Jeremy Fox in his ‘Dynamic Ecology’ Blog has raised the eternal question of what have been the big ideas in ecology and were they successful, and this has stimulated me to write about the Chitty Hypothesis and its history since 1952. I will write this from my personal observations which can be faulty, and I will not bother to put in many references since this is a blog and not a formal paper.
In 1952 when Dennis Chitty at Oxford finished his thesis on vole cycles in Wales, he was considered a relatively young heretic because he did not see any evidence in favour of the two dominant paradigms of population dynamics – that populations rose and fell because of food shortage or predation. David Lack vetoed the publication of his Ph.D. paper because he did not agree with Chitty’s findings (Lack believed that food supplies explained all population changes). His 1952 thesis paper was published only because of the intervention of Peter Medawar. Chitty could see no evidence of these two factors in his vole populations and he began to suspect that social factors were involved in population cycles. He tested Jack Christian’s ideas that social stress was a possible cause, since it was well known that some rodents were territorial and highly aggressive, but stress as measured by adrenal gland size did not fit the population trends very well. He then began to suspect that there might be genetic changes in fluctuating vole populations, and that population processes that occurred in voles and lemmings may occur in a wide variety of species, not just in the relatively small group of rodent species, which everyone could ignore as a special case of no generality. This culminated in his 1960 paper in the Canadian Journal of Zoology. This paper stimulated many field ecologists to begin experiments on population regulation in small mammals.
Chitty’s early work contained a ‘big idea’ that population dynamics and population genetics might have something to contribute to each other, and that one could not assume that every individual had equal properties. These ideas of course were not just his, and Bill Wellington had many of the same ideas in studying tent caterpillar population fluctuations. When Chitty suggested these ideas during the late 1950s he was told by several eminent geneticists who must remain nameless that his ideas were impossible, and that ecologists should stay out of genetics because the speed of natural selection was so slow that nothing could be achieved in ecological time. Clearly thinking has now changed on this general idea.
So if one could recognize these early beginnings as a ‘big idea’ it might be stated simply as ‘study individual behaviour, physiology, and genetics to understand population changes’, and it was instrumental in adding another page to the many discussions of population changes that had previously mostly included only predators, food supplies, and potentially disease. All this happened before the rise of behavioural ecology in the 1970s.
I leave others to judge the longer term effects of Chitty’s early suggestions. At present the evidence is largely against any rapid genetic changes in fluctuating populations of mammals and birds, and maternal effects now seem a strong candidate for non-genetic inheritance of traits that affect fitness in a variety of vertebrate species. And in a turn of fate, stress seems to be a strong candidate for at least some maternal effects, and we are back to the early ideas of Jack Christian and Hans Selye of the 1940s, but with greatly improved techniques of measurement of stress in field populations.
Dennis Chitty was a stickler for field experiments in ecology, a trend now long established, and he made many predictions from his ideas, often rejected later but always leading to more insights of what might be happening in field populations. He was a champion of discussing mechanisms of population change, and found little use for the dominant paradigm of the density dependent regulation of populations. Was he successful? I think so, from my biased viewpoint. I note he had less recognition in his lifetime than he deserved because he offended the powers that be. For example, he was never elected to the Royal Society, a victim of the insularity and politics of British science. But that is another story.
Chitty, D. (1952) Mortality among voles (Microtus agrestis) at Lake Vyrnwy, Montgomeryshire in 1936-9. Philosophical Transactions of the Royal Society of London, 236, 505-552.
Chitty, D. (1960) Population processes in the vole and their relevance to general theory. Canadian Journal of Zoology, 38, 99-113.
Whenever I think of Dennis Chitty, I think of the Chitty-grid at Kluane where I trapped arctic ground-squirrels back in 1996 (the summer of the Fence Party), when working on my Bachelor project from Aberystwyth University, via Prof. Boonstra. Thank you for this insightful blog and bringing back some good memories of hard working, problem solving scientists and a very memorable first summer in the Yukon. (Many have offended the Royal Society, no doubt. No comment needed there!)
Even if Chitty’s ideas, from the present perspective, were too simple and also wrong to some extent, they have been very influential. They have inspired a lot of research. Kids of the moderns days may wonder why there were “vole wars” (expression by Rick Ostfeld) in 60’s to 80’s. There were ideas but the tools to test genetics etc, in those days were not existing, as we have today. Also the data sets were very limited.
I would like to highlight an early Finnish ecologist, Paavo Voipio, who presented a kind of Chitty’s theory, but in a more diverse form, already in 1950. (see refs in my commentary below) Voipio’s idea was a physiological – genetic change in the course of the cycle.. Even if Voipio’s ideas were reviewed in some leading ecological journals in those days, they were forgotten soon. Not too uncommon for a remote journal from a small country. And maybe he was ahead of his time.
In their introductory chapters in the lemming book Stenseth and Ims (see below) reviewed ideas on population cycles, and with my modest help mentioned also Voipio’s ideas. That led Chitty to contact Voipio; Chitty had not known of Voipio’s papers, but he gave full acknowledgement to Voipio. Both gentlemen (in those days in their late 80’s I suppose) seemed to be happy.
Henttonen, H., 1998. A story around cycles – a commentary. – Ann. Zool. Fennici 35:129-130.
Stenseth, N.C. and Ims, R.A. (eds), The Biology of Lemmings. Academic Press for the Linnean Society of London,