Abstract: A genetical mathematical model is described which allows for interactions between relatives on one another's fitness. Making use of Wright's Coefficient of Relationship as the measure of the proportion of replica genes in a relative, a quantity is found which incorporates the maximizing property of Darwinian fitness. This quantity is named “inclusive fitness”. Species following the model should tend to evolve behaviour such that each organism appears to be attempting to maximize its inclusive fitness. This implies a limited restraint on selfish competitive behaviour and possibility of limited self-sacrifices.
Special cases of the model are used to show (a) that selection in the social situations newly covered tends to be slower than classical selection, (b) how in populations of rather non-dispersive organisms the model may apply to genes affecting dispersion, and (c) how it may apply approximately to competition between relatives, for example, within sibships. Some artificialities of the model are discussed.

Abstract: An altruistic individual has to gamble on cooperation to a stranger because it does not know whether the stranger is trustworthy before direct interaction. Nowak and Sigmund (Nature 393 (1998a) 573; J. Theor. Biol. 194 (1998b) 561) presented a new theoretical framework of indirect reciprocal altruism by image scoring game where all individuals are informed about a partner's behavior from its image score without direct interaction. Interestingly, in a simplified version of the image scoring game, the evolutionarily stability condition for altruism became a similar form of Hamilton's rule, i.e. inequality that the probability of getting correct information is more than the ratio of cost to benefit. Since the Hamilton's rule was derived by evolutionarily stable analysis, the evolutionary meaning of the probability of getting correct information has not been clearly examined in terms of kin and group selection. In this study, we applied covariance analysis to the two-score model for deriving the Hamilton's rule. We confirmed that the probability of getting correct information was proportional to the bias of altruistic interactions caused by using information about a partner's image score. The Hamilton's rule was dependent on the number of game bouts even though the information reduced the risk of cooperation to selfish one at the first encounter. In addition, we incorporated group structure to the two-score model to examine whether the probability of getting correct information affect selection for altruism by group selection. We calculated a Hamilton's rule of group selection by contextual analysis. Group selection is very effective when either the probability of getting correct information or that of future interaction, or both are low. The two Hamilton's rules derived by covariance and contextual analyses demonstrated the effects of information and group structure on the evolution of altruism. We inferred that information about a partner's behavior and group structure can produce flexible pathways for the evolution of altruism.

Abstract: Dominance relationships in primate societies are generally inferred by analyses of agonistic interactions. This aspect of social organization is so striking in macaque and baboon societies that many theoreticians have postulated selective mechanisms operating on the genetic attributes which contribute to high dominance rank. Alpha males were hypothesized to increase their genetic fitness by successfully competing with other males for access to ovulating females. Evidence relevant to these speculations has been mixed. Whereas some investigators found alpha males had near exclusive sexual access to females, others failed to confirm preferential access to ovulating females. Indeed, considerable variability in competition for females existed not only among species, but also among troops of the same species living in different habitats. Further, partner selection was not an exclusive male prerogative; females proved to express active preferences for particular males as sexual partners, and these preferences were not related to high male aggressivity. Alpha males, however, were noted to maintain their positions through social skills as members of a central core or alliance, and high rank was related primarily to seniority. Moreover, alpha males responded actively to challenges to the troop and were judged to contribute significantly to the survival of infants. It was therefore hypothesized that increased genetic fitness related to the increased survival of immature animals in the troop, most of which would already be the offspring of senior (and hence alpha) males. Selection would then be for the social skills leading to successful alliances in troop defense. Such skills might also relate to female partner preferences thus increasing the reproductive effectiveness of alpha males at any point in their careers, including years prior to and following their assumption of alpha rank.

Abstract: In many different species it is common for animals to spend large portions of their lives in groups. Such groups need to divide available resources amongst the individuals they contain and this is often achieved by means of a dominance hierarchy. Sometimes hierarchies are stable over a long period of time and new individuals slot into pre-determined positions, but there are many situations where this is not so and a hierarchy is formed out of a group of individuals meeting for the first time. There are several different models both of the formation of such dominance hierarchies and of already existing hierarchies. These models often treat the two phases as entirely separate, whereas in reality, if there is a genuine formation phase to the hierarchy, behaviour in this phase will be governed by the rewards available, which in turn depends upon how the hierarchy operates once it has been formed. This paper describes a method of unifying models of these two distinct phases, assuming that the hierarchy formed is stable. In particular a framework is introduced which allows a variety of different models of each of the two parts to be used in conjunction with each other, thus enabling a wide range of situations to be modelled. Some examples are given to show how this works in practice.

Abstract: Observation of foal development shows that the appearance of adult-type motor grazing behaviour, selection of grass vs. non-grass and the avoidance of poisonous plants occur concurrently between the ages of 4 and 6 weeks. Suckling behaviour and close association of foal with dam change with time but show no particular coincidence with grazing behavioural changes. Coprophagy of the foal on maternal faeces does, however, correspond chronologically with the foal learning to graze selectively. This correspondence suggests that, as well as other uses, in domestic horses coprophagy may function to imprint on the foal the food-selective values of its dam.