Ó Indian Academy of Sciences PERSPECTIVES William Bateson: a biologist ahead of his time PATRICK BATESON* Sub-Department of Animal Behaviour, University of Cambridge, Madingley, Cambridge CB3 8AA, UK William Bateson coined the term genetics and, more than son in a letter to Adam Sedgwick in 1905 when he hoped anybody else, championed the principles of heredity dis- to be appointed to a new chair (B. Bateson 1928). covered by Gregor Mendel. Nevertheless, his reputation William Bateson was the most vigorous promoter of is soured by the positions he took about the discontinui- Mendel’s ideas at the beginning of the twentieth century ties in inheritance that might precede formation of a new and effectively launched the modern subject of genetics. species and by his reluctance to accept, in its full- Historians of biology acknowledge the importance of this blooded form, the view of chromosomes as the control- contribution but criticise his ideas on sudden changes in lers of individual development. Growing evidence sug- evolution leading to the origin of new species and his gests that both of these positions have been vindicated. questioning of the role of chromosomes (Mayr 1982). In New species are now thought to arise as the result of this article I re-examine these criticisms of Bateson in the genetic interactions, chromosomal rearrangements, or light of modern advances in biology. both, that render hybrids less viable or sterile. Chromo- Bateson was born on 8 August 1861. He was raised in somes are the sites of genes but genes move between a comfortable home and had an eminent father who was chromosomes much more readily than had been previ- for 24 years Master of St John’s College, Cambridge. ously believed and chromosomes are not causal in indi- Family life was filled with conversation and William and vidual development. Development, like speciation, requires his siblings grew up to be strong-willed, disputatious and an understanding of the interactions between genes and highly intellectual. William obtained a first class in natural the interplay between the individual and its environment. sciences at Cambridge, specializing in zoology. His first major study was on the embryology of Balanoglossus living on tidal mudflats; it looks like a worm but is now Introduction regarded as a vertebrate ancestor. It was particularly common in Chesapeake Bay on the east coast of the William Bateson is credited with coining the term ‘gene- United States and it was there that Bateson got to know a tics’ as it is currently used as a noun. The adjective brilliant American zoologist, William K. Brooks, who ‘genetic’ had been introduced into the English language was bringing out a book about heredity at the time in the 1830s. When Charles Darwin was nominated for (Brooks 1883). Although few biologists doubted that the Royal Society’s highest award, the Copley Medal, his Charles Darwin had provided the most coherent and supporters referred, among other things, to his contribu- complete explanation for adaptation by the process of tion to ‘genetic biology’. Darwin was eventually awarded natural selection, the necessary conditions for one species the Copley Medal in 1864, but mention of this contribu- to become distinct from another remained a source of tion was omitted as grounds for making the award by a dispute. timorous Council of the Royal Society because On the origin of species was still so controversial (Burkhardt Discontinuous variation 2001). It is clear in the usage applied to Darwin’s then disputed contribution that ‘genetic’ meant ‘pertaining to origins’. Anyhow, genetics as a noun, meaning the study Darwin’s mechanism for evolutionary change consisted of heredity and variation, was first used by William Bate- of three crucial steps. Each step must have been in place if adaptation by the organism to the environment occur- red in the course of biological evolution. First, variation *E-mail: [email protected]. Correspondence to Prof. Patrick Bateson, must have existed. Second, some variants must have sur- The Provost’s Lodge, King’s College, Cambridge CB2 1ST, UK. vived more readily than others. Third, the variation must T Journal of Genetics, Vol. 81, No. 2, August 2002 49 Patrick Bateson have been inherited. While Darwin’s proposal provided a discontinuities could provide evidence for steps that powerful and plausible mechanism for generating adapta- might lead to the appearance of a new species. This inter- tions, it was less obvious that it would provide what was est prepared him for the rediscovery of Mendel’s work needed for the formation of a new species. His friends which provided the rules for how the qualitative differ- Thomas Henry Huxley and Francis Galton were doubtful. ences between members of the same species could be In his excellent book, Donald Forsdyke (2001) describes inherited. Bateson was sometimes thought to be anti- how George Romanes developed a physiological theory Darwinian because of his search for discontinuities in for hybrid sterility (e.g. Romanes 1886). The theory was natural variation (Bowler 1983). Mark Ridley (1996) has intended to solve the speciation problem, but it was argued that Darwin himself was only a gradualist as far roundly attacked by Darwin’s defenders and Romanes’s as adaptation was concerned and that he had not nailed contributions to evolutionary theory were forgotten, his colours to the mast over speciation. Anyhow, Bateson although his book Animal intelligence is read to this day. was never opposed to the Darwinian proposal for the William Brooks was also doubtful whether speciation evolution of adaptations, although he did not share the could be explained by natural selection. Much of this belief, usually attributed to Darwin, that the evolution of scepticism rubbed off on Bateson in the course of their a new species always involved continuous and gradual long discussions after work in Chesapeake Bay. modification. In the first phase of his life’s work, Bateson wanted to know what variation within a species might look like. He was particularly interested in finding major discontinui- Principles of inheritance ties in characters. He amassed a great quantity of examples in his book Materials for the study of variation which Having established that discontinuities were to be found appeared in 1894 (W. Bateson 1894). He believed that such in nature, the next step was to discover what happened to Figure 1. William Bateson (1861–1926). 50 Journal of Genetics, Vol. 81, No. 2, August 2002 William Bateson: a biologist ahead of his time such discontinuities from one generation to the next. stained deeply by the black urine. The critical compound Bateson set to work on the experimental breeding of in the urine of people with alkaptonuria, as the condition animals and plants to find out how variations might be is called, is homogentisic acid, of which 2.5 to 6.0 grams inherited. Before the rediscovery of Mendel’s work, the is produced each day. Alkaptonuria is much more com- best known principle was the Law of Ancestral Heredity, mon in men than in women and, though always rare, is promulgated in the nineteenth century most actively by particularly likely to occur in the offspring of first cousin Francis Galton (1897). Those of a mathematical bent marriages. Bateson got to hear of this and in December liked it because it meant that a prediction about the char- 1901 he and Saunders reported Garrod’s finding to the acteristics of an individual could be derived probabilisti- Evolution Committee of the Royal Society. They wrote: cally from the characteristics of the ancestors. Galton ‘. the mating of first cousins gives exactly the condi- produced two forms of the Law of Ancestral Heredity, tions most likely to enable a rare and usually recessive but these were mathematically inconsistent (Provine character to show itself. If the bearer of such a gamete 1971). Karl Pearson subsequently cleaned up the mathe- mates with individuals not bearing it, the character would matics (Pearson 1898) but did not help matters when he hardly ever be seen; but first cousins will frequently be called his revision ‘Galton’s theory’. Confusion reigned. bearers of similar gametes, which may in such unions The independent rediscovery of Mendel’s work by De meet each other, and thus lead to the manifestation of the Vries, Tschermak and Correns changed all that. An peculiar recessive characters in the zygote’. (Royal Society. enjoyable account is given by Henig (2000). When Bateson Reports to the Evolution Committee 1902) became aware of Mendel’s work he realized, supposedly Alexander Bearn, biographer of Garrod (Bearn 1993), in a flash, that here were the principles that would make found a long letter from Garrod to Bateson dated 11 sense of heredity. Supposedly, he was reading a paper by January 1902 beginning: ‘It was a great pleasure to De Vries on a train when he was travelling from Cam- receive your letter and to learn that you are interested in bridge to London to address the Royal Horticultural Society. the family occurrence of alkaptonuria.’ This suggests that It is worth noting that Olby (1987) has expressed some Bateson initiated the correspondence. Anyway, Garrod scepticism about both the suddenness and the site of this was quick to see the significance of Mendelism for con- realization. genital human conditions and referred explicitly to this Mendel’s discoveries may be summarized as follows. insight in his next paper (Garrod 1902). No case of two Inherited factors influencing the characteristics of an alkaptonurics having children together was known. Garrod organism come in pairs; while this is usually true, sex had both male and female alkaptonuric patients and he linkage (arising from unequal pairings of the sex chro- used to get them into the ward at the same time in the mosome in male mammals and female birds) was only hope that they might become fond of each other, marry discovered later.