Colin Russell Austin 1914–2004

Roger V. Short

Colin Russel

Colin Russell Austin, English by birth, initially graduated in Veterinary Science from the University of Sydney in 1936. The Second World War limited his career options, but he was fortunate to be employed by the CSIRO Division of Animal Health in Sydney. In 1954 he was invited to join the staff of the Medical Research Council’s laboratory in Mill Hill, London to study fertilization and early embryonic development in rats and rabbits. As a result, in 1962 he was asked to teach Fertilization and Gamete Physiology at theMarine Biological Laboratory,Woods Hole, Massachusetts, and subse- quently became Professor of Embryology in the Medical School at Tulane University, New Orleans. This alerted the University of Cambridge to his potential and they created a special Charles Darwin Chair for him in 1967. This enabled him to support the work of his young student Robert Edwards on human in vitro fertilization and embryonic development that culminated in the award of the Nobel Prize to Edwards and Patrick Steptoe in 2010. Austin also devoted a great deal of his time to editing the 13-volume Cambridge University Press series of textbooks, Reproduction in Mammals, completing the series from his retirement home in Buderim, Queensland in 1986.

Colin Russell Austin, known from childhood as ‘Bunny’, was born in Sydney on 12 September 1914 and spent his early days in India where his father, a Lieutenant Colonel in the British Army, was stationed during the First World War. At the end of the war, the family returned to England and then emigrated to Australia when Bunny was 15. After finishing his secondary education he enrolled at the University of Sydney, where he obtained a Bachelor of Veterinary Sci- ence degree in 1936. Since veterinary practice did not appeal, he continued at University, completing a Bachelor of Science degree in 1938 and then working towards a Master of Science degree in Biochemistry that he was awarded in 1940. In that year, he became a member of the research staff of the Division of Animal Health of Australia’s Council for Scientific and Industrial Research (CSIR, later CSIRO), where he stayed on the payroll until 1954 except for a couple of years, 1941–3, during the Second World War when he was seconded to the Dried Fruits Section of the Council’s Division of Food Preservation and Transport, working on Army diets and nutrition. It was there that he met Patricia, a CSIR librarian, and they married in 1941. They had two sons, Mark and Richard. In 1947,CSIR sent Bunny towork at the Medical Research Council laboratory in Mill Hill, London, where he spent a year before returning to Australia. He clearly made a very favourable impression, because in 1954 he was appointed to a permanent post in the newly created National Institute for Medical Research at Mill Hill. In the same year, he was awarded a Doctor of Science degree by the University of Sydney for work on fertilization and associated phenomena in mammals. At Mill Hill, Bunny eventually became Head of the Laboratory Animals Division from 1958 to 1961. The Austin family lived in Hadley Wood, North London, for ten years, and it was during this period that his scientific career took off, building on the study of fertilization and early embryonic development in rats and rabbits that he had started in Australia in collaboration with Dr A. W. H. Braden. Between 1948 and 1956, Bunny published ten papers in Nature (8, 10, 15, 16, 21, 22, 28, 30, 39, 42), and there were more to follow. Perhaps he will be best remembered for his 1952 paper in Nature (21). He showed that neither rabbit nor rat spermatozoa can fertilize their respective ova without a period of maturation in the female reproductive tract, a process he described as capacitation. He also found the time to write a landmark book, The Mammalian Egg, summarizing all these early findings. This was published by Blackwell, Oxford in 1961. Bunny’s research in due course brought him transatlantic recognition. In 1962, he was made a F. R. Lillie Memorial Fellow and a member of the teaching staff of the Fertilization and Gamete Physiology Training Program at the Marine Biological Laboratory, Woods Hole,Massachusetts, USA, an appointment that lasted until 1968. For the first three years, he and Pat spent the months of June, July and August in that inspiring academic environment. But he also inspired those around him and so it was that he became Head of the Genetic and Developmental Disorders Research Program at the Delta Regional Primate Research Center, Covington, Louisiana and also Professor of Embryology in the Medical School at Tulane University, New Orleans, 1964–7. The availability of primates enabled him to start working on spermatozoa in the epididymis of monkeys (104), the liquefaction of primate semen (106), the preservation of primate spermatozoa by freezing (107), the use of a rectal probe for electroejaculation of apes and monkeys (111, 114) and, most importantly, the use of human postmenopausal gonadotrophin to stimulate ovarian follicular and oocyte development in monkeys (115). So humans must be next on the list! Thus it was that his most important career move came in 1967, when he returned to England to take up a Chair that had been specially created for him at the University of Cambridge, the Charles Darwin Professorship in Animal Embryology in the Department of Physiology, together with a Fellowship at Fitzwilliam College. Here he was able to provide the perfect research environment for an up-and-coming young zoologist, Robert Edwards, who Bunny took into his department to develop, with his clinician colleague Patrick Steptoe, the contentious subject of human in vitro fertilization and embryo transfer—work that would eventually earn Edwards the 2010 Nobel Prize in Physiology or Medicine. Edwards and Bunny published their first paper together in 1959, on the induction of oestrus and ovulation in rats (67), but their most significant joint publication was in 1972 (131) on ‘Initiation of human development in vitro and transfer of early embryos’, presented at a UNESCO conference in Paris. In 1986, Bob Edwards had this to say of Bunny: ‘I would like to stress my own deep debt to him during themost difficult period of human in-vitro fertilization, when he was a clear thinking supporter of the work, prepared to defend it publicly when there were few others around’. This refers to the fact that several senior Cambridge academics, including a Nobel Laureate, were deeply opposed to human in vitro fertilization and would have torpedoed all the efforts of Edwards and Steptoe, had it not been for Bunny’s unwavering support as Charles Darwin Professor. Darwin would have been proud of Bunny! Bunny retired in 1981 and afterwards he and Pat settled in Buderim, Queensland. In 1987 he was elected a Fellow of the Australian Academy of Science, a fitting recognition of a great scientific career. 

Scientific achievements

In 1986, Professor Ryuzo Yanagimachi, Professor of Anatomy and Reproductive Biology at the University of Hawaii and an extremely distinguished gamete biologist, summarized Bunny’s scientific contributions in the following words: ‘I, and I believe all reproductive biologists working on mammalian fertilization today, consider Dr Austin as the founder of the modern study of mammalian fertilization. Between 1948 and 1955, when he was in Australia, Dr Austin published 34 research papers on mammalian fertilization. His outstanding contributions to the field of mammalian reproduction are: the codiscovery of the phenomenon of sperm capacitation; the first description of the acrosome reaction of mammalian spermatozoa; the discovery of the zona reaction; and the first detailed descriptionofnormal andabnormal fertilization. Without the pioneering studies of Dr Austin we would be far behind where we are today in reproductive biology. If it were not for Dr Austin, the success of human in vitro fertilization, for example, would have been set back at least a decade, or perhaps never occurred. ’That is praise indeed.

the egg by bunny austin

Figure 1.Figure 1. Bunny Austin’s chapter ‘The Egg’ in Reproduction in Mammals, Second Edition, 1982,Volume 1, has on page 61 this beautiful drawing by John Fuller, taken from the frontispiece of William Harvey’s classic book De generatione animalium (1651). It shows the hands of Jove holding apart the two halves of an egg, from which are emerging a plant, an insect, an amphibian, a reptile, a bird, a ruminant and even a human being, beautifully summarized in the words ‘Ex ovo omnia’ that might have been Bunny’s motto.

Reproduction in Mammals

There can be no doubt that one of Bunny’s great- est achievements was as the Senior Editor and a major contributor to the Cambridge Univer- sity Press series Reproduction in Mammals that he and I brought together. When planning this series, we decided first and foremost to make the books highly readable to undergraduates, to intersperse the text with excellent illustrations drawn by our Cambridge colleague John Fuller and, if possible, to avoid using tables since these break up the flow of the argument. 
The first volume, Germ Cells and Fertiliza- tion, was published in 1972. The other volumes were:  2.  Embryonic  and  Fetal  Development, 3. Hormones in Reproduction, 4. Reproductive Patterns, 5. Artificial Control of Reproduction, 6. The Evolution of Reproduction, 7. Mechanisms of Hormone Action, and 8. Human Sexuality. These were all published between 1972 and 1980. To write the chapters, we hand-picked scientists from around the world whom we knew personally and we never had a refusal! Either Bunny or I, and sometimes both of us, had a chapter in each volume, thereby giving us a sense of ownership of the series. It met with a very warm reception from both teachers and students. 
Scientifically, however, things were changing fast, and so we decided to produce a completely new edition of Volumes 1–5 between 1982 and 1986, in response to requests from our readers for a more up-to-date and detailed treatment of the subjects. As a result, the volumes doubled in size and a few tables crept in, but John Fuller’s beautiful drawings continued to enlighten the text (Fig. 1). There were a few title changes, with Volume 3 becoming Hormonal Control of Repro- duction, Volume 4, Reproductive Fitness, and Volume 5, Manipulating Reproduction. Much of the work for the new edition was done in Australia, with Bunny now living in Buderim while I had moved from Cambridge via the University of Edinburgh to Monash University in Melbourne, meaning that we were still able to keep in touch. Appropriately, Bunny had the last word because he wrote the final chapter in the new Volume 5, which he entitled ‘Barriers to Population Control’—a prescient theme in a world in which human population growth is one of the major problems facing mankind. 


  • Books (written)
  • The Mammalian Egg. Blackwell Scientific Publica- tions, Oxford. (1961).
  • Fertilization. Prentice-Hall Inc.,Englewood Cliffs, NJ. (1965).
  • Ultrastructure of Fertilization. Holt, Rinehart and Winston, New York. (1968).

Books (edited)

  • Sex Differentiation and Development. Memoirs of the Society for Endocrinology No 7. Cambridge, at the University Press. (1960).
  • Cell Mechanisms in Hormone Production and Action. Memoirs of the Society for Endocrinology No 8. Cambridge, at the University Press. (1961). With
  • P. C. Williams.
  • A Symposium on Agents Affecting Fertility. L. &
  • A.Churchill, London. (1965). With J. S. Perry.
  • Reproduction in Mammals. Cambridge University Press. With R. V. Short.
  • Book 1. Germ Cells and Fertilization. (1972).
  • Book 2. Embryonic and Fetal Development. (1972).
  • Book 3. Hormones in Reproduction. (1972).
  • Book 4. Reproductive Patterns. (1972).
  • Book 5. Artificial Control of Reproduction. (1972).
  • Book 6. The Evolution of Reproduction. (1976).
  • Book 7.Mechanisms of Hormone Action. (1979).
  • Book 8. Human Sexuality. (1980).
  • The Mammalian Fetus in Vitro. Chapman and Hall, London. (1973).
  • Mechanisms of Sex Differentiation in Animals and Man. Academic Press, London and New York. (1981). With R. G. Edwards.
  • Reproduction in Mammals, 2nd Edition. Cambridge University Press. With R. V. Short.
  • Book 1. Germ Cells and Fertilization. (1982).
  • Book 2. Embryonic and Fetal Development. (1982).
  • Book 3. Hormonal Control of Reproduction. (1984).
  • Book 4. Reproductive Fitness. (1985).
  • Book 5. Manipulating Reproduction. (1986).

Research Reports, Abstracts, Reviews, Chapters, etc

  1. The clinical examination of urine. Lab. J. Aus-tralasia, June, pp. 1–14. (1941). With J. H. Rofe.
  2. The thiamine (vitamin B1) content of the urine of Trichosurus vulpecular. J. Proc. R. Soc. NSW.,75, 118–121. (1941). With A. Bolliger.
  3. The reproductive hormones: a review of chemicaland physical aspects. Aust. Vet. J., 17, 222–228. (1941).
  4. The determination of carotene: a critical eval-uation. J. CSIR, 17, 115–126. (1944). With J. Shipton.
  5. Endocrinology and animal production.Aust. Vet. J., 22, 48–54. (1946).
  6. The metabolism of thiamine in the sheep. Aust. J. Exp. Biol. Med. Sci., 25, 147–155. (1947).
  7. The effect of hexoestrol on the food intake of sheep. Aust. J. Exp. Biol. Med. Sci., 25, 343–346. (1947). With W. K. Whitten, M. C. Franklin and R. L. Reid.
  8. Function of hyaluronidase in fertilization. Nature, 162, 534. (1948).
  9. Phase-contrast microscopy in the study of fertilization and early development of the rat egg. J. R. Micr. Soc., 68, 13–19. (1948). With J. Smiles.
  10. Number of sperms required for fertilization. Nature, 162, 534. (1948).
  11. Fertilization and the transport of gametes in thepseudo-pregnant rabbit. J. Endocr., 6, 63–70. (1949).
  12. The fragmentation of eggs following induced ovulation in immature rats. J. Endocr., 6, 104–110. (1949).
  13. The functions of the endocrine system in preg- nancy. Aust. Vet. J., August, pp. 190–193. (1949).
  14. The fecundity of the immature rat following induced super-ovulation. J. Endocr., 6, 293–301. (1950).
  15. Fertilization of the rat egg. Nature, 166, 407. (1950).
  16. Activation and the correlation between male and female elements in fertilization. Nature, 168, 558. (1951).
  17. Observations on the penetration of the sperm intothe mammalian egg. Aust. J. Sci. Res., Ser. B, 4,581–596. (1951).
  18. The formation, growth and conjugation of the pronuclei in the rat egg. J. R. Micr. Soc., 71, 295–306. (1951).
  19. The development of pronuclei in the rat egg, with particular reference to quantitative relations. Aust. J. Sci. Res., Ser. B, 5, 354–365. (1952).
  20. The development of the rat spermatid. J. R.Micr. Soc., 71, 397–406. (1952). With C. Sapsford.
  21. The ‘capacitation’ of mammalian sperm.Nature, 170, 326. (1952).
  22. Passage of the sperm and the penetration of the egg in mammals. Nature, 170, 919. (1952). With A.W. H. Braden.
  23. Nucleic acids associated with the nucleoli of living segmented rat eggs. Exp. Cell Res., 4,249–251. (1953).
  24. The distribution of nucleic acids in rats eggs infertilization and early segmentation. I. Studieson living eggs by ultraviolet microscopy. Aust.J. Biol. Sci., 6, 324–333. (1953). With A. W. H. Braden.
  25. The distribution of nucleic acids in rats eggs infertilization and early segmentation. II. Histo-chemical studies. Aust. J. Biol. Sci., 6, 665–673. (1953). With A. W. H. Braden.
  26. Fertilization and fertility in mammals. Aust. J.Vet., May, pp. 129–132. (1953). With A. W. H. Braden.
  27. The growth of knowledge on mammalian fertil- ization. Aust. J. Vet., July, pp. 191–198. (1953).
  28. Polyspermy in mammals. Nature, 172, 82. (1953). With A. W. H. Braden.
  29. An investigation of polyspermy in the rat and rab-bit. Aust. J. Biol. Sci., 6, 674–692. (1953). With A.W. H. Braden.
  30. Nucleus formation and cleavage induced in unfertilized rat eggs. Nature, 173, 999. (1954). With A. W. H. Braden.
  31. Reactions of unfertilized mouse eggs to someexperimental stimuli. Exp. Cell Res., 7, 277–280. (1954). With A. W. H. Braden.
  32. Time relations and their significance in the ovulation and penetration of eggs in rats and rabbits. Aust. J. Biol. Sci., 7, 179–194. (1954). With A.W. H. Braden.
  33. Induction and inhibition of the second polar division in the rat egg and subsequent fertilization. Aust. J. Biol. Sci., 7, 195–210.(1954).
  34. The reaction of the zona pellucid to sperm penetration. Aust. J. Biol. Sci., 7, 391–409. (1954). With A. W. H. Braden and H. A. David.
  35. Anomalies in rat, mouse and rabbit eggs. Aust. J. Biol. Sci., 7, 537–542. (1954). With A. W. H. Braden.
  36. The number of sperms about the eggs in mammals and its significance for normal fertilization.Aust. J. Biol. Sci., 7, 543–551. (1954). With A.W. H. Braden.
  37. Fertilization of the mouse egg and the effect ofdelayed coitus and of hot-shock treatment. Aust.J. Biol. Sci., 7, 552–565. (1954). With A. W. H. Braden.
  38. The fertile life of mouse and rat eggs. Science, 120, no. 3120. (1954). With A. W. H. Braden.
  39. Polyspermy after induced hyperthermia in rats. Nature, 175, 1038. (1955).
  40. Acquisition de la capacite fertilisatrice desspermatozoids (“capacitation”) dans les voiesgenitales femelles. In: La Fonction Tubaire, pp. 22–27.Masson et Cie, Paris. (1955).
  41. Observations on nuclear size and form in living rat and mouse eggs. Exp. Cell Res., 8, 163–172. (1955). With A. W. H. Braden.
  42. Study of fertility. Nature, 178, 185–187. (1956).
  43. An attempt to produce the Hertwig effect by X-irradiation of male mice. Studies in Fertility, 8, 121–131. (1956). With H. M. Bruce.
  44. Cortical granules in hamster eggs.Exp. Cell Res., 10, 533–540. (1956).
  45. Ovultation, fertilization and early cleavage in thehamster (Mesocricetus auratus). J. R. Micr. Soc.,75, 141–154. (1956).
  46. Effect of continuous oestrogen administrationon oestrus, ovulation and fertilization in rats and mice. J. Endocr., 13, 376–383. (1956). With H. M. Bruce.
  47. Activation of eggs by hypothermia in rats andhamsters. J. Exp. Biol., 33, 338–347. (1956).
  48. Effects of hypothermia and hyperthermia on fer-tilization in rat eggs. J. Exp. Biol., 33, 348–357. (1956).
  49. Early reactions of the rodent egg to sperma-tozoon penetration. J. Exp. Biol., 33, 358–365. (1956). With A. W. H. Braden.
  50. Environmental modification of oestrus in thevole. Nature, 179, 592–593. (1957). With H. Chitty.
  51. Preliminaries to fertilization in mammals. In:The Beginnings of Embryonic Development, pp. 71–107. Am. Assoc. Adv. Sci., Washington, DC. (1957). With M. W. H. Bishop.
  52. Sec chromatin in early cat embryos. Exp. Cell Res., 13, 419–421. (1957). With E. C.Amoroso.
  53. Fertilization in mammals. Biol. Rev., 32,296–349. (1957).
  54. Fertilization, early cleavage and associated phe-nomena in the field vole (Microtus agrestis).J. Anat., 91, 1–11. (1957).
  55. Fate of spermatozoa in the uterus if the mouseand rat. J. Endocr., 14, 335–342. (1957).
  56. Oestrus and ovulation in the field vole (Microtusagrestis). J. Endocr., 15, iv. (1957).
  57. Mammalian spermatozoa. Endeavour, 16, 137–150. (1957).
  58. Capacitation of mammalian spermatozoa. Nature, 181, 851. (1958). With M. W. H. Bishop.
  59. Research within the laboratory-animal division. In: Symposium. Organized by the Laboratory Animals Centre of MRC. Royal Society of Medicine, 5 May. (1958).
  60. Permeability of rabbit, rat and hamster egg mem-branes. Exp. Cell Res., 15, 260–261.(1958). With J. E. Lovelock.
  61. Some features of the acrosome and perfora- torium in mammalian spermatozoa. Proc. R.Soc., B, 149, 234–240. (1958). With M. W. H. Bishop.
  62. Role of the rodent acrosome and perforatorium in fertilization. Proc. R. Soc., B, 149, 241–248. (1958). With M. W. H. Bishop.
  63. Entry of spermatozoa into the Fallopian-tube mucosa. Nature, 183, 908–909. (1959).
  64. Differential fluorescence in living rat eggs treated with acridine orange. Exp. Cell Res., 17,35–43. (1959). With M. W. H. Bishop.
  65. The role of fertilization. Perspectives Biol. Med., 3, 44–54. (1959).
  66. Presence of spermatozoa in the uterine-tubemucosa of bats. J. Endocr., 18, viii–ix. (1959). With M. W. H. Bishop.
  67. Induction of oestrus and ovulation in adult rats. J. Endocr., 18, vii–viii. (1959). With R. G.Edwards.
  68. Prospective experimental animals for medicalresearch. J. Anim. Tech. Assoc., 10, 1–6. (1959).
  69. The mammalian egg. Endeavour, 18, 130–143. (1959). With E. C.Amoroso.
  70. Fertilization and development of the egg. In:Reproduction in Domestic Animals, eds. H. H.Cole and P. T. Cupps. Academic Press; New York and London. 1st Ed.Chap. 12; 2nd Ed. Chap. 13.
  71. Syngamy in a mammalian egg. Study by phase- contrast microscopy. Med. Biol. Illustr., 10, 62–63. (1960).
  72. Fate of spermatozoa in the female genital tract. J. Reprod. Fert., 1, 151–156. (1960).
  73. Capactitation and the release of hyaluronidase from spermatozoa. J. Reprod. Fert., 3, 310–311. (1960).
  74. Fertilization. In:Marshall’s Physiology of Repro- duction, 3rd edition, ed. A. S. Parkes, vol. 1, pt.2, chap.10. Longmans, Green; London. (1960). With A. Walton.
  75. Anomalies of fertilization leading to triploidy. J. Cell Comp. Physiol.,56, suppl. 1, 1–15. (1960).
  76. Egg. In: Encyclopedia of Biological Sciences,ed. P. Gray, pp. 327–328. Reinhold; New York.(1961).
  77. Significance of sperm capacitation. In: Proc. IVInt. Congr. Anim. Reprod., Hague. (1961).
  78. Fertilization of mammalian eggsin vitro. Int. Rev. Cytol., 12, 337–359. (1961).
  79. Sex chromatin in embryonic and fetal tissue. ActaCytol., 6, 61–68. (1962).
  80. Evidence against participation of a jelly-spittingagent in sperm penetration ofArbacia eggs. Biol.Bull., 123, 470. (1962). With J. Piatigorsky.
  81. Action of neuraminidase on Arbacia sperma-tozoa. Biol. Bull., 123, 471–472. (1962). With R.L. Brinster.
  82. Relationship of fertilizin to the acrosome reac- tion in Arbacia. Biol. Bull., 123, 473. (1962). With J. Piatigorsky.
  83. Passage of spermatozoa through the chorion of Ciona eggs. Biol. Bull., 123, 472. (1962). With S.D. Ezell, jr.
  84. Axial body and filament formation in oystersperms. Biol. Bull., 123, 474–475. (1962). WithD. H. Spoon and A. Forer.
  85. Introducing new animals to the laboratory. NewScientist, 17, 117–120. (1962).
  86. Fertilization in Pectinaria (=Cistenides) gouldii.Biol. Bull., 124, 115–124. (1963).
  87. Acrosome loss from the rabbit spermatozoon in relation to entry into the egg. J. Reprod. Fert., 6,313–314. (1963).
  88. Sperm morphology of Emerita talpoida. Biol.Bull., 125, 361–362. (1963). With K. R. Barker.
  89. Ultrastructure of Pectinaria gouldii gametes.Biol. Bull., 125, 364. (1963). With R. Lambson.
  90. Fine structure of Nereis limbata spermatozoa.Biol. Bull., 125, 362. (1963). With J. F. Fallon.

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About this memoir

This memoir was originally published in Historical Records of Australian Science, vol. 25, no.2, 2014. It was written by Roger V. Short Faculty of Medicine, Dentistry and Health Sciences, University of Melbourne.

© 2018 Australian Academy of Science