Mirror, Mirror on the Wall / Who is the Fittest of Us All?
The question, starkly posed, could be the following: Which country, India or Pakistan, has the better chance of survival, and why?
In fact, the question is just an artifact to extend a discussion we have been having on this blog about the relationship of tolerance to survival. Our engagement with the issue has been at the very basic level of understanding but the very fact that we have been debating it leads us on to better and more sophisticated arguments. This, I strongly believe, is the beneficial outcome of discussions and conversations on a blog like this.
It is only because of our discussion that I paused to read more carefully an article that in the normal course I might have skipped over. Our discussion provided the hook that made me look for material that supported or qualified the arguments we had been making at our lay level.
The article, in the latest issue of the New York Review of Books (October 14, 2010), is a review (Is Goodness in Your Genes?) by H. Allen Orr of a book (The Price of Altruism: George Price and the Search for the Origins of Kindness) by Oren Harman. The book is about George Price, “a brilliant figure who performed fundamental work in the 1970s on several problems in evolutionary biology, including altruism.”
The review begins by posing the fundamental question that puzzles all who start to grapple with Darwin’s notion of the survival of the fittest. If we substitute ‘tolerance’ for ‘altruism,’ the question gets to the heart of the discussion that has been progressing on this blog:
If animals, including human beings, evolved by natural selection—a merciless process in which organism struggles against organism and all that matters is outcompeting everyone else—how could altruism arise? How could natural selection promote, or even allow, behavior that is costly to the individual that performs it but that benefits someone else?
The answer out of this dilemma has been posited before and the notion is the same that we have touched upon in our discussion: that we should look at evolution not just at the level of the individual but also at the level of a group:
In the twentieth century, two big developments marked attempts to understand the biology of altruism. In the first, V.C. Wynne-Edwards, author of Animal Dispersion in Relation to Social Behavior (1962), argued that evolutionary biologists had partly misconstrued the biological level at which natural selection acts. Though selection might often involve competition among individual organisms, Wynne-Edwards maintained that it often involves competition among entire populations of organisms. Under this so-called group selection view, altruism seems easily explained. Within a group, selfish individuals (ones who, say, reproduce rapidly even when resources are limited) might outcompete altruistic individuals (ones who slow reproduction when times are tough). But among populations, things are different. A population comprised of selfish individuals will likely ultimately deplete its resources and crash to extinction, whereas a population comprised of altruists will not deplete its resources and will survive to another day. Natural selection acting at the group level might therefore sustain altruism in animals. Though many biologists were skeptical of group selection—especially as the models offered were mostly verbal, not mathematical—no fully convincing alternative seemed available.
The second big theory (that I will gloss over in this post) was that of ‘inclusive fitness’ in which W.D. Hamilton argued that:
what matters under natural selection is not the fitness of this or that individual (where fitness is measured roughly by the number of progeny an individual produces) but the fitness of an individual averaged together, in a certain way, with the fitness of its relatives. Inclusive fitness could explain all manner of previously puzzling acts of animal altruism.
In search for an explanation for the existence of altruism, the first major contribution had shifted the focus from the individual up to the group; the second major contribution shifted it down from the individual to the gene.
The contribution of George Price in the 1970s was to show (via the Price Equation, a mathematically rigorous formulation) that it is possible to:
partition evolutionary change in a trait into the possible effects of natural selection acting simultaneously at multiple biological levels, e.g., the species, population, organism, gamete (sperm or egg), and gene.
The reviewer comments that the result of the Price Equation is at the same time very simple and very surprising – “quite a miracle.”
One can also use the Price Equation to study animal behavior, including altruistic behavior. Doing so reveals that accounting for altruism biologically does not require one to choose between believing that natural selection acts within groups or between groups. Instead, selection might act simultaneously at both levels, with selection within groups favoring selfishness (individuals that are selfish will out-compete those that are saintly) and selection between groups favoring altruism (groups including many cooperative individuals will do better, as a group, than those including many uncooperative individuals). Whether or not altruism evolves depends on the relative strengths of these conflicting forms of selection.
Crucially, the Price Equation showed that group selection arguments for altruism need not be offered tentatively or apologetically; they could be offered with mathematical precision.
It was at this point that the India-Pakistan comparison suggested itself in the context of long-term survival. The hypothetical extension of the logic discussed above is as follows:
It is quite likely (As Ahmed Kamran has been arguing in his series on the orientation of Islam in Pakistan) that the most intolerant group within Pakistan, the Taliban, would survive and come out on top as the fittest organism in that environment. But this struggle for supremacy would so enfeeble the organism that its survival as a group would be at stake.
This argument presumes, of course, the existence of ‘other’ groups that may or may not be favorably disposed towards the first. If we take India to be that ‘Other,’ as many Pakistanis insist it is, then we can examine the implications of the fact that India is a more tolerant organism than Pakistan. No one would dispute this although there could be arguments about how much more tolerant it is. This is irrelevant to our discussion because in the evolutionary perspective even marginal differences would be crucial to the outcome.
Hence the conclusion that is now open for discussion: A viciously intolerant group would survive and emerge on top in Pakistan weakening it to a degree that it would be unable to compete while India, either forced to be more tolerant because of peculiar initial conditions or lucky to be so because of wiser leadership, would ultimately dominate the region. Is there an exit ramp off this highway for Pakistan?
The book review is accessible in full only to subscribers but NYRB should be available in many good libraries. Excerpts in italics are quotes from the review. Another review of the book, with full access, is now available here in the Nation.