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   Manual of Human Embryology II 1912: Nervous System | Chromaffin Organs and Suprarenal Bodies | Sense-Organs | Digestive Tract and Respiration | Vascular System | Urinogenital Organs | Figures 2 | Manual of Human Embryology 1 | Figures 1 | Manual of Human Embryology 2 | Figures 2 | Franz Keibel | Franklin Mall | Embryology History
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XX. The Interdependence of the Various Developmental Processes

By Franz Keibel. Freiburg i. Br.

Except in the chapters in which the earlier stages of development are considered and in that in which a review of the entire development and of the elaboration of the external form is given, the development of the various organs and systems of organs has been so far presented as if it took place for the most part individually and independently for each organ or system. We have now to consider how the development of the individual organs combines to produce the development of the whole. It is clear that the normal fully developed organism cannot be produced without a certain regular succession in the development of the organs and a regular interdependence of the individual developmental processes, but it is open to question whether this interdependence is the result of the individual and independent development of each organ taking place in such a way that it fits into that of the others or whether it is due to the individual anlagen of an organism mutually influencing one another during development so as to cause the formation of a normal organism. Both these views have their supporters. Roux (1893) advocates the theory that the various cell complexes of the ovum differentiate independently even from the segmentation stages (Mosaictheory).


Mehnert (1895) says: " The study of the individual time differences in the differentiation of the same organ within the same species shows that a constant correlation of the organ development in the same embryo does not exist." " The frequently striking lack of correlation in the development of organs shows that each developmental process in an organ is to a certain extent an independent process. The development of a vertebrate embryo consists of a series of successive individual processes, regulated only by phylogenetic relations. Only the form and position of any organ can be modified by the environment, ' : Also, so worthy and thoughtful an observer as Born (1897), in his investigations in regeneration in Amphibian larvae, comes to the conclusion that the development from its beginning (that is to say, from larvae in which the medullary canal had just closed and the tail-bud had begun to grow out) 1 depends essentially on the self -differentiation of the individual parts. ' ' A correlation influence of the neighboring parts or of the whole caD never be perceived — neither a negative nor a positive one. ,; " The development corresponds throughout to the mosaic theory of Roux." On the other hand, His has incorrectly been claimed as an advocate of this view. He says (1874) in the sixteenth letter of " Unsere Korperform": "The orderly connection of all the processes underlying the development of the body is a principle which the theory of descent must take into consideration in the future to a greater extent than it has up to the present. So long as investigators of phylogenetic problems have been content to sketch out special histories of individual organs or parts of organs, so long have they perceived the task to be accomplished only in what is certainly a very limited portion of its actual extent; for the development of each individual organ is always merely a dependent partial phenomenon of a huge total process, which links itself up in all directions.' 1 He expresses himself similarly in his paper on " Die Entwicklung der menschlichen und tierischen Physiognomien " (1892), and in his treatise on "Der Prinzip der organbildenden Keimbezirke und die Verwandtschaften der Gewebe " (1901) he assumes the existence not only of spatial and temporal but also of chemical influences ; consequently the principle of " organ forming germinal areas " that His has established is not to be interpreted in the sense of Roux's mosaic theory.


A standpoint similar to that of His is also taken by 0. Hertwig. In various places in his writings he maintains that all the individual parts of the embryo always develop in relation to one another and the development of each part is dependent on the development of the whole (compare 0. Hertwig, 1892 and 1906). This is the view that I also hold, and in what follows I shall endeavor to supply further reasons in support of it. I am of the opinion that the individual anlagen of an organism mutually influence one another in the development so as to contribute to the production of a normal organism. I do not in this dispute the fact that a relatively great individuality, varying in amount in different species of animals, occurs in the development of individual organs and organ complexes. That this is so is shown by the investigations of Born, already referred to, and also by numerous malformations, of man as well as of other animals; in these, individual organs or parts of organs may be developed relatively normally, while others may be abnormal to a high degree or may even be wanting. But neither by the experiments of Born and others, among which those of W. H. Lewis (1907) may be especially mentioned, nor yet by such malformations is it disproved that the individual developmental processes influence one another.


  • 1 For earlier stages there are similar investigations by W. H. Lewis, who transplanted pieces of the dorsal and lateral lip of the blastopore of Bana palustria and was able to show that even the different parts of the blastopore lip possessed in a high degree the power of self-differentiation.


Fischel (1896) has been led to assume such an influence from the measurements he has made of duck embryos with from 1 to 20 primitive segments. He finds that the older the embryo the more regular is the relation of the individual portions of the body one to another, and assumes that during the development regulating influences make themselves felt and bring it about that gradually a distinctly orderly structure of the body supervenes, whereby variations become slighter and rarer." He regards the correlation of the developing organs as such a regulating influence. It may be remarked that long ago Karl Ernst von Baer (1828) arrived at similar conclusions. Furthermore, one may appeal to experimental observations for evidence of the existence of a correlation between the individual parts of a developing ovum, as, for instance, the fundamental experiments in which several embryos are obtained from a single ovum, the experiments of Lewis (1904) and Spemann (1901, 1903) on the relation of the development of the lens to the optic cup and numerous experiments in regeneration.


In many of these experiments one may imagine a pure contact action, in others one must assume action at a distance. Undoubtedly action at a distance obtains when the development is influenced by the reproductive gland. I do not intend to enter into details here as to the effects of castration, but will merely recall the influence that it has unquestionably been shown to exert on the formation and growth of the bones. The epiphyses persist for a longer time and so the growth also lasts longer (compare Sellheim, 1899). Very interesting for their bearings on the doctrine of action at a distance in development are the investigations of Paton and Goodall (1904) and Paton (1904) on guineaKins, and especially those of Hammar (1909) and his pupils, Soederlund and Backman (1909), Jonson (1909) and Syk (1909) on the development of the thymus in the rabbit and man. The human thymus and also that of the mammalia grows until the onset of sexual maturity and then undergoes a rapid involution. Also the occurrence of an age-involution of the lymphoid tissue (Berry and Lack, 1906) may be mentioned. Furthermore, Hammar's pupil Lindberg has found that at about the time of puberty the number of the blood lymphocytes distinctly diminishes. To what extent the suprarenal bodies, as well as the functioning reproductive glands, inhibit a development of the thymus, acting as thymus depressors, and to what extent the thyreoid, the hypophysis, and the parathyreoids favor it, acting as thymus excitators, will not be further considered here. These are investigations that have invaded new territories and still need much enlargement.


The investigation of the interdependence of the various developmental processes has been the purpose of the publication by Keibel (since 1897) of the Normentafel zur Entwicklungsgeschichte der Wirbeltiere. A Normentafel zur Entwicklungsgeschichte des Menschen has been published by Keibel and Elze (1908) and some of the tables given in this may find place here.


Note - Embryos are currently being formatted as separate tables. Original tables spanned 2 pages.

Klb

Human Embryo Pfannenstiel “Klb
Kroemer-Pfannenstiel. Collection of Prof. Pfannenstiel, Giessen. Carnegie stage 10
Desig. Size Age Body form Primitive streak Primitive segments Chorda Nervous system Eye Ear Nose Hypophysis Mouth Digestive trac, liver and pancreas Branchial pouches, threoid thymus, trachea and lungs Urogenital system Heart and vessels Integument Skeleton Extremities Amnion Allantois Remarks
3 Human embryo Klb. N.T. Fig. 3a and 3d; Text-fig 5a to 5n. 138 sections of 10 μ = 1.38 mm. The age is estimated by Born at from 10 to 14 days. No dorsal bend. Head end bent down at right angles and cut by 24 sections of 10 μ. Still indications of a neurenteric canal. Short primitive streak. CIoacal membrane. 5-6 pairs of primitive segments. The chorda throughout is contained in the entoderm. The medullary canal is everywhere wide open, but the brain portion is aire ady marked off from the cord and shows beginning segmentation. Optic anlage not yet evident. Primary pharyngeal membrane No oral sinus. The 1st branchial pouch is formed but does not reach the ectoderm. Heart ventral, but still paired. Paired aortse. 1st branchial arch artery. Aa. umbilicales. Yv. omphalo-mesentericae. Vessels on the yolk sack full of blood corpuscles. No amniotic duct evident. Allantoic duct Obtained by laparotomy.
Size

Kroemer gives the following data: "The greatest length of the embryonic anlage from the anterior edge of the amnion of the head cap to the chorion end of the belly-stalk 1.95 mm, the length of the embryo without the belly-stalk from the head to the tail 1.8 mm., the greatest width of the yolk sack barely 1.2 mm, the width of the embryonic disk at the boundary between the amnion and yolk sack (measured in the head view) 0.9 mm. The measurements of the yolk sack were 1.1 mm, (height), 1.4 mm (width), 1.5 mm. (length)."

Remarks

Fixation: Mailer's fluid. Stain: alum carmine; paraffin. Sections: 10 micron Transverse

Literature: Pfannenstiel in Handbuch der Geburtshilfe, published by Winckel, Wiesbaden 1903. — Kroemer, Wachsmodell eines jungen menschl. Embryo. Verhandl. d. Ges. f. Gynakologie, 1903. The wide pericardial cavity is not connected with the peripheral ccelom. The embryonic ccelom is being formed caudally. On the chorion the layer of Langhans cells and syncytium.

Reference: Keibel F. and Elze C. Normal Plates of the Development of the Human Embryo (Homo sapiens). (1908) Vol. 8 in series by Keibel F. Normal plates of the development of vertebrates (Normentafeln zur Entwicklungsgeschichte der Wirbelthiere) Fisher, Jena., Germany.
Cited in: 1912 Human Embryology
1908 Embryo Tables: Klb (stage 10) | Pfannenstiel III (stage 11) | Meyer 300 (stage 12) | Strahl 4mm (stage 13) | Hertwig G31 (stage 14)

Pfannenstiel III

Human Embryo Pfannenstiel III
Collection of Prof. Pfannenstiel, Giessen. Carnegie stage 11
Desig. Size Age Body form Primitive streak Primitive segments Chorda Nervous system Eye Ear Nose Hypophysis Mouth Digestive trac, liver and pancreas Branchial pouches, threoid thymus, trachea and lungs Urogenital system Heart and vessels Integument Skeleton Extremities Amnion Allantois Remarks
6 Pfannenstiel III Fig. Vr and Vv. Textfig. 6a to 6w. Gr. L. about 2.6 mm. Embryo bent over the ventral surface and slightly bent spirally. Tail bud, on its ventral side doubtful remains of primitive streak. 13-14 pairs of primitive segments. Chorda emerging from entoderm, Cranially is still entoderm, caudally it is probably primarily independent of the entoderm and in this region it is no longer included in the entoderm. In brain region medullary canal is open to caudal to the region of the optic vesicle, similarly the caudal end. Anlagen of neuromeres already present. Primary optic vesicles. They are close to the ectoderm; in their region the medullary canal is wide open. Anlage of auditory vesicle recognizable as a thickened and at first but little depressed plate of ectoderm. Hypophysis just indicated. Primary pharyngeal membrane still closed. Oral sinus. Wide hepatic bay just cranial to the intestinal umbilicus. No trace yet of hepatic trabeculae. The two first branchial pouches reach the ectoderm, the 3rd is formed. Quite rudimentary "pronephric anlage" in 8th, 9th and 10 pairs of primitive segments. No trace yet of a Wolffian duct. Segmental vesicles in the 11th, 12th and 13th pairs of primitive segments. Heart S-shaped. Posterior mesocardium through a few sections. Aorta paired throughout. Allantoic duct Extirpation of uterus on account of carcinoma.
Remarks - Fixation formalin Muller's fluid. Stain Paracarmine. Sections 10 μ. Recent mitoses. Septum transverse. No ventral connection between pericardial and peritoneal cavities.
Reference: Keibel F. and Elze C. Normal Plates of the Development of the Human Embryo (Homo sapiens). (1908) Vol. 8 in series by Keibel F. Normal plates of the development of vertebrates (Normentafeln zur Entwicklungsgeschichte der Wirbelthiere) Fisher, Jena., Germany.
Cited in: 1912 Human Embryology and

Low A. Description of a human embryo of 13-14 mesodermic somites. (1908) J Anat Physiol. 42(3): 237-51. PMID 17232769 | PMC1289161

1908 Embryo Tables: Klb (stage 10) | Pfannenstiel III (stage 11) | Meyer 300 (stage 12) | Strahl 4mm (stage 13) | Hertwig G31 (stage 14)

Meyer 300

Human Embryo No. 300
Collection of Professor R. Meyer, Berlin. Carnegie stage 12
Desig. Size Age Body form Primitive streak Primitive segments Chorda Nervous system Eye Ear Nose Hypophysis Mouth Digestive trac, liver and pancreas Branchial pouches, threoid thymus, trachea and lungs Urogenital system Heart and vessels Integument Skeleton Extremities Amnion Allantois Remarks
7
N.T. Fig. VI 1, VI r ind VI v.
Gr. L. 2.5 mm. Remains of primitive streak. Cloacal membrane. 23 pairs of primitive segments. Chorda separated from entoderm, Anterior neuropore closed. but its position still recognizable. Medullary canal stili wide open for a stretch caudally. Roof of 4th vent, beginning to thin out. Neuromeres. Trigeminus and acustico-facialis ganglia distinct. Optic vesicles. Mesoderm between ectoderm and optic vesicle, Auditory vesicle almost closed (open through 4 or 5 sections of 5μ). Ductus endolymphaticus not yet visible. Anlage doubtful. Pharyngeal membrane just torn, still abundant remains of it. Intestine still communicates widely with the yolk sack. Liver a thick-walled sack from which the trabeculse are beginning to bud. The 3 anterior branchial pouches reacts the ectoderm; the 4th, though formed, does not. Thyreoidea mediana formed. Pulmotracheal groove. The paired condition of the pulmonary anlage already indicated. Rudimentary "pronephric anlage" Heart S-shaped. Allantoic duct The embryo was obtained by operation.
Remarks

Abundant mitoses in the embryo. Fixation: ? Stain: Borax carmine. Sections: 5 μ. The embryo was modelled by Dr. Peter Thompson (1907) Also see Meyer (1904)

Reference: Keibel F. and Elze C. Normal Plates of the Development of the Human Embryo (Homo sapiens). (1908) Vol. 8 in series by Keibel F. Normal plates of the development of vertebrates (Normentafeln zur Entwicklungsgeschichte der Wirbelthiere) Fisher, Jena., Germany.
Cited in: 1912 Human Embryology and

Thompson P. Description of a human embryo of twenty-three paired somites. (1907) J Anat Physiol, 41(3):159-71. PMID 17232726; Meyer, Rob., Ueber die Beziehung der Umierenkanalchen zum Coelomepithel, etc., Anat. Anz., Vol. 25, 1904

1908 Embryo Tables: Klb (stage 10) | Pfannenstiel III (stage 11) | Meyer 300 (stage 12) | Strahl 4mm (stage 13) | Hertwig G31 (stage 14)

Strahl 4 mm

Human Embryo 4 mm
Collection of Prof. Strahl, Giessen. Carnegie stage 13
Desig. Size Age Body form Primitive streak Primitive segments Chorda Nervous system Eye Ear Nose Hypophysis Mouth Digestive trac, liver and pancreas Branchial pouches, threoid thymus, trachea and lungs Urogenital system Heart and vessels Integument Skeleton Extremities Amnion Allantois Remarks
7
N.T. Text-fig 9a-9s.
Gr. L. 4 mm. Between Figs. 5 and 7 of His" Normenta- fel. (The embryo of Fig. 6 of His' Nor- mentafel is probably pathological.) Vertex bend complete. Nape bend has begun. Chorda separated from entoderm, but in the cranial part, through 19 sections, it has as yet no mesoderm below it. Optic vesicles. Mesoderm between ectoderm and optic vesicle,
Remarks -
Reference: Keibel F. and Elze C. Normal Plates of the Development of the Human Embryo (Homo sapiens). (1908) Vol. 8 in series by Keibel F. Normal plates of the development of vertebrates (Normentafeln zur Entwicklungsgeschichte der Wirbelthiere) Fisher, Jena., Germany.
Cited in: 1912 Human Embryology
1908 Embryo Tables: Klb (stage 10) | Pfannenstiel III (stage 11) | Meyer 300 (stage 12) | Strahl 4mm (stage 13) | Hertwig G31 (stage 14)

Hertwig G31

Human Embryo G31
Collection of Prof. O. Hertwig, Anat-biol. Institute, Berlin.Carnegie stage 14
Desig. Size Age Body form Primitive streak Primitive segments Chorda Nervous system Eye Ear Nose Hypophysis Mouth Digestive trac, liver and pancreas Branchial pouches, threoid thymus, trachea and lungs Urogenital system Heart and vessels Integument Skeleton Extremities Amnion Allantois Remarks
14 Human embryo G 31. N.T. Fig. VIII. Text-fig. 12a-12p. Gr. L. = nape-breech length, 4.9 mm. vertex-breech length 4.7 mm. Greatest diameter of yolk sack 0.58 mm Supposed to be 4 weeks. Between Figs. 6 and 7 of His' Normentafel, nearer Fig. 7. Vanished up to the tail bud. 35 pairs of primitive segments, the most anterior quite rudimentary. The anterior end terminates close to the hypophysis. Very thin chord sheath. Infundibulum. Roof of the 4th ventricle thin. 7 distinct neuromeres in 4th ventricle region. No dorsal columns in spinal cord as yet. Neuroenteric cord. Transition from optic vesicle to otic cup. Lens represented by thickened area of ectoderm,. Scattered mesenchyme cells between lens and otic vesicle. Auditory vesicle just constricted off. The point of constriction still visible at the ectoderm. Ductus endolymphaticus just formed. Distinct convex olfactory areas. Very broad hypophyseal pouch. No remains of pharyngeal membrane. Tuberculum impar. Stomach anlage already slightly twisted. Tail gut. Abundant trabeculae in liver anlage. Gall-bladder formed. Distinct anlage of dorsal pancreas. Early anlage of ventral pancreas. (Jankelowitz 1895 describes 2 ventral pancreas anlagen. Itis doubtful if there are really two; it seemed to us that there was only one.) 4 branchial pouches reach the ectoderm. The solid, two-lobed anlage of the median thyreoid is connected with the floor of the Trachea constricted off. Undivided bronchial buds. Wolffian duct interrupted cranially, free glomeruli right and left ("pronephric remains"). In the mesonephros anlage of glomeruli, caudally segmental vesicles, partly with rudimentary nephrostomes. Wolffian ducts have reached the cloaca, but do not yet open into it. Very early anlagen of the metanephric buds as enlargements of the Wolffian ducts. Nephrogenic cord. Cloaca with bladder and intestinal bay. Primitive germ cells. Atrial and ventricular septum formed, also the right valvular venous. Primary origin of the aa. umbilicales. The aa. omphalo- mesentericse form a ring around the in- testine. Vv. om- phalo-mesenteri- cae connected by an anastomosis dorsal to the intestine. Upper extremity plate-like, lower one like a swelling. Numerous fresh mitoses.
Remarks - Fixation: (according to Herr, 1893) sublimate- acetic acid ; (according to Jankelowitz, 1895 picric-sublimate-acetic acid. Stain: Borax carmine- aurantia. Sections: 10 μ, trans. The embryo was modelled by Dr. Ingalls, compare Irgalls (1907). Literature: Herr, Beitrag zur Entwicklungageschichte des menschl. Auges. Dissert. Berlin, 1893. Jankelowitz, Zur Entwicklung der Bauch- speicheldruse. Dissert. Berlin, 1895. Also, Ein junger menschlichen Embryo, etc. Arch, f mikr. Anat., Vol. 46, 1895.—O.Hertwig, , LehrbuchderEntwicklungsgeschichte. Also, Elemente der Entwick- lungslehre, 3rd ed., 1907. —Ingalls, N. W., Beschreibung eines menschl.

Embryo, etc., Arch. f. mikr. Anat., Vol. 70. 1907. Bursa omentalis, with foramen of Winslow, is formed.

Reference: Keibel F. and Elze C. Normal Plates of the Development of the Human Embryo (Homo sapiens). (1908) Vol. 8 in series by Keibel F. Normal plates of the development of vertebrates (Normentafeln zur Entwicklungsgeschichte der Wirbelthiere) Fisher, Jena., Germany.
Cited in: 1912 Human Embryology and

Ingalls NW. Description of a human embryo of 4.9 mm (Beschreibung eines menschlichen Embryos von 4.9 mm). (1907) Arch. f. mik. Anat., 70: 506-576.

1908 Embryo Tables: Klb (stage 10) | Pfannenstiel III (stage 11) | Meyer 300 (stage 12) | Strahl 4mm (stage 13) | Hertwig G31 (stage 14)

Strahl 6.75 mm

Template:Human embryo Strahl 6.75mm table

Hochstetter Chr 1

Template:Human embryo Hochstetter Chr 1 table

Meyer 250

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Keibel 1446

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Meyer 321

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These tables will suffice to give a general idea of the time relations of the development of the various organs. The practical value of the tables is also plainly evident. The investigator who wishes to study some particular organ may quickly determine from the Normentafel what stages he requires for his investigation. And I would place especial importance on the fact that the Normentafel make it possible to estimate the proper value of isolated observations and to assign them readily to their proper jolace.


Furthermore, as may be readily seen when one examines the entire series of Normentafel, in which numerous closely succeeding forms are shown, the tables furnish a measure for the determination of individual variations in the embryonic development of man. Such variations undoubtedly occur, but, as in other amniotes that have been studied, they are not very important. However, the tables also furnish information on more general questions. Even in the preparation of the first Normentafel, that of the pig (published 1897), Keibel (1895) was able to demonstrate that the modification of the time relations, indeed, one may say that overlapping in the development of the various organs is so extensive, that a satisfactory division of the entire development into stages corresponding to a praepiscme form, a fish, a terrestrial animal and an amniote, such as Oppel (1891), for example, has assumed and which must be assumed if the biogenetic law is strictly applicable, such a division is impossible. This was then found to be true for other amniotes that were studied in this way, and especially for man. We see a regular succession of the individual ontogenetic stages only when the preceding stage is the necessary condition for the succeeding one. " The unicellular organism," says 0. Hertwig (1906 2 ), " from its nature, can be transformed into a multicellular one only by cell-division. Consequently in all animals the ontogenesis must begin with a division of the ovum. An organism with definitely arranged celllayers and cell-groups can only be formed from a mass of cells when the cells, during their division, begin to arrange themselves into firm unions. ' : In this sense one may speak, with Oppel (1891), of indispensable stages of development. Otherwise, however, we find in the time succession of the organs no indication whatever of the succession in which they were acquired phylogenetically. It is sufficient to point out that the embryonic organs, the amnion, chorion, and allantoic, appear before other organ anlagen are recognizable. Keibel (1895) has shown that this precocious appearance of various organ anlagen is probably associated with the necessity for their earlier or later activity. That definite traces of the path taken by phylogeny are retained by heredity is, nevertheless, not to be denied, but for the time sucession heredity lias proved to be a force of little moment. That an actual recapitulation of the phylogeny in the ontogeny, and, therefore, the validity of the so-called biogenetic law, is impossible for purely logical reasons Keibel (1893) has already pointed out, when he states that "already in the first member of a developmental series the last one " is determined, and that, consequently, the first member of a series must appear modified, when compared with the first member of another series " which may have had the same phylogenetic beginning, but was completed at a lower stage of development." 3 0. Hertwig (1906?) has termed " this relative dependence between the ovum condition on the one hand and the course and final result of the ontogenesis on the other hand, the ontogenetic law of causation and the parallelism of anlage and anlage product. ' : It follows from this law that all intermediate members will show modifications. These questions, however, need not be further discussed in this place, but I would refer the reader to Keibel (1903) and O. Hertwig (1906 x 2 ). I would call the attention of those who may be interested in comparing the interdependence of the different developmental processes of the human embryo with those of ape embryos to the 9th part of Emil Selenka's Menschenaffen, in which F. Keibel (1906) discusses the external form and degree of development of the organs in ape embryos and has given tables showing the development of the apes in the same way that Keibel and Elze (1908) have shown that of man in the Normentafel zur Entwicklungsgeschichte des Menschen.



Literature

Berry RJA. and Lack LAH. The vermiform appendix of man and the structural changes therein coincident with age. (1906) J. Anat. and Phys. 40: 247-256. with 11 figs.

Born. G. : Leber Verwachsungsversuche mit Amphibienlarven. Arch, fur Entw. Mech., vol. 4, 1897. (Also published in book form.)

Fischel. A. : Leber Variability und Wachstum des embryonalen Korpers. Morph. Jahrb., vol. 24, 1896.

Hammar, J. A. : Der gegenwartige Stand der Morphologie und Physiologie der Thymusdruse. Verhandl. d. 16 ten Internat. Med. Kongr. (Ofen-Pesth), 1st section, p. 4-29, 1909. (In this review references are made to results, as yet unpublished, by Hammar's pupils, Hellmann, Kaellmark and Lindberg. )

Hertwig, O.: Aeltere und neuere Entwicklungstheorien. Address delivered at the anniversary of the foundation of the Militar-arztliehen Bildungsanstalten, Aug. 2. 1S92. Berlin. 1892.

Hertwig. O. : Allgemeine Biologie. 2nd edition of the text-book Die Zelle und die Gewebe. Jena, 1906 1 . Compare also Keibel (1898).


Hertwig, 0.: Ueber die Stellung der vergleichenden Entwicklungslehre zur vergleichenden Anatomie, zur Systematik und Descetidenz-theorie (das biogenetische Grundgesetz, Palingenese und Cenogenese). In 0. Hertwig's Handbuch, Jena, 1906 3 .

His, W. : Unsere Korperf orm und das pbysiologische Problem ibrer Entstebung. Leipzig, 1874.

His, W. : Die Entwicklung der menscblicben und tieriscben Physiognomien. Areb. f. Anat. u. Pbys. Anat. Abth., 1392.

His, W. : Das Prinzip der organbildenden Keimbezirke und die Verwandtschaften der Gewebe. Ibid., 1901.

Jonson, A. : Studien iiber die Tbymusinvolution. Die akzidentelle Involution bei Hunger. Arcb. f. mikr. Anat., vol. 73, p. 390-443, witb 12 plates and 11 text-figs., 1909.

Keibel, F. : Studien zur Entwieklungsgeschiehte des Schweines (Sus serofa domestieus) II. Scbwalbe's Morph. Arb., vol. v, 1895.

Keibel, F. : Das biogenetisebe Grundgesetz und die Cenogenese. Merkel and Bonnet's Ergbn. der Anat. und Entwicklungsgesch., vol. 7, 1898.

Keibel, F. : Ueber den Entwieklungsgrad der Organe in den verscbiedenen Stadien der embryonalen Entwicklung der Wirbeltiere. In Hertwig's Handbuch, 1906 (appeared in 1903).

Keibel, F. : Die aussere Korperf orm und der Entwieklungsgrad der Organe bei Affenembryonen. In Emil Selenka's Menschenaffen, continued by Hubrecbt, Strabl and Keibel. Wiesbaden, 1906.

Keibel F. and Elze C. Normal Plates of the Development of the Human Embryo (Homo sapiens). (1908) Vol. 8 in series by Keibel F. Normal plates of the development of vertebrates (Normentafeln zur Entwicklungsgeschichte der Wirbelthiere) Fisher, Jena., Germany. witb 6 plates and 44 figs, in text, Jena, 1908.

Lewis, W. H. : Experimental Studies on tbe Development of tbe Eye in Ampbibia. 1. On tbe Origin of tbe Lens. Amer. Journ. of Anat., vol. 3, p. 505-536, 1904.

Lewis, W. H. : Transplantation of tbe Lips of the Blastopore in Rana palustris. Amer. Journ. of Anat., vol. 7, 1907.

Mehnert, E. : Die individuelle Variation des Wirbeltierembryo. Scbwalbe's Morpb. Arb., vol. 5, 1895.

Oppel, A. : Vergleiebung des Entwicklungsgrades der Organe, Jena, 1891.

Paton, N. D. : Tbe Relationship of the Thymus to the Sexual Organ. II. The Influence of the Removal of the Thymus on the Growth of the Sexual Organs. Journ. of Pbys., Cambridge, vol. 32, 1904.

Paton, N. D., and Goodall, A.: Contribution to tbe Physiology of the Thymus. Journ. of Phys., Cambridge, vol. 31, 1904.

Rous, W. : Beitrage zur Entwicklungsmechanik des Embryo. VII. Ueber Mosaik arbeit und neuere Entwicklungshypothesen. Anat. Hefte, vol. 2, Parts 6-7, p. 279-330, 1893. (Alsp Gesammelte Abhandlungen iiber Entwicklungsmech., vol. 2, p. 818, Leipzig, 1895.)

Sellheim, H. : Kastration und Knochenwacbstum, Hegar's Beitrage zur Geburtsh. u. Gynakol, vol. 2, 1899.

Soederlund, G., and Backhan, A. : Studien iiber die Tbymusinvolution. Die Altersveranderungen der Thymusdriisen beim Kaninchen. Arch. f. mikr. Anat., vol. 73, p. 699-725, with 1 plate and 6 text-figs., 1909.

Spemann, H. : Ueber Korrelationen in der Entwicklung des Auges. Verhandl. Anat. Ges. (Bonn), 1901. Suppl. to Anat. Anz. vol. 19, 1901.

Spemann, H. : Ueber Linsenbildung bei def ekter Augenblase. Anat. Anz., vol. 23, 1903.

Syk, I. : Ueber Altersveranderungen in der Anzahl der Hassal'schen Korper nebst einem Beitrag zurn Studium der Mengenverhaltnisse der Mitosen in der Kaninchentbymus. Anat. Anz., vol. 34, p. 560-567, 1909.


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