Oscar Riddle was a scientist who came good. After an inauspicious and tedious early research career he is remembered for discovering the hormone prolactin. Prolactin from the pituitary gland is essential for lactation as well as being involved in a whole host of physiological processes in vertebrates, acting both through the blood stream as a hormone and as a local controlling factor within tissues and organs.
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| Oscar Riddle from here |
Working on something mammalian was a major diversion for Riddle. He had for over 20 years worked almost exclusively on pigeons, particularly in the 1920s on metabolism, reproduction and endocrinology. He was therefore well prepared to ask the simple question, is the secretion of crop ‘milk’ in pigeons also stimulated by his newly-extracted hormone. Yes, was the answer and Riddle, Bates and Dykshorn soon developed the pigeon crop sac as a bioassay for prolactin activity. The lasting importance of this discovery is sometimes overlooked by modern endocrinologists who all seem to be based in the increasingly inward looking sub-discipline of clinical endocrinology. Because they had to wait until the 1970s for a reliable method of measuring the concentrations of prolactin in blood, the bioassay is of no interest. However, it continued to be used, certainly into the 1980s, by those looking for prolactin-like hormones produced in other tissues, the physiologically very important placental lactogens produced by the placenta, for example.
On 26 December 1932, shortly after the publication of their work, Simon Dykshorn, variously described then as a research student or research fellow, was found shot dead in woods, where he had gone hunting, about a mile west of the Cold Spring Harbor Laboratory on Long Island, New York, where Riddle had worked since 1912. Dykshorn was 30.
After this tragedy, with Bates and Ernest Lester Lahr (1897-1982), Riddle went on to further purify their prolactin preparation and to identify it as a protein.
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| Riddle's Prolactin Team From Family Trees on ancestry.com |
The reason Riddle had spent much of his time studying pigeons and making little headway for some years was that he completed his PhD in Chicago under the supervision of Charles Otis Whitman. Whitman was trying to find the cause of the pattern of alternate light and dark bars on the feathers of many species of birds. This question arose out of Whitman’s long interest in the evolution of birds. There were many dead ends and misinterpretations, not helped by Whitman’s rejection of both Mendelian genetics and natural selection. His devotion to the study of pigeons was such that he worked increasingly at home where for a period of 15 years he usually had around 500 individuals of 40 species in dovecotes around his house. He hybridised 40 species. Foy years Whitman moved his pigeons each summer to Marine Laboratory at Woods Hole in Massachusetts (where had had been its director) by rail, and then moved them back to Chicago in September. Initially the University of Chicago paid for the pigeons to be moved but then refused and thereafter Whitman and his wife paid.
Riddle valued highly the companionship and guidance of Whitman. He wrote: ‘Whitman became nearer to being a father to me than anyone I have known.’ Whitman died suddenly in 1910, aged 57. Riddle undertook to write up Whitman’s work. However, the new regime in Chicago, not impressed by Whitman’s research, did not want his disciple. Whitman though had been a influential figure in American zoology and In 1912, the Carnegie Institute of Washington took on Riddle as a research associate, together with Whitman’s pigeons and papers. By 1913 all were ensconced at Cold Spring Harbor. There is something odd in that the director of the lab, Charles Benedict Davenport (1866-1944), must have acquiesced to Riddle’s appointment—perhaps he had no choice—but did nothing to provide him with adequate facilities. Whitman’s views were the exact opposite of the main thrust of work, Mendelian genetics, at the lab and it must have come as some embarrassment when Whitman’s work was eventually pulled together over the next 6 years and published in three volumes, when the title of the first was Orthogenetic Evolution in Pigeons. Whitman really believed he had demonstrated orthogenetic evolution or evolution along pre-determined pathways. However, Davenport wrote an extensive obituary of Whitman. Perhaps they were friends and Davenport felt under some obligation to provide some means of concluding Whitman’s work while not appreciating the direction Whitman was heading. In the meantime Davenport was earning opprobrium of his own as one of the leading American exponents of eugenics. His views were so extreme and his notions from Mendelian genetics so warped that other eugenicists disowned him. He was, and remained, a supporter of the Nazi party in Germany before and during the Second World War. Perhaps then there is a simpler explanation for Davenport’s lack of support for Riddle: was neither a Mendelian geneticist nor a eugenicist who did not fit into Davenport’s organisation.
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| Whitman with his pigeons From here |
Most of the immense amount of work to sort and write up Whitman’s notes fell to Riddle. But even this labour of love ran into difficulties. For some reason, Whitman’s widow did not allow Riddle access to all of her husband’s papers. The very long and arduous episode in Riddle’s life must be one of the best illustrations of ‘no good deed goes unpunished’. The publications were not well received, particularly the one on orthogenesis. Reviewers of that volume pointed out that many of his ideas had already been, in kinder words, shot down in flames. The three volumes can be found online and even a quick look provides some appreciation of the amount of work and the level of detail involved. Two artists were engaged to provide the colour plates and other illustrations. The cost to the Carnegie Institute must have been enormous.
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| Volume 1 of the posthumous works of Whitman |
Eventually, Riddle, by developing interests in how birds work and into the burgeoning fields of metabolism and endocrinology, pulled away from Whitman’s ideas on evolution. He was by 1930 very well equipped to discover something really important.
And it was Whitman who provides the link I referred to in the title of this article. Lord Walter Rothschild in his 1907 book, Extinct Birds, quoted from notes sent to him by James H. Fleming of Toronto”
For all practical purposes, the close of the Nineteenth Century saw the final extinction of the Passenger Pigeon in a wild state, and there remained only the small flock, numbering in 1903 not more than a dozen, that had been bred in captivity by Prof. C. O. Whitman, of Chicago; these birds are the descendants of a single pair, and have long ago ceased to breed. It was in an effort to obtain fresh blood for this flock that I started a newspaper enquiry that brought many replies, none of which could be substantiated as records of the Passenger Pigeon, and many referred to the Mourning Dove. I am aware that there has been lately wide-spread and persistent rumours of the return of the pigeons, but no rumour has borne investigation, and I feel that Prof. Whitman's small flock, now reduced (in 1906) to five birds, are the last representatives of a species around whose disappearance mystery and fable will always gather.
Rothschild’s correspondent seems to have overlooked the Passenger Pigeons at Cincinnati Zoo, where they had also bred and where the last one, ‘Martha’, died in 1914. The entry for Whitman on Wikipedia states that he sent Martha to Cincinnati in 1902. However, some contemporary evidence indicates that was not the case while another account—from the same source—suggests that the last living Passenger Pigeon was one received from Whitman. I will return to Martha and her origins in another article.
Oscar Riddle was elected to the U.S. National Academy of Sciences in 1939 for his work on prolactin. It was as a result of his earlier career with Whitman that he realised the importance of studying the effect of prolactin on the pigeon’s crop. The results of that second discovery were then developed into a vital bioassay that was used—and could well be still used—throughout the world for studying the biological activity prolactin-like compounds in all vertebrates.
Riddle was a major player in his later life in promoting the study and teaching of evolution in the USA where belief in the supernatural then—and still does—trumps rationality. He really did, after a shaky start, come good.
Oscar Riddle died on 29 November 1968, aged 91.
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| Female Passenger Pigeon by K. Hayashi K. Hayashi and K. Toda were the artists commissioned to provide the illustrations to Whitman's posthumous works. This is from Volume 2 |
Corner GW. 1974. Oscar Riddle 1877-1968. National Academy of Sciences. Biographical Memoirs: Volume 45. p 448-488. Washington, DC: The National Academies Press. doi.org/10.17226/568
Davenport, CB. 1917. The personality, heredity and work of Charles Otis Whitman. 1843-1910. American Naturalist 51, 5-30.
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