Goldschmidt, Richard Benedict

(b. Frankfurt am Main, Germany, 12 April 1878; d. Berkeley, California, 24 April 1958)

zoology, general biology.

Goldschmidt belonged to a very old German-Jewish family that had included scientists, artists, bankers, and industrialists. His father managed a coffeehouse combined with a wine trade and a confectionery. The young Goldschmidt, who had a wide circle of friends, lived in a well-to-do milieu. He attended the Gymnasium in Frankfurt and planned from the first year of secondary school to study the natural sciences: his native city possessed the famous Senckenberg Museum, a powerful attraction; and his teacher F. C. Noll was a zoologist.

Goldschmidt entered the University of Heidelberg in 1896, and among his professors were the great Otto Bütschli and Karl Gegenbaur. In 1898 he continued his studies at the University of Munich, where Richard Hertwig was teaching. In 1902, at Heidelberg, he defended his thesis on the maturation, fertilization, and embryonic development of the worm Polystomum integerrimum. A year later Goldschmidt became Hertwig’s assistant and in 1904, a Privatdozent; he remained at Munich until 1913. In that year Theodor Boveri and Carl Correns organized the Kaiser Wilhelm Institute for Biology in Berlin; Goldschmidt was appointed director of the genetics department, a post that he was to hold until 1935.

Having received a grant from the Club Autour du Monde Goldschmidt went to Japan in order to continue his research. When World War I broke out he was in Honolulu and went from there to San Francisco. He was detained in the United States, where he worked at various universities. In 1917 he was placed in an internment camp and finally was repatriated to Germany. In 1935, when conditions for Jewish scientists had become impossible under the Nazi regime, Goldschmidt decided to leave Germany. He received offers from England and Turkey, but he accepted a professorship at the University of California and left for America in July 1936. He began a new life in Berkeley, where he remained until his death. He rapidly organized a laboratory and formed a group of students and friends.

A zoologist, biologist, and geneticist of exceptional ability, an original thinker, a great traveler, and an indefatigable worker, Goldschmidt produced more than 250 memoirs and articles and about twenty books; and it is difficult to present a comprehensive view of his total output. Very broad general knowledge combined with great specialization enabled him to interpret new facts and to study individual problems thoroughly. Three orientations emerge quite clearly. He was interested at first in morphological problems and in the cytology, fertilization, meiosis, histology, comparative anatomy, and embryology of the trematodes, nematodes (Ascaris), and the Acrania. From the time of his appointment at Munich he was concerned with many students who were preparing dissertations. In 1906 he provided them with a vehicle for publication by founding a new journal, Archiv für Zellforschung.

During the same period Goldschmidt undertook a series of researches on moths of the genus Lymantria. The work lasted for twenty-five years. He was interested in a problem of microevolution: industrial melanism. Through recognizing that the melanic mutant possesses a selective advantage, he became one of the pioneers of population genetics. Goldschmidt experimented on the genetics of sex determination from 1911 to 1920 and pointed out the existence of intersexuality (a term coined in 1915), which he distinguished from gynandromorphism. He succeeded in obtaining at will all the degrees of intersexuality, up to a complete inversion of the genetic sex into the opposite sex. To account for these appearances, he constructed a coherent theory in which a quantitative balance intervenes between the male and female sex factors.

Since the Lymantria also presented enormous geographic variation, Goldschmidt undertook (1918-1933) an analysis of the genetics of geographic variation. He accepted the neo-Darwinian proposal that a geographic race represents a nascent species, but the numerous crossings carried out among geographic races from all parts of the world modified his opinion. While on a visit in Ithaca, New York (1933), Goldschmidt realized that geographic variation entails only a microevolution in the species and that it could not be the source of true evolution. He postulated the existence of macromutants, produced by alteration of the early embryonic processes; these he called “hopeful monsters.” In 1940 he pursued his critique of neo-Darwinian conceptions and argued for the existence of macroevolution carried out by means of macromutations.

The Lymantria became the subject of a new series of investigations on the theory of the gene. As early as 1916 Goldschmidt had fashioned a physiological theory of heredity (one gene, one enzyme). He published it only in 1920 in a book which marks the beginning of physiological genetics. According to this theory it should be possible, by modifying the speeds of the chains of reactions, to produce insects among which the nonhereditary phenotype copies the phenotype of the mutations; these are the phenocopies (1935). With his students Goldschmidt was able to produce varied phenocopies. At this time he left the Lymantria, which had been widely studied, and selected the Drosophila for examination.

With this change in material the third major period of research began. Goldschmidt studied the physiological genetics of the Drosophila and established that in the vestigial series the genetically controlled clipping of the wings can be influenced by the introduction of dominance modifiers. He then proposed an unorthodox theory concerning the nature of the gene, rejecting its corpuscularity (1938). This position aroused violent reactions, but in 1951 Goldschmidt opened the symposium on the gene at Cold Spring Harbor, New York, with an exposition of his ideas. His last works concerned the podoptera effect; the homoeotic mutants of Drosophila melanogaster, Podoptera and Tetraltera, hold great morphological, genetic, and evolutionary interest.

An amateur of the history of science, Goldschmidt wrote biographies of biologists he had known throughout the world. In addition, he wrote popular articles and books on science. Goldschmidt had, in addition, refined and discriminating taste; he particularly loved music and oriental art.

BIBLIOGRAPHY

I. Original Works. Goldschmidt’s articles include the following: “Zur Entwicklungsgeschichte der Echinococcusköpfchen,” in Zoologische Jahrbücher, Anatomie, 13 (1900), 467-494; “Untersuchungen über die Eireifung, Befruchtung and Zelltheilung bei Polystomum integerrimum,” in Zeitschrift für wissenschaftliche Zoologie, 71 (1902), 397-444; “Der Chromidialapparat lebhaftfunktionierender Gewebzellen,” in Zoologische Jahrbücher, Anatomie, 21 (1904), 1-100; “Amphioxides,” in Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition auf dem Dampfer “Valdivia” 1898-1899, 12 (1905), 1-92; “Lebensgeschichte der Mastigamöben Mastigella uitrea n. sp. und Mastigina setosa n. sp.,” in Archiv für Protistenkunde, supp. 1 (1907), 83-168; “Das Nervensystem von Ascaris lumbricoides und megalocephala,” in Zestschrift für wissenschaftliche Zoologie, 90 (1908), 73-136, and 92 (1909), 306-357, repr. in Festschrift R. Hertwig, II (Jena, 1910), 254-354; “Die cytologische Untersuchungen über Vererbung und Bestimmung des Geschlechtes,” in Die Vererbung und Bestimmung des Geschlechtes (Berlin, 1913), pp. 73-149; “A Preliminary Report on Some Genetic Experiments Concerning Evolution,” in American Naturalist, 52 (1918), 28-50; “Untersuchungen über Intersexualität,” in Zeitschrift für induktive Abstammungs- und Vererbungslehre, 23 (1920), 1-199; 29 (1922), 145-185; 31 (1923), 100-133; 49 (1929), 168-242; 56 (1930), 275-301; 67 (1934), 1-40; “Untersuchungen zur Genetik der geographischen Variation,” in Archiv für Entwicklungsmechanik der Organismen, 101 (1924), 92-337; 116 (1929), 136-201; 126 (1932), 277-324, 591-612, 674-768; 130 (1933), 266-339, 562-615; “Lymantria,” in Bibliotheca genetica, 11 (1934), 1-185; “The Time Law of Intersexuality,” in Genetica, 20 (1938), 1-50; “The Structure of Podoptera, a Homoeotic Mutant of Drosophila melanogaster,” in Journal of Morphology, 77 (1945), 71-103; “Ecotype, Ecospecies and Macroevolution,” in Experientia, 4 (1948), 465-472; “Fifty Years of Genetics,” in American Naturalist, 84 (1950), 313-340; “The Maternal Effect in the Production of the Beaded-Minute Intersexes in Drosophila melanogaster,” in Journal of Experimental Zoology, 117 (1951), 75-110; “The Podoptera Effect in Drosophila melanogaster,” in University of California Publications in Zoology, 55 (1951), 67-294, written with A. Hannah and L. K. Piternick: “Homoeotic Mutants and Evolution,” in Acta biotheoretica, 10 (1952), 87-104; “Heredity Within a Sex Controlled Structure of Drosophila,” in Journal of Experimental Zoology, 122 (1953), 53-96; “Materials for the Study of Dominant Personality Traits,” in Folia hereditaria et pathologica, 2 (1953), 267-295; “The Genetic Background of Chemically Induced Phenocopies in Drosophila,” in Journal of Experimental Zoology, 135 (1957), 127-202, and 136 (1957), 201-228, written with L. K. Piternick.

His books include Zoologissches Taschenbuch für Studierende (Leipzig, 1907; 6th ed., 1912), written with E. Selenka; Einführung in die Verebungswissenschaft (Berlin, 1911; 5th ed., 1928); Die quantitativen Grundlagen von Vererbung und Artbildung (Berlin, 1920); Mechanismus und Physiologie der Geschlechtsbestimmung (Berlin, 1920); Der Mendelismus (Berlin, 1920); Physiologische Theorie der Vererbung (Berlin, 1927); Die Lehre von der Vererbung (Berlin, 1927; 4th ed., 1953); Les problemes de la sexualité (Paris, 1932); Physiological Genetics (New York, 1938); The Material Basis of Evolution (New Haven, Conn., 1940); Understanding Heredity: An Introduction to Genetics (New York, 1952); Theoretical Genetics (Berkeley, Calif., 1955); Portraits From Memory: Recollections of a Zoologist (Seattle, Wash., 1956); and In and Out of the Ivory Tower. The Autobiography of Richard B. Goldschmidt (Seattle, Wash., 1960), with twenty-seven plates and a complete bibliography.

II. Secondary Literature. On Goldschmidt or his work, see A. Kühn, “Zum 70. Geburtstag Richard Goldschmidt am 12. April 1948,” in Experientia, 4 (1948), 239-240; “R. B. Goldschmidt 1878-1958. Zoologo, geneticista, evolucionista,” in Revista de la Sociedad mexicana de historia natural, 20 (1959), 185-193, with photo.; and a special vol. devoted to him of Portugaliae acta biologica, ser. A (1949-1951), published to honor his seventieth birthday: it contains a biography by A. Quintanilha and twenty-seven memoirs contributed by European and American students and friends.

AndrÉe TÉtry