Paper - Maturation of the ovum in swine (1917)

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Corner GW. Maturation of the ovum in swine . (1917) Anat. Rec. 13(2): 109-112.

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This 1917 paper by George Corner describes pig implantation.

Also by this author: Corner GW. and Amsbaugh AE. Oestrus and ovulation in swine. (1917) Anat. Rec. 12: 287-.

Modern Notes: estrous cycle | pig
See also: Corner GW. (1921). THE OVARIAN CYCLE OF SWINE. Science , 53, 420-1. PMID: 17773390 DOI.

George Corner Reference List 
Embryology References

Corner GW. Maturation of the ovum in swine . (1917) Anat. Rec. 13(2): 109-112.

Corner GW. Cyclic changes in the ovaries and uterus of swine, and their relations to the mechanism of implantation. (1921) Contrib. Embryol., Carnegie Inst. Wash. Publ. 394, :117-146.

Corner GW. Ovulation and menstruation in Macacus rhesus. (1923) Contributions to Embryology, vol. 15, Carnegie Inst. Washington Pub. no. 332, 75-101.

Corner GW. A well-preserved human embryo of 10 somites. (1929) Carnegie Instn. Wash. Publ. 394, Contrib. Embryol., Carnegie Inst. Wash. 20: 81-102.

Corner GW. The hormone of the corpus luteum. (1937) Trans Edinb Obstet Soc. 57: 61-80. PMID 29612342

Corner GW. Etymology and pronunciation of the word "oestrus" and its derivatives. (1937) Science. 85(2199):197-198. PMID 17844622

Corner GW. The events of the primate ovarian cycle. (1952) Br Med J. 2(4781): 403-409.PMID 14944840

Corner GW. The observed embryology of human single-ovum twins and other multiple births. (1955) Am. J. Obstet. Gynecol., 70(5); 933-51 PMID 13258680

Ramsey EM. Corner GW. Jr. Donner MW. and Stran HM. Radioangiographic studies of circulation in the maternal placenta of the rhesus monkey: preliminary report. (1960) Proc. Natl. Acad. Sci. U.S.A., 46(7): 1003-8 PMID 16590693

PubMed References

CORNER GW. (1955). The observed embryology of human single-ovum twins and other multiple births. Am. J. Obstet. Gynecol. , 70, 933-51. PMID: 13258680

CORNER GW & NESBITT RE. (1954). Pregnancy and pulmonary resection. Am. J. Obstet. Gynecol. , 68, 903-15. PMID: 13188923

NESBITT RE & CORNER GW. (1956). Torsion of the human pregnant uterus. Obstet Gynecol Surv , 11, 311-32. PMID: 13335050

CORNER GW & STRAN HM. (1957). A fetal heart ratemeter. Am. J. Obstet. Gynecol. , 73, 190-5. PMID: 13381811

CORNER GW & STRAN HM. (1957). A recording sphygmomanometer. Am. J. Obstet. Gynecol. , 73, 196-9. PMID: 13381812

CORNER GW, KISTNER RW & WALL RL. (1951). The relationship of prolonged labor to fetal mortality. Am. J. Obstet. Gynecol. , 62, 1086-92. PMID: 14885290

RAMSEY EM, CORNER GW, LONG WN & STRAN HM. (1959). Studies of amniotic fluid and intervillous space pressures in the rhesus monkey. Am. J. Obstet. Gynecol. , 77, 1016-27. PMID: 13649772

CORNER GW, RAMSEY EM & STRAN H. (1963). Patterns of myometrial activity in the rhesus monkey in pregnancy. Am. J. Obstet. Gynecol. , 85, 179-85. PMID: 14023069

RAMSEY EM, CORNER GW & DONNER MW. (1963). Serial and cineradioangiographic visualization of maternal circulation in the primate (hemochorial) placenta. Am. J. Obstet. Gynecol. , 86, 213-25. PMID: 13990708


CORNER GW. (1963). Exploring the placental maze. The development of our knowledge of the relation between the bloodstreams of mother and infant in utero. Am. J. Obstet. Gynecol. , 86, 408-18. PMID: 14040479

CORNER GW. (1963). The fetal and maternal circulation of the placenta. Clin Obstet Gynecol , 6, 17-25. PMID: 14023070


CORNER GW, FARRIS EJ & CORNER GW. (1950). The dating of ovulation and other ovarian crises by histological examination in comparison with the Farris test. Am. J. Obstet. Gynecol. , 59, 514-28. PMID: 15410346

CORNER GW, FARRIS EJ & CORNER GW. (1950). The dating of ovulation and other ovarian crises by histological examination in comparison with the Farris test. Am. J. Obstet. Gynecol. , 59, 514-28. PMID: 15410346

BOWYER TS & CORNER GW. (1947). Premature quadruplets. Am. J. Obstet. Gynecol. , 54, 1033-7. PMID: 20272310

CORNER GW. (1959). Endocrine factors in the etiology of spontaneous abortion. Clin Obstet Gynecol , 2, 36-44. PMID: 13639313

Corner GW & Allen WM. (2005). Physiology of the corpus luteum. 1929. Am. J. Obstet. Gynecol. , 193, 1574; discussion 1575. PMID: 16202759 DOI.

Corner GW & Harris BA. (1979). Sterilization by mini-laparotomy. Female Patient , 4, 51-3. PMID: 12261211

Corner GW. (1974). Herbert McLean Evans. Biogr Mem Natl Acad Sci , 45, 153-92. PMID: 11615647

Amoroso EC & Corner GW. (1972). Herbert McLean Evans, 1882-1971. Biogr Mem Fellows R Soc , 18, 83-186. PMID: 11615755


CORNER GW. (1963). GEORGE HOYT WHIPPLE, NOBEL PRIZE PATHOLOGIST. Trans Stud Coll Physicians Phila , 31, 40-1. PMID: 14044626

CORNER GW. (1962). [The hospital of the Rockefeller Institute]. Scalpel (Brux) , 115, 18-21. PMID: 13881387

CORNER GW. (1960). The past of anatomy in the United States. Anat. Rec. , 137, 179-82. PMID: 13812017

CORNER GW. (1958). The role of anatomy in medical education. J Med Educ , 33, 1-9. PMID: 13491981

Hoffmann D, Adams JD, Lisk D, Fisenne I & Brunnemann KD. (1987). Toxic and carcinogenic agents in dry and moist snuff. J. Natl. Cancer Inst. , 79, 1281-6. PMID: 3480379

CORNER GW. (1956). The histological dating of the human corpus luteum of menstruation. Am. J. Anat. , 98, 377-401. PMID: 13362120 DOI.

CORNER GW. (1954). Anatomical terminology. Science , 120, 124-5. PMID: 13178670

PALLIE W, CORNER GW & WEDDELL G. (1954). Nerve terminations in the myometrium of the rabbit. Anat. Rec. , 118, 789-811. PMID: 13158877

CORNER GW & GOODWIN WE. (1953). Benjamin Franklin's bladder stone. J Hist Med Allied Sci , 8, 359-77. PMID: 13109181

CORNER GW & CSAPO A. (1953). Action of the ovarian hormones on uterine muscle. Br Med J , 1, 687-93. PMID: 13032456

CSAPO AI & CORNER GW. (1953). The effect of estrogen on the isometric tension of rabbit uterine strips. Science , 117, 162-4. PMID: 13038473

CORNER GW. (1953). The discovery of smooth muscle in the uterus. Acta Physiol Lat Am , 3, 67-70. PMID: 13138239

CSAPO AI & CORNER GW. (1952). The antagonistic effects of estrogen and progesterone on the staircase phenomenon in uterine muscle. Endocrinology , 51, 378-85. PMID: 13010191 DOI.

CORNER GW. (1952). The events of the primate ovarian cycle. Br Med J , 2, 403-9. PMID: 14944840

CSAPO A & CORNER GW. (1951). Shortening of uterine muscle at different temperatures. Proc. Soc. Exp. Biol. Med. , 78, 266-9. PMID: 14891987

CSAPO A & CORNER GW. (1951). In vitro contraception of pseudopregnant uterine muscle contrasted with estrous motility. Endocrinology , 49, 349-68. PMID: 14887648 DOI.

CORNER GW. (1951). Our knowledge of the menstrual cycle, 1910-1950. Lancet , 1, 919-23. PMID: 14825858

CORNER GW. (1950). The relation of the ovary to the menstrual cycle; notes on the history of a belated discovery. An Fac Med Univ Repub Montev Urug , 35, 758-66. PMID: 14771519

CHAPMAN EM & CORNER GW. (1948). The collection of radioactive iodine by the human fetal thyroid. J. Clin. Endocrinol. Metab. , 8, 717-20. PMID: 18880940 DOI.

CORNER GW. (1947). Medical treasures in the library of the American Philosophical Society. Science , 106, 120. PMID: 20344397

HARTMAN CG & CORNER GW. (1947). Removal of the corpus luteum and of the ovaries of the rhesus monkey during pregnancy; observations and cautions. Anat. Rec. , 98, 539-46. PMID: 20255047

BARTELMEZ GW, CORNER GW & HARTMAN CG. (1946). Phases of the menstrual cycle in the macaque monkey. Anat. Rec. , 94, 512. PMID: 21020638


Corner GW. (1943). SPELLING OF THE ADJECTIVE "HYPOPHYSEAL". Science , 97, 67-8. PMID: 17799311 DOI.

Boyden EA, Clark SL, Danforth CH, Greulich WW & Corner GW. (1942). COMMITTEE ON ANATOMICAL NOMENCLATURE. Science , 96, 116. PMID: 17809984 DOI.

Lewis FT & Corner GW. (1938). THE FIFTIETH ANNIVERSARY OF THE AMERICAN ASSOCIATION OF ANATOMISTS 1888-1938. Science , 87, 531-4. PMID: 17746961 DOI.

Corner GW. (1937). ANATOMICAL NOMENCLATURE. Science , 85, 428. PMID: 17814219 DOI.


Allen WM, Butenandt A, Corner GW & Slotta KH. (1935). NOMENCLATURE OF CORPUS LUTEUM HORMONE. Science , 82, 153. PMID: 17811944 DOI.

Corner GW. (1921). THE OVARIAN CYCLE OF SWINE. Science , 53, 420-1. PMID: 17773390 DOI.

Search Pubmed: GW (Author)

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  Historic Papers: 1894 Blastodermic Vesicle | 1903 12mm Pig | 1903 Pig Adrenal | 1905 Thymus | 1906 Testis | 1908 Pancreas | 1908 Pharyngeal Pouches | 1908 Intestinal Diverticula | 1910 Hypoglossal Ganglia | 1911 Prenatal Growth | 1911 Embryo 7.8 mm | 1916 Colon | 1916 Yolk Sac | 1918 Wolffian body | 1919 Corpus Luteum | 1919 Postnatal Thyroid | 1919 Placental Cord | 1921 Estrous and Implantation | 1922 Limb Arteries | 1924 Pig | 1937 Coronary Circulatory | 1938 Abnormal Brain
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Maturation of the Ovum in Swine

George Corner (1889 - 1981)

By George W. Corner

From the Anatnmical Laboratory, University of California


Since the first description of maturation of the ovum of the rabbit by Van Beneden in 1875, the study of this remarkable phenomenon has boon extended to certain other niannnals. with the result that practically all observers now agree that two polar bodies are formed by every ovum, the first of which is extruded from the egg while still in the ovary, but immeiliately before rupture of the follicle. Formation of the second jwlar body then proceeds to the formation of a division-spindle, but goes no farther until the egg is discharged and fertilized. .Vfter the entrance of the spermatozoon the .seconil polar body is completed and extrudcil. In the event that fertilization does not occur, the ovum degenerates in the state in which it was freed from the ovary.

Full and comprehensive observations of the proc(>ss of maturation have been made in the rat (Sobotta and Hunkhard '10), guinea-pig (Rubaschkin '()o), opossum (Hartman 'IG), a bat, Vesperugo noctula (Van der Stricht '08), the domestic cat (Longley '11), and the mouse. The studies of Sobotta ('95) and others, lead them to believe that in the last named species many eggs form but one polar body. The interest excited by this statement has caused the mouse to be more fully investigated than any other, with tlie result that the views ciuoted have been proven ineoiTeet. The reader is referred to the work of I>ong and Mark Cll) for details of the question together with the most comprehensive account of maturation yet presenteti in any nuimmal.

The final disproof of this supposed exceptional case raises the question as to whether the process may not be identical in all manunals. To answer this a wider search will be necessary, for the species named above comprise but three rodents, one marsupial, one carnivore, and one chiropter. There are also old or undetailed accounts or preliniinary notes ujion the ova of the dog (O. \ an der Stricht 'OS), tlie rabbit (Van Beneden 75, Heape '05), the niarsuiual Dasyurus viverrinus or Australian "native eat" (Hill '10). an insectivore. Tupaia javanica (Hubrecht '95), and of a lemuroid ape. Tarsius spectrum (Hubrecht '02), and thus of the twelve orders of mammals there are four in which the process of maturation is well known in one species or more, two in which it is very obscure, anil six in which ripening ova have never been seen.

Among the last are the ungulates, in which the species are either rare and inaccessible, or if conunon, are so large that the search for the ova is very difficult. For these reasons it will be of interest to describe a small series consisting of 15 ova from 7 sows (sus scropha domesticus). I am indebted to Mr. Ralston B. Brown, Sui)eriiiten(lent of the Oakland Meat and Packing Company of Oakland. California, for the opportunity to collect the material, and to Mr. A. E. Amsbaugh and Mr. Felix H. Hurni for a.ssistance in its preparation. The sows were selected and the tubal ova found by the method given in a previous paper (Corner and Amsbaugh '17). Ovarian ova wer6 studied in serial sections of the follicles (celloidin) or by cutting hardened follicles into sliees, locating the discus proligerus and sectioning it in paraffin.


  1. Killed during oestrus, probably on the first day. Unruptured follicles. Three ovarian ova sectioned, two show germinative vesicles, the other shows the first polar body and the second polar spindle.
  2. First or second day of oestrus. One follicle ruptured, its ovum not found in the tubes. An ovarian ovum shows the first polar body and the second spindle.
  3. Second or third day of oestrus. All follicles ruptured. Four imfertilized tubal ova sectioned: all show the first polar body and the second spindle.
  4. History unknown. Five unfertilized tubal ova found, of which four (studied fresh) show one polar body extruded.
  5. Probably socoml day of oostrus. Copulation observed 24 hours before killing. One tubal ovum found, covered with spermatozoa. In sections, the male and female pronuclei are in contact. Two polar bodies.
  6. Second or third day of oostrus. Copulation observed 10 hours before killing. Two ova found in the tubes; both were covered by spermatozoa. One of them, studied fresh, showed two polar bodies. The other was sectioned, and showed the pronuclei in contact, but unfortunately the polar bodies were obscured by damage to the sections.
  7. Tubes contained segmenting ova of 2, 4. and G blastomeres. One of the two-celled embryos (studied fresh) showed two polar bodies; a favorable view of the others could not be obtained.


The first ova of an ungulate mammal to be studied indicate that the sequence of maturation is the same in swine as in previously stuilictl forms of other orders, the first polar body being extrudetl and the second polar division proceeding as far as spindle formation before fertilization, the second polar body being cut off only after the entrance of the spermatozoon.

Literature Cited

VAN Benkden, E. 1S7.3 La maturation de I'oeuf. la fecondation, Cetc.) chez le Lapin. l$iill. do I'.Vcad. Royale de Bclpiqup. T. 40, no. I'J, p. fiSfi.

CoUNER, G. W. AM) .Vmsuaui;!!, a. E. 1917 Oestrus and ovulation in swine. Anat. Rec, vol. 12, p. 287.

Hakt.max, v. O. 191C Studies in the development of the opossum, Didelphy virjiiniana L. Jour. Morph., vol. 27, p. 1.

Heape, W. 1905 Ovulation and deffcneration of ova in the rabbit. Proc. Royal Soeiety, London, vol. 71), B, p. 2()0.

Hli.l,, J. P. 1910 The early development of the marsupialia with especial reference to the native cat I Dasyurus viverrinusi. Quart. Jour. Micros. Sci., vol. 5f), p. 1.

HuBRECHT. .\. A. W. 1,S95 Die Phylogenose des .^mniros und die Bedeutung des Trophoblasts. Verhandelingen der Kon. .\kad. van Wctenschap pcn. te .\ni8terdam. Tweedo .'^ectie, Dec!. 4, No. 5.

1902 Furrhung und Keimblattbildung bei Tarsius spectrum. Ibid., Dccl 8, No. 6.

Long, J. A. and Mahk, E. L. ISMl Tin- imitiirntion of tlif fn\i of llie nio\i8C. Puliliriilion N<i. 142 of the CariicKii' Iiuslitiition of WaKhiiiKton.

LoNfiLEY, W. H. mil Till- iiiiktiiratioii of the ccfc nnd ovulution in the (lumestir cut. Am. Jour. Anat.. vol. V2. p. 13'J.

RrBASciiKiN. \V. 10(15 I'i'Imt <lir Ki'ifuiiR.s — iiiul I)<>fru(-htiiii):i<|)rozes8c dcs Mccrsrhweinclu'iK'icK. Anat. Ilcflc, Bd. 29, S. 507.

SoDOTTA. J. 1895 Die HefriirhtunK und Furchung dcs Eic8 dcr Mans. Arch. f. Mikr. Anat., Hd. J.i. S. 15.

SoBOTTA, J. AND BrHrHKAiiD. (i. L. 1910 Kcifung und Bofruchtiing dcs Eies

dcr Wcissi-ii Ilattr. Aiml. Hcftc. Bd. 42, S. 43.3. VA.N DEK i^THKiiT. O. 1!K)S La Structure <ic I'ocuf chcz la chioniic. C. R.

dc I'Assoc. dcs .Vnatoiiiists. Dixicmc Hounion. Marseille.

1910 La structure <le I'oeuf des inaniiniferes, (C'hauve-souris, Vcspc rugo noctulai. Troisieme partie. Memoire presentc a la classc des isciences, Academic Hoyalc de Bel|{i(|ue, 2 ""^ Serie, Tome 2.

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