Richard Goldschmidt
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Richard Benedict Goldschmidt (April 12, 1878 – April 24, 1958) was a German
geneticist A geneticist is a biologist or physician who studies genetics, the science of genes, heredity, and variation of organisms. A geneticist can be employed as a scientist or a lecturer. Geneticists may perform general research on genetic process ...
. He is considered the first to attempt to integrate genetics, development, and evolution. He pioneered understanding of reaction norms,
genetic assimilation Genetic assimilation is a process described by Conrad H. Waddington by which a phenotype originally produced in response to an environmental condition, such as exposure to a teratogen, later becomes genetically encoded via artificial selection ...
, dynamical genetics, sex determination, and
heterochrony In evolutionary developmental biology, heterochrony is any genetically controlled difference in the timing, rate, or duration of a Developmental biology, developmental process in an organism compared to its ancestors or other organisms. This lea ...
.Dietrich, Michael R. (2003)
Richard Goldschmidt: hopeful monsters and other 'heresies.'
''Nature Reviews Genetics'' 4 (Jan.): 68-74.
Controversially, Goldschmidt advanced a model of
macroevolution Macroevolution comprises the evolutionary processes and patterns which occur at and above the species level. In contrast, microevolution is evolution occurring within the population(s) of a single species. In other words, microevolution is the ...
through macromutations popularly known as the "Hopeful Monster" hypothesis. Goldschmidt also described the nervous system of the
nematode The nematodes ( or ; ; ), roundworms or eelworms constitute the phylum Nematoda. Species in the phylum inhabit a broad range of environments. Most species are free-living, feeding on microorganisms, but many are parasitic. Parasitic worms (h ...
, a piece of work that influenced
Sydney Brenner Sydney Brenner (13 January 1927 – 5 April 2019) was a South African biologist. In 2002, he shared the Nobel Prize in Physiology or Medicine with H. Robert Horvitz and Sir John E. Sulston. Brenner made significant contributions to wo ...
to study the "wiring diagram" of ''
Caenorhabditis elegans ''Caenorhabditis elegans'' () is a free-living transparent nematode about 1 mm in length that lives in temperate soil environments. It is the type species of its genus. The name is a Hybrid word, blend of the Greek ''caeno-'' (recent), ''r ...
'', winning Brenner and his colleagues the
Nobel Prize The Nobel Prizes ( ; ; ) are awards administered by the Nobel Foundation and granted in accordance with the principle of "for the greatest benefit to humankind". The prizes were first awarded in 1901, marking the fifth anniversary of Alfred N ...
in 2002.


Childhood and education

Goldschmidt was born in
Frankfurt-am-Main Frankfurt am Main () is the most populous city in the States of Germany, German state of Hesse. Its 773,068 inhabitants as of 2022 make it the List of cities in Germany by population, fifth-most populous city in Germany. Located in the forela ...
, Germany to upper-middle class parents of
Ashkenazi Jew Ashkenazi Jews ( ; also known as Ashkenazic Jews or Ashkenazim) form a distinct subgroup of the Jewish diaspora, that Ethnogenesis, emerged in the Holy Roman Empire around the end of the first millennium Common era, CE. They traditionally spe ...
ish heritage. He had a classical education and entered the
University of Heidelberg Heidelberg University, officially the Ruprecht Karl University of Heidelberg (; ), is a public university, public research university in Heidelberg, Baden-Württemberg, Germany. Founded in 1386 on instruction of Pope Urban VI, Heidelberg is List ...
in 1896, where he became interested in
natural history Natural history is a domain of inquiry involving organisms, including animals, fungi, and plants, in their natural environment, leaning more towards observational than experimental methods of study. A person who studies natural history is cal ...
. From 1899 Goldschmidt studied anatomy and zoology at the
University of Heidelberg Heidelberg University, officially the Ruprecht Karl University of Heidelberg (; ), is a public university, public research university in Heidelberg, Baden-Württemberg, Germany. Founded in 1386 on instruction of Pope Urban VI, Heidelberg is List ...
with Otto Bütschli and Carl Gegenbaur. He received his Ph.D. under Bütschli in 1902, studying development of the
trematode Trematoda is a Class (biology), class of flatworms known as trematodes, and commonly as flukes. They are obligate parasite, obligate Endoparasites, internal parasites with a complex biological life cycle, life cycle requiring at least two Host ( ...
''Polystomum''.


Career

In 1903 Goldschmidt began working as an assistant to Richard Hertwig at the
University of Munich The Ludwig Maximilian University of Munich (simply University of Munich, LMU or LMU Munich; ) is a public university, public research university in Munich, Bavaria, Germany. Originally established as the University of Ingolstadt in 1472 by Duke ...
, where he continued his work on nematodes and their
histology Histology, also known as microscopic anatomy or microanatomy, is the branch of biology that studies the microscopic anatomy of biological tissue (biology), tissues. Histology is the microscopic counterpart to gross anatomy, which looks at large ...
, including studies of the nervous system development of '' Ascaris'' and the anatomy of ''
Amphioxus The lancelets ( ), also known as amphioxi (: amphioxus ), consist of 32 described species of somewhat fish-like benthic filter feeding chordates in the subphylum Cephalochordata, class Leptocardii, and family Branchiostomatidae. Lancelets div ...
''. He founded the histology journal ''Archiv für Zellforschung'' while working in Hertwig's laboratory. Under Hertwig's influence, he also began to take an interest in
chromosome A chromosome is a package of DNA containing part or all of the genetic material of an organism. In most chromosomes, the very long thin DNA fibers are coated with nucleosome-forming packaging proteins; in eukaryotic cells, the most import ...
behavior and the new field of genetics. In 1909 Goldschmidt became professor at the
University of Munich The Ludwig Maximilian University of Munich (simply University of Munich, LMU or LMU Munich; ) is a public university, public research university in Munich, Bavaria, Germany. Originally established as the University of Ingolstadt in 1472 by Duke ...
and, inspired by
Wilhelm Johannsen Wilhelm Johannsen (3 February 1857 – 11 November 1927) was a Danish pharmacist, botanist, plant physiologist, and geneticist. He is best known for coining the terms gene, phenotype and genotype, and for his 1903 "pure line" experiments in ...
's genetics treatise ''Elemente der exakten Erblichkeitslehre'', began to study sex determination and other aspects of the genetics of
Lymantria dispar ''Lymantria dispar'', also known as the gypsy moth or the spongy moth, is a species of Lepidoptera, moth in the family Erebidae native to Europe and Asia. ''Lymantria dispar'' is subdivided into several subspecies, with subspecies such as ''Ly ...
, the gypsy moth, of which he was
crossbreed A crossbreed is an organism with purebred parents of two different breeds, varieties, or populations. A domestic animal of unknown ancestry, where the breed status of only one parent or grandparent is known, may also be called a crossbreed though ...
ing different races. He observed various stages of their sexual development, and found that some of the animals were neither
male Male (Planet symbols, symbol: ♂) is the sex of an organism that produces the gamete (sex cell) known as sperm, which fuses with the larger female gamete, or Egg cell, ovum, in the process of fertilisation. A male organism cannot sexual repro ...
, nor
female An organism's sex is female ( symbol: ♀) if it produces the ovum (egg cell), the type of gamete (sex cell) that fuses with the male gamete (sperm cell) during sexual reproduction. A female has larger gametes than a male. Females and ...
, nor
hermaphrodite A hermaphrodite () is a sexually reproducing organism that produces both male and female gametes. Animal species in which individuals are either male or female are gonochoric, which is the opposite of hermaphroditic. The individuals of many ...
s, but represented a whole spectrum of gynandromorphism. He named them '
intersex Intersex people are those born with any of several sex characteristics, including chromosome patterns, gonads, or genitals that, according to the Office of the United Nations High Commissioner for Human Rights, "do not fit typical binar ...
', and the phenomenon accordingly 'intersexuality' (''Intersexualität''). His studies of the gypsy moth, which culminated in his 1934 monograph ''Lymantria'', became the basis for his theory of sex determination, which he worked on from 1911 until 1931. Goldschmidt left Munich in 1914 for the position as head of the genetics section of the newly founded Kaiser Wilhelm Institute for Biology in Berlin. During a field trip to Japan in 1914, he was unable to return to Germany due to the outbreak of the
First World War World War I or the First World War (28 July 1914 – 11 November 1918), also known as the Great War, was a World war, global conflict between two coalitions: the Allies of World War I, Allies (or Entente) and the Central Powers. Fighting to ...
, and was detained as an
enemy alien In customary international law, an enemy alien is any alien native, citizen, denizen or subject of any foreign nation or government with which a domestic nation or government is in conflict and who is liable to be apprehended, restrained, secur ...
in the United States. He was placed in an
internment camp Internment is the imprisonment of people, commonly in large groups, without Criminal charge, charges or Indictment, intent to file charges. The term is especially used for the confinement "of enemy citizens in wartime or of terrorism suspects ...
in Fort Oglethorpe, Georgia for "dangerous Germans". 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 the
University of California, Berkeley The University of California, Berkeley (UC Berkeley, Berkeley, Cal, or California), is a Public university, public Land-grant university, land-grant research university in Berkeley, California, United States. Founded in 1868 and named after t ...
. During World War II, the Nazi party published a propaganda poster entitled "Jewish World Domination" displaying the Goldschmidt family tree.


Evolution

Goldschmidt was the first scientist to use the term " hopeful monster". He thought that small gradual changes could not bridge the divide between microevolution and macroevolution. In his book ''The Material Basis of Evolution'' (1940), he wrote Goldschmidt believed the large changes in
evolution Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
were caused by macromutations (large mutations). His ideas about macromutations became known as the "hopeful monster" hypothesis, a type of
saltational evolution Mutationism is one of several alternatives to evolution by natural selection that have existed both before and after the publication of Charles Darwin's 1859 book ''On the Origin of Species''. In the theory, mutation was the source of novelty, cr ...
, and attracted widespread ridicule. 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". Goldschmidt believed that the neo-Darwinian view of
gradual The gradual ( or ) is a certain chant or hymn in liturgical Christian worship. It is practiced in the Catholic Mass, Lutheran Divine Service, Anglican service and other traditions. It gets its name from the Latin (meaning "step") because i ...
accumulation of small
mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, ...
s was important but could account for variation only within species (
microevolution Microevolution is the change in allele frequencies that occurs over time within a population. This change is due to four different processes: mutation, selection ( natural and artificial), gene flow and genetic drift. This change happens over ...
) and was not a powerful enough source of evolutionary novelty to explain new
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
. Instead he believed that large genetic differences between species required profound "macro-mutations", a source for large genetic changes (
macroevolution Macroevolution comprises the evolutionary processes and patterns which occur at and above the species level. In contrast, microevolution is evolution occurring within the population(s) of a single species. In other words, microevolution is the ...
) which once in a while could occur as a "hopeful monster." 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 the synthetic theory only in his belief that a new species develops suddenly through discontinuous variation, or macromutation. Goldschmidt presented his hypothesis when
neo-Darwinism Neo-Darwinism is generally used to describe any integration of Charles Darwin's theory of evolution by natural selection with Gregor Mendel's theory of genetics. It mostly refers to evolutionary theory from either 1895 (for the combinations of D ...
was becoming dominant in the 1940s and 1950s, and strongly protested against the strict
gradualism Gradualism, from the Latin ("step"), is a hypothesis, a theory or a tenet assuming that change comes about gradually or that variation is gradual in nature and happens over time as opposed to in large steps. Uniformitarianism, incrementalism, and ...
of neo-Darwinian theorists. His ideas were accordingly seen as highly unorthodox by most scientists and were subjected to ridicule and scorn. However, there has been a recent interest in the ideas of Goldschmidt in the field of
evolutionary developmental biology Evolutionary developmental biology, informally known as evo-devo, is a field of biological research that compares the developmental biology, developmental processes of different organisms to infer how developmental processes evolution, evolved. ...
, as some scientists, such as Günter Theißen and Scott F. Gilbert, are convinced he was not entirely wrong. Goldschmidt presented two mechanisms by which hopeful monsters might work. One mechanism, involving "systemic mutations", rejected the classical
gene In biology, the word gene has two meanings. The Mendelian gene is a basic unit of heredity. The molecular gene is a sequence of nucleotides in DNA that is transcribed to produce a functional RNA. There are two types of molecular genes: protei ...
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.


Selected bibliography

* * Goldschmidt, R. B. (1923). ''The Mechanism and Physiology of Sex Determination'', Methuen & Co., London. (Translated by William Dakin) * * Goldschmidt, R. B. (1931). ''Die sexuellen Zwischenstufen'', Springer, Berlin. * * Goldschmidt, R. B. (1940). ''The Material Basis of Evolution'', New Haven CT: Yale Univ.Press. * * Goldschmidt, R. B. (1960) ''In and Out of the Ivory Tower'', Univ. of Washington Press, Seattle. * * * * * * * * * * * * * *


References


External links


Stephen Jay Gould on Richard Goldschmidt




by Edward Goldsmith
Guide to the Richard Goldschmidt Papers
at The Bancroft Library
National Academy of Sciences Biographical Memoir
{{DEFAULTSORT:Goldschmidt, Richard Benedikt 1878 births 1958 deaths Nematologists 20th-century German biologists American geneticists Emigrants from Nazi Germany to the United States German geneticists Internments in the United States Intersex healthcare Jewish American scientists Max Planck Institute directors Mutationism Scientists from Frankfurt