Difference between revisions of "The Works of Francis Balfour 3-19"

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Vide also Kolliker (No. 298), especially for the human and mammalian skull;  
Vide also Kolliker (No. 298), especially for the human and mammalian skull;  
Gotte (No. 296).  
Gotte (No. 296).
TJie Pectoral girdle.
Pisces. Amongst Fishes the pectoral girdle presents itself
in its simplest form in Elasmobranchii, where it consists of a
bent band of cartilage on each side of the body, of somewhat
variable form, meeting and generally uniting with its fellow
ventrally. Its anterior border is in close proximity with the
last visceral arch, and a transverse ridge on its outer and
posterior border, forming the articular surface for the skeleton
of the limb, divides it into a dorsal part, which may be called
the scapula, and a ventral part which may be called the
In all the remaining groups of Fishes there is added to the
cartilaginous band, which may wholly or partially ossify, an
osseous support composed of a series of membrane bones.
In the types with such membrane bones the cartilaginous
parts do not continue to meet ventrally, except in the Dipnoi
where there is a ventral piece of cartilage, distinct from that
bearing the articulation of the limb. The cartilage is moreover
produced into two ventral processes, an anterior and a posterior,
below the articulation of the limb ; which may be called, in
accordance with Gegenbaur's nomenclature, the praecoracoid
and coracoid. Of these the praecoracoid is far the most
prominent, and in the majority of cases the coracoid can hardly
be recognised. The coracoid process is however well developed
in the Selachioid Ganoids, and the Siluroid Teleostei. In
Teleostei the scapular region often ossifies in two parts, the
smaller of which is named by Parker praecoracoid, though it is
quite distinct from Gegenbaur's praecoracoid. The membrane
bones, as they present themselves in their most primitive state
in Acipenser and the Siluroids, are dermal scutes embracing the
anterior edge of the cartilaginous girdle. In Acipenser there
are three scutes on each side. A dorsal scute known as the
supra-clavicle, connected above with the skull by the posttemporal ; a middle piece or clavicle, and a ventral or infraclavicle (inter-clavicle), which meets its fellow below.
In most Fishes the primitive dermal scutes have become
subdermal membrane bones, and the infra-clavicle is usually not
distinct, but the two clavicles form the most important part of
the membranous elements of the girdle. Additional membrane bones (post-clavicles) are often present behind the main
The development of these parts in Fishes has been but little
In Scyllium, amongst the Elasmobranchii, I find that each
half of the pectoral girdle develops as a vertical bar of cartilage
at the front border of the rudimentary fin, and externally to the
Before the tissue forming the pectoral girdle has acquired
the character of true cartilage, the bars of the two sides meet
ventrally by a differentiation in situ of the mesoblastic cells, so
that, when the girdle is converted into cartilage, it forms an
undivided arc, girthing the ventral side of the body. There is
developed in continuity with the posterior border of this arc on
the level of the fin a horizontal bar of cartilage, which is
continued backwards along the insertion of the fin, and, as will
be shewn in the sequel, becomes the metapterygium of the adult
(figs. 344, bp and 348, mp). With this bar the remaining skeletal
elements of the fin are also continuous.
The foramina of the pectoral girdle are not in the first
instance formed by absorption, but by the non-development of
the cartilage in the region of pre-existing nerves and vessels.
The development of these parts in Teleostei has been recently investigated
by 'Swirski (No. 472) who finds in the Pike (Esox) that the cartilaginous
pectoral girdle is at first continuous with the skeleton of the fin. It forms
a rod with a dorsal scapular and ventral coracoid process. An independent
mass of cartilage gives rise to a prascoracoid, which unites with the main
mass, forming a triradiate bar like that of Acipenser or the Siluroids.
The coracoid process becomes in the course of development gradually
'Swirski concludes that the so-called praecoracoid bar is to some extent
a secondary element, and that the coracoid bar corresponds to the whole of
the ventral part of the girdle of Elasmobranchii, but his investigations do
not appear to me to be as complete as is desirable.
Amphibia and Amniota. The pectoral girdle contains a
more or less constant series of elements throughout the
Amphibia and Amniota ; and the differences in structure
between the shoulder girdle of these groups and that of Fishes
are so great that it is only possible to make certain general
statements respecting the homologies of the parts in the two
sets of types.
The generally accepted view, founded on the researches of
Parker, Huxley, and Gegenbaur, is to the effect that there is a
primitively cartilaginous coraco-scapular plate, homologous with
that in Fishes, and that the membrane bones in Fishes are
represented by the clavicle and inter-clavicle in the Sauropsida
and Mammalia, which are however usually admitted to be
absent in Amphibia. These views have recently been challenged
by Gotte (No. 466) and Hoffmann (No. 467), on the ground of
a series of careful embryological observations ; and until the
whole subject has been worked over by other observers it does
not seem possible to decide satisfactorily between the conflicting
views. It is on all hands admitted that the scapulo-coracoid
elements of the shoulder girdle are formed as a pair of cartilaginous plates, one on each side of the body. The dorsal half
of each plate becomes the scapula, which may subsequently
become divided into a supra-scapula and scapula proper ; while
the ventral half forms the coracoid, which is not always separated
from the scapula, and is usually divided into a coracoid proper,
a praecoracoid, and an epicoracoid. By the conversion of parts
of the primitive cartilaginous plates into membranous tissue
various fenestrae may be formed in the cartilage, and the bars
bounding these fenestrae both in the scapula and coracoid
regions have received special names ; the anterior bar of the
coracoid region, forming the praecoracoid, being especially
important. At the boundary between the scapula and the
coracoid, on the hinder border of the plate, is placed the glenoid
articular cavity to carry the head of the humerus.
The grounds of difference between Gotte and Hoffmann and
other anatomists concern especially the clavicle and inter-clavicle.
The clavicle is usually regarded as a membrane bone which may
become to some extent cartilaginous. By. the above anatomists,
and by Rathke also, it is held to be at first united with the
coraco-scapular plate, of which it forms the anterior limb, free
ventrally, but united dorsally with the main part of the plate ;
and Gotte and Hoffmann hold that it is essentially a cartilage
bone, which however in the majority of the Reptilia ossifies
directly without passing through the condition of cartilage.
The interclavicle (episternum) is held by Gotte to be
developed from a paired formation at the free ventral ends of
the clavicles, but he holds views which are in many respects
original as to its homologies in Mammalia and Amphibia. Even
if Gotte's facts are admitted, it does not appear to me necessarily
to follow that his deductions are correct. The most important
of these is to the effect that the dermal clavicle of Pisces has no
homologue in the higher types. Granting that the clavicle in
these groups is in its first stage continuous with the coracoscapular plate, and that it may become in some forms cartilaginous before ossifying, yet it seems to me all the same quite
possible that it is genetically derived from the clavicle of Pisces,
but that it has to a great extent lost even in development its
primitive characters, though these characters are still partially
indicated in the fact that it usually ossifies very early and
partially at least as a membrane bone 1 .
In treating the development of the pectoral girdle systematically it will be
convenient to begin with the Amniota, which may be considered to fix the
nomenclature of the elements of the shoulder girdle.
1 The fact of the clavicle going out of its way, so to speak, to become cartilaginous
before being ossified, may perhaps be explained by supposing that its close connection
with the other parts of the shoulder girdle has caused, by a kind of infection, a change
in its histological characters.
Lacertilia. The shoulder girdle is formed as two membranous plates,
from the dorsal part of the anterior border of each of which a bar projects
(Rathke, Gotte), which is free at its ventral end. This bar, which is usually
(Gegenbaur, Parker) held to be independent of the remaining part of the
shoulder girdle, gives rise to the clavicle and interclavicle. The scapulocoracoid plate soon becomes cartilaginous, while at the same time the clavicular bar ossifies directly from the membranous state. The ventral ends
of the two clavicular bars enlarge to form two longitudinally placed plates,
which unite together and ossify as the interclavicle.
Parker gives a very different account of the interclavicle in Anguis. He
states that it is formed of two pairs of bones 'strapped on to the antero-inferior part of the prassternum,' which subsequently unite into one.
Chelonia. The shoulder girdle of the Chelonia is formed (Rathke) of
a triradiate cartilage on each side, with one dorsal and two ventral limbs.
It is admitted on all hands that the dorsal limb is the scapular element,
and the posterior ventral limb the coracoid ; but, while the anterior ventral
limb is usually held to be the praecoracoid, Gotte and Hoffmann maintain
that, in spite of its being formed of cartilage, it is homologous with the
anterior bar of the primitive shoulder-plates of Lacertilia, and therefore the
homologue of the clavicle.
Parker and Huxley (doubtfully) hold that the three anterior elements of
the ventral plastron (entoplastron and epiplastra) are homologous with the
interclavicle and clavicles, but considering that these plates appear to belong
to a secondary system of dermal ossifications peculiar to the Chelonia, this
homology does not appear to me probable.
Aves. There are very great differences of view as to the development
of the pectoral arch of Aves.
About the presence in typical forms of the coraco-scapular plate and two
independent clavicular bars all authors are agreed. With reference to the
clavicle and interclavicle Parker (No. 468) finds that the scapular end of the
clavicle attaches itself to and ossifies a mass of cartilage, which he regards
as the mesoscapula, while the interclavicle is formed of a mass of tissue between the ends of the clavicles where they meet ventrally, which becomes
the dilated plate at their junction.
Gegenbaur holds that the two primitive clavicular bars are simply clavicles, without any element of the scapula ; and states that the clavicles are
not entirely ossified from membrane, but that a delicate band of cartilage
precedes the osseous bars. He finds no interclavicle.
Gotte and Rathke both state that the clavicle is at first continuous with
the coraco-scapular plate, but becomes early separated, and ossifies entirely
as a membrane bone. Gotte further states that the interclavicles are formed
as outgrowths of the median ends of the clavicles, which extend themselves
at an early period of development along the inner edges of the two halves of
the sternum. They soon separate from the clavicles, which subsequently
meet to form the furculum ; while the interclavicular rudiments give rise, on
the junction of the two halves of the sternum, to its keel, and to the ligament
connecting the furculum with the sternum. The observations of Gotte,
which tend to shew the keel of the sternum is really an interclavicle, appear
to me of great importance.
A prascoracoid, partially separated from the coracoid by a space, is present in Struthio. It is formed by a fenestration of a primitively continuous
cartilaginous coracoid plate (Hoffmann). In Dromaeus and Casuarius clavicles are present (fused with the scapula in the adult Dromaeus), though
absent in other Ratitae (Parker, etc.).
Mammalia. The coracoid element of the coraco-scapular plate is
much reduced in Mammalia, forming at most a simple process (except in the
Ornithodelphia) which ossifies however separately 1 .
With reference to the clavicles the same divergencies of opinion met with
in other types are found here also.
The clavicle is stated by Rathke to be at first continuous with the coracoscapular plate. It is however soon separated, and ossifies very early, in the
human embryo before any other bone. Gegenbaur however shewed that
the human clavicle is provided with a central axis of cartilage, and this observation has been confirmed by Kolliker, and extended to other Mammalia by
Gotte. The mode of ossification is nevertheless in many respects intermediate between that of a true cartilage bone and a membrane bone. The
ends of the clavicles remain for some time, or even permanently, cartilaginous, and have been interpreted by Parker, it appears to me on hardly
sufficient grounds, as parts of the mesoscapula and praecoracoid. Parker's
so-called mesoscapula may ossify separately. The homologies of the episternum are much disputed. Gotte, who has worked out the development of the
parts more fully than any other anatomist, finds that paired interclavicular
elements grow out backwards from the ventral ends of the clavicles, and
uniting together form a somewhat T-shaped interclavicle overlying the front
end of the sternum. This condition is permanent in the Ornithodelphia,
except that the anterior part of the sternum undergoes atrophy. But in the
higher forms the interclavicle becomes almost at once divided into three
parts, of which the two lateral remain distinct, while the median element
fuses with the subjacent part of the sternum and constitutes with it the presternum (manubrium sterni). If Gotte' s facts are to be trusted, and they
have been to a large extent confirmed by Hoffmann, his homologies appear to
be satisfactorily established. As mentioned on p. 563 Ruge (No. 438) holds
that Gotte is mistaken as to the origin of the presternum.
Gegenbaur admits the lateral elements as parts of the interclavicle, while
Parker holds that they are not parts of an interclavicle but are homologous
with the omosternum of the Frog, which is however held by Gotte to be a
true interclavicle.
1 This process, known as the coracoid process, is held by Sabatier to be the
pnecoracoid ; while this author also holds that the upper third of the glenoid cavity,
which ossifies by a special nucleus, is the true coracoid. The absence of a praecoracoid in the Ornithodelphia is to my mind a serious difficulty in the way of
Sabatier's view.
Amphibia. In Amphibia the two halves of the shoulder girdle are
each formed as a continuous plate, the ventral or coracoid part of which is
forked, and is composed of a larger posterior and a smaller anterior bar-like
process, united dorsally. In the Urodela the two remain permanently free
at their ventral ends, but in the Anura they become united, and the space
between them then forms a fenestra. The anterior process is usually (Gegenbaur, Parker) regarded as the praecoracoid, but Gotte has pointed out that
in its mode of development it strongly resembles the clavicle of the higher
forms, and behaves quite differently to the so-called praecoracoid of Lizards.
It is however to be noticed that it differs from the clavicle in the fact that it
is never segmented off from the coraco-scapular plate, a condition which has
its only parallel in the equally doubtful case of the Chelonia. Parker holds
that there is no clavicle present in the Amphibia, while Gegenbaur maintains
that an ossification which appears in many of the Anura (though not in the
Urodela) in the perichondrium on the anterior border of the cartilaginous
bar above mentioned is the representative of the clavicle. Gotte's observations on the ossification of this bone throw doubt upon this view of Gegenbaur ; while the fact that the cartilaginous bar may be completely enclosed
by the bone in question renders Gegenbaur's view, that there is present both
a clavicle and prsecoracoid, highly improbable.
No interclavicle is present in Urodela, but in this group and in a number
of the Anura, a process grows out from the end of each of the bars (praecoracoids) which Gotte holds to be the clavicles. The two processes unite
in the median line, and give rise in front to the anterior unpaired element of
the shoulder girdle (omosternum of Parker). They sometimes overlap the
epicoracoids behind, and fusing with them bind them together in the median
line. Parker who has described the paired origin of the so-called omosternum,
holds that it is not homologous with the interclavicle, but compares it with
his omosternum in Mammals.
(463) Bruch. " Ueber die Entwicklung der Clavicula und die Farbe des
Blutes. " Zeit.f. wiss. Zool., \\. 1853.
(464) A. Duges. " Recherches sur 1'osteologie et la myologie des Batraciens a
leurs differens ages." Memoires des savants etrang. Academic royale des sciences de
Finstitut de France^ Vol. vi. 1835.
(465) C. Gegenbaur. Untersuchungen zur vergleichenden Anatomie der Wirbelthiere, 2 Heft. Schultergiirtel der Wirbelthiere. Bmstflosse der Fische. Leipzig,
(466) A. Gotte. "Beitrage z. vergleich. Morphol. d. Skeletsystems d. Wirbelthiere : Brustbien u. Schultergiirtel." Archivf. mikr, Anat. Vol. xiv. 1877.
(467) C. K. Hoffmann. "Beitrage z. vergleichenden Anatomic d. Wirbelthiere." Niederlandisches Archivf. ZooL,Vol.v. 1879.
(468) W. K. Parker. "A Monograph on the Structure and Development of the
Shoulder-girdle and Sternum in the Vertebrata." Ray Society, 1868.
(469) H. Rathke. Ueber die Entwicklung der Schildkrbten. Braunschweig,
(470) H. Rathke. Ueber den Bau und die Entwicklung des Brustbeins der
Saurier, 1853.
(471) A. Sabatier. Comparaison des ceinfures et des membres antMeurs et posttrtturs d. la Serie d. Vertttrh. Montpellier, 1880.
(472) Georg 'Swirski. Untersuch. iib. d. Entwick. d. Schultergiirtels n. d.
Skelets d. Brustflosse d. Hechts. Inaug. Diss. Dorpat, 1880.
Pelvic girdle.
Pisces. The pelvic girdle of Fishes is formed of a cartilaginous band, to the outer and posterior side of which the basal
element of the pelvic fin is usually articulated. This articulation
divides it into a dorsal iliac, and ventral pubic section. The iliac
section never articulates with the vertebral column.
In Elasmobranchii the two girdles unite ventrally, but the
iliac section is only slightly developed. In Chimaera there is a
well developed iliac process, but the pubic parts of the girdle
are only united by connective tissue.
In the cartilaginous Ganoids the pelvic girdle is hardly to be
separated from the skeleton of the fin. It is not united with its
fellow, and is represented by a plate with slightly developed
pubic and iliac processes.
In the Dipnoi there is a simple median cartilage, articulated
with the limb, but not provided with an iliac process. In bony
Ganoids and Teleostei there is on each side a bone meeting its
fellow in the ventral line, which is usually held to be the rudiment of the pelvic girdle ; while Davidoff attempts to shew that
it is the basal element of the fin, and that, except in Polypterus,
a true pelvic girdle is absent in these types.
From my own observations I find that the mode of development of the pelvic girdle in Scyllium is very similar to that of
the pectoral girdle. There is a bar on each side, continuous on
its posterior border with the basal element of the fin (figs. 345
and 347). This bar meets and unites with its fellow ventrally
before becoming converted into true cartilage, and though the
iliac process (il) is never very considerable, yet it is better developed in the embryo than in the adult, and is at first directed
nearly horizontally forwards.
Amphibia and Amniota. The primitive cartilaginous pelvic
girdle of the higher types exhibits the same division as that of
Pisces into a dorsal and a ventral section, which meet to form
the articular cavity for the femur, known as the acetabulum.
The dorsal section is always single, and is attached by means
of rudimentary ribs to the sacral region of the vertebral column,
and sometimes to vertebrae of the adjoining lumbar or caudal
regions. It always ossifies as the ilium.
The ventral section is usually formed of two more or less
separated parts, an anterior which ossifies as the pubis, and a
posterior which ossifies as the ischium. The space between them
is known as the obturator foramen. In the Amphibia the two
parts are not separated, and resemble in this respect the pelvic
girdle of Fishes. They generally meet the corresponding elements
of the opposite side ventrally, and form a symphysis with them.
The symphysis pubis, and symphysis ischii may be continuous
(Mammalia, Amphibia).
The observations on the development of the pelvic girdle in
the Amphibia and Amniota are nearly as scanty as on those of
Amphibia. In the Amphibia (Bunge, No. 473) the two halves of
the pelvic girdle are formed as independent masses of cartilage, which
subsequently unite in the ventral line.
In the Urodelous Amphibia (Triton) each mass is a simple plate of
cartilage divided into a dorsal and ventral section by the acetabulum.
The ventral parts, which are not divided into two regions, unite in a
symphysis comparatively late.
The dorsal section ossifies as the ilium. The ventral usually contains
a single ossification in its posterior part which forms the ischium ; while
the anterior part, which may be considered as representing the pubis,
usually remains cartilaginous ; though Huxley (No. 475) states that it has
a separate centre of ossification in Salamander, which however does not
appear to be always present (Bunge). There is a small obturator foramen
between the ischium and pubis, which gives passage to the obturator nerve.
It is formed by the part of the tissue where the nerve is placed not becoming converted into cartilage.
There is a peculiar cartilage in the ventral median line in front of the
pubis, which is developed independently of and much later than the true
parts of the pelvic girdle. It may be called the praepubic cartilage.
Reptilia. In Lacertilia the pelvic girdle is formed as a somewhat
triradiate mass of cartilage on each side, with a dorsal (iliac) process, and two
ventral (pubic and ischiad) processes. The acetabulum is placed on the
outer side at the junction of the three processes, each of which may be
considered to have a share in forming it. The distal ends of the pubis
and ischium are close together when first formed, but subsequently separate.
Each of them unites at a late stage with the corresponding process of the
opposite side in a ventral symphysis. A centre of ossification appears in
each of the three processes of the primitive cartilage.
Aves. In Birds the parts of the pelvic girdle no longer develop as a
continuous cartilage (Bunge). Either the pubis may be distinct, or, as in the
Uuck, all the elements. The ilium early exhibits a short anterior process,
but the pubis and ischium are at first placed with their long axes at right
angles to that of the ilium, but gradually become rotated so as to lie parallel with it, their distal ends pointing backwards, and not uniting ventrally
excepting in one or two Struthious forms.
Mammalia. In Mammalia the pelvic girdle is formed in cartilage
as in the lower forms, but in Man at any rate the pubic part of the cartilage is formed independently of the remainder (Rosenberg). There are
the usual three centres of ossification, which unite eventually into a single
bone the innominate bone. The pubis and ischium of each side unite with
each other ventrally, so as completely to enclose the obturator foramen.
Huxley holds that the so-called marsupial bones of Monotremes and
Marsupials, which as shewn by Gegenbaur (No. 474) are performed in cartilage, are homologous with the praepubis of the Urodela ; but considering
the great gap between the Urodela and Mammalia this homology can only
be regarded as tentative. He further holds that the anterior prolongations
of the cartilaginous ventral ends of the pubis of Crocodilia are also structures of the same nature.
(473) A. Bunge. Untersuch. z, Entwick. d. Beckengiirtels d. Amphibien,
Reptilien u. Vogel, Inaug. Diss. Dorpat, 1880.
(474) C. Gegenbaur. " Ueber d. Ausschluss des Schambeins von d. Pfanne
d. Hiiftgelenkes." Morph. Jahrbuch, Vol. II. 1876.
(475) Th. H. Huxley. "The characters of the Pelvis in Mammalia, etc."
Proc. of Roy. Soc., Vol. xxvm. 1879.
(476) A. Sabatier. Comparaison des ceintures et des membres anterieurs et
posterieurs dans la Serie d. Vertebrcs. Montpellier, 1880.
Comparison of Pectoral and Pelvic girdles.
Throughout the Vertebrata a more or less complete serial
homology may be observed between the pectoral and pelvic
In the cartilaginous Fishes each girdle consists of a continuous
band, a dorsal and ventral part being indicated by the articulation
of the fin ; the former being relatively undeveloped in the pelvic
LIMBS. 609
girdle, while in the pectoral it may articulate with the vertebral
column. In the case of the pectoral girdle secondary membrane
bones become added to the primitive cartilage in most Fishes,
which are not developed in the case of the pelvic girdle.
In the Amphibia and Amniota the ventral section of each
girdle becomes divided into an anterior and a posterior part, the
former constituting the praecoracoid and pubis, and the latter the
coracoid and ischium ; these parts are however very imperfectly
differentiated in the pelvic girdle of the Urodela. The ventral
portions of the pelvic girdle usually unite below in a symphysis.
They also meet each other ventrally in the case of the pectoral
girdle in Amphibia, but in most other types are separated by
the sternum, which has no homologue in the pelvic region, unless
the praepubic cartilage is to be regarded as such. The dorsal or
scapular section of the pectoral girdle remains free ; but that of
the pelvic girdle acquires a firm articulation with the vertebral
If the clavicle of the higher types is derived from the membrane bones of the pectoral girdle of Fishes, it has no homologue
in the pelvic girdle ; but if, as Gotte and Hoffmann suppose, it is
a part of the primitive cartilaginous girdle, the ordinary view as
to the serial homologies of the ventral sections of the two girdles
in the higher types will need to be reconsidered.
It will be convenient to describe in this place not only the
development of the skeleton of the limbs but also that of the
limbs themselves. The limbs of Fishes are moreover so different
from those of the Amphibia and Amniota that the development
of the two types of limb may advantageously be treated separately.
In Fishes the first rudiments of the limbs appear as slight
longitudinal ridge-like thickenings of the epiblast, which closely
resemble the first rudiments of the unpaired fins.
These ridges are two in number on each side, an anterior
immediately behind the last visceral fold, and a posterior on the
level of the cloaca. In most Fishes they are in no way connected, but in some Elasmobranch embryos, more especially in
Torpedo, they are connected together at their first development
B. in. 39
by a line of columnar epiblast cells 1 . This connecting line of
columnar epiblast is a very transitory structure, and after its
disappearance the rudimentary fins become more prominent,
consisting (fig. 343, &) of a projecting ridge both of epiblast and
mesoblast, at the outer edge of which is a fold of epiblast only,
which soon reaches considerable dimensions. At a later stage
the mesoblast penetrates into this fold and the fin becomes a
simple ridge of mesoblast, covered
by epiblast. The pectoral fins
are usually considerably ahead
of the pelvic fins in development.
For the remaining history it
is necessary to confine ourselves
to Scylliurn as the only type
which has been adequately
The direction of the original
ridge which connects the two fins
of each side is nearly though not
quite longitudinal, sloping somewhat obliquely downwards. It
thus comes about that the attachment of each pair of limbs is
somewhat on a slant, and that
the pelvic pair nearly meet each
other in the median ventral line
a little way behind the anus.
The elongated ridge, forming
the rudiment of each fin, gradually projects more and more, and
so becomes broader in proportion to its length, but at the same
time its actual attachment to the side of the body becomes
shortened from behind forwards, so that what was originally the
attached border becomes in part converted into the posterior
border. This process is much more completely carried out in
the case of the pectoral fins than in that of the pelvic, and the
changes of form undergone by the pectoral fin in its development may be gathered from figs. 344 and 348.
b. pectoral fin ; ao. dorsal aorta ;
cav. cardinal vein ; ua. vitelline artery ; u.v, vitelline vein ; al. duodenum ; /. liver ; sd. opening of segmented duct into the body cavity ;
mp. muscle plate ; ;. umbilical
1 I''. M. I'alfour. Monograph on Elasmobranfh l-'hhes, pp. 1012.
LIMBS. 6ll
Before proceeding to the development of the skeleton of
the fin it may be pointed out that the connection of the two
rudimentary fins by a continuous epithelial line suggests the
hypothesis that they are the remnants of two continuous lateral
fins 1 .
Shortly after the view that the paired fins were remnants of
continuous lateral fins had been put forward in my memoir on
Elasmobranch Fishes, two very interesting papers were published
by Thacker (No. 489) and Mivart (No. 484) advocating this
view on the entirely independent grounds of the adult structure
of the skeleton of the paired fins in comparison with that of the
unpaired fins 2 .
The development of the skeleton has unfortunately not been
as yet very fully studied. I have however made some investigations on this subject on Scyllium, and 'Swirski has also made
some on the Pike.
In Scyllium the development of both the pectoral and pelvic
fins is very similar.
In both fins the skeleton in its earliest stage consists of a bar
springing from the posterior side of the pectoral or pelvic girdle,
and running backwards parallel to the long axis of the body.
The outer side of this bar is continued into a plate which
1 Both Maclise arid Humphry {Journal of Anat. and Pkys., Vol. v.) had
previously suggested that the paired fins were related to the unpaired fins.
2 Davidoff in a Memoir (No. 477) which forms an important contribution to our
knowledge of the structure of the pelvic fins has attempted from his observations to
deduce certain arguments against the lateral fin theory of the limbs. His main
argument is based on the fact that a variable but often considerable number of the
spinal nerves in front of the pelvic fin are united, by a longitudinal commissure, with
the true plexus of the nerves supplying the fin. From this he concludes that the pelvic
fin has shifted its position, and that it may once therefore have been situated close
behind the visceral arches. If this is the strongest argument which can be brought
against the theory advocated in the text, there is I trust a considerable chance of its
being generally accepted. For even granting that Davidoff's deduction from the
character of the pelvic plexus is correct, there is, so far as I see, no reason in the
nature of the lateral fin theory why the pelvic fins should not have shifted, and on the
other hand the longitudinal cord connecting some of the spinal nerves in front of the
pelvic fin may have another explanation. It might for instance be a remnant of the
time when the pelvic fin had a more elongated form than at present, and accordingly
extended further forwards.
In any case our knowledge of the nature and origin of nervous plexuses is far too
imperfect to found upon their character such conclusions as those of Davidoff.
extends into the fin, and which becomes very early segmented
into a series of parallel rays at right angles to the longitudinal
In other words, the primitive skeleton of both the fins
consists of a longitudinal bar running along the base of the fin,
The skeleton of the fin was still in the condition of embryonic cartilage.
b.p. basipterygium (eventual metapterygium) ; fr. fin rays; p.g. pectoral girdle in
transverse section; /. foramen in pectoral girdle; pc. wall of peritoneal cavity.
and giving off at right angles series of rays which pass into the
fin. The longitudinal bar, which may be called the basipterygium, is moreover continuous in front with the pectoral or
pelvic girdle as the case may be.
The primitive skeleton of the pectoral fin is shewn in
longitudinal section in fig. 344, and that of the pelvic fin at a
slightly later stage in fig. 345.
A transverse section shewing the basipterygium (inpi) of the
pectoral fin, and the plate passing from it into the fin, is shewn
in fig. 346.
Before proceeding to describe the later history of the two
fins it may be well to point out that their embryonic structure
completely supports the view which has been arrived at from
the consideration of the soft parts of the fin.
My observations shew that the embryonic skeleton of the
paired fin consists of a series of parallel rays similar to those
of the unpaired fins. These rays support the soft part of the fin
which has the form of a longitudinal ridge, and are continuous
at their base with a longitudinal bar, which may very probably
be due to secondary development. As pointed out by Mivart, a
longitudinal bar is also occasionally formed to support the
cartilaginous rays of unpaired
fins. The longitudinal bar of
the paired fins is believed by
both Thacker and Mivart to
be due to the coalescence of
the bases of primitively independent rays, of which they
believe the fin to have been
originally composed. This
view is probable enough in
itself, but there is no trace
bb. basipterygium ; pu. pubic process
of pelvic girdle ; il. iliac process of pelvic
in the embryo of the bar in question being formed by the
coalesceace of rays, though the fact of its being perfectly
continuous with the bases of the rays is somewhat in favour
of this view 1 .
A point may be noticed here which may perhaps appear to be a
difficulty, viz. that to a considerable extent in the pectoral, and to some
extent in the pelvic fin the embryonic cartilage from which the fin-rays
are developed is at first a continuous lamina, which subsequently segments
into rays. I am however inclined to regard this merely as a result of the
mode of conversion of the indifferent mesoblast into cartilage ; and in any
case no conclusion adverse to the above view can be drawn from it, since
I find that the rays of the unpaired fin are similarly segmented from a
continuous lamina. In all cases the segmentation of the rays is to a large
extent completed before the tissue in question is sufficiently differentiated
to be called cartilage by an histologist.
Thacker and Mivart both hold that the pectoral and pelvic
girdles have been evolved by ventral and dorsal growths of the
anterior end of the longitudinal bar supporting the fin-rays.
There is, so far as I see, no theoretical objection to be taken
to this view, and the fact of the pectoral and pelvic girdles
originating continuously, and long remaining united with the
1 Thacker more especially founds his view on the adult form of the pelvic fins in
the cartilaginous Ganoids ; Polyodon, in which the part which constitutes the basal
plate in other forms is divided into separate segments, being mainly relied on. It is
possible that the segmentation of this plate, as maintained by Gegenbaur and Davidoff,
is secondary, but Thacker's view that the segmentation is a primitive character seems
to me, in the absence of definite evidence to the reverse, the more natural one.
longitudinal bars of their respective fins is in favour of rather
than against this view. The same may be said of the fact that
the first part of each girdle to be formed is that in the neighbourhood of the longitudinal bar (basipterygium) of the fin, the
dorsal and ventral prolongations being subsequent growths.
The later development of the skeleton of the two fins is more
conveniently treated separately.
The pelvic fin. The changes in the pelvic fin are comparatively slight. The fin remains through life as a nearly horizontal
lateral projection of the body, and the longitudinal bar the
mpt. basipterygial bar (metapterygium) ; fr. fin ray; m. muscles; hf. horny fibres.
basipterygium at its base always remains as such. It is for a
considerable period attached to the pelvic girdle, but eventually
becomes segmented from it. Of the fin rays the anterior
remains directly articulated with the pelvic girdle on the separation of the basipterygium (fig. 347), and the remaining rays
finally become segmented from the basipterygium, though they
remain articulated with it. They also become to some extent
transversely segmented. The posterior end of the basipterygial
bar also becomes segmented off as the terminal ray.
The pelvic fin thus retains in all essential points its primitive
6l 5
The pectoral fin. The earliest stage of the pectoral fin
bp. basipterygium ; m.o. process of basipterygium continued into clasper; il. iliac
process of pectoral girdle ; pit. pubis.
differs from that of the pelvic fin only in minor points,
is the same longitudinal
or basipterygial bar to
which the fin-rays are
attached, whose position
at the base of the fin is
clearly seen in the transverse section (fig. 346,
mpf). In front the bar is
continuous with the pectoral girdle (figs. 344 and
The changes which
take place in the course of
the further development
are however very much
more considerable in the
case of the pectoral than
in that of the pelvic fin. "' 3+8. F^OJJL ,,, v.
By the process spoken m p t me tapterygium (basipterygium of earlier
stage); me.p. rudiment of future pro- and mesopterygium ; sc. cut surface of scapular process ;
cr. coracoid process;/;', foramen;/, horny fibres.
of above, by which the
attachment of the pec
toral fin to the body wall becomes shortened from behind
forwards, the basipterygial bar is gradually rotated outwards,
its anterior end remaining attached to the pectoral girdle.
In this way this bar comes to form the posterior border of the
skeleton of the fin (figs. 348 and 349, mp], constituting what
Gegenbaur called the metapterygium, and eventually becomes
segmented off from the pectoral girdle, simply articulating
with its hinder edge.
The plate of cartilage, which is continued outwards from the
basipterygium, or as we may now call it, the metapterygium,
into the fin, is not nearly so completely divided up into fin-rays
as in the case of the pelvic fin, and this is especially the case
with the basal part of the plate. This basal part becomes
in fact at first only divided into two parts (fig. 348) a small
anterior part at the front end (me.p), and a larger posterior along
the base of the remainder of the fin. The anterior part directly
joins the pectoral girdle at its base, resembling in this respect
the anterior fin-ray of the pelvic girdle. It constitutes the
rudiment of the mesopterygium and propterygium of Gegenbaur.
It bears four fin-rays at its extremity, the anterior not being
well marked. The remaining fin-rays are borne by the edge of
the plate continuous with the metapterygium.
The further changes in the cartilages of the limb are not
important, and are easily understood by reference to fig. 349
representing the limb of a nearly full-grown embryo. The
front end of the anterior basal cartilage becomes segmented
off as a propterygium, bearing a single fin-ray, leaving the
remainder of the cartilage as a mesopterygium. The remainder
of the now considerably segmented fin-rays are borne by the
The mode of development of the pectoral fin demonstrates
that, as supposed by Mivart, the metapterygium is the homologue of the basal cartilage of the pelvic fin.
From the mode of development of the fins of Scyllium conclusions
may be drawn adverse to the views recently put forward on the structure of the fin by Gegenbaur and Huxley, both of whom consider the
primitive type of fin to be most nearly retained in Ceratodus, and to
consist of a central multisegmented axis with numerous rays. Gegenbaur
derives the Elasmobranch pectoral fin from a form which he calls the
archipterygium, nearly like that of Ceratodus, with a median axis and two
6I 7
rows of rays ; but holds that in addition to the rays attached to the median
axis, which are alone found in Ceratodus, there were other rays directly
articulated to the shoulder-girdle. He considers that in the Elasmobranch
fin the majority of the lateral rays on the posterior (median or inner
according to his view of the position of the limb) side have become
aborted, and that the central axis is represented by the metapterygium ;
while the pro- and mesopterygium and their rays are, he believes, derived
from those rays of the archipterygium which originally articulated directly
with the shoulder-girdle.
Gegenbaur's view appears to me to be absolutely negatived by the facts
of development of the pectoral fin in Scyllium ; not so much because the
pectoral fin in this form is necessarily to be regarded as primitive, but
because what Gegenbaur holds to be the primitive axis of the biserial fin
is demonstrated to be really the base, and it is only in the adult that it is
conceivable that a second set of lateral rays could have existed on the
posterior side of the metapterygium. If Gegenbaur's view were correct
we should expect to find in the embryo, if anywhere, traces of the second
set of lateral rays ; but the fact is that, as may easily be seen by an inspection of figs. 344 and 346, such a second set of lateral rays could not possibly have existed in a type .
of fin like that found in the
embryo 1 . With this view of
Gegenbaur's it appears to
me that the theory held by
this anatomist to the effect
that the limbs are modified
gill arches also falls ; in
that his method of deriving
the limbs from gill arches
ceases to be admissible,
while it is not easy to see
how a limb, formed on the
type of the embryonic limb
of Elasmobranchs, could be
derived from a visceral arch
with its branchial rays 2 .
Gegenbaur's older view
m.p. metapterygium ; me.p. mesopterygium ;
//. propterygium ; cr. coracoid process.
1 If, which I very much doubt, Gegenbaur is right in regarding certain rays found
in some Elasmobranch pectoral fins as rudiments of a second set of rays on the
posterior side of the metapterygium, these rays will have to be regarded as structures
in the act of being evolved, and not as persisting traces of a biserial fin.
2 Some arguments in favour of Gegenbaur's theory adduced by Wiedersheim as
a result of his researches on Protopterus are interesting. The attachment which he
describes between the external gills and the pectoral girdle is no doubt remarkable,
but I would suggest that the observations we have on the vascular supply of these
gills demonstrate that this attachment is secondary.
that the Elasmobranch fin retains a primitive uniserial type appears to me
to be nearer the truth than his more recent view on this subject ; though I
hold that the fundamental point established by the development of these
parts in Scyllium is that the posterior border of the adult Elasmobranch fin
is the primitive base line, i.e. the line of attachment of the fin to the side of
the body.
Huxley holds that the mesopterygium is the proximal piece of the axial
skeleton of the limb of Ceratodus, and derives the Elasmobranch fin from
that of Ceratodus by the shortening of its axis and the coalescence of some
of its elements. The secondary character of the mesopterygium, and its
total absence in the embryo Scyllium, appears to me as conclusive against
Huxley's view, as the character of the embryonic fin is against that of
Gegenbaur ; and I should be much more inclined to hold that the fin of
Ceratodus has been derived from a fin like that of the Elasmobranchii by
a series of steps similar to those which Huxley supposes to have led to the
establishment of the Elasmobranch fin, but in exactly the reverse order.
With reference to the development of the pectoral fin in the Teleostei
there are some observations of 'Swirski (No. 488) which unfortunately do
not throw very much light upon the nature of the limb.
'Swirski finds that in the Pike the skeleton of the limb is formed of a
plate of cartilage, continuous with the pectoral girdle ; which soon becomes
divided into a proximal and a distal portion. The former is subsequently
segmented into five basal rays, and the latter into twelve parts, the number
of which subsequently becomes reduced.
These investigations might be regarded as tending to shew that the
basipterygium of Elasmobranchii is not represented in Teleostei, owing to
the fin rays not having united into a continuous basal bar, but the observations are not sufficiently complete to admit of this conclusion being
founded upon them with any certainty.
Tlie ckeiropterygium.
Observations on the early development of the pentadactyloid
limbs of the higher Vertebrata are comparatively scanty.
The limbs arise as simple outgrowths of the sides of the
body, formed both of epiblast and mesoblast. In the Amniota,
at all events, they are processes of a special longitudinal ridge
known as the Wolffian ridge. In the Amniota they also bear
at their extremity a thickened cap of epiblast, which may be
compared with the epiblastic fold at the apex of the Elasmobranch fin.
Both limbs have at first a precisely similar position, both
being directed backwards and being parallel to the surface of
the body.
In the Urodela (Gotte) the ulnar and fibular sides are
primitively dorsal, and the radial and tibial ventral : in Mammalia however Kolliker states that the radial and tibial edges
are from the first anterior.
The exact changes of position undergone by the limbs in the
course of development are not fully understood. To suit a
terrestrial mode of life the flexures of the two limbs become
gradually more and more opposite, till in Mammalia the corresponding joints of the two limbs are turned in completely
opposite directions.
Within the mesoblast of the limbs a continuous blastema
becomes formed, which constitutes the first trace of the skeleton
of the limb. The corresponding elements of the two limbs,
viz. the humerus and femur, radius and tibia, ulna and fibula,
carpal and tarsal bones, metacarpals and metatarsals, and
digits, become differentiated within this, by the conversion
of definite regions into cartilage, which may either be completely
distinct or be at first united. These cartilaginous elements
subsequently ossify.
The later development of the parts, more especially of the carpus and
tarsus, has been made the subject of considerable study ; and important
results have been thereby obtained as to the homology of the various
carpal and tarsal bones throughout the Vertebrata ; but this subject is too
special to be treated of here. The early development, including the succession of the growth of the different parts, and the extent of continuity
primitively obtaining between them, has on the other hand been but little
investigated ; recently however the development of the limbs in the Urodela has been worked out in this way by two anatomists, Gotte (No. 482)
and Strasser (No. 487), and their results, though not on all points in complete harmony, are of considerable interest, more especially in their bearing
on the derivation of the pentadactyloid limb from the piscine fin. Till
however further investigations of the same nature have been made upon
other types, the conclusions to be drawn from Gotte and Strasser's observations must be regarded as somewhat provisional, the actual interpretation
of various ontological processes being very uncertain.
The forms investigated are Triton and Salamandra. We may remind
the reader that the hand of the Urodela has four digits, and the foot five,
the fifth digit being absent in the hand 1 . In Triton the proximal row of
carpal bones consists (using Gegenbaur's nomenclature) of (i) a radiale, and
(2 and 3) an intermedium and ulnare, partially united. The distal row is
formed of four carpals, of which the first often does not support the first
1 This seems to me clearly to follow from Gotte and Strasser's observations.
metacarpal ; while the second articulates with both the first and second
metacarpals. In the foot the proximal row of tarsals consists of a tibiale,
an intermedium and a fibulare. The distal row is formed of four tarsals, the
first, like that in the hand, often not articulating with the first metatarsal,
the second supporting the first and second metatarsals ; and the fourth the
fourth and fifth metatarsals.
The mode of development of the hand and foot is almost the same. The
most remarkable feature of development is the order of succession of the
digits. The two anterior (radial or tibial) are formed in the first instance,
and then the third, fourth and fifth in succession.
As to the actual development of the skeleton Strasser, whose observations
were made by means of sections, has arrived at the following results.
The humerus with the radius and ulna, and the corresponding parts in
the hind limb, are the first parts to be differentiated in the continuous plate
of tissue from which the skeleton of the limb is formed. Somewhat later a
cartilaginous centre appears at the base of the first and second fingers
(which have already appeared as prominences at the end of the limb) in the
situation of the permanent second carpal of the distal row of carpals ; and
the process of chondrification spreads from this centre into the fingers and
into the remainder of the carpus. In this way a continuous carpal plate
of cartilage is established, which is on the one hand continuous with the
cartilage of the two metacarpals, and on the other with the radius and ulna.
In the cartilage of the carpus two special columns may be noticed, the
one on the radial side, most advanced in development, being continuous with
the radius ; the other less developed column on the side of the ulna being
continuous both with the ulna and with the radius. The ulna and radius are
not united with the humerus.
In the further growth the third and fourth digits, and in the foot the fifth
digit also, gradually sprout out in succession from the ulnar side of the
continuous carpal plate. The carpal plate itself becomes segmented from the
radius and ulna, and divided up into the carpal bones.
The original radial column is divided into three elements, a proximal the
radiale, a middle element the first carpal, and a distal the second carpal
already spoken of. The first carpal is thus situated between the basal cartilage of the second digit and the radiale, and would therefore appear
to be the representative of a primitive middle row of carpal
bones, of which the centrale is also another representative.
The centrale and intermedium are the middle and proximal products of
the segmentation of the ulnar column of the primitive carpus, the distal
second carpal being common both to this column and to the radial column.
The ulnar or fibular side of the carpus or tarsus becomes divided into a
proximal element the ulnare or fibulare the ulnare remaining partially
united with the intermedium. There are also formed from this plate two
carpals to articulate with digits 3 and 4 ; while in the foot the corresponding
elements articulate respectively with the third digit, and with the fourth and
fifth digits.
Gotte, whose observations were made in a somewhat different method to
those of Strasser, is at variance with him on several points. He finds that
the primitive skeleton of the limb consists of a basal portion, the humerus,
continued into a radial and an ulnar ray, which are respectively prolonged
into the two first digits. The two rays next coalesce at the base of the
fingers to form the carpus, and thus the division of the limb into the brachium,
antebrachium and manus is effected.
The ulna, which is primitively prolonged into the second digit, is
subsequently separated from it and is prolonged into the third ; from the side
of the part of the carpus connecting the ulna with the third digit the fourth
digit is eventually budded out, and in the foot the fourth and fifth digits arise
from the corresponding region. Each of the three columns connected
respectively with the first, second, and third digits becomes divided into three
successive carpal bones, so that Gotte holds the skeleton of the hand or foot
to be formed of a proximal, a middle, and a distal row of carpal bones each
containing potentially three elements. The proximal row is formed of the
radiale, intermedium and ulnare ; the middle row of carpal i, the centrale
and carpal 4, and the distal of carpal 2 (consisting according to Gotte of two
coalesced elements) and carpal 3.
The derivation of the cheiropterygium from the ichthyoptcrygium. All
anatomists are agreed that the limbs of the higher Vertebrata are derived
from those of Fishes, but the gulf between the two types of limbs is so great
that there is room for a very great diversity of opinion as to the mode of
evolution of the cheiropterygium. The most important speculations on the
subject are those of Gegenbaur and Huxley.
Gegenbaur holds that the cheiropterygium is derived from a uniserial
piscine limb, and that it consists of a primitive stem, to which a series of
lateral rays are attached on one (the radial) side ; while Huxley holds that the
cheiropterygium is derived from a biserial piscine limb by the "lengthening of the axial skeleton, accompanied by the removal of its distal
elements further away from the shoulder-girdle and by a diminution in the
number of the rays."
Neither of these theories is founded upon ontology, and the only ontological evidence we have which bears on this question is that above recorded
with reference to the development of the Urodele limb.
Without holding that this evidence can be considered as in any way
conclusive, its tendency would appear to me to be in favour of regarding the
cheiropterygium as derived from a uniserial type of fin. The humerus or
femur would appear to be the basipterygial bars (metapterygium), which
have become directed outwards instead of retaining their original position
parallel to the length of the body at the base of the fin. The anterior
(proximal) fin-rays and the pro- and mesopterygium must be supposed to
have become aborted, while the radius or ulna, and tibia or fibula are two
posterior fin-rays (probably each representing several coalesced rays like the
pro- and mesopterygium) which support at their distal extremities more
numerous fin-rays consisting of the rows of carpal and tarsal bones.
This view of the cheiropterygium corresponds in some respects with that
put forward by Gotte as a result of his investigations on the development of
the Urodele limbs, though in other respects it is very different. A difficulty
of this view is the fact that it involves our supposing that the radial edge of
the limb corresponds with the metapterygial edge of the piscine fin. The
difficulties of this position have been clearly pointed out by Huxley, but the
fact that in the primitive position of the Urodele limbs the radius is ventral
and the ulna dorsal shews that this difficulty is not insuperable, in that it is
easy to conceive the radial border of the fin to have become rotated from its
primitive Elasmobranch position into the vertical position it occupies in the
embryos of the Urodela, and then to have been further rotated from this
position into that which it occupies in the adult Urodela and in all higher
BIBLIOGRAPHY of the Limbs.
(477) M. v. Davidoff. "Beitrage z. vergleich. Anat. d. hinteren Gliedmaassen
d. Fische I." Morphol. Jahrbuch, Vol. v. 1879.
(478) C. Gegenbaur. Untersuckungen z. vergleich. Anat. d. Wirbelthiere.
Leipzig, 1864 5. Erstes Heft. Carpus u. Tarsus. Zweites Heft. Brustflosse d.
(479) C. Gegenbaur. "Ueb. d. Skelet d. Gliedmaassen d. Wirbelthiere im
Allgemeinen u. d. Hintergliedmaassen d. Selachier insbesondere." Jenaische Zeitsckrift, Vol. V. 1870.
(480) C. Gegenbaur. " Ueb. d. Archipterygium." Jenaische Zeitschrift, Vol.
vii. 1873.
(481) C. Gegenbaur. "Zur Morphologic d. Gliedmaassen d. Wirbelthiere."
Morphologisches Jahrbuch, Vol. II. 1876.
(482) A. Gotte. Ueb. Entivick. u. Regeneration d. Gliedmaassenskelets d. Molche.
Leipzig, 1879.
(483) T. H. Huxley. "On Ceratodus Forsteri, with some observations on the
classification of Fishes." Proc. Zool. Soc. 1876.
(484) St George Mivart. "On the Fins of Elasmobranchii." Zoological
Trans., Vol. x.
(485) A. Rosenberg. "Ueb. d. Entwick. d. Extremitaten-Skelets bei einigen
d. Reduction ihrer Gliedmaassen charakterisirten Wirbelthieren." Zeil.f. iviss. Zool.,
Vol. xxin. 1873.
(486) E. Rosenberg. "Ueb. d. Entwick. d. Wirbelsaule u. d. centrale carpi
d. Menschen. " Morphologisches Jahrbuch, Vol. I. 1875.
(487) H. Strasser. "Z. Entwick. d. Extremitatenknorpel bei Salamandern u.
Tritonen." Morphologisches Jahrbuch, Vol. V. 1879.
(488) G. 'S wirski. Untersitch. iib. d. Entwick. d. Schultergitrtels u. d. Skelcls d.
Brustflosse d. Hechts. Inaug. Diss. Dorpat, 1880.
(489) J. K. Thacker. "Median and paired fins. A contribution to the history
of the Vertebrate limbs." Trans, of the Connecticut Acad., Vol. ill. 1877.
(490) J. K. Thacker. "Ventral fins of Ganoids." Trans, of the Connecticut
Acad., Vol. iv. 1877.
The Body cavity.
IN the Ccelenterata no body cavity as distinct from the
alimentary cavity is present ; but in the remaining Invertebrata
the body cavity may (i) take the form of a wide space separating
the wall of the gut from the body wall, or (2) may be present in
a more or less reduced form as a number of serous spaces, or
(3) only be represented by irregular channels between the
muscular and connective-tissue cells filling up the interior of the
body. The body cavity, in whatever form it presents itself, is
probably filled with fluid, and the fluid in it may contain special
cellular elements. A well developed body cavity may coexist
with an independent system of serous spaces, as in the Vertebrata and the Echinodermata ; the perihaemal section of the
body cavity of the latter probably representing the system of
serous spaces.
In several of the types with a well developed body cavity it
has been established that this cavity originates in the embryo
from a pair of alimentary diverticula, and the cavities resulting
from the formation of these diverticula may remain distinct, the
adjacent walls of the two cavities fusing to form a dorsal and a
ventral mesentery.
It is fairly certain that some groups, e.g. the Tracheata, with
imperfectly developed body cavities are descended from ancestors
which were provided with well developed body cavities, but how
far this is universally the case cannot as yet be definitely
decided, and for additional information on this subject the
reader is referred to pp. 355 360 and to the literature there
referred to.
In the Chaetopoda and the Tracheata the body cavity arises
as a series of paired compartments in the somites of mesoblast
(fig. 350) which have at first a very restricted extension on the
ventral side of the body, but eventually extend dorsalwards and
vcntralwards till each cavity is a half circle investing the
alimentary tract ; on the dorsal side the walls separating the two
The section is taken slightly to one side of the middle line so as to shew the relation of the mesoblastic somites to the limbs. In the interior are seen the yolk
segments and their nuclei.
i 16. the segments ; pr.l. procephalic lobe ; do. dorsal integument.
half cavities usually remain as the dorsal mesentery, while
ventrally they are in most instances absorbed. The transverse
walls, separating the successive compartments of the body
cavity, generally become more or less perforated.
Chordata. In the Chordata the primitive body cavity is
cither directly formed from a pair of alimentary diverticula
(Cephalochorda) (fig. 3) or as a pair of spaces in the mesoblastic
plates of the two sides of the body (fig. 20).
As already explained (pp. 294 300) the walls of the dorsal
sections of the primitive body cavity soon become separated
from those of the ventral, and becoming segmented constitute
the muscle plates, while the cavity within them becomes
obliterated : they are dealt with in a separate chapter. The
ventral part of the primitive cavity alone constitutes the
permanent body cavity.
The primitive body cavity in the lower Vertebrata is at first
continued forwards into the region of the head, but on the
formation of the visceral clefts the cephalic section of the body
cavity becomes divided into a series of separate compartments.
Subsequently these sections of the body cavity become obliterated ; and, since their walls give rise to muscles, they may
probably be looked upon as equivalent to the dorsal sections of
the body cavity in the trunk, and will be treated of in connection
with the muscular system.
As a result of its mode of origin the body cavity in
trunk is at first divided into two
lateral halves ; and part of the mesoblast lining it soon becomes distinguished as a special layer of epithelium, known as the peritoneal epithelium, of which the part bounding the
outer wall forms the somatic layer,
and that bounding the inner wall the
splanchnic layer. Between the two
splanchnic layers is placed the gut.
On the ventral side, in the region of
the permanent gut, the two halves
of the body cavity soon coalesce,
the septum between them becoming
absorbed, and the splanchnic layers
of epithelium of the two sides uniting
at the ventral side of the gut, and
the somatic layers at the median
ventral line of the body wall (fig.
In the lower Vertebrata the body
cavity is originally present even in
the post-anal region of the trunk, but
usually atrophies early, frequently
before the two halves coalesce.
On the dorsal side of the gut the
28 F.
sp.c. spinal canal ; W. white
matter of spinal cord ; pr. posterior nerve-roots ; cA. notochord ;
x. sub-notochordal rod ; ao. aorta ;
nip. muscle-plate ; nip 1 , inner layer
of muscle-plate already converted
into muscles; Vr. rudiment of
vertebral body ; si. segmental
tube ; sd. segmental duct ; sp.v.
spiral valve ; v. subintestinal vein ;
p.o. primitive generative cells.
two halves of the body cavity never coalesce, but eventually the
splanchnic layers of epithelium of the two sides, together with a
thin layer of interposed mesoblast, form a delicate membrane,
known as the mesentery, which suspends the gut from the dorsal
wall of the body (figs. 119 and 351). On the dorsal side the
epithelium lining of the body cavity is usually more columnar
than elsewhere (fig. 351), and its cells partly form a covering for
the generative organs, and partly give rise to the primitive
germinal cells. This part of the epithelium is often known as
the germinal epithelium.
Over the greater part of the body cavity the lining epithelium becomes in the adult intimately united with a layer of the
subjacent connective tissue, and constitutes with it a special
lining membrane for the body cavity, known as the peritoneal
Abdominal pores. In the Cyclostomata, the majority of the Elasmobranchii, the Ganoidei, a few Teleostei, the Dipnoi, and some Sauropsida
(Chelonia and Crocodilia) the body cavity is in communication with the
exterior by a pair of pores, known as abdominal pores, the external
openings of which are usually situated in the cloaca 1 .
The ontogeny of these pores has as yet been but very slightly investigated.
In the Lamprey they are formed as apertures leading from the body cavity
into the excretory section of the primitive cloaca. This section would
appear from Scott's (No. 87) observations to be derived from part of the
hypoblastic cloacal section of the alimentary tract.
In all other cases they are formed in a region which appears to belong
to the epiblastic region of the cloaca ; and from my observations on Elasmobranchs it may be certainly concluded that they are formed there
in this group. They may appear as perforations (i) at the apices of
papilliform prolongations of the body cavity, or (2) at the ends of cloacal
pits directed from the exterior towards the body cavity, or (3) as simple
slit-like openings.
Considering the difference in development between the abdominal pores
of most types, and those of the Cyclostomata, it is open to doubt whether
these two types of pores are strictly homologous.
In the Cyclostomata they serve for the passage outwards of the generative products, and they also have this function in some of the few Teleostei
in which they are found ; and Gegenbaur and Bridge hold that the primitive
mode of exit of the generative products, prior to the development of the
Miillerian ducts, was probably by means of these pores. I have elsewhere
1 For a full account of these structures the reader is referred to T. W. Bridge,
"Pori Abdominales of Vertebrata. " Journal of Anat. and Physiol. , Vol. XIV., 1879.
suggested that the abdominal pores are perhaps remnants of the openings
of segmental tubes ; there does not however appear to be any definite
evidence in favour of this view, and it is more probable that they may have
arisen as simple perforations of the body wall.
Pericardial cavity, pleural cavities, and diaphragm.
In all Vertebrata the heart is at first
placed in the body cavity (fig. 353 A),
but the part of the body cavity containing it afterwards becomes separated as
a distinct cavity known as the pericardial cavity. In Elasmobranchii, Acipenser, etc. a passage is however left
between the pericardial cavity and the
body cavity ; and in the Lamprey a
separation between the two cavities does
not occur during the Ammoccete stage.
In Elasmobranchii the pericardial
cavity becomes established as a distinct
space in front of the body cavity in the
following way. When the two ductus
Cuvieri, leading transversely from the
sinus venosus to the cardinal veins, become developed, a horizontal septum,
shewn on the right side in fig. 352, is
formed to support them, stretching
across from the splanchnic to the somatic side of the body cavity, and
dividing the body cavity (fig. 352) in
this part into (i) a dorsal section formed
of a right and left division constituting
the true body cavity (pp), and (2) a
ventral part the pericardial cavity (pc).
The septum is at first of a very small
longitudinal extent, so that both in
front and behind it (fig. 352 on the left
side) the dorsal and ventral sections of the body cavity are in
free communication. The septum soon however becomes prolonged, and ceasing to be quite horizontal, is directed obliquely
upwards and forwards till it meets the dorsal wall of the body
40 2
The figure shews the separation of the body cavity from
the pericardial cavity by a
horizontal septum in which
runs the ductus Cuvieri ; on
the left side is seen the narrow
passage which remains connecting the two cavities.
sp.c. spinal canal ; w. white
matter of spinal cord ; pr.
commissure connecting the
posterior nerve-roots ; ch. notochord ; x. sub-notochordal
rod ; ao. aorta ; sv. sinus venosus ; cav. cardinal vein ; ht.
heart ; pp. body cavity ; pc.
pericardial cavity ; as. solid
oesophagus ; /. liver ; nip. muscle-plate.
Anteriorly all communication is thus early shut off between the
body cavity and the pericardial cavity, but the two cavities still
open freely into each other behind.
The front part of the body cavity, lying dorsal to the pericardial cavity, becomes gradually narrowed, and is wholly
obliterated long before the close of embryonic life, so that in
adult Elasmobranch Fishes there is no section of the body cavity
dorsal to the pericardial cavity. The septum dividing the body
cavity from the pericardial cavity is prolonged backwards, till it
meets the ventral wall of the body at the point where the liver
is attached by its ventral mesentery (falciform ligament). In
this way the pericardial cavity becomes completely shut off from
the body cavity, except, it would seem, for the narrow communications found in the adult. The origin of these communications
has not however been satisfactorily worked out.
The septum between the pericardial cavity and the body
cavity is attached on its dorsal aspect to the liver. It is at first
nearly horizontal, but gradually assumes a more vertical position,
and then, owing to the obliteration of the primitive anterior
part of the body cavity, appears to mark the front boundary of
the body cavity. The above description of the mode of formation of the pericardial cavity, and the explanation of its relations
to the body cavity, probably holds true for Fishes generally.
In the higher types the earlier changes are precisely the
same as those in Elasmobranch Fishes. The heart is at first
placed within the body cavity attached to the ventral wall of
the gut by a mesocardium (fig. 353 A). A horizontal septum is
then formed, in which the ductus Cuvieri are placed, dividing
the body cavity for a short distance into a dorsal (/./) and
ventral (p.c) section (fig. 353 B). In Birds and Mammals, and
probably also in Reptilia, the ventral and dorsal parts of the
body cavity are at first in free communication both in front of
and behind this septum. This is shewn for the Chick in
fig- 353 A an d B, which are sections of the same chick, A being
a little in front of B. The septum is soon continued forwards
so as completely to separate the ventral pericardial and the
dorsal body cavity in front, the pericardial cavity extending at
this period considerably further forwards than the body cavity.
Since the horizontal septum, by its mode of origin, is
necessarily attached to the ventral side of the gut, the dorsal
part of the primitive body space is divided into two halves by a
median vertical septum formed of the gut and its mesentery
(fig- 353 B). Posteriorly the horizontal septum grows in a
slightly ventral direction along the under surface of the liver
(fig- 354)j till it meets the abdominal wall of the body at the
insertion of the falciform ligament, and thus completely shuts
off the pericardial cavity from the body cavity. The horizontal
septum forms, as is obvious from the above description, the
dorsal wall of the pericardial cavity 1 .
A. B.
p.p. body cavity; p.c. pericardial cavity; al. alimentary cavity ; au. auricle; v. ventricle; s.v. sinus venosus; d.c. ductus Cuvieri ; ao. aorta; nip. muscle-plate; me.
medullary cord.
With the complete separation of the pericardial cavity from
the body cavity, the first period in the development of these
parts is completed, and the relations of the body cavity to the
1 Kolliker's account of this septum, which he calls the mesocardium laterale (No.
298, p. 295), would seem to imply that in Mammals it is completed posteriorly even
before the formation of the liver. I doubt whether this takes place quite so early as
he implies, but have not yet determined its exact period by my own observations.
pericardial cavity become precisely those found in the embryos
of Elasmobranchii. The later changes are however very different. Whereas in Fishes the right and left sections of the body
cavity dorsal to the pericardial cavity soon atrophy, in the
higher types, in correlation with the relatively backward situation of the heart, they rapidly become larger, and receive the
lungs which soon sprout out from the throat.
The diverticula which form the lungs grow out into the
splanchnic mesoblast, in front of
the body cavity ; but as they
grow, they extend into the two
anterior compartments of the body
cavity, each attached by its mesentery to the mesentery of the
gut (fig. 354, lg). They soon moreover extend beyond the region of
the pericardium into the undivided
body cavity behind. This holds
not only for the embryos of the
Amphibia and Sauropsida, but
also for those of Mammalia.
To understand the further
rrianfrps in rhp nerirardial ravitv FlG> 354- SECTION THROUGH
it is necessary to bear in mind its OF LACERTA MURALIS OF 9 MM. TO
relations to the adjoining parts. THE PERICARDIAL CAVITY.
It lies at this period completely
ventral to the two anterior pro
ht. heart ; pc. pericardial cavity ;
al. alimentary tract; lg. lung; /.
liver ; pp. body cavity ; md. open
longations of the body Cavity COn- end of Mullerian duct ; wd. Wolffian
. . duct; vc. vena cava inferior; ao.
taming the lungs (fig. 354). Its aorta; ch. notochord; me. medullary
dorsal wall is attached to the gut, cord>
and is continuous with the mesentery of the gut passing to the
dorsal abdominal wall, forming the posterior mediastinum of
human anatomy.
The changes which next ensue consist essentially in the
enlargement of the sections of the body cavity dorsal to the
pericardial cavity. This enlargement takes place partly by the
elongation of the posterior mediastinum, but still more by the
two divisions of the body cavity which contain the lungs
extending themselves ventrally round the outside of the peri
cardial cavity. This process is illustrated by fig. 355, taken
from an embryo Rabbit. The two dorsal sections of the body
cavity (pl.p] finally extend so as completely to envelope the
pericardial cavity (pc\ remaining however separated from each
other below by a lamina extending from the ventral wall of the
pericardial cavity to the body wall, which forms the anterior
mediastinum of human anatomy.
By these changes the pericardial cavity is converted into a
closed bag, completely surrounded at its sides by the two lateral
halves of the body cavity, which were primitively placed
SJ3. C.
ht. heart; pc. pericardial cavity; //./ pleural cavity; Ig. lung; al. alimentary
tract; ao. dorsal aorta; ch. notochord; rp. rib; st. sternum; sp.c. spinal cord.
dorsally to it. These two sections of the body cavity, which in
Amphibia and Sauropsida remain in free communication with
the undivided peritoneal cavity behind, may, from the fact of
their containing the lungs, be called the pleural cavities.
In Mammalia a further change takes place, in that, by the
formation of a vertical partition across the body cavity, known
as the diaphragm, the pleural cavities, containing the lungs,
become isolated from the remainder of the body or peritoneal
cavity. As shewn by their development the so-called pleurae or
pleural sacks are simply the peritoneal linings of the anterior
divisions of the body cavity, shut off from the remainder of
the body cavity by the diaphragm.
The exact mode of formation of the diaphragm is not fully
made out ; the account of it recently given by Cadiat (No. 491)
not being in my opinion completely satisfactory.
(491) M. Cadiat. "Du developpement de la partie cephalothoracique de 1'embryon, de la formation du diaphragme, des pleures, du pericarde, du pharynx et de
1'cesophage." Journal de F Anatomic et de la Physiologic, Vol. xiv. 1878.
Vascular System.
The actual observations bearing on the origin of the vascular
system, using the term to include the lymphatic system, are
very scanty. It seems probable, mainly it must be admitted on
d priori grounds, that vascular and lymphatic systems have
originated from the conversion of indefinite spaces, primitively
situated in the general connective tissue, into definite channels.
It is quite certain that vascular systems have arisen independently in many types ; a very striking case of the kind being
the development in certain parasitic Copepoda of a closed
system of vessels with a red non-corpusculated blood (E. van
Beneden, Heider), not found in any other Crustacea. Parts of
vascular systems appear to have arisen in some cases by a
canalization of cells.
The blood systems may either be closed or communicate
with the body cavity. In cases where the primitive body cavity
is atrophied or partially broken up into separate compartments
(Insecta, Mollusca, Discophora, etc.) a free communication
between the vascular system and the body cavity is usually
present ; but in these cases the communication is no doubt
secondary. On the whole it would seem probable that the
vascular system has in most instances arisen independently of
the body cavity, at least in types where the body cavity is
present in a well-developed condition. As pointed out by the
Hertwigs, a vascular system is always absent where there is not
a considerable development of connective tissue.
As to the ontogeny of the vascular channels there is still much to be
made out both in Vertebrates and Invertebrates.
The smaller channels often rise by a canalization of cells. This process
has been satisfactorily studied by Lankester in the Leech 1 , and may easily
be observed in the blastoderm of the Chick or in the epiploon of a newlyborn Rabbit (Schafer, Ranvier). In either case the vessels arise from a network of cells, the superficial protoplasm and part of the nuclei giving rise
to the walls, and the blood-corpuscles being derived either from nucleated
masses set free within the vessels (the Chick) or from blood-corpuscles
directly differentiated in the axes of the cells (Mammals).
Larger vessels would seem to be formed from solid cords of cells, the
central cells becoming converted into the corpuscles, and the peripheral cells
constituting the walls. This mode of formation has been observed by
myself in the case of the Spider's heart, and by other observers in other
Invertebrata. In the Vertebrata a more or less similar mode of formation
appears to hold good for the larger vessels, but further investigations are
still required on this subject. Gotte finds that in the Frog the larger vessels
are formed as longitudinal spaces, and that the walls are derived from the
indifferent cells bounding these spaces, which become flattened and united
into a continuous layer.
The early formation of vessels in the Vertebrata takes place in the
splanchnic mesoblast ; but this appears due to the fact that the circulation
is at first mainly confined to the vitelline region, which is covered by
splanchnic mesoblast.
The Heart.
The heart is essentially formed as a tubular cavity in the
splanchnic mesoblast, on the ventral side of the throat, immediately behind the region of the visceral clefts. The walls of this
cavity are formed of two layers, an outer thicker layer, which has
at first only the form of a half tube, being incomplete on its
dorsal side; and an inner lamina formed of delicate flattened
cells. The latter is the epithelioid lining of the heart, and the
cavity it contains the true cavity of the heart. The outer layer
gives rise to the muscular wall and peritoneal covering of the
heart. Though at first it has only the form of a half tube (fig.
1 "Connective and vasifactive tissues of the Leech." Quart. J. of Micr. Science,
Vol. XX. 1880.
356), it soon becomes folded in on the dorsal side so as to form
for the heart a complete muscular wall. Its two sides, after thus
meeting to complete the tube of
the heart, remain at first continuous
with the splanchnic mesoblast surrounding the throat, and form a provisional mesentery the mesocardium which attaches the heart to
the ventral wall of the throat. The
superficial stratum of the wall of
the heart differentiates itself as the
peritoneal covering. The inner epithelioid tube takes its origin at the
time when the general cavity of the
heart is being formed by the separation of the splanchnicmesoblastfrom
the hypoblast. During this process
(fig. 357) a layer of mesoblast remains close to the hypoblast, but connected with the main mass
al. alimentary tract ; sp. splanchnic mesoblast ; so. somatic mesoblast ; ht. heart.
hb. hind-brain; vg. vagus nerve; ep. epiblast; ch. notochorcl ; x. thickening of
hypoblast (possibly a rudiment of the sub-notochordal rod) ; al. throat; ht. heart;
//. body cavity; so. somatic mesoblast; sf. splanchnic mesoblast; Ay. hypoblast.
of the mesoblast by protoplasmic processes. A second layer
next becomes split from the splanchnic mesoblast, connected
with the first layer by the above-mentioned protoplasmic
processes. These two layers form together the epithelioid lining
of the heart ; between them is the cavity of the heart, which soon
loses the protoplasmic trabeculae which at first traverse it. The
cavity of the heart may thus be described as being formed by a
hollowing out of the splanchnic mesoblast, and resembles in its
mode of origin that of other large vascular trunks.
The above description applies only to the development of
the heart in those types in which it is formed at a period after
the throat has become a closed tube (Elasmobranchii, Amphibia,
Cyclostomata, Ganoids (?)). In a number of other cases, in
which the heart is formed before the conversion of the throat
into a closed tube, of which the most notable is that of Mammals
(Hensen, Gotte, Kolliker), the heart arises as two independent
mes fir
SAME AGE AS FIG. 144 B. (From Kolliker.)
B is a more highly magnified representation of part of A.
rf. medullary groove; mp. medullary plate; riv. medullary fold; h. epiblast ;
dd. hypoblast; dd' . notochordal thickening of hypoblast; sp. undivided mesoblast;
^.somatic mesoblast; dfp. splanchnic mesoblast; ph. pericardial section of body
cavity; ahh. muscular wall of heart; ihh. epithelioid layer of heart; vies, lateral
undivided mesoblast ; sw. part of the hypoblast which will form the ventral wall of
the pharynx.
tubes (fig. 358), which eventually coalesce into an unpaired
In Mammals the two tubes out of which the heart is formed appear at
the sides of the cephalic plates, opposite the region of the mid- and hindbrain (fig. 358). They arise at a time when the lateral folds which form
the ventral wall of the throat are only just becoming visible. Each half of
the heart originates in the same way as the whole heart in Elasmobranchii,
etc. ; and the layer of the splanchnic mesoblast, which forms the muscular
wall for each part (ahh), has at first the form of a half tube open below to
the hypoblast.
On the formation of the lateral folds of the splanchnic walls, the two
halves of the heart become carried inwards and downwards, and eventually
fib. hind-brain ; nc. notochord ; E. epiblast ; so. somatopleure ; sp. splanchnopleure ; d. alimentary tract ; hy. hypoblast ; hs. heart ; of. vitelline veins.
meet on the ventral side of the throat. For a short time they here remain
distinct, but soon coalesce into a single tube.
In Birds, as in Mammals, the heart makes its appearance as two tubes,
but arises at a period when the formation of the throat is very much more
advanced than in the case of Mammals. The heart arises immediately
behind the point up to which the ventral wall of the throat is established
and thus has at first a A -shaped form. At the apex of the A , which forms
the anterior end of the heart, the two halves are in contact (fig. 357),
though they have not coalesced; while behind they diverge to be continued
as the vitelline veins. As the folding in of the throat is continued backwards the two limbs of the heart are brought together and soon coalesce
from before backwards into a single structure. Fig. 359 A and B shews the
heart during this process. The two halves have coalesced anteriorly (A)
but are still widely separated behind (B). In Teleostei the heart is formed
as in Birds and Mammals by the coalescence of two tubes, and it arises
before the formation of the throat.
The fact that the heart arises in so many instances as a
double tube might lead to the supposition that the ancestral
Vertebrate had two tubes in the place of the present unpaired
The following considerations appear to me to prove that this
conclusion cannot be accepted. If the folding in of the splanchnopleure to form the throat were deferred relatively to the
formation of the heart, it is clear that a modification in the
development of the heart would occur, in that the two halves of
the heart would necessarily be formed widely apart, and only
eventually united on the folding in of the wall of the throat. It
is therefore possible to explain the double formation of the heart
without having recourse to the above hypothesis of an ancestral
Vertebrate with two hearts. If the explanation just suggested
is the true one the heart should only be formed as two tubes
when it arises prior to the formation of the throat, and as a single
tube when formed after the formation of the throat. Since this
is invariably found to be so, it may be safely concluded that the
formation of the heart as two cavities is a secondary mode of
development, which has been brought about by variations in the
period of the closing in of the wall of the throat.
The heart arises continuously with the sinus venosus, which in
the Amniotic Vertebrata is directly continued into the vitelline
veins. Though at first it ends blindly in front, it is very soon
connected with the foremost aortic arches.
The simple tubular heart, connected as above described, grows
more rapidly than the chamber in which it is contained, and is
soon doubled upon itself, acquiring in this way an S-shaped
curvature, the posterior portion being placed dorsally, and the
anterior ventrally. A constriction soon appears between the
dorsal and ventral portions.
The dorsal section becomes partially divided off behind from
the sinus venosus, and constitutes the relatively thin-walled
auricular section of the heart; while the ventral portion, after
becoming distinct anteriorly from a portion continued forwards
from it to the origin of the branchial arteries, which may be called
the truncus arteriosus, acquires very thick spongy muscular
walls, and becomes the ventricular division of the heart.
The further changes in the heart are but slight in the case of the Pisces.
A pair of simple membranous valves becomes established at the auriculoventricular orifice, and further changes take place in the truncus arteriosus.
This part becomes divided in Elasmobranchii, Ganoidei, and Dipnoi into a
posterior section, called the conus arteriosus, provided with a series of
transverse rows of valves, and an anterior section, called the bulb us
arteriosus, not provided with valves, and leading into the branchial
arteries. In most Teleostei (except Butirinus and a few other forms) the
conus arteriosus is all but obliterated, and the anterior row of its valves
alone preserved ; and the bulbus is very much enlarged 1 .
In the Dipnoi important changes in the heart are effected, as compared
with other Fishes, by the development of true lungs. Both the auricular
and ventricular chamber may be imperfectly divided into two, and in the
conus a partial longitudinal septum is developed in connection with a
longitudinal row of valves 2 .
In Amphibia the heart is in many respects similar to that of the Dipnoi.
Its curvature is rather that of a screw than of a simple S. The truncus
arteriosus lies to the left, and is continued into the ventricle which lies
ventrally and more to the right, and this again into the dorsally placed
auricular section.
After the heart has reached the piscine stage, the auricular section
(Bombinator) becomes prolonged into a right and left auricular appendage^
A septum next grows from the roof of the auricular portion of the heart
1 Vide Gegenbaur, "Zur vergleich. Anat. d. Herzens." Jenaische Zeit., Vol. n.
1866, and for recent important observations, J. E. V. Boas, "Ueb. Herz u. Arterienbogenbei Ceratodenu. Protopterus," and " Ueber d. Conus arter. b. Butirinus, etc.,"
Morphol. Jahrb., Vol. VI. 1880.
2 Boas holds that the longitudinal septum is formed by the coalescence of a row of
longitudinal valves, but this is opposed to Lankester's statements, "On the hearts of
Ceratodus, Protopterus and Chimaera, etc. Zool. Trans. Vol. x. 1879.
obliquely backwards and towards the left, and divides it in two chambers ;
the right one of which remains continuous with the sinus venosus, while
the left one is completely shut off from the sinus, though it soon enters
into communication with the newly established pulmonary veins. The
truncus arteriosus 1 is divided into a posterior conus arteriosus (pylangium)
and an anterior bulbus (synangium). The former is provided with a
proximal row of valves at its ventricular end, and a distal row at its anterior
end near the bulbus. It is also provided with a longitudinal septum, which
is no doubt homologous with the septum in the conus arteriosus of the
Dipnoi. The bulbus is well developed in many Urodela, but hardly exists
in the Anura.
In the Amniota further changes take place in the heart,
resulting in the abortion of the distal rows of valves of the conus
arteriosus 2 , and in the splitting up of the whole truncus arteriosus
into three vessels in Reptilia, and two in Birds and Mammals,
each opening into the ventricular section of the heart, and
provided with a special set of valves at its commencement. In
Birds and Mammals the ventricle becomes moreover completely
divided into two chambers, each communicating with one of the
divisions of the primitive truncus, known in the higher types
as the systemic and pulmonary aortae. The character of the
development of the heart in the Amniota will be best understood
from a description of what takes place in the Chick.
In Birds the originally straight heart (fig. 109) soon becomes doubled up
upon itself. The ventricular portion becomes placed on the ventral and
right side, while the auricular section is dorsal and to the left. The two
parts are separated from each other by a slight constriction known as the
canalis auricularis. Anteriorly the ventricular cavity is continued into the
truncus, and the venous or auricular portion of the heart is similarly connected behind with the sinus venosus. The auricular appendages grow out
from the auricle at a very early period. The general appearance of the
heart, as seen from the ventral side on the fourth day, is shewn in fig. 360.
Although the external divisions of the heart are well marked even before
this stage, it is not till the end of the third day that the internal partitions
become apparent ; and, contrary to what might have been anticipated from
the evolution of these parts in the lower types, the ventricular septum is the
first to be established.
1 For a good description of the adult heart vide Huxley, Article "Amphibia," in
the Encyclopedia Britannic a.
2 It is just possible that the reverse may be true, vide note on p. 640. If however,
as is most probable, the statement in the text is correct, the valves at the mouth of
the ventricle in Teleostei are not homologous with those of the Amniota ; the former
being the distal rov/ of the valves of the conus, the latter the proximal.
It commences on the third day as a crescentic ridge or fold springing
from the convex or ventral side of the rounded ventricular portion of the
heart, and on the fourth day grows rapidly across the ventricular cavity
towards the concave or dorsal side. It thus forms an incomplete longitudinal partition, extending from the canalis auricularis to the commencement
of the truncus arteriosus, and dividing the twisted ventricular tube into
two somewhat curved canals, one more
to the left and above, the other to
the right and below. These commu- A ^) ) CA
nicate with each other, above the free
edge of the partition, along its whole
Externally the ventricular portion
as yet shews no division into two parts.
By the fifth day the venous end of
the heart, though still lying somewhat
to the left and above, is placed as far FIG. 360. HEART OF A CHICK ON
forwards as the arterial end, the whole THE FOURTH DAY OF INCUBATION
organ appearing to be drawn together.
The ventricular septum is complete. L ?.- lef t a , uricular appendage; C.A.
, e .. , . , , canahs auricularis ; v. ventricle ; b. trun
The apex of the ventricles becomes cus arteriosus.
more and more pointed. In the auricular portion a small longitudinal fold appears as the rudiment of the
auricular septum, while in the canalis auricularis, which is now at its greatest
length, there is also to be seen a commencement of the valvular structures
tending to separate the cavity of the auricles from those of the ventricles.
About the io6th hour, a septum begins to make its appearance in the
truncus arteriosus in the form of a longitudinal fold, which according to
Tonge (No. 495) starts at the end of the truncus furthest removed from the
heart. It takes origin from the wall of the truncus between the fourth and
fifth pairs of arches, and grows downwards in such a manner as to divide the
truncus into two channels, one of which leads from the heart to the third and
fourth pairs of arches, and the other to the fifth pair. Its course downwards
is not straight but spiral, and thus the two channels into which it divides
the truncus arteriosus wind spirally the one round the other.
At the time when the septum is first formed, the opening of the truncus
arteriosus into the ventricles is narrow or slit-like, apparently in order to
prevent the flow of the blood back into the heart. Soon after the appearance
of the septum, however, semilunar valves (Tonge, No. 495) are developed
from the wall of that portion of the truncus which lies between the free edge
of the septum and the cavity of the ventricles 1 .
1 If Tonge is correct in his statement that the semilunar valves develop at some
distance from the mouth of the ventricle, it would seem possible that the portion of
the truncus between them and the ventricle ought to be regarded as the embryonic
conus arteriosus, and that the distal row of valves of the conus (and not the proximal
as suggested above, p. 639) has been preserved in the higher types.
The ventral and the dorsal pairs of valves are the first to appear : the
former as two small solid prominences separated from each other by a
narrow groove ; the latter as a single ridge, in the centre of which is a
prominence indicating the point where the ridge will subsequently become
divided into two. The outer valves appear opposite each other, at a
considerably later period.
As the septum grows downwards towards the heart, it finally reaches
the position of these valves. One of its edges then passes between the two
ventral valves, and the other unites with the prominence on the dorsal
valve-ridge. At the same time the growth of all the parts causes the valves
to appear to approach the heart, and thus to be placed quite at the top
of the ventricular cavities. The free edge of the septum of the truncus now
A. B.
A. from the ventral, B. from the dorsal side.
La. left auricular appendage; r.a. right auricular appendage ; r.v. right ventricle;
l.v. left ventricle; b. truncus arteriosus.
fuses with the ventricular septum, and thus the division of the truncus into
two separate channels, each provided with three valves, and each communicating with a separate side of the heart, is complete ; the position of
the valves not being very different from that in the adult heart.
That division of the truncus which opens into the fifth pair of arches is
the one which communicates with the right ventricle, while that which
opens into the third and fourth pairs communicates with the left ventricle.
The former becomes the pulmonary artery, the latter the commencement of
the systemic aorta.
The external constriction actually dividing the truncus into two vessels
does not begin to appear till the septum has extended some way back
towards the heart.
The semilunar valves become pocketed at a period considerably later
than their first formation (from the H7th to the,i65th hour) in the order of
their appearance.
At the end of the sixth day, and even on the fifth day (figs. 361 and 362),
the appearance of the heart itself, without reference to the vessels which
come from it, is not very dissimilar from that of the adult. The original
4 1
La. left auricular appendage ;
r,a. right auricular appendage ;
r.v. right ventricle ; l.v. left ventricle ; b. truncus arteriosus.
protuberance to the right now forms the apex of the ventricles, and the
two auricular appendages are placed at the anterior extremity of the heart.
The most noticeable difference (in the ventral
view) is the still externally undivided condition of the truncus arteriosus.
The subsequent changes which the heart
undergoes are concerned more with its internal structure than with its external shape.
Indeed, during the next three days, viz. the
eighth, ninth, and tenth, the external form of
the heart remains nearly unaltered.
In the auricular portion, however, the
septum which commenced on the fifth day
becomes now more conspicuous. It is placed
vertically, and arises from the ventral wall ;
commencing at the canalis auricularis and
proceeding towards the opening into the
sinus venosus.
This latter structure gradually becomes
reduced so as to become a special appendage
of the right auricle. The inferior vena cava
enters the sinus obliquely from the right, so that its blood has a tendency to
flow towards the left auricle of the heart, which is at this time the larger of
the two.
The valves between the ventricles and auricles are now well developed,
and it is about this time that the division of the truncus arteriosus into the
aorta and pulmonary artery becomes visible from the exterior.
By the eleventh to the thirteenth day the right auricle has become as
large as the left, and the auricular septum much more complete, though
there is still a small opening, the foramen ovale, by which the two cavities
communicate with each other.
The most important feature in which the development of the Reptilian
heart differs from that of Birds is the division of the truncus into three
vessels, instead of two. The three vessels remain bound up in a common
sheath, and appear externally as a single trunk. The vessel not represented
in Birds is that which is continued into the left aortic arch.
In Mammals the early stages in the development of the heart present no
important points of difference from those of Aves. The septa in the truncus,
in the ventricular, and in the auricular cavities are formed, so far as
is known, in the same way and at the same relative periods in both groups.
In the embryo Man, the Rabbit, and other Mammals the division of
the ventricles is made apparent externally by a deep cleft, which, though
evanescent in these forms, is permanent in the Dugong.
The attachment of the auriculo-ventricular valves to the wall of the
ventricle, and the similar attachment of the left auriculo-ventricular valves
in Birds, have been especially studied by Gegenbaur and Bernays (No. 492),
and deserve to be noticed. In the primitive state the ventricular walls
have throughout a spongy character ; and the auriculo-ventricular valves are
simple membranous projections like the auriculo-ventricular valves of Fishes.
Soon however the spongy muscular tissue of both the ventricular and
auricular walls, which at first pass uninterruptedly the one into the other,
grows into the bases of the valves, which thus become in the main muscular
projections of the walls of the heart. As the wall of the ventricle thickens,
the muscular trabeculas, connected at one end with the valves, remain at the
other end united with the ventricular wall, and form special bands passing
between the two. The valves on the other hand lose their muscular
attachment to the auricular walls. This is the condition permanent in
Ornithorhynchus. In higher Mammalia the ends of the muscular bands
inserted into the valves become fibrous, from the development of intermuscular connective tissue, and the atrophy of the muscular elements.
The fibrous parts now form the chordae tendinea?, and the muscular the
musculi papillares.
The sinus venosus in Mammals becomes completely merged into the
right auricle, and the systemic division of the truncus arteriosus is apparently not homologous with that in Birds.
In the embryos of all the Craniata the heart is situated very
far forwards in the region of the head. This position is retained
in Pisces. In Amphibia the heart is moved further back, while
in all the Amniota it gradually shifts its position first of all into
the region of the neck and finally passes completely within the
thoracic cavity. The steps in the change of position may be
gathered from figs. 109, in, and 118.
BIBLIOGRAPHY of the Heart.
(492) A. C. Bernays. " Entwicklungsgeschichte d. Atrioventricularklappen."
Morphol. Jahrbuch,^o\. II. 1876.
(493) E. Gasser. " Ueber d. Entstehung d. Herzens beim Hiihn." Archiv f.
mikr. Anat., Vol. xiv.
(494) A. Thomson. "On the development of the vascular system of the foetus
of Vertebrated Animals." Edinb. New Phil. Journal, Vol. ix. 1830 and 1831.
(495) M. Tonge. "Observations on the development of the semilunar valves
of the aorta and pulmonary artery of the heart of the Chick." Phil. Trans. CLIX.
Vide also Von Baer (291), Rathke (300), Hensen (182), Kolliker (298), Gotte (296),
and Balfour (292).
Arterial System.
In the embryos of Vertebrata the arterial system consists of
a forward continuation of the truncus arteriosus, on the ventral
41 2
side of the throat (figs. 363, abr, and 364, a), which, with a few
exceptions to be noticed below, divides into as many branches on
each side as there are visceral arches. These branches, after
traversing the visceral arches, unite on the dorsal side of the
throat into a common trunk on each side. This trunk (figs. 363
and 364) after giving off one (or more) vessels to the head (c and
c] turns backv/ards, and bends in towards the middle line, close
to its fellow, immediately below the notochord (figs. 21 and 116)
and runs backwards in this situation towards the end of the tail.
The two parallel trunks below the notochord fuse very early into
a single trunk, the dorsal aorta (figs. 363, ad, and 364, a"}.
ttbr v "a,
a. auricle ; v. ventricle ; abr. branchial artery ; c'. carotid ; ad. dorsal aorta ;
s. branchial clefts; sv. sinus venosus; dc. ductus Cuvieri; n. nasal pit
There is given off from each collecting trunk from the visceral
arches, or from the commencement of the dorsal aorta, a subclavian
artery to each of the anterior limbs ; from near the anterior end
of the dorsal aorta a vitelline artery (or before the dorsal aortae
have united a pair of arteries fig. 125, R of A and L of A) to the
yolk-sack, which subsequently becomes the main visceral artery 1 ;
and from the dorsal aorta opposite the hind limbs one (or two)
arteries on each side the iliac arteries to the hind limbs ; from
these arteries the allantoic arteries are given off in the higher
types, which remain as the hypogastric arteries after the
disappearance of the allantois.
The primitive arrangement of the arterial trunks is with a
few modifications retained in Fishes. With the development of
the gills the vessels to the arches become divided into two parts
connected by a capillary system in the gill folds, viz. into the
1 In Mammalia the superior inesenteric artery arises from the vitelline artery,
which may probably be regarded as a primitive crclinco-mescnteric artery.
branchial arteries bringing the blood to the gills from the truncus
arteriosus, and the branchial veins transporting it to the dorsal
aorta. The branchial vessels to those arches which do not bear
gills, either wholly or partially atrophy; thus in Elasmobranchii
the mandibular trunk, which is fully developed in the embryo
(fig. 193, \av}, atrophies, except for a small remnant bringing
blood to the rudimentary gill of the spiracle from the branchial
vein of the hyoid arch. In Ganoids the mandibular artery
atrophies, but the hyoid is usually preserved. In Teleostei both
mandibular 1 and hyoid arteries are absent in the adult, except
that there is usually left a rudiment of the hyoid, supplying the
pseudobranch, which is similar to the rudiment of the mandibular
artery in Elasmobranchii. In Dipnoi the mandibular artery
atrophies, but the hyoid is sometimes preserved (Protopterus),
and sometimes lost.
In Fishes provided with a well developed air-bladder this
organ receives arteries, which arise sometimes from the dorsal
aorta, sometimes from the caeliac arteries, and sometimes from
the dorsal section of the last (fourth) branchial trunk. The
latter origin is found in Polypterus and Amia, and seems to have
been inherited by the Dipnoi where the air-bladder forms a true
The pulmonary artery of all the air-breathing Vertebrata is derived from the pulmonary artery of the
In all the types above Fishes considerable changes are
effected in the primitive arrangement of the arteries in the
visceral arches.
In Amphibia the piscine condition is most nearly retained 2 .
The mandibular artery is never developed, and the hyoid artery
is imperfect, being only connected with the cephalic vessels and
never directly joining the dorsal aorta. It is moreover developed
later than the arteries of the true branchial arches behind. The
subclavian arteries spring from the common trunks which unite
to form the dorsal aorta.
In the Urodela there are developed, in addition to the hyoid,
1 The mandibular artery is stated by Gotte never to be developed in Teleostei, but
is distinctly figured in Lereboullet (No. 71).
2 In my account of the Amphibia, Gotte (No. 296) has been followed.
four branchial arteries. The three foremost of these at first
supply gills, and in the Perennibranchiate forms continue to do
so through life. The fourth does not supply a gill, and very
early gives off, as in the Dipnoi, a pulmonary branch.
The hyoid artery soon sends forward a lingual artery from its
ventral end, and is at first continued to the carotid which grows
forward from the dorsal part of the first branchial vessel.
In the Caducibranchiata, where the gills atrophy, the following
changes take place. The remnant of the hyoid is continued
entirely into the lingual artery. The first branchial is mainly
continued into the carotid and other cephalic branches, but a
narrow remnant of the trunk, which originally connected it with
the dorsal aorta, remains, forming what is known as a ductus
Botalli. A rete mirabile on its course is the remnant of the
original gill.
The second and third branchial arches are continued as
simple trunks into the dorsal aorta, and the blood from the fourth
arch mainly passes to the lungs, but a narrow ductus Botalli still
connects this arch with the dorsal aorta.
In the Anura the same number of arches is present in the
embryo as in the Urodela, all four branchial arteries supplying
branchiae, but the arrangement of the two posterior trunks is
different from that in the Urodela. The third arch becomes at a
very early period continued into a pulmonary vessel, a relativelynarrow branch connecting it with the second arch. The fourth
arch joins the pulmonary branch of the third. At the metamorphosis the hyoid artery loses its connection with the carotid, and
the only part of it which persists is the root of the lingual artery.
The first branchial artery ceases to join the dorsal aorta, and
forms the root of the carotid : the so-called carotid gland placed
on its course is the remnant of the gill supplied by it before the
The second artery forms a root of the dorsal aorta. The
third, as in all the Amniota, now supplies the lungs, and also
sends off a cutaneous branch. The fourth disappears. The
connection of the pulmonary artery with both the third and
fourth branchial arches in the embryo appears to me clearly to
indicate that this artery was primitively derived from the fonrtli
arc/i as in the Urodela, and that its permanent connection
with the third arch in the Anura and in all the Amniota is
In the Amniota the metamorphosis of the arteries is in all
cases very similar. Five arches, viz. the mandibular, hyoid, and
three branchial arches are always developed (fig. 364), but, owing
to the absence of branchiae,
never function as branchial arteries. Of these the main parts of
the first two, connecting the truncus arteriosus with the collecting
trunk into which the arterial
arches fall, always disappear, usually before the complete development of the arteries in the posterior arches.
The anterior part of the collecting trunk into which these
vessels fall is not obliterated
when they disappear, but is on
the contrary continued forwards
as a vessel supplying the brain,
homologous with that found in
Fishes. It constitutes the internal
carotid. Similarly the anterior
part of the trunk from which the mandibular and hyoid arteries
sprang is continued forwards as a small vessel 1 , which at first
passes to the oral region and constitutes in Reptiles the lingual
artery, homologous with the lingual artery of the Amphibia ; but
in Birds and Mammals becomes more important, and is then
known as the external carotid (fig. 125). By these changes the
roots of the external and internal carotids spring respectively
from the ventral and dorsal ends of the primitive third artery,
i.e. the artery of the first branchial arch (fig. 365, c and c'} ; and
thus this arterial arch persists in all types as the common carotid,
AMNIOTA. (From Gegenbaur ; after
a. ventral aorta; a", dorsal aorta;
' 2 > 3> 4> 5- arterial arches ; c. carotid
1 His (No. 232) describes in Man two ventral continuations of the truncus arteriosus, one derived from the mandibular artery, forming the external maxillary artery,
and one from the hyoid artery, forming the lingual artery. The vessel from which
they spring is the external carotid. These observations of His will very probably be
found to hold true for other types.
6 4 8
and the basal part of the internal carotid. The trunk connecting
the third arterial arch with the system of the dorsal aorta persists
in some Reptiles (Lacertilia, fig. 366 A) as a ductus Botalli, but
is lost in the remaining Reptiles and in Birds and Mammals (fig.
366 B, C, D). It disappears earliest in Mammals (fig. 365 C),
later in Birds (fig. 365 B), and still later in the majority of
The fourth arch always continues to give rise, as in the Anura,
to the system of the dorsal aorta.
In all Reptiles it persists on both sides (fig. 366 A and B),
but with the division of the truncus arteriosus into three vessels
OF A. A LIZARD ; B. THE COMMON FOWL; C. THE PIG. (From Gegenbaur; after
The first two arches have disappeared in all three. In A and B the last three are
still complete, but in C the last two are alone complete.
/. pulmonary artery springing from the fifth arch, but still connected with the
system of the dorsal aorta by a ductus Botalli; c. external carotid; <'. internal
carotid; ad. dorsal aorta; a. auricle; v. ventricle; n. nasal pit; m, rudiment of
one of these, i.e. that opening furthest to the left side of the
ventricle (e and d), is continuous with the right fourth arch, and
also with the common carotid arteries (c) ; while a second
springing from the right side of the ventricle is continuous with
the left fourth arch (Ji and f). The right and left divisions of the
fourth arch meet however on the dorsal side of the oesophagus to
give origin to the dorsal aorta (g).
In Birds (fig. 366 C) the left fourth arch (h) loses its connection with the dorsal aorta, though the ventral part remains as
the root of the left subclavian. The truncus arteriosus is moreover only divided into two parts, one of which is continuous
with all the systemic arteries. Thus it comes about that in
Birds the right fourth arch (e) alone gives rise to the dorsal
In Mammals (fig. 366 D) the truncus arteriosus is only
divided into two, but the left fourth arch (>), instead of the right,
is that continuous with the dorsal aorta, and the right fourth
arch (/) is only continued into the right vertebral and right
subclavian arteries.
The fifth arch always gives origin to the pulmonary artery
(fig. 365, /) and is continuous with one of the divisions of the
truncus arteriosus. In Lizards (fig. 366 A, i), Chelonians and
Birds (fig. 366 C, i] and probably in Crocodilia, the right and
left pulmonary arteries spring respectively from the right and
left fifth arches, and during the greater part of embryonic life
the parts of the fifth arches between the origins of the pulmonary
arteries and the system of the dorsal aorta are preserved as
ductus Botalli. These ductus Botalli persist for life in the
Chelonia. In Ophidia (fig. 366 B, Ji) and Mammalia (fig.
366 D, m) only one of the fifth arches gives origin to the two
pulmonary arteries, viz. that on the right side in Ophidia, and
the left in Mammalia.
The ductus Botalli of the fifth arch (known in Man as the
ductus arteriosus) of the side on which the pulmonary arteries
are formed, may remain (e.g. in Man) as a solid cord connecting
the common stem of the pulmonary aorta with the systemic
The main history of the arterial arches in the Amniota has
been sufficiently dealt with, and the diagram, fig. 366, copied
from Rathke, shews at a glance the character of the metamorphosis these arches undergo in the different types. It merely
remains for me to say a few words about the subclavian and
vertebral arteries.
The subclavian arteries in Fishes usually spring from the
trunks connecting the branchial veins with the dorsal aorta.
This origin, which is also found in Amphibia, is typically found
in the embryos of the Amniota. In the Lizards this origin
persists through life, but both subclavians spring from the right
side. In most other types the origin of the subclavians is
carried upwards, so that they usually spring from a trunk
common to them and the carotids (arteria anonyma) (Birds and
some Mammals); or the left one, as in Man and some other
Mammals, arises from the systemic aorta just beyond the
carotids. Various further modifications in the origin of the
subclavians of the same general nature are found in Mammalia,
A 13
Mivart ; after Rathke.)
A. a. internal carotid; b. external carotid ; c. common carotid; d. ductus Botalli
between the third and fourth arches ; e. right aortic trunk ; /. subclavian ; g. dorsal
aorta; h. left aortic trunk; i. pulmonary artery; k. rudiment of ductus Botalli
between the pulmonary artery and the system of the dorsal aorta.
B. a. internal carotid; b. external carotid; c. common carotid; d. right aortic
trunk; e. vertebral artery;/, left aortic trunk of dorsal aorta; h. pulmonary artery ;
i. ductus Botalli of pulmonary artery.
C. a. internal carotid ; b. external carotid ; c. common carotid ; d. systemic
aorta; e. fourth arch of right side (root of dorsal aorta);/, right subclavian; g. dorsal
aorta; h, left subclavian (fourth arch of left side); i. pulmonary artery; k. and /.
right and left ductus Botalli of pulmonary arteries.
D. a. internal carotid; b. external carotid; c. common carotid; d. systemic aorta;
c. fourth arch of left side (root of dorsal aorta);/ dorsal aorta; g. left vertebral
artery; h. left subclavian artery; i. right subclavian (fourth arch of right side); k.
right vertebral; /. continuation of right subclavian; in. pulmonary artery; n. ductus
Botalli of pulmonary artery.
6 5 I
but they need not be specified in detail. The vertebral arteries
usually arise in close connection with the subclavians, but in
Birds they arise from the common carotids.
BIBLIOGRAPHY of the Arterial System.
(496) H. Rathke. " Ueb. d. Entwick. d. Arterien vv. bei d. Saugethiere von
d. Bogen d. Aorta ausgehen." Miiller's Archiv, 1843.
(-197) H. Rathke. " Untersuchungen lib. d. Aortenwurzeln d. Saurier."
Denkschriften d. k. Akad. Wien, Vol. XIII. 1857.
Vide also His (No. 232) and general works on Vertebrate Embryology.
TJie Venous System,.
The venous system, as it is found in the embryos of Fishes,
consists in its earliest condition of a single large trunk, which
traverses the splanchnic mesoblast investing the part of the
alimentary tract behind the heart. This trunk is directly continuous in front with the heart, and underlies the alimentary
canal through both its praeanal and postanal sections. It is
shewn in section in fig. 367, v, and may be called the subintestinal vein. This vein has been found in the embryos of
Teleostei, Ganoidei, Elasmobranchii and Cyclostomata, and runs
parallel to the dorsal aorta above, into which it is sometimes
continued behind (Teleostei, Ganoidei, etc.).
In Elasmobranch embryos the subintestinal vein terminates,
as may be gathered from sections (fig. 368, v.cau), shortly before
the end of the tail. The same series of sections also shews that
at the cloaca, where the gut enlarges and comes in contact with
the skin, this vein bifurcates, the two branches uniting into a
single vein both in front of and behind the cloaca.
In most Fishes the anterior part of this vein atrophies, the
caudal section alone remaining, but the anterior section of it
persists in the fold of the intestine in Petromyzon, and also
remains in the spiral valve of some Elasmobranchii. In
Amphioxus, moreover, it forms, as in the embryos of higher
types, the main venous trunk, though even here it is usually
broken up into two or three parallel vessels.
It no doubt represents one of the primitive longitudinal trunks of the
vermiform ancestors of the Chordata. The heart and the branchial artery
constitute a specially modified anterior continuation of this vein. The
dilated portal sinus of Myxine is probably also part of it ; and if this is
really rhythmically contractile 1 the fact would be interesting as shewing that
this quality, which is now localised in the heart, was once probably common
to the subintestinal vessel for its whole length.
On the development of the cardinal veins (to be described
below) considerable changes are
effected in the subintestinal vein.
Its postanal section, which is known
in the adult as the caudal vein,
unites with the cardinal veins. On
this junction being effected retrogressive changes take place in the
praeanal section of the original subintestinal vessel. It breaks up in
front into a number of smaller
vessels, the most important of which
is a special vein, which lies in the
fold of the spiral valve, and which is
more conspicuous in some Elasmobranchii than in Scyllium, in which
the development of the vessel has
been mainly studied. The lesser of
the two branches connecting it
round the cloaca with the caudal
vein first vanishes, and then the
larger ; and the two posterior cardinals are left as the sole forward
continuations of the caudal vein.
The latter then becomes prolonged
forwards, so that the two cardinals
open into it some little distance in
front of the hind end of the kidneys.
By these changes, and by the disappearance of the postanal section of the gut, the caudal vein is
made to appear as a supraintestinal and not, as it really is, a
subintestinal vessel.
From the subintestinal vein there is given off a branch which
supplies the yolk-sack. This leaves the subintestinal vein close
1 J. Miiller holds that this sack is not rhythmically contractile.
THAN 28 F.
sp.c. spinal canal; W. white
matter of spinal cord ; pr. posterior nerve-roots; ch. notochord ;
x. sub-notochordal rod ; ao. aorta ;
mp. muscle plate; ;;//'. inner layer
of muscle-plate already converted
into muscles; Vr. rudiment of
vertebral body; st. segmental
tube ; sd. segmental duct ; sp.v.
spiral valve; v. subintestinal vein ;
p.o. primitive generative cells.
to the liver. The liver, on its development, embraces the
subintestinal vein, which then breaks up into a capillary system
in the liver, the main part of its blood coming at this period
from the yolk-sack.
The portal system is thus established from the subintestinal
vein ; but is eventually joined by the various visceral, and sometimes by the genital, veins as they become successively developed.
The blood from the liver is brought back to the sinus venosus by veins known as the hepatic veins, which, like the hepatic
capillary system, are derivatives of the subintestinal vessel.
There join the portal system in Myxinoids and many
Teleostei a number of veins from the anterior abdominal walls,
representing a commencement of the anterior abdominal or
epigastric vein of higher types 1 .
In the higher Vertebrates the original subintestinal vessel never attains a
full development, even in the embryo. It is represented by (i) the ductus
A. is the posterior section.
nc. neural canal; al. post-anal gut; alv. caudal vesicle of post-anal gut; x.
subnotochordal rod; mp. muscle-plate; c/i. notochord; cl.al. cloaca; ao. aorta;
v.cait. caudal vein.
1 Stannius, Vergleich. Anat., p. 251.
venosus, which, like the true subintestinal vein, gives origin (in the Amniota)
to the vitelline veins to the yolk-sack, and (2) by the caudal vein. Whether
the partial atrophy of the subintestinal vessel was primitively caused by the
development of the cardinal veins, or for some other reason, it is at any rate
a fact that in all existing Fishes the cardinal veins form the main venous
channels of the trunk.
Their later development than the subintestinal vessel as well as their
absence in Amphioxus, probably indicate that they became evolved, at any
rate in their present form, within the Vertebrate phylum.
The embryonic condition of the venous system, with a single
large subintestinal vein is, as has been stated, always modified
by the development of a paired system of vessels, known as the
cardinal veins, which bring to the heart the greater part of the
blood from the trunk.
The cardinal veins appear in Fishes as four paired longitudinal trunks (figs. 363 and 369), two anterior (/) and two
posterior (c). They unite into two transverse trunks on either
side, known as the ductus Cuvieri (dc), which fall into the sinus
venosus, passing from the body wall to the sinus by a lateral
mesentery of the heart already spoken of (p. 627, fig. 352). The
anterior pair, known as the anterior cardinal or jugular veins,
bring to the heart the blood from the head and neck. They
are placed one on each side above the level
of the branchial arches (fig. 299, a.cv). The
posterior cardinal veins lie immediately dorsal to the mesonephros (Wolfifian body), and
are mainly supplied by the blood from this
organ and from the walls of the body (fig.
275, c.a.v). In many forms (Cyclostomata,
Elasmobranchii and many Teleostei) they
unite posteriorly with the caudal veins in
the manner already described, and in a large
number of instances the connecting branch
between the two systems, in its passage through
the mesonephros, breaks up into a capillary
network, and so gives rise to a renal portal
The vein from the anterior pair of fins
(subclavian) usually unites with the anterior
jugular vein.
OF A FISH. (From
Gegenbaur. )
j. jugular vein
(anterior cardinal
vein) ; c. posterior
cardinal vein; //. hepatic veins ; sv. sinus
venosus ; dc. ductus
The venous system of the Amphibia and Amniota always
differs from that of Fishes in the presence of a new vessel, the
vena cava inferior, which replaces the posterior cardinal veins;
the latter only being present, in their piscine form, during
embryonic life. It further differs from that of all Fishes, except
the Dipnoi, in the presence of pulmonary veins bringing back
the blood directly from the lungs.
In the embryos of all the higher forms the general characters
of the venous system are at first the same as in Fishes, but with
the development of the vena cava inferior the front sections of
the posterior cardinal veins atrophy, and the ductus Cuvieri,
remaining solely connected with the anterior cardinals and their
derivatives, constitute the superior venae cavae. The inferior
cava receives the hepatic veins.
Apart from the non-development of the subintestinal vein
the visceral section of the venous system is very similar to that
in Fishes.
The further changes in the venous system must be dealt
with separately for each group.
Amphibia. In Amphibia (Gotte, No. 296) the anterior and posterior
cardinal veins arise as in Pisces. From the former the internal jugular vein
arises as a branch ; the external jugular constituting the main stem. The
subclavian with its large cutaneous branch also springs from the system of
the anterior cardinal. The common trunk formed by the junction of these
three veins falls into the ductus Cuvieri.
The posterior cardinal veins occupy the same position as in Pisces, and
unite behind with the caudal veins, which Gotte has shewn to be originally
situated below the post-anal gut. The iliac veins unite with the posterior
cardinal veins, where the latter fall into the caudal vein. The original
piscine condition of the veins is not long retained. It is first of all disturbed
by the development of the anterior part of the important unpaired venous
trunk which forms in the adult the vena cava inferior. This is developed
independently, but unites behind with the right posterior cardinal. From
this point backwards the two cardinal veins coalesce for some distance, to
give rise to the posterior section of the vena cava inferior, situated between
the kidneys 1 . The anterior sections of the cardinal veins subsequently
atrophy. The posterior part of the cardinal veins, from their junction with
the vena cava inferior to the caudal veins, forms a rhomboidal figure. The
iliac vein joins the outer angle of this figure, and is thus in direct communication with the inferior vena cava, but it is also connected with a longitu
1 This statement of Gotte's is opposed to that of Rathke for the Amniota, and
cannot be considered as completely established.
dinal vessel on the outer border of the kidneys, which receives transverse
vertebral veins and transmits their blood to the kidneys, thus forming a
renal portal system. The anterior limbs of the rhomboid formed by the
cardinal veins soon atrophy, so that the blood from the hind limbs can only
pass to the inferior vena cava through the renal portal system. The
posterior parts of the two cardinal veins (uniting in the Urodela directly
with the unpaired caudal vein) still persist. The iliac veins also become
directly connected with a new vein, the anterior abdominal vein, which
has meanwhile become developed. Thus the iliac veins become united
with the system of the vena cava inferior through the vena renalis advehens
on the outer border of the kidney, and with the anterior abdominal veins by
the epigastric veins.
The visceral venous system begins with the development of two vitelline
veins, which at first join the sinus venosus directly. They soon become
enveloped in the liver, where they break up into a capillary system, which
is also joined by the other veins from the viscera. The hepatic system has
in fact the same relations as in Fishes. Into this system the anterior
abdominal vein also pours itself in the adult. This vein is originally
formed of two vessels, which at first fall directly into the sinus venosus,
uniting close to their opening into the sinus with a vein from the truncus
arteriosus. They become prolonged backwards, and after receiving the
epigastric veins above mentioned from the iliac veins, and also veins from
the allantoic bladder, unite behind into a single vessel. Anteriorly the
right vein atrophies and the left continues forward the unpaired posterior
A secondary connection becomes established between the anterior abdominal vein and the portal system ; so that the blood originally transported
by the former vein to the heart becomes diverted so as to fall into the liver.
A remnant of the primitive connection is still retained in the adult in the
form of a small vein, the so-called vena bulbi posterior, which brings the
blood from the walls of the truncus arteriosus directly into the anterior
abdominal vein.
The pulmonary veins grow directly from the heart to the lungs.
For our knowledge of the development of the venous system of the
Amniota we are mainly indebted to Rathke.
Reptilia. As an example of the Reptilia the Snake may be selected,
its venous system having been fully worked out by Rathke in his important
memoir on its development (No. 300).
The anterior (external jugular) and posterior cardinal veins are formed in
the embryo as in all other types (fig. 370, vj and vc] ; and the anterior
cardinal, after giving rise to the anterior vertebral and to the cephalic veins,
persists with but slight modifications in the adult ; while the two ductus
Cuvieri constitute the superior venos cavas.
The two posterior cardinals unite behind with the caudal veins. They
are placed in the usual situation on the dorsal and outer border of the
SNAKE. (From Gegenbaur;
after Rathke.)
vc. posterior cardinal
vein; vj. jugular vein; DC.
ductus Cuvieri ; vu. allantoic vein ; v. ventricle ; ba.
truncus arteriosus ; a. visceral clefts ; /. auditory
With the development of the vena cava inferior, to be described below,
the blood from the kidneys becomes mainly
transported by this vessel to the heart ; and the
section of the posterior cardinals opening into
the ductus Cuvieri gradually atrophies, their
posterior parts remaining however on the outer
border of the kidneys as the vena? renales
advehentes 1 .
While the front part of the posterior cardinal
veins is undergoing atrophy, the intercostal veins,
which originally poured their blood into the
posterior cardinal veins, become also connected
with two longitudinal veins the posterior vertebral veins which are homologous with the
azygos and hemiazygos veins of Man ; and bear
the same relation to the anterior vertebral veins
that the anterior and posterior cardinals do to
each other.
These veins are at first connected by trans
verse anastomoses with the posterior cardinals,
but, on the disappearance of the front part of the
latter, the whole of the blood from the intercostal veins falls into the
posterior vertebral veins. They are united in front with the anterior vertebral veins, and the common trunk of the two veins on each side falls into
the jugular vein.
The posterior vertebral veins are at first symmetrical, but after becoming
connected by transverse anastomoses, the right becomes the more important
of the two.
The vena cava inferior, though considerably later in its development
than the cardinals, arises fairly early. It constitutes in front an unpaired
trunk, at first very small, opening into the right allantoic vein, close to the
heart. Posteriorly it is continuous with two veins placed on the inner
border of the kidneys 2 .
The vena cava inferior passes through the dorsal part of the liver, and in
doing so receives the hepatic veins.
The portal system is at first constituted by the vitelline vein, which is
directly continuous with the venous end of the heart, and at first receives
the two ductus Cuvieri, but at a later period unites with the left ductus.
1 Rathke's account of the vena renalis advehens is thus entirely opposed to that
which Gotte gives for the Frog, but my own observations on the Lizard incline me to
accept Rathke's statements, for the Amniota at any rate.
2 The vena cava inferior does not according to Rathke's account unite behind with
the posterior cardinal veins, as it is stated by Gotte to do in the Anura. Gb'tte
questions the accuracy of Rathke's statements on this head, but my own observations
are entirely in favour of Rathke's observations, and lend no support whatever to
Gotte's views.
It soon receives a mesenteric vein bringing the blood from the viscera,
which is small at first but rapidly increases in importance.
The common trunk of the vitelline and mesenteric veins, which may be
called the portal vein, becomes early enveloped by the liver, and gives off
branches to this organ, the blood from which passes by the hepatic veins
to the vena cava inferior. As the branches in the liver become more
important, less and less blood is directly transported to the heart, and finally
the part of the original vitelline vein in front of the liver is absorbed, and the
whole of the blood from the portal system passes from the liver into the
vena cava inferior.
The last section of the venous system to be dealt with is that of the
anterior abdominal vein. There are originally, as in the Anura, two veins
belonging to this system, which owing to the precocious development of the
bladder to form the allantois, constitute the allantoic veins (fig. 370, vu}.
These veins, running along the anterior abdominal wall, are formed
somewhat later than the vitelline vein, and fall into the two ductus Cuvieri.
They unite with two epigastric veins (homologous with those in the Anura),
which connect them with the system of the posterior cardinal veins. The
left of the two eventually atrophies, so that there is formed an unpaired
allantoic vein. This vein at first receives the vena cava inferior close to the
heart, but eventually the junction of the two takes place in the region of the
liver, and finally the anterior abdominal vein (as it comes to be after the
atrophy of the allantois) joins the portal system and breaks up into capillaries
in the liver 1 .
In Lizards the iliac veins join the posterior cardinals, and so pour part of
their blood into the kidneys ; they also become connected by the epigastric
veins with the system of the anterior abdominal or allantoic vein. The
subclavian veins join the system of the superior venae cavas.
The venous system of Birds and Mammals differs in two important
points from that of Reptilia and Amphibia. Firstly the anterior abdominal
vein is only a foetal vessel, forming during foetal life the allantoic vein ;
and secondly a direct connection is established between the vena cava
inferior and the veins of the hind limbs and posterior parts of the cardinal
veins, so that there is no renal portal system.
Aves. The Chick may be taken to illustrate the development of the
venous system in Birds.
On the third day, nearly the whole of the venous blood from the body
of the embryo is carried back to the heart by two main venous trunks,
the anterior (fig. 125, S.Ca.V) and posterior (V.Ca) cardinal veins, joining on
each side to form the short transverse ductus Cuvieri (DC), both of which
unite with the sinus venosus close to the heart. As the head and neck
continue to enlarge, and the wings become developed, the single anterior
1 The junction between the portal system and the anterior abdominal vein is
apparently denied by Rathke (No. 300, p. 173), hut this must he an error on
his part.
cardinal or jugular vein (fig. 371, /), of each side, is joined by two new
veins : the vertebral vein, bringing back blood from the head and neck, and
the subclavian vein from the wing (W\
On the third day the posterior cardinal veins are the only veins which
return the blood from the hinder part of the body of the embryo.
About the fourth or fifth day, however, the vena cava inferior (fig. 371,
V.C.L) makes its appearance. This, starting
from the sinus venosus not far from the heart,
is on the fifth day a short trunk running backward in the middle line below the aorta, and
speedily losing itself in the tissues of the
Wolffian bodies. When the true kidneys are
formed it also receives blood from them, and
thenceforward enlarging rapidly becomes the
channel by which the greater part of the blood
from the hinder part of the body finds its way
to the heart. In proportion as the vena cava
inferior increases in size, the posterior cardinal
veins diminish.
The blood originally coming to them from
the posterior part of the spinal cord and trunk
is transported into two posterior vertebral veins,
similar to those in Reptilia, which are however
placed dorsally to the heads of the ribs, and
join the anterior vertebral veins. With their
appearance the anterior parts of the posterior
cardinals disappear. The blood from the hind
limbs becomes transported directly through the
kidney into the vena cava inferior, without
forming a renal portal system 1 .
On the third day the course of the vessels from the yolk-sack is very
simple. The two vitelline veins, of which the right is already the smaller,
form the ductus venosus, from which, as it passes through the liver on its
way to the heart, are given off the two sets of vena advehentes and vena
revehentes (fig. 371).
With the appearance of the allantois on the fourth day, a new feature is
introduced. From the ductus venosus there is given off a vein which
quickly divides into two branches. These, running along the ventral walls
of the body from which they receive some amount of blood, pass to the
allantois. They are the allantoic veins (fig. 371, U] homologous with the
anterior abdominal vein of the lower types. They unite in front to form a
single vein, which becomes, by reason of the rapid growth of the allantois,
very long. The right branch soon diminishes in size and finally disappears.
Meanwhile the left on reaching the allantois bifurcates ; and, its two
H. heart ; d. c. ductus Cuvieri. Into the ductus Cuvieri
of each side fall/, the jugular
vein, W. the vein from the
wing, and c. the inferior cardinal vein ; S. V. sinus venosus ;
Of. vitelline vein ; U. allantoic vein, which at this stage
gives off branches to the bodywalls ; V.C.l. inferior vena
cava ; /. liver.
The mode in which this is effected requires further investigation.
42 2
branches becoming large and conspicuous, there still appear to be two
main allantoic veins. At its first appearance the allantoic vein seems to be
but a small branch of the vitelline, but as the allantois grows rapidly,
and the yolk-sack dwindles, this state of things is reversed, and the less conspicuous vitelline appears as a branch of the larger allantoic vein.
On the third day the blood returning from the walls of the intestine is
insignificant in amount. As however the
intestine becomes more and more developed, it acquires a distinct venous system,
and its blood is returned by veins which
form a trunk, the mesenteric vein (fig. 372,
M") falling into the vitelline vein at its
junction with the allantoic vein.
These three great veins, in fact, form a
large common trunk, which enters at once
into the liver, and which we may now call
the portal vein (fig. 372, P. V}. This, at its
entrance into the liver, partly breaks up
into the vena advehentes, and partly continues as the ductus venosus (D.V}
straight through the liver, emerging from
which it joins the vena cava inferior. Before
the establishment of the vena cava inferior,
the venas revehentes, carrying back the
blood which circulates through the hepatic
capillaries, join the ductus venosus close to
its exit from the liver. By the time however that the vena cava has become a large
and important vessel it is found that the
venae revehentes, or as we may now call
them the hepatic veins, have shifted their
embouchment, and now fall directly into
that vein, the ductus venosus making a separate junction rather higher up (fig. 372).
This state of things continues with but slight changes till near the end
of incubation, when the chick begins to breathe the air in the air-chamber
of the shell, and respiration is no longer carried on by the allantois. Blood
then ceases to flow along the allantoic vessels ; they become obliterated.
The vitelline vein, which as the yolk becomes gradually absorbed proportionately diminishes in size and importance, comes to appear as a mere
branch of the portal vein. The ductus venosus becomes obliterated ; and
hence the whole of the blood coming through the portal vein flows into the
substance of the liver, and so by the hepatic veins into the vena cava.
Although the allantoic (anterior abdominal) vein is obliterated in the
adult, there is nevertheless established an anastomosis between the portal
system and the veins bringing the blood from the limbs to the vena cava
H. heart ; V.S.R. right vena
cava superior; V.S.L. left vena cava
superior. The two venas cavrc
superiores are the original 'ductus
Cuvieri,' they open into the sinus
venosus. J. jugular vein; Su.V.
anterior vertebral vein ; In. V. inferior vertebral vein ; W. subclavian; V.C.I, vena cava inferior;
D. V. ductus venosus ; P. V. portal
vein ; M. mesenteric vein bringing
blood from the intestines into the
portal vein ; O.f. vitelline vein ; U.
allantoic vein. The three last mentioned veins unite together to form
the portal vein ; /. liver.
inferior, in that the caudal vein and posterior pelvic veins open into a
vessel, known as the coccygeo-mesenteric vein, which joins the portal
vein ; while at the same time the posterior pelvic veins are connected with
the common iliac veins by a vessel which unites with them close to their
junction with the coccygeo-mesenteric vein.
Mammalia. In Mammals the same venous trunks are developed in
the embryo as in other types (fig. 373 A). The anterior cardinals or
external jugulars form the primitive veins of the anterior part of the body,
and the internal jugulars and anterior vertebrals are subsequently formed.
The subclavians (fig. 373 A, j), developed on the formation of the anterior
limbs, also pour their blood into these primitive trunks. In the lower
Mammalia (Monotremata, Marsupialia, Insectivora, some Rodentia, etc.,
the two ductus Cuvieri remain as the two superior venae cavae, but more
usually an anastomosis arises between the right and left innominate veins,
and eventually the whole of the blood of the left superior cava is carried to
the right side, and there is left only a single superior cava (fig. 373 B and C).
MAMMALS (MAN). (From Gegenbaur.)
j. jugular vein ; cs. vena cava superior; s. subclavian veins; c. posterior cardinal
vein ; v. vertebral vein ; az. azygos vein ; cor. coronary vein.
A. Stage in which the cardinal veins have already disappeared. Their position
is indicated by dotted lines.
B. Later stage when the blood from the left jugular vein is carried into the right
to form the single vena cava superior ; a remnant of the left superior cava being however still left.
C. Stage after the left vertebral vein has disappeared; the right vertebral
remaining as the azygos vein. The coronary vein remains as the last remnant of the
left superior vena cava.
A small rudiment of the left superior cava remains however as the sinus
coronartus and receives the coronary vein from the heart (figs. 373 C,
cor and 374, cs).
The posterior cardinal veins form at first the only veins receiving the
blood from the posterior part of the trunk and kidneys ; and on the
development of the hind limbs receive the blood from them also.
As in the types already described
an unpaired vena cava inferior becomes
eventually developed, and gradually
carries off a larger and larger portion
of the blood originally returned by the
posterior cardinals. It unites with the
common stem of the allantoic and
vitelline veins in front of the liver.
At a later period a pair of trunks
is established bringing the blood from
the posterior part of the cardinal veins
and the crural veins directly into the
vena cava inferior (fig. 374, il}. These
vessels, whose development has not
been adequately investigated, form the
common iliac veins, while the posterior
ends of the cardinal veins which join
them become the hypogastric veins (fig.
374, hy). Owing to the development of
the common iliac veins there is no renal
portal system like that of the Reptilia
and Amphibia.
Posterior vertebral veins, similar to
those of Reptilia and Birds, are established in connection with the intercostal
and lumbar veins, and unite anteriorly
with the front part of the posterior
cs. vena cava superior ; s. subclavian vein ; ji. internal jugular ; je.
external jugular ; az. azygos vein ; ha.
hemiazygos vein ; c. clotted line shewing previous position of cardinal veins ;
ci. vena cava inferior ; r. renal veins ;
il. iliac ; hy. hypogastric veins ; h.
hepatic veins.
The dotted lines shew the position
of embryonic vessels aborted in the
cardinal veins (fig. 373 A) 1 .
On the formation of the posterior vertebral veins, and as the inferior
vena cava becomes more important, the middle part of the posterior cardinals becomes completely aborted (fig. 374, f), the anterior and posterior
parts still persisting, the former as the continuations of the posterior
vertebrals into the anterior vena cava (az\ the latter as the hypogastric veins
Though in a few Mammalia both the posterior vertebrals persist, a
transverse connection is usually established between them, and the one (the
right) becoming the more important constitutes the azygos vein (fig. 374, az),
the persisting part of the left forming the hemiazygos vein (ha}.
The remainder of the venous system is formed in the embryo of the
vitelline and allantoic veins, the former being eventually joined by the
mesenteric vein so as to constitute the portal vein.
1 Rathke, as mentioned above, holds that in the Snake the front part of the
posterior cardinals completely aborts. Further investigations are required to shew
whether there really is a difference between Mammalia and Reptilia in this matter.
The vitelline vein is the first part of this system established, and divides
near the heart into two veins bringing back the blood from the yolk-sack
(umbilical vesicle). The right vein soon however aborts.
The allantoic (anterior abdominal) veins are originally paired. They
are developed very early, and at first course along the still widely open
somatic walls of the body, and fall into the single vitelline trunk in front.
The right allantoic vein disappears before long, and the common trunk
formed by the junction of the vitelline and allantoic veins becomes considerably elongated. This trunk is soon enveloped by the liver.
The succeeding changes have been somewhat differently described by
Kolliker and Rathke. According to the former the common trunk of the
allantoic and vitelline veins in its passage through the liver gives off
branches to the liver, and also receives branches from this organ near its
anterior exit. The main trunk is however never completely aborted, as in
the embryos of other types, but remains as the ductus venosus Arantii.
With the development of the placenta the allantoic vein becomes the
main source of the ductus venosus, and the vitelline or portal vein, as it may
perhaps be now conveniently called, ceases to join it directly, but falls into
one of its branches in the liver.
The vena cava inferior joins the continuation of the ductus venosus in
front of the liver, and, as it becomes more important, it receives directly
the hepatic veins which originally brought back blood into the ductus
venosus. The ductus venosus becomes moreover merely a small branch of
the vena cava.
At the close of foetal life the allantoic vein becomes obliterated up to its
place of entrance into the liver ; the ductus venosus becomes a solid cord
the so-called round ligament and the whole of the venous blood is brought
to the liver by the portal vein 1 .
Owing to the allantoic (anterior abdominal) vein having merely a fcetal
existence an anastomosis between the iliac veins and the portal system by
means of the anterior abdominal vein is not established.
BIBLIOGRAPHY of the Venous System.
(498) J. Marshall. "On the development of the great anterior veins." Phil.
Trans., 1859.
(499) H. Rathke. " Ueb. d. Bildung d. Pfortader u. d. Lebervenen b. Saugethieren." MeckeVs Archiv, 1830.
(500) H. Rathke. "Ueb. d. Bau u. d. Entwick. d. Venensystems d. Wirbelthiere." Bericht. Jib. d. natttrh. Seminar, d. Univ. Konigsberg, 1838.
Vide also Von Baer (No. 291), Gotte (No. 296), Kolliker (No. 298), and Rathke
(Nos. 299, 300, and 301).
1 According to Rathke the original trunk connecting the allantoic vein directly
with the heart through the liver is aborted, and the ductus venosus Arantii is a
secondary connection established in the latter part of foetal life.
Lymphatic System.
The lymphatic system arises from spaces in the general parenchyma of
the body, independent in their origin of the true body cavity, though communicating both with this cavity and with the vascular system.
In all the true Vertebrata certain parts of the system form definite trunks
communicating with the venous system ; and in the higher types the walls of
the main lymphatic trunks become quite distinct.
But little is known with reference to the ontogeny of the lymphatic vessels,
but they originate late in larval life, and have at first the form of simple
intercellular spaces.
The lymphatic glands appear to originate from lymphatic plexuses, the
cells of which produce lymph corpuscles. It is only in Birds and Mammals,
and especially in the latter, that the lymphatic glands form definite structures.
The Spleen. The spleen, from its structure, must be classed with the
lymphatic glands, though it has definite relations to the vascular system.
It is developed in the mesoblast of the mesogastrium, usually about the
same time and in close connection with the pancreas.
According to Miiller and Peremeschko the mass of mesoblast which
forms the spleen becomes early separated by a groove on the one side from
the pancreas and on the other from the mesentery. Some of its cells
become elongated, and send out processes which uniting with like processes
from other cells form the trabecular system. From the remainder of the
tissue are derived the cells of the spleen pulp, which frequently contain more
than one nucleus. Especial accumulations of these cells take place at a
later period to form the so-called Malpighian corpuscles of the spleen.
(501) W. Miiller. "The Spleen." Strieker's Histology.
(502) Peremeschko. " Ueb. d. Entwick. d. Milz." Sitz. d. Wuti. Akad.
Wiss., Vol. LVI. 1867.
Suprarenal ^bodies.
In Elasmobranch Fishes two distinct sets of structures are found, both of
which have been called suprarenal bodies. As shewn in the sequel both of
these structures probably unite in the higher types to form the suprarenal
One of them consists of a series of paired bodies, situated on the
branches of the dorsal aorta, segmentally arranged, and forming a chain
extending from close behind the heart to the hinder end of the body cavity.
Each body is formed of a series of lobes, and exhibits a well-marked
distinction into a cortical layer of columnar cells, and a medullary substance
formed of irregular polygonal cells. As first shewn by Leydig, they are
closely connected with the sympathetic ganglia, and usually contain numerous
ganglion cells distributed amongst the proper cells of the body.
The second body consists of an unpaired column of cells placed between
the dorsal aorta and unpaired caudal vein, and bounded on each side by the
posterior parts of the kidney. I propose to call it the interrenal body.
In front it overlaps the paired suprarenal bodies, but does not unite with
them. It is formed of a series of well-marked lobules, etc. In the fresh
state Leydig (No. 506) finds that "fat molecules form the chief mass of the
body, and one finds freely imbedded in them clear vesicular nuclei." As
may easily be made out from hardened specimens it is invested by a tunica
propria, which gives off septa dividing it into well-marked areas filled with
polygonal cells. These cells constitute the true parenchyma of the body.
By the ordinary methods of hardening, the oil globules, with which they are
filled in the fresh state, completely disappear.
The paired suprarenal bodies (Balfour, No. 292, pp. 242 244) are developed from the sympathetic ganglia. These ganglia, shewn in an early
stage in fig. 380, sy.g, become gradually divided into a ganglionic part and a
glandular part. The former constitutes the sympathetic ganglia of the adult ;
the latter the true paired suprarenal bodies. The interrenal body is however
developed (Balfour, No. 292, pp. 245 247) from indifferent mesoblast cells
between the two kidneys, in the same situation as in the adult.
The development of the suprarenal bodies in the Amniota has been most
fully studied by Braun (No. 503) in the Reptilia.
In Lacertilia they consist of a pair of elongated yellowish bodies, placed
between the vena renalis revehens and the generative glands.
They are formed of two constituents, viz. (i) masses of brown cells placed
on the dorsal side of the organ, which stain deeply with chromic acid, like
certain of the cells of the suprarenals of Mammalia, and (2) irregular cords,
in part provided with a lumen, filled with fat-like globules l , amongst which
are nuclei. On treatment with chromic acid the fat globules disappear, and
the cords break up into bodies resembling columnar cells.
The dorsal masses of brown cells are developed from the sympathetic
ganglia in the same way as the paired suprarenal bodies of the Elasmobranchii, while the cords filled with fat-like globules are formed of indifferent
mesoblast cells as a thickening in the lateral walls of the inferior vena cava,
and the cardinal veins continuous with it. The observations of Brunn (No.
504) on the Chick, and Kolliker (No. 298, pp. 953955) n the Mammal,
add but little to those of Braun. They shew that the greater part of the
gland (the cortical substance) in these two types is derived from the mesoblast,
and that the glands are closely connected with sympathetic ganglia ; while
Kolliker also states that the posterior part of the organ is unpaired in the
embryo rabbit of 1 6 or 17 days.
The structure and development of what I have called the interrenal body
1 These globules are not formed of a true fatty substance, and this is also probably
true for the similar globules of the interrenal bodies of Elasmobranchii.
in Elasmobranchii so closely correspond with that of the mesoblastic part of
the suprarenal bodies of the Reptilia, that I have very little hesitation in
regarding them as homologous 1 ; while the paired bodies in Elasmobranchii,
derived from the sympathetic ganglia, clearly correspond with the part of the
suprarenals of Reptilia having a similar origin ; although the anterior parts
of the paired suprarenal bodies of Fishes have clearly become aborted in the
higher types.
In Elasmobranch Fishes we thus have (i) a series of paired
bodies, derived from the sympathetic ganglia, and (2) an unpaired body of mesoblastic origin. In the Amniota these bodies
unite to form the compound suprarenal bodies, the two constituents of which remain, however, distinct in their development.
The mesoblastic constituent appears to form the cortical part of
the adult suprarenal body, and the nervous constituent the
medullary part.
BIBLIOGRAPHY of the Suprarenal bodies,
(503) M. Braun. "Bau u. Entwick. d. Nebennieren bei Reptilien. " Arbeit,
a. d. zool.-zoot. Institut Wurzlttrg, Vol. V. 1879.
(504) A. v. Brunn. "Ein Beitrag z. Kenntniss d. feinern Baues u. d. Entwick.
d. Nebennieren." Archiv f. mikr. Anat., Vol. VIII. 1872.
(505) Fr. Leydig. Untersiich. iib. Fische u. fieptilten. Berlin, 1853.
(506) Fr. Leydig. Rochen u. Haie. Leipzig, 1852.
Vide also F. M. Balfour (No. 292), Kolliker (No. 298), Remak (No. 302), etc.
1 The fact of the organ being unpaired in Elasmobranchii and paired in the
Amniota is of no importance, as is shewn by the fact that part of the organ is unpaired
in the Rabbit.
IN all the Ccelenterata, except the Ctenophora, the contractile elements of the body wall consist of filiform processes of
ectodermal or entodermal epithelial cells (figs. 375 and 376 B).
The elements provided with these processes, which were first
discovered by Kleinenberg, are known as myo-epithelial
cells. Their contractile parts may either be striated (fig. 376)
or non-striated (fig. 375). In some
instances the epithelial part of the
cell may nearly abort, its nucleus
alone remaining (fig. 376 A) ; and
in this way a layer of muscles lying
completely below the surface may
be established.
There is embryological evidence
of the derivation of the voluntary
muscular system of a large number of types from myo-epithelial
cells of this kind. The more important of these groups are the
Chaetopoda, the Gephyrea, the Chaetognatha, the Nematoda, and
the Vertebrata 1 .
While there is clear evidence that the muscular system of a
large number of types is composed of cells which had their
origin in myo-epithelial cells, the mode of evolution of the
1 If recent statements of Metschnikoff are to be trusted, the Echinodermata must
be added to these groups. The amoeboid cells stated in the first volume of this
treatise to form the muscles in this group, on the authority of Selenka, give rise,
according to Metschnikoff, only to the cutis, while the same naturalist states the
epithelial cells of the vasoperitoneal vesicles are provided with muscular tails.
CELLS OF HYDRA. (From Gegenbaur ; after Kleinenberg.)
m. contractile fibres.
muscular system of other types is still very obscure. The
muscles may arise in the embryo from amoeboid or indifferent
cells, and the Hertwigs 1 hold that in many of these instances the
muscles have also phylogenetically taken their origin from
indifferent connective-tissue cells. The subject is however beset
with very serious difficulties, and to discuss it here would carry
me too far into the region of pure histology.
The voluntary muscular system of the CJiordata.
The muscular fibres. The muscular elements of the
Chordata undoubtedly belong to the myo-epithelial type. The
embryonic muscle-cells are at first simple epithelial cells, but
FIG. 376. MUSCLE-CELLS OF LIZZIA KOLLIKERI. (From Lankester ; after
O. and R. Hertwig.)
A. Muscle-cell from the circular fibres of the subumbrella.
B. Myo-epithelial cells from the base of a tentacle.
soon become spindle-shaped : part of their protoplasm becomes
differentiated into longitudinally placed striated muscular fibrils,
while part, enclosing the nucleus, remains indifferent, and constitutes the epithelial element of the cells. The muscular
fibrils are either placed at one side of the epithelial part of the
cell, or in other instances (the Lamprey, the Newt, the Sturgeon,
the Rabbit) surround it. The latter arrangement is shewn for
the Sturgeon in fig. 57.
The number of the fibrils of each cell gradually increases,
and the protoplasm diminishes, so that eventually only the
nucleus, or nuclei resulting from its division, are left. The
products of each cell probably give rise, in conjunction with a
further division of the nucleus, to a primitive bundle, which,
1 O. and R. Hertwig, Die Calomthcorie. Jena, 1881.
except in Amphioxus, Petromyzon, etc., is surrounded by a
special investment of sarcolemma.
The voluntary muscular system. For the purposes of
description the muscular system of the Vertebrata may conveniently be divided into two sections, viz. that of the head and
that of the trunk. The main part, if
not the whole, of the muscular system
of the trunk is derived from certain
structures, known as the muscle-plates,
which take their origin from part of
the primitive mesoblastic somites.
It has already been stated (pp.
292 ^296) that the mesoblastic somites
are derived from the dorsal segmented
part of the primitive mesoblastic plates.
Since the history of these bodies is
presented in its simplest form in Elasmobranchii it will be convenient to
commence with this group. Each
somite is composed of two layers a
somatic and a splanchnic both formed
of a single row of columnar cells.
Between these two layers is a cavity,
which is at first directly continuous
with the general body cavity, of which
indeed it merely forms a specialised
part (fig. 377). Before long the cavity
becomes however completely constricted off from the permanent body cavity.
Very early (fig. 377) the inner or splanchnic wall of the
somites loses its simple constitution, owing to the middle part of
it undergoing peculiar changes. The meaning of the changes is
at once shewn by longitudinal horizontal sections, which prove
(% 378) that the cells in this situation (mp') have become
extended in a longitudinal direction, and, in fact, form typical
spindle-shaped embryonic muscle-cells, each with a large
nucleus. Every muscle-cell extends for the whole length of a
somite. The inner layer of each somite, immediately within
the muscle-band just described, begins to proliferate, and produce
nc. neural canal ; pr. posterior root of spinal nerve ; x.
subnotochordal rod ; ao. aorta ;
sc. somatic mesoblast ; sf>.
splanchnic mesoblast ; mp.
muscle-plate ; mp', portion of
muscle-plate converted into
muscle ; Vr. portion of the
vertebral plate which will give
rise to the vertebral bodies ; al.
alimentary tract.
a mass of cells, placed between the muscles and the notochord
( Vr\ These cells form the commencing vertebral bodies, and
have at first (fig. 378) the same segmentation as the somites
from which they sprang.
After the separation of the vertebral bodies from the somites
the remaining parts of the somites may be called muscle-plates ;
since they become directly converted into the whole voluntary
muscular system of the trunk (fig. 379, mp}.
According to the statements of Bambeke and Go'tte, the Amphibians
present some noticeable peculiarities in the development of their muscular
system, in that such distinct muscle-plates as those of other vertebrate types
are not developed. Each side-plate of mesoblast is divided into a somatic
and a splanchnic layer, continuous throughout the vertebral and parietal
portions of the plate. The vertebral portions (somites) of the plates soon
become separated from the parietal, and form independent masses of cells
constituted of two layers, which were originally continuous with the
somatic and splanchnic layers of the parietal plates (fig. 79). The outer or
somatic layer of the vertebral plates is formed of a single row of cells, but
the inner or splanchnic layer is made up of a kernel of cells on the side of
the somatic layer and an inner layer. The kernel of the splanchnic layer
and the outer or somatic layer together correspond to a muscle- plate of other
Vertebrata, and exhibit a similar segmentation.
Osseous Fishes are stated to agree with Amphibians in the development
of their somites and muscular
system 1 , but further observations
on this point are required.
In Birds the horizontal splitting of the mesoblast extends at
first to the dorsal summit of the
mesoblastic plates, but after the
isolation of the somites the split
between the somatic and splanchnic layers becomes to a large extent obliterated, though in the anterior somites it appears in part
to persist. The somites on the
second day, as seen in a transverse section (fig. 115, P.?'.), are
somewhat quadrilateral in form
but broader than they are deep.
Each at that time consists of
a somewhat thick cortex of radi
The section is taken at the level of the
notochord, and shews the separation of the
cells to form the vertebral bodies from the
ch. notochord ; ep. epiblast ; Vr, rudiment
of vertebral body ; mp. muscle- plate ; mp' .
portion of muscle-plate already differentiated
into longitudinal muscles.
1 Ehrlich, " Ueber den peripher. Theil d. Urwirbel." Archiv f. mikr. Anal.,
Vol. XI.
ating rather granular columnar cells, enclosing a small kernel of spherical
cells. They are not, as may be seen in the above figure, completely
separated from the ventral (or lateral as they are at this period) parts of the
mesoblastic plate, and the dorsal and outer layer of the cortex of the
somites is continuous with the somatic layer of mesoblast, the remainder of
the cortex, with the central kernel, being continuous with the splanchnic
layer. Towards the end of the second and beginning of the third day the
upper and outer layer of the cortex, together probably with some of the
central cells of the kernel, becomes separated off as a muscle-plate (fig. 1 16).
The muscle-plate when formed (fig. 117) is found to consist of two layers,
an inner and an outer, which enclose between them an almost obliterated
central cavity ; and no sooner is the muscle-plate formed than the middle
portion of the inner layer becomes converted into longitudinal muscles.
The avian muscle-plates have, in fact, precisely the same constitution as
those of Elasmobranchii. The central space is clearly a remnant of the
vertebral portion of the body cavity, which, though it wholly or partially
disappears in a previous stage, reappears again on the formation of the
The remainder of the somite, after the formation of the muscle-plate,
is of very considerable bulk ; the cells of the cortex belonging to it lose
their distinctive characters, and the major part of it becomes the vertebral
In Mammalia the history appears to be generally the same as in Elasmobranchii. The split which gives rise to the body cavity is continued to
the dorsal summit of the mesoblastic plates, and the dorsal portions of the
plates with their contained cavities become divided into somites, and are
then separated off from the ventral. The later development of the somites
has not been worked out with the requisite care, but it would seem that they
form somewhat cubical bodies in which all trace of the primitive slit is lost.
The further development resembles that in Birds.
The first changes of the mesoblastic somites and the formation of the muscle-plates do not, according to existing statements,
take place on quite the same type throughout the Vertebrata,
yet the comparison which has been instituted between Elasmobranchs and other Vertebrates appears to prove that there are
important common features in their development, which may be
regarded as primitive, and as having been inherited from the
ancestors of Vertebrates. These features are (i) the extension
of the body cavity into the vertebral plates, and subsequent
enclosure of this cavity between the two layers of the muscleplates ; (2) the primitive division of the vertebral plate into an
outer (somatic) and an inner (splanchnic) layer, and the formation
of a large part of the voluntary muscular system out of the inner
layer, which in all cases is converted into muscles earlier than
the outer layer.
The conversion of the muscle-plates into muscles. It
will be convenient to commence this subject with a description
of the changes which take place in
such a simple type as that of the
At the time when the muscleplates have become independent
structures they form flat two-layered
oblong bodies enclosing a slit-like
central cavity (fig. 379, mp). The
outer or somatic wall is formed of
simple epithelial -like cells. The
inner or splanchnic wall has however a somewhat complicated structure. It is composed dorsally and
ventrally of a columnar epithelium,
but in its middle portion of the
muscle-cells previously spoken of.
Between these and the central cavity
of the plates the epithelium forming
the remainder of the layer commences to insert itself; so that between the first-formed muscle and
the cavity of the muscle-plate there
appears a thin layer of cells, not
however continuous throughout.
When first formed the muscleplates, as viewed from the exterior,
have nearly straight edges ; soon
however they become bent in the middle, so that the edges have
an obtusely angular form, the apex of the angle being directed
forwards. They are so arranged that the anterior edge of the
one plate fits into the posterior edge of the one in front. In the
lines of junction between the plates layers of connective-tissue
cells appear, which form the commencements of the intermuscular
The growth of the plates is very rapid, and their upper ends
28 F.
sp.c. spinal canal ; W. white
matter of spinal cord ; pr. posterior nerve-roots ; ch. notochord ;
x. sub-notochordal rod ; ao. aorta ;
mp. muscle-plate; mp' . inner layer
of muscle-plate already converted
into muscles ; Vr. rudiment of
vertebral body ; si. segmental
tube ; sd. segmental duct ; sp.v.
spiral valve ; z/. subintestinal vein ;
P.O. primitive generative cells.
soon extend to the summit of the neural canal, and their lower
ones nearly meet in the median ventral line. The original band
of muscles, whose growth at first is very slow, now increases
with great rapidity, and forms the nucleus of the whole voluntary muscular system (fig. 380, mp'). It extends upwards and
downwards by the continuous conversion of fresh cells of the
splanchnic layer into muscle-cells. At the same time it grows
rapidly in thickness by the addition of fresh spindle-shaped
muscle-cells from the somatic layer as well as by the division of
the already existing cells.
Thus both layers of the muscle-plate are concerned in forming
the great longitudinal lateral muscles, though the splanchnic layer
is converted into muscles very much sooner than the somatic 1 .
Each muscle-plate is at first a continuous structure, extending
from the dorsal to the ventral surface, but after a time it becomes
divided by a layer of connective tissue, which becomes developed
nearly on a level with the lateral line, into a dorso-lateral and
a ventro-lateral section. The ends of the muscle-plates
continue for a long time to be formed of undifferentiated
columnar cells. The complicated outlines of the inter-muscular
septa become gradually established during the later stages of
development, causing the well-known appearances of the muscles
in transverse sections, which require no special notice here.
The muscles of the limbs. The limb muscles are formed
in Elasmobranchii, coincidently with the cartilaginous skeleton,
as two bands of longitudinal fibres on the dorsal and ventral
surfaces of the limbs (fig. 346). The cells, from which these
muscles originate, are derived from the muscle-plates. When
the ends of the muscle-plates reach the level of the limbs they
bend outwards and enter the tissue of the limbs (fig. 380).
Small portions of several muscle-plates (m.pl) come in this way
to be situated within the limbs, and are very soon segmented
off from the remainder of the muscle-plates. The portions of
the muscle-plates thus introduced soon lose their original dis
1 The brothers Hertwig have recently maintained that only the inner layer of the
muscle-plates is converted into muscles. In the Elasmobranchs it is easy to demonstrate the incorrectness of this view, and in Acipenser (vide fig. 57, mp) the two layers
of the muscle-plate retain their original relations after the cells of both of them have
become converted into muscles.
B. in. 43
The section is diagrammatic in so far that the anterior nerve-roots have been
inserted for the whole length ; whereas they join the spinal cord half-way between
two posterior roots.
sp.c. spinal cord; sp.g. ganglion of posterior root; ar. anterior root; dn. dorsally
directed nerve springing from posterior root; nip. muscle-plate; mp'. part of muscleplate already converted into muscles; vi.pl. part of muscle-plate which gives rise to
the muscles of the limbs; /. nervus lateralis; ao. aorta; ch. notochord; sy.g. sympathetic ganglion; ca.v. cardinal vein; sp.n. spinal nerve; sd. segmental (archinephric)
duct; st. segmental tube; du. duodenum; pan. pancreas; hp.d. point of junction of
hepatic duct with duodenum ; umc. umbilical canal.
tinctness. There can however be but little doubt that they
supply the tissue for the muscles of the limbs. The muscleplates themselves, after giving off buds to the limbs, grow
downwards, and soon cease to shew any trace of having given
off these buds.
In addition to the longitudinal muscles of the trunk just described,
which are generally characteristic of Fishes, there is found in Amphioxus a
peculiar transverse abdominal muscle, extending from the mouth to the
abdominal pore, the origin of which has not been made out.
It has already been shewn that in all the higher Vertebrata
muscle-plates appear, which closely resemble those in Elasmobranchii; so that all the higher Vertebrata pass through, with
reference to their muscular system, a fish- like stage. The
middle portion of the inner layers of their muscle-plates becomes, as in Elasmobranchii, converted into muscles at a very
early period, and the outer layer for a long time remains formed
of indifferent cells. That these muscle-plates give rise to the
main muscular system of the trunk, at any rate to the episkeletal
muscles of Huxley, is practically certain, but the details of the
process have not been made out.
In the Perennibranchiata the fish-like arrangement of muscles is retained through life in the tail and in the dorso-lateral parts of the trunk.
In the tail of the Amniotic Vertebrata the primitive arrangement is also
more or less retained, and the same holds good for the dorso-lateral trunk
muscles of the Lacertilia. In the other Amniota and the Anura the
dorso-lateral muscles have become divided up into a series of separate
muscles, which are arranged in two main layers. It is probable that the
intercostal muscles belong to the same group as the dorso-lateral muscles.
The abdominal muscles of the trunk, even in the lowest Amphibia,
exhibit a division into several layers. The recti abdominis are the least
altered part of this system, and usually retain indications of the primitive
inter-muscular septa, which in many Amphibia and Lacertilia are also
to some extent preserved in the other abdominal muscles.
In the Amniotic Vertebrates there is formed underneath the vertebral
column and the transverse processes a system of muscles, forming part
of the hyposkeletal system of Huxley, and called by Gegenbaur the subvertebral muscles. The development of this system has not been worked
out, but on the whole I am inclined to believe that it is derived from
the muscle-plates. Kolliker, Huxley and other embryologists believe
however that these muscles are independent of the muscle-plates in their
Whether the muscle of the diaphragm is to be placed in the same
category as the hyposkeletal muscles has not been made out.
It is probable that the cutaneous muscles of the trunk are derived
from the cells given off from the muscle-plates. Kolliker however believes
that they have an independent origin.
The limb-muscles, both extrinsic and intrinsic, as may be concluded
from their development in Elasmobranchii, are derived from the muscleplates. Kleinenberg found in Lacertilia a growth of the muscle-plates
into the limbs, and in Amphibia Gotte finds that the outer layer of the
muscle-plates gives rise to the muscles of the limbs.
In the higher Vertebrata on the other hand the entrance of the muscleplates into the limbs has not been made out (Kolliker). It seems therefore
probable that by an embryological modification, of which instances are so
frequent, the cells which give rise to the muscles of the limbs in the higher
Vertebrata can no longer be traced into a direct connection with the muscleplates.
TJte Somites and muscular system of the head.
The extension of the somites to the anterior end of the body
in Amphioxus clearly proves that somites, similar to those of
the trunk, were originally present in a region, which in the
higher Vertebrata has become differentiated into the head. In
the adult condition no true Vertebrate exhibits indications of
such somites, but in the embryos of several of the lower Vertebrata structures have been found, which are probably equivalent
to the somites of the trunk : they have been frequently alluded
to in the previous chapters of this volume. These structures
have been most fully worked out in Elasmobranchii.
The mesoblast in Elasmobranch embryos becomes first split
into somatic and splanchnic layers in the region of the head ;
and between these layers there are formed two cavities, one on
each side, which end in front opposite the blind anterior extremity of the alimentary canal ; and are continuous behind
with the general body-cavity (fig. 20 A, vp}. I propose calling
them the head-cavities. The cavities of the two sides have
no communication with each other.
Coincidently with the formation of an outgrowth from the
throat to form the first visceral cleft, the head-cavity on each
side becomes divided into a section in front of the cleft and a
section behind the cleft ; and at a later period it becomes, owing
to the formation of a second cleft, divided into three sections :
(i) a section in front of the first or hyomandibular cleft; (2) a
section in the hyoid arch between the hyomandibular cleft and
the hyobranchial or first branchial cleft ; (3) a section behind
the first branchial cleft.
The front section of the head-cavity grows forward, and soon
becomes divided, without the intervention of a visceral cleft, into
an anterior and posterior division.
The anterior lies close to the eye,
and in front of the commencing
mouth involution. The posterior
part lies completely within the mandibular arch.
As the rudiments of the successive visceral clefts are formed, the
posterior part of the head-cavity becomes divided into successive sections, there being one section for
each arch. Thus the whole headcavity becomes on each side divided
into (i) a premandibular section ; (2)
a mandibular section (vide fig. 29 A,
PP] > (3) a hyoid section ; (4) sections
in each of the branchial arches.
The first of these divisions forms
a space of a considerable size, with
epithelial walls of somewhat short
columnar cells (fig. 381, ipp}. It is
situated close to the eye, and presents a rounded or sometimes a
triangular figure in section. The
two halves of the cavity are prolonged ventralwards, and meet below
the base of the fore-brain. The
connection between them appears to last for a considerable time.
These two cavities are the only parts of the body-cavity within
the head which unite ventrally. The section of the head-cavity
just described is so similar to the remaining sections that it
must be considered as serially homologous with them.
The next division of the head-cavity, which from its position
The section, owing to the cranial flexure, cuts both the foreand the hind-brain. It shews the
premandibular and mandibular
head-cavities ipp and ipp, etc.
The section is moreover somewhat
oblique from side to side.
fb. fore-brain ; /. lens of eye ;
m. mouth ; pt. upper end of mouth,
forming pituitary involution; lao.
mandibular aortic arch; ipp. and
ipp. first and second head-cavities;
\vc. first visceral cleft; V. fifth
nerve ; aim. auditory nerve ; VII.
seventh nerve ; aa. dorsal aorta ;
acv. anterior cardinal vein ; ch,
may be called the mandibular cavity, presents a spatulate shape,
being dilated dorsally, and produced ventrally into a long thin
process parallel to the hyomandibular gill-cleft (fig. 20, pp}.
Like the previous space it is lined by a short columnar epithelium.
The mandibular aortic arch is situated close to its inner side
(fig. 381, 2pp). After becoming separated from the lower part
(Marshall), the upper part of the cavity atrophies about the time
of the appearance of the external gills. Its lower part also
becomes much narrowed, but its walls of columnar cells persist.
The outer or somatic wall becomes very thin indeed, the
splanchnic wall, on the other hand, thickens and forms a layer
of several rows of elongated cells. In each of the remaining
arches there is a segment of the original body-cavity fundamentally similar to that in the mandibular arch (fig. 382). A dorsal
dilated portion appears, however, to be present in the third or
hyoid section alone (fig. 20), and even
there disappears very soon, after being
segmented off from the lower part
(Marshall). The cavities in the posterior parts of the head become much
reduced like those in its anterior part,
though at rather a later period. FlG . 382 . HORIZONTAL
sions of the body-cavity in the head, AN EMBRYO OF PRISTIURUS.
with the exception of the anterior, e p. epiblast; vc. pouch of
early become atrophied, not so how- hypoblast which will form the
walls of a visceral cleit ; //.
CVer their walls. The cells forming segment of body-cavity in vis
the walls both of the dorsal and ven- ceral arch ; aa ' aortic arch '
tral sections of these cavities become elongated, and finally
become converted into muscles. Their exact history has not
been followed in its details, but they almost unquestionably
become the musculus contrictor superficialis and musculus interbranchialis 1 ; and probably also musculus levator mandibuli and
other muscles of the front part of the head.
The anterior cavity close to the eye remains unaltered much
longer than the remaining cavities.
1 Vide Vetter, " Die Kiemen und Kiefermusculatur d. Fische." Jenaische Zcltschrift, Vol. vn.
Its further history is very interesting. In my original account
of this cavity (No. 292, p. 208) I stated my belief that its walls
gave rise to the eye-muscles, and the history of this process has
been to some extent worked out by Marshall in his important
memoir (No. 509).
Marshall finds that the ventral portion of this cavity, where
its two halves meet, becomes separated from the remainder.
The eventual fate of this part has not however been followed.
Each dorsal section acquires a cup-like form, investing the
posterior and inner surface of the eye. The cells of its outer
wall subsequently give rise to three sets of muscles. The middle
of these, partly also derived from the inner walls of the cup,
becomes the rectus internus of the eye, the dorsal set forms the
rectus superior, and the ventral the rectus inferior. The obliquus
inferior appears also to be in part developed from the walls of
this cavity.
Marshall brings evidence to shew that the rectus externus (as
might be anticipated from its nerve supply) has no connection
with the walls of the premandibular head-cavity, and finds that
it arises close to the position originally occupied by the second
and third cavities. Marshall has not satisfactorily made out the
mode of development of the obliquus superior.
The walls of the cavities, whose history has just been recorded, have definite relations with the cranial nerves, an account
of which has already been given at p. 461.
Head-cavities, in the main similar to those of Elasmobranchii, have been found in the embryo of Petromyzon (fig. 45,
/ic\ the Newt (Osborn and Scott), and various Reptilia (Parker).
(507) G.M.Humphry. " Muscles in Vertebrate Animals." Journ. of Anat.
and Phys., Vol. vi. 1872.
(508) J. Miiller. " Vergleichende Anatomic d. Myxinoiden. Part I. Osteologie
u. Myologie." Akad. Wiss., Berlin, 1834.
(509) A. M. Marshall. "On the head cavities and associated nerves of
Elasmobranchs." Quart. J. of Micr. Science, Vol. xxi. 1881.
(510) A. Schneider. " Anat. u. Entwick. d. Muskelsystems d. Wirbelthierc."
Silz. d. Oberhessischen Gesellschaft, 1873.
(511) A. Schneider. Beitrdge z. vergleich. Anat. . Entwick. d. Wirbelthiere.
Berlin, 1879.
Vide 2^0 Gotte (No. 296), Kolliker (N o. 298), Balfour (No. 292), Huxley, etc.
EXCRETORY organs consist of coiled or branched and often
ciliated tubes, with an excretory pore opening on the outer surface
of the body, and as a rule an internal ciliated orifice placed in the
body-cavity. In forms provided with a true vascular system,
there is a special development of capillaries around the glandular
part of the excretory organs. In many instances the glandular
cells of the organs are filled with concretions of uric acid or some
similar product of nitrogenous waste.
There is a very great morphological and physiological similarity between almost all the forms of excretory organ found in
the animal kingdom, but although there is not a little to be said
for holding all these organs to be derived from some common
prototype, the attempt to establish definite homologies between
them is beset with very great difficulties.
Platyelminthes. Throughout the whole of the Platyelminthes these organs are constructed on a well-defined type, and
in the Rotifera excretory organs of a similar form to those of the
Platyelminthes are also present.
These organs (Fraipont, No. 513) are more or less distinctly
paired, and consist of a system of wide canals, often united into a
network, which open on the one hand into a pair of large tubes
leading to the exterior, and on the other into fine canals which
terminate by ciliated openings, either in spaces between the
connective-tissue cells (Platyelminthes), or in the body-cavity
(Rotifera). The fine canals open directly into the larger ones,
without first uniting into canals of an intermediate size.
68 1
The two large tubes open to the exterior, either by means of
a median posteriorly placed contractile vesicle, or by a pair of
vesicles, which have a ventral and anterior position. The former
type is characteristic of the majority of the Trematoda, Cestoda.
and Rotifera, and the latter of the Nemertea and some Trematoda.
In the Turbellaria the position of the external openings of the
system is variable, and in a few Cestoda (Wagner) there are
lateral openings on each of the successive proglottides, in addition
to the terminal openings. The mode of development of these
organs is unfortunately not known.
Mollusca. In the Mollusca there are usually present two
independent pairs of excretory organs one found in a certain
number of forms during early larval life only 1 , and the other
always present in the adult.
The larval excretory organ has been found in the pulmonate
Gasteropoda (Gegenbaur, Fol 2 , Rabl), in Teredo (Hatschek), and
possibly also in Paludina. It is placed in the anterior region of
the body, and opens ventrally on each side, a short way behind
the velum. It is purely a larval organ, disappearing before the
close of the veliger stage. In the aquatic Pulmonata, where it is
best developed, it consists on each side of a V-shaped tube, with
a dorsally-placed apex, containing an enlargement of the lumen.
There is a ciliated cephalic limb, lined by cells with concretions,
and terminating by an internal opening near the eye, and a nonciliated pedal limb opening to the exterior 3 .
Two irreconcilable views are held as to the development of
this system. Rabl (Vol. II. No. 268) and Hatschek hold that it
is developed in the mesoblast ; and Rabl states that in Planorbis
it is formed from the anterior mesoblast cells of the mesoblastic
bands. A special mesoblast cell on each side elongates into two
processes, the commencing limbs of the future organ. A lumen
is developed in this cell, which is continued into each limb, while
1 I leave out of consideration an external renal organ found in many marine
Gasteropod larvte, vide Vol. II. p. 280.
2 H. Fol, "Etudes sur le devel. d. Mollusques. " Mem. Hi. Archiv d. Zool.
exfJr. et gener., Vol. VIII.
3 The careful observations of Fol seem to me nearly conclusive in favour of this
limb having an external opening, and the statement to the reverse effect on p. 280 of
Vol. ii. of this treatise, made on the authority of Rabl and Biitschli, must probably be
the continuations of the two limbs are formed by perforated
mesoblast cells.
According to Fol these organs originate in aquatic Pulmonata
as a pair of invaginations of the epiblast, slightly behind the
mouth. Each invagination grows in a dorsal direction, and after
a time suddenly bends on itself, and grows ventralwards and
forwards. It thus acquires its V-shaped form.
In the terrestrial Pulmonata the provisional excretory organs
are, according to Fol, formed as epiblastic invaginations, in the
same way as those in the aquatic Pulmonata, but have the form
of simple non-ciliated sacks, without internal openings.
The permanent renal organ of the Mollusca consists typically
of a pair of tubes, although in the majority of the Gasteropoda
one of the two tubes is not developed. It is placed considerably
behind the provisional renal organ.
Each tube, in its most typical form, opens by a ciliated funnel
into the pericardial cavity, and has its external opening at the
side of the foot. The pericardial funnel leads into a glandular
section of the organ, the lining cells of which are filled with
concretions. This section is followed by a ciliated section, from
which a narrow duct leads to the exterior.
As to the development of this organ the same divergence of
opinion exists as in the case of the provisional renal organ.
Rabl's careful observations on Planorbis (Vol. II. No. 268) tend
to shew that it is developed from a mass of mesoblast cells, near
the end of the intestine. The mass becomes hollow, and,
attaching itself to the epiblast on the left side of the anus,
acquires an opening to the exterior. Its internal opening is not
established till after the formation of the heart. Fol gives an
equally precise account, but states that the first rudiment of the
organ arises as a solid mass of epiblast cells. Lankester finds
that this organ is developed as a paired invagination of the.
epiblast in Pisidium, and Bobretzky also derives it from the
epiblast in marine Prosobranchiata. In Cephalopoda on the
other hand Bobretzky's observations (I conclude this from his
figures) indicate that the excretory sacks of the renal organs are
derived from the mesoblast.
Polyzoa. Simple excretory organs, consisting of a pair of
ciliated canals, opening between the mouth and the anus, have
68 3
been found by Hatschek and Joliet in the Entoproctous Polyzoa,
and are developed, according to Hatschek, by whom they were
first found in the larva, from the mesoblast
Brachiopoda. One or rarely two (Rhynchonella) pairs of
canals, with both peritoneal and external openings, are found in
the Brachiopoda. They undoubtedly serve as genital ducts, but
from their structure are clearly of the same nature as the
excretory organs of the Chaetopoda described below. Their
development has not been worked out.
Chaetopoda. Two forms of excretory organ have been met
with in the Chaetopoda. The one form is universally or nearly
universally present in the adult, and typically consists of a pair
of coiled tubes repeated in every segment. Each tube has an
internal opening, placed as a rule in the segment in front of that
in which the greater part of the organ and the external opening
are situated.
There are great variations in the structure of these organs,
which cannot be dealt with here. It may be noted however that
the internal opening may be absent, and that there may be
several internal openings for each organ (Polynoe). In the
Capitellidae moreover several pairs of excretory tubes have been
shewn by Eisig (No. 512) to be present in each of the posterior
The second form of excretory organ has as yet only been
found in the larva of Polygordius, and will be more conveniently
dealt with in connection with the development of the excretory
system of this form.
There is still considerable doubt as to the mode of formation
of the excretory tubes of the Chaetopoda. Kowalevsky (No. 277),
from his observations on the Oligochasta, holds that they develop
as outgrowths of the epithelial layer covering the posterior side
of the dissepiments, and secondarily become connected with the
Hatschek finds that in Criodrilus they arise from a continuous
linear thickening of the somatic mesoblast, immediately beneath
the epidermis, and dorsal to the ventral band of longitudinal
muscles. They break up into S-shaped cords, the anterior end
of each of which is situated in front of a dissepiment, and is
formed at first of a single large cell, while the posterior part is
continued into the segment behind. The cords are covered by
a peritoneal lining, which still envelopes them, when in the
succeeding stage they are carried into the body-cavity. They
subsequently become hollow, and their hinder ends acquire
openings to the exterior. The formation of their internal
openings has not been followed.
Kleinenberg is inclined to believe that the excretory tubes
take their origin from the epiblast, but states that he has not
satisfactorily worked out their development.
The observations of Risig (No. 512) on the Capitellidae
support Kowalevsky's view that the excretory tubes originate
from the lining of the peritoneal cavity.
Hatschek (No. 514) has given a very interesting account of
the development of the excretory system in Polygordius.
The excretory system begins to be formed, while the larva is
still in the trochospere stage (fig. 383, npli), and consists of a
provisional excretory organ, which is placed in front of the future
segmented part of the body, and occupies a position very
similar to that of the provisional excretory organ found in some Molluscan
larvae (vide p. 68 1).
Hatschek, with some shew of reason, holds that the provisional excretory organs of Polygordius are homologous with those of the Mollusca.
In its earliest stage the provisional
excretory organ of Polygordius consists of a pair of simple ciliated tubes, FIG. 383. POLYOORDIUS
, . , r 11-1 LARVA. (After Hatschek.)
each with an anterior funnel-like open- m _ moulh . ^ supraKBSO .
ing situated in the midst of the meSO- phageal ganglion ; nph. nephri11 11 . , dion ; ine.p. mesoblastic band;
blast cells, and a posterior external an _ anus 5 oL stomach .
opening. The latter is placed immediately in front of what afterwards becomes the segmented region
of the embryo. While the larva is still unsegmented, a second
internal opening is formed for each tube (fig. 383, np/i) and the
two openings so formed may eventually become divided into
five (fig. 384 A), all communicating by a single pore with the
When the posterior region of the embryo becomes segmented,
paired excretory organs are formed in each of the posterior
segments, but the account of their development, as given by
Hatschek, is so remarkable that I do not think it can be
definitely accepted without further confirmation.
From the point of junction of the two main branches of the
larval kidney there grows backwards (fig. 384 B), to the hind
end of the first segment, a very delicate tube, only indicated by
its ciliated lumen, its walls not being differentiated. Near the
front end of this tube a funnel, leading into the larval body
cavity of the head, is formed, and subsequently the posterior end
of the tube acquires an external opening, and the tube distinct
walls. The communication with the provisional excretory organ
is then lost, and thus the excretory tube of the first segment is
The excretory tubes in the second and succeeding segments
are formed in the same way as in the first, i.e. by the continuation of the lumen of the hind end of the excretory tube from
the preceding segment, and the subsequent separation of this
part as a separate tube.
The tube may be continued with a sinuous course through
several segments without a distinct wall. The external and
internal openings of the permanent excretory tubes are thus
secondarily acquired. The internal openings communicate with
the permanent body-cavity. The development of the perma
nent excretory tubes is diagrammatically represented in fig.
384 C and D.
The provisional excretory organ atrophies during larval life.
If Hatschek's account of the development of the excretory system of
Polygordius is correct, it is clear that important secondary modifications
must have taken place in it, because his description implies that there sprouts
from the anterior excretory organ, while it has its own external opening, a
posterior duct, which does not communicate either with the exterior or with
the body-cavity! Such a duct could have no function. It is intelligible
either (i) that the anterior excretory organ should lead into a longitudinal
duct, opening posteriorly ; that then a series of secondary openings into the
body-cavity should attach themselves to this, that for each internal opening
an external should subsequently arise, and the whole break up into separate
tubes ; or (2) that behind an anterior provisional excretory organ a series of
secondary independent segmental tubes should be formed. But from Hatschek's account neither of these modes of evolution can be deduced.
Gephyrea. The Gephyrea may have three forms of excretory organs, two of which are found in the adult, and one,
similar in position and sometimes also in structure, to the
provisional excretory organ of Polygordius, has so far only been
found in the larvae of Echiurus and Bonellia.
In all the Gephyrea the so-called 'brown tubes' are
apparently homologous with the segmented excretory tubes of
Chaetopods. Their main function appears to be the transportation of the generative products to the exterior. There is but a
single highly modified tube in Bonellia, forming the oviduct and
uterus ; a pair of tubes in the Gephyrea inermia, and two or
three pairs in most Gephyrea armata, except Bonellia. Their
development has not been studied.
In the Gephyrea armata there is always present a pair of
posteriorly placed excretory organs, opening in the adult into
the anal extremity of the alimentary tract, and provided with
numerous ciliated peritoneal funnels. These organs were stated
by Spengel to arise in Bonellia as outgrowths of the gut ; but in
Echinrus Hatschek (No. 515) finds that they are developed from
the somatic mesoblast of the terminal part of the trunk. They
soon become hollow, and after attaching themselves to the
epiblast on each side of the anus, acquire external openings.
They are not at first provided with peritoneal funnels, but these
parts of the organs become developed from a ring of cells at
their inner extremities ; and there is at first but a single funnel
for each vesicle. The mode of increase of the funnels has not
been observed, nor has it been made out how the organs themselves become attached to the hind-gut.
The provisional excretory organ of Echiurus is developed at
an early larval stage, and is functional during the whole of
larval life. It at first forms a ciliated tube on each side, placed
in front of that part of the larva which becomes the trunk of the
adult. It opens to the exterior by a fine pore on the ventral
side, immediately in front of one of the mesoblastic bands, and
appears to be formed of perforated cells. It terminates internally in a slight swelling, which represents the normal internal
ciliated funnel. The primitively simple excretory organ becomes
eventually highly complex by the formation of numerous
branches, each ending in a slightly swollen extremity. These
branches, in the later larval stages, actually form a network, and
the inner end of each main branch divides into a bunch of fine
tubes. The whole organ resembles in many respects the excretory organ of the Platyelminthes.
In the larva of Bonellia Spengel has described a pair of
provisional excretory tubes, opening near the anterior end of
the body, which are probably homologous with the provisional
excretory organs of Echiurus (vide Vol. II., fig. 162 C, se).
Discophora. As in many of the types already spoken of,
permanent and provisional excretory organs may be present in
the Discophora. The former are usually segmentally arranged,
and resemble in many respects the excretory tubes of the
Chaetopoda. They may either be provided with a peritoneal
funnel (Nephelis, Clepsine) or have no internal opening
Bourne 1 has shewn that the cells surrounding the main duct
in the medicinal Leech are perforated by a very remarkable
network of ductules, and the structure of these organs in the
Leech is so peculiar that it is permissible to state with due reserve
their homology with the excretory organs of the Chaetopoda.
The excretory tubes of Clepsine are held by Whitman to be
developed in the mesoblast.
1 "On the Structure of the Nephridia of the Medicinal Leech." Quart. J. of
Micr. Science, Vol. XX. 1880.
There are found in the embryos of Nephelis and Hirudo
certain remarkable provisional excretory organs the origin and
history of which are not yet fully made out. In Nephelis they
appear as one (according to Robin), or (according to Biitschli)
as two successive pairs of convoluted tubes on the dorsal side of
the embryo, which are stated by the latter author to develop
from the scattered mesoblast cells underneath the skin. At
their fullest development they extend, according to Robin, from
close to the head to near the ventral sucker. Each of them is
U-shaped, with the open end of the U forwards, each limb of the
U being formed by two tubes united in front. No external
opening has been clearly made out. Fiirbringer is inclined from
his own researches to believe that they open laterally. They
contain a clear fluid.
In Hirudo, Leuckart has described three similar pairs of
organs, the structure of which he has fully elucidated. They
are situated in the posterior part of the body, and each of them
commences with an enlargement, from which a convoluted tube
is continued for some distance backwards; the tube then turns
forwards again, and after bending again upon itself opens to the
exterior. The anterior part is broken up into a kind of
labyrinthic network.
The provisional excretory organs of the Leeches cannot be
identified with the anterior provisional organs of Polygordius
and Echiurus.
Arthropoda. Amongst the Arthropoda Peripatus is the
only form with excretory organs of the type of the segmental
excretory organs of the Chsetopoda 1 .
These organs are placed at the bases of the feet, in the
lateral divisions of the body-cavity, shut off from the main
median division of the body-cavity by longitudinal septa of
transverse muscles.
Each fully developed organ consists of three parts :
(i) A dilated vesicle opening externally at the base of a
foot. (2) A coiled glandular tube connected with this, and
subdivided again into several minor divisions. (3) A short
terminal portion opening at one extremity into the coiled tube
1 Vide F. M. Balfour, " On some points in the Anatomy of Peripatus Capensis."
Quart. J, of Micr. Science, Vol. XIX. 1879.
and at the other, as I believe, into the body cavity. This
section becomes very conspicuous, in stained preparations, by
the intensity with which the nuclei of its walls absorb the
colouring matter.
In the majority of the Tracheata the excretory organs have
the form of the so-called Malpighian tubes, which always (vide
Vol. II.) originate as a pair of outgrowths of the epiblastic
proctodaeum. From their mode of development they admit of
comparison with the anal vesicles of the Gephyrea, though in
the present state of our knowledge this comparison must be
regarded as somewhat hypothetical.
The antennary and shell-glands of the Crustacea, and
possibly also the so-called dorsal organ of various Crustacean
larvae appear to be excretory, and the two former have been
regarded by Claus and Grobben as belonging to the same
system as the segmental excretory tubes of the Chaetopoda.
Nematoda. Paired excretory tubes, running for the whole
length of the body in the so-called lateral line, and opening in
front by a common ventral pore, are present in the Nematoda.
They do not appear to communicate with the body cavity, and
their development has not been studied.
Very little is known with reference either to the structure or
development of excretory organs in the Echinodermata and the
other Invertebrate types of which no mention has been so far
made in this Chapter.
Excretory organs and generative ducts of the Craniata.
Although it would be convenient to separate, if possible, the
history of the excretory organs from that of the generative
ducts, yet these parts are so closely related in the Vertebrata, in
some cases the same duct having at once a generative and a
urinary function, that it is not possible to do so.
The excretory organs of the Vertebrata consist of three
distinct glandular bodies and of their ducts. These are (i) a
small glandular body, usually with one or more ciliated funnels
opening into the body cavity, near the opening of which there
projects into the body cavity a vascular glomerulus. It is
situated very far forwards, and is usually known as the head
kidney, though it may perhaps be more suitably called, adopting
Lankester's nomenclature, the pronepliros. Its duct, which forms
the basis for the generative and urinary ducts, will be called the
segmented duct.
(2) The Wolffian body, which may be also called the
mesonepJiros. It consists of a series of, at first, segmentally
(with a few exceptions) arranged glandular canals (segmental
tubes) primitively opening at one extremity by funnel-shaped
apertures into the body cavity, and at the other into the
segmental duct. This duct becomes in many forms divided
longitudinally into two parts, one of which then remains
attached to the segmental tubes and forms the Wolffian or
mesonepJiric duct, while the other is known as the Milllerian
(3) The kidney proper or metanephros. This organ is only
found in a completely differentiated form in the amniotic Vertebrata. Its duct is an outgrowth from the Wolrfian duct.
The above parts do not coexist in full activity in any living
adult member of the Vertebrata, though all of them are found
together in certain embryos. They are so intimately connected
that they cannot be satisfactorily dealt with separately.
Elasmobranchii. The excretory system of the Elasmobranchii is by no means the most primitive known, but at the
same time it forms a convenient starting point for studying the
modifications of the system in other groups. The most remarkable peculiarity it presents is the absence of a pronephros.
The development of the Elasmobranch excretory system has
been mainly studied by Semper and myself.
The first trace of the system makes its appearance as a knob
of mesoblast, springing from the intermediate cell-mass near the
level of the hind end of the heart (fig. 385 K,pd). This knob is
the rudiment of the abdominal opening of the segmental duct,
and from it there grows backwards to the level of the anus a
solid column of cells, which constitutes the rudiment of the
segmental duct itself (fig. 385 B, pd). The knob projects
towards the epiblast, and the column connected with it lies
between the mesoblast and epiblast. The knob and column do
not long remain solid, but the former acquires an opening into
the body cavity (fig. 421, sd) continuous with a lumen, which
makes its appearance in the column (fig. 386, sd). The knob
forms the only structure which can be regarded as a rudiment of
the pronephros.
The sections illustrate the development of the segmental duct (pd) or primitive
duct of the pronephros. In A (the anterior of the two sections) this appears as a
solid knob (pd) projecting towards the epiblast. In B is seen a section of the column
which has grown backwards from the knob in A.
spn. rudiment of a spinal nerve; me. medullary canal; ch. notochord; X. subnotochordal rod; mp. muscle-plate; mp' . specially developed portion of muscle-plate;
ao. dorsal aorta ; pd. segmental duct ; so. somatopleure ; sp. splanchnopleure ; //.
body cavity; ep. epiblast; al. alimentary canal.
While the lumen is gradually being formed, the segmental
tubes of the mesonephros become established. They appear to
arise as differentiations of the parts of the primitive lateral plates
of mesoblast, placed between the dorsal end of the body cavity
and the muscle-plate (fig. 386, st) 1 , which are usually known as
the intermediate cell-masses.
The lumen of the segmental tubes, though at first very small,
soon becomes of a considerable size. It appears to be established
in the position of the section of the body cavity in the intermediate cell-mass, which at first unites the part of the body
cavity in the muscle-plates with the permanent body cavity.
The lumen of each tube opens at its lower end into the dorsal
part of the body cavity (fig. 386, st}, and each tube curls obliquely
1 In my original account of the development I held these tubes to be invaginations
of the peritoneal epithelium. Sedgwick (No. 549) was led to doubt the accuracy of
my original statement from his investigations on the chick ; and from a re-examination of my specimens he arrived at the results stated above, and which I am now
myself inclined to adopt.
backwards round the inner and dorsal side of the segmental
duct, near which it at first ends blindly.
One segmental tube makes its
appearance for each somite (fig. 265),
commencing with that immediately
behind the abdominal opening of the
segmental duct, the last tube being
situated a few segments behind the
anus. Soon after their formation
the blind ends of the segmental tubes
come in contact with, and open into
the segmental duct, and each of them
becomes divided into four parts.
These are (i) a section carrying the
peritoneal opening, known as the
peritoneal funnel, (2) a dilated vesicle
into which this opens, (3) a coiled
tubulus proceeding from (2), and
terminating in (4) a wider portion
opening into the segmental duct. At
the same time, or shortly before this,
each segmental duct unites with and
opens into one of the horns of the
cloaca, and also retires from its
primitive position between the epiblast and mesoblast, and assumes a
position close to the epithelium lining
the body cavity (fig. 380, sd}. The
general features of the excretory
organs at this period are diagrammatically represented in the
woodcut (fig. 387). In this fig. pd is the segmental duct and
o its abdominal opening; s.t points to the segmental tubes,
the finer details of whose structure are not represented in the
diagram. The mesonephros thus forms at this period an elongated gland composed of a series of isolated coiled tubes, one
extremity of each of which opens into the body cavity, and the
other into the segmental duct, which forms the only duct of the
system, and communicates at its front end with the body cavity,
and behind with the cloaca.
28 F.
sp.c. spinal canal; W. white
matter of spinal cord ; pr. posterior nerve-roots ; ch. notochord ;
x. sub-notochordal rod ; ao. aorta ;
nip, muscle-plate ; nip', inner layer
of muscle-plate already converted
into muscles ; Vr, rudiment of
vertebral body ; st. segmental
tube; sd. segmental duct; sp.v.
spiral valve ; v. subintestinal vein ;
p.o. primitive generative cells.
The next important change concerns the segmental duct,
which becomes longitudinally split into two complete ducts in
the female, and one complete duct and parts of a second duct in
the male. The manner in which this takes place is diagrammatically represented in fig. 387 by the clear line x, and in
transverse section in figs. 388 and 389. The resulting ducts are
(i) the Wolffian duct or mesonephric duct (wd\ dorsally, which
remains continuous with the excretory tubules of the mesonephros, and ventrally (2) the oviduct or Miillerian duct in the
female, and the rudiments of this duct in the male. In the
pd. segmental duct. It opens at o into the body cavity and at its other extremity
into the cloaca; x. line along which the division appears which separates the segmental
duct into the Wolffian duct above and the Miillerian duct below; s.t. segmental
tubes. They open at one end into the body cavity, and at the other into the segmental duct.
female the formation of these ducts takes place (fig. 389) by a
nearly solid rod of cells being gradually split off from the
ventral side of all but the foremost part of the original segmental
duct. This nearly solid cord is the Miillerian duct (pd}. A
very small portion of the lumen of the original segmental duct
is perhaps continued into it, but in any case it very soon acquires
a wide lumen (fig. 389 A). The anterior part of the segmental
duct is not divided, but remains continuous with the Mullerian
duct, of which its anterior pore forms the permanent peritoneal
opening 1 (fig. 387). The remainder of the segmental duct (after
the loss of its anterior section, and the part split off from its
ventral side) forms the Wolffian duct. The process of formation
of these ducts in the male differs from that in the female chiefly
1 Five or six segmental tubes belong to the region of the undivided anterior part
of the segmental duct, which forms the front end of the Mullerian duct ; but they appear to atrophy very early, without acquiring a definite attachment to the segmental
in the fact of the anterior undivided part of the segmental duct,
which forms the front end of the Miillerian duct, being shorter,
The figure shews how the
segmental duct becomes split
into the Wolffian or mesonephric duct above, and Miillerian duct or oviduct below.
wd. Wolffian or mesonephric duct; od. Miillerian
duct or oviduct ; sd. segmental duct.
me. medullary canal; mp. muscle-plate;
ch. notochord; ao. aorta; cav. cardinal
vein; st. segmental tube. On the left side
the section passes through the opening of
a segmental tube into the body cavity. On
the right this opening is represented by
dotted lines, and the opening of the segmental tube into the Wolffian duct has
been cut through; iv.d. Wolffian duct;
m.d. Miillerian duct. The section is taken
through the point where the segmental
duct and Wolffian duct have just become
separate; gr. the germinal ridge with the
thickened germinal epithelium ; /. liver ;
i. intestine with spiral valve.
and in the column of cells with which it is continuous being
from the first incomplete.
The segmental tubes of the mesonephros undergo further
important changes. The vesicle at the termination of each peritoneal funnel sends a bud forwards towards the preceding
tubulus, which joins the fourth section of it close to the opening
into the Wolffian duct (fig. 390, px). The remainder of the
vesicle becomes converted
into a Malpighian body (mg}.
By the first of these changes 10^-4 M @W>f
a tube is established connecting each pair of segments
of the mesonephros, and
though this tube is in part
aborted (or only represented
by a fibrous band) in the
anterior part of the excretory
organs in the adult, and most
probably in the hinder part,
yet it seems almost certain
that the secondary and tertiary Malpighian bodies of
the majority of segments are
developed from its persisting
blind end. Each of these
The figure contains two examples of the
budding of the vesicle of a segmental tube
(which forms a Malpighian body in its own
segment) to unite with the tubulus in the
preceding segment close to its opening into
the Wolffian (mesonephric) duct.
ge. epithelium of body-cavity; st. peritoneal funnel of segmental tube with its
peritoneal opening; mg. Malpighian body;
px. bud from Malphigian body uniting with
preceding segment.
secondary and tertiary Malpighian bodies is connected with a
convoluted tubulus (fig. 391, a.mg), which is also developed from
the tube connecting each pair of segmental tubes, and therefore
falls into the primary tubulus close to its junction with the
The figure shews a fibrous band passing from the primary to the secondary Malpighian bodies in two segments, which is the remains of the outgrowth from the
primary Malpighian body.
sf.o. peritoneal funnel; p. ing. primary Malpighian body; a.mg. accessory Malpighian body; w.d. mesonephric (Wolffian) duct.
segmental duct. Owing to the formation of the accessory tubuli
the segments of the mesonephros acquire a compound character.
The third section of each tubulus becomes by continuous
growth, especially in the hinder segments, very bulky and
The general character of a slightly developed segment of
the mesonephros at its full growth may be gathered from fig.
391. It commences with (i) a peritoneal opening, somewhat
oval in form (st.d) and leading directly into (2) a narrow tube,
the segmental tube, which takes a more or less oblique course
backwards, and, passing superficially to the Wolffian duct (w.d},
opens into (3) a Malpighian body (p.mg) at the anterior extremity of an isolated coil of glandular tubuli. This coil forms
the third section of each segment, and starts from the Malpighian body. It consists of a considerable number of rather
definite convolutions, and after uniting with tubuli from one,
two, or more (according to the size of the segment) accessory
Malpighian bodies (a.mg) smaller than the one into which the
segmental tube falls, eventually opens by (4) a narrowish
collecting tube into the Wolffian duct at the posterior end of
the segment. Each segment is probably completely isolated
from the adjoining segments, and never has more than one
peritoneal funnel and one communication with the Wolffian duct.
Up to this time there has been no distinction between the
anterior and posterior tubuli of the mesonephros, which alike
open into the Wolffian duct. The collecting tubes of a considerable number of the hindermost tubuli (ten or eleven in
Scyllium canicula), either in some species elongate, overlap,
while at the same time their openings travel backward so that
they eventually open by apertures (not usually so numerous as
the separate tubes), on nearly the same level, into the hindermost section of the Wolffian duct in the female, or into the
urinogenital cloaca, formed by the coalesced terminal parts of
the Wolffian ducts, in the male; or in other species become
modified, by a peculiar process of splitting from the Wolnian
duct, so as to pour their secretion into a single duct on each
side, which opens in a position corresponding with the numerous
ducts of the other species (fig. 392). In both cases the modified
posterior kidney-segments are probably equivalent to the per
manent kidney or metanephros of the amniotic Vertebrates, and
for this reason the numerous collecting tubes or single collecting
tube, as the case may be, will be spoken of as ureters. The
anterior tubuli of the primitive excretory organ retain their early
relation to the Wolffian duct, and form the permanent Wolffian
body or mesonephros.
The originally separate terminal extremities of the Wolffian
ducts always coalesce, and form a urinal cloaca, opening by a
single aperture, situated at the extremity of the median papilla
behind the anus. Some of the peritoneal openings of the segmental tubes in Scyllium, or in other cases all the openings,
become obliterated.
In the male the anterior segmental tubes undergo remarkable modifications, and become connected with the testes.
Branches appear to grow from the first three or four or more of
them (though probably not from their peritoneal openings),
which pass to the base of the testis, and there uniting into a
longitudinal canal, form a network, and receive the secretion of
the testicular ampullae (fig. 393, nf). These ducts, the vasa
efferent ia, carry the semen to the Wolffian body, but before
opening into the tubuli of this body they unite into a canal
known as the longitudinal canal of the Wolffian body (l.c\ from
which pass off ducts equal in number to the vasa efferentia,
each of which normally ends in a Malpighian corpuscle. From
the Malpighian corpuscles so connected there spring the convoluted tubuli, forming the generative segments of the Wolffian
body, along which the semen is conveyed to the Wolffian duct
(v.d). The Wolffian duct itself becomes much contorted and
acts as vas deferens.
Figs. 392 and 393 are diagrammatic representations of the
chief constituents of the adult urinogenital organs in the two
sexes. In the adult female (fig. 392), there are present the
following parts :
(1) The oviduct or Mullerian duct (m.d) split off from the
segmental duct of the kidneys. Each oviduct opens at its
anterior extremity into the body cavity, and behind the two
oviducts have independent communications with the general
(2) The mesonephric ducts (w.d), the other product of the
segmental ducts of the kidneys. They end in front by becoming continuous with the tubulus of the anterior persisting
segment of the mesonephros on each side, and unite behind to
m.d. Miillerian duct; w.d. Wolffian duct; s.t. segmental tubes; five of them are
represented with openings into the body cavity, the posterior segmental tubes form
the mesonephros ; ov. ovary.
open by a common papilla into the cloaca. The mesonephric
duct receives the secretion of the anterior tubuli of the primitive
(3) The ureter which carries off the secretion of the kidney