Narrow Roads of Gene Land, The Collected Papers of W.D. Hamilton, Volume I: Evolution of Social Behavior

American Zoologist, Feb 1997 by Allen J Moore

Narrow Roads of Gene Land, The Collected Papers of W. D. Hamilton. Volume 1: Evolution of Social Behavior. W. H. FREEMAN. 1996. 552 pages, 58 illustrations. $25.95 paperback. (ISBN: 0-71674530-5).

Since the 1960's the study of evolutionary aspects of social behavior has enjoyed a stunning rise in popularity, in large part because of W. D. Hamilton. Prior to his contributions, explanations for the evolution of social behavior presented special problems because it was not clear how many acts expressed within animal societies benefit the individual as well as the group. Explaining acts such as altruism, a behavior that is taxonomically widespread, was even worse; such behavior seemed to hurt individuals while benefiting the group. George Williams and John Maynard Smith, and later Richard Dawkins, convinced most animal behaviorists that "group thinking" should be replaced by "individual advantage," but what possible individual advantage could be associated with self-sacrifice, celibacy, or providing protection to unrelated individuals? This problem stumped Darwin; Fisher and Wright didn't even provide impenetrable mathematical models; Haldane came close, but didn't get it right (Hamilton, 1975). Not until the publication of Hamilton's seminal papers (Hamilton, 1964a,b) was the issue resolved. These two papers, which gave us kin selection, inclusive fitness, and an inordinate fondness for the social insects, provided empiricists with a testable theoretical framework. However, Hamilton is more than the father of kin selection. This collection of his papers through 1980 shows us why we should pay attention to whatever he has to say and why his ideas are considerably more diverse and fascinating than the 1964 papers alone.

Part I of Hamilton's collected papers provides us with his publications on the evolution of social behavior. (His work on sex and parasites since 1980 will be collected into part II and is to be published in 1997.) Purchasing collected works has always struck me as somewhat odd, thanks to the Xerox corporation, but I can heartily recommend this volume on several levels.

First, minor errors that appeared in the original works have been corrected. Thus, we get an accurate account of what Hamilton meant to say rather than continuing to perpetuate errors. While this may depreciate the historical value of the collection, it makes it extremely valuable for those who might want to "discover" Hamilton for the first time. Second, such a collection provides us with easy access to some of his more obscure (but still important) works. I, for one, had never read his 1971 contribution on the `Prisoner's Dilemma' or his chapter on the innate social aptitudes of humans (Hamilton, 1975), even though both contain ideas that are directly related to my own research. Finally, Hamilton makes this an invaluable volume for anyone interested in the social context of science by providing each chapter (paper) with a description of how and what he was thinking at the time the papers were written. This book is a bargain.

The autobiographical introduction to each paper is not just a few pages that set the story. Rather, some are fairly long and many are well-referenced (one has 48 notes, most of which are references to related literature or work that has occurred since the original paper was published). It is here that Hamilton alerts us to any corrections (typos, minor errors in calculations or formulas, mistakes regarding social insects) that have been made. But it is also in these introductions that we learn how his collaborations came about, how he was influenced by others (especially Price and May), and how personal events shaped his work. The personal events make for fascinating reading. For example, we learn that Hamilton's first published paper (Hamilton, 1963) was actually written after his betterknown 1964 papers. Further, we are told that this paper was submitted to the American Naturalist (after the requisite rejection from Nature) because he felt that the "hope of receiving my Ph.D. at the time seeming to founder" and he needed a publication to justify his time in graduate school. So we also learn that Hamilton is not always correct. In several of the introductions Hamilton relates portions of the tragic fate of his friend George Price. Before Price committed suicide he invented the covariance approach to selection, was the first to apply game theory to social interactions, and stimulated much of Hamilton's own work in the 1970's. Other introductions provide insights into the politics of science, and we learn how to get a paper published in Nature (collusion) and get the job of our choice (fame). All of the introductions make each paper more personal and easier to follow and made me want to read even those that I had read before.

Hamilton joins Darwin, Fisher, and Wright as one whose works are more often cited than read (something that has even been documented in Hamilton's case; see Seger & Harvey, 1980.). This can now be corrected. Hamilton has much to say on social behavior, social insects, senescence, sex, figs, and dispersal. Even more is to come on sex and parasites. This volume should be on the shelf of every graduate student interested in evolution, population genetics, or behavior. Hamilton's works provide a road map not only for some of the most exciting topics (many still under investigated) in evolutionary studies of animal behavior, but also show how ideas develop and should be pursued despite advisors and committees. For those of us no longer in graduate school, we too should own this book. And this time we should actually read the papers.