
| ear, all the primordia are now established. The otic vesicle with
| |
| its endolymphatic duct, representing the inner ear, is in close
| |
| juxtaposition to the first visceral pouch (hyomandibular) which
| |
| will give rise to the auditory tube and chamber of the middle ear;
| |
| the external auditory meatus, or outer ear, will arise from the
| |
| first or hyomandibular groove.
| |
| | |
| REFERENCES
| |
| | |
| Arey, L. B. 1934. Developmental Anatomy, 3rd Ed., Chap. 19.
| |
| | |
| “Boyden, E. A. 1933. A Laboratory Atlas of the Pig Embryo.
| |
| | |
| Keibel, I’. 1897. Normaltafeln, I, des Schweines (Sus scrofa domesticus).
| |
| | |
| Lewis, F. T. 1902. The gross anatomy of a 12 mm. pig, Am. Jour. Anat., Vol. 2,
| |
| pp. 211-226.
| |
| | |
| ‘Minot, C.S. 1911. A Laboratory Textbook of Embryology, 2nd Ed.
| |
| | |
| Patten, B. M. 1931. The Embryology of the Pig, 2nd Ed.
| |
| | |
| Wallin, E. 1917. A teaching model of a 10 mm. pig embryo, Anat. Rec., Vol. 5,
| |
| pp. 17-45.
| |
| PART V
| |
| MICROSCOPICAL TECHNIQUE
| |
| | |
| CHAPTER XIV
| |
| PREPARATION OF EMBRYOLOGICAL MATERIAL
| |
| | |
| A method much employed in the study of comparative embryology is that of cutting a preserved egg or embryo into a series
| |
| of extremely thin slices, and arranging these in order upon a
| |
| glass slide, so that they may be examined under the microscope.
| |
| The older embryologists, however, were limited to the study of
| |
| entire embryos and of minute dissections. These methods are
| |
| still of great value in supplementing the study of serial sections,
| |
| for it is a difficult mental exercise to translate sections into terms
| |
| of the whole embryo. The single section, especially, is meaningless except when interpreted as a part of the complete series. It
| |
| is very helpful, therefore, when facilities permit, for each student
| |
| to prepare for himself a whole mount and a series of sections
| |
| through one of the embryos he is to study.
| |
| | |
| A. COLLECTION AND REARING OF EMBRYOS
| |
| | |
| Although preserved embryos of the more important laboratory
| |
| types may be obtained from the biological supply houses, it is
| |
| often desirable to collect and rear live embryos.
| |
| | |
| THE FROG. — There are some sixty species of tailless Amphibia
| |
| within the continental limits of the United States. Although
| |
| the capture of adults in a pond where eggs are found is strong
| |
| circumstantial evidence as to the species of the eggs, even this
| |
| evidence is often lacking, so that the ability to identify the
| |
| eggs or larvae from their own characteristics is highly desirable.
| |
| A key to the eggs and larvae of some of the common Eastern frogs
| |
| and toads is found in Wright’s ‘‘ Life History of the Anura of
| |
| Ithaca, N. Y.” For the Pacific slope fauna, see Storer, “A
| |
| Synopsis of the Amphibia of California.”’ The eggs of the salamander, Ambystoma, are laid at the same time and in the same
| |
| localities as those of the early frogs, but may be distinguished
| |
| from them by the greater proportion of jelly to the eggs in the
| |
| | |
| mass of spawn.
| |
| 331
| |
| 332 PREPARATION OF EMBRYOLOGICAL MATERIAL
| |
| | |
| Experiments dealing with the effect of pituitary hormones have
| |
| led to the discovery that one of these hormones will induce
| |
| ovulation in the female frog, and the drive to amplexus in the
| |
| male, out of the breeding season. Rugh! (1934) has described in
| |
| detail a technique for inducing ovulation and bringing about
| |
| artificial fertilization which has been since used in several laboratories, including the author’s, with complete success.
| |
| | |
| The rate of development of the frog’s egg depends upon the
| |
| temperature of the water. In the laboratory, the eggs will hatch
| |
| in about one week after laying, at the ordinary room temperature. The egg masses should be kept in clean glass containers
| |
| with at least ten times as much water. The water should not be
| |
| changed until after hatching, when the larvae should be transferred to fresh water with aquatic plants. After the assumption
| |
| of the tadpole form, they should be fed small pieces of finely
| |
| ground meat. Metamorphosis may be hastened by feeding fresh
| |
| or desiccated thyroid tissue.
| |
| | |
| Artificial fertilization is the best method of obtaining the
| |
| earliest stages of development. The testes and vasa deferentia
| |
| of the male are teased out in a watch glass of water. The eggs
| |
| from the distal portions of the oviducts are placed in this water for
| |
| five minutes and then removed to glass containers with not more
| |
| than four inches of water.
| |
| | |
| THE CHICK. — In collecting hens’ eggs for incubation, it is a
| |
| truism that they must be fresh and fertile. The best results are
| |
| obtained from trap-nested eggs in the spring semester. The egg
| |
| is normally laid in the gastrula stage (Chapter II), but in those
| |
| cases where the egg does not reach the distal end of the oviduct
| |
| by 4 P.M., it is retained till the following morning and undergoes
| |
| further development. After laying, the egg cools and development ceases until incubation is commenced. The fertilized egg
| |
| is viable for five weeks at a temperature of 8°-10° C. The time
| |
| of hatching, as in the frog’s egg, is dependent upon the temperature. The minimum temperature at which development will
| |
| take place is about 25° C.; the optimum is 37° C., at which
| |
| temperature the egg will hatch in twenty-one days; the maximum
| |
| temperature is about 41° C. In incubating eggs, care must be
| |
| | |
| 1R. Rugh. Induced Ovulation and Artificial Fertilization in the Frog, Biol.
| |
| Bull. 66, 22-29.
| |
| PRESERVATION OF MATERIAL 333
| |
| | |
| taken to keep the air in the incubator moist and to rotate the
| |
| eggs once a day.
| |
| | |
| Instructive demonstrations may be made by opening the shell
| |
| and shell membranes under aseptic conditions and removing a
| |
| bit of the albumen. <A window of celloidin placed over the opening and carefully sealed will permit of observations on the development of the embryo for several days. An alternative method
| |
| is that of opening the egg and placing the contents in a sterilized
| |
| small stender dish. A glass ring is placed on the yolk to keep it
| |
| beneath the surface of the albumen, and the dish is covered and
| |
| placed in the incubator. If this operation is carried on under
| |
| aseptic conditions, development will continue for two or three days.
| |
| | |
| THE PIG. — The early stages of development in any mammal
| |
| are valuable. The larger embryos are visible as protuberances
| |
| on the inner side of the uterine tubes. The tube should be slit
| |
| open and the embryos exposed by cutting open the embryonic
| |
| membranes which surround them. Smaller stages are obtained
| |
| by washing out the contents of the tube with normal salt solution
| |
| or preserving it entire.
| |
| | |
| Pig embryos may be obtained in quantities from any good-sized
| |
| packing house. As many as eighteen may be found in a single
| |
| female, but the average number is eight. The period of gestation
| |
| in the pig is 121 days. Pig embryos of 10 mm. body length are
| |
| the most useful in the elementary course. Later stages are of
| |
| value in the detailed study of organogeny.
| |
| | |
| B. PRESERVATION OF MATERIAL
| |
| | |
| The preliminary preparation of material for microscopical work
| |
| involves three distinct operations: killing, fixing, and preservation. In practice, two or three of these operations are performed
| |
| by a single reagent known as a “ fixing fluid.” Such a reagent
| |
| should kill the embryo so rapidly that it will undergo the minimum
| |
| of post-mortem changes; it should preserve the structures of the
| |
| embryo with as life-like an appearance as possible; and it should
| |
| harden the soft parts so that they may undergo the later processes
| |
| of technique without loss of form or structure. Some fixing fluids,
| |
| such as alcohol or formalin, may be used indefinitely as preservatives, but the majority are used for a particular optimum period,
| |
| and then washed out and replaced by alcohol.
| |
| 334 PREPARATION OF EMBRYOLOGICAL MATERIAL
| |
| | |
| THE FROG. — The frog’s egg, before hatching, is best fixed by
| |
| Smith’s fluid.
| |
| | |
| Potassium bichromate.......... cece eee eee eee 0.5 gram
| |
| Glacial acetic aCid. . 6... ce cece cece cece eee eee 2.5 ce.
| |
| Formalin. 2.0... ccc ccc cece cee cece cence eeaee 10.0 ce.
| |
| | |
| Distilled water... 2... . cece eee eee eee eee 75.0 cc.
| |
| | |
| 1. Cut the egg masses into small pieces of about twenty-five
| |
| eggs each, and submerge them in a dish of Smith’s fluid for
| |
| twenty-four hours. A quantity equal to ten times the volume
| |
| of the eggs should be used.
| |
| | |
| 2. Rinse the eggs in water and wash with a 5 per cent aqueous
| |
| solution of formalin until no more free color comes out. The
| |
| eggs may be kept indefinitely in this fluid. If it is desired to
| |
| remove the egg membranes, proceed as follows:
| |
| | |
| 3. Wash in water for twenty-four hours, changing the water
| |
| several times.
| |
| | |
| 4. Place the eggs in eau de Javelle, diluted with three time its
| |
| volume of water, and shake gently from time to time during a
| |
| period of 15 to 30 minutes until the membranes are almost
| |
| dissolved and will shake off.
| |
| | |
| 5. Rinse in water and run through 50 per cent and 70 per cent
| |
| alcohol, an hour to a day each, and preserve in 80 per cent alcohol.
| |
| | |
| After hatching, larvae are best fixed in Bouin’s fluid.
| |
| | |
| Picric acid, saturated aqueous solution................ 75 cc.
| |
| Formalin. 2.0... cece cece eee een eeeeees 25 cc.
| |
| Glacial acetic acid... .. ec cee cece eeeeees 5 ce.
| |
| | |
| 1. Larvae are left in this fluid from one to eighteen hours,
| |
| according to size.
| |
| | |
| 2. After rinsing in 50 per cent alcohol, wash in 70 per cent
| |
| alcohol, to which has been added a few drops of lithium carbonate, saturated aqueous solution, until the yellow color is
| |
| extracted, and preserve in 80 per cent alcohol.
| |
| | |
| THE CHICK. — The chick embryo must be removed from the
| |
| shell, albumen, and yolk before fixation. As the early stages
| |
| are more difficult to handle, it is advisable to practice this operation on embryos of seventy-two hours’ incubation and then work
| |
| backward toward the stages of the first day.
| |
| THE CHICK 335
| |
| | |
| 1. Place the egg in a dish 3 inches high and 6 inches in diameter, two-thirds full of normal saline solution, warmed to 40° C.
| |
| | |
| 2. Crack the shell at the broad end with the flat of the scalpel,
| |
| and pick away the pieces of shell until an opening slightly larger
| |
| than a half dollar has been made. Remove the outer and inner
| |
| shell membranes. Invert egg beneath the surface of the salt
| |
| solution and allow the contents to flow out. The blastoderm,
| |
| containing the embryo, will rotate until it is uppermost. With
| |
| fine-pointed scissors, cut rapidly a circle of blastoderm, about the
| |
| size of a quarter, with the embryo at the center. With blunted
| |
| forceps, pull the blastoderm and adherent vitelline membrane
| |
| away from the yolk and albumen, waving it gently beneath the
| |
| surface of the salt solution to remove all yolk.
| |
| | |
| 3. Submerge a syracuse watch glass in the salt solution and
| |
| float the embryo into this. Remove the watch glass carefully
| |
| from the large dish and examine the embryo with a dissecting lens.
| |
| If the vitelline membrane has not yet separated from the blastoderm, it should be removed at this time with fine-pointed forceps
| |
| and needles. Make sure that the embryo lies dorsal side up, as
| |
| it did when the egg was opened.
| |
| | |
| 4, Slide a cover glass under the embryo, and remove all salt
| |
| solution: with a pipette, taking care that the embryo lies in the
| |
| center of the cover glass. Lift the cover glass by one corner so
| |
| that the overhanging edges of the blastoderm fold under, and
| |
| place it in a dry watch glass on a piece of thin absorbent tissue
| |
| paper and add fixing fluid at once. While the embryo is becoming
| |
| attached to the cover glass, remove the yolk, albumen, and pieces
| |
| of shell from the dish of salt solution to a slop jar, reheat the salt
| |
| solution to 40° C., and prepare another embryo. Three embryos
| |
| of each stage are to be prepared.
| |
| | |
| 5. After five minutes, drop the cover glass, embryo side up,
| |
| into a small stender dish of Bouin’s fluid and leave from two to
| |
| four hours.
| |
| | |
| 6. Rinse in 50 per cent alcohol, wash for two days in 70 per
| |
| cent alcohol to which lithium carbonate has been added or until
| |
| the yellow color is extracted from the embryo, and preserve in
| |
| 80 per cent alcohol.
| |
| 336° PREPARATION OF EMBRYOLOGICAL MATERIAL
| |
| | |
| THE PIG. — Embryos of 6 mm. body length and over are easily
| |
| located in the uterine wall. Slit open the uterus and remove the
| |
| embryo with fine-pointed forceps and a horn spoon, taking pains
| |
| not to rupture the membranes. Place at once in Bouin’s fluid.
| |
| Embryos of 10 mm. body length should be fixed for four hours.
| |
| Rinsing and preserving are done as for the frog or chick. Larger
| |
| embryos should have the body cavity slit open to admit the fixing fluid. Fetal pigs of 6 inches or more should be injected
| |
| through the umbilical artery with formalin (20 per cent aqueous
| |
| solution). This solution is also injected into the body cavity and
| |
| cranium, after which the fetus is submerged in the same medium
| |
| for a week and preserved in 6 per cent formalin.
| |
| | |
| “°C. WHOLE MOUNTS
| |
| | |
| It is very helpful to have some embryos mounted entire for
| |
| comparison with the serial sections. In making these whole
| |
| mounts, the embryos are stained, cléared, and mounted, i.e.,
| |
| transferred to a final medium for preservation and examination
| |
| on the slide beneath a cover glass.
| |
| | |
| THE FROG. — Frog eggs and embryos may be mounted as opaque
| |
| objects with the natural pigmentation, or they may be cleared
| |
| and stained as transparent mounts.
| |
| | |
| Opaque mounts. —
| |
| | |
| 1. Prepare a saturated aqueous solution of thymol. Filter
| |
| the solution, and add gelatin until saturated. Remove the
| |
| supernatant liquid.
| |
| | |
| 2. Liquefy the gelatin by immersing a small quantity, in a
| |
| test tube, in a dish of hot water. Fill a hollow-ground depression
| |
| slide with gelatin and allow to cool.
| |
| | |
| 3. With a hot needle, melt a small hole in the gelatin, sufficiently large to hold the embryo. Place the embryo in the
| |
| desired position and hold it in place until the gelatin has cooled.
| |
| | |
| 4, Add a drop of gelatin just warm enough to be liquid and
| |
| cover with a cover glass which has been slightly warmed. When
| |
| the gelatin has cooled, any surplus may be removed from the
| |
| edges of the cover glass with a toothpick wrapped in moist cotton.
| |
| In order to prevent the later formation of bubbles, the edges
| |
| of the cover glass should be painted with gold size or Valspar.
| |
| | |
| Free-hand sections and dissections are admirably mounted by
| |
| THE CHICK 337
| |
| | |
| this method, but great care must be exercised to prevent the
| |
| formation of air bubbles through cracks in the gold size.
| |
| | |
| Transparent stained mounts. —
| |
| | |
| 1. Bleach the embryo, until white, in hydrogen peroxide.
| |
| About one week is required for this purpose. Embryos that have
| |
| been preserved in 80 per cent alcohol should first be passed
| |
| through 70 and 50 per cent alcohol to water, an hour or more in
| |
| each fluid. Embryos in formalin must be rinsed in water for
| |
| one hour.
| |
| | |
| 2. Stain in dilute borax carmine four days or more.
| |
| | |
| Borax, 4 per cent aqueous solution. .................. 100 ce.
| |
| Carmine. . 0.0.2... ccc cece eee eeneee 1 gr.
| |
| Boil until dissolved and add alcohol, 70 per cent....... 100 ce.
| |
| | |
| To dilute, take 5 cc. of the borax carmine and 95 ce. of 35 per cent
| |
| alcohol and add a crystal of thymol.
| |
| | |
| 3. If overstained, remove the surplus color with hydrochloric
| |
| acid (1 per cent solution in 70 per cent alcohol) after passing
| |
| through water and 50 per cent alcohol, an hour each.
| |
| | |
| 4, Run up through 80, 95, and 100 per cent alcohol, an hour
| |
| each, and place in xylene (xylol) until transparent.
| |
| | |
| 5. Prepare a mounting diagram by drawing an outline of a
| |
| slide on a piece of cardboard and in this laying off an outline of
| |
| the cover glass to be used. Place a clean slide on the diagram,
| |
| and, just’ inside the right and left margins of the cover-glass outline, attach a thin strip of celluloid, 15/1000 of an inch in thickness, by means of a drop of acetone. Greater thicknesses may
| |
| be obtained by attaching other strips as necessary. When these
| |
| supports are dry, place a few drops of Canada balsam, dissolved
| |
| in xylene, between the supports, place the embryo in position,
| |
| and lower a clean cover glass gently. Try to avoid the formation
| |
| of air bubbles. If these appear later they may be removed by a
| |
| needle which has been heated or dipped in xylene. A little fresh
| |
| balsam may be run into the cavity.
| |
| | |
| THE CHICK. — Total mounts may be stained either with the
| |
| borax carmine or with Conklin’s modification of Delafield’s
| |
| hematoxylin. Delafield’s hematoxylin, which gives a blue color
| |
| to the embryo, is made as follows:
| |
| 338 PREPARATION OF EMBRYOLOGICAL MATERIAL
| |
| | |
| Hematoxylin (16 per cent solution in 100 per cent al
| |
| COMO]... cc ee een eee nes 25 ce.
| |
| Ammonia alum (saturated aqueous solution).......... 400 ce.
| |
| Hydrogen peroxide, neutralized. . 0... 6. cee eee ee eee 25 cc.
| |
| Glycerin. 20... cc cece eet ee eee eens 100 ce.
| |
| Alcohol methyl... 0.0.0... ccc cence nens 100 ce.
| |
| | |
| Conklin’s modification consists of diluting the stain with four
| |
| times the volume of distilled water and adding to each 100 ce. of
| |
| the dilute stain 1 ec. of picrosulphuric acid, prepared by adding
| |
| 2 cc. of sulphuric acid to 98 cc. of picric acid (saturated aqueous
| |
| solution).
| |
| | |
| 1. Run the embryo from the 80 per cent alcohol down to
| |
| water through changes of 70 and 50 per cent alcohol, an hour
| |
| each.
| |
| | |
| 2. Stain in borax carmine, undiluted, over night, or in hematoxylin from one to three hours. Either stain may be diluted
| |
| still further and the staining period prolonged. In the author’s
| |
| laboratory the schedule demands a four-day staining period and
| |
| the borax carmine is diluted 5 x, the hematoxylin 20 x.
| |
| | |
| 3. Destain, if necessary, in acid alcohol until the desired color
| |
| is obtained. Embryos stained with hematoxylin will turn red
| |
| in the acid alcohol, and the blue color must be restored by washing them in running water or, after washing in neutral 70 per
| |
| cent alcohol, placing them in alkaline alcohol (1 per cent ammonia
| |
| in 80 per cent alcohol).
| |
| | |
| 4. Run up the alcohols, 80, 95, and 100 per cent, half an hour
| |
| each. Pour off half the 100 per cent alcohol and add an equal
| |
| amount of xylene. When the diffusion currents disappear, transfer to pure xylene and leave until the embryo is transparent. In
| |
| rainy weather, or when 100 per cent alcohol cannot be obtained,
| |
| phenol-xylene (phenol crystals, 25 gr. and xylene 75 cc.) may be
| |
| substituted.
| |
| | |
| 5. Remove the embryo from the cover glass (if it has not already detached itself) and trim the surrounding blastoderm to
| |
| the form of an oblong or circle. Arrange a clean slide on the
| |
| mounting diagram, as described for the frog, attach celluloid
| |
| support, and mount the embryo in Canada balsam with the same
| |
| side uppermost as when the egg was opened. Put the slide away
| |
| where it may lie flat and free from dust until the balsam has
| |
| hardened. This will take at least a week, after which the slide
| |
| EMBEDDING IN PARAFFIN 339
| |
| | |
| may be cautiously cleaned and studied. The process may be
| |
| hastened by drying the slide in the paraffin oven.
| |
| | |
| THE PIG. — Embryos up to 10 mm. body length may be prepared as whole mounts by staining in dilute borax carmine, destaining until only a trace of color persists, and mounting in
| |
| Canada balsam. The time spent in each alcohol should be at
| |
| least an hour for the larger embryos.
| |
| | |
| D. SERIAL SECTIONS
| |
| | |
| In the preparation of serial sections of an embryo, the fixed
| |
| material is (1) embedded in a suitable matrix and (2) sliced into
| |
| extremely thin sections, which are (3) mounted in serial order
| |
| upon slides. The embryo may be stained before or after
| |
| sectioning.
| |
| | |
| Embedding. — There are two principal methods of embedding,
| |
| in paraffin or in celloidin. For especially delicate objects, the
| |
| best results are obtained by a combination of these methods,
| |
| the embryo being first impregnated with celloidin in order to
| |
| avoid the shrinkage (about 10 per cent) caused by paraffin embedding, and the block of celloidin then immersed in paraffin so
| |
| that ribbons of serial sections may be cut.
| |
| | |
| Embedding in paraffin. — In preparing the first few embryos
| |
| for sectioning, it is advisable to stain, dehydrate, dealcoholize,
| |
| and clear as if for a total mount. Later, the staining may be
| |
| omitted until after the sections are affixed to the slide.
| |
| | |
| 1. After clearing in xylene, which should be done in a warm
| |
| place, for example, the low-temperature oven at about 40° C.,
| |
| pour off half the xylene and add an equal amount of paraffin chips.
| |
| In the author’s laboratory a paraffin of about 55° melting point,
| |
| obtained by mixing commercial paraffin with parawax, is used.
| |
| The parawax, unfortunately, varies in melting point, so that the
| |
| formula is empirical. The embryo may be left in this xylene
| |
| paraffin for two days.
| |
| | |
| 2. If the mixture has hardened it should again be melted in
| |
| the low-temperature oven. Fill a clean stender dish with melted
| |
| paraffin, transfer the embryo to this, and place in the high-temperature oven at about 56° C. (or one degree above the melting
| |
| point of the paraffin used) for not more than two hours. The
| |
| xylene paraffin should be thrown in the slop jar. Take care not
| |
| 340 PREPARATION OF EMBRYOLOGICAL MATERIAL
| |
| | |
| to get any xylene in the high-temperature oven or paraffin used
| |
| for the final embedding.
| |
| | |
| 3. Smear the interior of a small watch glass with a 10 per cent
| |
| aqueous solution of glycerin (or vaseline), and fill with fresh
| |
| melted paraffin. Transfer the embryo to this, making any necessary adjustments in position with a heated needle. Place the
| |
| embryo dorsal side up, and note the position of the head. Cool
| |
| the surface of the paraffin by blowing on it gently until it is congealed. Then plunge it immediately into a dish of cold water or
| |
| waste alcohol and leave it there for five minutes. Mark the block
| |
| for identification. Objects may be left in paraffin indefinitely.
| |
| | |
| 4. On removing the block of paraffin from its container,
| |
| examine for the following flaws:
| |
| | |
| a. Air bubbles, if they are not near the embryo, may be removed with a hot needle. Otherwise it is better to trim the
| |
| block close to the embryo, put it into melted paraffin, and
| |
| re-embed.
| |
| | |
| b. Milky streaks are due to the presence of xylene. These
| |
| will crumble during sectioning, so that it is best to re-embed if
| |
| they occur near the embryo.
| |
| | |
| c. If the paraffin has “ fallen ” in the center, it is because the
| |
| surface was cooled too long before the block was immersed in the
| |
| water. If any part of the embryo is exposed, it must be reembedded.
| |
| | |
| Sectioning after paraffin embedding. — Before sectioning your
| |
| first embryo, be sure you understand the mechanism of the
| |
| microtome (there are many varieties, of which the rotary type is
| |
| best adapted to beginning students), and have practised the
| |
| technique on a block of paraffin. There are three standard planes
| |
| of sectioning corresponding to the axes of the body (Fig. 238).
| |
| Transverse sections are obtained by cutting the cephalic end of
| |
| the body first, with the knife entering the left side. Sagittal
| |
| sections are made by cutting the right side first, with the knife
| |
| entering the ventral surface. Frontal sections are made by
| |
| commencing at the ventral surface, the knife entering the left
| |
| side. It is best to begin with transverse sections.
| |
| | |
| 1. Attach the paraffin block to the object-carrier of the microtome in the proper manner to obtain the type of section desired.
| |
| This is done by heating the surface of the carrier until it will just
| |
| SECTIONING AFTER PARAFFIN EMBEDDING 341
| |
| | |
| melt paraffin, pressing the block against it in the desired orientation, and lowering into a dish of cold water. A little melted
| |
| paraffin may be poured around the base of the block and this
| |
| again cooled to secure additional support.
| |
| | |
| 2. Place the object-carrier in the microtome and, after orienting the block with respect to the knife, trim it so that the end
| |
| of the block is a perfect rectangle with one of the longer sides
| |
| parallel to the knife edge. If one of the angles is cut off slightly
| |
| there will be a series of indentations in the ribbon which will
| |
| assist in orienting the sections on the slide.
| |
| | |
| 3. If microtome knives are not available, place a new safetyrazor blade (Autostrop type) in the holder provided, allowing the
| |
| | |
|
| |
| | |
|
| |
| | |
| Transverse Sagittal
| |
| | |
| Fig. 238. — Diagram to show method of orienting embryo with reference to microtome knife according to type of section desired.
| |
| | |
| edge to project between a sixteenth and an eighth of an inch.
| |
| Screw the holder in the knife-carrier so that the edge of the blade
| |
| is tilted inward about 10° from the perpendicular.
| |
| | |
| 4. Set the regulator for 20 microns (thousandths of a
| |
| millimeter). .
| |
| | |
| 5. Run the feed screw as far back as it runs freely; do not
| |
| force it.
| |
| | |
| 6. Advance the knife-carrier until the edge of the blade just
| |
| clears the block.
| |
| | |
| 7. Release safety catch and turn the wheel steadily until the
| |
| knife begins to cut the block. Cut slowly, making necessary
| |
| adjustments to the block and knife until you are cutting a perfectly straight ribbon without wrinkles or splits. The principal
| |
| causes of trouble and their remedies are as follows:
| |
| 342 PREPARATION OF EMBRYOLOGICAL MATERIAL
| |
| | |
| a. The ribbon curls to right or left. This happens because (1)
| |
| the block is thicker on the side away from which the ribbon
| |
| curls, or (2) the knife is duller on the side toward which the ribbon
| |
| curls. Remedy: (1) trim the sides of the block parallel; (2)
| |
| shift the knife to one side.
| |
| | |
| b. The sections curl and the ribbon is not continuous. This
| |
| is due to (1) too much tilt of the knife, (2) too hard a grade of
| |
| paraffin, or (3) too cold a room. Remedy: (1) lessen tilt of
| |
| knife; (2) re-embed in softer paraffin; (3) move microtome to
| |
| warmer place, light an electric light or micro-bunsen burner
| |
| near microtome, or cut thinner sections.
| |
| | |
| c. The ribbon wrinkles badly. This is caused by (1) too little
| |
| tilt to the knife, (2) too soft a grade of paraffin, (3) too warm a
| |
| room, or (4) a dull or dirty knife. Remedy: (1) increase the
| |
| tilt of the knife; (2) re-embed in harder paraffin; (8) move to a
| |
| cooler room, or cool the knife and block by dropping alcohol on
| |
| them and blowing vigorously, or cut thicker sections; (4) clean
| |
| knife edge with cloth moistened in xylene or shift to a new place
| |
| on the knife.
| |
| | |
| d. The ribbon splits lengthwise. This is due to (1) a nick in
| |
| the knife, (2) a bubble in the paraffin, or (3) dirt on the knife
| |
| edge or side of the block. Remedy: (1) shift to new cutting edge;
| |
| (2) paint surface with thin celloidin; (3) clean knife edge and block.
| |
| | |
| e. The sections refuse to ribbon; they fly apart or cling to
| |
| the knife or the block. This is due to the electrification of the
| |
| sections caused by unfavorable atmospheric conditions. Many
| |
| remedies have been suggested; the best is to ground the microtome to a water pipe. Usually it is advisable to wait for more
| |
| favorable conditions.
| |
| | |
| 8. Remove the ribbon in 6 inch lengths with a camel’s hair
| |
| brush and arrange these in order, shiny side down, in a cardboard
| |
| box cover. Avoid air currents of all kinds. The ribbons may be
| |
| put away in a dust-free place if the room is not too warm. It is
| |
| better to affix them to slides as soon as possible.
| |
| | |
| Affixing paraffin sections to the slide. — 1. Prepare a mounting
| |
| diagram by laying off the outline of a slide as before, but enclose
| |
| in this the outline of a long cover glass (25 by 50 mm. approximately) and leave space for a label on the right-hand side.
| |
| | |
| 2. Clean a slide thoroughly by washing with acid alcohol
| |
| EMBEDDING IN CELLOIDIN 343
| |
| | |
| followed by distilled water. Place this over the mounting diagram and brush over the surface above the outline of the cover
| |
| glass with the following dilute solution of egg albumen:
| |
| | |
| Egg albumen, beaten and skimmed.................. 50 ce.
| |
| Glycerin... 0... cece cece eee eee neeeeeeees 50 ce.
| |
| Filter and add Thymol.............. 0. ccc cee ee ee ees a crystal
| |
| Dilute 2 drops of this to distilled water............... 25 ce.
| |
| | |
| 3. Cut the ribbon into lengths about 2 per cent shorter than
| |
| the length of the cover glass. Using the wet brush from which
| |
| most of the albumen solution has been squeezed, pick up these
| |
| lengths and arrange them on the albumenized slide so that the
| |
| sections will follow each other like the words on a printed page.
| |
| The shiny side of the ribbon should be next to the slide. Great
| |
| care should be taken to lower the ribbon slowly so as to prevent
| |
| the formation of air bubbles beneath it.
| |
| | |
| 4. Carefully warm the slides on a warming plate or a piece of
| |
| plate glass, previously heated in the paraffin oven, until the sections are expanded and perfectly smooth. If bubbles appear
| |
| beneath the ribbon, prick them with a hot needle while the ribbon
| |
| is still soft and hot. Drain off the surplus water, carefully realign
| |
| the sections, mark the slides with a glass-marking crayon, and set
| |
| them away in the low-temperature oven to dry, at least two days.
| |
| They may be kept indefinitely in this condition if not exposed
| |
| to dust. .
| |
| | |
| Embedding in celloidin. — This method is preferred by some
| |
| technicians as no heat is used in the process and the shrinkage is
| |
| less than that resulting from the paraffin method. However,
| |
| thin sections are not so easy to obtain and the sections must be
| |
| handled individually.
| |
| | |
| 1. Embryos are dehydrated as for the paraffin method. Leave
| |
| in absolute alcohol one day.
| |
| | |
| 2. Absolute alcohol and ether, equal parts, one day.
| |
| | |
| 3. Thin celloidin, three days to one week.
| |
| | |
| Alcohol, 100 per cent. ... 0... eee eee eee eee eee 100 ce
| |
| | |
| Ether... 0.2... cece ccc eee cence eee eeeeee 100 ce
| |
| | |
| Celloidin. £0... ccc ccc cece eee e eee e eee eaes 5 gr.
| |
| 4, Thick celloidin, two days to two weeks.
| |
| | |
| Alcohol, 100 per cent... 1... 0.0... ccc cece eee 100 ce
| |
| | |
| BO 6) a 100 ce
| |
| 344 PREPARATION OF EMBRYOLOGICAL MATERIAL
| |
| | |
| 5. Remove the embryo to a small watch glass and pour thick
| |
| celloidin over it. Cover lightly, or place under a bell jar until
| |
| the celloidin is hard enough to cut with a scalpel.
| |
| | |
| 6. Dip a block of vulcanized fiber in thick celloidin. Cut
| |
| out a block of celloidin containing the embryo from the watch
| |
| glass and, after moistening the end by which it is to be attached
| |
| in ether alcohol, press it firmly against the prepared fiber block.
| |
| | |
| 7. Pour a little chloroform into a stender dish, add the block
| |
| and embryo, cover tightly, and allow the celloidin to harden in
| |
| the fumes for thirty minutes.
| |
| | |
| 8. Fill the stender dish with chloroform and cover. Leave for
| |
| thirty minutes.
| |
| | |
| 9. Pour off half the chloroform and add an equal amount of
| |
| cedar oil. Leave for one hour.
| |
| | |
| 10. Transfer to pure cedar oil where it may remain indefinitely.
| |
| | |
| Sectioning after celloidin embedding. — Celloidin sections are
| |
| usually cut with some form of sliding microtome. Be sure to
| |
| study the mechanism and cut a piece of hardened celloidin before
| |
| proceeding further.
| |
| | |
| 1. Set the knife with a little more tilt than would be used for
| |
| paraffin, and obliquely to the object so that at least half the
| |
| cutting edge will be drawn through the block.
| |
| | |
| 2. Orient the block upon the object-holder so that the desired
| |
| type of sections may be obtained. The long side of the block
| |
| should be parallel to the edge of the knife.
| |
| | |
| 3. Cut sections 20 » or more in thickness, using a steady
| |
| drawing cut. Mount sections as they are cut.
| |
| | |
| Affixing celloidin sections to the slide. — This is best done as
| |
| the sections are cut.
| |
| | |
| 1. Using the mounting diagram as before, rub on a thin film
| |
| of undiluted albumen solution to cover the areas of the cover
| |
| glass. Rub in well with the ball of the finger.
| |
| | |
| 2. Arrange the sections in order on this area. When this is
| |
| filled, lay a cigarette paper over the sections and press gently
| |
| with another slide. The slides may be kept in a dust-free
| |
| container.
| |
| | |
| Double embedding in celloidin and paraffin. — This process,
| |
| although tedious, combines the best points of the two methods
| |
| already given.
| |
| AFTER STAINING IN BULK 345
| |
| | |
| 1. Embed in celloidin according to the method above, omitting
| |
| step 6.
| |
| | |
| 2. Trim the celloidin block close to the embryo and wash out
| |
| the cedar oil with xylene, three changes in two hours.
| |
| | |
| 3. Embed in paraffin as described above, commencing at step 2.
| |
| | |
| 4. Section according to the method given for paraffin.
| |
| | |
| 5. Affix to the slide according to the method given for paraffin
| |
| sections.
| |
| | |
| Staining serial sections. — When the embryo has been stained
| |
| before sectioning, it is only necessary to remove the paraffin (or
| |
| celloidin), replace with Canada balsam, and cover, if the stain
| |
| proves to be satisfactory. Sometimes, however, it is advisable
| |
| to strengthen or weaken the stain or to add a contrasting dye.
| |
| | |
| After staining in bulk. —
| |
| | |
| 1. Paraffin sections on the slide should be put in a Coplin
| |
| staining jar of xylene and left until the paraffin is dissolved, up
| |
| to fifteen minutes.
| |
| | |
| 2. Transfer to a mixture of xylene and 100 per cent alcohol,
| |
| equal parts, five minutes.
| |
| | |
| 3. Transfer to 100 per cent alcohol, five minutes.
| |
| | |
| 4, Examine slide rapidly under microscope after wiping the
| |
| back of the slide.
| |
| | |
| a. If the stain is satisfactory:
| |
| | |
| 5a. Absolute alcohol and xylene, five minutes.
| |
| 6a. Xylene, ten minutes.
| |
| 7a. Mount in balsam under cover glass.
| |
| | |
| b. If the stain is too intense:
| |
| 5b. Ninety-five and 85 per cent alcohol, one minute each.
| |
| 6b. Acid 70 per cent alcohol, until stain is correct.
| |
| 7b. Sections stained in hematoxylin should have the blue
| |
| color restored in alkaline 85 per cent alcohol.
| |
| | |
| 8b. Eighty-five, 95, and 100 per cent, one minute each.
| |
| 9b. Absolute alcohol and xylene, five minutes.
| |
| | |
| 10b. Xylene, ten minutes.
| |
| | |
| 11b. Mount in balsam.
| |
| | |
| c. If the stain is too light:
| |
| 5c. Ninety-five, 85, 70, and 50 per cent alcohol, one minute
| |
| each,
| |
| 346 PREPARATION OF EMBRYOLOGICAL MATERIAL
| |
| | |
| 6c. Stain until desired effect is secured.
| |
| 7c. Distilled water, five minutes.
| |
| 8c. Fifty, 70, 85, 95, 100 per cent alcohol, one minute each.
| |
| 9c. Absolute alcohol and xylene, five minutes.
| |
| 10c. Xylene, ten minutes.
| |
| llc. Mount in balsam.
| |
| | |
| Celloidin sections on the slide should be exposed to the fumes
| |
| of the aleohol-ether mixture for half a minute, dried for one minute, and placed in a staining jar of 95 per cent aleohol. All other
| |
| operations may be carried on as above except that phenol-xylene
| |
| should be substituted for 100 per cent alcohol.
| |
| | |
| Counterstaining after staining in bulk. — In order to differentiate the parts of the embryo more sharply, it is often desirable
| |
| to add a second stain contrasting with the first. The stains that
| |
| have been employed in the previous exercises are nuclear dyes;
| |
| that is, when extracting by acid alcohol, the color will persist
| |
| in the nucleus after it has been washed out of the cytoplasm.
| |
| The second stains affect the cytoplasm and should contrast in
| |
| color with the nuclear stain employed. After borax carmine, a
| |
| 0.5 per cent solution of anilin (Lyons) blue in 95 per cent alcohol
| |
| is employed; after hematoxylin, a similar solution of cosin should
| |
| be used.
| |
| | |
| 1. Proceed as in the preceding section as far as 60.
| |
| | |
| 2. Destain in acid alcohol until the color persists only in the
| |
| nuclei.
| |
| | |
| 3. Restore the blue color to hematoxylin-stained sections in
| |
| alkaline 80 per cent alcohol.
| |
| | |
| 4, Eighty and 95 per cent alcohol, one minute each.
| |
| | |
| 5. Counterstain lightly, dipping the slide into the solution
| |
| repeatedly until a light color persists in the sections, one-half to
| |
| one minute.
| |
| | |
| 6. Rinse in 95 per cent alcohol, dehydrate with 100 per cent
| |
| alcohol, followed by xylene-absolute, clear in xylene, and mount.
| |
| | |
| Staining with Delafield and eosin on the slide. — Follow
| |
| directions given for sections stained in bulk (where stain is too
| |
| light), as far as step 6c, and follow with directions for counterstaining as given above.
| |
| | |
| Staining with Heidenhain’s hematoxylin. — This is one of the
| |
| most important embryological stains.
| |
| OPPEL’S POLYCHROMATIC STAIN 347
| |
| | |
| 1. Remove the paraffin from the sections and run down the
| |
| alcohols to distilled water.
| |
| | |
| 2. Four per cent aqueous solution of iron alum, one hour to
| |
| over night.
| |
| | |
| 3. Rinse in distilled water and place in 0.5 per cent aqueous
| |
| solution of hematoxylin, same time as in the iron alum.
| |
| | |
| 4. Rinse in distilled water and return to the iron alum until
| |
| sections are a pale gray. Check from time to time by rinsing in
| |
| distilled water and examining under microscope to see that the
| |
| desired structures are still visible.
| |
| | |
| 5. When sufficiently destained, wash in running water for
| |
| twenty minutes, or in distilled water, with frequent changes, for
| |
| two hours.
| |
| | |
| 6. Run up the alcohols, clear, and mount.
| |
| | |
| Fuchsin and picro-indigo-carmine. — This polychromatic stain
| |
| is especially fine for organogeny.
| |
| | |
| 1. Remove the paraffin and run down the alcohols to distilled
| |
| water.
| |
| | |
| 2. Stain in basic fuchsin, saturated aqueous solution, twenty
| |
| minutes.
| |
| | |
| 3. Rinse in distilled water and place in picro-indigo-carmine
| |
| for five minutes.
| |
| | |
| Picric acid, saturated aqueous solution................ 50 ce.
| |
| Indigo-carmine, saturated aqucous solution............ 50 ce.
| |
| | |
| 4. Pass rapidly through 70, 95, and absolute alcohol into
| |
| xylene-alcohol. The green dye is extracted most rapidly by the
| |
| 70 per cent alcohol, the red by the absolute. Only experience
| |
| will teach the right time allowance for each alcohol.
| |
| | |
| 5. Clear in xylene and mount.
| |
| | |
| Oppel’s polychromatic stain. — This gives beautiful effects with
| |
| older embryos and larvae.
| |
| | |
| 1. Fix in Bouin.
| |
| | |
| 2. Stain in bulk with undiluted borax-carmine, one to two
| |
| days. Destain for the same period.
| |
| | |
| Embed, preferably by the double method.
| |
| Cut sections, 15-20 u.
| |
| | |
| Run down the alcohols to water.
| |
| | |
| Stain in picro-indigo-carmine, 14 minutes.
| |
| Stain in picro-fuchsin, one minute.
| |
| | |
| NES OUR
| |
| 348 PREPARATION OF EMBRYOLOGICAL MATERIAL
| |
| | |
| Picric acid, saturated aqueous solution................ 50 ce.
| |
| Acid fuchsin, saturated aqueous solution.............. 50 ce.
| |
| 8. Wash in distilled water, changed repeatedly, five minutes.
| |
| 9. Ninety-five per cent alcohol, two minutes.
| |
| 10. Phenol-xylene, xylene, and mount.
| |
| | |
| E. TECHNICAL RECORDS
| |
| | |
| Not the least important part of technique is the keeping of
| |
| exact records covering every technical operation. For each
| |
| embryo there should be a card, giving the following data:
| |
| Kind of embryo and stage of development.
| |
| Method of fixation, time and date.
| |
| Bulk staining, time and date.
| |
| Method of embedding, time and date.
| |
| Plane and thickness of sections, and date.
| |
| Slide staining, time and date.
| |
| Method of mounting, and date.
| |
| Name of preparator.
| |
| | |
| PN OOP Wh Ee
| |
| | |
| F. OUTLINE OF TECHNICAL PROCEDURE FOR CHICK EMBRYOS
| |
| | |
| 1. Remove embryo from egg in warm normal salt solution.
| |
| | |
| 2. Fix for two hours in Bouin’s fluid.
| |
| | |
| 3. Wash in 70 per cent alcohol (plus lithium carbonate), at
| |
| least one change, for two days.
| |
| | |
| 4. Pass through 50 per cent alcohol and water, one hour each.
| |
| | |
| 5. Stain in dilute borax-carmine or Delafield’s alum-hematoxylin, four days.
| |
| | |
| 6. Destain in acid 70 per cent alcohol until desired effect is
| |
| obtained.
| |
| | |
| 7. Wash in neutral 85 per cent alcohol. (The hematoxylinstained specimen is transferred to alkaline 85 per cent alcohol
| |
| until blue color is restored.) Two days.
| |
| | |
| 8. Dehydrate and clear: 95 per cent, 100 per cent alcohol,
| |
| absolute alcohol-xylene, xylene, twenty minutes each.
| |
| | |
| Mount in Canada balsam
| |
| OR
| |
| | |
| 9. Prepare for embedding by pouring off half the xylene and
| |
| adding an equal amount of paraffin chips. Keep in warm place
| |
| up to four days.
| |
| REFERENCES 349
| |
| | |
| 10. Continue by transferring embryo to melted paraffin and
| |
| place in paraffin oven for an hour and a half.
| |
| 11. Embed in fresh paraffin and cool in water. Make blocks.
| |
| 12. Cut transverse sections 20 u in thickness on microtome.
| |
| 13. Prepare clean albumenized slide, float sections on this in
| |
| order, warm until sections are expanded, remove surplus water.
| |
| Dry for at least two days.
| |
| 14. Remove paraffin with xylene, and
| |
| A. Mount in balsam, or
| |
| B. Run down alcohols to 70 per cent and destain. Run up
| |
| the alcohols, through absolute alcohol and xylene and
| |
| xylene, mount in balsam, or
| |
| C. Run down alcohols to water and restain, dehydrate, clear
| |
| and mount, or
| |
| D. To 95 per cent and counterstain for one minute. Dehydrate, clear, and mount.
| |
| | |
| REFERENCES
| |
| | |
| Baker, J. R. 1933. Cytological Technique.
| |
| | |
| Ballentyne, F. M. 1928. An Introduction to the Technique of Section Cutting.
| |
| | |
| Carleton, H. M. 1926. Histological Technique.
| |
| | |
| Gage, S. H. 1925. The Microscope, 14th Ed.
| |
| | |
| Guyer, M. F. 1917. Animal Micrology, 2nd Ed.
| |
| | |
| Lee, A. B. 1929. The Microtomist’s Vade-Mecum, 9th Ed.
| |
| | |
| McClung, Ch. 1929. Handbook of Microscopical Technique.
| |
| | |
| Oppel, A. 1914. Embryologisches Practikum und Entwicklungslehre.
| |
| | |
| Rugh, R. 1934. Induced Ovulation and Artificial Fertilization in the Frog.
| |
| Biol. Bull. 66, 22-29.
| |
| | |
| Shumway, W. 1926. Fuchsin and Picro-indigo-carmine, a Polychromatic Stain
| |
| for Vertebrate Organogeny. Stain Technology I, 1.
| |
| CHAPTER XV
| |
| STUDY OF EMBRYOLOGICAL PREPARATIONS
| |
| | |
| During the carly stages of development, embryos are too small
| |
| to be studied with the unaided eye. Some observations, to
| |
| be sure, may be made with the dissecting lens, but most embryological work requires the use of the compound microscope.
| |
| Although the student may be familiar with the use of the microscope from the elementary course in biology, he should nevertheless review this subject before proceeding further. In addition, he should at this time familiarize himself with the simpler
| |
| methods of measuring objects with the aid of the microscope,
| |
| as embryological drawings require a strict accuracy as to proportions. A great convenience in embryological work is the camera
| |
| lucida or some other device by means of which accurate outlines
| |
| may be traced. Finally, we must consider the methods by which
| |
| the embryo may be reconstructed in magnified form from serial
| |
| sections, thus returning, in a sense, to the point where the study
| |
| of embryological technique was begun.
| |
| | |
| A. THE USE OF THE MICROSCOPE
| |
| | |
| Nomenclature of the microscope. — The separate parts of the
| |
| microscope (Fig. 239) may be grouped into two systems, the
| |
| mechanical parts, and the optical parts. The principal mechanical parts are the base, from which arises the pillar, attached to
| |
| which is the arm, which may be inclined at the joint. Attached
| |
| to the arm, just above the joint, is the stage, upon which the
| |
| slide is placed for examination, and beneath this, the movable
| |
| sub-stage equipment, consisting of a condenser-sleeve, and one
| |
| or two iris-diaphragms, by means of which the amount of light
| |
| to be used is regulated. At the base of the arm is the mirror, a
| |
| silvered double mirror, with a plane surface on one side and a
| |
| concave surface on the other. At the upper end of the arm are
| |
| two screws, the coarse and fine adjustments, by means of which
| |
| | |
| the barrel of the microscope may be raised or lowered either
| |
| 300
| |
| THE OBJECTIVES 351
| |
| | |
| rapidly or very slowly. The barrel is composed of the bodytube, connected to the arm by a rack and pinion, in the upper
| |
| end of which is enclosed an inner tube, the draw-tube, on which
| |
| is a graduated scale of millimeters representing the tube length
| |
| exclusive of the revolving nose-piece at the lower end. The
| |
| optical parts of the microscope
| |
| are systems of lenses, the condenser, placed in the condensersleeve, the objectives, attached
| |
| to the revolving nose-piece, and
| |
| the oculars, one of which is placed
| |
| at the upper end of the drawtube.
| |
| | |
| The condenser. — This is a
| |
| system of lenses which increases
| |
| the amount of illumination O
| |
| thrown upon the object, and is BJECTIVE
| |
| required only with the higherpower objectives.
| |
| | |
| The objectives. — These are
| |
| systems of lenses which produce
| |
| an enlarged and inverted image
| |
| of the object under proper conditions. Objectives were formerly marked by arbitrary letters V1¢. 239.— Diagram showing parts of
| |
| or numbers, with the lowest-pow- ae compound microscope. (From
| |
| er objectives beginning the series. age.)
| |
| | |
| To-day they are usually indicated by the equivalent focal length
| |
| (E. F.), that is, the focal length of a simple lens at 250 mm. or 10
| |
| inches, or else by the actual magnification (x) at 160 mm.
| |
| (Leitz microscopes, 170 mm.). In some of the older microscopes
| |
| the tube lengths indicated on the draw-tube were calibrated
| |
| without including the length of the revolving nose-piece, then an
| |
| accessory part. When setting up these instruments the length of
| |
| the nose-piece (Leitz, 18 mm.) must be deducted and the drawtube set at the reduced length (Leitz, 152 mm.). The most
| |
| useful objectives for general embryological purposes are the
| |
| 25-mm. or 6 X, which will hereafter be spoken of as the lowerpower objective; the 16-mm. or 10 X, which will be called the
| |
| | |
|
| |
|
| |
| | |
| Coarse
| |
| Adjustment
| |
| | |
| 4yBuayz-aqny
| |
| 352 STUDY OF EMBRYOLOGICAL PREPARATIONS
| |
| | |
| medium-power objective; and the 4-mm. or 40 x, known as the
| |
| high-power objective. For the study of the germ cells, an oilimmersion objective, of which the front lens must be in contact
| |
| with the cover glass by means of a drop of cedar oil, is necessary.
| |
| The most generally used immersion objective is that of 1.9 mm.
| |
| E. F. or approximately 95 xX.
| |
| | |
| Oculars. — These are systems of lenses which magnify the
| |
| real image formed by the objective. Like objectives, these
| |
| were, in the past, usually numbered or lettered, beginning with
| |
| that of the lowest power, but now are marked with the E. F. at
| |
| 250 mm. or the actual magnification at 160 mm. (Leitz oculars,
| |
| 170 mm.). The most useful oculars are the 50-mm. (5 X) or
| |
| low-power ocular, and the 25-mm. (10 x) or high-power ocular.
| |
| When used with the objectives given above, a range of magnification from 30 xX to 450 * may be obtained. A method of obtaining the exact magnification will be described in connection
| |
| with the directions for reconstruction given below.
| |
| | |
| The use of the microscope. —
| |
| | |
| 1. Place the microscope squarely in front of you with the pillar
| |
| toward you and the stage horizontal.
| |
| | |
| 2. Place the low-power ocular in the draw-tube, and adjust
| |
| this to a length of 160 mm. (170 mm. for Leitz instruments) as
| |
| indicated on the millimeter scale. Swing the low-power objective into position. Place the mirror bar in the median line
| |
| and adjust the mirror to secure an even illumination. Use the
| |
| plane side of the mirror. The concave side is employed only
| |
| when the condenser is not in use.
| |
| | |
| 3. Place the slide on the stage so that the object to be examined
| |
| is in the center of the stage aperture, and fasten it down with the
| |
| spring clips provided. With the coarse adjustment, lower the
| |
| body-tube until the objective nearly touches the cover glass.
| |
| Then, with the eye at the ocular, slowly raise the body-tube until
| |
| the object comes into plain view. With the fine adjustment,
| |
| raise and lower the body-tube a little at a time until the point
| |
| at which the smallest details show clearly is discovered. This
| |
| is the focal point.
| |
| | |
| 4, When using the low-power and medium-power objectives, the
| |
| condenser should be lowered until the illumination is evenly distributed. With the high-power objective, the condenser should
| |
| MICROMETRY 353
| |
| | |
| be raised almost to the level of the stage. The iris diaphragm
| |
| should be open sufficiently to illuminate about three-quarters of
| |
| the aperture of the objective. In other words, it is more widely
| |
| open for the low-power objective than for the high-power objective.
| |
| | |
| 5. If a greater magnification is desired, change to the highpower ocular, which will double the magnification. If this is not
| |
| sufficient, return to the low-power ocular and swing the mediumpower objective into position, and so on. On most modern
| |
| instruments, the objectives are par-focal; that is to say, the
| |
| lengths of the objectives are such that when another objective
| |
| is swung into place the object will still be visible. If, however,
| |
| the object is not in focus, it is best to lower the body-tube until
| |
| the new objective almost touches the cover glass, and focus up
| |
| until the object comes into view. If the oil-immersion objective
| |
| is to be used, lower the condenser and place a drop of oil on its
| |
| upper surface; then raise it until it touches the bottom of the
| |
| slide. Place another drop immediately over the object on the
| |
| cover glass and lower the body-tube with great care until the
| |
| front lens of the objective touches the oil. Focus by means of
| |
| the fine adjustment only. .
| |
| | |
| 6. All optical parts of the microscope must be cleaned with
| |
| lens-paper. After the oil-immersion objective has been used,
| |
| the front lens, condenser, and slide should be wiped with a bit
| |
| of lens-paper dipped in xylene and then dried with a fresh piece.
| |
| Never separate any of the optical parts. The microscope should
| |
| be lifted by the pillar unless a special grip is provided to the arm.
| |
| The microscope should be kept in the case when not in use. One
| |
| of the oculars should be left in the draw-tube at all times to
| |
| prevent dust getting on the upper lenses of the objectives. Beginners should try to avoid the error of closing the eye that is not
| |
| in use. Practice will enable the microscopist to work with both
| |
| eyes open and even to alternate the right and left eye at the
| |
| ocular.
| |
| | |
| Micrometry. — The unit of measurement in microscopy is the
| |
| micron (x). It is the one-thousandth part of a millimeter.
| |
| Measurement of microscopic objects is performed with the aid of
| |
| micrometers, of which there are two types, the stage micrometer
| |
| and the ocular micrometer. The former is a glass slide, in the
| |
| center of which, under a cover glass, is a line, usually 2 mm. long,
| |
| 354 STUDY OF EMBRYOLOGICAL PREPARATIONS
| |
| | |
| divided into 200 equal parts, each of which, therefore, is equivalent
| |
| to 10 wu. The ocular micrometer is a glass disc, placed in an
| |
| ocular at the level of the ocular diaphragm, on which is engraved
| |
| a scale, with arbitrary subdivisions. Some oculars are furnished
| |
| with a draw-tube so that the upper lens of the system may be
| |
| focused more sharply upon the scale. The value of the divisions
| |
| indicated on the scale varies according to the amount of magnification of the real image, and so must be obtained for each objective independently, according to the following method:
| |
| | |
| 1. Arrange the microscope as before, taking particular care to
| |
| secure the proper tube-length.
| |
| | |
| 2. Focus the eye-lens on the ocular micrometer scale by means
| |
| of the ocular draw-tube. Focus the objective on the stage micrometer.
| |
| | |
| 3. Make the lines of the stage micrometer parallel with those
| |
| of the ocular micrometer, and determine the value of the divisions
| |
| of the ocular micrometer in terms of those of the stage micrometer. Thus, if it requires 10 spaces of the ocular micrometer,
| |
| and the latter is equal to 0.1 mm., then the value of a single
| |
| space of the ocular micrometer for that particular objective and
| |
| at that particular tube-length is 0.01 mm. or 10 ». Determine
| |
| the value of the ocular micrometer for each objective in the same
| |
| way.
| |
| | |
| B. EMBRYOLOGICAL DRAWINGS
| |
| | |
| Free-hand drawings of microscopic objects can only approximate an accurate representation. However, great pains should
| |
| be taken to secure at least accurate proportions, neat and cleancut lines, and complete labels. Accurate outlines can be secured
| |
| by the aid of the camera lucida, various types of projection
| |
| apparatus, or microphotography.
| |
| | |
| Equipment. — The student will need a hard lead pencil (4H),
| |
| a medium pencil (HB), and blue, red, and yellow colored pencils,
| |
| an eraser, and bond paper to fit the note-book cover used in
| |
| earlier courses.
| |
| | |
| Free-hand drawing. —
| |
| | |
| 1. Lay off the space to be occupied by the drawing, by placing
| |
| four dots at the corners. Rule in two lines, intersecting at the
| |
| center of this space. These will represent the dorso-ventral
| |
| ABBE CAMERA LUCIDA 355
| |
| | |
| and the dextro-sinistral axes, if the drawing is to be of a transverse
| |
| section.
| |
| | |
| 2. Measure the corresponding axes of the sections by means
| |
| of the ocular micrometer, multiply by the desired magnification
| |
| of the drawing, and lay off these magnified measurements on the
| |
| cross lines already drawn. The following magnifications are
| |
| recommended: for the twenty-four hour chick, 100 x; for the
| |
| thirty-three hour chick, 75 x; for the forty-eight hour chick,
| |
| 50 xX; for the seventy-two hour chick, 30 x; for the 10 mm.
| |
| pig, 20 x.
| |
| | |
| 3. Draw in a careful outline of the section and of the internal
| |
| structures, paying particular attention to the proportions, which
| |
| should be measured with the ocular micrometer and laid off on
| |
| the axes at the proper magnification.
| |
| | |
| 4, On one side of the dorso-ventral axis, all structures should
| |
| be colored with the crayons in accordance with the following
| |
| scheme: ectoderm, blue; mesoderm, red; and endoderm, yellow.
| |
| | |
| 5. Label all structures represented in the section, using broken
| |
| lines at right angles to the long axis of the paper to connect the
| |
| label with the structure indicated.
| |
| | |
| 6. Identify the drawing fully, by means of a serial number,
| |
| the species, and stage of development, the number given to the
| |
| series, slide, and section, the type of sections, and the amount of
| |
| magnification. Example: No. 23, Chick, 48 hours, Series 1102,
| |
| Slide 2, section 28, transverse section 50 x. If a drawing has
| |
| already been made of the total embryo or a total mount, indicate
| |
| on this, by means of a heavy ruled line, the position of the section
| |
| just drawn, and number this line with the serial number of the
| |
| section.
| |
| | |
| Abbé camera lucida. — This is an attachment which reflects
| |
| the light from the drawing board, by means of a mirror, to a
| |
| silvered prism, whence the light is reflected to the eye, superimposed on the image of the object which is transmitted through
| |
| a small hole in the silvered surface of the prism directly above the
| |
| ocular of the microscope (Fig. 240).
| |
| | |
| 1. Attach the camera to the draw-tube of the microscope in
| |
| such a way that the mirror projects to the right, and the opening
| |
| in the prism lies above the center of the ocular.
| |
| | |
| 2. Extend the mirror arm to its greatest length and set the
| |
| 356 STUDY OF EMBRYOLOGICAL PREPARATIONS
| |
| | |
| mirror at an angle of 45°. The mirror arm must be parallel to
| |
| the drawing board.
| |
| | |
| 3. Try various combinations of objectives and oculars until an
| |
| image of the desired magnification appears on the paper. Magnifications intermediate to those obtainable in this way may be secured
| |
| by varying the tube-length or by
| |
| raising or lowering the drawing
| |
| board. If the stage of the microscope interferes with the drawing, the
| |
| mirror should be set at an angle of
| |
| 40° or 35° and the drawing board
| |
| tilted toward the microscope at an
| |
| angle of 10° or 20°, respectively, by
| |
| means of wooden images. If the
| |
| image is stronger than the reflection
| |
| of the pencil point, a smoked glass
| |
| may be placed beneath the prism, or
| |
| the aperture of the iris diaphragm
| |
| may be reduced. If the reflection of
| |
| the pencil is stronger than the
| |
| image, smoked glass may be placed
| |
| . at the side of the prism or the amount
| |
| Fig. 240. — Diagram showing prin- Of light falling on the paper reduced
| |
| | |
| ciple of the Abbé camera lucida. by means of a screen.
| |
| | |
| Path of image seen in microscope 4, Draw in the outlines of the sec
| |
| shown in broken lines, that on, .
| |
| | |
| drawing paper shown in unbroken tions and the larger internal struc
| |
| lines. (From Gage.) tures. The details may be added
| |
| | |
| free-hand.
| |
| | |
| 5. Remove the slide and substitute a stage micrometer. Trace
| |
| in part of the scale by means of which both the magnification
| |
| of the drawing and the absolute size of the object may be computed
| |
| readily.
| |
| | |
| Projection apparatus. — Where many drawings are to be made,
| |
| as in the case of reconstructions, some form of apparatus by
| |
| means of which the image of the section may be projected
| |
| directly upon the paper is very helpful. There are many types of
| |
| projection apparatus, directions for the use of which may be
| |
| obtained with the instruments.
| |
| THE GRAPHIC METHOD (OF HIS) 357
| |
| | |
| Microphotography. — The photography of minute objects with
| |
| the aid of the microscope is of great assistance in embryology.
| |
| However, the methods are so difficult, the apparatus so complex,
| |
| expensive, and delicate, and the process requires so much technical knowledge and skill, that microphotography has been considered a field too advanced for the beginning student, although
| |
| a method described by Headland seems to overcome these
| |
| difficulties to a large extent. In recent years the motion-picture
| |
| camera has been adapted for use with the microscope, and
| |
| excellent results have already been obtained.
| |
| | |
| C. RECONSTRUCTION
| |
| | |
| After an embryo has been sectioned, it is sometimes necessary
| |
| to reconstruct some part of it from the sections. There are two
| |
| important methods: graphic reconstruction, in which a geometric projection of a sagittal section, for example, might be
| |
| made from transverse sections; and plastic reconstruction, in
| |
| which magnified replicas of each section are made of wax and
| |
| piled together so as to make an enlarged model of the object to
| |
| be studied. A complete series of sections of uniform thickness
| |
| and accurate orientation is required for either type of reconstruction, and an outline drawing of the embryo before sectioning is
| |
| of great assistance.
| |
| | |
| The graphic method (of His).— This method can best be
| |
| described by giving practical directions for a particular problem,
| |
| e.g., to prepare a geometrical sagittal projection 20 x of the
| |
| neural tube of a 10 mm. pig embryo from a series of transverse
| |
| sections 20 pz in thickness.
| |
| | |
| 1. From the lateral view of the embryo drawn before sectioning,
| |
| make an outline drawing 20 x.
| |
| | |
| 2. Draw a median line corresponding to the cephalo-caudal
| |
| axis, the length of which, in this case, should be 200 mm.
| |
| | |
| 3. Count the number of sections in the series, in this case, 500.
| |
| | |
| 4. Locate the position of each transverse section which you
| |
| have drawn on the median line of the outline. Thus if the most
| |
| anterior section drawn was the fiftieth of the series of 500 sections, it would be located at a point 1/10 of the total length of
| |
| the axis (200 »), or 20 mm. from the cephalic end.
| |
| | |
| 5. Theoretically, each section is at right angles to the median
| |
| 358 STUDY OF EMBRYOLOGICAL PREPARATIONS
| |
| | |
| line, but this angle may be greater or less as a result of variations
| |
| in technique. Study each drawing of a cross section in connection with the ‘drawing of the total embryo and determine the
| |
| angle made by that section with the cephalo-caudal axis of the
| |
| embryo. Draw in, at each point located on the median line, a
| |
| cross line at the proper angle so determined. These lines represent the dorso-ventral axes of the transverse sections. Their
| |
| lengths should correspond with those of the dorso-ventral axes
| |
| of the drawings of the transverse sections previously made at
| |
| the same magnification, 20 x.
| |
| | |
| 6. Plot in on each section-plane line (dorso-ventral axis) the
| |
| dorsal and ventral boundaries of the neural tube as determined
| |
| from measurements of the drawings already made. Interpolate
| |
| by direct measurement and magnification of these points on
| |
| intervening sections.
| |
| | |
| 7. Sketch in the contours of the neural tube by connecting up
| |
| the points which have just been plotted. Compare the drawing
| |
| with a sagittal section of an embryo in the same stage of development.
| |
| | |
| Plastic reconstruction. — This method also will be indicated
| |
| by practical directions for the reconstruction of a particular
| |
| organ, in this case, a model 50 x of the heart of a 10 mm. pig,
| |
| from a series of transverse section, 20 u in thickness.
| |
| | |
| 1. Prepare a number of wax plates of the proper thickness.
| |
| In this case, if every section is to be reconstructed, the thickness
| |
| of the plates must be 50 X 20 u,orlmm. Nearly as good results
| |
| can be obtained by reconstructing every second section and
| |
| making the plates twice as thick. The wax is prepared according
| |
| to the following formula:
| |
| | |
| Beeswax... 0... ccc cc cece eee eee e eee eeeeee 6 parts
| |
| Paraffin, 56° C. mp... ccc eee cceeec eee eeeeeee 4 parts
| |
| Rosin, white lump. ...... 0. cee eee cece ee eee eee 2 parts
| |
| | |
| Mix and melt.
| |
| | |
| Pour 130 grams of this wax into a pan with an inside measurement of 500 X 280 mm., into which boiling water has been poured
| |
| to a depth of 15 mm. This amount of wax will make a plate
| |
| 1 mm. in thickness. Bubbles in the wax may be removed by
| |
| playing the flame of a bunsen burner over the surface as it is cooling. As the surface hardens, cut the edges free from the sides of
| |
| PLASTIC RECONSTRUCTION 359
| |
| | |
| the pan. When the wax has set put is still malleable, roll up
| |
| the plate and remove it to a soapstone slab, where it is unrolled
| |
| and allowed to cool.
| |
| | |
| 2. With the help of a camera lucida or projection apparatus,
| |
| prepare outlines 50 x of the heart in all the sections in which it
| |
| isfound. Number the drawings consecutively and note the serial
| |
| number of the sections drawn, so that it will be possible to
| |
| check the drawings later if necessary. Note also whether the
| |
| right and left sides of the drawing actually correspond with the
| |
| right and left sides of the embryo or whether this condition is
| |
| reversed. This is very important, as a mistake at this point
| |
| would render the reconstruction valueless.
| |
| | |
| 3. Transfer the drawings to the wax plates by means of carbon
| |
| paper. Place the wax plates on a sheet of glass, and cut out the
| |
| parts to be preserved with a sharp scalpel, leaving bridges of wax
| |
| to connect the parts which would otherwise be separated. These
| |
| bridges are best made in the form of arches bending towards the
| |
| outside of the section.
| |
| | |
| 4. Pile the sections in order, taking care to avoid the reversal
| |
| of right and left sides, and to get an accurate fit. It is best to
| |
| group the sections in piles of ten. A steady pressure of the hand
| |
| will be sufficient to cause the sections to adhere to each other.
| |
| The bridges may be cut away and stout pieces of wire substituted. Heat the wire at each end and press into position.
| |
| After the wire is set, the wax bridges are cut away and the edges
| |
| of the piece smoothed with a heated scalpel or aluminum modeling
| |
| tool.
| |
| | |
| 5. When all the sections have been combined in groups of ten,
| |
| these groups should be united and the completed model smoothed
| |
| in the same way. Such models may be painted or dissected,
| |
| and mounted on wooden supports as desired. They are quite
| |
| permanent if not exposed to high temperatures. Plaster of
| |
| Paris molds and casts may be made from them in the customary
| |
| manner.
| |
| | |
| REFERENCES
| |
| | |
| Belling, J. 1930. The Use of the Microscope.
| |
| Cage, S. H. 1932. The Microscope, 15th Ed.
| |
| Guyer, M. F. 1917. Animal Micrology, 2nd Ed:
| |
| 360 STUDY OF EMBRYOLOGICAL PREPARATIONS
| |
| | |
| Headland, C. I. 1924. A Simple and Rapid Photomicrograph for Embryological
| |
| Sections. Anatomical Record, XVII, 2.
| |
| | |
| Lee, A. B. 1929. The Microtomist’s Vade-Mecum, 9th Ed.
| |
| | |
| Mueller, J. F. 1935. A Manual of Drawing for Science Students.
| |
| | |
| Norman, J. R. 1923. Methods and Technique of Reconstruction. Journal of the
| |
| Royal Microscopical Society.
| |
Shumway W. Introduction to Vertebrate Embryology. (1935) John Wiley & Sons, New York
- Shumway (1935): Preface - Contents | Part I. Introduction | Part II. Early Embryology | Part III. Organogeny | Part IV. Anatomy of Vertebrate Embryos | Part V. Embryological Technique
Historic Disclaimer - information about historic embryology pages
|
Pages where the terms "Historic" (textbooks, papers, people, recommendations) appear on this site, and sections within pages where this disclaimer appears, indicate that the content and scientific understanding are specific to the time of publication. This means that while some scientific descriptions are still accurate, the terminology and interpretation of the developmental mechanisms reflect the understanding at the time of original publication and those of the preceding periods, these terms, interpretations and recommendations may not reflect our current scientific understanding. (More? Embryology History | Historic Embryology Papers)
|
Introduction to Vertebrate Embryology (1935)
Part II. Early Embryology
Chapter III The Germ Cells
The germ with which the development of the vertebrate commences is the fertilized egg, or zygote. Before discussing the
development of the zygote, it is advisable to examine the gametes,
egg and sperm, whose union results in its existence. We shall
proceed first to the description of the gametes, comparing them
with each other and with a generalized cell. Next we shall consider the way in which the germ cells originate and become
mature. Thereafter we shall turn to the study of fertilization.
A. THE GAMETES
Vertebrates are characterized by the bisexual method of reproduction, in which there are two distinct sexes: the female, or
egg-producing individuals; and the male, or sperm-producing
individuals. Among the protochordates (tunicates) we find
groups in which the same individual produces both eggs and
sperms. Such individuals are called hermaphrodites. This phenomenon is rare among the vertebrates and is not typical of any
species.
The two kinds of gametes, eggs and sperms, differ from each
other in appearance, size, and structure. These differences will
be more apparent after a brief review of cell structure in general.
TABLE 3
STRUCTURE OF THE CELL
A. Nucleus (composed of karyoplasm).
1. Reticulum (composed of chromatin).
2. Karyolymph (nuclear sap).
3. Nucleolus (plasmasome).
4. Nuclear membrane.
B. Cytosome (composed of cytoplasm).
. Hyaloplasm (ground-protoplasm).
. Centrosomes (centrioles).
. Mitochondria (chondriogomes).
. Golgi bodies (dictyosomes).
. Plastids.
. Metaplasm (relatively lifeless accumulations).
Plasma membrane.
C. Envelopes or matrix (cell wall).
The cell. — The familiar definition of a cell (Fig. 5) is, “ a mass
of protoplasm, containing a nucleus, both of which have arisen by
the division of the corresponding elements of a preéxisting cell.”
Protoplasm in this sense refers to the living substance of the cell,
including both the material inside the nucleus and that in the
cell body or cytosome. It is customary to use the term karyoplasm (nucleoplasm) for the nuclear protoplasm, and the word
cytoplasm for the protoplasm of the cell body. Some writers employ only the words nucleus and cytoplasm to distinguish
between nucleus and cell body.
Fig. 5. — Diagram of a composite cell. (After Wilson.)
The nucleus. — The cell nucleus is generally a rounded body
separated from the eytosome by a delicate nuclear membrane.
Within this is a transparent ground substance known as the karyolymph or nuclear sap. But the characteristic substance of the
nucleus is its chromatin, a substance staining sharply with basic
dyes, and arranged usually in a network of threads called the
reticulum (Sharp). Sometimes swellings, chromomeres, are apparent at the nodes of the network. The nucleus usually contains a smaller body known as the nucleolus, a droplet of some
THE CYTOSOME 33
material heavier than the nuclear sap, but staining with acid
dyes. Its staining properties alter during cell division.
The nucleus may fragment to form polynuclear cells. It may
also divide, often many times, while the cell body remains undivided, resulting in the formation of a syncytium. Sometimes
the nucleus may be ejected to leave enucleate cells such as the red
blood corpuscles of mammals. But in general every cell has one
nucleus.
The type of nucleus here described is known as the vesicular
type. There is distinguished also the massive or compact type of
nucleus, in which the chromatin forms apparently a solid mass,
as in the sperm cell. Then there is a diffuse type, in which the
nuclear membrane is absent and the chromatin is scattered
through the cell body in granules called chromidia.
The cytosome. — The cytoplasm of the cell body includes an
outer delicate semipermeable membrane known as the plasma
membrane. This is the surface at which the protoplasm of the
cell is in contact with its environment. Within this is the liquid
ground substance or hyaloplasm, in which are distributed a
number of differentiated bodies. Of these cytoplasmic inclusions
the more important seem to be the centrosomes, mitochondria,
and Golgi bodies, all of which appear to have the properties of
independent growth and division.
The centrosomes (centrioles), small spherical bodies, one or two
in number, lie near the nucleus. They seem to be concerned in
the process of cell division. In cells with locomotor organs, like
the tail of the sperm, the centrosomes are connected with the
contractile element of the cell.
The mitochondria (chondriosomes) are small rods, or granules,
very numerous and scattered through the cytoplasm. They are
dissolved by many common methods of preparing cells for observation, but can be demonstrated in the living cell by a stain
called Janus Green B. They are preserved by special chemicals,
e.g., osmium tetroxide.
The Golgi bodies (dictyosomes) are sometimes scattered through
the cytoplasm but often aggregated into a network, the Golgi apparatus. Some authors deny that there is a real structure of
Golgi bodies and speak therefore of the Golgi material or the
Golgi zone. Other investigators have sought to identify these
34 THE GERM CELLS
bodies with the plastids, cytoplasmic elements which are found in
plant cells. Golgi bodies are hard to identify in living cells but
can be demonstrated by special techniques involving the use of
osmium tetroxide or silver nitrate. Their function is doubtful,
but there is some reason to believe that they are concerned with
the elaboration of substances within the cell such as enzymes.
Still another type of inclusion in the cytosome is represented
by the plastids. These bodies are found more frequently in plant
cells, e.g., chloroplasts, the chlorophyll bodies, which appear to
have the capacity of independent growth and division.
Metaplasm is the name given to all those bodies in the cytoplasm which clearly do not possess the properties of independent
growth and division. These may be aggregated in vacuoles or
distributed in tiny droplets, granules, etc. Among these are
such bodies as secretory granules, intermediate stages in the production of cell secretions (enzymes, etc.). Storage granules are
end stages in the accumulation of reserve food materials such as
yolk, oil, starch, etc. Here also we may include the minute
pigment granules. Embryologists sometimes use the term deutoplasm for reserve food materials in the cell.
The cell wall. — In concluding this brief review of cell structures we must recall that the cell may secrete a wall around itself
such as the vitelline membrane. In some tissues these cell walls
unite to form a matrix such as the intercellular substance of
cartilage or bone.
The sperm.— The male germ cell of vertebrates is a very
minute flagellate cell ranging in size from 20 microns (crocodile)
to more than 2 mm. (Discoglossus, an amphibian). The general
shape is that of a tadpole with an excessively long tail, but there
are sufficient differences among these tiny cells for them to be
identified by specialists.
The sperm (spermatozoon) consists of a head and a tail (Fig. 6).
The head contains the nucleus, which is compact and stains very
deeply with basic dyes. Here also is the acrosome, usually at the
apical end, originating from Golgi bodies, possibly connected with
the production of some secretion involved in fertilization. The
head is surrounded with a delicate plasma membrane.
The tail consists of three divisions: middle-piece, main-piece,
and end-piece. The middle-piece contains two centrosomes.
Yolk. — The bulk of the egg is due to the presence of metaplasm in the form of yolk. This substance contains the principal
foodstuffs for the developing embryo. Studies on the yolk of the
hen’s egg indicate that it contains water (50 per cent), proteins,
fats, carbohydrates, inorganic salts, vitamins, pigments, and
enzymes (Needham).
The yolk is present in the form of spheres, ovoids, or discs, which
stain usually with basic dyes. The yolk tends to accumulate in
one hemisphere of the egg, forcing the nucleus into the other.
Since the yolk is heavier than the other constituents of the egg, the
yolk-laden hemisphere is the lower one when the egg is suspended
in water. In large-yolked (macrolecithal, megalecithal) eggs,
such as those of the frog and chick, the accumulation of the yolk
in one region is so marked that they are known as telolecithal eggs.
In small-yolked (microlecithal, oligolecithal) eggs, like those of the
amphioxus and of man, the yolk is distributed more generally and
they are called isolecithal (homolecithal).
Polarity. — Even in isolecithal eggs there is a visible distinction
between the two hemispheres of the egg, so that an axis exists
from the center of one hemisphere to that of the other. This,
known as the polar axis, is the earliest indication of a differentiation in the egg. The two ends of the axis are known as the poles.
The polar bodies, referred to in the preceding chapter, are formed
at one of these which is known as the animal (apical) pole. It is
sometimes called simply the pole. The other is called the vegetal
(vegetative, abapical) pole, sometimes the antipole. The nucleus
always lies in the polar axis, more or less towards the animal
pole. The yolk shows a gradation from the animal towards the
vegetal pole. We shall observe in later chapters that the animal
pole marks the anterior end of the developing embryo and the
vegetal pole marks the posterior end. There is also reason to
believe that the polar axis, in addition to being the first expression
of symmetry in the egg, marks a gradient of metabolism (Child).
By this is meant that metabolic processes are accelerated at the
animal pole and progressively retarded towards the vegetal pole.
A considerable body of evidence shows that the animal pole of
the egg is the one which was most active in physiological exchange
with its environment while still in the ovary. It is the pole of the
egg which is attached to the ovary in the amphioxus (Conklin) and the chick (Conklin). It has been suggested that in the frog
the animal pole of the egg is the one lying nearest the arterial
blood supply (Bellamy).
Egg envelopes. — The ovum usually possesses, in addition to
the plasma membrane, a variety of protective envelopes which
are divided into three classes according to the mode of their formation. Primary envelopes are those formed by the egg itself,
such as the delicate vitelline membrane. The secondary envelopes are those formed by the follicle cells which immediately surround the egg in the ovary. A good example is the so-called
“chorion” of one of the cyclostomes, Myzine (Fig. 9). It is
usually quite difficult to distinguish primary from secondary envelopes, and it is probable that many vitelline membranes are
compound in origin. In those vertebrates in which fertilization
is external, such as the cyclostomes and bony fish, the primary
and secondary envelopes are often perforated by openings called
micropyles through which the sperm may have access to the egg.
The tertiary envelopes include all those formed by the walls of the
oviduct during the passage of the egg. Examples are the egg
albumen, shell membranes, and shells of such groups as the reptiles, birds, and the egg-laying mammals; the egg capsules of the
elasmobranchs, and the egg jelly of the amphibia and many bony fish. These envelopes are not formed until after fertilization,
except in the case of the egg jelly, and this does not attain its
final thickness until after the entrance of the sperm, when it
swells by the absorption of water.
Fig. 9. — Egg of Myxine, showing “chorion” and micropyle (after Dean).
THE EGG OF THE AMPHIOXUS. — The eggs (Fig. 10A) are 0.1
mm. in diameter. Before maturation the large nucleus is roughly
0.05 mm. in diameter displaced well towards the animal pole.
The cytoplasm consists of a thin outer layer relatively free from
Fig. 10. — Typical eggs. A, amphioxus, approx. X70 (after Wilson in Willey).
B, frog X8. C, hen X? (after Duval). D, human X250 (after Allen in Arey).
yolk, and probably containing mitochondria. The rest of the
cytoplasm contains yolk. There are no egg envelopes except
perhaps a vitelline membrane. The egg is classed as isolecithal.
THE EGG OF THE FROG. — The diameter of the egg (Fig. 10B)
is 1.7 mm. (R. pipiens, Wright), with a large nucleus before
maturation. There is a thin outer layer of cytoplasm, containing
granules of pigment in the animal hemisphere. Pigment is also
found around the nucleus. The yolk is distributed with fewer
and smaller platelets in the animal hemisphere grading down to
more and larger platelets in the vegetal hemisphere. There are
a vitelline membrane (primary), ‘‘ chorion ”’ (secondary), and one
to three layers of egg jelly (tertiary). The eggs are discharged
40 THE GERM CELLS
in large masses which adhere to each other by means of this jelly.
The eggs are classified as telolecithal.
THE EGG OF THE CHICK.— The hen’s egg (Fig. 10C) is extremely telolecithal. The cytoplasm, with the nucleus in its
center, forms a small germinal disc upon the great mass of yolk.
This yolk is arranged in
wei e, Albumen or concentric layers of
Outer shell yellow and white mateshen Yial around a_ central
x
membrane
Inner shell
membrane mass of white yolk,
chamber called the latebra (Fig.
11). From this latebra
a stalk of white yolk (the
neck of the latebra) exFia. 11. — Diagram of hen’s egg sectioned tends up ward. The
(after Lillie). germinal disc rests on
this isthmus. The yolk
and germinal disc are surrounded by a delicate vitelline membrane
(primary). This in turn is surrounded by the albumen, a viscous
tertiary membrane twisted spirally about the egg from left to
right, starting from the broad end of the egg. The albumen
next to the vitelline membrane is denser than the rest and is
prolonged into two spirally twisted cords, the chalazae, one
at cither end of the egg. The albumen is in turn surrounded
by two parchment-like shell membranes, of which the inner
one is the thinner. These two are separated at the blunt end
of the egg, thus forming the air chamber. The egg shell is a
calcareous deposit upon the outer shell membrane. Its color is
due to bile pigments of the hen. The germinal dise is about
4 mm. in diameter, the yolk about 40 mm. The size of the egg
as a whole varies largely, depending on the amount of albumen
deposited around the yolk.
Giant and dwarf eggs are sometimes recorded. In the hen’s
egg, double- and triple-yolked eggs are known, as well as those
which have no yolk at all. A very strange abnormality is known
as the “‘ ovum in ovo,” where one egg is formed around another.
The eggs of birds are either male-producing or female-producing,
a statement, based solely on the evidence of genetics as no visible
differences have been observed.
THE EGG OF MAN. — The human egg (Fig. 10D) is extremely
smal}. The yolk granules are concentrated about the nucleus,
which is slightly excentric. It is not positively known whether
a vitelline membrane is present. But the egg is enclosed in a
thick capsule with radial striations (canals?), the zona pellucida.
It is not clear whether this is a primary or secondary envelope.
At the time the egg leaves the ovary it is still surrounded by a
layer of follicle cells which
make up the so-called A ge \4- Zona pellucida
corona radiata (Fig. 12). sacs Rp ie
The egg may be termed ee NE Saas
isolecithal. Its diameter ae :
is about 0.13 mm.
Eggs and their environment. — Needham has
recently pointed out that
eggs differ from one another in respect to the
physico-chemical constitution of the unfertilized
egg, and the possibility
of obtaining necessary material from the environment. The
marine egg, exemplified by the amphioxus, develops in a medium
containing oxygen and inorganic salts. The egg is organized in
such a manner as to facilitate the exchange of materials with the
environment, and the yolk is small in amount and (to judge from
analyses made on marine fish) relatively poor in fats and inorganic
salts. Development is rapid up to the hatching stage, but thereafter the larva takes a long time to attain its full size and sexual
maturity.
The egg which develops in fresh water, like that of the frog, does
not have a medium so rich in salts as the marine egg. It is therefore originally equipped with a larger store of this material. But
the aqueous medium still affords facilities for the exchange of
carbon dioxide and oxygen and for the disposal of nitrogenous
wastes. The jelly with which the frog’s egg is provided consists
almost entirely of protein and water. Diffusion takes place
through it readily, and it affords protection against mechanical injury and bacterial infection, as well as furnishing a source of
nourishment immediately after hatching.
Via. 12.— Human egg (ovarian) X200 (after Waldeyer).
The terrestrial (cleidoic) egg, such as that of the hen, stands easily first in respect to the amount of yolk present. The ratio
of fat to protein in the yolk is also the highest. It is obvious that
the egg must contain all the material necessary for growth except
free oxygen and water, for these are the only substances passing
from the atmosphere through the protective envelopes of the egg.
Hence, as pointed out by Milnes-Marshall, except in the earliest
stages the chick develops more rapidly than the amphioxus and
attains its adult form in a much shorter time. The egg albumen
also a source of food is a watery solution of protein with some
carbohydrates. As we shall see in later chapters, the relative
isolation of the embryo in the cleidoic egg is correlated with the
development of its extra-embryonic sacs, i.e., the amnion or water
bath, and the allantois which serves in the first instance to store
nitrogenous wastes.
_ The uterine egg, typical of the mammals, is characterized by
little yolk, for, from a very early period, its nourishment is derived
exclusively from the body of the mother. Accordingly there is a
precocious separation of a special layer, the trophoblast, concerned with implantation, and later the development of a special
organ of interchange, the placenta.
Comparison of the egg and the sperm. — Both gametes are
morphologically complete cells. Each has a nucleus and a cytosome containing representatives of the centrosomes, mitochondria,
and Golgi bodies. Each hasa plasma membrane. Yet neither is
capable of independent, continued existence, for physiologically
they are unbalanced. The egg is large, inert, and contains a vast
store of metaplasm, is protected by egg envelopes, and has lost
the power of continued division. The sperm is small, highly
motile, contains little cytoplasm and no metaplasm, is devoid of
protective membranes, and in itself has lost the power of continued division. We shall now turn to the study of the development of the germ cells and see how the structural differences, at
least, arise.
TABLE 4 CoMPARISON OF THE VERTEBRATE EGG AND SPERM
Ovum
Ceil structures
Sperm
Large amount
One, disappears in maturation
Diffuse
Cytoplasm
Centrosomes
Mitochondria
Small amount
Two, retained in maturation
Spiral coil
Diffuse Golgi bodies Acrosome
Present Plasma membrane | Present
Vesicular Nucleus Compact
Present Nucleolus Indistinguishable
Present Nuclear membrane | Present
Other differences
Large Size Small
Quiescent Movement Swims actively
None Motile organs Flagellum
Egg envelopes Protection None
Spheroid Shape Tadpole
Few to many Numbers produced | Very many
B. GAMETOGENESIS
Gametogenesis is the term applied to the history of the gametes
— their origin and development (Fig. 13).
The special history
of the male gametes is called spermatogenesis, that of the female
gametes odgenesis.
The origin of the germ cells. — Weismann is responsible for a
theory that the germ cells separate completely from all the other
cells of the body (soma cells) ata very early stage in development.
There is some evidence for this-in the embryology of a few invertebrate animals such as Ascaris, a parasitic roundworm. In
the very first cleavage of the fertilized egg, the two daughter cells
show a-striking difference, for when one of the daughter cells
divides it retains all the chromatin of its nucleus whereas the other
gives up a portion of this material to the cytosome. This phenomenon has been called chromatin diminution, and the cell
showing this characteristic becomes a soma cell. The other is
THE GERM CELLS
Soma Cells
Meiosis <
st
\
7
o———> o—-e0-+83-|=
ow-~
a
o
Zygote
Fig. 13. — Diagram of gametogenesis, male on left, female on right (after Wilson).
THE ORIGIN OF THE GERM CELLS 45
known as a stem cell (Fig. 14), and in its division it produces in
turn one cell which will be a soma cell and one which will be a
germ cell. Eventually a stem cell gives rise to two identical
cells, both of which are germ cells. These are known as primor
cg
Fia. 14. — Origin of stem cells in Ascarts. A, first cleavage. B, C and D, second
cleavage. P; and Ps are stem cells. 8S, (which gives rise to A and B) and 8; are
soma cells. (From Richards after Boveri.)
dial germ cells, and, from this time on, they and their descendants
produce germ cells only.
This theory of the distinction between germ cells and soma
cells has held an important place in the history of biology because
46 THE GERM CELLS
it seemed to deny the possibility of the inheritance of characteristics acquired after fertilization. In other words, the characteristics would be acquired by the soma cells whereas inheritance is
transmitted ‘by the germ cells which are entirely distinct. Now
that we know that the nuclei of all cells are identical, whether
they are germinal or somatic, the theory of the continuity of the
germ cells has less theoretical importance.
Primordial germ cells. — In all vertebrates, so far as is known,
the germ cells are first recognizable in the lining of the gut at a
very early stage of development. These primordial germ cells,
as they are called, are distinguishable by their large size, clear
cytoplasm, and heavily staining nucleus (Fig. 15). From the gut
Mesonephric duct
Postcaval vein
ee ee = - ¥ — “ P me a i * , q
Else sé: \ \__ Primordial
8 5D a 4% germ cell
Dorsal mesentery
Fia. 15. — Primordial germ cells in the frog (Rana sylvatica). Part of transverse
section through 10 mm. larva, showing coelomic roof, X375. (After Witschi,
1929.)
wall they migrate into the mesentery suspending the gut from the
roof of the coelom, and thence to the wall of the coelom. Here
they multiply rapidly and produce two longitudinal ridges, which
are the primordia of the sex glands, or gonads.
The ’gonia. — There are two opinions concerning the fate of
the primordial germ cells:i in vertebrates: one that they give rise
to all the later generations of germ cells; the other that they
degenerate and the later germ cells arise independently from the
tissue of the gonads. In any case, the germ cells which continue
to multiply actively in the gonads are known as ’gonia: spermatogonia i if they are to give rise to sperm, odgonia ifthey givé Tise
to eggs.
THE MATURATION DIVISIONS 47
The ’cytes. — When the individual becomes sexually mature,
individual ’gonia undergo a period of growth by means of which
they become transformed into ’cytes (auxocytes, meiocytes):
spermatocytes if male, odcytes if female. The ’cyte (Fig. 16) is
a large cell with a vesicular nucleus, two centrosomes surrounded
by a clear area sometimes known as the sphere substance, which
is in turn surrounded by a layer A
of Golgi bodies, and a cloud of
mitochondria.
The maturation divisions. —
Each ’cyte gives rise to four iS
daughter cells or gones (Sharp) by __ diosome Golgi bodies
means of two cell divisions. i ,
These divisions are unique because 4,, 1g — Diagram of an early cyte
of certain internal phenomena and (auxocyte). (After Wilson.)
are known as the maturation
divisions. The nature of these divisions will be discussed in more
detail in a later chapter (page 64). Meantime we note that the
spermatocyte gives rise to four cells of equal size, the spermatids,
each of which will be transformed into a sperm. The odcyte on
the contrary gives rise, by the first maturation division, to two
cells one of which is very minute, the first polar body (polocyte I).
The larger cell undergoes a second unequal division, resulting in
the production of a second polar body (polocyte IT) and the mature
egg or ovum. It will be recalled that among the vertebrates the
TABLE 5
Stages IN GAMETOGENESIS
Nucleus
Mitochondria
Spermatogenesis General Odgenesis
(Stem cells) ¢
Primordial germ cells *
Period of migration ~
Spermatogonia ’Gonia O&gonia
Period of multiplication
Spermatocytes ’Cytes. Odcytes
Period of maturation coe
Spermatids Gones (Sharp) Ovum and polocytes (Odtids)
Period of metamorphosis
Sperms
®
48 THE GERM CELLS
sperm enters the egg before the production of the second polar
body. Sometimes the first polar body also divides so that -four
cells (odtids) may be produced by the odcyte. .
Spermatogenesis. — The male ’cyte (primary spermatocyte) is
a large cell containing a large vesicular nucleus, more or less
excentric. Near the nucleus, and in the center of the thicker
layer of cytoplasm surrounding it, are to be seen one or two
centrosomes, surrounded by a clear substance known as the sphere
substance. This compound body is known as the idiosome, and
with it are often associated the Golgi bodies, sometimes so closely
connected as to form an investing reticulum or even a shell.
Around the idiosome are also grouped the mitochondria forming
a cap which sometimes includes the nucleus as well.
The primary spermatocyte divides, giving rise to two secondary
spermatocytes, which divide again, often without intermission,
each forming two spermatids. The four spermatids thus produced from the primary spermatocyte are later transformed into
the sperms.
During the two divisions mentioned above, the chromatin of
the spermatocyte nucleus is distributed to the spermatids in such
a way that they will differ from each other in respect to the
nuclear contents. The details will be discussed later (page 64).
The centrosomes divide at each cell division so that each spermatid has a centrosome. The Golgi bodies, each with a small
amount of sphere substance, are divided among the four spermatids, in each of which they aggregate to form an idiosome. The
mitochondria are divided with almost exact evenness among the
spermatids, in each of which they assemble to form a paranucleus
(nebenkern). A plasma membrane is present.
In the transformation of the spermatid into the mature sperm
(Fig. 17), the nucleus, having previously extruded a large amount
of material, condenses into a deeply staining mass which elongates
into its final shape. The centrosome divides, and the two centrosomes take up a position which marks the posterior end of the
future sperm, one centrosome (proximal) lying against the
nucleus, the other (distal) posterior to the first. The paranucleus
also takes a posterior position while the idiosome moves around
the nucleus to the anterior end. The greater part of the cyto
plasm is sloughed off.
SEMINATION 51
Ovulation. — Within the ovary the vertebrate egg is surrounded by nurse cells which make up a nest or follicle (Fig. 19).
Within this it enlarges and may undergo its first maturation
division. Periodically, varying from once a year in most vertebrates to once a month in the human species, or daily in the
domestic fowl, eggs are discharged from the ovary. In
numbers this discharge
varies from a single egg as
in man or the fowl to
thousands in the frog or
millions in many fish.
The factors bringing
about ovulation are diverse.
In the frog it has been
shown by Rugh! that ovulation is brought about by
the contraction of a thin Fia. 19. — Transverse section through part of
muscular layer in each fol- frog ovary. X95.
licle, plus the action of an
enzyme which digests the outer wall of each follicle and thereby
weakens it. In mammals a follicular fluid is secreted about
the egg, enlarging the follicle until it protrudes from the surface. Finally the outer wall of the follicle, now very thin,
ruptures, owing perhaps to factors similar to those acting on the
frog’s egg. It has been shown in many vertebrates that ovulation
can be induced at any time by means of a hormone secreted by
the anterior lobe of the pituitary gland (page 332).
From the ovary the eggs are caught up in the open end of the
oviduct, down which they pass to the exterior. In many aquatic
forms they are discharged directly. In others they accumulate
in an enlarged portion of the oviduct known as the uterus, awaiting discharge from the body; such animals are known as oviparous
(amphioxus, frog, chick). In still others the egg remains in the
uterus until development has reached an advanced stage; these
are the viviparous animals (man, etc.).
Semination. — This term is applied to the discharge of the
sperms. These cells remain in the testis (Fig. 20) until mature,
1 Jour. Exp. Zool. In press.
Wall of ovary
Follicle cells
Odcyte
Oégonium
52 THE GERM CELLS
often attached to nurse cells. When discharged they pass through
tubules of the testis which lead directly to a sperm duct. They
become motile upon reaching the medium in which fertilization
takes place. Enormous numbers are produced at a single discharge (over 200,000,000 in man).
In aquatic animals such as the amphioxus and fish the two
sexes congregate together at the breeding season, and eggs and
sperms are discharged together. In some cases even aquatic
animals have copulatory organs which introduce the sperms into
the oviduct, bringing about internal fertilization. In the frog,
Tubule
Interstitial
cells
Spermatocytes
Fig. 20. — Section through part of frog testis. 200.
the males and females unite in pairs (amplexus), thus ensuring
that the sperms are discharged simultaneously with the eggs so
that fertilization, although external, is regulated. In all terrestrial vertebrates fertilization is internal.
Fertilization. — The actual fertilization of the egg (syngamy)
has been observed in the amphioxus and the frog, but our detailed
knowledge of the process is obtained from the study of such
marine invertebrates as the sea urchin. The essential act in fertilization is the entrance of a single sperm into the egg and the
coming together of the two nuclei (pronuclei) (Fig. 21). But this
phenomenon is preceded by other events concerned in bringing
the sperm to the egg.
Attraction. — One of the factors believed to bring the sperm
towards the egg is an attraction (chemotaxis) caused by the emission of some chemical substance by ‘the egg or the female sex
PENETRATION 53
organs. It is known also that sperms swim in a spiral path, and
it has been suggested that when they come in contact with a solid
object they remain in contact with it (thigmotaxis). If the spiral
brings a sperm obliquely towards the egg the contact flattens out
the spiral, causing the sperm to remain in contact without penetration. But if the sperm arrives at the egg in a radial direction
penetration is facilitated. Lillie has shown that the sea-urchin
egg emits a secretion (fertilizin), which brings about a temporary
and reversible adhesion of the sperm heads in clusters (agglutination). Fertilizin is produced only after the egg is mature and
before it is fertilized.
Penetration. — The sperm bores its way through the egg
envelope but then apparently comes to rest against the plasma
Q Copulation path
On.
Fia. 21. — Diagram to show fertilization of the egg. A, fertilization cone. B, penetration path. C, female copulation path, and rotation of sperm head. D, male
copulation path. E, cleavage path. F, first cleavage.
membrane. Meantime there appears at the surface of the egg a
cone or even a long filament of cytoplasm which comes in contact
with the sperm head. It then retracts drawing the sperm to the
egg and engulfing it (Fig. 21A, B). Thereafter, and commencing
at the point where the sperm head was engulfed, a thin layer of
surface protoplasm is elevated to form what is known as a fertilization membrane (vitelline membrane?). In older days, when
it was thought that the sperm bored its way into the egg, it was
believed also that the fertilization membrane acted as a bar to
54 THE GERM CELLS
other sperms. Apparently the elevation of the membrane is due
to the secretion of some fluid from the egg, which decreases in
diameter at the same time. Okkelberg describes a loss of 14 per
cent in the volume of the egg of the brook lamprey. The formation of this membrane with its perivitelline fluid underneath
marks the successful fertilization of the egg. For example, the
fertilized frog’s egg will rotate within this membrane.
The pronuclei. — After the second maturation division, which
does not take place in vertebrates until after the entrance of the
sperm, the nucleus of the ovum (female pronucleus) is near the
periphery at the animal pole, while the nucleus of the sperm
(male pronucleus) is at the periphery near the point of penetration. The sperm head rotates 180° so that the male pronucleus
now lies distal to the middle-piece containing the centrosome
(Fig. 21C). The two pronuclei come together (Fig. 21B, C) by
a route which may be analyzed into the following components:
(1) the sperm penetration path, which is usually the radius of the
egg at which the sperm entered; (2) the sperm copulation path,
which is directed towards the point at which the pronuclei will
meet and is often at a considerable angle to the penetration path;
(3) the egg copulation path, along which the female pronucleus
moves towards the meeting point; and (4) the cleavage path
(Fig. 21E), along which the two pronuclei move to their final
position on the egg axis, often slightly nearer the animal pole.
The two pronuclei may unite to form a common reticulum, or
they may remain close together contributing independently to
the first division of the zygote (Fig. 21F). See also page 156.
The centrosome of the egg disappears after the second maturation division. The centrosome of the zygote, therefore, is either
the centrosome of the sperm or, as it is believed in some cases, a
new one developed in the egg cytoplasm near the engulfed sperm
head.
The mitochondrial material of the sperm fragments and is distributed throughout the cytoplasm of the zygote. The later
history of the acrosome has not been followed.
There is much divergence among different kinds of animals
with respect to those parts of the sperm which are actually
engulfed in the egg.- In some, it is the entire sperm; in others,
only the sperm head.
PARTHENOGENESIS 55
Presumptive organ regions. — In many different kinds of eggs,
the student of cellular embryology has been able to recognize
different regions by differences in the cytoplasm, such as the
presence or absence of pigment, mitochondria, yolk, etc., and to
trace the distributions of these materials into the different
daughter cells as cleavage takes place. These presumptive organ
regions, as they may be called, are usually more easily demonstrated after fertilization. For example, before fertilization the
living egg of the tunicate Styela (Cynthia), according to Conklin
(1905), has orange pigment 6) Ground
granules uniformly distributed protoplasm
in its outer layer of cytoplasm. ais
During fertilization, following
an intricate series of stream
movements, the orange pigment
(Fig. 22) is concentrated in a
crescentic area at what will later
be the posterior surface. Immediately above this is a similar
area of clear protoplasm. On
the opposite side of the egg at @) Grey yolk
what will become the anterior ,,, 22, — Presumptive organ regions
surface is a gray crescent. Be- (organ-forming substances) in egg of
low these crescents the vegetal Styela after fertilization, viewed from
hemisphere is marked by the im) approx. X250. (After Conk:
presence of gray yolk. In later
cleavage, the yellow crescent will be distributed to the cells which
form the mesoderm, the gray crescent to the cells which form the
notochord and neural plate, the gray yolk to the cells of the endoderm, and the remainder of the egg goes to the cells of the
epidermis. The materials contained in the different presumptive
organ regions are frequently called organ-forming substances.
The regions themselves are called presumptive endoderm, presumptive mesoderm, etc.
Parthenogenesis. — This term is applied to the development
of a new individual from an unfertilized egg. It does not occur
naturally among the vertebrates but may be illustrated by the
honey bee, in which the unfertilized egg develops into a male
56 THE GERM CELLS
(drone) and the fertilized egg becomes a female (either queen or
worker according to the type of food supplied).
Artificial parthenogenesis has been produced in the frog’s egg
by slight punctures with a finely pointed glass needle. Most of
the parthenogenetic eggs do not go far in development, but Loeb
was able to raise, out of many thousands of treated eggs, a few
adult frogs (15 males, 3 females, 2 doubtful).
FERTILIZATION OF THE AMPHIOXUS EGG. — In the case of the
amphioxus, fertilization is external. The males and females leave
the sands to swarm in the shallow waters during late afternoons
of spring and summer. Eggs and sperms are discharged, from
the segmental gonads in which they develop, into the cavity
of the atrium, and escape to the exterior through the atriopore.
The first polocyte is given off before fertilization. Immediately
after fertilization the vitelline, now the fertilization, membrane
Anterior
™ Posterior
Presumptive
Presumptive
mesoderm
endoderm
Fig. 23. — Presumptive organ regions in egg of the amphioxus, one hour after fertilization. Sagittal section, approx. X220. (After Conklin 1932.)
expands greatly, leaving the egg in a large perivitelline space
(Figs. 23 and 10A). The second polocyte given off after the formation of the fertilization membrane remains attached to the
egg while the first is usually lost to view.
The fertilized egg of the amphioxus (Conklin 1932) shows in
sections (Fig. 23) a crescent of more deeply staining protoplasm
on the side of the egg which will give rise to the posterior part of
the body. This crescent will form the cells of the mesoderm
(compare the orange crescent of Styela). Opposite this is a less
clearly defined crescentic area from which the cells of the notochord and neural plate will be formed (compare the gray crescent
FERTILIZATION OF THE FROG’S EGG 57
of Styela and of the frog). The material of the vegetal hemisphere, bounded above by these crescents, will form the endoderm; the material of the animal hemisphere above the crescents
will form the epidermis.
FERTILIZATION OF THE FROG’S EGG. — The fertilization of the
frog’s egg is external, but the sperm are brought into close proximity to the eggs during the sexual embrace or amplexus. During
the breeding season the males embrace the females with the
fore-legs, at which time the germ cells of each are extruded. The
sperms make their way through the egg jelly before this envelope
Vitelline Polocytes
membrane
Jelly < Pigmented
hemisphere
S Gray
Sperm 4 crescent
Unpigmented
hemisphere
Fig. 24. — Diagram of frog’s egg after fertilization to show gray crescent. The line
immediately external to the vitelline membrane and polocytes represents the
“chorion.” X10.
has attained its final thickness. The entire sperm enters the egg
usually within 40° of the apical pole. The vitelline membrane
is thrown off as the fertilization membrane, leaving a perivitelline
space within which the egg may rotate. The second maturation
division then occurs, followed by the conjugation of the pronuclei.
The penetration and sperm copulation paths are marked by a
trail of pigment dragged in with the sperm head. A single sperm
enters the egg. Immediately upon fertilization the cortical cytoplasm of the egg rushes towards the point of penetration, carrying
with it the black pigment (melanin) of the animal hemisphere.
Upon the side of the egg opposite the point of penetration there
appears a crescent-shaped area in which the pigment is less dense
58 THE GERM CELLS
and which is therefore known as the gray crescent (Fig. 24).
The region gives rise to the notochord and neural plate.
FERTILIZATION OF THE HEN’S EGG. — In the fowl, fertilization
is internal. The sperms, introduced into the cloaca of the female during copulation, make their way to the upper end of the oviduct,
where fertilization takes place. Five or six sperms enter the
germinal disc, where they remain inactive until after the second
maturation division. One of them then moves inward until it
comes in contact with the female pronucleus, which has itself
moved downward from the surface of the germinal disc. The
supernumerary sperms move outward to the border of the disc,
where, after a few divisions, they degenerate. The fertilized egg
moves slowly down the oviduct while the tertiary envelopes are
forming about it.
FERTILIZATION OF THE HUMAN EGG. — Fertilization is internal
and occurs at the upper end of the oviduct (Fallopian tube). It
Fig. 25. — Fertilization of the guinea pig egg. Three stages following that shown
in Fig. 20A. (After Lams.)
is probable that a single sperm enters the egg after the first
maturation division. Further details are lacking, as no direct
observations have been r recorded. Figure 25 illustrates fertilization in the egg of the guinea pig.
SUMMARY
The gametes are atypical cells, the egg and sperm differing both
from each other and from a composite cell. The egg most resembles a composite cell, from which it differs in the absence of a
REFERENCES 59
centrosome after it has become mature. It is large, quiescent,
and protected by envelopes. The sperm, almost devoid of cytoplasm, is small, active, and naked.
The gametes are derived from primordial germ cells which are
first recognizable in the wall of the gut. Thence they migrate
into the roof of the coelom where they multiply rapidly giving rise
to the gonad. In the gonad multiplication continues until, when
the individual is attaining maturity, some of the ’gonia enlarge
to become ’cytes, each of which will undergo two meiotic divisions.
The spermatocyte gives rise to four spermatids each of which will
be transformed into asperm. The odcyte, on the contrary, gives
rise to an ovum and two or three polocytes.
The zygote, or fertilized egg, arises from the union of an egg
and a sperm. This union is preceded by the discharge of eggs
from the ovary (ovulation) and sperms from the testis (semination). The sperm, attracted to the egg, enters it due to the
mutual action of the two gametes, and the nuclei of the two
gametes come together, each to contribute to the first division of
the fertilized egg.
After fertilization, and sometimes even before, it can be seen
that the egg has a definite organization, manifest in its polarity
(seen even in ovarian eggs) and, in especially favorable material,
evident in presumptive organ regions.
REFERENCES
Allen, E. (ed.) 1932. Sex and Internal Secretions, Chap. 14.
Brachet, A. 1921. Traité d’embryologie, Book 1.
Cowdry, E. V. (ed.) 1924. General Cytology, Sections I, V-VIII.
Hertwig, O. (ed.) 1906. Handbuch, etc., I, Chaps. 1 and 2.
Jenkinson, J. W. 1913. Vertebrate Embryology, Chaps. 3 and 4.
Kellicott, W. E. 1913. General Embryology, Chaps. 2 to 5.
Kerr, J. G. 1919. Textbook of Embryology, I, Chap. 1.
Aorschelt, E., and Heider, K. 1902. Lehrbuch, etc. Chaps. 4 to 6.
Lillie, F.R. 1919. Problems of Fertilization.
Loeb, J. 1916. The Organism as a Whole from a Physicochemical Viewpoint.
Meisenheimer, J. 1921. Geschlecht und Geschlechter im Tierreiche.
organ, T. H. 1927. Experimental Embryology.
Wilson, E. B. 1925. The Cell, etc. Chaps. 1-6, 9-12.
Sharp, L. W. 1934. Introduction to Cytology, Chaps. 1-16.
CHAPTER IV
THE CHROMOSOMES AND THE GENES
The germ cells are really cells detached from the bodies of the
parents. When they unite in fertilization they bring together
material from both parents. Herein lies the explanation of the
inheritance of parental characteristics, of the fact that the fertilized egg develops in a way characteristic of the species and the
fact that individuals differ from one another. In the following
paragraphs we shall review the theory that the individual units
of heredity are the genes, borne in the chromosomes, distributed
in the maturation divisions, and brought together in fertilization.
A. THE CHROMOSOMES
It will be necessary first to describe the chromosomes as they
behave in ordinary (somatic) cell division, then to point out the
peculiar features of the maturation (meiotic) divisions and of
fertilization, and finally to indicate how this behavior of the
chromosomes fits the known laws of heredity.
The chromosomes in mitosis. — The division of most cells is
accompanied by the formation and longitudinal division of threads
of chromatin, called chromosomes, in the nucleus. This type of
cell division is known as mitosis (Fig. 26). Some cells, however,
divide without the formation of chromosomes (amitosis), and the
daughter cells are thereafter incapable of mitotic division. For
the sake of convenience we may use the terms karyokinesis for
the division of the nucleus in mitosis and- cytokinesis for the
division of the cytosome.
Karyokinesis. — Before cell division the metabolic (“‘ resting ’’)
nucleus is a reticulum of chromatin lying in the fluid karyolymph
with a nucleolus, the whole surrounded with a nuclear membrane
. (Fig. 26A). In mitosis we distinguish four stages, prophase,
metaphase, anaphase, and telophase.
In the prophase the reticulum separates into its constituent
threads, chromonemata, by the breaking down of the smaller
60
KARYOKINESIS 61
threads connecting them. Very early it can be seen that these
threads are double or split longitudinally (Fig. 26B). Soon
thereafter a matrix is visible about the two chromonemata.
This compound structure, consisting of the two chromonemata
T Kutul deb
Fia. 26. — Diagrams of somatic mitosis. A, metabolic (‘‘resting”’) stage. B, early
prophase showing chromonemata and attachment points. C, middle prophase,
matrix appearing. D, late prophase, chromonemata obscured. E, metaphase.
F, anaphase. G, early telophase, matrix disappearing. H, middle telophase,
nuclear membrane forming. I, late telophase, reticulum developing. (Based on
a diagram by Sharp.)
and the surrounding matrix, is a chromosome (Fig. 26C). The
number of chromosomes so formed is the same in every cell of
every individual belonging to any particular species. (This
statement is subject to exceptions. See pages 69 ff.) Towards
the end of the prophase the chromonemata are usually invisible.
62 THE CHROMOSOMES AND THE GENES
Finally the nuclear membrane disappears, and the karyolymph
assumes the form of a double cone or spindle (Fig. 26D).
In the metaphase (Fig. 26E), the chromosomes line up in an
equatorial plane through the spindle. Each has a definite attachment region lying in the equatorial plane even though the
ends of the chromosomes may lie outside of the plane.
In the anaphase (Fig. 26F), the chromosomes separate into two
longitudinal portions each containing one of the original chromonemata with surrounding matrix. Preceded always by its
attachment region each daughter chromosome moves towards a
pole of the spindle. Carothers (1934) describes the growth of a
fiber from the attachment region of each daughter chromosome
to the nearest pole of the spindle. Eventually two equivalent
sets of chromosomes are formed, one in the vicinity of either pole,
each set containing a daughter chromosome from each of the
original chromosomes formed in the prophase.
In the telophase (Fig. 26G, H, I), each set of chromosomes assumes the metabolic condition. The matrix loses its staining
capacity and the chromonemata reappear, often already split
longitudinally. The nuclear membrane is formed about each
group, the chromonemata are united by tiny cross-strands, the
nucleolus reappears, and the nucleus is seen to be filled with
karyolymph. The cell now contains two daughter nuclei each
identical with the other and with the parent nucleus.
Cytokinesis. — Other striking events are taking place in the
cytosome during mitosis. During the prophase the centrosome,
if not already divided, separates into two daughter centrosomes
which move apart. About each of them is a spherical mass of
protoplasm, often containing radial striations, known as the aster.
Between them is a central spindle apparently containing fibers.
Cytologists distinguish three types of fibers: (1) primary or continuous fibers extending from centrosome to centrosome, (2) half
spindle components extending from chromosome to centrosome,
and (3) interzonal connections extending between the separating
daughter chromosomes (Schrader). The centrosomes reach the
opposite sides of the nucleus just as the nuclear membrane disappears. The karyolymph apparently unites with the material
between the two centrosomes to form the mitotic spindle along
which the chromosomes move in the anaphase. In the telophase,
DISTRIBUTION OF THE CHROMOSOMES 63
asters and spindle disappear and the centrosomes alone remain in
the positions they occupied at the poles of the mitotic spindle.
Sometimes they divide in anticipation of the next mitosis.
The mitochondria usually divide en masse (Fig. 27A). This
division of the mitochondria is approximately an equal one, and
there is some reason to believe that the individual mitochondria
divide during mitosis or just prior to it.
The Golgi bodies, even when aggregated into a Golgi apparatus,
separate during mitosis and are segregated into the daughter cells,
usually associating themselves with the two centrosomes (Fig.
27B). It is uncertain whether each Golgi body divides individ
Mitochondria
Spindle
fibres
Golgi bodies
Fia. 27.— The mitochondria and Golgi bodies in mitosis. A, mitochondria.
B, Golgi bodies. (After Bowen.)
ually at mitosis, but some evidence has been brought forward to
support this contention.
In animal cells the cytosome as a whole divides by construction.
In this process the cell elongates in the direction of the spindle
during the anaphase and telophase. Following the reconstruction
of the daughter nuclei in the telophase, a furrow appears at the
periphery of the cell, around the equatorial belt, and at right
angles to the axis of elongation. This furrow advances towards
the center of the cell until the cell is completely divided.
Distribution of the chromosomes. — Each daughter cell has
approximately half of the cytoplasm proper, half of the mitochondria and Golgi bodies, a centrosome derived from that of the
64 THE CHROMOSOMES AND THE GENES
parent cell, and a nucleus built up from a set of chromosomes,
each of which was produced by the division of a chromosome in
the parent cell. It is apparent from the foregoing account that
the key to the complexities of mitosis is the division of the chromosomes. The achromatic figure is the framework upon which this
division takes place. The division of the mitochondria and Golgi
bodies is still too little understood. But the chromosomes, appearing in the prophase, halved with such accuracy in metaphase
and anaphase, and disappearing again in the telophase, are characterized by a constancy in number, an individuality evinced in
form and behavior, and a persistence from generation to generation. In some favorable material it has even been possible to
demonstrate that the chromonemata arise in the prophase exactly
as they merged into a reticulum in the previous telophase. From
the statements above, it is not unreasonable to draw the conclusion that the chromosomes are directly concerned with inheritance
in cell reproduction.
The chromosomes in meiosis. — During the two maturation
divisions by which the gametes are formed, the number of chromosomes is reduced to one-half the number characteristic of the
species. Since in the ordinary somatic mitosis the number of
chromosomes given to each daughter cell is exactly the same as
that of the parent, it is evident that we are dealing with a peculiar
type of mitosis (Fig. 28). The name meiosis is frequently applied
to the maturation divisions.
First meiotic division. — The essential feature in which the
first meiotic division differs from the ordinary (somatic) mitosis
is that during the prophase the chromosomes unite in pairs
(Fig. 29, 2). This is synapsis and occurs only in the first meiotic
division. Since each of the chromosomes always divides during
the prophase also (Fig. 29, 4), the net result is that at the end of
the prophase there are only half the number of chromosomes seen
in somatic mitosis, but each of these consists of four parts (chromatids) instead of two (Fig. 29, 5). These compound bodies consisting of four chromatids are called tetrads (Fig. 29, 6). The
quadripartite nature of the tetrad may be expressed by the
formula 42 Tn where A represents one of the synaptic mates and a
the other.
DIAGRAM COMPARING
SOMATIC MITOSIS WITH
THE MEIOTIC MITOSES
First columns Equational fo
division of a diploid somatic °
chromosome complement, \
Second column: The meiotic be ABe
divisions, changing the diplaid °
to the monoploid state.
Fig. 28. — Comparison of somatic and meiotic mitosis. (From Sharp.) (5)
66 THE CHROMOSOMES AND THE GENES
es
9
Fig. 29.— Diagram of meiosis. 1, first meiotic division, prophase, (leptonema
stage). 2, do. showing synapsis (zygonema stage). 3, do. showing thickening of
the chromosomes (pachynema stage). 4, do. showing forma tion of tetrads (diplonema stage). 6, do. showing condensation of matrix (diakinesis stage). 6, metaphase I. 7, anaphase I showing dyads. 8, telophase I. 9, second meiotic
division, prophase showing dyads united at attachment points. 10, metaphase IT.
11, anaphase II showing the separation of the chromatids which composed the
dyads. 12, telophase II. Each of the four germ cells now has the haploid number of chromatids (chromosomes). (From Sharp.)
DISTRIBUTION OF THE CHROMOSOMES 67
In the anaphase (Fig. 29, 7), the daughter chromosomes each
possess two chromatids and are known as dyads. But there are
two different ways of dividing a tetrad. In one case the two
chromatids derived from one of the synaptic mates (4) might be
separated from those derived from the other mate (2) in a reduc
tion (disjunction) division. In the other, each dyad might contain one chromatid from each of the synaptic mates (A : a) as the
result of an equation division.
The telophase (and prophase of the second meiotic division)
sometimes is omitted if the second division succeeds the first
immediately.
Second meiotic division. — If these omissions take place, each
of the daughter ’cytes divides immediately, the chromosomes,
still in the dyad condition, lining up on the spindles for the metaphase of the second meiotic division. But even if the telophase
of the first and prophase of the second meiotic divisions are not
omitted (Fig. 29, 8), it is obvious that the chromosomes arising in
the prophase (Fig. 29, 9) are dyads and that they undergo no
other longitudinal split. The anaphase of the second meiotic
division (Fig. 29, 11) merely separates the two chromatids of each
dyad from each other. The final result is that the four cells produced by the meiotic divisions (Fig. 29, 12) each have one chroma*
tid from each tetrad or one-half the number of chromosomes found
before meiosis took place. This is expressed in another way by
saying that the number of chromosomes has been reduced from
the diploid to the haploid (monoploid) number.
Here we must note that it makes no difference whether the first
meiotic division divided a tetrad reductionally or equationally.
The second division always distributes the two chromatids of each
dyad into different cells. Each of the four daughter cells has one
chromatid from each tetrad, and therefore one representative fro
either one of the two synaptic mates (A or a), but not from both!
Distribution of the chromosomes. — As each tetrad orients
itself independently upon the spindle it is evident that it is a
matter of chance which half of a tetrad, or of a dyad, goes to
either pole of the spindle. Accordingly, if we had eight chromosomes, A, a, B, b, C, c, D, and d, these would unite in synapsis to
68 THE CHROMOSOMES AND THE GENES
form four double chromosomes, Aa, Bb, Cc, and Dd. These
A:a B:6b C: D:d
would form the four tetrads, A: 7 Bb’ CO: s, and Ded’ Fol
lowing the two meiotic divisions (equation and reduction, regardless of their order), the mature germ cells would have four
chromosomes (the haploid number), but only one representative
of each synaptic pair. The possible combinations are 2‘ or 16,
namely, ABCD, ABCd, ABcd, Abcd, ABcD, AbCD, AbCd,
aBCD, aBCd, aBcd, abcd, aBcD, abcD, abCD, and abCd (Fig. 30).
Fia. 30. — Showing the distribution of the chromosomes in fertilization and the
following meiotic divisions. (After Wilson.)
Accordingly the number of different types of gametes which may
be formed can be determined from the formula 2” when n is the
haploid number of chromosomes characteristic of the species.
The chromosomes in fertilization. — Evidently when the egg
and sperm unite in fertilization, the pronucleus contributed by
each contains the haploid number of chromosomes. In this way ©
the diploid number characteristic of the species is restored. It is
obvious that, unless the number had been reduced by meiosis, it
would be doubled in each new generation.
In the second place, it is clear that each germ cell contributes a
homologous set of chromosomes, and that in synapsis the chromosomes unite in homologous pairs. In the example referred to
SEX CHROMOSOMES, X-O TYPE 69
above, chromosomes A, B, C’, D came from one parent and a, b, c,
d came from the other. We can now visualize each synaptic pair
as consisting of one paternal and one maternal chromosome.
During meiosis the paternal and maternal chromosomes are
sorted out into different assortments in the different germ cells.
During fertilization these different assortments are brought together in random recombinations. We have said that in an
animal with 8 chromosomes we might have 2‘ or 16 different
classes of gametes. In random fertilization this number would be
squared, so that there would be 4‘ or 256 possible combinations.
Many of these would be duplicates, so that the exact number of
different classes of zygotes according to their assortment of
chromosomes would be 34 or 81.
ants >
Ss eee @@
e
Fie. 31. — Chromosomes of Protenor. A, A’, male diploid group. B, B’, female
diploid group. The X-chromosomes are indicated by X. (After Wilson.)
Sex chromosomes, X-O type. — In many animals, such as the
insect Protenor, the male has one chromosome less than the
female, the numbers in Protenor being 13 and 14, respectively
(Fig. 31). If the synaptic pairs are assembled, it is clear that the
male has six pairs of ordinary chromosomes (autosomes) and an
extra one, the X-chromosome. The female has six pairs of autosomes and a pair of X-chromosomes. In the female the X70 THE CHROMOSOMES AND THE GENES
chromosomes unite in synapsis, form a tetrad, and are segregated
in the meiotic divisions so that every egg has a complete set of
autosomes and one X-chromosome (A + X). In the male, on
the other hand, the single X-chromosome has no synaptic mate
and so goes on the spindle of the first meiotic division as a dyad,
which is carried to one pole of the spindle entire. In the second
meiotic division the dyad is divided as usual. The end result is
that only half the spermatids receive an X-chromosome, and two
classes of sperms are formed, either with or without an X-chromosome (A + X or A +0). If a sperm with an X-chromosome
fertilizes the egg, the female combination (2A + 2X) is restored.
oO,
o°
OO \ a 2yg0te
Polocyte
Meiotic Sperm
Divisions
Meiotic
a 0 gO Divisions 8
- <—
($e) ‘ Oy, { ti rd
Spermatogonium
Ovum Odgonium
Sperm
r
CORO | D Zygot6e
Fig. 32. — Diagram showing history of the X-chromosome during meiosis and
fertilization. (After Wilson.)
If a sperm without the X-chromosome penetrates the egg, the
male combination (2A + X) is formed (Fig. 32).
Sex chromosomes, X-Y type. — But the sexes do not always
differ in chromosome number, for in many animals, like the insect
Lygaeus (Fig. 33), the X-chromosome of the male is furnished
with a synaptic mate which differs from it in size, form, and
probably composition, and is therefore known as the Y-chromo
.
some. The male forms a tetrad ae and the sperms therefore
have either an X-chromosome or a Y-chromosome. Fertilization
by a sperm bearing the X-chromosome results in the development of a female (2A + 2X), whereas if a sperm bearing a
CHROMOSOMES OF THE FROG 71
Y-chromosome enters the egg the embryo will give rise to a male
(2A + XY).
Sex chromosomes, W-Z type. — As an exception to the general
rule among the vertebrates, the birds have dissimilar sex chromosomes in the female. The cytological details are difficult to interpret but the theoretical explanation is that the female has two
dissimilar sex chromosomes known as W and Z, while the male
Core o¢¢ee eos
er eoceeee
Fig. 33. — Chromosomes of Lygaeus. A, A’, male diploid group. B, B’, female
diploid group. X and Y indicate the X- and Y-chromosomes respectively. (After
Wilson.)
possesses two similar sex chromosomes of the Z type (Fig. 34B).
In the meiosis of the odcyte, therefore, a tetrad wis is formed,
and the ovum receives either a W-chromosome or a Z-chromosome. The spermatocyte forms a tetrad a and all sperms
carry one Z-chromosome. In this group, therefore, it is the ovum
which determines the sex of the embryo rather than the sperm.
This explanation agrees with the data obtained from genetics.
CHROMOSOMES OF THE AMPHIOxuUS.— The diploid number
is 24,
CHROMOSOMES OF THE FROG.— The diploid number is 26,
and the sex chromosomes of the male are of the X-Y type
(Fig. 34A).
72 THE CHROMOSOMES AND THE GENES
CHROMOSOMES OF THE CHICK. — The diploid number is 35 or
36. The sex chromosomes have not been positively identified,
but genetic evidence indicates that the sex chromosomes of the
female are of the O-Z or*the W-Z type (Fig. 34B).
CHROMOSOMES OF MAN. — The diploid number, according to
the most recent researches, is 48. The sex chromosomes are
of the X-Y type (Fig. 34C). It is interesting to note that
with 48 chromosomes the possible types of oe number 274
Fig. 34. — Metaphase plates of male diploid chromosome groups. A, frog (after
Witschi). B, chick (after Hance). C, man (after Painter).
or 16,777,300, and that from these 3% zygote-recombinations are
possible.
B. THE GENES
It has already been said that the behavior of the chromosomes
itself might suggest that these bodies are concerned with the
transmission of hereditary characters. We shall now turn our
attention to the laws of heredity as worked out by plant and
animal breeders and learn how the data of genetics agree with
the data of cytology.
The unit of genetics is the gene. These genes are arranged in
linear order in the chromosomes, presumably bound together by
the chromonemata, and possibly identified with the chromomeres.
They exist in great numbers; in the fruitfly Drosophila it is
estimated that there are between 2000 and 3000. Ordinarily
ultramicroscopic, it has been reported recently by Belling (1930)
and by Bridges (1934) that they have been able to identify these
units in material of exceptionally favorable nature. The genes
are known by the effects their presence induces, and named according to the most obvious of these effects. Thus the Drosophila
SEGREGATION 73
has a gene for (or a gene which induces among other effects) the
normal type of wing. But there have arisen, among the millions
of fruitflies raised by geneticists, some with abnormal types of
wings, such as a vestigial wing. In thisease there is said to be a
gene for (or a gene which induces among other effects) the vestigial
type of wing.
Dominance. — Among the original discoveries of Mendel was
the fact that, if two organisms with alternative characters were
mated, the offspring would show either one or the other of the
characters concerned. This is known as the law of dominance.
When a Drosophila with normal wings is mated to one with
vestigial wings, all the offspring have the normal type of wing
(Fig. 35). Therefore the gene for normal is said to be dominant
to the gene for vestigial, which, conversely, is said to be recessive
to the gene for normal.
It is customary among geneticists to use the initial letter of the
name for the abnormal character as a symbol for the gene inducing
its appearance, as well as a symbol for the gene inducing the
alternative (allelomorphic) normal character. The two are distinguished by using a capital letter for the dominant gene, a
lower-case letter for the recessive gene. In this case, then, the
symbol of the gene for vestigial is v, and the symbol of the gene
for normal is V.
Every adult has two haploid sets of chromosomes, and therefore
a pair of every kind of chromosome. If both members of a pair
have the same gene (w or VV) they are said to be homozygous;
but if one chromosome has the dominant, and the other has the
recessive gene (Vv), they are said to be heterozygous.
The individuals that are mated together in-the first instance
are known as the parental generation (P,); their offspring are
known as the first filial generation (F1); the next generation is
the second filial generation (Ff); and so on.
Segregation. — In the experiment where a normal long-winged
fly was mated with a vestigial-winged fly, the long-winged parent
must have had two chromosomes each containing the dominant
gene V, for all the offspring (F'1) showed this dominant character.
The vestigial-winged parent must have had two chromosomes
containing the recessive gene v. In the maturation of the gametes
all the sperms received V, while the eggs all received »,
74 THI CHROMOSOMES AND THE GENES
Fia. 35. — Diagram to show the effects of crossing two flies differing in respect to one
pair of genes. V is used for the dominant gene for the normal character long
wings; v is used for the recessive mutant gene for vestigial wing. (From Curtis
and Guthrie, after Morgan et al.)
The F, flies have the genetic constitution Vv, that is to say,
they are heterozygous. When they are mated to each other the
eggs will receive a chromosome containing the gene V or the gene
v, and the same is true of the sperm.
ASSORTMENT 75
The F, flies will consist of three genetic groups (genotypes)
because of random fertilization, namely, homozygous long-winged
flies (VV), heterozygous long-winged flies (Vv), and vestigialwinged flies (wv). From Fig. 36, it will be seen that the ratio
will be one homozygous long-winged fly to two heterozygous
long-winged flies, and to one vestigial-winged fly. Or one may
say that there are two recognizable classes of adults (phenotypes),
in the ratio of three long-winged flies to one vestigial-winged fly.
This is the famous Mendelian ratio applied to the inheritance of
&
\
Cag
= (68)
Fia. 36. — Diagram to show the segregation of the genes caused by the distribution
of the chromosomes to the gametes and zygotes of the Fi and F; generations.
Y, and v as before.
Gametes as
one pair of allelomorphic characters, or as we should say today,
to one pair of genes.
Evidently Mendel’s law of segregation may be stated in terms
of the gene theory as follows: allelomorphic genes are segregated
during maturation into different gametes.
Assortment. — It is an amazing coincidence that Mendel studied the inheritance of seven pairs of allelomorphic characters in
76 THE CHROMOSOMES AND THE GENES
the edible pea, a species which has seven pairs of chromosomes,
and that the genes for each pair of characters were located in a
different pair of chromosomes.
When a Drosophila with vestigial wings and normal gray body
color is mated to a fly with normal long wings and ebony body
color, the F, flies are gray-bodied and long-winged. Evidently
the gene for gray body (£) is dominant to the gene for ebony
body (e). That the genes for these characters are independent
of those affecting wing length is shown when the hybrid F, flies
are mated together. Four classes of phenotypes result in the F,
generation: 9 long-winged, gray-bodied; 3 long-winged, ebonybodied; 3 vestigial-winged, gray-bodied; and 1 vestigial-winged,
ebony-bodied. This ratio of 9: 3:3 :1 breaks down to 3 long
to 1 vestigial, and 3 gray to 1 ebony, demonstrating mathematically that two pairs of factors are involved.
It is evident that the problem involves the segregation of two
pairs of chromosomes (Fig.37). The genetic constitutions of the P;
flies were wHE and V Vee, respectively. The gametes receive one
chromosome from each pair of synaptic mates, so the genetic constitution of the eggs is v# and that of the sperms Ve (or vice versa).
The F;, flies have the formula VvKe, and their gametes, because
the chromosome pairs are assorted independently, will belong to
four classes: VE, Ve, vE, and ve.
The F, flies as seen from the checkerboard diagram will fall into
16 combinations, which by canceling the duplicates reduce to 9
genotypes (VVEE, VVEe, VvEE, VuEe, VVee, Vee, wKE,
vvEe, and vvee), and 4 phenotypes as listed above.
Mendel’s law of assortment may be phrased in terms of gene
theory as follows: different pairs of allelomorphic genes when
located in different pairs of chromosomes are assorted independently
during maturation into different gametes.
It may be noted that if n stands for the number of pairs of
genes located in different pairs of chromosomes, then 2” represents
the number of gamete classes formed by the F: generation; 2"
represents the number of phenotypes in the F, generation; 3” the
number of genotypes in the F, generation; and 4" the number of
combinations in the Punnett square. The number of individuals
in each phenotype is obtained by expanding the 3 : 1 formula as
follows: (8:1), (9:3:3:1), (27:9:9:9:3:3:3:1)....
ASSORTMENT 77
Zygotes from
which P,
developed
Gametes of P,
to long-winged,
gray~bodied flies
©) Zygotes of F,
that give rise
a
C<_)
axe |(HB)|(HO)| (HH) | (HO
VVEB VVEe VvEBD VvEe Zygotes of F,
long long long long
gray . gray gray gray
9 long-winged,
VVEe VVee VvEe Vvee one veth ed
©) long long long long gray"
bo: :
gray ebony gray ebony 8 long-winged,
bony-bodi
VvEE | vVvEe wEE vvEe ebony-bodied
long long vestigial vestigial 8 vestigial-winged,
gray gray gray gray gray-bodied
VvEe ‘Vvee vvEe vvee 1 vestigial-winged,
ly long long vestigial vestigial ebony-bodied
gray ebony gray ebony
Fig. 37. — Diagram to show the assortment of two pairs of genes due to the distribution of two pairs of chromosomes. E, gene for gray body; e for ebony body;
V, and v as before. (From Curtis and Guthrie.) .
78 THE CHROMOSOMES AND THE GENES
Linkage. — The characters with which Mendel worked segregated freely, showing that their genes were not borne in the same
chromosome. Later studies have shown that some characters do
not segregate, and this leads to the assumption that their genes
are carried in the same chromosome and therefore are inherited
together.
When a Drosophila with gray body color and long wings is
mated to one with black body color and vestigial wings, the F1
flies are gray-bodied and long-winged. Note that the gene for
black (6) will act very differently from the gene for the similar
color ebony (e). If the F; flies are bred together a very confusing
ratio appears in the F, generation: practically all the flies are
gray-bodied and long-winged or black-bodied and vestigial-winged
like the P, generation, but there are only a few individuals representing the other classes we might expect under Mendel’s law of
assortment. If we make a reciprocal cross between a long-winged
black-bodied fly and a vestigial-winged gray-bodied fly, the F,
flies are all of these two (P;) types with few exceptions. This
continued association of two genes through several generations is
called linkage and suggests that the associated genes are located
in the same chromosome.
This theory may be tested by back-crossing (Fig. 38) a male of
the F; generation (BbVv) to a double recessive female (bbw).
All her eggs will have the recessive genes (bv). We can then test
the constitution of the sperm by examining the progeny of this
cross (here called F, for convenience), for all the F, flies must have
the genes (bv) from the mother. The flies of this generation are
either gray-bodied and long-winged (BbVv) or black-bodied and
vestigial-winged (bbvv). This seems to show that the genes B and
V were located in one chromosome while b and v were located in
the synaptic mate.
Crossing over. — Now for the exceptional (cross-over) flies
noted above. There is no crossing over in the maturation of
the male F, fly, but how about the female? When we mate
(Fig. 39) a female F, fly (BbVv) to a double recessive male (bbvv),
the progeny (F,) fall into four classes: 41} per cent gray-bodied
and long-winged (BbVv), 413 per cent black-bodied and vestigialwinged (bbvv), 84 per cent gray-bodied and vestigial-winged
(Bbvv), and 83 per cent black-bodied and long-winged (bbV»).
CROSSING OVER 79
Obviously there has been an exchange of some sort between the
chromosomes of the female F, fly. Both cytological and experi
Fig. 38. — Diagram to show the inheritance of two pairs of genes when located in
one pair of chromosomes, (linkage). In this case the male /; fly is back-crossed
to a double recessive female. B, gene for gray body; }, for black body; V, and v
as before. (After Morgan.)
mental evidence seem to indicate that this crossing over takes
place in the prophase of the first meiotic division (Fig. 29, 4).
Although there are still difficulties in determining exactly how
the crossing over takes place between the four strands, it is gener80 THE CHROMOSOMES AND THE GENES
ally agreed that the actual crossing over takes place between two
of them. The idea of linkage between genes in the same chromo
41.5 85 85 41.5
Fig. 39. — Diagram to show the inheritance of two pairs of genes when located in
one pair of chromosomes between which crossing over takes place. In this case
the female F; fly is back-crossed to a double recessive male. Symbols as in Fig.
38. The figures at the bottom of the illustration indicate the percentage of each
phenotype in the entire hatch. (After Morgan.)
some suggested the idea that the genes form a longitudinal series
in each chromosome. This is supported by the behavior of the
SEX-LINKED INHERITANCE 81
chromosomes in ordinary somatic mitosis, in synapsis, and in
crossing over (Fig. 40).
Finally Sturtevant (1913) suggested that the percentage of
crossing over between two pairs of linked genes might represent
a function of the distance between the loci of the genes in
the chromosome. Accordingly, maps have been constructed, by ©
Morgan and his co-workers, on the general assumption that one
per cent of cross-overs is represented on the map by a distance of
one unit between the genes involved. Without going into further
details of the methods used in constructing these maps, for there
A B C
_—" c i ~ c ~*~ —~
Fig. 40. — Diagram to illustrate: A, splitting of a chromosome in somatic mitosis;
B, union of two chromosomes in synapsis; C, union of two chromosomes in synapsis accompanied by crossing over. (After Wilson.)
are many complicating factors, a glance at the accompanying
chart (Fig. 41) will show the progress that has been made in this
direction.
Sex-linked inheritance. — One of the most striking evidences
that genes are borne in the chromosomes is afforded by what is
known as sex-linked (criss-cross) inheritance. This is illustrated
in Drosophila by the inheritance of white eye color, an allelomorph
of red, the normal eye color. If a white-eyed male is mated with
a red-eyed female (Fig. 42), the F, flies of both sexes will have red
eyes. But if these F, flies are bred together the F, generation
will be made up of red-eyed females, (50 per cent) red-eyed males
(25 per cent), and white-eyed males (25 per cent). It looks at
first like an ordinary 3 to 1 Mendelian ratio, except for this
curious distribution of eye color in the two sexes.
100) a m
3 eltow (B) 7 0 telegraph(W) 7 0. roughoid (E)
¥ Wes airy win 2. Star (E) |
t ‘ vhs Pethet cay T 34 aristaless (8)
F So broad (W) + 6.4 expanded (Ww)
+ |. prune Ce) :
fT. (8 white(E)
Ve facet te) 4 12+ Gull WW)
\ 134 Notch CE Yt Truncate (W)
45 Abnormal(B) + !4.t dachsous (B)
T.\55 echinus(E) +416. Streak ()
+\ 169 bifid (W)
Ty 75 ruby CE)
SB eres 0)
Tt clu + 1
+ N72 Geltex cw) 20. divergent (w)
20. cut (W)
21. singed (H) aot @)
275 tan (B) + 3. achs (8 4 26. sepia CE
F 217 lozenge (E) yT 265 hairy &
T 35. Ski- I (w)
T 33. vermillion(E)
361 miniature(w) +41. Jammed(W) 4 35. rose (CE)
+ 362 dusky (W) + 362 cream-M (E)
J 38% furrowed (E) 1464 Minute-e (H) 401 Minute-h (H)
148.5 black (B) $ 402 tilt (W)
+ 43. sable @) 48.1 jaunty (Ww) 404 Dichaete (H)
+ 444 garnet (E) 422 thread (8
_T 546 purple (Ee) + 44. scarlet ()
5 4575 cinnabar(E)~. 48. pink (E)
4.2 small win . 44
+ 545 redmentangtW) 02 safranin(E) Ff, sn maroon (e)
i 366 forked a BO. curled (W)
F 57. Ba . . L
¥ be srall ° 7 O44 pinkwingw) T B48 Bairy win supr
r 59. fuse F 58.5 spineless(H)
4d 596 +67. vestigial(W) = P
62. Beadex CW) + 68+ telescope W) F..587 bithorax (B)
| “59.5 bithorax-b
+ 65. cleft (W) die Lobe Ce) tT. 62. stripe (B)
- hope | 63.1 glass C)
| 474% gap Ww) 662 Delta WW)
a> +4 10. bobbed(H) +755 Curved (w) 4+ 695 hairless CH)
+ 107 ebon
F 337 peony
4d ‘757 cardinal (E)
4835 fringed) T 76.2 white ocelli )
—>!
+90.
x \ “ 9 humpy (8)
+ 914 rough @)
v 4+ 995 arc (W) 4 93. crumpled Ww)
+ 1005 lexus Ww) + 938 Beaded Ww.
T 102+ fethal-ta 94.1 Painted (W)
I 05. brown (E
mW 05+ blistered(w) | 1007 glare ©)
« 4 106. purpleoid, {e) F101. Minute CH)
gid Teeckt)
spec
F075 balloon ew) + 106.2 Minute-g (H)
Vv
bent (W)
shaven (B)
eyeless (E)
rotated (8)
Minute-I¥ (H)
male fertility
“Long bristled
male fertility’
|
Fia. 41.— The chromosomes of Drosophila melanogaster and map showing the
positions of many genes as determined from cross over ratios.
theses indicate part of body affected: B, body; E, eye; H, hair; W, wing: -’ Arrows
indicate position of attachment point.
exact position of genes in Y still undetermined.
1925 and Stern 1929.)
Letters in paren
All genes in IV are closely linked. The
(From Sharp after Morgan et al.
(82)
SEX-LINKED INHERITANCE 83
In the reciprocal mating, a red-eyed male to a white-eyed
female (Fig. 43), the F: generation is made up of red-eyed females
Fig. 42. — Diagram to show the inheritance of one pair of genes when located in the
X-chromosome (sex-linkage). W, gene for red eye (dominant, normal); w, gene
for white eye (recessive, mutant). The empty hook-shaped chromosome represents the Y-chromosome. N. B. In the text the X-chromosome, when bearing
w, the gene for white eyes, is designated by a small x. In this cross, red-eyed
female is mated to a white-eyed male. (After Morgan et al.)
and white-eyed males (criss-cross inheritance). When these F,
flies are bred together, there are four classes of flies in the F,
generation: red-eyed males and females, and white-eyed males
84 THE CHROMOSOMES AND THE GENES
and females (25 per cent in each class). This is not a Mendelian
ratio, but it can be explained on the assumption that the gene for
Ina. 43. — The reciprocal cross to that shown in Fig. 42. A white-eyed female is
mated to a red-eyed male. Symbols as in Fig. 42. (After Morgan et al.)
white eye color (and its allelomorphs, of which there are several)
is located in the X-chromosome.
Let us use the symbol X for an X-chromosome bearing a
gene for red, x for an X-chromosome bearing a gene for white,
and Y for the Y-chromosome. In the first genetic experiment,
formulas for the parental generation are X X (red female) and
x Y (white male). All the eggs receive an X, the sperms either
SEX-LINKED INHERITANCE 85
zor Y. Consequently the Ff: generation is made up of flies with
the formula Xx (heterozygous red female) and XY (red male).
The eggs of this generation receive X or x, the sperms X or Y.
A
Two pairs of homologous chromosomes
showing positions of allelomorphic genes.
uae Re
Sab
a
Crossing over: The chromosomes of the
pair shown in A may twist about one another
as in C and break in the plane of the dotted
line so that comparable sections are exchanged as shown in D.
ac wR
bank
oe Ob
bath
RQ
Deletion: One member of the chromosome
pair shown in A may twist on itself as in K
and break in the plane of the dotted line so e
that an internal section containing gene c is
lost, or deleted, as shown in F.
~
bab >
ae
9 Qa 86 >
Inversion: One member of the chromosome
pair shown in A may twist on itself as in G
and break in the plane of the dotted line so
that the section containing genes B and C is
inverted as shown in H.
bs &Q >
a 8° &@ &
Duplication and Deficiency: If one member
of the chromosome pair shown in A comes to
lie across the other as shown in I and a break
occurs in the plane of the dotted line, the
chromosome on the left in J will have a
duplication and contain both gene d and
gen D, while the chromosome on the right
will have a deficiency of the section contain- I J
ing gene D.
tea a @&
Q
Translocation: One member of the chromosome pair shown in A may come to lie
across one member of the chromosome pair
shown in B, as seenin K. If a break occurs
in the plane of the dotted line, sectiotis of
non-homologous chromosomes are exchanged,
or translocated, as shown in L. K L
wy o>
aaecs
an0° 6 8
Fia. 44.— Diagrams to show crossing over and various chromosomal aberrations.
(From Curtis and Guthrie.)
86 THE CHROMOSOMES AND THE GENES
So the F, generation is composed of flies with the following combinations: XX (homozygous red females), Xx (heterozygous red
females), XY (red males), and xY (white males).
In the other experiment the parental formulas are xx (white
female) and XY (red male). The eggs receive an x, the sperm X
or Y. Hence there are two classes in the F; generation, xX
(heterozygous red females) and xY (white males). The eggs receive either an x or an X, the sperms receive either zor Y. The
four combinations possible in the F, generation are xX (heterozygous red female), xx (white female), xY (white male), and XY
(red male).
Chromosomal aberrations. — Crossing over takes place between
the two X-chromosomes, but apparently not between the X-chro
Oocytes
) ) (44) Disjunctional
Division
Ovum Body Ovum Body Ovum Body
oe N Male
Sperm ormal Female &) (X from Father) Zygotes
Normal Male Female Dies
(both X’s from Mother)
Fia. 45. — Diagrams showing I, normal disjunction of X-chromosomes in odgenesis,
and fertilization by two types of sperms; II, non-disjunction, both X-chromosomes
remaining in egg; III, non-disjunction, both X-chromosomes passing to polocyte.
A, one haploid set of autosomes. (From Curtis and Guthrie.)
mosome and the Y-chromosome in Drosophila. We have already
noted the fact that in this little fruitfly crossing over does not
take place in the male.!' But crossing over by no means exhausts
the possibility of effecting new combinations of genes by the behavior of the chromosomes during the maturation of the germ
cells. Exact as the mechanism of meiosis may seem, many possi
1 There is some recent evidence to show that such crossing over can be induced
by high temperatures.
NON-DISJUNCTION 87
bilities of disturbance have been discovered by genetic and cytological methods.
The accompanying diagram (Fig. 44) illustrates graphically
some of the aberrations which may take place during meiosis.
These result in the appearance of unexpected individuals with
new combinations of genes ,
.
ne
a
ARN
A
t
‘\,
PRLS
poe
ry
Fia. 46. — Intersexes and supersexes in Drosophila, occurring in the progeny triploid
females. A, female-type intersex. B, male-type intersex. C, superfemale.
D, supermale. a, b, and c are the chromosome groups characteristic of A, B,
and C respectively. ‘ (From Curtis and Guthrie, after Morgan et al.)
Non-disjunction. — A special type of chromosomal aberration
is one in which the two members of the synaptic pair may fail to
separate during the meiotic divisions, so that one egg receives, for
example, two X-chromosomes (A + 2X), while another receives
none (A) (Fig. 45). When fertilized by a sperm with an Xchromosome, the egg with two X-chromosomes, if it develops
into an adult, will be a superfemale (2A + 3X) differing markedly
from her sisters (Fig. 46C). When fertilized by a sperm with a
88 THE CHROMOSOMES AND THE GENES
Y-chromosome, the egg without any X-chromosomes (2A + Y)
dies. The other possible combinations are shown in the diagram.
In some cases all the chromosomes fail to disjoin so that an egg
receives a diploid set of chromosomes (2A + 2X). When fertilized by an A + X sperm it becomes a triploid female (8A + 3X).
The eggs formed by these triploid females may have the formula
2A + XorA+2X. If an egg of the first type (2A + X) is
fertilized by sperm carrying an X-chromosome (A + X), the
zygote will have the formula 3A + 2X. Such a zygote develops
into an abnormal fly known as an intersex (Fig. 46A), male in
some respects and female in others. Superfemales (2A + 3X)
may also arise from the egg of the second type (A + 2X) being
fertilized by an A + X sperm. Supermales (3A + XY), on the
other hand, arise from the fertilization of a 2A + X egg by an
A + Y sperm (Fig. 46D). It would appear from these formulas
as though the determination of sex depended on some sort of ratio
between the genes in the X-chromosomes and the autosomes, and
Bridges (1921) has formulated a theory of
genic balance to account for the observed
results.
Gynandromorphs. — Intersexes must not
be confused with gynandromorphs, which
are individuals with one part of the body
male and the rest female. Bilateral gynandromorphs in Drosophila (Fig. 47) arise
from female zygotes (2A + 2X), but during
nat the first cleavage division one of the XFig. 47. —Gynandromorph_ ghromosomes is lost on the mitotic spindle.
in Drosophila melanogas- :
ter. Note eosin eye and Lhe result is that one of the daughter cells
miniature wing on right has the female complex (2A + 2X) while
as compared to red eye the other has the male complex (2A + X).
and long wing on left. . °
This fly is male on the S0Metimes such an aberration takes place
right side and female on in a later cleavage division so that there
the left. (After Morgan ig only a small area of male cells.
and Bridges.) Teratology. — All students of embryology are familiar with the fact that development does not always
proceed normally. Abnormal embryos are known as monsters,
and their study forms the subject matter of the embryological
subscience known as teratology. It is clear from the sections
SUMMARY 89
just preceding that many of these monsters must be due to
chromosomal aberrations with consequent disturbance of the
genic balance. Others, as will be noted in Chapter VII, are due
to environmental factors.
Mutations. — So far we have considered the genes as though
they were immutable. But the question naturally arises as to
the origin of the genes which are allelomorphic to the so-called
normal genes. In Drosophila the abnormal genes, or mutants as
they are called, arose in laboratory cultures. It has been discovered that the rate of mutation, i.e., the number of mutants
arising in a given number of flies, may be increased by high temperatures (Plough) and by irradiation (Miller). When one of
these genes is altered in any way to become a mutant, the course
of development is disturbed. Most mutant genes disturb the
course of development so greatly as to cause death (lethal
mutants). A smaller number produce visible changes when
present in each chromosome of the synaptic mates (recessive
mutants). A few produce visible changes if contained in a single
chromosome (dominant mutants). Accordingly, every species
of animals contains a certain number of mutant genes (400 in
Drosophila). As these enter into new genetic combinations according to the behavior of the chromosomes in meiosis and fertilization, they give rise to individual differences in development.
But the greater number of stable or non-mutant genes holds
development true to the specific type.
One of the outstanding problems in experimental embryology
still awaiting solution is the question how the genes actually
determine the course of development. But the modern student
of embryology accepts the general theory that it is the complement of genes, from the egg and sperm respectively, which
initiates and largely controls the development of the individual.
SUMMARY
The egg and sperm are the material contributions of the parents
to the new individual. The equivalent structures of the egg and
the sperm are their nuclei. Each nucleus contains the haploid
number of chromosomes. The fertilized egg has two haploid
sets, or the diploid number. In somatic mitosis the chromosomes
are split longitudinally and divided equally among the daughter
90 THE CHROMOSOMES AND THE GENES
cells, so that each daughter cell contains an assortment precisely
equivalent to that of its sister cell and the mother cell. In the
course of the meiotic divisions the diploid number of chromosomes
is reduced to one haploid set. This is accomplished through the
union of the homologous members of the two sets in synapsis.
Each synaptic pair forms a tetrad of four chromatids, the members
of which are distributed independently among the mature germ
cells. In this way different classes of gametes are formed with
varying chromosomal complexes.
The chromosome is built up from a thin thread, the chromonema, which binds together the genes, the units of heredity,
provisionally located at nodes of the chromonema called chromomeres. These genes, ordinarily ultra-microscopic, are self-reproducing units which seem to accelerate definite chemical reactions
without losing any of their own substance in the process. The
course of development is largely controlled by the activities of
these genes. These activities may be disturbed during meiosis
by chromosomal aberrations, thus altering the genic balance and
modifying the course of development, in some cases so much as to
cause death. The genic balance may also be altered by point
mutations or changes in the constitution of an individual gene
recognizable through the effects produced.
Either aberrations or point mutations when not lethal may be
transmitted in heredity. The distribution of these aberrant
chromosomes or mutant genes in meiosis and fertilization is the
material basis for heritable differences arising in the course of
development of individuals belonging to the same species.
REFERENCES
Cowdry, E. V. (ed.) 1924. General Cytology, Sections X, XI.
Morgan, T. H. 1913. Heredity and Sex.
—— 1919. The Physical Basis of Heredity.
“—— 1922. The Mechanism of Mendelian Inheritance, 2nd Ed.
A— 1934. Embryology and Genetics.
fA and others. 1928. The Theory of the Gene.
Sharp, L. W. 1934. Introduction to Cytology, Chaps. 17-24,
Wilson, E. B. 1925. The Cell, etc., Chaps. 9-12.
CHAPTER V
CLEAVAGE AND THE GERM LAYERS
The fertilized egg (zygote) is a complete and balanced cell.
It has two entire sets of chromosomes, each with a full comple
“ment of genes, one set from each parent. These nuclear elements
are contained in a cell body whose cytoplasm is principally
maternal in origin and which has a definite organization as indicated by its polarity. We are now to examine the way in
which the embryo develops from the fertilized egg.
It is customary to distinguish three steps in the early development of the embryo. First is the period of cleavage in which the
egg undergoes a number of mitotic cell divisions at each of which
the number of cells (blastomeres) increases while the size of the
cells decreases. The period ends with the embryo in the form of
a blastula, a sphere or disc in which the blastomeres are not
stratified into different layers.
Second comes the period of gastrulation in which the blastomeres arrange themselves into an outer and inner layer of cells,
known as ectoderm and endoderm, respectively. This twolayered embryo is called a gastrula.
’ Third is the period in which a middle layer, including the mesoderm and the notochord, is formed between the ectoderm and
endoderm. Although this layer sometimes develops during gastrulation, it is customary to distinguish a period of mesoderm
(chorda-mesoderm) formation. This distinction is not always
valid, nor is it important, for, as will be seen, the material which
is to form the middle germ layer can sometimes be distinguished
in gastrulation, cleavage, or even in the fertilized egg.
A. CLEAVAGE
As there are different types of eggs according to the amount
and distribution of the yolk, so there are different types of
cleavage according to the pattern formed by the dividing egg.
Rules of cleavage. — Certain rules have been formulated to
express the simpler geometrical relationships of the blastomeres.
91
- 92 CLEAVAGE AND THE GERM LAYERS
The first are those of Sachs: (1) cells typically tend to divide into
equal parts; (2) each new plane of division tends to intersect the
preceding one at right angles. Sachs’s rules are supplemented,
and to some extent explained, by those of Hertwig: (1) the
typical position of the nucleus (and hence of the mitotic figure)
tends towards the center of its sphere of influence, i.e., of the protoplasmic mass in which it lies; (2) the axis of the spindle typically lies in the longest axis of the protoplasmic mass, and division
therefore tends to cut this axis transversely.
Methods of cleavage. — The rate of division is governed by
the rule of Balfour: the rate of cleavage is inversely proportional
to the amount of yolk present. This leads to a distinction between two types of cleavage. In the first type the cleavage
A B Cc
Fia. 48. — Diagram to show main types of cleavage in vertebrates. A, equal holoblastic. B, unequal holoblastic. C. meroblastic.
planes divide the egg completely into separate blastomeres.
This is known as holoblastic cleavage, and is characteristic of
isolecithal and moderately telolecithal eggs. In the second type
the cleavage planes do not pass through the yolk and so the
separate blastomeres come to lie upon a mass of undivided yolk.
This is known as meroblastic cleavage and is typical of extremely
telolecithal eggs. It is generally true that isolecithal eggs have
equal holoblastic cleavage (Fig. 48A). Moderately telolecithal
eggs have unequal holoblastic cleavage (Fig. 48B), and extremely
telolecithal eggs have meroblastic cleavage (Fig. 48C).
Cell lineage. — It must not be thought that cleavage results in
a mass of identical blastomeres. Painstaking examination of
dividing eggs has shown that in the normal development of
favorable material the origin and fate of every blastomere can
be determined accurately. The genealogical history of the blastoCELL LINEAGE 93
meres is known appropriately as cell lineage. One of the most
clean-cut examples, in forms allied to the vertebrates, is the cell
lineage of the tunicate Styela (Cynthia), worked out by Conklin
in 1905. The accompanying diagram (Table 6) shows the cell
lineage up to the 32-cell stage with the ultimate fate of each of
the blastomeres.
In reading this chart the student should understand the system
used in naming the blastomeres, which is illustrated most easily
Fia. 49. — Cleavage of Styela (Cynthia) egg. A, 4-cell stage from left side. B, same
stage from animal pole. C, 8-cell stage from left side. D, same stage from animal
pole. For explanation of lettering see text. (Irom Richards, after Conklin.)
by means of the 8-cell stage (Fig. 49). The blastomeres which
will give rise to structures on the right side of the embryo are
underlined. The blastomeres formed at the animal hemisphere
are in lower-case letters; those at the vegetal hemisphere are in
capital letters. Those formed at the antero-dorsal side of the
embryo are given the designation A or a; those at the posteroventral side are named Borb. The first exponent is the number
of the cell generation, counting the fertilized egg as the first
94 CLEAVAGE AND THE GERM LAYERS
generation, the blastomeres of the first cleavage as the second
generation, etc. The exponent after the decimal point indicates
whether the cell is in the first, second, third, etc., row from the
vegetal pole. Thus the cell labelled A‘! is antero-dorsal, left
side, vegetal hemisphere, of the fourth generation, and in the row
next to the vegetal pole.
TABLE 6
Ceuvt Lineage or Styela (Cynthia) arteR Conknin (1905)
(Ist) (2nd) (8rd) (4th) (5th) (6th) Generation
(1) (2) (4) (8) (16) (32) Number of cells
5 a®8 Tctoderm (epidermis)
a ao7 “ “
4.2
a 98-6 “ “
5.3
a5 “ (neural plate)
A®4 Chorda-neural plate
A&3 _Endoderm
A®2 Chorda-neural plate
A®! Hindoderm
b&8 Ectoderm
b®7 “
bé-6 “cc;
bé5 “cs
B&+ Mesoderm (gray crescent)
Be3 “cc 6c “cc
Be “ (yellow “ )
Be! Endoderm
|
|
| _
Eaa | ats Ectoderm (epidermis)
|
|
Ab.2
Abt
AB?
(Left) ( ps4
b®3
B3
Bs2
Be
5.4
a g8-7
4.2
a 6“ ‘6
353 ass
= ass “ (neural plate)
A®4 Chorda-neural plate
A®3 Endoderm
A’? Chorda-neural plate
A®! Endoderm
b*8 Ectoderm
b&7 “
bes “
bes “
Bé+ Mesoderm (gray crescent)
Bes ‘“ 6 ‘“
Be “ (yellow “ )
Be! Endoderm
A52
Atl ~
~~ Aba
AB! ~
b*2
b53
Bs
Bt
B51
The first cleavage is bilateral; i.e., it divides the egg, with its
presumptive organ regions, into a right blastomere (AB?) and a
left blastomere (AB’). At the second cleavage each of these is
CLEAVAGE: THE AMPHIOXUS 95
divided into an antero-dorsal blastomere (A? and A‘) and a
postero-ventral blastomere (B? and B*). The third cleavage
plane (Fig. 49C, D) separates the smaller cells of the animal hemisphere (a*?, b*”, a*?, b*?) from the larger cells of the vegetal hemisphere (A4}, Bt, Atl, B*?).
By the sixth generation (32-cell stage) the organ-forming regions have been segregated into different blastomeres as follows:
Animal hemisphere:
14 Ectoderm, epidermis.
2 Ectoderm, neural plate.
Vegetal hemisphere:
4 Ictoderm and mesoderm, chorda-neural plate.
4 Mesoderm, gray crescent.
2 Mesoderm, yellow crescent.
6 Endoderm cells.
32
The cell lineage of many types of invertebrates has been investigated in a similar manner, and as a result it is now generally
recognized that during cleavage the successive generations of
blastomeres show a progressive differentiation. Larlier or later,
the presumptive organ regions of the fertilized egg are segregated
into different groups of blastomeres, each group forming a presumptive organ region of the blastula (page 102).
Later (Chapter VII), experiments will be described which indicate that individual blastomeres may, under different conditions,
give rise to parts of the embryo other than those which they
produce in the normal course of development.
CLEAVAGE: THE AMPHIOXxUS. — In the egg of the amphioxus
(Fig. 50), which is isolecithal, cleavage is holoblastic and almost
equal. The first cleavage commences as a depression at the
animal pole, which later assumes a groove-like form and elongates
until it becomes a wide meridional furrow extending around the
egg. This constriction deepens until the two hemispheres are
completely divided, when each blastomere rounds up into a
spherical shape. The second cleavage also commences at the
animal pole and is meridional but at right angles to the first,
following the second rule of Sachs. The third plane of cleavage
96 CLEAVAGE AND THE GERM LAYERS
is at right angles to both the first and second and hence would be
equatorial if the egg were completely isolecithal. But as the yolk
is a little concentrated at the vegetal pole, the nucleus, following
Hertwig’s first rule, is in the center of the protoplasm, i.e., on the
egg axis slightly nearer the animal pole. So the third cleavage
plane is nearer the animal pole and accordingly is latitudinal.
The quartette of cells in the animal hemisphere is therefore smaller
than those in the vegetal hemisphere. The smaller cells are called
micromeres; the larger ones, macromeres. The fourth division
Anterior
Presumptive
Anterior —~ mesoderm
Fig. 50. — Cleavage of the amphioxus egg. A, before cleavage. B, commencing
first cleavage, from posterior side. C, second cleavage, from vegetal pole. D,
third cleavage, from left side. KE, fourth cleavage, from vegetal pole. F, fifth
cleavage, side view, segmentation cavity indicated by dotted lines. 166.
(After Conklin, 1932.)
divides each of the eight existing blastomeres in two. There are
two planes of cleavage, each meridional, at right angles to the
third, and also at right angles to each other. Sometimes the
cleavage planes of the fourth division are parallel to each other
instead of being at right angles. This makes the bilateral
symmetry of the dividing egg quite obvious. In the fifth cleavage
32 cells are produced, again by two planes of cleavage, at right
angles to the planes of the fourth, but this time latitudinal and
parallel to each other. From this time on cleavage becomes more
and more irregular. The early cleavages have been fairly regular;
CLEAVAGE: THE FROG 97
each has divided the entire egg mass; and the blastomeres, with
the exceptions noted, have been almost equal. The blastomeres
round up as each cleavage is completed, and a jelly is secreted
between them. In this way a small cavity called the segmentation cavity or blastocoel is formed.
Conklin (1933) states that comparison of the cleavage of the
amphioxus with that of the tunicates shows a general resemblance
between the two in the distribution of the organ-forming substances to the blastomeres, in the generally bilateral type of
cleavage, and the order of division; but in all respects the tunicate
egg is the more precise and the more precocious in differentiation.
CLEAVAGE: THE FROG. — The frog’s egg (Fig. 51) is telolecithal
with holoblastic unequal cleavage. Here the first division com
Fig. 51. — Cleavage of the frog’s egg. A, third cleavage. B, fourth cleavage (12
cells). C, fifth cleavage. D, sixth cleavage. 1, I’, later stages. (After Morgan.)
mences as a depression at the animal pole, which elongates and
extends around the egg as a shallow furrow until the ends meet
at the vegetal pole. The constriction extends inwards and eventually bisects the egg into two blastomeres which round up very
slightly. The plane of second division is also meridional and
through the animal pole but at right angles to the first. The first
two cleavage planes intersect each other at the animal pole; but
as the blastomeres round up, the planes no longer form a cross,
98 CLEAVAGE AND THE GERM LAYERS
but two blastomeres are pushed away from each other, while the
other two are in contact forming a short polar furrow between
them. The third cleavage is latitudinal, about 20° above the
equator, and the micromeres are considerably smaller than the
macromeres. Theoretically the fourth and fifth planes of cleavage bear the same relationships to the earlier ones as do those of
Amphioxus, but actually they are more irregular. The two planes
of the fourth cleavage often fail to pass through the vegetal pole
and hence become vertical rather than true meridional planes.
As these planes originate in the animal hemisphere, the micromeres are divided before the macromeres, so that a 12-cell stage
intervenes between the 8-cell and 16-cell stages. Similarly, following Balfour’s rule, the latitudinal cleavage plane in the animal
hemisphere of the he fifth division appears before the corresponding
“a plane in the vegetal hemisphere,
so that there is a 24-cell stage
before the 32-cell stage is attained.
The cell lineage of the frog’s egg
has not been followed in detail as
A B has that of the tunicate or amphi
Fie. 62.—The gray crescent of the Oxus. It is known, however, that
frog’s egg in early cleavage. A, first the first cleavage plane ordinarily
cleavage, posterior view. W}.third divides the gray crescent into two
cleavage, from left side. dagen trical hal .
diagrammatic. bs symmetrical halves (Fig. 52A), so
that cleavage is normally bilaterally symmetrical from the outset. The blastomeres receiving the
gray crescent material will give rise to notochord and neural
plate in later development.
CLEAVAGE: THE CHICK. — In telolecithal eggs with meroblastic
cleavage such as that of the fowl, only the protoplasm of the egg,
i.e., the blastodisc, is divided, and the cleavage planes do not segment the yolk (Fig. 53). The first furrow commences at the
animal pole and extends outwards towards the edges of the blastodisc. The second is formed by two furrows, at right angles to
the first, one in each blastomere, which grow towards the first furrow and also towards the edge of the blastodisc. They may join
the first furrow at approximately the same point or at separate
points, in which case a polar furrow is formed. These four cells
are incomplete, as the furrows do not extend all the way to the
CLEAVAGE: MAN AND OTHER MAMMALS 99
"y
yolk nor to the edge of the blastodise, but remain connected both
below and at their margins. From this point on, tleavage is
irregular. Some cleavage planes are circular and cut off central
cells from marginal. These may be compared with the latitudinal
planes of the holoblastic type. Others are radial, like the first
Cc
Fia. 53.— Cleavage of the hen’s egg. A, first cleavage. B, second cleavage.
C, third cleavage. D, later cleavage. All from animal pole. Approx. X12.
(A, B, D, after Kélliker; C, after Patterson.)
and second. Still others are tangential and divide the central
cells into upper and lower layers, as in the frog’s egg.
CLEAVAGE: MAN AND OTHER MAMMALS. — The cleavage of the
human ovum has not yet been observed, but in the egg of the
monkey (Fig. 54) and rabbit (Fig. 55) the cleavage is clearly of the
equal holoblastic type. In the rabbit the first cleavage takes place
100 CLEAVAGE AND THE GERM LAYERS
about: 22} hours after coitus. It is equal and complete. The
second cleavage follows in about 3 hours. Here the two cleavage
spindles frequently lie at right angles to each other so that the four
blastomeres assume the form of a cross. Cleavage is now irregular, 5-, 6-, 7-, and 8-cell stages appearing in order. The 8-cell
Fia. 54. — Cleavage of the monkey’s egg. A, first cleavage. B, second cleavage.
C, third cleavage. 170. (After Lewis and Hartman in Arey.)
Vitelline
Fig. 55. — Cleavage of the rabbit’s egg. A, fertilized egg (note albumen layer).
B, first cleavage. C, second cleavage. D, third cleavage. I, fourth cleavage.
F, fifth cleavage. 180. (After Gregory.)
stage is attained about 32 hours after coitus. There is now considerable difference in size, the largest blastomere being almost
twice the size of the smallest. The 16-cell stage is reached in
another hour and a half. In reaching this stage the cleavage of
one blastomere is tangential so that there is always one cell completely enclosed. In later cleavages more tangential cleavages
THE BLASTULA 101
occur, and this, with the shifting of the blastomeres upon each
other, results in a solid mass of cells called a morula.
The blastula. — The period of cleavage terminates in the appearance of the blastula, but this does not mean that cell division
comes to anend. The blastula is generally defined as_a hollow
sphere of blastomeres surrounding a cavity, the blastocog]. But
Blastocoel
Fig. 56.— Diagrams of vertebrate blastulae. A, coeloblastula following holoblastic equal cleavage (amphioxus). , coeloblastula following holoblastic unequal cleavage (frog). CC, discoblastula following meroblastic cleavage (chick).
D, blastocyst (mammals.)
this definition does not fit the blastulae formed by meroblastic
cleavage. So we shall distinguish three classes of blastulae. The
first is of the hollow sphere type (coeloblastula) and is the result
of holoblastic equal cleavage*(Fig. 56A). A variety of this type,
in which the blastocoel is displaced towards the animal pole, is the
result of holoblastic unequal cleavage (Fig. 56B).
The second type of blastula (discoblastula) is the result of
meroblastic cleavage in which the blastomeres rest in a flat disc,
102 CLEAVAGE AND THE GERM LAYERS
the blastoderm, on the undivided yolk mass (Fig. 56C). A segmentation cavity later combines with a yolk cavity, formed by the
digestion of the yolk underlying the blastoderm, to form a blastocoel. Such a blastocoel is roofed with cells but has a floor of yolk.
The third type of blastula is found only in mammals and is
called a blastocyst (Fig. 56D). The solid morula forms a blastocoel which enlarges until it almost separates an outer layer of
cells (trophoblast) from an inner cell mass (the embryonic knob).
Presumptive organ regions of the blastula. — As might be inferred from the results of cell-lineage studies, the regions of the
blastula will give rise to different parts of the embryo in normal
development. In the tunicate and amphioxus, Conklin has
mapped out the presumptive organ regions of the blastula, and
Vogt and his students, by means of a most ingenious technique,
have accomplished the same result for the amphibian blastula.
Experimental evidence (Chapter VII) indicates that in the tunicate and amphioxus the organ-forming regions are definitely determined whereas in amphibians, the regions have a greater plasticity and may give rise to parts of the embryo quite different
from those formed in normal development.
BLASTULA OF THE AMPHIOXxuS. — In the development of the
amphioxus we find a good example of the coeloblastula (Fig. 57).
The blastomeres are arranged
in a single layer around the enlarged blastocoel which is entirely cut off from the exterior.
The blastomeres at the animal
pole are micromeres; those at
the vegetal pole are macroPresumptive — Mees ; the cells at the equa- .
torial belt are transitional in
Presumptive type. ;
Fia. 57. — Blastula of the amphioxus. Sag- The cells which are to form
ittal section. X220. (After Conklin.) the mesoderm are rounded and
in active mitosis. They are
arranged on a crescent on one side of the egg while those which
will form the chorda-mesoderm make up a corresponding crescent
on the other. The endoderm cells are the larger cells of the
vegetal hemisphere.
Presumptive
ectoderm
Ses
BLASTULA OF THE FROG
103
BLASTULA OF THE FROG. — The blastula of the frog (Fig. 58)
resembles that of the amphioxus in all essential characters, but
shows minor differences due
largely to the greater amount of
yolk present. In the first place,
the blastoderm is no longer one
layer of cells in thickness. Tangential divisions have increased
the number of cells so that at the
animal pole the blastoderm may
be approximately four cells deep.
Furthermore, the greater differ
ence in size between the micro- Fr
meres of the animal pole and the
macromeres of the vegetal pole
Dorsal lip
A of blastopore 4 3 B 2
\, Blastopore
IG.
Fig. 59. — Diagrams of the Triton egg showing
movement of surface areas stained with nile
blue and neutral red during gastrulation.
Areas on surface shown with sharp outline,
those on interior without outline.
Vogt.)
(After
Micromeres
Blastocoel
UR 6
58. —Blastula of the frog. Vertieal section. (After Brachet.)
result in the blastocoel’s occupying an eccentric position entirely within the
limits of the animal hemisphere.
The blastula of the frog
shows certain regional differentiations. Thus the
cells of the animal hemisphere are smaller than
those of the vegetal hemisphere. Morgan has pointed out that those arising in
the region of the gray crescent are definitely smaller,
i.e., dividing more rapidly,
than those in any other
meridian.
Vogt has demonstrated
the fate of different regions of the blastula in normal development
by marking them with such harmless dyes as nile blue and neutral
red. The stain persists long enough so that the migration of the
dyed cell groups can be traced through gastrulation and even later
(Fig. 59).
He has succeeded in mapping out the surface of the
Sucker
Sucker
Eye Epidermis
Limit of neural plate
Neural fold
Limit of inturned
material
Notochord
Somites
Anterior limb bud
Visceral pouches
Lip of blastopore
Fig. 60. — Diagrams to show presumptive organ regions of the frog blastula. A, from left side. B, from dorsal surface.
The cross indicates the position of the vegetal pole. (After Vogt, 1929.)
104
CLE
AVAGE
AND THE GERM LAYERS
~
BLASTULA OF THI CHICK 105
blastula into presumptive organ regions, as seen in the diagram
(Fig. 60).
BLASTULA OF THE CHICK. — The blastula of the chick is a discoblastula. The blastoderm consists of an inner mass of micromeres completely separated from one another by cleavage planes,
and an outer ring of macromeres which are partially separated
from one another by incomplete radial cleavage planes only.
These latter cells are in direct protoplasmic continuity by means
of an outer ring of undivided cytoplasm and a thin lower layer of
undivided cytoplasm passing beneath the inner mass (Fig. 61).
This undivided cytoplasm is called the periblast. The micromeres of the inner mass are separated from the underlying un
Blastocoel Blastomeres Periblast
Fig. 61. — Section of early chick blastula. Compare Fig. 53D. (After Patterson.)
divided periblast by means of a thin cleft which is the original
blastocoel.
The blastoderm expands over the yolk, new cells being added
to the inner cell mass from the outer ring of cells. The periblast,
contributing its cytoplasm to the formation of new cells in the
outer ring, soon uses up all the material contained in the thin
lower layer. Meantime its outer ring, now called the germ wall,
expands outward. With the disappearance of the lower layer of
periblast, the cells of the inner mass form the roof of a cavity
which includes the original blastocoel plus the space originally
occupied by the lower layer of periblast. These cells form an area
known as the area pellucida because it can be detached from the
yolk without carrying any yolk particles and hence appears more
transparent. The cells of the outer ring and the germ wall make
up the area opaca, so-called because particles of yolk adhere to
them when removed from the egg and render them less transparent.
106 CLEAVAGE AND THE GERM LAYERS
BLASTULA OF MAN AND OTHER MAMMALS. — No human embryo in the blastula stage has been recorded, so a description of
the blastocyst of the rabbit will be given in its place. About 75
hours after coitus and while the egg is still in the oviduct, a cleft,
the blastocoel, appears in
Embryonic knob the morula apparently
Albumen Embryonic duc to the formation of
some fluid. This extends
rapidly until an outer
layer of cells, the trophoblast, is separated from
an inner cell mass, the
embryonic knob. The
Blastocoel
ae separation is almost comTrophoblast ’ . ,
B plete (Fig. 62A), extend. °
Fia. 62. — Sections of rabbit blastocysts. 200. ing about 270 of the
(After Gregory.) possible 360°. By this
- time the blastocyst has
reached the uterus and the secretion of fluid is greatly increased,
expanding the blastocoel and stretching the trophoblast cells.
The embryonic knob flattens against one pole (dorsal) of the trophoblast, and the entire blastocyst increases greatly in size (Fig.
62B). This flattening of the embryonic knob is not characteristic
of all mammalian blastocysts.
B. GASTRULATION
The vertebrate blastula becomes converted into a two-layered
embryo, or gastrula, through the migration of cells from the
exterior to the interior of the embryo. In so doing the blastocoel
is obliterated and replaced by a new cavity, the gastrocoel
(archenteron), which communicates to the exterior by means of an
opening, the blastopore. The cells left on the exterior form the
outer germ layer commonly known as ectoderm (ectoblast, epiblast). Those on the inside, lining the gastrocoel, form the inner
germ layer, usually called the endoderm (entoderm, entoblast,
hypoblast). But, as will be seen later, they may also include
cells which will give rise to the middle germ layer, the chordamesoderm, consisting of the mesoderm (mesoblast) and notochord
(chorda dorsalis). In such cases the inner layer may be called
GASTRULATION 107
mesendoderm (see page 115). The different types of blastulae
resulting from different types of cleavage naturally give rise to
different types of gastrulae (Fig. 63) according to the means by
which the endoderm is segregated from the ectoderm.
Students of gastrulation distinguish five types of cell migrations which will be
described briefly here, and developed more fully in later paragraphs.
1. Invagination (Fig. 638A). Typical of the cocloblastula resulting from equal
holoblastic cleavage. The cells of the animal hemisphere move inward in a con
Gastrocoel
Blastocoel
Fig. 63. — Diagrams of vertebrate gastrulation. A, by invagination (amphioxus).
B, by epiboly and involution (frog). C, by involution (chick). D, by delamination (mammal).
tinuous shect, obliterating the blastococl, until they come to rest against the cells of
the animal hemisphere, thus giving rise to a new cavity, the gastrocoel, which opens
to the exterior by means of the blastopore. This process is made possible by the
continued growth of cells at the lip of the blastopore which roll inward (involution,
see 3) as invagination continucs.
2. Epiboly (Fig. 63B). Typical of the coeloblastula resulting from unequal
holoblastic cleavage. The cells of the animal hemisphere grow over the cells of the
vegetal hemisphere, creating a gradually narrowing circular fold, the lip of the
blastopore. This process also involves the growth and rolling inward of cells at the
moving lip (involution, sec 3) to form the roof of the gastrocoel.
3. Involution (Fig. 63B, C). Typical of the discoblastula resulting from meroblastic cleavage. The cells at one region of the disc roll inward and spread out
under the disc to form the roof of a gastrocoel. The region where involution takes
place is the dorsal lip of the blastula. Involution also accompanies invagination
and epiboly (sec 1 and 2).
4. Delamination (Fig. 63D). Typical of the blastocyst in mammals. The lower
108 CLEAVAGE AND THE GERM LAYERS
cells of the embryonic knob split off as a loose layer which later reorganizes itself to
enclose a gastrocoel.
5. Concrescence (Fig. 64). As the blastopore narrows, cells which originally lay
along the right and Iecft halves of the dorsal lip converge towards each other. And,
since the dorsal lip is also growing backward, these cells will form the right and
left sides of an axial (antero-posterior) streak.
43 34
B
4391234
44
Cc
Fig. 64. — Diagrams showing four stages in the process of concrescence.
(After Lillie.)
GASTRULATION IN THE AMPHIOXuS. — The first indication of
gastrulation is a flattening of the macromeres of the vegetal hemisphere (Fig. 65A). These cells divide less frequently and become
more columnar, while the others divide more frequently and become more cubical or spherical in shape. This change in the
shape and rate of division, says Conklin (1932), is apparently the
principal cause of invagination (Fig. 65B), although it may be
due also in part to the resorption of material from the blastocoel
jelly, or to exosmosis, for the contents of the blastocoel become
less viscous as gastrulation proceeds.
In later stages of gastrulation the gastrula increases in length,
owing to the backward growth of the lips of the blastopore (Fig.
65C). While this process is taking place cells are being rolled
from the exterior to the interior (involution). The lips of the
blastopore grow unevenly, the ventral lip finally turning upward
GASTRULATION IN THE FROG 109
to reduce the blastopore to a very small opening (Fig. 65D).
Conklin expressly denies that this narrowing of the blastopore is
caused by the growing together of the right and left halves of the
dorsal lip (concrescence). The cells left on the exterior after
gastrulation is complete are ectoderm. Those which have been
carried to the interior are endoderm, and presumptive chorda
Presumptive ectoderm
| Presumptive
7 chordax neural plate
Presumptive
mesoderm Presumptive
endoderm
Fig. 65. — Sections of amphioxus embryos during gastrulation. A, blastula (6
hours after fertilization). B, gastrula (93 hours). C, gastrula (12 hours).
D, gastrula (14 hours). Animal pole indicated by presence of polocyte. Anteroposterior axis shown by arrow. All sagittal sections. 180. (After Conklin,
1932.)
mesoderm. The segregation of the notochord and mesoderm cells
is discussed in Section C of this chapter.
In late gastrulation the cells of the ectoderm develop cilia, by
means of which the embryo rotates within its fertilization
membrane.
GASTRULATION IN THE FROG. — The first stage in the gastrulation of the frog is the formation of a groove on the dorsal side of
the embryo in the region of the gray crescent (Fig. 66A). Along
this groove, cells are pushed into the interior (involution), while
at the same time the cells immediately above the groove are
growing down over the surface of the embryo to cover them
(epiboly). In this way a two-layered fold is produced, the dorsal
lip of the blastopore (Fig. 66D).
110 CLEAVAGE AND THE GERM LAYERS
ahs
Roof of
gastrocoel
Dorsal
lip
Neural plate
Notochord
Endoderm
Fia. 66. — Three stages in the gastrulation of the frog’s egg. A, dorsal lip stage, from
vegetal pole. B, do., sagittal section. C, lateral lip stage, from posterior surface.
D, do., sagittal section E, ventral lip (yolk-plug) stage, from posterior surface.
¥, do., sagittal section. (B, D, I, after Brachet.)
GASTRULATION IN THE CHICK AND PIGEON 111
As the two-layered fold grows down over the cells of the vegetal
hemisphere, it extends laterally, thus forming the lateral lips of
the blastopore (Fig. 66B). And, since it is covering a spherical
surface, the ends of the fold eventually meet to form the ventral
lip (Fig. 66C). Epiboly and involution take place at all points
on the lip of the blastopore, but chiefly at the dorsal lip, which
moves approximately 90° around the egg. At this time the egg
presents the appearance of a black sphere with a small white
circular area, known as the yolk plug (Fig. 66C).
Within the egg, two distinct phenomena have been taking place.
First, the cells turned inward by involution at the dorsal lip have
spread out to form the roof of a wide but shallow cavity, the
gastrocoel. Second, small cells have arisen from the large yolkladen cells of the vegetal hemisphere, and these form the floor of
the gastrocoel. They join the cells resulting from involution at
the anterior end of the gastrocoel (Fig. 66E).
There is now an extensive displacement of the interior cells, resulting from the growth forward of the gastrocoel, and the consequent thinning of its floor. It is still uncertain whether the floor
is pushed across the blastocoel, thereby obliterating it, or whether
the thin floor is ruptured so that the blastocoel is added to the
enlarging gastrocoel (Fig. 66F). In either event the center of
gravity in the egg is now altered so that it rotates about a horizontal axis in such a way that the blastopore is carried back to a
point a little beyond its starting point, 100°.
The blastopore is now in its definitive position and marks the
posterior end of the embryo. The dorsal side, already marked by
the appearance of the dorsal lip, is uppermost. In the concluding
stages of gastrulation the blastopore narrows to a small slit.
This narrowing is brought about by the growing together of the
right and left halves of the dorsal lip (concrescence) as epiboly
and involution continue.
The cells of the inner layer during later stages of gastrulation appear to be split into two separate layers. The one of these which
lines the gastrocoel is endoderm. The other lying between the
endoderm and the ectoderm is the chorda-mesoderm. The mode
of origin of the latter will be described in the following section.
GASTRULATION IN THE CHICK AND PIGEON. — The blastula of
the chick is a disc of blastomeres lying over the undivided yolk.
112 CLEAVAGE AND THE GERM LAYERS
It is divided into an interior area pellucida and an outer area
opaca. This outer area is extending itself in all directions over
the undivided yolk (epiboly).
The account which follows
is based on gastrulation in
the pigeon.
Three zones are distinguishable in the area opaca.
First, there is a margin of
overgrowth where the cells
are completely separated
from the yolk. Second
comes a zone of junction,
whose deeper cells are not
separated from the yolk.
The third division is the inner zone, whose cells, completely separate from the
yolk, are being added to the
_ area pellucida.
The first indication of
gastrulation is the thinning
of the blastoderm at the
posterior end and the complete separation of the cells
from the yolk at that region
(Fig. 67A). In other words,
there is a crescentic area,
almost a quarter of the cir
Endoderm cymference, of the blastoderm which lacks the zone
of junction completely.
Fra. 67. — Surface views showing three stages Here the cells roll inward
in the gastrulation of the hen’s egg, from ,. luti Fie. 68 d
the animal pole. (After Patterson.) (invo ution) (Fig. ) an
multiply until they have
spread completely under the upper layer to roof in the old blastocoel and convert it into the new gastrocoel, whose floor is made
up of undivided yolk. The slit-like opening where the zone of
Dorsal lip
Dorsal lip
GASTRULATION IN MAN AND OTHER MAMMALS 113°
junction disappeared is the blastopore, and the rim along which
involution took place is the dorsal lip.
There is very little overgrowth at the dorsal lip while involution
is taking place, and consequently the edges of the blastoderm on
either side swing around to enclose the lip region in the advancing
Fia. 68. — Sagittal section through early gastrula of pigeon (36 hours after fertilization). Posterior half of section only. d.b., dorsal lip of blastopore. (J*‘rom
Richards after Patterson.)
germ wall. In this way the blastopore is compresscd laterally and
concrescence takes place.
GASTRULATION IN MAN AND OTHER MAMMALS.— No human
embryo nas been observed before the separation of the germ
layers. The account which follows is based on the pig. From
the lower surface of the embryonic knob, individual cells detach
themselves to form a sheet (Fig. 69) which rapidly establishes
Fia. 69. — Section to show an early stage in the gastrulation of the bat’segg. (After
Van Beneden.)
itself as a layer immediately inside the trophoblast, enclosing
most of the old blastocoel. We may now consider the trophoblast
and the remainder of the embryonic knob as ectoderm and the
inner layer as endoderm. The cavity which it encloses is com114 CLEAVAGE AND THE GERM LAYERS
parable to the gastrocoel plus the yolk sac of the egg-laying
mammals.
The cells of the trophoblast immediately overlying the embryonic knob (Rauber’s cells) now disappear, and the embryonic
knob flattens out to become the embryonic disc. This disc lies
at the surface and constitutes part of the wall of the blastocyst.
In the primates, judging from studies on the lemur, Tarsius,
and from the appearance of the earliest human embryo (Fig. 70),
the endoderm does not grow out around the entire trophoblast,
Endoderm
Amniotic
cavity
Fia. 70. — Diagrams to show three stages in the gastrulation of the human egg
during implantation. The uterine wall indicated by hatching. (Hypothetical
based on Teacher; the embryo in C based on Miller.)
but forms a very small vesicle immediately under the embryonic
y y
knob. The cavity of this vesicle may be considered a gastrocoel
but is more generally known as the ‘‘yolk sac.”
C. THE MIDDLE GERM LAYER (CHORDA-MESODERM)
During or immediately following gastrulation a third germ
layer appears between the ectoderm and endoderm. This layer
consists of the notochord (chorda dorsalis), an axial supporting
rod found only in the vertebrates and their allies the protochordates, and two sheets of mesoderm on each side of the notochord. Later wandering ameboid cells, originating from the
mesoderm and known collectively as the mesenchyme, make their
appearance.
The student should note that in many elementary texts the
middle germ layer is called the mesoderm and that the notochord
is variously derived from mesoderm, endoderm (amphioxus and
frog), or ectoderm (chick and mammals). This terminology dates
back to the phylogenetic period of-embryology (Chapter I), when
THE LATER HISTORY OF THE GERM LAYERS 115
it was supposed that a blastula composed of undifferentiated
blastomeres gave rise to a gastrula with two separate (primary)
layers, and that the mesoderm and the notochord arose separately
from one or the other of the so-called primary layers, primitively
from the endoderm. Today it is generally recognized that the
notochord arises in the same manner and at the same time as the
mesoderm. To avoid the clumsy phrase, mesoderm and notochord, many writers are now employing the term chorda-mesoderm for the middle germ layer, and restricting the term mesoderm
to the middle germ layer exclusive of the notochord, a usage
employed in this text. The compound word mesendoderm
(mesentoderm) is now used by many writers to include both the
endoderm and the chorda-mesoderm when these layers lie beneath the ectoderm but have not yet segregated from each
other.
In collateral reading the student will sometimes encounter the
word endo-mesoderm used in connection with mesoderm “ originating from” or, better, associated with, endoderm in early
development. Similarly the word ecto-mesoderm is employed
to designate mesoderm “ originating from,” or associated with,
ectoderm in early development. Other writers use the terms
peristomial mesoderm, meaning mesoderm appearing in the region
of the blastopore, and gastral mesoderm for mesoderm appearing
to arise from the invaginated endoderm. But inasmuch as the
middle germ layer can often be traced to definite blastomeres
during early cleavage, this distinction is of small importance.
It is well established, however, that among the vertebrates the
movement of the presumptive chorda-mesoderm to its definitive
position in the roof of the gastrocoel is intimately associated with
the formation and closure of the blastopore. This is true no
matter whether the blastopore is a large circular opening as in the
amphioxus and the frog, or reduced ‘to a primitive streak by
concrescence as in the chick and man.
The later history of the germ layers. — With the segregation
of the three germ layers, the presumptive organ regions are now
located in one or another of the three. But it must not be supposed that the organs of the adult are exclusively ectodermal,
endodermal, or mesodermal. On the contrary, most of them contain material from at least two, and sometimes all three. In Part III will be found an account of the development of the different organ systems, classified according to the germ layer from
which arise the tissues associated with their special functions.
Meantime the following table is presented.
TABLE 7
DERIVATIVES OF GERM LAYERS
Eetoderm Chorda-mesoderm Tendoderm
A Notoehord
2B) Mesoderm
1. Epidermis of skin and 1. Epithelium of coclom 1. Mpithelia of digesall openings into the body | and exocoel tive tube, including thy2. Epithelia of eye, ear, 2. Nephric (exeretory) | mus gland, thyroid gland,
and nose system parathyroid gland, in3. Nervous system, in- 3. Genital (reproduc- | ternal respiratory — orcluding interrenal glands, | tive) system gans, volk sac, and alpituitary gland (in part), 4. Suprarenal gland lantois
pineal gland 5. Blood-vaseular sys4. Epithelium of amnion | tem
and chorion 6. Connective — tissue
including skeleton
7. Musculature
8. Dermis of skin
THE MIDDLE GERM LAYER IN THE AMPHIOXUS. — \s mentioned
in earlier sections, Conklin (1933) has been able to distinguish the
mesoderm cells in the amphioxus in the blastula stage (ig. 57),
where they form a crescent of small rounded blastomeres in the
region where the ventral and lateral lips of the blastopore will
form. The notochord cells, associated with those which will later
give rise to the neural plate, occupy a corresponding chordaneural crescent at the dorsal lip. After the invagination of the
endoderm the cells of the mesoderm and notochord form the lip
of the blastopore, the notochord cells at the dorsal lip, mesoderm
at the ventral and lateral lips. As the lips of the blastopore grow
backward, these cells are carried to the interior by involution
(Fig. 65).
When the ventral lip grows upward, the mesodermal cresent is
tilted up behind so that its arms run in an antero-posterior direction to form the angles between the roof and sides of the
gastrocoel (Fig. 71). In the meantime the notochord cells, also
carried into the interior, form a flat plate between the two arms
of the mesoderm. Thus the roof of the gastrocoel is composed
THE MIDDLE GERM LAYER IN THE AMPHIOXUS 117°
of three strips of chorda-mesoderm, mesoderm on each side,
notochord in the middle.
A longitudinal groove in the notochord plate deepens, and the
folds on either side come together to form a solid cord separate
Neural plate
KEL
Ae ; STR avoocs
bEES Hy Y Kerk Mesodermal
BS ae ; cr Ve ie S groove
fees : pes * ©
Gastrocoel ay ae CS SEL Gastrocoel
Ho Ye: = : : a
Mesodermal sae x Ha Blastopore
Notochord ey :
Neural hse { <
plate Years A
Gs B
Blastopore” “O
A
Fig. 71. — Optical hemi-sections of amphioxus gastrula (14 hours after fertilization).
A, left inside. B, posterior to show notochord and mesodermal groove inside.
166. (After Conklin, 1932.)
from the ectoderm above and the mesoderm on either side. The
mesodermal grooves (Fig. 72) also become deeper. Transverse
constrictions meantime appear in the lateral grooves, which
Anterior
neuropore >\*:
Level of B_|:
Fig. 72. — Sections of amphioxus embryo (19 hours after fertilization). A, sagittal
section. B, transverse section. 166. (After Conklin.)
divide them into a series of pouches (enterocoels). Finally these
pouches are constricted off from the gastrocoel and become the
paired somites (Fig. 73).
The endoderm, which formerly occupied the floor and anterior
end of the gastrocoel, extends to form new sides and a new roof.
118 CLEAVAGE AND THE GERM LAYERS
The gastrocoel, now for the first time completely lined with
endoderm, is the primordium of the digestive tube.
The cells of the chorda-neural crescent remaining on the exterior of the embryo give rise to the neural plate on the dorsal
surface. They are covered by the ventral lip of the gastrula as it
grows over the dorsal side of the embryo. Beneath this covering
Neural plat
Enterocoel cura’ plate
Fia. 73. — Sections of amphioxus larva (244 hours after fertilization). A, frontal
section. B, transverse section. X166. (After Conklin.)
there appears a longitudinal groove with a fold on either side.
These folds arch up and meet in the ventral line to form the neural
tube.
THE MIDDLE GERM LAYER IN THE FROG. — As noted in earlier
sections, we owe to Vogt (1929) the identification of the various
regions on the amphibian blastula. This identification was accomplished by staining small regions of the blastula surface with
harmless dyes and tracing their movements during and after
gastrulation (Fig. 59). He finds that the material first to be
turned in at the dorsal lip is endoderm. Immediately anterior
and dorsal to this is a crescent-shaped area which will give rise to
the notochord. On either side of this are the horns of a crescent
extending from the other side of the blastula which will become
mesoderm. Immediately anterior to the chorda crescent is the
crescent-shaped area of the neural plate, the two together being
equivalent to the chorda-neural crescent of the amphioxus. The
THE MIDDLE GERM LAYER IN THE FROG 119.
mesodermal crescent also corresponds to the mesodermal crescent
of the amphioxus except that its arms already extend dorsally.
In the gastrulation of the tailed amphibia (urodeles), the material turned in at the dorsal lip is notochord and mesoderm, so that
the roof of the gastrocoel is chorda-mesoderm as it is in the
amphioxus, and endoderm cells must grow up from the sides and
floor to form a new roof.
In the frog, however, the first material to roll in at the dorsal
lip of the blastopore is endoderm and notochord (Fig. 60). When
the material from the mesodermal crescent rolls in, instead
of following the endoderm, it
wedges in between the endoderm and ectoderm (Fig. 66F),
so giving the appearance of
splitting off from the endoderm
in the roof of the gastrocoel.
The roof and sides of the gastrocoel are, therefore, endodermal except for a narrow dorsal
strip represented by the notochord (and a narrow strip beneath it, the hypochord).
When the notochord (and hypochord) separate from the roof,
this small gap is closed by endoderm and the roof is completely
endodermal.
As the endoderm, notochord,
and mesodermal regions are
turned in around the lips of the
blastopore the overgrowth of
the lips covers the large yolkladen cells from which the floor
of the gastrocoel is produced.
Meantime the expanding cells
from the ectodermal region of
the blastula occupy the region
formerly held by the material which has been turned in.
Fa. 74. — Diagrams showing direction of
displacements during amphibian gastrulation. A, from posterior surface. B,
from left side. Thick lines on exterior
surface. Thin lineson interior. (After
Vogt, 1929.)
Now the
120 CLEAVAGE AND THE GERM LAYERS
dorsal lip of the blastopore is the one at which epiboly and involution take place most rapidly. Consequently materials on the
right and left of the mid-dorsal region are stretched towards the
medial line to take the place of the material lost by involution
(Fig. 74). In this way the two arms of the mesodermal crescent
move together to form parallel strips on either side of the noto
Notochord
Mesoderm
Fig. 75. — Transverse sections to show three stages in the origin of the notochord
and mesoderm in the frog embryo. (After Brachet.)
chord. Similarly the two horns of the neural crescent move together to form parallel strips which eventually enclose the blastopore at the posterior end, while the neural plate itself occupies a
longitudinal dorsal position on the gastrula. All the rest of the
surface is now material
which will form the epidermoet . Mis of the skin.
( myocoel ) The mesoderm continues
Intermediate its growth between the ec
mesoderm
(nephrocoel) toderm and endoderm
Lateral (Fig. 75) until it forms a
Cecoleen} continuous sheet except at
the blastopore. The material on either side of the
notochord is separated by
Fie. 76. — Diagram of a transverse section of transverse constrictions
vertebrate embry 0 te show the regions of the into blocks or somites, corresponding to the somites
of the amphioxus. Next comes an intermediate zone from which
the gonad and kidney will arise. The remainder splits into an
outside (somatic) layer closely applied to the ectoderm, and an
THE MIDDLE GERM LAYER IN THE CHICK 121
inner (splanchnic) layer similarly applied to the*endoderm. The
space between (Fig. 76) is the coelom.
The neural plate develops a longitudinal groove, surrounded at
the anterior end and sides by ridges known as the neural folds.
The embryo has now reached the stage known as the neurula
(Fig. 112).
THE MIDDLE GERM LAYER IN THE CHICK.— Mesoderm formation in the chick takes place after the egg has been laid and incubation begun. At about the sixteenth hour (Patten) the blasto- por ee
derm is considerably lengthened Pa ser,
in an antero-posterior direction, {4
and has an axial thickening known 3
as the primitive streak (Fig. 77). 3
This streak represents the dorsal
lip of the blastopore laterally compressed through concrescence as
explained on page 108. The germ
wall has grown together behind
the primitive streak and is ad- Vie. 77. — Blastoderm of the chick at
- 15 hours of incubation. (After
vancing out over the yolk. Ina puyar)
more advanced embryo the primitive streak is differentiated into a primitive groove in the
middle, primitive folds on either sides, a primitive pit at the
_ Primitive
- streak
Mesoderm
5,
45;
4 mon OTs
Mesoderm Ectoderm Primitive streak
Notochord
Primitive streak
at Endoderm B Notochord
Ectoderm
Dorsal li
pea P «.. Ventral lip
sane a
Endoderm Notochord
Fie. 78. — Blastoderm of chick to show early stage in development of notochord.
A, surface view at 20 hours (after Duval). B, transverse section, left half only.
C, sagittal section. (B, C, after Lillie.)
anterior end of the groove, and a primitive (Hensen’s) node
in front of the pit where the primitive folds unite (Fig. 78).
122 CLEAVAGE AND THE GERM LAYERS
Sections reveal that from the sides and posterior end of the
primitive groove, cells are growing outward, between the ectoderm and the endoderm, to form a sheet of mesoderm. At the
anterior end a narrow strip of cells grows forward to form the
notochord.
During the remainder of the first day of incubation the area
pellucida increases in length, particularly in the region directly
in front of the primitive streak. This appears to displace the
primitive streak rearwards, and during this time the streak
actually shortens.
The mesoderm growing out to the sides is carried forward in
this movement and so comes to lie close to the advancing notochord. Furthermore, two horns of mesoderm grow forward, later
to curve in and meet in front of an area which contains ectoderm
and endoderm only (proamnion). The mesoderm on either side
of the notochord thickens to form a segmental zone, so called
because it will shortly be divided by transverse constriction into
somites, exactly as in the frog. Six pairs of somites are present
at the end of the first day (Duval). There is a zone of intermediate mesoderm. The remaining or lateral mesoderm, growing out into the area opaca, splits tangentially into an outer
somatic and an inner splanchnic layer, as in the frog. In the
splanchnic mesoderm, thickenings appear in the inner region of
the area opaca. They mark the primordium of the area vasculosa
(Fig. 79).
The ectoderm and endoderm of the clear area give rise to a
crescentic fold at the anterior end which is called the head fold
as it is the primordium of the head of the embryo. It contains
a pocket of endoderm known as the fore-gut, distinguished by
the possession of a cellular floor. There is an opening known as
the anterior intestinal portal between the fore-gut and the midgut, whose floor is the undivided yolk.
The ectoderm in front and to either side of the notochord is the
neural plate. It develops a groove and folds shortly before the
end of the first day, and at 24 hours of incubation the folds have
met in the region of the brain to form a tube but have not as
yet fused together.
THE MIDDLE GERM LAYER IN MAN. — The earliest human embryo is the ‘ Miller’ ovum (Fig. 69). This specimen, supposed
THE MIDDLE GERM LAYER IN MAN 123.
to be about 13-14 days old, consists of an outer vesicle, the
trophoblast, containing two smaller vesicles, one of which, lined
with endoderm, represents a small gastrocoel (yolk sac) the
- Area opaca ~ .
. os A : . : : : .
. . : . ° ° °
. oo. 3 - 3 ° .
. co. 3 Ss. & .
s = g- ¥ . .
“6° 8 3 .
- +3: 2 4: : :
SE 2
o° °
Sig: 2 . ;
. we < er °
i< I} .
. ¢ an >
i +¢-(Isthmus) °
a * “Zone of junction
Margin of overgrowth
Fia. 79. — Diagram showing embryonic and extra-embryonic areas of chick embryo
at 24 hours of incubation. Above, surface view; below, transverse section.
other of ectoderm surrounds a cavity (the amnion, Chapter V).
Where the two vesicles are in contact a circular disc of ectoderm
and endoderm pressed together represents the embryonic disc.
In later specimens this embryonic disc develops a primitive streak,
.124 CLEAVAGE AND THE GERM LAYERS
quite as in the chick blastoderm (Fig. 80). Notochord and
mesoderm develop in much the same way, somites appearing at
the end of the first month. A
head fold and neural groove ap
Cut edge of Hear in similar fashion.
amnion
SUMMARY
During cleavage the fertilized
egg is divided into a large number
Notochord of daughter cells or blastomeres
. which arrange themselves about
Primitive a cavity to form the blastula.
The pattern of cleavage and the
form of the blastula vary accord
ay ing to the amount and distribution
Fia. 80. — Surface view of embryonic of the yolk in the fertilized egg.
dise in human embr i 4 ‘ :
has been cut away, wo. * After The presumptive organ Tegions of
Heuser.) the fertilized egg are segregated
into different groups of cells which
compose the presumptive organ regions of the blastula.
During gastrulation, the blastomeres are reorganized into different strata or germ layers about a new cavity, thus forming a
gastrula. The method of gastrulation varies according to the
type of blastula formed after cleavage. The two layers segregated during gastrulation are usually known as the ectoderm
and endoderm, but it must be recognized that one or the other
of these so-called primary layers includes the presumptive
mesoderm as well.
In the concluding period of germ-layer formation, the middle
germ layer or chorda-mesoderm, including the notochord and the
mesoderm proper, is segregated from the other germ layers to
occupy a middle position between them.
While the germ layers are being segregated from each other the
primordia of certain organs are arising from their respective presumptive regions. Thus the notochord is separated from the
mesoderm proper, the neural plate from the presumptive epidermis. In the mesoderm proper, the somites begin to take form,
and the somatic layer separates from the splanchnic to form the
coelom.
REFERENCES 125
REFERENCES
Brachet, A. 1921. Traité d’embryologic, Books 3, 4, and 5.
Conklin, E. G. 1905. The Organization and Cell Lineage of the Ascidian Egg.
Jour. Acad. Nat. Sci. Phila., 2nd Series, Vol. XIII.
—— 1932. The Embryology of Amphioxus. Jour. Morph. 54:69-151.
—— 1933. The Development of Isolated and Partially Separated Blastomeres of
Amphioxus. Jour. Exp. Zool. 64:303-375.
Cowdry, E. V. (ed.) 1924. Gencral Cytology, Section 9.
Gregory, P. W. 1930. The Early Embryology of the Rabbit. Publ. Carnegie
Inst. Wash. 407:141~-168.
Hertwig, O. (ed.) 1906. Handbuch, etc., I, Chaps. 2 and 3.
‘Huxley, J. S., and de Beer, G. R. 1934. The Elements of Experimental Embryology, Chap. 2.
Jenkinson, J. W. 1913. Vertebrate Embryology, Chaps. 5 and 6.
Kellicott, W. E. 1913. General Embryology, Chaps. 7 and 8.
Kerr, J. G. 1919. Textbook of Embryology, II, Chap. 1.
Korschelt, E., and Heider, K. 1902-1910. Lehrbuch, etc., Chaps. 7 and 8.
Lillie, F. R. 1919. The Development of the Chick, 2nd Ed.
MacBride, E. W. 1914. Textbook of Embryology, I, Chap. 17.
Patten, B. M. 1929. The Early Embryology of the Chick, 3rd Ed.
1931. The Embryology of the Pig, 2nd Ed.
Wilson, E. B. 1925. The Cell, etc., Chaps. 13 and 14.
CHAPTER VI
EMBRYONIC FORM AND EXTRA-EMBRYONIC
STRUCTURES
After the germ layers have been segregated, the primordia of
several great organ systems are already localized. Before proceeding to an account of the way in which the organ systems develop from the different germ layers (organogeny), we must
examine the way in which the vertebrate body assumes its form.
This is found to be closely connected with certain structures
(adnexa) which develop also from the germ layers and play an
important part in embryonic (and fetal) life, but which are discarded before hatching (or birth). These extra-embryonic structures are the yolk sac, the amnion, chorion, and allantois, as well
as a structure found only in the mammals, the placenta.
A. THE FORM OF THE BODY
The general form of the vertebrate body is cylindrical, while
the form of the vertebrate egg is spherical. There are in general
_-_Nearal tube two methods of growth
( &_ Notochord Amniotic by means of which the
cae cylindrical shape is at
.. tained. In the first,
y.’ characteristic of smallyolked eggs with a
spherical gastrula, the
Fia. 81. — Diagrammatic transverse sections show- main factor is growth in
ing effects of yolk on form of embryo. A, small length, along the anteroyolked embryo (frog). B, large yolked embryo posterior (cephalo-cau
(chick). (After Assheton.) dal) axis. Inthesecond
type, which is characteristic of large-yolked eggs, the embryo is
modeled from a flat disc into the form of a cylinder connected with
a great yolk sac by some sort of pedestal or stalk. Much of this
modeling is done by the outgrowth of the head and the tail re
spectively, especially among the anamniote vertebrates, but there
126
GENERAL PLAN OF THE BODY 127
is also some actual undercutting, especially evident among the
Amniota. This undercutting is accompanied by the formation
of amniotic folds, as will be seen in the development of the chick.
diagram of cross-sections through the body of a small-yolked
embryo (Fig. 81A) and a large-yolked embryo (Fig. 81B) will
make clear the difference between the cylindrical embryo and the
plate-like embryo before it has been remoulded. In the amniote
vertebrates with a large-yolked egg the embryo develops from
e)
B c
Fig. 82. — Diagrams to show growth in length by concrescence. Arrows indicate
direction of growth. (After Assheton.)
material at the edge of the blastoderm, and as this is rolled together
in concrescence the embryo increases in length (Fig. 82).
vy General plan of the body. — The body of the vertebrate is
basically a tube within a tube, i.e., a digestive tube within a body
tube (Fig. 76).
The digestive tube is endodermal in origin and originates from
the gastrocoel. Here again the small-yolked form has a tubular —
intestine from the beginning. It is only necessary to form anterior and posterior openings, for the blastopore either closes or
is roofed in by the neural folds. The new openings arise from
ectodermal pits, the stomodgum at the anterior end, the proctodeum at the posterior end. In general these openings are not
completed until after the yolk has been wholly consumed. The
gastrocoel of large-yolked embryos has only a roof and sides of
endoderm, for the floor is composed of the yolk. Hence the rolling in or undercutting of the body commencing at the head end,
and later at the tail end, forms a pocket at each end, the fore-gut
and hind-gut respectively. The mid-gut is the remainder of the
open gastrocoel connected with the developing yolk sac by means
of the yolk stalk.
128 EMBRYONIC FORM AND EXTRA-EMBRYONIC STRUCTURES
Between the two tubes lies the mesoderm. The ventral mesoderm of small-yolked embryos (lateral of large-yolked forms)
splits into a somatic and splanchnic layer. The first of these is
closely applied to the ectoderm to form the somatopleure; the
second is associated with the endoderm to form the splanchnopleure. The space between is the coelom or body cavity. Other
and lesser antero-posterior tubes such as the neural tube, formed
from ectoderm, and axial blood vessels, e.g., the aorta, formed
from mesoderm, are indicated in the figure and will be discussed
in later chapters.
Metamerism. — With growth in length is associated a second
factor in the development of the vertebrate body, that of metamerism. This is first indicated by the appearance of metameres
Oral gland
Pronephric
elevation
Somites
Fig. 83. — Diagrams of early embryos to show similarities in body form. A, frog
(after W. Patten). B, chick (after Kerr). C, man (after His).
such as the enterocoels in the amphioxus or somites in the true
vertebrates. In later organogeny are found further evidences of.
metamerism in the nervous system, nephric system, vascular system, and others. However, the primary metamerism of the body
is shown in the mesoderm. The somites are formed successively,
commencing at the anterior end and therefore affording a basis
of classifying the early embryos of any species by the number of
these units present (Fig. 83).
BODY FORM OF THE FROG 129
The head. — The vertebrate body is distinguished by a wellmarked region at the anterior end, containing the mouth, visceral
arches, special sense organs (nose, eye, and ear) and the highly
developed brain. Herein the amphioxus differs from the vertebrates, for it has so little head that some zoélogists make a special
group (Acraniata) to contain it.
The anterior end of the body is already determined in the vertebrate egg (animal pole). It is the surface opposite that of the
blastopore, or in front of the primitive streak. It is the region
where the neural folds first arise and where they first meet. It
is the first part of the body to be freed of the yolk in the largeyolked embryos. A glance at the diagrams of carly embryos
(Fig. 83) will suffice to prove that this is the most highly differentiated part of the body.
In the Amniota the head is inclined ventrally at the region of
the branchial arches. This cervical flexure causes a constriction
(Fig. 83) which is the primordium of the neck, a region found only
in reptiles, birds, and mammals.
The tail. — All vertebrate embryos, even those of species in
which the adult is tailless (frog, man), develop a well-marked tail
in early development. This region is characterized by the absence of a digestive tube and coelom. It develops early in the
anamniotes, where it is of great use to the free-swimming larva,
but more slowly in the amniotes.
The appendages. — The paired appendages of vertebrates arise
as buds (Fig. 83C) which later develop into fins or limbs. Limb
buds do not appear in the amphioxus or the cyclostomes. In all
other vertebrates which do not possess paired appendages in the
adult condition, it is said that limb buds appear in the embryonic
life and are resorbed later.
BODY FORM OF THE FROG. — The spherical egg of the frog,
being only moderately telolecithal, is converted into the cylindrical shape of the embryo principally through the growth of the head
and of the tail.
In the head region the neural plate is much wider than elsewhere, and when the neural folds close in to form the neural tube
the brain will be larger than the spinal cord. On either side of
the head the optic vesicles, the primordia of the eyes, push out
from the brain and make well-marked bulges. The ectoderm im130 EMBRYONIC FORM AND EXTRA-EMBRYONIC STRUCTURES
mediately external to each optic cup will later give rise to the lens
of the eye. Anterior to each eye is a depression in the ectoderm,
the nasal (olfactory) pit. These pits are the primordia of the
nose. Posterior to each eye a similar otic (auditory, acoustic)
pit originates, the primordium of the inner car. On the ventral
side, folds of ectoderm give rise to the ventral sucker (mucous
gland) in the form of the letter V. Between the limbs of the V
there appears an ectodermal pit called the stomodeum or primordium of the mouth. On the ventral side of the body, just ante
Fig. 84. — Growth of the frog embryo. A, late neurula, 2.4mm. B, embryo of 3
mm. C, embryo of 6 mm., just hatched. D, young larva, external gill stage,
9mm. H#, larva, internal gill stage, 11 mm. (Measured alive and drawn after
preservation. X10.)
rior to the base of the tail, a similar pit, the proctodeum, is the
primordium of the cloacal opening.
On the sides of the head five dorso-ventral grooves appear
(in the order I, V, II, III, IV). These are the visceral (branchial,
- gill ”?) grooves, some of which will later break through into corresponding outpushings from the fore-gut, the visceral (pharyngeal, “ gill’) pouches, to form the visceral (pharyngeal, “ gill ’’)
clefts. For the present we need simply note that they separate
six transverse bars or ridges which are known as the visceral
BODY FORM OF THE FROG 131
arches. Each visceral arch contains an aortic arch. (See Table
8.) Arch I (mandibular) contributes to the formation of the
jaws. Arch II (hyoid) contributes to the gill cover (operculum)
and to the support of the tongue. Arches III, IV, and V are often
known as branchials 1, 2, and 3, respectively. On arches III,
TV, and V develop outgrowths which become the external gills
TABLE 8
PHARYNGEAL DerivaTIVES
Pouches Aortic Grooves
TL Arches ‘lefts : ;
(From endoderm) Arehes Clefts Arches — |(From ectoderm)
Visceral Aortic
arch I arch [
(mandibular)
Visceral Visceral Visceral
pouch I cleft I groove I
(hyomandibular) (spiracle of
clasmobranchs)
Visceral Aortic
arch IT arch II
(hyoid)
Visceral Visceral Visceral
pouch II cleft IT groove II
Visceral Aortic
arch LIT arch IIT
(1st branchial)
Visceral Visceral Visceral
pouch ITI cleft IIL groove II]
Visceral Aortic
arch 1V arch 1V
(2nd branchial)
Visceral Visceral Visceral
pouch IV cleft IV groove IV
Visceral Aortic
arch V arch V
(8rd branchial)
Visceral Visceral Visceral
pouch V cleft V groove V
Visceral Aortic
arch VI arch VI
(4th branchial)
Visceral Visceral Visceral
pouch VI cleft VI groove VI
(vestigial in frog) (lacking in frog) (lacking in frog)
132. EMBRYONIC FORM AND EXTRA-EMBRYONIC STRUCTURES
(branchiae). That on V is rudimentary. Later a fold grows
from arch II to cover the external gills completely on the right,
but with an opening on the left known as the atriopore (“ spiracle”’). While this is taking place the grooves between arches
II, III, IV, V, and VI break through into the corresponding
visceral pouches to form the visceral clefts. Internal gills (demibranchs) develop in the clefts, and the external gills disappear.
Meantime the mouth has opened and developed horny jaws.
The tail arises by the backward growth of the tissue in the neural
folds (Bijtel) at the point where they united over the blastopore.
The notochord and neural tube grow backward, carrying epidermis and muscle-forming material with them. Dorsal and
ventral folds make the tail fin.
The paired limbs arise as limb buds. The anterior buds arise
first but are concealed beneath the operculum. The one on the
left side appears first, pushing through the atriopore.
BODY FORM IN THE CHICK. — The body of the chick is cut off
from the blastoderm by the outgrowth of a head fold accompanied
by an undercut, the subcephalic pocket, which appears during
the first day of incubation. This fold extends backward in the
form of an inverted U as the lateral folds arise. These are
also accompanied by undercuts known as the lateral sulci.
Finally there is a posterior tail fold accompanied by a subcaudal
pocket appearing on the third day. Outgrowth at the folds with
some undercutting as well causes the body of the embryo to stand
up from the surrounding blastoderm to which it is attached by a
short pedestal, the umbilical stalk. The head bends down sharply
at the cephalic flexure, but pressing against the yolk, it turns
or twists toward the right so that the left side of the head rests
on the yolk. The ventral bend is known as flexure, the dextral
twist is known as torsion. Flexure and torsion commence in the
middle of the second day of incubation, and continue in a caudal
direction until, at the end of the fourth day, the chick lies completely on its left side.
The primordia of the brain and sense organs arise much as they
do in the frog. A stomodeum appears early in the third day of
incubation, the proctodeum during the fourth day. Four visceral
grooves (in the order I, IJ, III, IV) and five arches appear between
the end of the second and beginning of the fourth day of incubaet Te teta mene em mena
BODY FORM IN THE CHICK 133
‘
.
‘
‘
D
A, 25 hours of incubation. B, 38 hours of
D, 68 hours of incubation. Compare
C
Fig. 85. — Growth of the chick embryo.
incubation. C, 48 hours of incubation.
Figs. 200, 206, 212, 218, respectively. A, B, approx. <9; C, D, approx. X4.
(After Duval.)
184 EMBRYONIC FORM AND EXTRA-EMBRYONIC STRUCTURES
tion. Only the first three clefts actually open into the fore-gut,
and these are soon closed again.
The tail arises from the backward growth of the tail fold but
never attains any great length.
The limb buds appear during the third day of incubation.
BODY FORM IN MAN. — Human embryologists distinguish three
periods during intra-uterine development: the period of the
ovum, from fertilization to germ-layer formation, two weeks;
the period of the embryo, until the embryo has assumed a definitely human appearance, the end of the second month; and
the period of the fetus. It is the second of these with which we
are concerned.
By the end of the third week the head fold is formed, and at the
fifth the tail fold is developed. Neural folds are formed and unite
. A B C
Fia. 86. — Growth of the humanembryo. A, neural folds (after Ingalls). B, neural
tube commencing, seven somites (after Payne). C, ten somites, (after Corner).
much as in the chick (Fig. 86). The primordia of eye, ear, and
nose-also_originate in a similar manner. Five visceral grooves
are formed, by the end of the fifth week, separating six visceral
_arches, but although the visceral pouches appear and unite with
the grooves, true visceral clefts are not formed. By the end of
the seventh week, the visceral grooves have disappeared. A
cephalic flexure appears in the fifth week. The neck (cervical)
THE DISPOSITION OF THE YOLK IN THE FROG 135
flexure develops in the week following and accelerates the disappearance of the visceral grooves.
A tail is developed from the tail fold which is quite prominent
during the six and seventh weeks of development but is overgrown
and resorbed during the eighth.
Limb buds make their appearance toward the end of the fifth
week.
B. THE YOLK SAC
Yolk sacs are found in the development of all large-yolked eggs,
among both anamniotes and amniotes. As the name implies,
this structure is a larger or smaller bag protruding from the body
and connected with it by a yolk stalk.
Origin and development. — The yolk sac develops from the
outer margin of the blastoderm which advances under the vitelline membrane and around the yolk mass until the yolk is completely enclosed (Fig. 82).
Function and fate. — It contains the yolk, which, in meroblastic cleavage, is not divided among the blastomeres. But it
plays a far more important part in development than simply
acting as a reservoir for food reserves. (It is lined with endoderm
just like that of the intestine, and is furnished with arteries, veins,
and capillaries, which make up the area vasculosa. The endodermal lining digests the yolk, and the vitelline veins carry the
digested food to the developing embryo. We may think of the
yolk sac as an extra-embryonic intestine. \ It is interesting to note
that in some viviparous elasmobranchs, like the dogfish, the yolk
sac continues to be of use, even after the yolk is consumed.
Pressed against the wall of the uterus it absorbs the uterine
“milk” which this organ secretes (much like a tertiary egg
envelope) and conveys it to the embryo through the vitelline
veins. A similar device is seen among the marsupials (page 144).
he yolk sac is usually drawn up into the body when the umbilicus
closes and is later resorbed.)
THE DISPOSITION OF THE YOLK IN THE FROG. — The frog has
no yolk sac, for the yolk is divided among the large blastomeres
which later make up the floor of the intestine. The mass of these
cells, however, creates a bulge on the ventral surface of the embryo (Fig. 84) which resembles externally a small sac.
136 EMBRYONIC FORM AND EXTRA-EMBRYONIC STRUCTURES
THE YOLK SAC OF THE CHICK. — The yolk sac of the chick is
formed by the advancing edge of the blastoderm. Looking down
on the blastoderm of the chick at the end of the first day of incubation (Fig. 79), one distinguishes a series of concentric rings.
Proceeding from the periphery inward, we note first the area vitellina externa, consisting of the margin of overgrowth and the zone
of junction (page 112). Then comes the area vitellina interna in
which we can distinguish the ectoderm and endoderm, the latter
closely applied to the yolk. Finally there is distinguished the
area vasculosa into which the mesoderm has pushed, splitting, as
it advances, into the somatic layer (next the ectoderm) and the
splanchnic layer (next the endoderm). Between the somatic and
splanchnic layers lies the exocoel (extra-embryonic coelom), as
the coelem is called when it extends beyond the boundaries of the
embryo. The blood vessels of the area vasculosa develop in the
splanchnic mesoderm. The exocoel separates the splanchnopleure (endoderm and splanchnic mesoderm) from the somatopleure (somatic mesoderm and ectoderm), so that it can be said
that the yolk sac of the chick consists of splanchnopleure. By the
end of the fourth day of incubation the yolk is completely covered except for a small area at the vegetal pole, known as the
yolk sac umbilicus (Fig. 89C, D). When the chick hatches, the
empty yolk sac still attached to the intestine is drawn into the
coelom and gradually disappears.
THE YOLK SAC OF MAN. — In man, as in other mammals, the
yolk sac arises in connection with gastrulation. The endoderm
growing out from the lower surface of the embryonic knob apparently reorganizes itself to form a very small gastrocoel or yolk
sac. The roof of this gastrocoel forms the roof of the digestive
canal; the anterior end is set off (with the head fold) to make the
fore-gut; the posterior end is set off (with the tail fold) to make
the hind-gut. The remainder constitutes the small yolk sac
(Fig. 86A). This sac is later squeezed between the amnion and
chorion (Fig. 90), and loses its connection with the intestine,
through the degeneration of the yolk stalk.
In other mammals (Fig. 68) the endoderm grows completely
around the interior of the trophoblast and forms a larger yolk sac.
In the mouse, where the embryonic knob hangs well down in the
cavity of the blastocyst, this results in the knob’s being covered
AMNION AND CHORION 141
Body
stalk
Amniotic
cavity f Allantois
Yolk
sac
Chorion
A
B
Chorion
laeve
~ Amniotic
cavity
Umbilical
cord
Allantoi¢e
stalk
“t)
Chorion ‘Placenta
frondosum D
Fig. 90. — Diagrams to show development of extra-embryonie structures in human
embryo. Four stages illustrated by sagittal sections. (After Corning.)
Trophoblast
Rauber’s cells
Ectoderm 4 — Endoderm
Amniotic
a folds
2 Primitive ,
streak — Betoderm
Fig. 91. — Amnion formation in the bat’s egg. A, primary amniotic cavity. B,
origin of amniotic folds. (After Van Beneden.)
142 EMBRYONIC FORM AND EXTRA-EMBRYONIC STRUCTURES
ate. The embryonic disc thus comes to form part of the
blastocyst wall.
The amnion and chorion are formed by amniotic folds (Fig. 91).
The internal limb of each fold is formed of somatopleure derived
from the embryonic disc and will form the amnion as in the chick.
The outer limb of each fold, however, is formed of ectoderm
derived from the trophoblast associated with somatic mesoderm
and gives rise to the chorion. The mesoderm growing out from
the primitive streak, and delaminating into somatic and splanchnic
layers, becomes the lining of the exocoel.
D. THE ALLANTOIS
The development of an amnion and chorion is always accompanied by the appearance of another sac, the allantois. This
extra-embryonic structure appears as an evagination from the
hind-gut and is therefore lined with splanchnopleure. It grows
out through the exocoel of the umbilical stalk into the exocoel of
the chorion, which it usually fills. It is filled with an allantoic
fluid which receives the nitrogenous wastes of the embryo in the
form of uric acid (Needham), and may be thought of in the first
instance as an extra-embryonic urinary bladder. As it fills the
chorion, its walls, being composed of splanchnic mesoderm in the
outer layer, easily fuse with the mesodermal layer of mesoderm
of the amnion, chorion, and yolk sac, whenever these structures
come together. Furthermore, it has an area vasculosa served by
the allantoic (umbilical) veins and arteries. This area vasculosa
when applied to the chorion is the region where the blood is nearest
to a source of atmospheric oxygen. Here an exchange of gases,
carbon dioxide for oxygen, takes place, and the allantois may be
considered as an extra-embryonic lung.
In the cleidoic egg of reptiles, birds, and egg-laying mammals,
the allantois also takes part in the formation of an albumen sac
wherein this material is digested. In the marsupials and placental
mammals it contributes to the formation of a placenta (hemiplacenta in marsupials) whereby digested food is obtained from the
maternal circulation. These functions of the placenta will be
discussed in the sections following.
ALLANTOIS OF THE CHICK. — The allantois (Fig. 92) arises towards the end of the third day as an evagination from the floor of
ALLANTOIS OF THE CHICK 143
the hind-gut. It grows out between the yolk and the wall of the
subcaudal pocket into the exocoel (Fig. 89B). Here it expands
greatly until by the end of the ninth day it has filled the entire
exocoel. Its outer wall unites with the chorion (Fig. 89C) to form
a chorio-allantois, its inner wall unites with the amnion above
and the yolk sac below.
Now the chorion, carrying with it an inner fold of allantois,
grows down beyond the yolk-sac umbilicus (page 136), and around
Yolk sac
Area
vasculosa
Allantois
Fia. 92. — The embryo chick and its extra-embryonic structures on the sixth day of
incubation. X13. (After Duval.)
the mass of albumen, which has become more viscous through the
loss of water and is displaced towards the lower side of the egg.
The albumen is enclosed in a double-walled sac of chorion with
the allantois between the two walls of the sac (Fig. 89D). The
layer next to the albumen is the ectoderm of the chorion, but
the mesoderm of the allantois supplies the blood vessels. It is
interesting to observe that it is the ectoderm of the albumen sac
which absorbs the albumen, whereas in the yolk sac it is the
endoderm which carries on this function.
By the twelfth day of incubation the albumen sac is closed
except at the yolk-sac umbilicus where it has an open connection
144. EMBRYONIC FORM AND EXTRA-EMBRYONIC STRUCTURES
with the yolk sac. On the sixteenth day the albumen is consumed.
On the seventeenth the yolk-sac umbilicus closes by the constriction of a ring of mesoderm derived from the old edge of the blastoderm. The yolk sae with the remains of the albumen sac still
attached is retracted into the body cavity of the chick on the
nineteenth day of incubation, aided by contractions of the amnion
and the inner wall of the allantois.
ALLANTOIS OF MAN AND OTHER MAMMALS. — In most of the
mammals there is a well-developed allantois, arising like that of
the chick, growing into the exocoel, and uniting with the chorion
to participate in the formation of the placenta, but the human
allantois is rudimentary. It arises as a minute tubular evagination which develops from the endodermal roof of the gastrocoel
even before the formation of the tail fold. It grows out into the
body stalk, a mass of mesoderm connecting the embryo with the
chorion (Fig. 90) for a short distance, but never gets so far as the
chorion. However, the allantoic (umbilical) blood vessels continue down the body stalk to the chorion where they form a chorionic area vasculosa in the region of the developing placenta.
THE PLACENTA
Before discussing the human placenta it will be helpful to review the different types of placentation recognized in mammals.
Two types are distinguished according to the degree of union between the trophoblast and the lining of the uterus (mucosa); a
second basis of distinction is whether the wall of the allantois
comes in contact with the chorion or not.
Indeciduate type. — The first_type of placenta is called indeciduate. In this type, found in several groups particularly the
ungulates, the trophoblast is closely applied to the mucosa but
both retain their integrity. The blood vessels of the placenta
absorb food material excreted by the mucosa and exchange carbon
dioxide for oxygen by diffusion.
Marsupials. — Among the marsupials are found both nonallantoic and allantoic hemiplacentae. In the opossum, Didelphys (Fig. 938A), the enlarged yolk sac is pressed against the
trophoblast, which in turn is closely applied to the mucosa,
forming folds which project into depressions in the uterine wall.
The absorbed nutriment is conveved to the embrvo bv means of
INDECIDUATE TYPE 145
the area vasculosa of the yolk sac. In Perameles (Fig. 93B), an
allantoic hemiplacenta is formed by the union of the allantoic
sac with the trophoblast. Where this hemiplacenta touches the
Allantoic
cavity
Exocoel
cavity
Fig. 93.— The extra-embryonic structures of marsupials. Diagrammatic. <A,
Didelphys. B, Perameles. (After Jenkinson.)
mucosa the epithelium of the latter thickens and is invaded by
maternal capillaries. The trophoblast is said to be resorbed so
that the capillaries of the allantois come into intimate connection
with those of the uterus. It should be mentioned in this connection that Perameles also possesses a well-developed area vasculosa
Allantoic cavity
Amniotic
Exocoel cavity
Wall of
uterus
Yolk sac
Allantoie cavity
stalk
Fig. 94. — Diagram of extra-embryonic structures in the pig. (After Smith.)
on the yolk sac. It is very probable, therefore, that both yolk sac
and allantoic circulations are concerned with the nutrition of the
developing young.
Ungulates. — In the ungulates there is a well-developed allantoic placenta of the indeciduate type (Fig. 94). The blastocyst
elongates, and over its surface appear projections of the tropho146 EMBRYONIC FORM AND EXTRA-EMBRYONIC STRUCTURES
blast which contain a core of mesoderm. These projections, villi,
grow into corresponding depressions of the uterine wall, called
crypts. The allantois meantime has filled the exocoel, and capillaries from the allantoic arteries and veins penetrate the mesodermal cores of the villi. These capillaries are brought very near
those of the uterine wall, but the blood remains separated from
that of the mother by (1) the endothelial lining of the maternal
capillaries, (2) the connective tissue of the mucosa, (3) the
epithelium of the mucosa, (4) the trophoblast, (5) the mesoderm
of the villi, and (6) the endothelial lining of the fetal capillaries
(Fig. 99A, B). At birth the villi are pulled out of the crypts, and
the placenta, with the remaining embryonic membranes, is discharged as the “ after-birth.”
Deciduate type. — The second type of placentation is called
deciduate. In this type the trophoblast attacks the mucosa and
Amniotic
cavity
Exocoel
Wall of uterus
Fia. 95. — Diagram of extra-embryonic structures in the dog. Sagittal section.
(After Jenkinson.)
erodes part of the lining.. It is characteristic of the majority of
the clawed mammals (unguiculates) and primates. In the first
group the placenta is allantoic; in the primates, non-allantoic.
Carnivores. — In the carnivores (Fig. 95) is found a deciduate
placenta of the allantoic type. The blastocyst elongates although
not to the extent it does in the ungulates. During this time
PLACENTA OF MAN 147
the epithelium of the uterus is cast off. At the circular zone of
the uterus which is in contact with the equator of the blastocyst
the epithelium of the uterus fails to regenerate. Into this
zonary area grow the villi of the trophoblast which become penetrated by the allantoic capillaries. The villi send out branched
processes, each with its capillaries, which surround the maternal
capillaries. Thus the maternal blood is separated from that of
the fetus by (1) the endothelium of the maternal capillaries, (2) a
varying amount of maternal connective tissue, (3) the trophoblast, (4) a varying amount of chorionic connective tissue, and
(5) the endothelial lining of the fetal capillaries (Fig. 99C). At
birth a certain amount of maternal tissue is torn away with the
placenta.
PLACENTA OF MAN. — In the human placenta there is the most
intimate contact between the maternal and fetal circulation.
Amniotic
cavity (. (9 3 Allantoic cavity
ARAN
ANG v bp
\ QS! NC Kes LP Yolk sac
WON p ZA cavity
Exocoel a f
Fig. 96. — Diagram of extra-embryonic structures in man. (After Kolliker.)
The placenta is non-allantoic. It will be recalled that the embryonic knob retains its connection with the trophoblast as the
body stalk. Into the body stalk grows the small evagination
from the hind-gut which represents the endodermal lining of the
allantois (Fig. 90). It:never comes in contact with the trophoblast and soon degenerates. The limiting sulci of the amnion
approach each other and become the walls of the umbilical cord.
148
EMBRYONIC FORM AND EXTRA-EMBRYONIC STRUCTURES
This encloses (Fig. 96) the body stalk, yolk stalk, allantoic stalk,
as well as the two umbilical arteries and two umbilical veins which
Fia. 97. — Diagram to show the uterine deciduae
(After JXollmann.)
(human).
vessels.
grow out from the body
of the embryo towards
the trophoblast. These
umbilical blood vessels
represent the allantoic
vessels of all other amniotes. Later the umbilical veins fuse, and
all this tissue assumes
common connective tissue characteristics with
the exception of the
walls. of the blood vessels.
The deciduae. — It
will be remembered that
the blastocyst burrows
into the uterine wall,
eroding epithelium, connective tissue, and blood
As the embryo increases in size, this erosion continues
and the embryo sinks into the compact layer of the mucosa and
comes in contact with the
spongy layer. The mucosa
grows around the burrowing
embryo, shutting it off from
the cavity of the uterus.
There may now be distinguished (Fig. 97) three
regions in the mucosa: (1)
the decidua basalis, to which
the blastocyst is attached;
(2) the decidua capsularis,
which cuts off the blastocyst
from the uterine cavity;
and (8) the decidua vera,
Chorion
car
eS Seite
Fig. 98. — Human embryo 11 mm. in length,
about 6 weeks old, to show extra-embryonic
structures.
x1}. (After Arey.)
including the remainder of the uterine lining.
THE CHORION 149
The chorion. — The trophoblast, while entering the uterine
wall, becomes differentiated into an outer syncytial layer and an
inner cellular layer. During the process of implantation, nutrition is obtained by the syncytial layer, which sends out projections
or false villi into the maternal tissue. Thereafter mesodermal
cores grow into the false villi converting them into the true villi
which later receive capillaries from the umbilical blood vessels.
Chorionic _—
epithelium ~yu gy Se
\ Chorionic
Uterine PS ») epithelium
epithelium Bt (| Uterine
. ay) E ~ epithelium
Uterine cs VN Uterine
capillary ~~" =~ + SS capillary
Chorionic
~~ epithelium
Uterine Villus
capillary
Lacuna
=— Decidua
Sy
Fig. 99. — Sections through placentae of A, pig; B, cow; C, cat; and D, human.
(Semi-diagrammatic after Grosser.)
Some of these bore into the uterine wall to become fixation villi.
The others, losing their syncytial layer, remain in the space between the trophoblast of the chorion and the maternal tissue as
nutrition villi (Fig. 98). These are bathed in maternal blood
which is brought into the intervillous space and carried thence
by the eroded uterine capillaries. Only those villi which are in
150 EMBRYONIC FORM AND EXTRA-EMBRYONIC STRUCTUBES
contact with the decidua basalis persist; the others degener ite,
thus differentiating the chorion into the chorion frondosum, with
villi, and the chorion laeve, devoid of the same. In the human
Amnion 6,
Villi of
chorion
Decidua
basalis
Fused
decidua
vera and
capsularis
Fia. 100. — Diagram of fetus (near term) to show relationships of extra-embryonic
structures and deciduae. (After Ahlfeld.)
placenta the maternal blood is separated from the fetal blood
stream by only (1) the cellular layer of the trophoblast, (2) the
chorionic connective tissue of the villi, and (3) the endothelia of
the fetal capillaries (Fig. 99D).
SUMMARY 151
Parturition. — The history of the extra-embryonic structures
as well as that of the deciduae is terminated by birth (parturition).
Owing to the absence of an allantoic sac the amnion enlarges to
fill the exocoel. Later, growth of the fetus results in pressing the
ch nfs eve and decidua capsularis against the decidua vera and
i, ating the uterine cavity (Fig. 100). At birth the placenta,
carrying with it the decidua basalis, and the attached membrane,
whfch represegee the fused amnion, chorion laeve, decidua capsularis, and decjghaa vera, are cast off as the caul or ‘‘after-birth.”
Y p
i SUMMARY
The method by which the external form of the vertebrate
embryo is assumed is closely connected on the one hand with the
shape of the gadtppla, and on the other with the presence or absence of certain extra-embryonic structures, the yolk sac, amnion,
chorion, and allantois.
With growth in length we associate the occurrence of metamerism, or the serial repetition of parts, and the formation of a head
and a tail. The paired limbs arise as buds.
The yolk sac is found only in embryos developing from extremely telolecithal eggs. It is lined with endoderm and functions as an extra-embryonic intestine. The splanchnic layer of
the mesoderm adjacent to it develops an area vasculosa which
conveys the digested yolk to the body of the embryo.
The amnion and the chorion arise typically from folds of somatopleure which fuse above the embryo, thus giving rise to an inner
membrane, the amnion, and an outer one, the chorion. The
amnion, lined with ectoderm internally, contains amniotic fluid
in which the embryo develops. The chorion, lined with somatic
mesoderm internally, contains the exocoel, a continuation of the
embryonic coelom. Neither of these membranes has any vascular
system of its own. They are found only in the development of
reptiles, birds, and mammals.
The allantois always develops in amniote embryos. It arises
as a ventral evagination of the hind-gut and typically grows out
into the exocoel which it completely fills. It functions as an
extra-embryonic bladder and lung, and because of its vascular
area may act (in connection with the chorion) as an organ of
nutrition, e.g., as an albumen sac.
- 152 EMBRYONIC FORM AND EXTRA-EMBRYONIC STRUCTURES
In mammals the blood vessels of the allantois invade the
chorion giving rise to the placenta, an organ where substances
may be exchanged by diffusion between the maternal and fetal
blood streams. The placenta is connected to the embryo by the
umbilical stalk, whose walls are formed by the amnion. In some
mammals, such as the carnivores and primates, parts of the
uterine wall, the deciduae, are concerned in the formation of the
placenta, and cast off with them at birth.
REFERENCES
Allen, FE. (ed.) 1932. Sex and Internal Seerctions.
Assheton, R. 1916. Growth in Length.
Hertwig, O. (ed.) 1906. Handbuch, cte., I, Chaps. 6-8.
Jenkinson, J. W. 1913. Vertebrate Embryology, Chaps. 8 and 9.
Kerr, J. GG. 1919. Textbook of Embryology, 1, Chaps. 7 and 8.
Lillie, F. R. 1916. The Development of the Chick, 2nd Ed., Chap. 7.
“Marshall, F. H. A. 1922. The Physiology of Reproduction, 2nd Td.
Meisenheimer, J. 1921--1930. Geschlecht und Geschlechter im Tierreiche.
Needham, J. 1931. Chemical !mbryology, III, Sections 20-22, 24, and Epilegomena.
CHAPTER VII
EXPERIMENTAL VERTEBRATE EMBRYOLOGY
Recent progress in vertebrate embryology has resulted so
largely from the application of the experimental method that even
the beginning student must acquaint himself with some of the
methods used and the results so far obtained. Within the limits
of this text only a few of the important fields in which the experimental method has been employed can be mentioned, and the
student must be referred to more extended treatises for further
information concerning this relatively new and important branch
of embryology.
The amphioxus and the frog have long been used by experimental embryologists, and more recently successful methods have
been devised for the experimental study of the developing egg of
the hen. Triton, in Germany, and Ambystoma, in this country,
are urodele amphibia whose eggs have been particularly favorable
for experimental embryology. The eggs of mammals, difficult to
obtain, and, so far, impossible to orient during the carly stages of
embryology, have been employed to a lesser extent.
The experimental embryologist alters the conditions under
which the egg develops in the hope of determining the factors involved in particular developmental processes. It is appropriate
that we conclude our study of carly embryology with a short
account of some of the experiments which bear directly on the
organization of the fertilized egg, on differentiation during cleavage and the formation of the germ layers, and on the direct
effects of environmental factors upon development.
A. THE ORGANIZATION OF THE FERTILIZED EGG
The fertilized egg, as we have seen, is the product resulting
from the union of two germ cells, the egg and the sperm. It
contains two pronuclei, of maternal and paternal origin, respectively, as well as a mass of cytoplasm which is almost ex
clusively maternal in origin. The nuclei contain the parental
153
154 EXPERIMENTAL VERTEBRATE EMBRYOLOGY
contributions of genes, the units which together determine the
hereditary characters of the developing individual. How the
genes produce their effects is not known, but it is certain that
they must act directly upon the cytoplasm. Accordingly we may
turn first to experiments dealing with the nuclei of the fertilized
egg, and second, to those concerned with the organization of the
cytosome itself.
1. Tue NuciueEar ORGANIZATION
The fact that the fertilized egg has the diploid number of
chromosomes and of genes, while the two gametes have the
haploid number, naturally leads to the question whether the
diploid number is necessary to continued development. A considerable number of experiments bear directly upon this question.
Artificial parthenogenesis. — The frog’s egg can be induced to
develop by puncture with a finely pointed glass needle (Loeb and
others). These artificially parthenogenetic eggs have given rise
to tadpoles and frogs. Apparently the number of chromosomes is
redoubled (diploid number), perhaps by a division of the chromosomes without a corresponding division of the cell. But the
genes are exclusively maternal in origin.
Irradiated sperm. — Sperms of the amphibian Triton, treated
to an appropriate dosage of radium emanations, have their nuclei
injured in such a way that they are unable to form normal pronuclei (Hertwig). But they retain their mobility and are able
to penetrate the egg and induce development. The sperm head
remains in the cytoplasm and passes to one or another of the
developing blastomeres but takes no part in mitosis and ultimately
degenerates. The number of chromosomes in the larval cells is
usually haploid, although redoubling may occur.
Irradiated eggs. — Eggs of Triton have also been irradiated to
kill the egg nucleus and then fertilized with normalsperms. These
eggs develop with the haploid number of chromosomes, showing
that either pronucleus, maternal or paternal, is adequate for
development.
Fertilization of enucleate eggs.— In some marine invertebrates, e.g., the sea urchin, the egg can be broken into fragments
by shaking. Naturally only one fragment will contain the
nucleus, but the enucleate fragments can be fertilized and will
FERTILIZATION OF ENUCLEATE EGGS 155
give rise to dwarf but otherwise normal larvae. This phenomenon
is known as merogony. A similar result can be obtained in
telolecithal vertebrate eggs such as those of Triton, where several
sperms normally enter the egg. After the entrance of the sperm
Egg nucleus
Sperm nuclei
Fig. 101. — The experimental production of haploid larvae in Triton. A, fertilized
egg with two sperm nuclei. B, same after constriction separating part of egg with
diploid nucleus (right) from part with haploid nucleus formed by supernumerary
sperm (left). C, showing relatively more advanced diploid embryo (right) and
less advanced haploid embryo (left). D, diploid larva. , haploid larva. (After
Spemann.)
it is possible to constrict the egg into two halves, by means of a
fine hair loop, in such a way that the female pronucleus lies in one
half (Fig. 101). This half will eventually have the diploid number of chromosomes, for a sperm pronucleus will conjugate with
156 EXPERIMENTAL VERTEBRATE EMBRYOLOGY
the egg pronucleus. The other half will have only the haploid
number. Both halves will develop into larvae, one of which will
have haploid and the other diploid nuclei.
Species hybrids. — Many experiments have been made in the
attempt to fertilize the egg of one species with a sperm from
another species. Often as in the teleost fish
f 5 \ (Moenkhaus), both pronuclei take part in the
oF i 2 subsequent cleavage, although frequently the
i Wry chromosomes from the two pronuclei (Fig. 102)
i i : form separate groups on the mitotic spindle
i il | | ; (gonomery). But in other cases Hertwig has
i | ft : shown that the male pronucleus takes no part in
i | § subsequent cleavages, so that the embryo reall
| : 1 y y
| Q develops parthenogenetically.
Jd! a i Natural interspecifie hybrids in both plant and
No animal kingdom are more common than for
Fie. 102. -—Chro- merly believed. Usually these interspecifie hymosomes in ana- brids are infertile, as the mule and many types
phase of first of hybrid bony fish, but they often grow to larger
cleavage of a hy- oto. ane . " : : . .
brid fish, Menidia 512° and are more active (hybrid vigor) than the
eggand Fundulus parents.
sperm, illustrat- |The equivalence of the pronuclei. — Although,
Chien eo Mncake as we have seen in Chapter IV, the pronuclei may
haus.) differ from one another in regard to individual
genes, the experiments mentioned above indi
cate that a single set of genes, paternal or maternal, is adequate
for the development of an egg. It must be recognized that the
experimental haploid animals are frequently less vigorous than
normal diploid forms.
2. THE ORGANIZATION OF THE CYTOPLASM
Polarity. — The primary expression of the egg’s organization
is the polarity already impressed upon it in the ovary (page 37).
That this polarity is itself not due to gravity is shown by the fact
that frog eggs which are kept in motion during early development
give rise to normal embryos (Morgan, Kathariner). But polarity
is not immutable, for many experiments in which the eggs of frogs
have been made to develop in an inverted position (Born, Pfliiger,
Morgan) show that the yolk streams down through the egg, and
ASYMMETRY 157
cleavage begins in the relatively yolk-free region which was
formerly the vegetal pole.
Gradient. — There scems to be good reason to suppose that the
polar axis represents a metabolic axial gradient (Child), for when
dilute solutions of lethal chemicals, e.g., potassium cyanide, are
applied to the frog’s egg (Bellamy), disintegration begins at the
animal pole and continues toward the vegetal pole, which is the
last part of the egg to be affected.
Cytoplasmic materials. — In some animals there seems to be a
definite stratification of materials in the egg along the polar axis,
but when this stratification is disturbed by whirling the eggs
about in a centrifuge, the eggs develop with the original polarity undisturbed. On the other hand, in telolecithal eggs like
that of the frog, centrifuging distorts the cytoplasmic framework
(Conklin).
Bilaterality. — The animal pole marks the anterior end of the
developing amphibian embryo. Its dorsal side is marked by the
gray crescent which appears on the side opposite the point of
entry of the sperm. Many observations (Jenkinson and others)
show that the point of entry marks a second dorso-ventral axis
and establishes the bilaterality of the developing embryo. But
in parthenogenetic eggs (when development is initiated by puncture) the point of entrance of the needle seems to have no constant
relation to the subsequent bilaterality of the egg. This would
indicate (Huxley and de Beer) that the egg has an underlying
bilaterality of its own which is not strong enough to withstand
the stronger stimulus afforded by the entrance of the sperm but
is apparent in parthenogenesis.
Bellamy has described a second axial gradient in the frog’s egg
shown by the action of potassium cyanide in which the high point
centers in the gray crescent. ‘This is the dorso-ventral axis of
the embryo, which is therefore normally determined by the
entrance point of the sperm.
Asymmetry. — The vertebrate embryo is not, strictly speaking,
bilaterally symmetrical. <A third axis or gradient from one side
to the other (usually left to right) is often apparent, as seen in the
development of the atriopore on the left side of the tadpole, the
fact that the heart of the chick develops on the right side, and the
fact that the head turns to the right in torsion. The stomach in
158 EXPERIMENTAL VERTEBRATE EMBRYOLOGY
all vertebrates is twisted to the left of the mid-line, and many
other examples might be mentioned. When this asymmetry is
reversed we have the phenomenon known as situs inversus, and
this condition can be reproduced experimentally by developing
the egg in a lateral temperature gradient and in other ways.
Thus the egg of the hen when overheated on the left side develops
situs inversus. It has been shown by Spemann that, when two
blastomeres which would ordinarily produce the right and left
sides of an embryo are separated by a hair loop, the left-hand
blastomere gives rise to a normally asymmetrical embryo, while
the right-hand blastomere gives rise either to an embryo with
normal asymmetry or to one with situs inversus.
These few examples of experiments on the fertilized egg indicate
that the egg is a complex system with a definite organization
indicated by its three axial gradients corresponding to its three
spatial dimensions, viz., an antero-posterior gradient (polarity),
a dorso-ventral gradient, and frequently a left-right gradient.
Furthermore, the system contains two complete sets of chromosomes and genes, either one of which is adequate in further
development.
B. ORGANIZATION OF THE EMBRYO DURING CLEAVAGE
Cell-lineage studies seemed to indicate that the dividing egg is
becoming a mosaic of blastomeres, cach set apart from the others
to form a specific portion of the embryo. Roux (1888) was the
first to realize that this might be tested experimentally. He
destroyed one of the 4-blastomeres of the frog’s egg and observed
that the other gave rise to a }-embryo, which later regenerated
the missing portion.
Later investigators devised a number of methods by which
blastomeres could be separated from each other, by shaking them,
cutting them apart with fine needles, constricting them with fine
threads, or placing them in artificial calcium-free sea water.
Blastomeres of marine eggs in this medium separate immediately,
and when returned to normal sea water continue their development without further separation (Herbst).
Regulation and mosaic eggs. — The results of their experimentation seemed to indicate that in some eggs, e.g., those of the
amphioxus (Fig. 103), either of the }-blastomeres might, when
REGULATION AND MOSAIC EGGS 159
separated, give rise to complete embryos (Wilson). These were
called regulation eggs and were said to have indeterminate cleavage. In others, such as Styela (Conklin) or the molluse Dentalium
(Wilson), the }-blastomeres give rise only to }-embryos (Fig. 103).
These were called mosaic eggs and were said to have determinate
cleavage.
Experiments on frog’s eggs had been inconclusive until recently
an improved technique has made it possible to separate blastomeres of the two-cell stage completely (Schmidt, 1980, 1933).
eee !
4
Fig. 103. — Diagram to show the fate of isolated blastomeres from mosaic and regulation eggs. A, mosaic egg of Dentalium. At left, a complete embryo produced
by entire egg: at right, partial embryos produced by the }-blastomeres when
artificially separated. B, regulation egg of Amphiorus. At left, embryo pro
duced by entire egg; at right, perfect dwarf embryos produced by 34-blastomeres.
(After Wilson.)
These experiments show that each of the 3-blastomeres can give
rise to a complete and perfect larva, provided only it contains
some of the gray crescent region. If, on the other hand, the egg
is so constricted that the first cleavage divides it into an animal
and a vegetal half, the animal half, containing the gray crescent,
Fig. 104. — Embryos arising from separated 3-blastomeres of the newt’segg. A, the
constriction separates the dorsal and ventral halves of the embryo. B, the constriction separates the right and left halves. C, perfect embryo arising from the
dorsal }-blastomere. D, mass of cells arising from ventral }-blastomere. E, two
perfect embryos arising from right and left 3-blastomeres respectively. (After
Spemann.) (160)
REGULATION AND MOSAIC EGGS 161
gives rise to a complete embryo, while the vegetal half, lacking
this region, is unable so to organize itself (Fig. 104). The importance of the gray crescent as the seat of the organizer is discussed on page 169. This seems to indicate that Roux’s results
were due to the presence of the injured blastomere inhibiting
complete development on the part of the uninjured blastomere.
In this connection it is interesting to note that Witschi (1927) has
Fig. 105. — Experiment demonstrating equality of nuclei formed during cleavage
(Triton). A ligature has been tied around the fertilized egg restricting the nucleus
to the right-hand portion. A, 16-cell stage, one nucleus passing into left-hand
portion. B, ligature tightened to separate the two portions. C, perfect embryos
formed by the separate portions. The nucleus of a ith-blastomere equivalent to
that of a complete zygote. (After Spemann.)
described a case in which two eggs were found in a single chorion.
Each of them was flattened on the side next to its neighbor and in
later development showed deficiencies in the corresponding region.
A beautiful demonstration that it is the cytoplasm and not the
nucleus which is concerned with differentiation during cleavage
is afforded by an instructive experiment of Spemann. If the egg
162 EXPERIMENTAL VERTEBRATE EMBRYOLOGY
is tied off before cleavage so that the nucleus is confined in one of
its halves (Fig. 105), all cleavage planes will be restricted to that’
half until eventually a cleavage plane, in this case at the fourth
cleavage, coincides with the plane of constriction. The nucleus
which enters the previously enucleate half is naturally one which
would serve a ;-blastomere. If the loop is now tightened until
the two haves are completely separated, the portion containing
this single nucleus will give rise to an embryo like the one from
the portion containing the fifteen nuclei and exactly like one
arising from a complete fertilized egg.
Pressure experiments. — Further examples of the regulative
power of some eggs may be seen in pressure experiments. If the
eggs of the frog are placed between glass plates during cleavage,
the third cleavage planes will be meridional instead of latitudinal,
and the fourth cleavage plane is latitudinal (Fig. 106). Now if
Fia. 106. — Diagram to show new relationship of blastomeres in frog’s egg resulting
from pressure during cleavage. A, normal 8-cell stage. B, 8-cell stage formed
under pressure. C, normal 16-cell stage. D, 16-cell stage formed under pressure.
Cells normally in animal hemisphere shown in stipple. (Suggested by a diagram
in Wells, Huxley and Wells.)
the eggs are released, their later development will be quite normal
even though the blastomeres are occupying positions unlike those
which they hold ordinarily.
Double embryos. — Still another example may be seen in the
eggs of Triton. If these are freed from the egg envelopes, the
blastomeres at the two-cell stage assume a dumb-bell shape.
Mangold discovered that, by placing one embryo in the two-cell
stage over another (Fig. 107), a double embryo resulted almost
exactly similar to a single embryo in the four-cell stage, and would
MONOVULAR TWINS AND MONSTERS 163
develop as such, provided only that the gray-crescent regions of
the two fell in the same plane. Otherwise double monsters resulted. We shall see the importance of the gray-crescent region
more clearly in a later section dealing
with the organizer which develops in
this region.
Chemo-differentiation. — It is
quite clear from these experiments
that the developing egg of the regulation type possesses a very great
plasticity in the early stages of development as compared to the mosaic
type illustrated by the egg of the
tunicates. It may be assumed that
the difference between these two lia. 107. — Double embryo arising
types lies in the time at which defi- from fusion of 2-cell stages of
. : Triton alpestris (above and below)
nite organ-forming substances are and Triton taeniatus (right and
segregated in the cytoplasm of the left) when laid over each other
egg. Conklin has demonstrated crosswise. Note that a new
that these rogions are segrogated {Hwee ip under ny al
after fertilization in the egg of the — geidel.)
tunicate, whereas in amphibian eggs
the only segregated region is that of the gray crescent. Huxley
(1924) has suggested the term chemo-differentiation for the segregation of organ-forming substances. A good example is seen in
the first division of the egg of Dentaliwm, the mollusc referred to
above where a polar lobe passes completely to one or the other of
the first 4-blastomeres. The cell receiving this lobe gives rise to
the apical organ, mesoderm, foot, and shell. Here the very first
division of the fertilized egg is determinate and dependent upon
the segregation of the organ-forming substance found in the polar
lobe (Fig. 108).
Monovular twins and monsters. — The extreme plasticity of
the vertebrate egg as seen by the fact that either two separate
individuals or duplicate monsters may be formed from the complete or partial separation of blastomeres suggests an explanation
of identical twins and the duplicate monsters which play so large
a part in the study of teratology. It is generally accepted that
identical, as distinguished from fraternal, twins are the product
164 EXPERIMENTAL VERTEBRATE EMBRYOLOGY
of a single fertilized egg which has divided completely during early
embryology, whereas the duplicate monsters, ranging from Siamese twins to monsters in which one individual is but a parasite
upon the body of the other, result from incomplete separation.
These identical twins are always of the same sex. Ordinary or
Ja Wy
Fig. 108. — Diagram to show possible distribution of organ-forming substances in
mosaic and regulation eggs. *A, immature egg. B, mature egg showing stratified
organ-forming substances. C, cleavage with equal division of chromosomes.
D, segregation of one organ-forming substance in left-hand 3-blastomere. EF,
equal division of organ-forming substances between the 3-blastomeres. (After
Wilson.)
fraternal twins (triplets, etc.) are supposed to be the product of
separate eggs which ovulated and were fertilized at about the
same time. Such twins are frequently of different sexes. In this
connection we might mention the free-martin, a sterile female
twinned with a male, not infrequent among cattle, and supposed
to result from one of two eggs which develop a common chorion
and therefore a common blood stream. It is supposed that a male
PLASTICITY (DEPENDENT DIFFERENTIATION) 165
hormone circulating in the common blood stream inhibits the
normal development of the female twin, so resulting in the
production of the sterile free-martin (Lillie).
C. ORGANIZATION OF THE EMBRYO DURING
GERM-LAYER FORMATION
The amphibian embryo is remarkably hardy and during the
early stages of development will endure very severe operations.
The work of Harrison in this country and of Spemann in Germany
has resulted in the perfection of a method of removing portions
of an embryo (micro-dissection) and grafting them into a new
environment, where they will continue development. The embryo from which the portion is removed is known as the donor;
the removed portion is called the graft (transplant); and, when
the portion removed is transplanted into another embryo, the
second embryo is termed the host.
The accompanying diagram (Fig. 109) will bring out some of
the methods which have been developed in transplantation experiments. Thus the graft may be transplanted into another portion
of the same embryo (homoplastic transplantation).!. It may be
transplanted into another embryo of the same species (heteroplastic transplantation). It may even be transplanted into an
embryo of another species or genus (xenoplastic transplantation).
Another method which has brought interesting results is to
transplant the removed portion into a nutrient medium and allow
it to develop there under sterile conditions (explantation). This
is also known as cultivation “ in vitro,” which means in glass.
Another ingenious technique is to transplant the graft into a
cavity of another embryo and allow it to develop there. The
example shown in the diagram is of a bit of embryonic tissue
transplanted into the eyeball of a tadpole, which acts as a nutrient
chamber. Hoadley and others have developed a technique of
grafting chick-embryo tissue from a donor to the chorio-allantois
of a host. Such a technique is called interplantation (implantation).
Plasticity (dependent differentiation). — In the amphibian egg,
which is of the regulation type, it has been demonstrated that the
1 Some" investigators use autoplastic=homoplastic; homoplastic=heteroplastic;
and heteroplastic = xenoplastic.
166 EXPERIMENTAL VERTEBRATE EMBRYOLOGY
presumptive organ regions of the blastula, (and until about the
middle of gastrulation) are quite plastic, i.e., can be transplanted
into other localities and will give rise to the organs appropriate
Cover glass
Fig. 109. — Diagrams to show different methods of transplantation, ete. A, homoplastic transplantation. B, heteroplastic transplantation (both donor and host
of same species). C, xenoplastic transplantation (donor and host of different
species). D, explantation (in vitro). E, interplantation. (Based on a diagram of
Dirken.)
to the new locality. Thus, material which is presumptive epidermis can be transplanted into a region where it will become
neural plate, mesoderm, orevenendoderm. Or on the other hand,
material which is presumptive endoderm can be made to develop
into ectoderm or mesoderm by transplantation. The only exception to this rule is the region where the dorsal lip is to form.
PLASTICITY (DEPENDENT DIFFERENTIATION) 167
This will never give rise to anything except dorsal lip and the
structures arising from the dorsal lip. This exception will receive
special attention later (page 169).
Very instructive experiments are those in which material is
transferred from a species with heavy pigmentation (Triton
taentatus) to one with light pigmentation (Triton cristatus).
Fig. 110. — Xenoplastic transplantation between Triton taeniatus (dark), the donor,
and Triton cristatus (light), the host to show early plasticity. A, immediately
after transplantation. B, the transplant developing in the gill region. C, the
gills of the transplant relatively more advanced. D, section through C in the gill
region. (After Spemann.)
Here the graft preserves its racial character of pigmentation while
otherwise conforming to the development of the host. Figure 110
illustrates such an example of xenoplastic transplantation. The
light-colored graft from T’. cristatus has developed into part of the
neural tube of the host, where it stands out by reason of its light
color. In the reciprocal transplantation (Fig. 111), the dark
graft from 7’. taeniatus has given rise to the right external gills of
the host.
168 EXPERIMENTAL VERTEBRATE EMBRYOLOGY
The loss of plasticity (self-differentiation). — After gastrulation
is well under way this plasticity seen in earlier stages is lost. The
various regions of the embryo have become determined and
A B
Fig. 111. — Reciprocal transplant to that shown in Fig. 110. Here T. cristatus
(light) is the donor and 7. taeniatus (dark) is the host. A, after transplantation.
B, the transplant developing in the neural plate (region of the brain). C, section
in later stage showing transplant developing in forebrain. (After Spemann.)
thenceforth will give rise only to the structures normally developing from them. In other words, the amphibian embryo does not
undergo chemo-differentiation until this time. From now on it
isa real mosaic. Figure 112 shows a neurula in which the various
Ear field Neural tube field
Eye field
Nose field
Lens field
Hindlimb field
Balancer field .
Heart jj,‘ Forelimb field
field fielg
Fig. 112. — Diagram of an amphibia neurula showing organ fields as determined by
transplantation experiments. (After Huxley and de Beer.)
organ fields are determined. If a bit of tissue is removed from
the eye-field and transferred to the flank of another neurula
(Fig. 113), it will give rise only to an eye, even in its new and
abnormal environment.
Similar experiments have been carried on with the chick (implantation on chorio-allantois), and it has been proved that the
THE ORGANIZER 169
eye-field, ear-field, limb-buds, and other regions will develop and
give rise only to the respective organs.
Very striking results have been obtained by implanting portions
of rat embryos on the chorio-allantois of the chick, and a considerable amount of self-differentiation has been demonstrated.
Donor Host
Fig. 113. — Self-differentiation in the toad Bombinator. A, donor in early neurula
stage showing region from which graft was taken. B, host in late neurula stage.
C, section through later embryo of host, showing graft forming optic cup in region
normally occupied by pronephros. (After Spemann.)
The organizer. — The loss of its early plasticity by the embryo
seems to be due to the presence of an organizer (organisator) as
discovered by Spemann. In the amphibian embryo this is the
dorsal-lip region, already mentioned. It will be recalled that this
region alone of the presumptive organ fields of early gastrulation
did not show the phenomenon of plasticity. Wherever it is
transplanted it will become dorsal lip. But the most striking
thing about this dorsal-lip region is that wherever it is transplanted
it will bring about involution, and will transform the surrounding
material into organ fields such as are ordinarily found about the
dorsal lip. In a word, the grafted dorsal lip organizes a new,
170 EXPERIMENTAL VERTEBRATE EMBRYOLOGY
secondary, embryo about itself, quite independent of the embryo
which is organized about the dorsal lip of the host (Fig. 114).
The organizer itself undergoes involution beneath the surface
of the host and becomes the notochord and the somite-mesoderm
of the secondary embryo. The other structures, such as neural
Primary ’ \ "2% es
a Yes notochord
Primary gut
Fig. 114. — Effect of transplanting organizer. A, dorsal view of host (Triton
taeniatus) in neurula stage. B, right side view at same stage showing secondary
neural plate induced by organizer (dorsal lip region) of the donor (T'’riton cristatus)
shown in white. C, later stage showing primary embryo in side view and secondary embryo in dorsal view. D, transverse section through C. (After Spemann
and Mangold.)
plate, eyes, ears, kidney, heart, etc., arise from host tissue which
has been brought under the influence of the organizer. Even
after gastrulation this influence is continued, as can be shown by
the following experiment. A bit of gastrocoel roof (notochord
and mesoderm, in the urodeles), when transplanted into the
GRAVITY (AND CENTRIFUGAL FORCE) 171
side of the gastrocoel, will induce the formation of neural folds
above it.
So great are the powers of induction possessed by the organizer
that it can cause presumptive ectoderm to become mesoderm or
endoderm, and conversely, presumptive mesoderm can be transformed into ectoderm.
It is noteworthy that the organizer can exert its influence even
in xenoplastic transplantation, e.g., the organizer from a toad can
induce the formation of a secondary embryo in.a newt. Apparently the effects of the organizer are physico-chemical in nature,
for the dorsal-lip region can be narcotized, boiled, or even dried,
and still induce the formation of a secondary embryo. It is
suggestive that bits of agar after being in contact for some time
with an organizer are themselves capable of producing induction.
There is reason to believe that the glycogen (animal starch)
content of the organizer has something to do with its effects, and
quite recently, it has been reported that cephalin will bring out
about the induction of a secondary embryo. Many parts of the
adult vertebrate are capable of bringing about induction, but in
the living embryo, the chemical substance responsible is found
only in the organizer itself.
D. ENVIRONMENTAL FACTORS IN DEVELOPMENT
Many experiments have been carried on in the attempt to find
the definite results produced on the developing embryo by
changes in the environment. These investigations have established normal limits of temperature, etc., within which development can be completed. Within these limits, although development may be altered as to rate, etc., it is nevertheless carried on
to a successful outcome. Beyond these limits the alterations are
so profound as to produce monsters or cause death. Among the
factors susceptible to experimental control are gravity, heat,
light, the chemical constitution of the environment, and food.
Gravity (and centrifugal force). — It has been remarked (page
156) that the original polarity of the egg is not due to any effect
of gravity. In telolecithal eggs, however, gravity may have some
effect on the course of development. Thus frog’s eggs when forcefully inverted may give rise to duplicate monsters. The hen’s egg
if not rotated at regular intervals fails to hatch. It has been
172 EXPERIMENTAL VERTEBRATE EMBRYOLOGY
shown (Dareste, 1877) that this is due to the failure of the yolk
sac to complete its development. It adheres to the allantois and
cannot be retracted into the body as in normal development.
The influence of gravity may be shown in an exaggerated
manner by prolonged centrifuging. It was found by O. Hertwig
that, if the frog’s egg is centrifuged during cleavage, the yolk is so
concentrated in the vegetal hemisphere that the cleavage planes fail to
cut through it and the end result is
meroblastic cleavage suggestive of
that seen in the chick (Fig. 115).
Undivided yolk Heat. — The rate of development
is directly affected by temperature.
Thus for the egg of the frog (Rana
fusca, Hertwig) the normal temperaFia. 115.— Vertical section ture is about 15°-16°C. From this
through blastula of a frog’s egg point up to about 20°—22° C., devel
following centrifuging. (After Hertwig.) opment continues normally; beyond this limit it is abnormal, death ensuing
rapidly at 30°C. Below 15° C., development is retarded progressively with the drop in temperature, and at 0°C. cleavage
ceases completely.
For the hen’s egg, Kaestner determined the optimum temperature for normal development to be between 35° and 39°C.
(95°-102° F.). The maximum temperature tolerated is 43°C.,
the minimum 28° C. (20°-21° C., Edwards).
Eggs of either frog or hen which have been exposed to extreme
heat or cold and then returned to the optimum temperature often
develop abnormally. A common type of monster is one in which
neural plate and notochord are split (spina bifida).
Very striking results have been obtained by subjecting the eggs
of the frog or the hen to a temperature gradient, that is, controlling the temperature so that one side is hotter or colder than the
other. If the gradient runs along the polar axis, and the greater
heat is applied to the animal pole, the result is that the embryos
and larvae have overlarge heads; if the higher temperature is applied to the vegetal pole, the head region is subnormal. When the
temperature gradient is applied laterally, the development of the
heated side proceeds more rapidly than that of the cooled side.
It may be concluded that, within the limits of toleration, development is accelerated by increased temperatures and retarded
by decreased temperatures.
Light and other forms of radiation. — In spite of a considerable
number of experiments designed to determine the effects of
definite intensities and wavelengths of light upon the developing
embryo, the results are as yet too inchoate to be discussed in an
elementary text.
Ultra-violet light, X-rays, and radium emanations in extreme
dosage cause the cessation of development. In smaller dosage,
they bring about anomalies (abnormalities in structure caused by
disturbances in development). It should be remembered that the
work of Miiller and others indicates that these agents accelerate
the rate of mutation of Drosophila genes, and so induce genetic
point mutations as well as developmental anomalies.
Chemical composition. — The chemical composition of the surrounding medium affects profoundly the nature of development.
The embryo cannot develop without oxygen, for it cannot live
without respiration. It has been pointed out by Morgan that
frog’s eggs in the very center of the egg mass often develop abnormally (spina bifida, etc.). And it has long been known that
the hen’s egg ceases development if the pores of the shell are
closed by water glass, varnish, or other agents.
Water, too, is an essential. The growth of the embryo depends upon the absorption of water, and all embryos must undergo
their development within a watery medium. Even the terrestrial
embryo has its private pond in the amnion. A slowing up in the
rate of development, accompanied by abnormalities and a large
percentage of deaths, results from incubating hen’s eggs in a
desiccator. The percentage of water in the frog’s egg increases
steadily during the first two weeks of development.
A very striking series of experiments was carried on by Herbst
on the development of the sea urchin in artificial sea waters which
had been made up omitting one after another of the elements
found in normal sea water. Jenkinson, summarizing the evidence
says:
“The experiments which we have been considering are unique
of their kind, and it is impossible to exaggerate their importance.
For, whatever may be the ultimate explanation of the facts, there can be no doubt whatever that the most complete demonstration
has been given of the absolute necessity of many of the elements
occurring in ordinary sea water, its normal environment, for the
proper growth and differentiation of the larva of the sea urchin.
Nor is this all. Some of the substances are necessary for one
part or phase of development, some for another, some from the
very beginning, others only later on. Thus potassium, magnesium, and a certain degree of alkalinity are essential for fertilization, chlorine and sodium for segmentation, calcium for the
adequate cohesion of the blastomeres, potassium, calcium and
the hydroxyl ion for securing the internal osmotic pressure
necessary for growth, while without the sulph-ion and magnesium
the due differentiation of the alimentary tract and the proper
formation of the skeleton cannot occur; the secretion of pigment
depends on the presence of some sulphate and alkalinity, the
skeleton requires calcium carbonate, cilia will only beat in an
alkaline medium containing potassium and magnesium, and
muscles will only contract when potassium and calcium are
there.”
The addition of chemicals to the medium has resulted in many
interesting disturbances in development. We can call attention
here to two only. In the sea urchin it was found that the addition of lithium salts to sea water caused the embryo to undergo
a very curious form of gastrulation, in which the endoderm and
mesoderm were evaginated instead of being invaginated (Herbst).
Such an embryo is called an exogastrula.
Quite recently, Holtfreter (1933) has induced exogastrulation
in the egg of Triton by removing the egg envelopes and placing
the developing egg in weak Ringer’s salt solution. In the cases
where development continued for some length of time (Fig. 116),
it was discovered that the embryo developed in two parts, an
ectodermal portion with no differentiation, connected by a narrow
isthmus to a mesendodermal portion in which differentiation
proceeded, but inan abnormal fashion. The embryo is inside-out.
The mesendodermal portion of the exo-embryo develops a typical
notochord, somites, kidney, gonad, a heart (empty), and a digestive tube, in which all the typical regions are indicated, including
visceral pouches. These results confirm those of transplantation
and explantation experiments discussed in an earlier section.
FOOD (INCLUDING HORMONES AND VITAMINS) 175
Food (including hormones and vitamins).— The amount
and kind of food supplied to the developing young naturally
affect the subsequent development. ‘Thus, if frog tadpoles are
fed on an exclusively vegetarian diet, the intestine becomes much
longer than when an exclusively meat diet is offered. Specific
foods often result in equally definite changes in the body. Thus Gudernatsch discovered that frog tadpoles fed on thyroid tissue
grew less rapidly but underwent metamorphosis much more
rapidly than the controls. Thymus-fed tadpoles, on the other
hand, had a retarded metamorphosis accompanied by excessive
growth. Later investigations indicate that the effects of thyroid
are due to a hormone formed by this gland (thyroxin), which is a
definite factor in bringing about amphibian metamorphosis.
Fig. 116. — Exogastrulation in Ambystoma. A, B, exogastrulae showing direction
of displacements during exogastrulation, compare Fig. 74. C, section of later
exo-embryo. (After Holtfreter in Huxley and de Beer.)
It is interesting to note that by the use of thyroid or thyroxin
the Mexican axolotl (Fig. 117) may be induced to undergo
metamorphosis, when it becomes a normal Ambystoma tigrinum.
Otherwise the axolotl becomes sexually mature in the larval
condition (neoteny), and was, therefore, long thought to be a
separate species.
In this connection we may refer briefly to the many experiments dealing with the effects of the various endocrine glands
when given as food or as transplants and the effects produced
when these glands are removed at their first appearance (extirpation). Without going into details, for the results of these experiments are sometimes ambiguous, we may say only that they
Fig. 117. — Metamorphosis in Ambystoma. A, neotenic larva (axolotl). B, metamorphosed adult. (After Diirken.)
indicate the importance of hormones in embryonic as well as in
adult life.
The rdle of the vitamins in the metabolism of the embryo is too
little understood at the present time for us to do more than allude
to this subject. Vitamin E is often called the anti-sterility
vitamin because its absence from the diet results in loss of the
reproductive power. Adamstone (1931) in this laboratory has
shown that the chick embryo produced by hens on a vitamin-Efree diet dies early in development following extensive disturbances in the blood-vascular system.
REFERENCES 177
SUMMARY
Experimental embryology demonstrates that development is
epigenetic. Given a suitable inheritance of genes, and a favorable
environment, development proceeds normally through stages of
increasing complexity. Any alteration, either in the genetic
complex or in the factors of the environment, will bring about
alterations in development.
The fertilized egg shows a definite organization as seen in its
polarity and symmetry. These seem to be the expression of
axial gradients. Sooner or later the cytoplasm of the egg undergoes chemo-differentiation and develops organ-forming substances
— sooner in mosaic eggs, later in regulation eggs.
Cleavage segregates the organ-forming substances as they are
differentiated, with the result that the isolated blastomeres of
mosaic eggs have a limited potency, those of regulation eggs have
a greater potency.
During germ-layer formation, the presumptive organ regions
are segregated into the different germ layers. Among the vertebrates this reorganization is effected by an organizer, which in
the frog is associated with the dorsal lip of the blastopore, and in
the chick with the homologous primitive streak.
Even in regulation eggs a mosaic stage is established during
germ-layer formation. The different organ fields are now determined, the earlier plasticity disappears, and each field is capable
only of self-differentiation.
REFERENCES
Allen, E. (ed.) 1932. Sex and Internal Secretion.
Bertalanffy, L. von, and Woodger, J. H. 1933. Modern Theories of Development.
Brachet, A. 1931. L’oeuf et les factors de l’ontogénése.
Brambell, F. W. R. 1930. The Development of Sex in Vertebrates.
de Beer, G. R. 1926. Introduction to Experimental Embryology.
Child, C. M. 1915. Individuality in Organisms.
Duesberg, J. 1926. L’oeuf et ses localisations germinales.
Dirken, B. 1932. Experimental Analysis of Development (trans).
Fauré-Fremiet, M. E. 1925. La cinétique du développement.
Huxley, J.S., and de Beer,G.R. 1934. The Elements of Experimental Embryology.
Jenkinson, J. W. 1909. Experimental Embryology.
- 1917. Three Lectures on Experimental Embryology.
Korschelt, E. 1927-1931. Regeneration and Transplantation.
Morgan, T. H. 1928. Experimental Embryology.
1934. Embryology and Genetics.
Needham, J. 1932. Chemical Embryology.
Newman, H.H. 1923. The Physiology of Twinning.
“Russell, E.S. 1930. The Interpretation of Development and Heredity.
Schleip, W. 1929. Die Determination der Primitiventwicklung.
Weiss, J. 1930. Entwicklungsphysiologie der Tiere.
Wilson, E. B. 1925. The Cell in Development and Heredity, 3rd Ed.
Cite this page: Hill, M.A. (2024, April 30) Embryology Book - Introduction to Vertebrate Embryology 1935-2. Retrieved from https://embryology.med.unsw.edu.au/embryology/index.php/Book_-_Introduction_to_Vertebrate_Embryology_1935-2
- What Links Here?
- © Dr Mark Hill 2024, UNSW Embryology ISBN: 978 0 7334 2609 4 - UNSW CRICOS Provider Code No. 00098G