In this essay, I shall try to establish precise criteria for accurately describing gene "behaviour" in the context of evolutionary theory as "selfish" and proceed in the series of essays that follow to show that there are no grounds in ethological studies and evolutionary genetics for conceiving of genes as "selfish" or postulating a gene centred theory of evolution.
I shall argue that the creators of the selfish gene theory/gene centered theory of evolution approached the body of ethological evidence with a strong atheistic bias that compelled them to cherry-pick the evidence in support of a bottom-up theoretical picture in line with their philosophical prejudice as atheists.
What exactly does it mean to say that genes are selfish?
According to the selfish gene or gene centred theory of evolution, genes have a tendency to seek to maximize copies of themselves in a gene pool, or as stated in the context of ethological studies, an animal seeks to maximize its inclusive fitness—the number of copies of its genes passed on globally (rather than by a particular individual).
Regardless of any definition of the term "selfish" that might have been suggested with reference to the "selfish" gene theory, genes can only be described correctly and with scientific precision as "selfish" if and only if it can be demonstrated that evolution consistently subordinates organism/group goals to the stated gene goal of maximizing copies of itself.
The last statement in italics is pivotal to understanding the argument, because even in the context of an organism/group centered view of evolution, allowance must still be made for expression of gene goals as asserted in the selfish gene theory. The selfish gene as the primary driving force in evolution is demonstrated only if it can be shown that in circumstances of conflict between gene goals and organism/group goals, gene goals consistently override organism/group goals. Conversely, gene "altruism" (for want of a better term!) is revealed in circumstances of conflict between gene goals and organism/group goals, as tendency for organism goals to override gene goals; that is, in the overall, where gene goals are found to be subsumed in organism goals (rather than vice-versa as "selfish" gene theory proposes).
An overall definition of gene "behaviour" as altruistic, therefore, allows, as already stated, that genes pursue "selfish" motives of their own only to the extent that they do not come into conflict with those of the organism or the group. Where there is a conflict, the synthetic perspective predicts that organism/group goals will override gene goals.
By way of illustration, the social altruism of the kamikaze pilot is demonstrated conclusively only at the moment he crashes his plane into enemy ship. The Muslim suicide bomber's altruism is revealed unquestionably only when he sacrifices his life for the promotion of the Islamic cause. The dominant male baboon who makes himself into a decoy to allow his troop, consisting of more genetically unrelated individuals than closely related individuals, escape danger behaves altruistically, even if moments before he had been throwing his weight around "selfishly" as the dominant male in the troop.
My essays will draw evidence mostly from higher social animals only for the reason that this is the area of my greater familiarity. The discussion of ethological evidence of evolution in the context of higher life forms takes us a level higher, from the organism centered to a group centered view of evolution, an idea that has for long been a taboo in the gene centered theory of evolution.
Kin selection strategy and the male social order
I shall begin in the next few posts with an overview of incidence of "altruistic" behavior in higher social animals that calls the assumptions of gene centered theory of evolution to question before proceeding to delve in greater detail on the significance of "altruistic" behaviour in both human and non-human social animals to evolutionary theory. Later, I shall be examining the body of clinching evidence against the gene centered theory of evolution in the mechanisms of sexual reproduction in higher animals.
The received wisdom in Darwinian evolutionary thought, in the explanation of incidence of what is considered apparent "altruistic" behavior in social animals, is contained in the theoretical speculations of kin selectionists. According to the theory of kin selection, individuals do not sacrifice themselves for the "social order" or for overall good of the group (or species) being primarily "selfish." Instances of apparent altruism ultimately turn out not to be "selfless," because the individual (according to the kin selection theory) who gives up his life or endangers himself does so not that natural selection may operate at the large scale of the social entity but only so as to promote copies of his genes in a son, brother or sister. Thus, the degree of relatedness of two individuals(percentage of genes held in common) becomes(all other things being equal) a reliable basis for prediction of "altruistic" behavioral disposition between individuals.
In spite of this argument, no serious ethologist denies the fact of what appears genuinely "altruistic" behavior in highly socialized higher animals (going by the mathematics of "degree of relatedness" as postulated in the kin selection theory) and the fact that too often the behavior of social animals reflects, in the rigidly hierarchical social status order, a tendency to genetic suicide, especially among males. Patterns of socially adjusted behavior in the typical highly socialized male animal which appear at cursory glance in enhancement of his genetic fitness often turn out on closer examination to be literally a "call to service" fundamentally inimical to the goal of maximizing global genetic fitness with respect to a genotype or phenotype, as technically defined. Examination of the behavior of highly socialized higher male animals is suggestive of a top-down imposition that predisposes the male social animal to patterns of "jingoistic behavior" tantamount to genetic suicide making nonsense of the "Dawkinian" refrain that animals always seek to maximize genetic fitness.
Examples abound in nature of indisputable cases of behavior in higher social animals that defy the postulates of kin selection Darwinian strategy: The male baboon in the large baboon troop who, on sighting a leopard approaching the troop, not only makes himself conspicuous but also attacks the leopard inefficaciously getting disemboweled in the process. The male bird who gives a warning cry at the appearance of the hawk thus enabling the entire flock fly away but makes himself conspicuous and at a high risk of being picked out by the hawk. Young men who enlist enthusiastically to die in useless, pointless wars. All three (male baboons, male birds and male humans) in their behavior frustrate the "selfish" aims of kin selection, which is to enhance genetic fitness, for they give up or endanger their lives for more unrelated persons (as evolutionary geneticists reckon "relatedness" in a population) than related persons; and usually the lives of close genetic relatives in the population are not in sufficient immediate danger to warrant the act of "self sacrifice."
The mass genetic suicide of pointless wars of imperial adventures hardly qualifies for kin selection Darwinian strategy maximizing of genetic fitness (try to construct a kin selectionist argument to explain Alexander the Great fighting all the way from Greece to India, and dying at 33 in a drunken brawl, childless!). Yet as Robert Audrey pointed out in his "African Genesis," war has been the primary cultural preoccupation of male humans in history.
Some evolutionary psychologists have attempted to dribble around this problem by pointing to the fact that a major motive in war among primitive tribes is to steal women, as part of individual strategy for maximizing genetic fitness. But it has been shown that jingoism and its associated suicidal behavioural tendencies are extraordinarily easy to evoke in male social animals both human and non-human, independent of the lure of scarce resource rewards or genetic fitness enhancing access to females. Steven Pinker (in his How the Miind Works) reports an experiment by Muzafar Sherif in which he randomly divided a group of socially well adjusted summer camp American boys in to two groups and set them to compete in sports and skits. Within a few days the competition had intensified to the point that the groups had gone into open warfare with sticks, bats and rocks forcing Sherif to hastily terminate the experiment.
Sherif's boys were not competing for scarce resources, they were not competing for access to females. Issues of "passing on their genes to the next generation" were not at stake. Yet, gang competition degenerated readily into mortal combats.
The evidence form Sherif's experiment and others observations of male social animal behavior, which I shall lay out plainly in subsequent essays, show that social animals, especially male social animals, do not compete primarily to maximize genetic fitness as selfish gene theory leads us to expect.