Paper - Report upon the collection of human embryos at the Johns Hopkins University (1911)
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Mall FP. Report upon the collection of human embryos at the Johns Hopkins University. (1911) Anat. Rec. 5(7): 343–357.
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Report upon the Collection of Human Embryos at the Johns Hopkins University
Professor of Anatomy, Johns Hopkins University, Baltimore
About twenty years ago I wrote a note for the American Naturalist asking for human embryological specimens. At that time my collection consisted of two embryos, one of which was in an excellent state of preservation. This I had carried as an offering to my teacher, Professor His, who would not accept it but returned it to me with several series from his own collection, including his well-known embryo No. XLIV (Bff). “These,” he said, “will make a nucleus for your own collection. It will give me immense pleasure to see publications coming from new sources.”
The misplaced note in The Naturalist brought no return nor was the reward from Minot’s legacy to write numerous articles in the Reference Handbook on various phases of human embryology any better. But fate was with me for I concentrated my energies upon embryo No. 2 which not only taught me the method of reconstruction in wax but also served as a basis for my career in anatomy. I am proud of No. 2. At that time its effigy was the most elaborate reconstruction of an embryo in existence; it was the first,reconstruction of an embryo made in America and I always think in human anatomy in relation to No. 2.
The numerous wax plates required in making the reconstruction of No. 2 and the form of the various organs as revealed in piling their pieces attracted the attention of various medical friends, who in the course of time supplied me with new embryos for study. They naturally brought their questions with the specimens. I was constantly asked: “How old is the embryo?” “What caused the abortion?” and “Could it have been prevented?” An attempt to answer these questions as well as other information served to satisfy the scientifically inclined, and soon I found a large number of physicians willing to cooperate with me. This number was enlarged through the publication of several papers outlining my field of work and numerous letters for specimens in many medical journals. Thus it came about that collaborators were secured and many pleasant hours have been passed in conversation and discussion with them. This interest is by no means conlined to physicians with whom I have personal acquaintance, but includes many whom I have never seen. The correspondence with these has often been extensive and long continued, their letters forming valuable notes which belong to the collection.
The making of a show collection will not satisfy the curiosity of thinking physicians very long, but they require rightly that their labors be rewarded by substantial contributions to embryological science. The anatomy of embryos will occupy much of our attention for some time to come. In case embryos studied are young the various structures of the whole specimen can be considered to advantage, but our powers are limited and only an individual system, organ, tissue or region can be followed in older embryos from stage to stage. The various embryos must be classified and standardized, the membranes must be studied anew, and abnormal specimens must be recognized and considered in relation to the cause which produced the deformity. So a single worker makes headway slowly and even a large number of experts with the aid of students and technicians cannot do justice to representative collections of specimens. Some aid has been secured by enlisting investigators in other institutions in studying this collection, and it is to be hoped that this may continue to a much greater extent in the future. As matters now are it will take a very long time before human anatomy rests entirely upon an embryological basis, and we should organize our forces in order to reduce the time as much as possible. His has proposed that an Embryological Institute be organized with this end in view, in which human embryologists cooperate in their work much as is done in the Brain Commission. The example in this direction set by the astronomers is still better and should be followed. My specimens would be placed at the disposal of a joint commission and would help to make a good beginning. As long as the methods of investigation in this subject remain as they are this collection will be of value; later it will still be of historical interest.
But I must come back from dreamland and state what can be done single-handed. At first we were fortunate if specimens came fairly well preserved in alcohol; too often they were injured or badly preserved. The introduction of formalin as a preservative gave us much better specimens, and now perfectly fresh and sometimes living embryos are brought to us. These are preserved by the more refined methods and often the blood vessels are injected. The collection now includes a representative series of embryos and foetuses, from 15 mm. upward, with the blood vessels well injected.
Although the collection is well illustrated with numerous unpublished drawings and photographs, an effort is being made to record more elaborately unusually important stages and perfect specimens with many careful drawings and studies before they are cut into serial sections. The histories, although extensive, are becoming better and better. These bear upon the cause of the abortion; the nature of the injury in case it is mechanical, and the embryo has been unwittingly experimented upon; and the disease of the uterus in case the specimen is pathological. Reactions between leucocytes and syncytium are found t o be pronounced and moles and monsters seem to bear relation to unskilled obstetricians as well as to the social evil. Studies in this direction must be made on a large scale to establish certain points of great value to medicine and to the race. A research institution or the cooperation of a group of obstetrical and gynecological clinics will be required to undertake the work before such questions can be answered. The routine work to be done is much too great for a single individual to undertake.
In the 533 specimens recorded under 500 numbers there are 50 normal embryos and 105 pathological under 8 mm in length, that is, under six weeks old. Between 9 and 25 mm. there are 133 normal embryos and 66 pathological. In the whole collection there are 335 normal specimens and 198 pathological. Among the abortions of the first six weeks of pregnancy nearly 68 per cent of the specimens are pathological, from the sixth to the end of the eighth, 34 per cent, and during the remaining seven months but 18 per cent. From these figures as well as from my experience in the study of tubal pregnancy the conclusion may be drawn that a much larger percentage of fertilized ova are not implanted, escape observation and are lost. It would be interesting in this connection to compare collections of ova obtained from communities free from venereal disease with those in which syphilis and gonorrhoea are quite general. In the latter case abortions should be more common and the percentage of monsters should be greater.
Since it is established that monsters arise from normal eggs in an unhealthy environment the hope that we may be able to diminish their number is not visionary. Moles, or pathological ova, which are of far more common occurrence might be eliminated altogether.
It is necessary to determine with much greater precision than has been done the norm of both the embryo and its membranes for each stage of development. Much of the material for this study is at hand but before this is undertaken it will be necessary to establish the various stages. We have been in the habit of classifying embryos according to age, which is by no means a satisfactory method. An index dependent upon the length of the embryo is used by embryologists but for well known reasons this also leads to confusion. It will be necessary to establish stages based upon the degree of development. For example, the embryos which are from 2 to 3 mm long and are believed to be from two to four weeks old, might arbitrarily be called Stage H, leaving the first letters of the alphabet to represent earlier stages. At this time the myotomes are appearing and an embryo with two myotomes could be called Stage HP, one with three myotomes Stage Ha, and so on. Later we could determine the degree of development of each organ and tissue and then refine the subdivision of stages still more. It may turn out that the organs are developed unequally in embryos of the same stage and this would be an observation of great importance.
When embryos have been standardized in this way we shall find that specimens of the same stage vary in length even when measured accurately in the fresh state. We may also find when the ages have been determined with greater precision, that the rapidity of development is not constant.
Since the unfertilized ovum has been observed in the uterine tube by Hyrtl and since an ovum 2mm. in diameter has been found by Teacher there is every reason to hope that all stages of development will soon be found. Operations which expose the pelvic organs are now so common that if the gynecologist will only show the proper interest and look with especial care when a fresh corpus luteum is seen, the desired specimens will not remain hidden much longer.
There is an abundance of material at hand to study the growth of foetuses. The establishment of stages will necessitate the extension of the physical anthropological measurements downward through the foetus to the embryo. We should have tables which will tell us, for example, the length of each bone, the length of the brain, the degree of the development of its cortex, etc., for each stage. It is almost needless to add that cooperative studies of this kind will necessitate the introduction of a uniform terminology to articulate with the BNA.
In recent years a large number of foetuses as well as embryos have been injected in order to study the blood vessels with much greater care. A number of the specimens came into our hands still alive, which made it possible to obtain complete vascular injection in embryos less than 20 mm long. In the catalogue they are marked [i]. It is now within our reach to give a satisfactory account of the development of the vascular tree of each organ, and in the course of time the exact extent of the lymphatic system in the human embryo will also be revealed by the injection method.
Many specimens have been clarified in caustic potash and glycerine to show the extent of ossification; these have been marked [k] in the catalogue. About 100 specimens less than 100 mm. long have been prepared in this way, and much more material is at hand for similar treatment. This work can be extended to show the extent of chondrification in embrios from which the organs have been removed for study. This brings me to the studies in organogeny which needs an abundance of whole organs to determine their form, as everything cannot be reconstructed. The tissues are well represented in serial sections of earlier stages and for older foetuses individual tissues from various stages of perfectly fresh specimens have been fixed and preserved by the best methods. A great field opens to us for the study of histogenesis which binds embryology to histology to make the foundation for scientific anatomy. Here the method of attack is also experimental and the horizon extends to include physiology, teratology and pathology.
Nearly 300 specimens were sent to me by physicians residing in Maryland. About 40 came from New York, 30 each from Ohio and the District of Columbia, and 25 from Massachusetts. Between 5 and 10 came from each of the following states: Iowa, Illinois, Wisconsin, California, New Jersey, Minnesota, Maine and Michigan, while smaller numbers were received from fifteen other states and countries.
One hundred publications are based, most of them entirely, upon this collection. The list of papers includes the chapters in the Manual of Human Embryology; this makes the data complete up to 1911. In a way this report may be considered a turning point in the history of the collection, as the Manual contains an inventory of the literature upon human embryology and is based upon this as well as upon other collections. The Manual may be viewed as a by-product, for it has been produced by investigators whose chief occupation has been teaching, and the volumes have been published generously by Hirzel of Leipzig and Lippincott of Philadelphia. It is not dependent upon a single research grant. If “to him that hath shall he given” applies to this effort we have every reason to hope for a far more elaborate display in the second edition. The “Institute of Human Embryology” must do this work; it should enlist the energies of all competent investigators in this subject, the necessary material should be placed at their disposal, and research funds should be available to aid them in their work. I firmly believe that such a cooperative organization would produce more valuable results than is possible if our efforts remain divided. Gross human anatomy as a science is bankrupt. It is made solvent through embryology, which alone illuminates it.
List of Embryos
NOTE. The abbreviations in parentheses are as follows:
- The first figure gives the length of the embryo from crown to rump in millimeters.
- The letter following indicates the direction of the section T means a transverse, S means a sagittal, and C means a coronal.
- The figure following the sign glves the thickness of the sectioning.
- The words good, fair or poor indicate the suallty of the series. In case the specimen Is pathological, abnormal or is treated In some special way this is marked in brackets: thus, [p]
Following the abbreviatlon af of each specimen is the name and address of the physlclan from whom it was obtalned. If no address is given Baltimore is understood. The figure after the name refer to papers with corresponding numbers in the bibliography based upon this specimen. In case a number is inltall ca it Indlcates that this speclmen is illustrated In that paper.
♂ = male; ♀ - female; a - abnormal; i = injected; p = pathological; k = cleared in KOH; t = tubal pregnancy
Following the abbreviations for each specimen is the name and address of the physician from whom it was obtained. If no address is given Baltimore is understood. The figures after the name refer to papers with corresponding numbers in the bibliography based upon this specimen. In case a number is in italics it indicates that this specimen is illustrated in that paper.
|List of Embryos (to be formatted)|
1 (4.5. T, 10, poor.) Dr. Gavin, 13, 21. 27, 40, 41, 63, 75, 83, 91.
2 (7, T 15, good.) Dr. C. 0. Miller,1,2, 4, 5, 6,7, 3, 9,10, 11.1£,13,14,15, 19, 21, 24, 27, 28, 29, 30, 32, 34, 36, 39. 40, 41, 45, 46, 47, 48, 49, 50, 52, 54, 66, 58. 62, 63, 68, 71, 73, 75, 76. 83……
3 (2.2, T, 10, poor.) Prof. His. Leipzig. Germany, 10, 30. 40. 63, 74, 9|.
4 (7, T, 10. fair.) Dr. Williams. 10, 30. 40, 41, 63. 91.
5 (18.5, T, 20,fair.) Dr. Kittredge, Nashua. N. 11., 10,13. 21, 27. 30, 40, 41, 63, 75, 83. 91.
6 (24, T, 20, good.female) Dr. C. O. Miller, 4, 10,1l,13,15,19, 21, £2, 27. 30, 40. 57, 63, 68. 75.82.83, 91,
7 (19 5. T, 40. poor.) Dr. Booker, 30, 40, 57, 63, 91.
8 (17, T, 0, poor.) Dr. Ritter, Brooklyn, N. Y., 10. 30. 40
9 (17.5, .20. fair. female) Dr.Eyelesh mer, Chicago,10,11 68,83,131
10 (24, T, 0, fair.) Dr. W. S. Miller, adison, Wis..1D,11,15, 21, 30, 40, 66. 57. 60, 63, 68, 75, 83, 91.
11. (0.8, S, 10, fair [p].) Dr. Kittredge, Nashua. N. H.. 3. 4, I0. 13. 91. 27, 30. 33. 40. 48, 63, .
12. (2.2. T, 10, good.) Dr. Ellis, Elkton, M.d., 4,10,11,12, 13, 15, 19, £1, 24, 27. 28, 50, 3!, 36. 39.40, (1,
13. (1.4, S, 10, fair [p].) Prof. His, Leipzig, Germany.10, 1. 30. 55. 37, 40, 63. 75. 91.
14. (1.5. 10, fair [pj.) Dr. Friedenwald. 10. £1, 30. 83, 40, 63. 76.
15. (6.) Dr. Hoge, Richmond. Va.. 63. 83.
16. (7.) Dr. Sherwood.10, 13, 21, 27, 40. 63. 75, 83.
17. (18. 50 & 100, poor.) Dr. Cottell Louisville, Ky.,10, 13, 21, 27,
18. (7, T, 20, fair.) Dr. Douglass, Nashville. Tenn.. 10, 13, 19. 21.
19. (5 T,20, poor.) Dr.Williams.10, 13,1.9,21,27.30,38, 40.41.63. 75,
20 (50, fair, [p].) Dr. Williams, 10, £1, 37, 40. 63. 76.
21. (5.5, T. 20, fair. [p].) Dr. Cullen, London. Canada. 10, 21, 33. 8
February, 20. 1911
February, 20. 1911
Tltles of papers based entlrely or In part upon this collectlon of human embryos.
Mall FP. Development of the lesser peritoneal cavity in birds and mammals. (1891) J of Morphology 5: 165-179.
Mall FP. A human embryo of the second week. (1893) Anat. Anz. 8: 630-633.
Mall FP. Early human embryos and the mode of their preservation. (1893) Johns Hopkins Hospital Bulletin 4: 115-121.
Mall FP. The heart. (1893) Ref. Handb. Med. Sci. Supplement, 9: 391-395.
1897a Development of the human coelom. Jour. Morph., vol. 12.
1897b Ueber die Entwlckelung des menschllchen Darmes und selner Lage belm Erwachaenen. Arch. f. Anat. u. Physlol.. Supplementband.
1898a Development of the ventral abdominal walls In man. Jour. Morph., MI. 14.
1898b The value of embryological speclmens. Maryland Medical Jour.
1898 Development of the internal mammary and deep eplgastric arteries In man. Johns Hopkina Hospital Bulletln.
1898d Development of the human lntestlnr and Its posltlon In the adult. Ibld.
1898 The development of the bile capillaries as revealed by Golgl’s method. Ibld.
Mall FP. Supplementary note on the development of the human intestine. (1899) Anat. Anz. 16: 492-495.
BARKEIR L. F. 1889 The nervous system. D. Appleton and Co., New York.
CLARK J. G. 1900 The orlgln. development and degeneratlon of the blood veasels of the human 18 19 20 February, 20. 1911.
MACCALLUM J.,B. 1898 On the histogenesis of the striated muscle fibre, and the growth of the sartorlua muecle. Ibld. ovary. Welch Festsohrlft. Johns Hopklns Hospital Reports, vol. 9. 21’
Mall FP. A contribution to the study of the pathology of early human embryos, (1900) Johns Hopkins Hosp. Rep., 9: 1-68.
BERRY J. M. 1900 On the development of the vllll of the human intestine. Anat. Am Bd. 17. 23
MACALLCUM. .B. 1900 On the muscular architecture and growth of the ventrlcles of the heart. Welch Festcshrlft. Johns Hopklns Hospital Reports, vol. 9. 24
LEWIS, W.H. 1901 Observntlons on the pectoralis major muscle In man. Ibld. 26
HARRISON R.G. 1901 On the occurrence of tails in man, with a description of a case reported by Dr. Watson. Ibld.
MALL, F. P. 1901 Age of human embryos. Ref. Handb. Med. Sci., 2nd ed., vol. 3. 28
BARDEEN C,.R. and LEWIS,W. H. Development of the llmbs. body-wall and back In man. Amer. Jour. Anat vol. 1. 29
MALL, F. P. 1901a Compiratlve development of the coelom. Ref. Handb. Med. Sci. vol. 3. 30
- 1901b Development of the human coelom. Ibid. 31
LEWIS W. H. 1902 The development of the arm in man. Amer. Jour. Anat.. vol. 1. 33
MALL F, .P. 1901 Pathologlcal human embryos. Ref. Handb. Med. Sci., 2nd. ed.. vol. 3.
MACCALLUM M.B. 1902 Notes on the Wolffian body of higher mammals. Amer.Jour.Anat.. vol. 1.
MALL F,.P. 1902 Development of the heart. Ref. Handb. Med. Scl., 2nd. ed., vol.4. 36
Sudler MT. The development of the nose and of the pharynx and its derivatives in man. (1902) Amer. J Anat. 1:391–416.
MALL, F. P. 1903 Second contribution to the study of the pathology of early human embryos. Contrlbution to Medical Research. Dedicated to Vlctor C. Vaughan, Ann Arbor.
WILLIAMS J. W. 1903 Obstetrics. D. Appelton and Co., New York. LEWISW'.H. 1903 Human embryology.. New International Encyclopedia New York.
MALL, F. P. 1903a Note on the collection of human embryos in the Anatomical Laboratory of Johns Hopklns Unlversity. Johns Hopkins Bulletin.
- 1903b On the transitory or artlficlal fissures of the human cerebrum. Amer. Jour. Anat. vol. 2.
PEARCER. M. 1903 On the development of the lslands of Langerhans in the human embryo. kmer. Jour. Anat., vol. 2.
GAGES,.P. 1903 Serial order of the segments In the fore-brain of three and four-weeks human embryos. Science, N. S., vol. 47.
- 1904 The mesonephros of a three-weeks human embryo. Amer. Jour. Anat., vol. 3. p. 6.
MALL, F.P. I904a Development of the thymus gland. Ref. Handb. Med. Sci., 2nd ed.. vol. 6.
- 1904b Development of the thyrold. Ibid.
LEWIS W. H.1904 Development of the foetus. Ibld.
MYER, A. W. 1904 On the structure of the human umbillcal vesicle. Amer. Jour. Anat., vol. 3.
STREETER, G. L. 1904 Peripheral development ofthe nerves Inthe occipital region of the human embryo. Amer. Jour. Anat., vol. 4.
MALL, F.P. 1904 On the development of tht blood-vesselsof the brain in the human embryo. Ibid.
POHLMAN. G. . 1904 Concerning the embryology of kidney anomalies. Amer. Med., vol. 7, P.987.
- 1905 The abnormalities In the form of the kidney and ureter dependent on the development of the renal bud. Johns Hopklns Bulletin.
- 1904 Has a persistance of the MUllerian duct any relation to the condition of cryptorchidism. Amer. Med. vol.8,q.1003.
- 1903A_ nztepfi the developmental relations of the kidney and ureter-human embryos, J. H. H. Bull
GAGES, .P. 1904 The total folds of the forebrain their orlgln and development to the third week In the human embryo. Amer. Jour. Anat. vol. 4 Ir.
BARDEEN C,. R. 1905a Studies of the development of this human skeleton. Amer. Jour. Anat.. vol. 4.
- 1904 Numerical vertebral varlatlons in the human adult and embryo. Anatom. Aneelger, Bd. 25.
- 1905b Development of the thoracic vertebrae 1n man. Amer. Jour. Anat., vol. 4.
GAGES, P. 1905 A three weeks human embryo with especial reference to the brain and the nephrlc system. Amer. Jour. Anat., vol. 4.
BRODELM, AX 1905 Anatomy of the sppendlx, Kelly's Vermlform Appendix, Saunders, Phlladelphis.
JACKEOCN.,M. 1905 Onthe topography of the pancreasInthe humen foetus. Anat. Anzelger, Bd. 27,p. 488.
STEEETER G. L. 1905-06 Concerning the development of the acowtle ganglion in the human embryo. Verhandl.d.Anatom. Gesell. and Amer. Jour. Anat.,vol.5,p.1.
MALL, F.P. 1904 Catalogue of the collectlor oi human embryos in the Anatomlcal Laboratory of the Johns Hopklns Unlverslty. Baltlmore Printed privately).
HILL, EBENC. 1906b On the Schdtze clearlng method as used In the Anatomical Laboratory of the Johns Hopklns University Johns Hopklns Hoepltal Bulletln. 1 W b On the embryonlc deveiopment of a case of fused kldneys. Johns Hopkins Hospits1 Bulletln.
STREETER G.L. 1906 Development of the membranous labyrinth and acoustic ganglion in the human embryo. Amer. Jour. Anat vol. 5 p. vil.
GAGE 9, .P. 1906 Total folds of the brain tube in'the embryo and their relatlon to definite structures. Amer. Jour. Anat vol. 5 p. Ix.
MALL F, . l906a A study of tce struciural unlt of the llvei. Amer. Jour. Anat., vol. 6. &b On Oeslficatlon centres In human embryos less than one hundred days old. Amer. Jour. Anat. vol. 5.
GAOE. S.P. 1906 The notochord of the head in human embryos of the third to the twelfth week, and comparisons with other vertebrates: Science, September 7.
STREETER G.L. 1907 Onthe development of the membranous labynlnth and the acoustic and facial nerves In the human embryo. Amer. Jour. Anat., vol. 6.
MALL, F.P. 1907 The collection of human embryos at the Johns Hopkins Unlverslty. Anat. Rec., no. 2, Amer. Jour. Anat.. vol. 6.
BARDEENC. R. 1907 Development and variation of the nerves and the musculature of the inferl6rextremlty and of the nefghborln regions of the trunk ln man. Amer.Jour.Anat.,vol.6.
STREETER G,. L. 1907 Development of % e inter forebraln commlesures In the human embryo. Anat. Rec.. no. 3,
MALL,F. P. 1908 A study of the causes underlylng the orlgfn of human monsters. (Third contribution to the study of the pathology of human embryos.) Jour.Morph. vol.19.
- Also monograph published by The Wlstsr Institute of Anatomy, Philadelphia.
BARDEEN C, R. 1908 Vertebral determinatlon in young human embryos. Anat. Rec., vol. 2.
STREETER G,.L. 1908 The nuclei of origin of the cranial nerves In a 10mm. human embryo. Anat. Rec.. vol. 2.
BARDEEN C. R. 1908 Early development of the cervical vertebrae and the base of the occipital bone in man. Amer. Jour. Anat., vol. 8.
RETZER ROBERT 1908 Some results of recent fnvestlgatlons on the mammalian heart. Anat. Rec. vol. 2.
EYANSH,. M. 1908 Onan Instance of two suhclavlan arteries to the early arm bud in man. Anat. Rec., vol. 2.
ESSICICC,.R. 1909 On the embryology of the corpus pontobulbare and its relation to the development of the pons. Anat. Rec., vol. 3.
SABIN F,. R. 1909 The Lymphatic system In human embryos with a conslderatlon of the morphology of the system as a whole. Amer. Jour. Anat.. vol. 9.
MALL F,.P. 1910a Die Alterbestlmmungvonmenschllchen Embryononen und Feten. Keibel Mall Handbuchder Ectw1cklungs-ge8chichtedes Menschen, Leipelg,Bd. 1; English Edltlon. Philadelphia vol. 1.
- 1910b Die Pathologic des menschlichs Eies. Ibid.
BARDEEN C,. R. 1910 Die Entwicklung des skeletts und des Blndegewebes. Ibid.
LEWIS W H. 1910 Die Entwickelung des Muskelsystems. Ibid.
MALL. F. P. 1910 CMon und Zwerehfell. Ibld.
DANDY, W.E.1910 A human embryo with seven palrs of somites measuring about 2mm. in length. Amer. Jour. Anat., vol. 10.
MINOTC. S. 1910 Laboratory textbook of embryology Phlladelphia.
STREETER., L. 1911. Die Histogenese des NervengeGebes. Keibel-Mall Handbuch, Bd. 2.
MALL F, EANHLPIN. 1910 A list of human embryos which have been cut Into serlal sections. Anat. Rec., VOl. 4.
WILSON L.B. and WILLIS B.C. 1911 A comparatlve study of the histology of the so called hyperhephromata and the embryology of the nephridlal and adrenal tissues. Jour. Medical Research, vol. 24.
LISSER, H. 1911 Studies on the development ofthe human larynx. Amer. Jour. Anat.. vol. 12.
LEWIS F. T. 1911 Die Entwicklune-des Darmes und der Atmunw-ore-ane. Kelbel-Mall Hand-buch, Bd. 2.
MINOT, C.S. 1911 Die Entwickelung des Blutes. Ihid.
EVANS,H. M. 1911 Die Entwicklung des Blutgefiiss-systems. Ibid.
SABIN, F.R. 1911 Die Entwicklung des LymphgefBss-s stems. Ibid.
Pohlman A.G. 1911 The development of the cloaca In human embryos. Amer. Jour. Anat., vol. 12.
WHITEHEAD. R.H. and WannELL .J. A. 1911 The early stages of the development of the mammalian sternum. Amer. Jour. Anat., vol. 12.
MALL, F.P. 1911 Report upon thee collection of human embryos at the Johns Hopkins University. Anat. Rec., vol. 5.
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- Carnegie Stages: 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 | 16 | 17 | 18 | 19 | 20 | 21 | 22 | 23 | About Stages | Timeline
Carnegie Stage Table
Weeks shown in the table below are embryonic post ovulation age, for clinical Gestational Age (GA) measured from last menstrual period, add 2 weeks.
(not to scale)
|fertilized oocyte, zygote, pronuclei|
|morula cell division with reduction in cytoplasmic volume, blastocyst formation of inner and outer cell mass|
|loss of zona pellucida, free blastocyst|
|extraembryonic mesoderm, primitive streak, gastrulation|
|gastrulation, notochordal process|
|primitive pit, notochordal canal|
|Somitogenesis Somite Number 1 - 3 neural folds, cardiac primordium, head fold|
|Somite Number 4 - 12 neural fold fuses|
|Somite Number 13 - 20 rostral neuropore closes|
|Somite Number 21 - 29 caudal neuropore closes|
|Somite Number 30 leg buds, lens placode, pharyngeal arches|
|lens pit, optic cup|
|lens vesicle, nasal pit, hand plate|
|nasal pits moved ventrally, auricular hillocks, foot plate|
|straightening of trunk|
|upper limbs longer and bent at elbow|
|hands and feet turned inward|
|eyelids, external ears|
|rounded head, body and limbs|
- Glossary: A | B | C | D | E | F | G | H | I | J | K | L | M | N | O | P | Q | R | S | T | U | V | W | X | Y | Z | Numbers | Symbols | Term Link
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