Boris Ivan Balinsky 10 September 1905 ? 1 September 1997 E.S. Grossman* BALINSKY WAS BORN IN KIEV, UKRAINE, THE elder of two sons of Ivan Balinsky, a his- tory teacher, and Elizabeth Radzimovsky, a biology teacher. His interest in zoology began during the summer holidays he spent in the village of Severinovka, 80 km southwest of Kiev, where his maternal grandfather, a Russian Orthodox priest, lived. The country life with beekeeping, farming and harvesting was instrumental in directing his passion for the outdoors, and a book, by Akasov on collecting butterflies which he received in 1916, marked the beginning of his interest in the natural sciences. It was fortunate that Boris?s curiosity about nature was encour- aged prior to his formal schooling, which started in 1917, the year of the Russian Revolution, as he found natural his- tory ?a great disappointment? at his first school. In 1923, he started his zoology studies under the famous evolutionary biologist, Professor I.I. Schmalhausen, at the Uni- versity of St Vladimir (subsequently the University of Kiev). All students were obliged to present a seminar as part of the course. At the suggestion of Schmal- hausen, Balinsky based his presentation on a recent paper on the specificity of ger- minal layers in Triton by Otto Mangold and this was to launch him on his career as an embryologist. A further conse- quence of the seminars was that Balinsky met his future wife, Katia Syngayevskaya, also a student, at these meetings. Balin- sky?s undergraduate experiments on the transplantation of the ear vesicle of amphibian (newt) embryos soon earned him a reputation as a promising scientist. The groundbreaking discovery of this work, the induction of supernumary limbs in newt embryos, was published as his first scientific paper, in 1925. It is rare indeed in the scientific world that a researcher?s first studies as an undergrad- uate establish a lifelong international reputation. He completed his course in 1926, but this was not concluded with the awarding of a degree: this was considered bourgeois at the time. Balinsky rose quickly to the status of professor of embryology at the university and deputy director of the Zoological Institute of the Ukrainian Academy of Sciences by 1933. In addition to his embryological studies, he examined development in inverte- brates such as sea urchins and ascidians. He revived his interest in entomology, focusing on stoneflies, which he ulti- mately collected and described in diverse locations such as the Caucasus, south- ern Germany, Scotland and southern Africa. Of more practical use, he received German lessons from an Austrian Serb, a language which was to stand him in good stead in the future. The Soviet government subsequently reversed its attitude towards academic qualifications and re-introduced scientific degrees. This enabled Balinsky to submit a monograph entitled ?Induction of the Limbs in Amphibia?, for which he obtained the doctorate of biological sci- ences from the Ukrainian Academy of Sciences in 1935. At that time the situation in the academy was very favourable, in part due to its president, academician Bogomoletz, who was held in high esteem by the Communist authorities. It was under his directorship that the acad- emy acquired control over the Marine Biological Station at Karadag in the Crimea, a move that was to play a decisive role in Balinsky?s life. Balinsky then moved on to study the development of the endoderm, which gives rise to the alimentary canal. By systematic marking and transplant exper- iments in amphibians, he was able to trace the origins of different organs arising from this inner germinal layer. This work turned out to be innovative, interesting and unexpected and led to several publi- cations. This productive period came to an end on 22 October 1937, when his wife Katia was arrested and sentenced to 10 years? confinement in a work rehabilita- tion camp for ?engaging in counter- revolutionary propaganda?, leaving Boris to care for their three-year-old son Ivan (Vania). Consequently, Balinsky was re- lieved of his lecturing duties, lost his posts of professor of embryology at the univer- sity and deputy director of the institute, although he was able to continue his work at the latter. When Katia was released 18 months later, with all charges withdrawn, his working life was restored to much the state that existed before her arrest, but nevertheless neither his professorship nor his post of deputy director was ever reinstated. Between Katia?s release and the war with Germany in 1941, Balinsky contin- ued with his work on the developing alimentary canal in amphibians, concen- trating on the interaction of events involved in the formation of the mouth. He also expanded his research to fish embryos, in particular the development of fins, using the goldfish as a model. In this way he hoped to shed some light on the transition from fins to legs in the course of evolution. Boris Balinsky?s standing in the scientific world also gradually improved after the setback it had received from Katia?s arrest. He was asked to edit two monographs, a long chapter on amphibian embryology and he started a textbook on comparative History of Science South African Journal of Science 101, May/June 2005 309 *MRC/University of the Witwatersrand Dental Research Institute, Private Bag 3, WITS 2050, South Africa. E-mail: grossmane@dentistry.wits.ac.za On 10 September 2005, we celebrate the centenary of the birth of Boris Ivan Balinsky, one of the best-known and most respected embryologists of the twentieth century. Through his remarkable and painstaking research, he laid the foundation of developmental biology as we know it today. Balinsky was a man whose research was shaped by his time, and for the first half of his working life this was dictated largely by the turbulence of the Russian Revolution and later by the instability of the Second World War. Soviet life depended on the whims of the man in power at any given time: thus, the course of Balinsky?s research was directed according to the situation in which he found himself and the facilities available to him. It is due to his insight, single-mindedness, ability to adapt, hands-on approach and meticulous technique that he managed to achieve the noteworthy research and groundbreaking findings in the years prior to, and after, his move to the West. Boris Balinsky as he was in 1975. embryology. This last had to be aborted by the beginning of the war with Ger- many. In December 1940, he received a formal distinction from the Academy of Sciences of the USSR in Moscow ? the Kawalewsky Prize, for his work on the determination of the endoderm in amphibian embryos. In 1941 he was appointed professor of zoology at the medical school in Kiev. The Germans attacked the Soviet Union on 22 June 1941. At the time the Balinsky family was at the Karadag Marine Biologi- cal Station in the Crimea, where Boris wished to combine a family holiday with some scientific work on marine animals and their embryos. With no immediate possibility of returning to Kiev, they managed to get to Kharkov, where he was able to regain contact with the Kiev Medi- cal School, which had been evacuated to the town. He also found employment with the railways as chief of the rat- catching division, a job requirement for this post being biological training! It was at this time that Balinsky made the ?most momentous decision? of his life. By avoid- ing evacuation to Siberia with the medical school and remaining in Kharkov, he opened the possibility of eventually moving to the West. Once he was able to return to Kiev, he was appointed to the staff of the Fisheries Institute, one of several research centres allowed under the German occupation. Boris?s work on goldfish embryos was instrumental in his appointment and he was able to expand these studies to include the developmen- tal stages of commercial fish, as well as identifying the larvae of the many species of fish in the waters adjoining the station. For this latter project he used the distribu- tion of pigment cells inside the transpar- ent body as a useful means of distinction. This work was published some six years later in Scotland. During the autumn and winter of 1942/3, Boris spent time writing up the results of experiments on the determination of parts of the endoderm in newts and salamanders. Using 5-watt lamps in his flat, he was able to set up an apparatus to make photomicrographs of sectioned material to illustrate the work which was subsequently published in Roux? Archiv after the war. He also prepared a manu- script on stoneflies that was published in England. It was during this time that Katia fell ill and died on 31 March 1943 after a period of poor health. By mid- September of that year it was evident that the German retreat from Kiev was only a matter of time and the institutes under German control, together with their staff, were to be evacuated to Posen in the west- ern part of Poland. Before the end of the month, Boris, his mother Elizabeth and Vania left for Poland, accompanied by three packing cases containing their personal belongings and all of Boris?s scientific records and research equip- ment. In Posen, Balinsky was able to section most of the embedded embryos prepared during 1941. In the spring of 1944 he was also able to obtain additional fresh material from fish with commercial value, such as the carp and roach, and was able to complete the developmental stages of these fish with every stage accurately drawn in black ink and with shading ? a painstaking task. He also designed what he came to regard as his ?personal coat of arms?, a five-legged newt, in recognition of his first success in experimental embry- ology. This he ultimately used as a frontis- piece in his book on embryology. In December 1944, the institute was evacu- ated yet again, this time to Marburg. As it turned out, the Fisheries Institute was never re-assembled in Marburg and Balinsky lost virtually all the results of five years of work including his drawings of fish stages. He found respite in the Zool- ogy Department at the University of T?bingen. The German scientific commu- nity recognized his lack of income and gave him laboratory space for research whilst paying him a stipend from the Notgemeinschaft der Deutschen Wissen- schaft. This was not to last. By December 1945 he moved to the American Occupa- tion Zone in Heidelberg to avoid enforced repatriation. This was a step which took him closer to his goal of working in the West. He then served as professor of histology and embryology and head of department at the Displaced Persons University, orga- nized in Munich under the auspices of the United Nations Repatriation and Rehabilitation Administration for so long as it existed (1945?47). A textbook on his- tology for students taking the course (Vorlesungen ?ber Histologie) serves as a tangible reminder of this time, for which Boris wrote and prepared 85 drawings. In Munich, he also met Elizabeth (Betty) Stengel, who was to become his second wife, on 15 March 1947. The post-war situation in Germany was desperate: and although Balinsky wished to emigrate and settle in the United States, their American intelligence screening was unfavourable. Argentina was also a possi- bility but that would have meant working as a farmer, something that did not appeal. Boris sent his CV around the Eng- lish-speaking world and had two ap- proaches: a tentative one from Professor van der Horst, head of the Department of Zoology at the University of the Wit- watersrand in Johannesburg, the other a firm offer from Professor Waddington at the Institute of Animal Genetics of the Agricultural Research Council in Edin- burgh, Scotland. Unfortunately, the con- dition was that he come to Scotland alone, without his family. Having no real choice, he went to Scotland in October 1947, with his mother, Betty and Vania joining him six months later. Boris?s initial work was to investigate the factors controlling the number and location of milk glands in mice. This was subsequently expanded to a study of the morpho-physiological mechanism of the early stages of develop- ment of the mammary glands in rabbits and cattle. During this time he conceived the layout for a book on embryology, which he followed some years later as An Introduction to Embryology. In 1949 he received an offer of a lectureship at the University of the Witwatersrand (Wits) which he accepted, arriving in Johannes- burg in September 1949 with his family, including his newly born daughter, Helen. Balinsky immediately started a new series of investigations on amphibians using the numerous and varied species of South African frogs. By hetero-trans- plants between embryos he gained infor- mation on factors determining the size and differentiation of organ rudiments between species. He also resumed his work on the embryology of marine inver- tebrates during visits to the Biological Marine Station on Inhaca Island, off the Mozambique coast. Within five years (1955), he was promoted to fill the chair of zoology and headship of the department at Wits. In 1956 Balinsky went to Yale Uni- versity for his sabbatical leave, where he was introduced to the two outstanding pioneers of biological electron micros- copy, Drs G.E. Palade and K.R. Porter of the Rockefeller Institute in New York. With growing excitement, he realized that the techniques of electron microscopy would help him solve a number of prob- lems in his work on limb induction. On his return to Johannesburg, he was delighted to learn that the university had in the mean time acquired a Siemens Elmiskop 1. This enabled him to use elec- tron microscopy to advance his embryo- logical studies, which he expanded to include sea urchins and insects. In doing so he was the first person to practise biological electron microscopy in this country. One of the great achievements of Boris Balinsky was the manner in which he pioneered the application of the electron microscope to the study of the ultra- structure of early development, how he foresaw the relevance of molecular biol- ogy to the understanding of development and the way in which he achieved a syn- thesis of these two approaches. This inter- est led him to become a founder member of the Electron Microscope Society of Southern Africa, of which he served as 310 South African Journal of Science 101, May/June 2005 History of Science president between 1962 and 1973. The Boris Balinsky Lecture is given at the society?s annual meetings to honour his contribution. In addition, he was an active member of the South African Association for the Advancement of Science, the Ento- mological Society of Southern Africa and the International Institute of Embryology. Boris Balinsky filled the chair of zoology and headship of the department until his retirement at the end of 1973. He was dean of the Faculty of Science from 1965 to 1967 and was awarded the degree of D.Sc. honoris causae by the University of the Witwatersrand in 1978. He was a dedi- cated and productive researcher until he died, publishing 133 research papers and several books, mainly in the fields of experimental embryology and entomol- ogy. A paper on the electron microscope investigation of frog development was celebrated as a ?citation classic? by the journal Current Contents in 1984. While Balinsky?s experimental work on the induction of supernumary limbs brought the greatest international recog- nition and acclaim, his textbook entitled Introduction to Embryology, which was first published in 1960, secured his interna- tional reputation. The book was based on courses given to students at the Univer- sity of the Witwatersrand. Such was its impact that it has been published in five English editions, two Japanese, two Italian and one in Spanish. It was cer- tainly the most widely used embryology textbook in the world and has influenced the education of countless numbers of students. Balinsky was a man of many talents, many interests and many achievements. His abiding interest in insects, which started in 1916, led to ongoing collecting of stoneflies, dragonflies, butterflies and moths. This resulted in his identifying, describing and naming several new species of the families Odonata and Plectoptera. On his retirement, he was ap- pointed professor emeritus and honorary research professorial fellow by Wits University in recognition of his many achievements. He worked on the classifi- cation of moths for the Transvaal Mu- seum and, in 1984, he was awarded the status of Associate Member of the mu- seum. Even in the last years of his life he engaged in yet another new field of study, the genetics of the butterfly Acraea horta. Among his other interests were music, piano-playing, oil-painting, gardening and astronomy. He is one of the few people who have observed Halley ?s comet twice. His deep love of animals and nature also led to extensive travels in the wilderness areas of southern Africa, with family, friends and students from Wits. Scientists who have achieved interna- tional recognition across a range of disciplines are rare indeed. Boris Ivan Balinsky was such a person, through his pioneering contributions in the fields of embryology, entomology, genetics, and herpetology. This article is based largely on Balinsky?s own account of events, courtesy of the Boris I. Balinsky Memoir, Record Series 15/35/57, in the University of Illinois at Urbana-Champaign Archives. It was prepared under the aegis of the Microscopy Society of Southern Africa. List of publications 1. Transplantation des Ohrbl?schens bei Triton. Roux? Archiv 105, 1925, 718?731. 2. Weiteres zur Frage der expetimentellen Induk- tion einer Extremit?tenanlage. Roux? Archiv 107, 1926, 679?683. 3. Experimental limb-induction. Travaux de l?Institute Biologique (Kiev) 1, 1926, 361?382. (in Ukrainian). 4. I. Schmalhausen and B.I. Balinsky. On the speci- ficity of the skeletogenous tissue after experi- ments of transplantation of the ear vesicle in Triton. Trans. 2nd Congress of Zoologists, Anatomists and Histologists, Moscow, 1927, 165?167. (in Rus- sian). 5. Xenoplastische Ohrbl?schentransplantation zur Frage der Induktion einer Extremit?tenanlage. Roux? Archiv 110, 1927, 63?70. 6. Ueber die experimentelle Induktion einer Extremit?tenanlage bei Triton mit besonderer Ber?cksichtigung der Innervation und Symme- trieverh?ltnisse derselben. Roux? Archiv 110, 1927, 71?88. 7. A new demonstration of the existence of limb induction with the aid of xenoplastic transplanta- tion and the factors that are involved therein. Travaux de l?Institut Biologique (Kiev) 2, 1927, 49? 73. (in Ukrainian) 8. Activation of the latent potencies of mesoderm. Trans. 3rd Congress of Zoologists, Anatomists and Histologists, Leningrad, 1928, 169?170. (in Rus- sian). 9. Ueber die Mesodermverschiebungen bei der Extremit?teninduktion. Roux? Archiv 116, 1929, 604?632. 10. Studies on the specificness of mesoderm I. On the shifting of mesodermal material in limb- induction. Travaux de l?Institut Biologique (Kiev) 3, 1929, 125?153. (in Ukrainian) 11. Extirpation of the pronephros in Triton. Travaux de l?Institut Biologique (Kiev) 4, 1929, 355?372. (in Ukrainian). 12. A case of dependent differentiation of the fibres of the lens in the absence of an eye cup. M?m. de las classe des Sc. Phys. et Math. de l?Acad. des Sc. de l?Ukraine 12(3), 1929, 311?321. (in Ukrainian). 13. Problems of mechanics of development. I. Me- chanics of morphogenesis in skeletogenous mesenchyme. J. Exp. Biol. (Moscow) 6(4), 1930, 395?407. (in Russian). 14. Ein Fall der abh?ngigen Entwicklung von Lisen- fasern bei vollst?ndigem Mengel eines Augen- bechers bei Triton. Roux? Archiv 122, 1930, 12?21. 15. Zur Dynamik der Extremit?tenknospenbildung. Roux? Archiv 123, 1931, 565?648. 16. Ueber den Teilungsrhythmus bei der Entwick- lung des Eies der Ascidie Ciona intestinalis. Roux? Archiv 125, 1931. 17. Mechanics of the first stages of morphogenesis in vertebrates. Trans. 4th Congress of Zoologists, Anat- omists and Histologists, Kiev, 1931, 102?103. 18. On the dynamics of development of the limb bud in Urodeles. Ibid., 1931, 103?104. 19. On the rhythm of cell divisions in the develop- ment of the tunicate Ciona intestinalis. Ibid., 1931, 104?105. 20. The role of different parts in the development of the limb bud. M?m. de la Cl. des Sc. Natur. et Techn. de l?Acad. des Sc. de l?Ukraine 8, 1931, 273?380. (in Ukrainian). 21. How the Embryo is Developed from the Egg. Mono- graph, State Medical Publishers, Kiev, 1932, 1?68. (in Ukrainian). 22. Gastrulation and the germinal layers in the light of recent experimental results. Biol. J., Moscow 1(5/6), 1932, 104?125. (in Russian). 23. Interaction of two heteropolar equipotential sys- tems studied by the method of conplantation of morulae of the sea urchin Strongylocentrotus droebachiensis. J. Bio-Zoological Cycle of the Ukrai- nian Acad. Sci. 1/2, 1932, 5?13. (in Ukrainian). 24. The role of location in the limb induction. Investi- gations on the Ontogenesis of Animals (Kiev) 6, 1933, 11?22. (in Ukrainian). 25. Studies on the specificness of mesoderm. II. Selfdifferentiation of the limb-mesoderm. Inves- tigations on the Ontogenesis of Animals (Kiev) 6, 1933, 23?34. (in Ukrainian). 26. Das Extremit?tenseitenfeld, seine Ausdehning und Beschaffenheit. Roux? Archiv 130, 1933, 704?746. 27. A qualitative and quantitative investigation of the limb-forming potency in the lateral region. Travaux de l?Institut de Zoologie et Biologie (Kiev) 1, 1934, 91?135. (in Ukrainian). 28. The developmental mechanics of the paired limbs in vertebrates. Uspekhi Sovrem. Biolog. (Mos- cow) 4, 1935, 21?45. (in Russian). 29. The growth and development. In The Growth of Animals, Moscow, 1935, 85?106. (in Russian). 30. Studies on the specificness of mesoderm. III. De- velopment of interplanted mesoderm of the limb-rudiment. Travaux de l?Institut de Zoologie et Biologie (Kiev) 6, 1935, 23?56. (in Ukrainian). 31. Dependent differentiation and self differentia- tion in the development of limbs. Trans. Union Conference of Histologists, Moscow, 1935, 60?64. (in Russian). 32. The significance of the stage of development for limb-induction. Bull. sc. de l?Univ. de Kiev 1, 1935, 237?249. (in Ukrainian). 33. Experimentelle Extremit?teninduktion und die Theorien des phyiogenetischen Ursprungs der paarigen Extremit?ten der Wirbeltiere. Anat. Anzeiger 80, 1935, 136?142. 34. Selbstdifferenzierung des Extremit?tenmeso- derms in Interplantat. Zool. Jahrb., Abt. f. allg. Zool. und Physiol. 54, 1935, 327?348. 35. Embryological researches in the University of Kiev. In The Development of Science in the State Uni- versity of Kiev during 100 Years, Kiev, 1935, 171?182. (in Ukrainian). 36. The First Stages of Morphogenesis in Animals. Mono- graph, State Medical Publishers, Kiev, 1936, 1?189. (in Ukrainian). 37. The Development of the Embryo. The Problem of De- termination in Embryonic Development. Mono- graph, State Medical and Biological Publishers, Moscow, Leningrad, 1936, 1?184. (in Russian). 38. Induction of Limbs in Amphibia. Monograph, Ukrainian Acad. of Sc., Kiev, 1936, 1?160. (in Ukrainian). 39. The basic principles of organogenesis. Nature (Leningrad) 1936, 27?39. 40. Formation de structures suppl?mentaires aux d?pens du transplant et sous son influence. Ex- perimental Medicine (Kharkov) 7, 1936, 63?68. 41. Zur Frage der Natur der extremit?tenindu- zierenden Wirkung. Roux? Archiv 136, 1937, 221? 249. 42. Ueber die zeitlichen Verh?ltnisse bei der Extremi- t?teninduktion. Roux? Archiv 136, 1937, 250?285. 43. On the specificity of inductors for different stages of development. Travaux de l?Institut de Recherches Sc. de Biol. de l?Univ. de Kiev 1, 1937, 33?42. (in Ukrainian). 44. The production of supernumerary balancers in deficiency experiments in Triton embryos. C.R. (Doklady) de l? Acad. des Sc. de l?URSS 17, 1937, 503? 504. 45. The work of the Laboratory for Mechanics of Development of the Zoological and Biological In- stitute of the Academy of Sciences of the Ukrai- nian S.S.R. during 15 years (1922?1937). Uspekhi Sovr. Biol. Moscow 8, 1938, 291?302. (in Russian). 46. The processes of induction in embryonic devel- opment. Bull. de l?Acad. des Sc. de l?URSS. Ser. Biol. 3, 1937, 941?954. (in Russian). 47. On the determination of the entodermal organs in Amphibia. Reports of the Institute of Zoology and Biology (Kiev) 12, 1938, 3?36. (in Ukrainian). 48. On anomalies in the development of balancers History of Science South African Journal of Science 101, May/June 2005 311 caused by deficiencies in the ento-mesoderm of Triton embryos. Reports of the Inst. of Zool. and Biol. (Kiev) 12, 1938, 73?98. (in Ukrainian). 49. Interrelation between the normal and induced development of limbs. Travaux de l?Institut de Zoologie et Biologie de l?Acad. Sc. de la RSS d?Ukraine 18, 1938, 23?50. (in Ukrainian). 50. On the determination of entodermal organs in Amphibia. C.R. (Doklady) Acad. Sc. USSR 20, 1938, 215?217. 51. On the competence for limb induction in differ- ent amphibians. Trans. Inst. Exper. Morphogenesis, Univ. of Moscow 6, 1938, 33?42. 52. Experiments on total extirpation of the whole entoderm in Triton embryos. C.R. (Doklady) de l?Acad. des. Sc. d?URSS 23, 1939, 196?198. 53. On the development of the ectodermal stomo- daeal invagination in amphibians. C.R. (Doklady) de l?Acad. des Sc. de l?URSS 23, 1939, 199?202. 54. Individual development of animals in the light of contemporary experimental research. In Biology for the Masses 5, Kiev, 1939, 51?64. 55. Determination in the three germinal layers. Uspekhi Sovr. Biol., Moscow 11, 1939, 396?419. 56. On the significance of the differentiation of the inductor for limb-induction. Travaux de l?Institut de rescherches sc. de Biologie de l?Univ. de Kiev 4, 1940, 125?148. 57. On the determination of the branchial and cloacal part of the gut in Amphibia Urodela. In- vestigations on the Ontogenesis of Animals 13, 1940, 3?49. 58. Zur Frage der Induktion der Medullarplatte durch Extrakte und synthetische Pr?parte (in col- laboration with B.I. Goldstein, R.I. Lirzman, and E.M. Schapiro). C.R. (Doklady) de l?Acad. des Sc. de l?URSS 27, 1940, 508?511. 59. Formierung des definitiven Darmkanals bei den Amphibien (nach Versuchen der vitalen Farb- markierung). C.R. (Doklady) de l?Acad. des Sc. de l?URSS 27, 1940, 512?515. 60. Die Verh?ltnisse zwischen verschiedenen Typen von Korrelationen in der Entwicklung der ekto- dermelen Mundeinst?lpung bei den Amphibien. C.R. (Doklady) de l?Acad des Sc. de l?URSS 28, 1940, 757?759. 61. The concept of determination in the mechanics of development. Uspekhi Sovr. Biol., Moscow 13(2), 1940, 327?346. (in Russian). 62. Vorlesungen ?ber Histologie. Akademischer Verlag der Ausl?ndischen Wissenschaftler, M?nchen, 1947, 1?239. 63. Kinematik des entodermalen Materials bei der Gestaltung der wichtigsten Teile des Darmkanals bei den Amphibien. W. Roux? Archiv f. Ent. Mech. der Organismen 143(1/2), 1947, 126?166. 64. Korrelationen in der Entwicklung der Mund- und Kiemenregion und des Darmkanals bei Amphibien. W. Roux? Archiv f. Entw. Mech. der Organismen 143(5/6), 1948, 365?395. 65. On the development of specific characters in cyprinid fishes. Proc. Zool. Soc. 118(2), 1948, 335?344. 66. On the prenatal growth of the mammary gland rudiment in the mouse. J. Anat. 84(3), 1950, 227? 235. 67. On the Placoptera of the Caucasus. Trans. R. Ent. Soc. London 101(2), 1950, 59?87. 68. On the developmental processes in mammary glands and other epidermal structures. Trans. R. Soc. Edinburgh 62(1), 1950, 1?31. 69. New researches on the chemical composition of cells. S. Afr. J. Sci. 47, 1951, 223?226. 70. On the eye-cup lens correlation in some South African amphibians. Experientia 7(5), 1951, 180. 71. Growth and cellular proliferation in the early ru- diments of the eye and the lens. Quart. J. Microsc. Sci. 93(3), 1952, 357?368. 72. On the breeding habits of the South African bull- frog, Pyxicephalus adspersus. S. Afr. J. Sci. 51, 1954, 55?58. (Jointly with J.B. Balinksy). 73. Histogenetic and organogenetic processes in the development of specific characters in some South African tadpoles. J. Embryol. Exp. Morph. 3(2), 1955, 93?120. 74. The problem of development. Inaugural lecture, Witwatersrand University Press publication, 1955, 1?13. Johannesburg. 75. In P.D. Nieuwkoop and J. Faber (eds): Normal table of Xenopus laevis (Daudin). Section: The develop- ment of the intestinal tract and glands up to stage 57. North Holland, Amsterdam, 1956, 150?159. 76. Reactivity of the epidermis in lens development. S. Afr. J. Sci. 52, 1956, 155. 77. A new species of Chlerolestes (Odonata) from Na- tal. Ann. Transvaal Mus. 22(4), 1956, 511?514. 78. A new theory of limb induction. Proc. Natl Acad. Sci. (Philadelphia) 42(10), 1956, 781?785. 79. On some stoneflies (Plecoptera) from the eastern parts of South Africa. J. Ent. Soc. S. Afr. 19, 1956, 289?301. 80. New experiments on the mode of action of the limb inductor. J. Exp. Zool. 134, 1957, 239?274. 81. On the factors determining the size of the lens rudiment in amphibian embryos. J. Exp. Zool. 135, 1957, 255?300. 82. South African Amphibia as material for biological research. S. Afr. J. Sci. 53, 1957, 383?391. 83. Classification of the females in the genus Pseudagrion (Odonata) based on thoracic struc- ture. J. Ent. Soc. S. Afr. 20, 1957, 280?294. 84. Origin of cells from non-cellular living matter. S. Afr. J. Sci. 54, 1958, 61?66. 85. Notes on the distribution of Odonata in South Africa. J. Ent. Soc. S. Afr. 21(1), 1958, 227?229. 86. On the factors controlling the size of the brain and eyes in anuran embryos. J. Exp. Zool. 139(3), 1958, 403?442. 87. An electron microscopic investigation of the mechanisms of adhesion of the cells in a sea urchin blastula and gastrula. Exp. Cell Res. 16(2), 1959, 429?433. 88. B.I. Balinsky and G.V. James. Explosive reproduc- tion of organisms in the Kariba Lake. S. Afr. J. Sci. 56, 1960, 101?104. 89. An Introduction to Embryology. Saunders, Philadel- phia; 1960. 90. Two expeditions to the Kariba Lake. Convocation Commentary, University of the Witwatersrand, Johannesburg, 3, 1960, 21?25. 91. The role of cortical granules in the formation of the fertilization membrane and the surface membranes of fertilized sea urchin eggs. Symp. on germ cells and development. Inst. Intern. d?Embryol. and Fondazione A. Baselli, Pavia, 1961, 205?219. 92. Ultrastructural mechanisms of gastrulation and neurulation. Ibid., 1961, 250?263. 93. B.I. Balinksy and H. Walther. The immigration of presumptive mesoblast from the primitive streak in the chick as studied with the electron micro- scope. Acta Embryol. et Morphol. Experimentalis 4, 1961, 261?283. 94. Observations of the dragonfly fauna of the coastal region of Zululand, with descriptions of three new species (Odonata). J. Ent. Soc. S. Afr. 24, 1961, 72?91. 95. Patterns of animal distribution on the African continent. Annals Cape Provincial Museums 2, 1962, 299?310. 96. Eine elektronemikroskopische Untersuchung der Differenzierung der Oocyten beim Krallenfrosch Xenopus laevis. Verhandl. d. 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