An overview of the life and work of the eminent German-American geneticist, Richard Goldschmidt, and the angry reaction he received from the defenders of Orthodox Darwinism as a result of the challenging theory of evolution expounded in his magnum opus The Material Basis of Evolution.
Richard Benedict Goldschmidt was born in Frankfurt in 1878 and studied classical morphology and embryology in Heidelberg and Munich, where he wrote his doctoral thesis on maturation, fertilisation, and embryonic development in the trematode – a parasitic flatworm – (polystomum integerrimum). In 1903 he became assistant to Professor R. Hertwig in Munich, where he remained until 1913, when he accepted an invitation to become Director of Genetics at the Kaiser Wilhelm Institute for Biology in Berlin.
He was teaching a course in the USA when the First World War broke out and ended up in an internment camp for enemy aliens. When the war came to an end he was repatriated to Germany, and resumed his position at the Kaiser Wilhelm Institute.
In the 1930′s his career was again interrupted by the rise of Nazism. He was a proud man and when confronted by a Nazi poster which described the genealogy of his family as testimony to the Jewish danger, he reacted by suggesting that “it could well be used as a chart demonstrating the effect of long selection of favourable hereditary traits upon the improvement of human families”. However, he was forced to relinquish his post in 1935 and returned to the USA, where, in 1936, he became a professor at Berkeley – a post he held until his retirement.
During the course of his career he wrote a number of books and a large number of articles in professional journals. However, he is best known for his seminal work The Material Basis of Evolution, that was first published in 1940 (Yale University Press) and was republished in 1982 with a long introduction by Professor Stephen Jay Gould, from which much of the information in this chapter is derived.
His work, especially after the publication of The Material Basis of Evolution, became increasingly unorthodox. The theory of evolution he proposed in that book was completely at odds with neo-Darwinism, also with its latest variant, ‘the modern synthesis’, that had just been formulated and was already gospel in the biological community of that period – as indeed it still is today.
The neo-Darwinians reacted savagely. His book was mercilessly ridiculed by those who read it, and even by those who did not bother to do so. The biologist Professor Frazzetta recalled that when he was a student, Goldschmidt was
always introduced as a kind of biological ‘in’ joke, and all we students laughed and sniggered dutifully to prove that we were not guilty of either ignorance or heresy.
In Goldschmidt’s own words “he had struck a hornets’ nest” and was seen to be “not only crazy but also criminal”. In spite of all the criticism of his work, Goldschmidt’s confidence in his theories remained unshaken, and in his biography he assures us that
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in twenty years time this book [The Material Basis of Evolution] which is now ignored, will be given an honourable place in the history of evolutionary thought.
Let us see exactly what was Richard Goldschmidt’s thesis, and why it was so controversial. To begin with, for Darwinians and neo-Darwinians alike, microevolution – the small changes that occur in populations isolated from each other, or in domestic animals bred by farmers to be more productive – is taken to be the model for macroevolution, that is, for the bigger changes that lead to the development of new species (speciation).
In other words, intraspecies change is taken to be the model for interspecies change. In the words of the famous neo-Darwinian biologist, Ernst Mayr,
adaptations to local conditions and evolutionary change are two aspects of the same genetic phenomenon, the continuous adjustment of an integrated gene complex to a changing environment.
This is totally consistent with the Darwinian and neo-Darwinian view that evolution proceeds by an accumulation of small and gradual changes.
Goldschmidt denied this. For him, microevolution and macroevolution were totally different processes. The former occurred as a result of small, gradual, changes, which, from the evolutionary point of view, led to ‘blind alleys’ or ‘dead ends’. In no case could it result in the development of a new species, for he saw species as separated from each other by what he referred to as a ‘bridgeless gap’. As he put it,
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. [The Material Basis of Evolution, pp.205/206.]
Such changes, he saw as occurring suddenly and as the result of a massive change or ‘macromutation’. This occurred when a gene – whose role he saw mainly as controlling the rate of chemical processes – mutated in response to an environmental stress, very early on in development, and thereby affecting the timing of the crucial early stages of this process. This, he saw as capable of engendering a veritable cascade of other effects, producing a large phenotypic (or physical) jump “in a single genetic step”.
However, even more heretical was his view that these abrupt macroevolutionary changes need not occur as a result of a gene mutation at all. Phenotypic changes, identical to those produced by genetic mutations, could occur in response to serious environmental stress. He referred to these as “phenocopies”. Later he came to the conclusion that these abrupt macroevolutionary changes were the result of a complete repatterning or “reshuffling” of the chromosomal architecture, which could have
a huge effect upon a series of developmental processes, leading at once to a new and stable form by diverging from the former.
In other words, he considered that changes acquired during the course of the life of living things – could be fixed genetically, thereby stating – in a perhaps more acceptable manner – Lamarck’s heretical thesis that acquired characteristics could be transmitted genetically.
Perhaps even more heretical was his view that these “systemic mutations”, as he referred to them, “rarely occurred by chance”. In other words, they were directive rather than random – again a notion that could not possibly be reconciled with the neo-Darwinian thesis. Later Goldschmidt went even so far as to deny the existence of what he referred to as the “corpuscular gene”, and came to view all genetic changes as alterations in pattern.
Goldschmidt referred to the product of these systemic mutations as “hopeful monsters”. If they were hopeful, i.e. if they could lead to viable and adaptive forms rather than “hopeless”, by leading to unadaptive monsters, it is that such changes were canalised or under phenotypic control – which means that, in a particular species, only certain types of changes could occur. To quote Goldschmidt,
a change in the hereditary type can only occur within the possibilities and limitations set by the normal process of control on development.
He continued with his work long after his retirement, and tells us in his autobiography that
it was his greatest intellectual happiness that he could still work in his laboratory and even make interesting discoveries in the field of chemically induced phenocopies.
At the same time he worked on his autobiography [In and Out of the Ivory Tower: the autobiography of Richard B. Goldschmidt, University of Washington Press, Seattle], which was published in 1960, two years after his death. He had many other interests as well. Stephen Jay Gould tells us that he was an erudite and highly cultivated man, and was, among other things, an expert on oriental art.Back to top
How then can we evaluate, 40 years later, Richard Goldschmidt’s work? To begin with one must realize that even though his ideas were at the time laughed at and generally derided, no one questioned his eminence in the fields of genetics, developmental biology and evolutionary biology.
Nor did all the members of his peer group reject his ideas. For instance, the great British geneticist and embryologist C. H. Waddington, himself a critic of neo-Darwinism as far back as 1952, described The Material Basis of Evolution as “one of the most important of recent contributions to the theory of evolution.”
Stephen Jay Gould describes him today as “one of the premier geneticists of our century”. He also notes that in the last ten years, many of the basic assumptions underlying the neo-Darwinian thesis have been seriously questioned and as a result there has been “a strong reawakening of interest in Goldschmidt’s views among evolutionary biologists.”Back to top
One of these basic assumptions is the denial of any feedback between behaviour and evolution – the two processes being seen by neo-Darwinians as independent of each other. As a result, evolutionary changes are not seen by neo-Darwinians as under phenotypic control, and any changes are possible if they are adaptive. As Gould notes,
if evolution is gradual and continuous, moving in any direction dictated by selective pressures of a changing environment, then developmental constraints play a little role, except as hindrances to be overcome by selection, or as generators of non-adaptive and non-important by-products of primary events.
This position is very difficult to maintain. It is clear that the evolutionary possibilities in any given species must be limited by morphological and physiological, and indeed by developmental features. In fact, the evolutionary process itself must be seen as largely determined by these factors, a point made very clearly as far back as 1960 by Lancelot Law Whyte in his famous little book Internal Factors in Evolution.
As Waddington notes,
Richard Goldschmidt is one of the first to look at evolution in what are, in effect, developmental terms – one of the first in fact to note the absurdity of the thesis that development and evolution (or ontogeny and phylogeny) can be understood in isolation from each other. [C. H. Waddington, The Evolution of an Evolutionist.]
The idea that evolution proceeds by massive and abrupt evolutionary changes as opposed to small and gradual changes, is also gaining greater credibility, especially as the result of the work of Gould, Eldridge, and Stanley, who between them developed the theory of ‘punctuated equilibrium’. Though they do not see evolutionary changes to be quite as abrupt as did Goldschmidt, they do not see them either as occurring via a continuous series of small changes. Gould sees his work and that of his colleagues as providing
strong support for Goldschmidt’s gut feeling that extrapolation of small-scale adaptive change within local populations would not encompass all of evolution.
However, they do not accept Goldschmidt’s view that macroevolutionary change can occur as the result of the repatterning of the chromosomes, though C. H. Waddington takes this view very much more seriously, considering “genetic recombination” as an important factor in evolutionary change.
Goldschmidt’s idea that phenotypic changes, adaptive to stressful conditions can eventually become fixed genetically has also gained ground in the past forty years. A mechanism is clearly required to explain how feedback occurs between development (or ontogeny) and evolution.
One such mechanism was proposed by the American psychologist J. M. Baldwin in 1896. The Russian biologist Ivan Schmalhausen proposed another in his book Factors of Evolution: the theory of stabilizing evolution in 1949, as did Waddington with his “Genetic Assimilation”. [See The Evolution of an Evolutionist, his collected essays.]
Significantly Jean Piaget saw the requisite feedback mechanism as provided by a modified version of Goldschmidt’s phenocopy. [See Jean Piaget, Le Comportement, Moteur de l'Evolution, Gallimard, Paris.]
With regards to Goldschmidt’s notion that systemic mutations are usually directive rather than random, this too has become more acceptable as a result of the experiments carried out by John Cairns and his associates, who found that many mutations are highly functional – which is even more difficult to reconcile with the Darwinian thesis.
Goldschmidt’s idea that the ‘corpuscular gene’ does not exist, is, on the other hand, difficult to maintain in the light of the deciphering of the genetic code by Francis Crick and James Watson. However, it has now become apparent that individual genes determine only superficial characteristics like eye colour and the wrinkles on a pea – (i.e. the sort of characteristics that Mendel was concerned with).
Important characteristics tend to be determined by whole constellations of different genes (polygeny) while each gene is now also seen as contributing to the characteristics of many different traits in the individual. This suggests that evolutionary change cannot be explained in terms of changes occurring to individual genes. Indeed the Harvard biologist, Richard Lewontin, has gone so far as to propose a “geneless” theory of evolution.Back to top
Of course, theoretical biology is still dominated by hard-line neo-Darwinians and sociobiologists, such as E. O. Wilson, John Maynard-Smith, Michael Ruse, and Richard Dawkins, who have chosen to ignore the obvious flaws in the theory they still defend so ardently.
However, it is unlikely that their dominance of this field will last for long. Neo-Darwinism cannot be reconciled with Jim Lovelock’s Gaia Theory, which is gaining credibility all the time, nor with the work of the structuralists such as Brian Goodwin at the Open University, and David Lambert at the University of Auckland, or with the work of other young biologists, who see evolution in more holistic terms, such as Mae Wan Ho, also at the Open University. The latter in particular, as she fully admits, has been very much influenced by the work of Richard Goldschmidt.
We may, in fact, be on the verge of a general paradigm shift, and in a few years it may well be the defenders of the neo-Darwinian orthodoxy whose turn it will be to be laughed at and derided.
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