Richard Goldschmidt
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Richard Benedict Goldschmidt (April 12, 1878 – April 24, 1958) was a German-born American
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 processe ...
. He is considered the first to attempt to integrate genetics, development, and evolution. He pioneered understanding of reaction norms, genetic assimilation, 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 process in an organism compared to its ancestors or other organisms. This leads to changes in the ...
.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 usually means the evolution of large-scale structures and traits that go significantly beyond the intraspecific variation found in microevolution (including speciation). In other words, macroevolution is the evolution of taxa abov ...
through macromutations popularly known as the "Hopeful Monster" hypothesis. Goldschmidt also described the nervous system of the nematode, 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 work ...
to study the wiring diagram of ''Caenorhabditis elegans'', winning Brenner and his colleagues the
Nobel Prize The Nobel Prizes ( ; sv, Nobelpriset ; no, Nobelprisen ) are five separate prizes that, according to Alfred Nobel's will of 1895, are awarded to "those who, during the preceding year, have conferred the greatest benefit to humankind." Alfr ...
in 2002.


Childhood and education

Goldschmidt was born in Frankfurt-am-Main, Germany to upper-middle class parents of
Ashkenazi Jew Ashkenazi Jews ( ; he, יְהוּדֵי אַשְׁכְּנַז, translit=Yehudei Ashkenaz, ; yi, אַשכּנזישע ייִדן, Ashkenazishe Yidn), also known as Ashkenazic Jews or ''Ashkenazim'',, Ashkenazi Hebrew pronunciation: , singu ...
ish heritage. He had a
classical education Classical education may refer to: *''Modern'', educational practices and educational movements: **An education in the Classics, especially in Ancient Greek and Latin **Classical education movement, based on the trivium (grammar, logic, rhetoric) an ...
and entered the
University of Heidelberg } Heidelberg University, officially the Ruprecht Karl University of Heidelberg, (german: Ruprecht-Karls-Universität Heidelberg; la, Universitas Ruperto Carola Heidelbergensis) is a public research university in Heidelberg, Baden-Württemberg, ...
in 1896, where he became interested in natural history. From 1899 Goldschmidt studied anatomy and zoology at the
University of Heidelberg } Heidelberg University, officially the Ruprecht Karl University of Heidelberg, (german: Ruprecht-Karls-Universität Heidelberg; la, Universitas Ruperto Carola Heidelbergensis) is a public research university in Heidelberg, Baden-Württemberg, ...
with
Otto Bütschli Johann Adam Otto Bütschli (3 May 1848 – 2 February 1920) was a German zoologist and professor at the University of Heidelberg. He specialized in invertebrates and insect development. Many of the groups of protists were first recognized by him. ...
and Carl Gegenbaur. He received his Ph.D. under Bütschli in 1902, studying development of the
trematode Trematoda is a class of flatworms known as flukes. They are obligate internal parasites with a complex life cycle requiring at least two hosts. The intermediate host, in which asexual reproduction occurs, is usually a snail. The definitive host ...
''Polystomum''.


Career

In 1903 Goldschmidt began working as an assistant to
Richard Hertwig Richard Wilhelm Karl Theodor Ritter von Hertwig (23 September 1850 in Friedberg, Hesse – 3 October 1937 in Schlederloh, Bavaria), also Richard Hertwig or Richard von Hertwig, was a German zoologist and professor of 50 years, notable as the fir ...
at the
University of Munich The Ludwig Maximilian University of Munich (simply University of Munich or LMU; german: Ludwig-Maximilians-Universität München) is a public research university in Munich, Germany. It is Germany's List of universities in Germany, sixth-oldest u ...
, where he continued his work on nematodes and their
histology Histology, also known as microscopic anatomy or microanatomy, is the branch of biology which studies the microscopic anatomy of biological tissues. Histology is the microscopic counterpart to gross anatomy, which looks at larger structures vis ...
, including studies of the nervous system development of ''
Ascaris ''Ascaris'' is a genus of parasite, parasitic nematode worms known as the "small intestinal roundworms", which is a type of parasitic worm. One species, ''Ascaris lumbricoides'', affects humans and causes the disease ascariasis. Another species ...
'' and the anatomy of ''
Amphioxus The lancelets ( or ), also known as amphioxi (singular: amphioxus ), consist of some 30 to 35 species of "fish-like" benthic filter feeding chordates in the order Amphioxiformes. They are the modern representatives of the subphylum Cephalochorda ...
''. 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 long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
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 or LMU; german: Ludwig-Maximilians-Universität München) is a public research university in Munich, Germany. It is Germany's List of universities in Germany, sixth-oldest u ...
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 g ...
's genetics treatise ''Elemente der exakten Erblichkeitslehre'', began to study sex determination and other aspects of the genetics of the
gypsy moth ''Lymantria dispar'', also known as the gypsy moth or the spongy moth, is a species of moth in the family Erebidae. ''Lymantria dispar'' is subdivided into several subspecies, with subspecies such as ''L. d. dispar'' and ''L. d. japonica'' bei ...
of which he was
crossbreed A crossbreed is an organism with purebred parents of two different breeds, varieties, or populations. ''Crossbreeding'', sometimes called "designer crossbreeding", is the process of breeding such an organism, While crossbreeding is used to main ...
ing different races. He observed different stages of their sexual development.  Some of the animals were neither
male Male (symbol: ♂) is the sex of an organism that produces the gamete (sex cell) known as sperm, which fuses with the larger female gamete, or ovum, in the process of fertilization. A male organism cannot reproduce sexually without access to ...
, nor
female Female (Venus symbol, symbol: ♀) is the sex of an organism that produces the large non-motile ovum, ova (egg cells), the type of gamete (sex cell) that fuses with the Sperm, male gamete during sexual reproduction. A female has larger gamet ...
, nor
hermaphrodite In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many Taxonomy (biology), taxonomic groups of animals (mostly invertebrate ...
s, but represented a whole spectrum of
gynandromorphism A gynandromorph is an organism that contains both male and female characteristics. The term comes from the Greek γυνή (''gynē'') 'female', ἀνήρ (''anēr'') 'male', and μορφή (''morphē'') 'form', and is used mainly in the field ...
. He named them '
intersex Intersex people are individuals 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 bina ...
', 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 developed 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. During a field trip to Japan in 1914 he was not able to return to Germany due to the outbreak of the First World War and got stranded in the United States. He ended up in an internment camp in Fort Oglethorpe, Georgia for "dangerous Germans". After his release in 1918 he returned to Germany in 1919 and worked at the Kaiser Wilhelm Institute. Sensing that it was unsafe for him to remain in Germany he emigrated to the United States in 1936, where he became professor at the
University of California, Berkeley The University of California, Berkeley (UC Berkeley, Berkeley, Cal, or California) is a public land-grant research university in Berkeley, California. Established in 1868 as the University of California, it is the state's first land-grant u ...
. During World War 2, 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 "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 in
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
were caused by macromutations (large mutations). His ideas about macromutations became known as the hopeful monster hypothesis, a type of saltational evolution, 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 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 Dar ...
view of
gradual The gradual ( la, graduale or ) is a chant or hymn in the Mass, the liturgical celebration of the Eucharist in the Catholic Church, and among some other Christians. It gets its name from the Latin (meaning "step") because it was once chanted ...
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, mi ...
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 a r ...
) and was not a powerful enough source of evolutionary novelty to explain new
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
. Instead he believed that large genetic differences between species required profound "macro-mutations", a source for large genetic changes (
macroevolution Macroevolution usually means the evolution of large-scale structures and traits that go significantly beyond the intraspecific variation found in microevolution (including speciation). In other words, macroevolution is the evolution of taxa abov ...
) 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 Dar ...
was becoming dominant in the 1940s and 1950s, and strongly protested against the strict
gradualism Gradualism, from the Latin ''gradus'' ("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, incrementa ...
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, as some scientists, such as
Günter Theißen Günter Theißen (born 16 January 1962 in Mönchengladbach) is a German geneticist. He holds the chair of genetics at the University of Jena. Life Günter Theißen studied biology at the University of Düsseldorf and did his PhD in 1991 wit ...
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 (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
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. * * Goldschmitdt, 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 Edward René David Goldsmith (8 November 1928 – 21 August 2009), widely known as Teddy Goldsmith, was an Anglo-French environmentalist, writer and philosopher. He was a member the prominent Goldsmith family. The eldest son of Major Fr ...

Guide to the Richard Goldschmidt Papers
at
The Bancroft Library The Bancroft Library in the center of the campus of the University of California, Berkeley, is the university's primary special-collections library. It was acquired from its founder, Hubert Howe Bancroft, in 1905, with the proviso that it retai ...

National Academy of Sciences Biographical Memoir
{{DEFAULTSORT:Goldschmidt, Richard Benedikt 1878 births Scientists from Frankfurt German emigrants to the United States 1958 deaths 20th-century German biologists Jewish American scientists American geneticists German geneticists Intersex and medicine Mutationism Internments in the United States Max Planck Institute directors