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 processes ...
. 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 Jewish 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 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 h ...
''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 sixth-oldest university in continuous operatio ...
, 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 vi ...
, including studies of the nervous system development of ''
Ascaris
''Ascaris'' is a genus of 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, ''Asc ...
'' 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 sixth-oldest university in continuous operatio ...
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 mai ...
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 ( symbol: ♀) is the sex of an organism that produces the large non-motile ova (egg cells), the type of gamete (sex cell) that fuses with the male gamete during sexual reproduction.
A female has larger gametes than a male. Females ...
, nor
hermaphrodites, 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 bin ...
', 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 replication, DNA or viral repl ...
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 ...
) 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 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 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 b ...
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.
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References
External links
Stephen Jay Gould on Richard Goldschmidt by
Edward Goldsmith
Guide to the Richard Goldschmidt Papersat
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