Richard Goldschmidt | |
|---|---|
 In his laboratory | |
| Born | (1878-04-12)April 12, 1878 Frankfurt am Main, Germany |
| Died | April 24, 1958(1958-04-24) (aged 80) |
| Alma mater | University of Heidelberg |
| Known for | "Hopeful monster" hypothesis |
| Scientific career | |
| Fields | genetics |
| Institutions | University of Munich,UC Berkeley |
| Doctoral advisor | Otto Bütschli |
| Other academic advisors | Richard Hertwig |
Richard Benedict Goldschmidt (April 12, 1878 – April 24, 1958) was a Germangeneticist. He is considered the first to attempt to integrate genetics, development, and evolution.[1] He pioneered understanding ofreaction norms,genetic assimilation,dynamical genetics,sex determination, andheterochrony.[2] Controversially, Goldschmidt advanced a model ofmacroevolution throughmacromutations popularly known as the "Hopeful Monster" hypothesis.[3]
Goldschmidt also described the nervous system of thenematode, a piece of work that influencedSydney Brenner to study the "wiring diagram" ofCaenorhabditis elegans,[4] winning Brenner and his colleagues theNobel Prize in 2002.
Goldschmidt was born inFrankfurt-am-Main, Germany to upper-middle class parents ofAshkenazi Jewish heritage.[5] He had aclassical education and entered theUniversity of Heidelberg in 1896, where he became interested innatural history. From 1899 Goldschmidt studied anatomy and zoology at theUniversity of Heidelberg withOtto Bütschli andCarl Gegenbaur. He received his Ph.D. under Bütschli in 1902, studying development of thetrematodePolystomum.[2]
In 1903 Goldschmidt began working as an assistant toRichard Hertwig at theUniversity of Munich, where he continued his work on nematodes and theirhistology, including studies of the nervous system development ofAscaris and the anatomy ofAmphioxus. He founded the histology journalArchiv für Zellforschung while working in Hertwig's laboratory. Under Hertwig's influence, he also began to take an interest inchromosome behavior and the new field of genetics.[2]
In 1909 Goldschmidt became professor at theUniversity of Munich and, inspired byWilhelm Johannsen's genetics treatiseElemente der exakten Erblichkeitslehre, began to studysex determination and other aspects of the genetics ofLymantria dispar, the gypsy moth, of which he wascrossbreeding different races. He observed various stages of their sexual development, and found that some of the animals were neithermale, norfemale, norhermaphrodites, but represented a whole spectrum ofgynandromorphism. He named them 'intersex', and the phenomenon accordingly 'intersexuality' (Intersexualität).[6] His studies of the gypsy moth, which culminated in his 1934 monographLymantria, became the basis for his theory of sex determination, which he worked on from 1911 until 1931.[2] Goldschmidt left Munich in 1914 for the position as head of the genetics section of the newly foundedKaiser Wilhelm Institute for Biology in Berlin.[7]
During a field trip to Japan in 1914, he was unable to return to Germany due to the outbreak of theFirst World War, and was detained as anenemy alien in the United States. He was placed in aninternment camp inFort Oglethorpe, Georgia for "dangerous Germans".[8] After his release in 1918, he returned to Germany in 1919 and resumed his work at the Kaiser Wilhelm Institute. Sensing that it was unsafe for him to remain in Germany, he emigrated in 1936 to the United States, where he became a professor at theUniversity of California, Berkeley. During World War II, the Nazi party published a propaganda poster entitled "Jewish World Domination" displaying the Goldschmidt family tree.[9]
Goldschmidt was the first scientist to use the term "hopeful monster". He thought thatsmall gradual changes could not bridge the divide between microevolution and macroevolution. In his bookThe Material Basis of Evolution (1940), he wrote
the change from species to species is not a change involving more and more additional atomistic changes, but a complete change of the primary pattern or reaction system into a new one, which afterwards may again produce intraspecific variation by micromutation.
Goldschmidt believed the large changes inevolution were caused bymacromutations (large mutations). His ideas about macromutations became known as the "hopeful monster" hypothesis, a type ofsaltational evolution, and attracted widespread ridicule.[10]
According to Goldschmidt, "biologists seem inclined to think that because they have not themselves seen a 'large' mutation, such a thing cannot be possible. But such a mutation need only be an event of the most extraordinary rarity to provide the world with the important material for evolution".[11] Goldschmidt believed that theneo-Darwinian view ofgradual accumulation of smallmutations was important but could account for variation only within species (microevolution) and was not a powerful enough source of evolutionary novelty to explain newspecies. Instead he believed that largegenetic differences between species required profound "macro-mutations", a source for large genetic changes (macroevolution) which once in a while could occur as a "hopeful monster."[12][13]
Goldschmidt is usually referred to as a "non-Darwinian"; however, he did not object to the general microevolutionary principles of the Darwinians. He veered from thesynthetic theory only in his belief that a new species develops suddenly throughdiscontinuous variation, or macromutation. Goldschmidt presented his hypothesis whenneo-Darwinism was becoming dominant in the 1940s and 1950s, and strongly protested against the strictgradualism of neo-Darwinian theorists. His ideas were accordingly seen as highly unorthodox by most scientists and were subjected to ridicule and scorn.[14] However, there has been a recent interest in the ideas of Goldschmidt in the field ofevolutionary developmental biology, as some scientists, such asGünter Theißen andScott F. Gilbert, are convinced he was not entirely wrong.[15][16] Goldschmidt presented two mechanisms by which hopeful monsters might work. One mechanism, involving "systemic mutations", rejected the classicalgene concept and is no longer considered by modern science; however, his second mechanism involved "developmental macromutations" in "rate genes" or "controlling genes" that change early development and thus cause large effects in the adult phenotype. These kinds of mutations are similar to those considered in contemporary evolutionary developmental biology.[17]
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