Course Materials

George C. Williams, Adaptation and Natural Selection, 1966, Princeton University Press (new ed. 1996)


Williams is one of the most respected theorists in the contemporary evolutionary biology. This eminence is due, among others, to his book Adaptation and Natural Selection. In the new Preface attached to the 1996 edition, Williams recalls:

My first awareness of a motive for Adaptation and Natural Selection came during the 1954-55 academic year while on a teaching fellowship at the University of Chicago. The triggering event may have been a lecture by A. E. Emerson, a renowned ecologist and terminte specialist. The lecture dealt with what Emerson termed beneficial death, an idea that included August Weismann's theory that senescence was evolved to cull the old and impaired from populations so that fitter youthful individuals could take their places. My reaction was that if Emerson's presentation was acceptable biology, I would prefer another calling. (ix, Uchii's italics)

Can you see the thrust of the italicized part?

I am going to retire very soon (March 2006), not due to scenescene but due to the institution called "retirement age", beneficial (I believe) to younger researchers. But is the biological world like our bureaucratic institution? If you find this question uninteresting, don't come to our seminar! If you are going to attend, begin with Introduction; some chapters will be skipped, but not those on "group selection".

Chapter 1. Introduction: pp. 3-19

Lateral Line / Anting of Birds / Rectal Gland / Cypress Knees

Chapter 2. Natural Selection, Adaptation, and Progress: pp. 20-35

Intermediate Generalizations / Selection Coefficient / Geological Time Divisions / Consequences of Natural Selection / Common Fallacies /

Chapter 3. Natural Selection, Ecology, and Morphogenesis: pp. 56-71; 87-91 [discussion of genetic assimilation and regeneration will be skipped]

Environment / Sickle-cell Anemia / Homology and Homoplasy /

Chapter 4. Group Selection: pp. 92-124

Group Selection vs. Gene Selection / Adaptation, organic and biotic / Fossil Record and its interpretations / Models of Group Selection / Meiotic Drive /

Chapter 5. Adaptations of Genetic System: pp. 125-157

Sexual Reproduction /

Chapter 6. Reproductive Physiology and Behavior: pp. 158-192


Chapter 7. Social Adaptations: pp. 193-220


References for the Beginner (for the background of evolutionary biology)


Darwin Seminar (course materials, 2002)

Brief History of Evolutionary Biology (And Consult an elementary textbook on evolutionary biology or population genetics)

Cyber Museum of Genetics(遺伝学電子博物館) 

Philosophy of Biology (course materials, 1999; see Evolution of altruism by group selection, in particular)


G. C. Williams, Official Site

The Structure of Adaptationism

Who is Dr. Pangloss?

Williams on Pony Fish

Hardy-Weinberg Law

Mark Ridley, Evolution, Tutorials (at Blackwell, useful for beginners)

Darwin on "Group Selection"



First Assignment // Second Assignment // Third Assignment

Last modified, Sept. 15, 2007. (c) Soshichi Uchii