Abstract: Recent evidence in natural and semi-natural settings has revealed a variety of left-right perceptual asymmetries among vertebrates. These include preferential use of the left or right visual hemifield during activities such as searching for food, agonistic responses, or escape from predators in animals as different as fish, amphibians, reptiles, birds, and mammals. There are obvious disadvantages in showing such directional asymmetries because relevant stimuli may be located to the animal's left or right at random; there is no a priori association between the meaning of a stimulus (e.g., its being a predator or a food item) and its being located to the animal's left or right. Moreover, other organisms (e.g., predators) could exploit the predictability of behavior that arises from population-level lateral biases. It might be argued that lateralization of function enhances cognitive capacity and efficiency of the brain, thus counteracting the ecological disadvantages of lateral biases in behavior. However, such an increase in brain efficiency could be obtained by each individual being lateralized without any need to align the direction of the asymmetry in the majority of the individuals of the population. Here we argue that the alignment of the direction of behavioral asymmetries at the population level arises as an “evolutionarily stable strategy” under “social” pressures occurring when individually asymmetrical organisms must coordinate their behavior with the behavior of other asymmetrical organisms of the same or different species.

Abstract: Gene expression affects social behavior only through changes in the excitabilities of neural circuits that govern the release of the relevant motor programs. In turn, social behavior affects gene expression only through patterns of sensory stimulation that produce significant activation of relevant portions of the nervous system. In crayfish, social interactions between pairs of animals lead to changes in behavior that mark the formation of a dominance hierarchy. Those changes in behavior result from changes in the excitability of specific neural circuits. In the new subordinate, circuits for offensive behavior become less excitable and those for defensive behavior become more excitable. Serotonin, which is implicated in mechanisms for social dominance in many animals, modulates circuits for escape and avoidance responses in crayfish. The modulatory effects of serotonin on the escape circuits have been found to change with social dominance, becoming excitatory in dominant crayfish and inhibitory in subordinates. These changes in serotonin's effects on escape affect the synaptic response to sensory input of a single cell, the lateral giant (LG) command neuron for escape. Moreover, these changes occur over a 2-week period and for the subordinate are reversible at any time following a reversal of the animal's status. The results have suggested that a persistent change in social status leads to a gradual change in the expression of serotonin receptors to a pattern that is more appropriate for the new status. To test that hypothesis, the expression patterns of crayfish serotonin receptors must be compared in dominant and subordinate animals. Two of potentially five serotonin receptors in crayfish have been cloned, sequenced, and pharmacologically characterized. Measurements of receptor expression in the whole CNS of dominant and subordinate crayfish have produced inconclusive results, probably because each receptor is widespread in the nervous system and is likely to experience opposite expression changes in different areas of the CNS. Both receptors have recently been found in identified neurons that mediate escape responses, and so the next step will be to measure their expression in these identified cells in dominant and subordinate animals.

Abstract: Theoreticians propose that trained winning and losing are important processes in creating linear animal dominance hierarchies, and experiments have shown that both processes can occur in animals, but their actual roles in creating natural hierarchies are unknown. We described agonism in 18 broods of three blue-footed boobies, Sula nebouxii, a species for which trained winning and losing have been demonstrated, to infer how these processes generate and maintain a natural hierarchy. Ranks in the linear hierarchy that emerged in every brood were initially assigned by asymmetries in age, size and maturity, which led to differences between broodmates in levels of expressed and received aggression and, consequently, to differences in the training of their aggressiveness and submissiveness. Later, ranks appeared to be maintained by the chicks' acquired aggressive and submissive tendencies combined with ongoing effects of persisting differences in size and maturity. Our results suggest that trained winning and trained losing are important in the construction of booby hierarchies but that these two axes of learning are largely independent. Increase in submissiveness occurs over a period of about 10-20 days, and the level of submissiveness reached varies with the amount of aggression received. After training, submissiveness is apparently maintained by a lower level of aggression and increasing use of threats. Threats become increasingly effective as chicks age, but are never as effective as attacks.

Abstract: This simulation study explores some conditions leading to transitivity within dominance orders. Combinations of three parameters were varied to study their consequences upon hierarchy formation and upon the degree of linearity of resultant structures. The factors studied were: (1) the importance of initial resource holding potentials (RHPs); (2) changes brought in RHPs by successive victories and defeats; and (3) accuracy of RHP assessment made by opponents. Results show that initial differences in RHP always lead to perfectly transitive chains whose rank order reflects the importance of initial differences. Even when simulated animals make important errors while assessing each other during round robin tournaments, emerging dominance structures are perfectly linear and ranks obtained in the structure are highly correlated with initial values in RHPs. Moreover, accumulated experiences of victory and/or defeat alone always lead to perfectly linear hierarchies. Their combination with initial individual differences in RHP led to the same conclusion. Even when assessment was far from being perfect, not only perfect chains were formed but initial values in RHPs significantly influenced rank order when the contribution of victory and defeat to RHP was relatively unimportant. The higher the importance of victory and defeat to RHP as compared to that of initial RHP values, the lower was the correlation between initial RHP values and the ranks order reached by individuals in the resultant hierarchies. In general also, the lower the variation within initial RHPs, the lower was the correlation between initial RHPs and ranks in the hierarchy. At a given level of initial RHP dispersion, increasing the contribution of victory and defeat to RHP diminished the correlation between initial RHP values and obtained ranks. In addition, inaccurate assessment reduced the overall correlation, especially when dispersion of initial RHP values was low and the contribution of victory and defeat relatively unimportant. These results shed some light on the controversy about the respective roles of initial individual attributes and that of patterns of resolution in the formation of animal hierarchies. We present the emergence of social order within closed systems as those simulated here as a case of self-organization.

Abstract: We spent two winters studying the social behaviour of wild woodland caribou (Rangifer tarandus caribou) at a time when their main food (ground lichens; Cladina sp.) is available only at snow craters dug by the animals. The competition for access to such craters was severe, the animals constantly trying to take over the craters of others. During a two-month period when a group maintained a constant size (20) and composition (all age-sex classes represented), we could rank the animals in a rather linear dominance hierarchy (Landau's index = 0.87). Rank was correlated with access to resources, percent of time spent active, and percent of time feeding in craters. It was also correlated with age and antler size. However, rank is not an attribute of individuals, but of a relationship between individuals. As such it is only an intervening variable between physical attributes and access to resources, a variable whose value has meaning only within a given group. Among the three attributes studied (age, sex, antler size), the latter was by far the best predictor of the occurrence and outcome of interactions. Between two individuals within any of the three age-sex classes studied (adult and yearling males and adult females), the one with larger antlers initiated significantly more often, escalated its aggression (to the point of hitting the target) less often, and enjoyed a higher success rate in obtaining resources. When their antlers were larger than those of an adult male target (i.e. males that had shed their antlers), adult females won almost all their interactions with adult males even though they escalated only one fourth of them. This clarifies the long-standing speculation that female caribou have antlers and shed them later than males, in order to overcome their sexual handicap in competition for food in the winter. We conclude that the link between rank and dominance of an individual on one hand, and some of its attributes on the other (e.g. sex, age, weight, antler size) is fundamentally realized by the animal itself through its active preference for targets it is likely to beat, i.e. targets with smaller antlers.