Richard Goldschmidt

German American geneticist

• Born: April 12, 1878; Frankfurt am Main, Germany
• Died: April 24, 1958; Berkeley, California

German-born geneticist Richard Goldschmidt provoked significant debate within the fields of evolutionary biology and genetics with his theory of “hopeful monsters.” His controversial 1940 book The Material Basis of Evolution argues that macromutations provide an explanation for large evolutionary changes.

Primary field: Biology

Specialties: Genetics; evolutionary biology

Early Life

Richard Benedict Goldschmidt was born in Frankfurt am Main in the German Empire. Growing up, he received a classical education. In 1896, Goldschmidt entered Heidelberg University, where he began studies in medicine and studied under renowned zoologist Otto Bütschli and anatomist Carl Gegenbaur. During these years, Goldschmidt developed a keen and enduring interest in natural history. In 1898, he moved to the University of Munich to study with zoologist Richard Hertwig. Goldschmidt eventually returned to Heidelberg and wrote his doctoral thesis on the embryological development and maturation of the trematode (parasitic flatworm) Polystomum integerrimum, earning his doctorate in 1902. His early work on embryonics and morphology ignited his lifelong passion for genetics and launched his later research into the relationship between genetics and evolution.

Life’s Work

In 1903, Goldschmidt took his first academic appointment, under Hertwig in Munich, carrying on his research into genetics and cytology. During this time, Goldschmidt continued his studies of parasitic worms, focusing on the development of the nervous system of the genus Ascaris (parasitic nematodes, also known as giant intestinal roundworms), and also conducted anatomical studies of the lancelets of the genus Branchiostoma, small marine invertebrates that resemble small, slender fish. He also founded the journal Archiv für Zellforschung (Archive for cell research). In his early work on parasitic worms, Goldschmidt continued to develop his ideas about the nature of chromosomal development, sex determination, and the relationship of these elements to the larger development of species. While working under Hertwig, Goldschmidt married Else Kühnlein, with whom he had two children.

In 1909, Goldschmidt became a professor at the University of Munich, where he began to study the genetics of Lymantria dispar, the gypsy moth. He left Munich in 1914 to take a position with the new Kaiser Wilhelm Institute for Biology in Berlin. While at the institute, he investigated the problem of sex determination in Lymantria, and after discovering that geographical variation could play a role in genetic development, he experimented with mating geographical variants of Lymantria with one another. Goldschmidt eventually proposed a model called physiological genetics, which contended that a gene’s expression of a particular phenotype (observable properties of the organism in question) is tied to that gene’s rate of production of a substance such as an enzyme. According to this theory, if the gene produces the substance at a faster or slower rate, the resulting phenotype will be different; therefore, one gene can produce a variety of different phenotypes.

Goldschmidt’s study of gypsy moths was interrupted for several years during World War I. In 1914, he traveled to Japan to collect samples of Lymantria, intending to return to Germany to carry out further research, but the British blockade of Germany prevented him from doing so. Goldschmidt lived in the United States throughout the remainder of the war, taking on temporary positions with institutions such as the University of California. He was detained in an internment camp for Germans in 1918 and released later that year, following the end of the war. He returned to Germany and in 1920 published his first major book on his work, Mechanismus und Physiologie der Geschlechtsbestimmung (The Mechanism and Physiology of Sex Determination, 1923). Continuing his work with Lymantria, he published a groundbreaking monograph on his findingsin 1934.

Following the advent of Nazism, Goldschmidt was forced to leave his post because of his Jewish heritage. He immigrated to the United States in 1936 and took a position in the Department of Zoology at the University of California, Berkeley, where he remained until his retirement. During his tenure at Berkeley, Goldschmidt published the significant and controversial book The Material Basis of Evolution (1940). The book, which advanced his ideas about the genetic nature of evolution, made him an object of ridicule among many biologists who regarded his theories as outlandish. Goldschmidt challenged prevailing ideas about the ways in which small-scale evolution (microevolution) affects and influences large-scale evolution (macroevolution).

For most Darwinists, microevolution provides a model for macroevolution; small changes in discrete populations become the model for larger changes. Goldschmidt disputed this idea, arguing that microevolution and macroevolution are in fact two different, unrelated processes. Because microevolution involves small and gradual changes, such a process often leads to dead ends rather than the development of new species. Goldschmidt contended that species are separated from each other by a “bridgeless gap” that can only be jumped in a single genetic step. He illustrated that such steps occur through substantial and immense changes, or macromutations. Such macroevolutionary changes are abrupt, the consequence of a complete rearrangement of the chromosomal structure of a species. Goldschmidt asserted that these structural elements guiding macroevolution are systemic mutations and famously referred to the resulting organisms as “hopeful monsters,” a phrase for which he was often ridiculed by his critics.

Goldschmidt continued this work after his retirement in 1948, turning his attention to the chromosomal structure and morphology of fruit flies (Drosophilia) as he maintained his investigations into sex determination and macroevolution. He was elected president of the Ninth International Congress of Genetics in 1953 and wrote an autobiography, In and Out of the Ivory Tower, published posthumously in 1960. Goldschmidt died in 1958 in Berkeley.

Impact

Many scientists have called Goldschmidt one of the most controversial geneticists of the twentieth century. Evolutionary biologist Ernst Mayr wrote his Systematics and the Origin of Species (1942) as a direct refutation of Goldschmidt’s Material Basis of Evolution, and many of Goldschmidt’s contemporaries either forcefully rejected his ideas or openly ridiculed them. Even so, some did praise his work. In the late twentieth century, the evolutionary biologist Stephen Jay Gould defended Goldschmidt’s work and argued that he was one of the twentieth-century’s premier geneticists.

Goldschmidt was instrumental in establishing a significant connection between development and evolution. For Darwinists and neo-Darwinists, evolution is gradual and continuous; it must proceed in one direction, and selection and adaption often depend on overcoming developmental changes. Goldschmidt, however, demonstrated that morphological, genetic, and physiological development cannot be isolated from evolution and that such developments provide the key to macroevolution. Goldschmidt’s theory that evolution is based on massive and abrupt changes challenged the idea that it proceeds in a slow, gradual fashion. His ideas influenced later evolutionary biologists who argued in favor of the theory of punctuated equilibrium, in which evolutionary change occurs not in a continuous fashion but in bursts amid periods of stasis. Although many scientists do not embrace his ideas about genetics, Goldschmidt pioneered the idea that chromosomal structure can be an integral factor in evolutionary change, and his ideas about the influence of genetics on evolutionary patterns have contributed significantly to theories concerning the genetic nature of evolution.

Bibliography

Dietrich, Michael R. “Reinventing Richard Goldschmidt: Reputation, Memory, and Biography.” Journal of the History of Biology 44.4 (2011): 693–712. Print. Provides a biography of Goldschmidt and evaluates his ideas about evolutionary biology.

Goldschmidt, Richard. The Material Basis of Evolution. New Haven: Yale UP, 1940. Print. Establishes Goldschmidt’s theories of systemic mutation, macroevolution, hopeful monsters, and physiological genetics.

Gould, Stephen Jay. The Structure of Evolutionary Theory. Cambridge: Harvard UP, 2002. Print. Defends Goldschmidt from critics and distinguishes his theory of hopeful monsters from the theory of punctuated equilibrium.

Mayr, Ernst. Systematics and the Origin of Species. New York: Columbia UP, 1942. Print. A direct refutation of Goldschmidt’s The Material Basis of Evolution by one of his fiercest critics.

Ruse, Michael, and Joseph Travis, eds. Evolution: The First Four Billion Years. Cambridge: Harvard UP, 2009. Print. An introduction to the theory of evolution that contains a brief overview of Goldschmidt and his work.