On dominance relations and the structure of animal societies: I. Effect of inherent characteristics

Societies are considered in which a non-transitive dominance relation exists between every pair of members, such as the peck-right in a flock of hens. A one-dimensional measure of the structure of such a society,h, is defined, withh=0 for equality andh=1 for the hierarchy. It is assumed that each member of the society is characterized by an ability vector whose components depend on individual characteristics such as size, concentration of sex hormone, etc., but not on social factors such as social rank. The distribution of abilities among members of the society is assumed to be given by a distribution function which is the same for all members, and the probability that one member dominates another is given by a function of the ability vectors of the two. On these assumptions formulas for the expected (mean) value and variance ofh are determined in terms of the distribution and dominance probability functions. Some special cases are calculated, especially that for normany distributed abilities and dominance probability given by the normal probability integral. Several conclusions are derived. If all members are of equal ability, so that dominance probability is 1/2, then any sizable society is much more likely to be near the equality than the hierarchy; and, as the size of the society increases, the probability that it will be near the hierarchy becomes vanishingly small. If the dominance probability is a weighted sum of several independent components, which make up the ability vector, then the society is less likely to be close to the hierarchy as the number of these components increases. The hierarchy is the prevalent structure only if unreasonably small differences in ability are decisive for dominance. From this it appears that the social factors, or psychological factors such as the previous history of dominance, which are not included in the present treatment, may be of great importance in explaining the observed prevalence of structures very close to the hierarchy in flocks of domestic hens.     

On dominance relations and the structure of animal societies: III The condition for a score structure

The necessary and sufficient condition is given forn integers to be the score structure of a society with a dominance relation. A proof is also given for a theorem showing that there are members who dominate every other member either directly or indirectly through a single intermediate member.     

Towards the integration of social dominance and spatial structure

My aim was to show how individual-oriented (or artificial life) models may provide an integrative background for the development of theories about dominance by including effects of spatial structure. Dominance interactions are thought to serve two different, contrasting functions: acquisition of high rank and reduction of aggression. The model I present consists of a homogeneous virtual world inhabited by artificial agents whose actions are restricted to grouping and dominance interactions in which the effects of winning and losing are self-reinforcing. The two functions are implemented as strategies to initiate dominance interactions and the intensity of aggression and dominance perception (direct or memory based) are varied experimentally. Behaviour is studied by recording the same behavioural units as in real animals. Ranks appear to differentiate more clearly at high than at low intensity of aggression and also more in the case of direct than of memory-based rank perception. Strong differentiation of rank produces a cascade of unexpected effects that differ depending on which function is implemented: for instance, a decline in aggression, spatial centrality of dominants and a correlation between rank and aggression. Insight into the origination of these self-organized patterns leads to new hypotheses for the study of the social behaviour of real animals. Copyright 2000 The Association for the Study of Animal Behaviour.     

Bystander effects and the structure of dominance hierarchies

Prior modeling work has found that pure winner and loser effects(i.e., changing the estimation of your own fighting abilityas a function of direct prior experience) can have importantconsequences for hierarchy formation. Here these models areextended to incorporate "bystander effects." When bystandereffects are in operation, observers (i.e., bystanders) of aggressiveinteractions change their assessment of the protagonists' fighting abilities (depending on who wins and who loses). Computer simulations demonstrate that when bystander winner effects alone are atplay, groups have a clear omega (bottom-ranking individual),while the relative position of other group members remainsdifficult to determine. When only bystander loser effects arein operation, wins and losses are randomly distributed throughout a group (i.e., no discernible hierarchy). When pure and bystanderwinner effects are jointly in place, a linear hierarchy, inwhich all positions (i.e., to when N = 4) are clearly defined,emerges. Joint pure and bystander loser effects produce thesame result. In principle one could test the predictions fromthe models developed here in a straightforward comparativestudy. Hopefully, the results of this model will spur on suchstudies in the future.     

Measuring social structure: a comparison of eight dominance indices

Measurement of social status is an important component of many behavioural studies. A variety of techniques have been developed and adopted, but while there have been some analyses of index properties using simulated data, the rationale for selecting a method remains poorly documented. As a first step in exploring the implications of index choice, we compared the characteristics of eight popular indices by applying each to the same data set from interactions between male fowl Gallus gallus, the system in which social hierarchies were first described. Data from eight social groups, observed over four successive breeding seasons, were analysed to determine whether different indices produced consistent dominance scores. These scores were then used in tests of the relation between social status and crowing to explore whether index choice affected the results obtained. We also examined the pattern of dominance index use over the last decade to infer whether this has likely been influenced by tradition, or by taxa of study animal. Overall agreement among methods was good when groups of birds had perfectly linear hierarchies, but results diverged when social structure was more complex, with either intransitive triads or reversals. While all regression analyses revealed a positive relationship between dominance and vocal behaviour, there were substantial differences in the amount of variance accounted for, even though the original data were identical in every case. Index selection can hence perturb estimates of the importance of dominance, relative to other factors. We also found that several methods have been adopted only by particular research teams, while the use of others has been taxonomically constrained, patterns implying that indices have not always been chosen solely upon their merits. Taken together, our results read as a cautionary tale. We suggest that selection of a dominance index requires careful consideration both of algorithm properties and of the factors affecting social status in the system of interest.     

Sociometrics of Macaca mulatta: II. Decoupling centrality and dominance in rhesus monkey social networks

The concept of the linear dominance hierarchy and a much less precise notion of a central/peripheral ordering of individuals have been prominent among the ideas about social organization of monkey groups. Although the latter has seldom been quantified, the ranks of individuals in the two orders are usually assumed to be correlated. This paper reports on a longitudinal study of a free-ranging group of rhesus monkeys. The individual histories of progression of dominance rank and an independently determined measure of centrality in the social grooming network are compared among a set of males. Centrality is not a static correlate of dominance rank as implied by the findings of short-term studies. Rather, centrality is a more sensitive indicator of status than is dominance rank, to which it is related in a dynamic fashion. Small changes in dominance rank may be followed by large changes in centrality. An increase in centrality may facilitate rise in dominance rank. These findings suggest a complex psychology of status, rather than a simple causative relation between the two variables.     

Winner and loser effects and the structure of dominance hierarchies

In the literature on dominance hierarchies, "winner" and "loser"effects usually are denned as an increased probability of winningat time T, bated on victories at time T-l, T-2, etc, and anincreased probability of losing at time T, based on losing atT-1, T-2, etc., respectively. Despite some early theoreticalwork on winner and loser effects, these factors and how theyaffect the structure of dominance hierarchies have not beenexamined in detail. I developed a computer simulation to examinewinner and loser effects when such effects are independent ofone another (as well as when they interact) and when combatantsassess each other's resource-holding power. When winner effectsalone were important, a hierarchy in which all individuals heldan unambiguous rank was found. When only loser effects wereimportant, a dear alpha individual always emerged, but the rankof others in the group was often unclear because of the scarcityof aggressive interactions. Increasing winner effects for agiven value of the loser effect increase the number of individualswith unambiguous positions in a hierarchy and the converse istrue for increasing the value of the loser effect for a givenwinner effect Although winner and loser effects have been documentedin a number of species, no study has documented both winnerand loser effects (using some controlled, pairwise testing system)and the detailed nature of behavioral interactions when individualsare in groups. I hope the results of this model will spur suchstudies in the future.     

Indirect genetic effects and the genetic bases of social dominance: evidence from cattle

Genetic studies of social behaviour have currently received new impetus from models including indirect genetic effects (IGEs) of social partners. This study aimed at investigating the contribution of conspecifics in social dominance, considered as response of dyadic interaction that is, winning (dominant individual) or losing (subordinate). A genetic correlation of −1 is expected between the attitude to win and the attitude to loose, and because a population always accounts for half winners and half losers, the heritability of the dominant status should be close to zero. Specifically, social dominance was studied in Aosta Chestnut and Aosta Black Pied (Bos taurus) breeds, alpine rustic cattle famous for traditional tournaments where pairs of cows assess dominant status in bloodless fights. The outcomes of 25 590 dyadic interactions performed by 8159 individuals in 11 years were analysed by applying a classical quantitative model and models including indirect effects. Data were analysed via Bayesian approach on a threshold trait. The assessment of variances revealed a genetic correlation of −0.976 between direct and indirect genetic components. The heritability measured on a liability scale was 0.122 for direct phenotype, but decreased to 0.014 when the total heritable variance (TBV) was considered. The trend of estimated breeding values showed that the total TBV was constant over the years, even though its direct component increased and the indirect part decreased. This result confirms the relevance of IGEs on social behaviour and the assumption that the mean individual social dominance cannot evolve within a population, due to the evolutionary constraints imposed by the ‘social environment''.     

Vocalizations of isolated piglets. II. Some environmental factors

The vocalizations of suckling piglets were studied during brief tests which involved social isolation, restriction of movement, and exposure to unfamiliar surroundings. Similar trends were shown in most cases by the three classes of call studied — closed mouth grunts, open mouth grunts, and squeals. The rate of calling was much lower when a piglet was in the home pen than when alone in an unfamiliar pen, and more calls were given in both situations when the animal's movement was restricted by caging. The presence of the dam and four littermates in unfamiliar surroundings caused a large reduction in calling, part of which was seen with only the dam present, and part with only the littermates. Provision of familiar bedding in the unfamiliar situation had no effect. Aspects of piglets' adaptation to environmental and social change are discussed.     

The structure of the incompatibility factors of Schizophyllum commune. Class II factors

Summary The possible causes for the lack of detectable recombinants of class II B factors were considered. The results of functional tests indicate that class II factors are constituted of two linked loci as all other B factors. The loci are very tightly linked and recombine at a frequency lower than 0.0002. The interrelations of the three classes of B factors are discussed.The research was supported by grants from the Atomic Energy Commission of the U.S. No. AT (30-1)-3875 and the National Institutes of Health No. AI 06124 during a post-doctoral training of the author at the Biological Laboratories, Harvard University, Cambridge, Mass., USA.     

Group size in folivorous primates: ecological constraints and the possible influence of social factors

The ecological-constraints model assumes that food items occur in depletable patches and proposes that an increase in group size leads to increased day range due to more rapid patch depletion. Smaller groups become advantageous when an increase in travel costs is not repaid by an increase in energy gained or some other fitness advantage. On the other hand, we also know that group size can be influenced by social factors. Here we contrast the diet and group size of red colobus (Procolobus badius) and black-and-white colobus (Colobus guereza) in Kibale National Park, Uganda to consider how ecological and social factors are affecting their group sizes. Subsequently, we examine whether the insights gained from this detailed comparison can provide an understanding of why the social organization and group size of mantled howlers (Alouatta palliata) and black howlers (A. pigra) differ. Two groups of red colobus and two groups of black-and-white colobus were studied over 10 months. Red colobus groups were larger (48 and 24) than black-and-white colobus groups (9 and 6). The two groups of red colobus overlap home ranges with the two groups of black-and-white colobus; 75% and 95% of their home ranges were within red colobuss home range. There was a great deal of similarity in the plant parts eaten by the two species and both species fed primarily on young leaves (red colobus 70%, black-and-white colobus 76%). In terms of the actual species consumed, again there was a great deal of similarity between species. The average dietary overlap among months for the two neighboring groups of red colobus was 37.3%, while the dietary overlap between the red colobus and the black-and-white colobus group that had its home range almost entirely within the home range of the red colobus groups averaged 43.2% among months. If ecological conditions were responsible for the difference in group size between the two colobine species, one would expect the density of food trees to be lower in the home ranges of the black-and-white colobus monkeys, since they have the smaller group size. We found the opposite to be true. Both black-and-white colobus groups had more food trees and the cumulative size of those trees was greater than those in the red colobuss home ranges. We quantify how these differences parallel differences in mantled and black howlers. The average group size for mantled howlers was 12.9 individuals, and for black howlers it was 5.3 individuals. We explore possible social constraints, such as infanticide, that prevent black-and-white colobus and black howlers from living in large groups.This revised version was published online in April 2005 with corrections to the cover date of the issue.     

Insect societies and the molecular biology of social behavior

This article outlines the rationale for a molecular genetic study of social behavior, and explains why social insects are good models. Summaries of research on brain and behavior in two species, honey bees and fire ants, are presented to illustrate the richness of the behavioral phenomena that can be addressed with social insects and to show how they are beginning to be used to study genes that influence social behavior. We conclude by considering the problems and potential of this emerging field.     

Crystal structure of Bacillus stearothermophilus alpha-amylase: possible factors determining the thermostability

The crystal structure of a thermostable alpha-amylase from Bacillus stearothermophilus (BSTA) has been determined at 2.0 A resolution. The main-chain fold is almost identical to that of the known crystal structure of Bacillus licheniformis alpha-amylase (BLA). BLA is known to be more stable than BSTA. A structural comparison between the crystal structures of BSTA and BLA showed significant differences that may account for the difference in their thermostabilities, as follows. (i) The two-residue insertion in BSTA, Ile181-Gly182, pushes away the spatially contacting region including Asp207, which corresponds to Ca(2+)-coordinating Asp204 in BLA. As a result, Asp207 cannot coordinate the Ca(2+). (ii) BSTA contains nine fewer hydrogen bonds than BLA, which costs about 12 kcal/mol. This tendency is prominent in the (beta/alpha)(8)-barrel, where 10 fewer hydrogen bonds were observed in BSTA. BLA forms a denser hydrogen bond network in the inter-helical region, which may stabilize alpha-helices in the barrel. (iii) A few small voids observed in the alpha-helical region of the (beta/alpha)(8)-barrel in BSTA decrease inter-helical compactness and hydrophobic interactions. (iv) The solvent-accessible surface area of charged residues in BLA is about two times larger than that in BSTA.     

Extrinsic effects, estimating opponents' RHP, and the structure of dominance hierarchies

We examined the impact of winner and loser effects on dominance hierarchy formation when individuals are capable of estimating their opponent's resource holding power (RHP). The accuracy of such estimates was a variable in our simulations, and we considered cases in which all individuals err within the same bounds, as well as cases in which some individuals consistently overestimate, while others consistently underestimate their opponent's fighting RHP. In all cases, we found a clearly defined linear hierarchy. In most simulations, the vast majority of interactions were 'attack-retreats', and the remainder of interactions were almost all 'fights'. Error rates had no effect on the linearity of the hierarchy or the basic attack-retreat nature of interactions, and consistent over and underestimation did not affect the ultimate position of an individual in a hierarchy.