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The Material Basis of Evolution
In his autobiography, published posthumously in 1960, Richard Goldschmidt “I am confident that in twenty years my [work], which is now ignored, will be given an honorable place in the history of evolutionary thought.” The publication of this edition, now reissued with a new introduction by Stephen Jay Gould, proves Goldschmidt’s prediction to be correct.
Goldschmidt, one of the world’s great geneticists, delivered the prestigious Silliman lectures at Yale University in 1939 and published his remarks in 1940 as The Material Basis of Evolution . His intent was to inquire into the types of hereditary differences that produce new species. Goldschmidt used a wide range of research to formulate his own picture of evolution. Contrary to most scientists, he insisted that the neo-Darwinist theory of micromutations was no longer tenable as a general theory of evolution. Instead, Goldschmidt claimed, macroevolution accounted for the larger steps in evolution.
Although Goldschmidt’s views were reviled by scientists of his day, some of his basic ideas are now gaining acceptance. As Gould writes in his “I do…believe that its general vision is uncannily correct (or at least highly fruitful at the moment) in several important areas where conventional Darwinian theory has become both hidebound and unproductive.”
Goldschmidt, one of the world’s great geneticists, delivered the prestigious Silliman lectures at Yale University in 1939 and published his remarks in 1940 as The Material Basis of Evolution . His intent was to inquire into the types of hereditary differences that produce new species. Goldschmidt used a wide range of research to formulate his own picture of evolution. Contrary to most scientists, he insisted that the neo-Darwinist theory of micromutations was no longer tenable as a general theory of evolution. Instead, Goldschmidt claimed, macroevolution accounted for the larger steps in evolution.
Although Goldschmidt’s views were reviled by scientists of his day, some of his basic ideas are now gaining acceptance. As Gould writes in his “I do…believe that its general vision is uncannily correct (or at least highly fruitful at the moment) in several important areas where conventional Darwinian theory has become both hidebound and unproductive.”
438 pages, Paperback
First published September 10, 1940
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About the author
Richard Goldschmidt
96 books4 followersRichard Benedict Goldschmidt (April 12, 1878 – April 24, 1958) was a German-born American geneticist. He is considered the first to integrate genetics, development, and evolution. He pioneered understanding of reaction norms, genetic assimilation, dynamical genetics, sex determination, and heterochrony. Controversially, Goldschmidt advanced a model of macroevolution through macromutations that is popularly known as the "Hopeful Monster" hypothesis.
Goldschmidt also described the nervous system of the nematode, a piece of work that later influenced Sydney Brenner to study the wiring diagram of Caenorhabditis elegans an achievement that later won Brenner and his colleagues the Nobel Prize in 2002.
Goldschmidt was born in Frankfurt-am-Main, Germany to upper-middle class parents. He had a classical education and entered the University of Heidelberg in 1896, where he became interested in natural history. From 1899 Goldschmidt studied anatomy and zoology at the University of Heidelberg with Otto Bütschli and Carl Gegenbaur. He received his Ph.D. under Bütschli in 1902, studying development of the trematode Polystomum.
In 1903 Goldschmidt began working as an assistant to Richard Hertwig at the University of Munich, where he continued his work on nematodes and their histology, including studies of the nervous system development of Ascaris and the anatomy of Amphioxus. 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 behavior and the new field of genetics.
In 1909 Goldschmidt became professor at the University of Munich and, inspired by Wilhelm Johannsen's genetics treatise Elemente der exakten Erblichkeitslehre, began to study sex determination and other aspects of the genetics of the gypsy moth. 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 for "dangerous Germans". After his release in 1918 he returned to Germany in 1919. Because he was Jewish he had to leave Germany in 1935 and emigrated to the United States, where he became professor at the University of California, Berkeley.
Goldschmidt also described the nervous system of the nematode, a piece of work that later influenced Sydney Brenner to study the wiring diagram of Caenorhabditis elegans an achievement that later won Brenner and his colleagues the Nobel Prize in 2002.
Goldschmidt was born in Frankfurt-am-Main, Germany to upper-middle class parents. He had a classical education and entered the University of Heidelberg in 1896, where he became interested in natural history. From 1899 Goldschmidt studied anatomy and zoology at the University of Heidelberg with Otto Bütschli and Carl Gegenbaur. He received his Ph.D. under Bütschli in 1902, studying development of the trematode Polystomum.
In 1903 Goldschmidt began working as an assistant to Richard Hertwig at the University of Munich, where he continued his work on nematodes and their histology, including studies of the nervous system development of Ascaris and the anatomy of Amphioxus. 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 behavior and the new field of genetics.
In 1909 Goldschmidt became professor at the University of Munich and, inspired by Wilhelm Johannsen's genetics treatise Elemente der exakten Erblichkeitslehre, began to study sex determination and other aspects of the genetics of the gypsy moth. 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 for "dangerous Germans". After his release in 1918 he returned to Germany in 1919. Because he was Jewish he had to leave Germany in 1935 and emigrated to the United States, where he became professor at the University of California, Berkeley.
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January 12, 2025
THE CONTROVERSIAL BOOK WHICH FIRST PROPOSED THE ‘HOPEFUL MONSTER’
Stephen Jay Gould wrote in his Introduction to this1940 book, “Richard Goldschmidt, one of the world’s great geneticists… wrote just as the Darwinian paradigm was coalescing into a confident general theory of evolution… Among this incipient chorus of consensus… Goldschmidt’s views injected an especially disharmonious note. He seemed bent on undoing this unity… to assert once again that macroevolution was a thing apart, abrupt in its occurrence, and unilluminated in principle by processes of microevolution that could be studied directly. He even invented an unforgettable and disturbing (if whimsical) name for his independent macroevolutionary agent---the ‘hopeful monster.’ … Goldschmidt claimed, ‘…The Neo-Darwinians reacted savagely.’ … The counterattack was successful… though Goldschmidt had largely himself to blame for burying his gems so deeply in unacceptable the overextended claims. Indeed, he suffered the worst fate of all: to be ridiculed AND unread.” (Pg. xiii-xiv)
Richard Goldschmidt (1878-1958) wrote in the opening chapter, “[I] challenge the adherents of the strictly Darwinian view… to try to explain the evolution of the following features by accumulation and selection of small mutants: hair in mammals, feathers in birds… the transformation of the gill arches in phylogeny… teeth, shells of mollusks… compound eyes, poison apparatus of snakes… and, finally, primary chemical differences like hemoglobin vs. hemocyanin, etc. Corresponding examples from plants could be given.” (Pg. 6-7)
He suggests, “there are two types of variation: the more or less haphazard… recombinations of the polymorphic Mendelian characters … and the orderly cline of different forms in a definitive direction for which isolation cannot account… Isolation… had not led to any larger deviations than the typical subspecific ones, linked by all intermediates, and the material in question does not help us to understand diversification beyond the limits of the species.” (Pg. 135)
He asserts, “Subspecies are actually… neither incipient species nor models for the origin of species. They are more or less diversified blind alleys within the species. The decisive step in evolution, the first step toward macroevolution, the step from one species to another, requires another evolutionary method than that of sheer accumulation of micromutations.” (Pg. 183)
He explains, “The facts … thus far have led… the conclusion that we must look for a genetic agency able to accomplish separation of species without slow accumulation of gene mutations… We must, therefore, find out whether a method of macroevolution has been discovered here which does not need the slow accumulation of minute steps by gene mutation… The discoveries just discussed… have led me to believe that a pattern change in the chromosomes, completely independent of gene mutations… will furnish this new method of macroevolution… One might call this different type of genetic change a ‘systemic mutation,’ though this does not have to occur in one step… These steps may be without visible effect until the patterning of the chromosomes… leads to a new stable pattern… a new chemical system. This… is so basically changed that a new phenotype emerges… separated from the old one by a bridgeless gap and an incompatible intrachromosomal pattern. ‘Emergent evolution’ but without mysticism!” (Pg. 205-206)
He outlines, “A conception of macroevolution by systemic mutations… requires a number of basic assumptions… (a) Macroevolution cannot be conceived on the basis of accumulation of micromutations… (b) Macroevolution is accompanied by repatterning of the chromosomes…. (c) an intrachromosomal pattern change may exert a considerable phenotypic effect independent of genic changes… (d) Such a thing exists as a complete change of the reaction system based upon a genetic change different from an accumulation of micromutants… (e) It is possible to produce immense phenotypic changes of a macroevolutionary order by relatively small systemic mutations not involving the creation of anything new within the germ plasm. (f) The classical atomistic theory of the gene… blocks progress in evolutionary thought… (g) Models are available which make it possible to visualize the systemic effect of pure pattern changes in the germ plasm.” (Pg. 209-210)
He contends, “I am firmly convinced that, except in microevolution, the facts already available today force us to drop completely from evolutionary thought the idea of the so-called gene mutation, whatever it turns out to be physically or chemically… we have concluded that the really decisive change of the genetic material involved in macroevolution is the change of chromosomal pattern. The problem thus left is to find out … whether this type of genetic change is able to produce the effects required.” (Pg. 242-243)
He summarizes, “I have come to the conclusion that all the recent developments of genetics tend to show that the classical theory of the gene … is no longer tenable; that the linear order of the loci in a chromosome is an internal pattern…which does not necessarily involve the existence of separate units of a molecular order; that the mutational change at a definite locus… does not prove that a particle is located at this point, a break, for example, being also able to account for the changed action. In spite of all the opposition which such a viewpoint is bound to encounter, I am fully convinced that geneticists will have to accept it eventually.” (Pg. 247)
He proposes, “The actually existing series of large anatomical differences between taxonomic groups does therefore not require an evolution by simultaneous selection of numerous small mutants … A single mutational step affecting the right process at the right moment can accomplish everything, providing that it is able to set in motion the every-present potentialities of embryonic regulation… this statement also contains the explanation of atavism as well as of the positive and negative features of embryonic recapitulation.” (Pg. 297)
He expresses doubts about the neo-Darwinian explanation in some cases, “as it always meets with the old difficulty that the original mutant would be wiped out before it could become established… This difficulty would, however, disappear in … our systemic mutation where the trial and error take place within the chromosome without external effect until the new pattern emerges which… controls the emergence of a complete… integrated change of morphogenesis. Unfortunately, no experimental attack upon this problem is at present apparent…” (Pg. 334) He gives an example: “the macroevolutionary wing rudimentation in Carabid beetles does not necessitate the assumption of a gradual accumulation of macromutations, but can be explained by single steps, which change the relative rates of processes of growth and differentiation at an earlier or later time in development.” (Pg. 361)
Later, he adds, “the time of stoppage or slowing down of differentiation in development is connected with the degree of rudimentation in the simple relation… A mutant causing these changes in early development, therefore, will produce the maximum effect in one step. We may recall the fact that the rudimentary eye of the blind newt … may develop into a normal eye, just as the wingless earwig…occasionally grows wings.” (Pg. 379) He asks of the neo-Darwinian interpretation, “I wonder if anybody could ever succeed in explaining the phylogeny of the auditory ossicles in this way…” (Pg. 386)
He continues, “If macroevolutionary changes proceed by mutations affecting the rate of embryogenetic processes at a definite time in development, the ontogeny of all descendants of the mutant form must continue along ancestral lines up the stage … first affected by the mutant… Obviously, the mechanics of development do not permit any other course. If the mutation which changed the long tail of the Archaeopteryx… into the rudimentary tail of birds with fanlike tail feathers… the present embryology of birds must necessarily contain an Archaeopteryx stage, which is actually the case.” (Pg. 389)
He proposes, “In a former paper … I used the term ‘hopeful monster’ to express the idea that mutants producing monstrosities may have played a considerable role in macroevolution. A monstrosity appearing in a single genetic step might permit the occupation of a new environmental niche and thus produce a new type in one step… Actually, the idea expressed in the … plastic term ‘hopeful monster’ is not a new one… Darwin … pointed out that under domestication monstrosities occur which resemble normal structures in widely different animals…” (Pg. 390-391) He adds, “Thus, the combined facts of genetics, embryology, and taxonomy demonstrate that the hopeful monster is one of the means of macroevolution by single large steps.” (Pg. 393)
He notes that Otto H. Schindewolf, “the most progressive investigator known to me… showed that the material presented by paleontology leads to exactly the same conclusions as derived in my writings, to which he refers… He elaborates the thesis that macroevolution on a higher level takes place in an explosive way within a short geological time, followed by a slower series of orthogenetic perfections. He realizes that the conception of preadaptation accounts completely for this type of evolution. He shows by examples from fossil material that the major evolutionary advances must have taken place in single large steps… He shows that many missing links in the paleontological record are sought for in vain because they never existed. ‘The first bird hatched from a reptilian egg.’ … It is gratifying that all the disciplines which furnish material for the understanding of evolution… supply ample and parallel evidence for a theory of evolution which is more plausible than the neo-Darwinian theory.” (Pg. 395)
Always highly controversial, Goldschmidt’s book will nevertheless be ‘must reading’ for anyone seriously studying evolutionary theory.
Stephen Jay Gould wrote in his Introduction to this1940 book, “Richard Goldschmidt, one of the world’s great geneticists… wrote just as the Darwinian paradigm was coalescing into a confident general theory of evolution… Among this incipient chorus of consensus… Goldschmidt’s views injected an especially disharmonious note. He seemed bent on undoing this unity… to assert once again that macroevolution was a thing apart, abrupt in its occurrence, and unilluminated in principle by processes of microevolution that could be studied directly. He even invented an unforgettable and disturbing (if whimsical) name for his independent macroevolutionary agent---the ‘hopeful monster.’ … Goldschmidt claimed, ‘…The Neo-Darwinians reacted savagely.’ … The counterattack was successful… though Goldschmidt had largely himself to blame for burying his gems so deeply in unacceptable the overextended claims. Indeed, he suffered the worst fate of all: to be ridiculed AND unread.” (Pg. xiii-xiv)
Richard Goldschmidt (1878-1958) wrote in the opening chapter, “[I] challenge the adherents of the strictly Darwinian view… to try to explain the evolution of the following features by accumulation and selection of small mutants: hair in mammals, feathers in birds… the transformation of the gill arches in phylogeny… teeth, shells of mollusks… compound eyes, poison apparatus of snakes… and, finally, primary chemical differences like hemoglobin vs. hemocyanin, etc. Corresponding examples from plants could be given.” (Pg. 6-7)
He suggests, “there are two types of variation: the more or less haphazard… recombinations of the polymorphic Mendelian characters … and the orderly cline of different forms in a definitive direction for which isolation cannot account… Isolation… had not led to any larger deviations than the typical subspecific ones, linked by all intermediates, and the material in question does not help us to understand diversification beyond the limits of the species.” (Pg. 135)
He asserts, “Subspecies are actually… neither incipient species nor models for the origin of species. They are more or less diversified blind alleys within the species. The decisive step in evolution, the first step toward macroevolution, the step from one species to another, requires another evolutionary method than that of sheer accumulation of micromutations.” (Pg. 183)
He explains, “The facts … thus far have led… the conclusion that we must look for a genetic agency able to accomplish separation of species without slow accumulation of gene mutations… We must, therefore, find out whether a method of macroevolution has been discovered here which does not need the slow accumulation of minute steps by gene mutation… The discoveries just discussed… have led me to believe that a pattern change in the chromosomes, completely independent of gene mutations… will furnish this new method of macroevolution… One might call this different type of genetic change a ‘systemic mutation,’ though this does not have to occur in one step… These steps may be without visible effect until the patterning of the chromosomes… leads to a new stable pattern… a new chemical system. This… is so basically changed that a new phenotype emerges… separated from the old one by a bridgeless gap and an incompatible intrachromosomal pattern. ‘Emergent evolution’ but without mysticism!” (Pg. 205-206)
He outlines, “A conception of macroevolution by systemic mutations… requires a number of basic assumptions… (a) Macroevolution cannot be conceived on the basis of accumulation of micromutations… (b) Macroevolution is accompanied by repatterning of the chromosomes…. (c) an intrachromosomal pattern change may exert a considerable phenotypic effect independent of genic changes… (d) Such a thing exists as a complete change of the reaction system based upon a genetic change different from an accumulation of micromutants… (e) It is possible to produce immense phenotypic changes of a macroevolutionary order by relatively small systemic mutations not involving the creation of anything new within the germ plasm. (f) The classical atomistic theory of the gene… blocks progress in evolutionary thought… (g) Models are available which make it possible to visualize the systemic effect of pure pattern changes in the germ plasm.” (Pg. 209-210)
He contends, “I am firmly convinced that, except in microevolution, the facts already available today force us to drop completely from evolutionary thought the idea of the so-called gene mutation, whatever it turns out to be physically or chemically… we have concluded that the really decisive change of the genetic material involved in macroevolution is the change of chromosomal pattern. The problem thus left is to find out … whether this type of genetic change is able to produce the effects required.” (Pg. 242-243)
He summarizes, “I have come to the conclusion that all the recent developments of genetics tend to show that the classical theory of the gene … is no longer tenable; that the linear order of the loci in a chromosome is an internal pattern…which does not necessarily involve the existence of separate units of a molecular order; that the mutational change at a definite locus… does not prove that a particle is located at this point, a break, for example, being also able to account for the changed action. In spite of all the opposition which such a viewpoint is bound to encounter, I am fully convinced that geneticists will have to accept it eventually.” (Pg. 247)
He proposes, “The actually existing series of large anatomical differences between taxonomic groups does therefore not require an evolution by simultaneous selection of numerous small mutants … A single mutational step affecting the right process at the right moment can accomplish everything, providing that it is able to set in motion the every-present potentialities of embryonic regulation… this statement also contains the explanation of atavism as well as of the positive and negative features of embryonic recapitulation.” (Pg. 297)
He expresses doubts about the neo-Darwinian explanation in some cases, “as it always meets with the old difficulty that the original mutant would be wiped out before it could become established… This difficulty would, however, disappear in … our systemic mutation where the trial and error take place within the chromosome without external effect until the new pattern emerges which… controls the emergence of a complete… integrated change of morphogenesis. Unfortunately, no experimental attack upon this problem is at present apparent…” (Pg. 334) He gives an example: “the macroevolutionary wing rudimentation in Carabid beetles does not necessitate the assumption of a gradual accumulation of macromutations, but can be explained by single steps, which change the relative rates of processes of growth and differentiation at an earlier or later time in development.” (Pg. 361)
Later, he adds, “the time of stoppage or slowing down of differentiation in development is connected with the degree of rudimentation in the simple relation… A mutant causing these changes in early development, therefore, will produce the maximum effect in one step. We may recall the fact that the rudimentary eye of the blind newt … may develop into a normal eye, just as the wingless earwig…occasionally grows wings.” (Pg. 379) He asks of the neo-Darwinian interpretation, “I wonder if anybody could ever succeed in explaining the phylogeny of the auditory ossicles in this way…” (Pg. 386)
He continues, “If macroevolutionary changes proceed by mutations affecting the rate of embryogenetic processes at a definite time in development, the ontogeny of all descendants of the mutant form must continue along ancestral lines up the stage … first affected by the mutant… Obviously, the mechanics of development do not permit any other course. If the mutation which changed the long tail of the Archaeopteryx… into the rudimentary tail of birds with fanlike tail feathers… the present embryology of birds must necessarily contain an Archaeopteryx stage, which is actually the case.” (Pg. 389)
He proposes, “In a former paper … I used the term ‘hopeful monster’ to express the idea that mutants producing monstrosities may have played a considerable role in macroevolution. A monstrosity appearing in a single genetic step might permit the occupation of a new environmental niche and thus produce a new type in one step… Actually, the idea expressed in the … plastic term ‘hopeful monster’ is not a new one… Darwin … pointed out that under domestication monstrosities occur which resemble normal structures in widely different animals…” (Pg. 390-391) He adds, “Thus, the combined facts of genetics, embryology, and taxonomy demonstrate that the hopeful monster is one of the means of macroevolution by single large steps.” (Pg. 393)
He notes that Otto H. Schindewolf, “the most progressive investigator known to me… showed that the material presented by paleontology leads to exactly the same conclusions as derived in my writings, to which he refers… He elaborates the thesis that macroevolution on a higher level takes place in an explosive way within a short geological time, followed by a slower series of orthogenetic perfections. He realizes that the conception of preadaptation accounts completely for this type of evolution. He shows by examples from fossil material that the major evolutionary advances must have taken place in single large steps… He shows that many missing links in the paleontological record are sought for in vain because they never existed. ‘The first bird hatched from a reptilian egg.’ … It is gratifying that all the disciplines which furnish material for the understanding of evolution… supply ample and parallel evidence for a theory of evolution which is more plausible than the neo-Darwinian theory.” (Pg. 395)
Always highly controversial, Goldschmidt’s book will nevertheless be ‘must reading’ for anyone seriously studying evolutionary theory.
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