Social traits,social networks and evolutionary biology

The social environment is both an important agent of selection for most organisms, and an emergent property of their interactions. As an aggregation of interactions among members of a population, the social environment is a product of many sets of relationships and so can be represented as a network or matrix. Social network analysis in animals has focused on why these networks possess the structure they do, and whether individuals’ network traits, representing some aspect of their social phenotype, relate to their fitness. Meanwhile, quantitative geneticists have demonstrated that traits expressed in a social context can depend on the phenotypes and genotypes of interacting partners, leading to influences of the social environment on the traits and fitness of individuals and the evolutionary trajectories of populations. Therefore, both fields are investigating similar topics, yet have arrived at these points relatively independently. We review how these approaches are diverged, and yet how they retain clear parallelism and so strong potential for complementarity. This demonstrates that, despite separate bodies of theory, advances in one might inform the other. Techniques in network analysis for quantifying social phenotypes, and for identifying community structure, should be useful for those studying the relationship between individual behaviour and group‐level phenotypes. Entering social association matrices into quantitative genetic models may also reduce bias in heritability estimates, and allow the estimation of the influence of social connectedness on trait expression. Current methods for measuring natural selection in a social context explicitly account for the fact that a trait is not necessarily the property of a single individual, something the network approaches have not yet considered when relating network metrics to individual fitness. Harnessing evolutionary models that consider traits affected by genes in other individuals (i.e. indirect genetic effects) provides the potential to understand how entire networks of social interactions in populations influence phenotypes and predict how these traits may evolve. By theoretical integration of social network analysis and quantitative genetics, we hope to identify areas of compatibility and incompatibility and to direct research efforts towards the most promising areas. Continuing this synthesis could provide important insights into the evolution of traits expressed in a social context and the evolutionary consequences of complex and nuanced social phenotypes.     

Social features of online networks: the strength of intermediary ties in online social media

An increasing fraction of today's social interactions occur using online social media as communication channels. Recent worldwide events, such as social movements in Spain or revolts in the Middle East, highlight their capacity to boost people's coordination. Online networks display in general a rich internal structure where users can choose among different types and intensity of interactions. Despite this, there are still open questions regarding the social value of online interactions. For example, the existence of users with millions of online friends sheds doubts on the relevance of these relations. In this work, we focus on Twitter, one of the most popular online social networks, and find that the network formed by the basic type of connections is organized in groups. The activity of the users conforms to the landscape determined by such groups. Furthermore, Twitter's distinction between different types of interactions allows us to establish a parallelism between online and offline social networks: personal interactions are more likely to occur on internal links to the groups (the weakness of strong ties); events transmitting new information go preferentially through links connecting different groups (the strength of weak ties) or even more through links connecting to users belonging to several groups that act as brokers (the strength of intermediary ties).     

Facultative development of courtship and communication in juvenile male cowbirds (Molothrus ater)

We investigated effects of naturally occurring variation inexperience with adult males on development of song and courtshipcompetence in captive juvenile cowbirds. We studied birds ingroups housed in large outdoor aviaries that allowed birdsto regulate access to social stimulation. In two aviaries, we housed juvenile males and females either with or withoutadult males. Birds remained in these conditions from September1999 through their breeding season. We documented social andvocal development of juvenile males in the two aviaries bymeasuring social assortment and patterns and frequencies of their song interactions. We then brought the juveniles fromthe two aviaries together to compete against each other foraccess to females. In addition, we recorded juveniles' songsfour times over the study and played back their breeding seasonsongs to females in sound-attenuating chambers to measure the effectiveness of songs in eliciting copulatory responses fromthe females. Compared to juvenile males housed with adult males,juvenile males housed without adult males developed atypicalbehavior patterns. They (1) displayed little intrasexual aggressionor near-neighbor associations and (2) exhibited different patternsof courtship and copulation, but (3) were as successful at competing for copulations. Furthermore, they developed stereotypedsongs sooner and developed more potent breeding season songs.These different outcomes could not be traced to one variablebut to a cascade of effects involving diverging patterns ofsong acquisition and social interaction. The patterns of socialskills that emerged indicate considerable plasticity in the mechanisms underlying acquisition of courtship competence.     

The context of social learning: association patterns in a captive flock of brown-headed cowbirds

Much work on social learning has involved behaviour transmission between pairs of individuals, but recently the need to examine the social context in which learning occurs has been recognized. Previous studies using small numbers of animals have shown social influence on the behavioural development of juvenile male brown-headed cowbirds, Molothrus ater. Here we looked at the larger social context that forms the framework for such influence in more natural settings. We allowed a captive group of over 70 cowbirds, comprising adult and juvenile males and females, to associate freely in a large complex of connected aviaries. Highly organized social assortment emerged in the group, with individuals associating with others based on similarity in age and sex. Juvenile males that associated more with adult males had higher courtship success. Juvenile males that associated more with females sang less over the year. These results indicate that the social context of social learning for juvenile males is not just random association with all other birds in the social group, but is a selective and structured pattern of interaction. Differences in navigating this social structure correlated with courtship success and vocalization, behaviour known to be affected by social learning. Studies such as this, using large groups with free assortment of individuals, are the first step towards understanding the effects of the larger social context surrounding social learning.     

Eggshell strength in cuckoos and cowbirds

Parasitic cuckoos of the genus Clamator and parasitic cowbirds Molothrus spp. lay eggs that appear to be stronger than those of their hosts. Similar differences in eggshell strength between the eggs of the Cuculus/Cacomantis/Chrysococcyx cuckoos and their hosts are not apparent. We suggest that the advantage of a strong eggshell in the Clamator and Molothrus species is to protect the parasite's egg from damage if the nest is multiply parasitized. This suggestion relies on the observation that Clamator and Molothrus species commonly lay more than one egg per nest and often remove or damage eggs in the process.     

Personality in the context of social networks

There is great interest in environmental effects on the development and evolution of animal personality traits. An important component of an individual's environment is its social environment. However, few studies look beyond dyadic relationships and try to place the personality of individuals in the context of a social network. Social network analysis provides us with many new metrics to characterize the social fine-structure of populations and, therefore, with an opportunity to gain an understanding of the role that different personalities play in groups, communities and populations regarding information or disease transmission or in terms of cooperation and policing of social conflicts. The network position of an individual is largely a consequence of its interactive strategies. However, the network position can also shape an individual's experiences (especially in the case of juveniles) and therefore can influence the way in which it interacts with others in future. Finally, over evolutionary time, the social fine-structure of animal populations (as quantified by social network analysis) can have important consequences for the evolution of personalities-an approach that goes beyond the conventional game-theoretic analyses that assumed random mixing of individuals in populations.     

The influence of social cues on the reproductive endocrinology of male brown-headed cowbirds: field and laboratory studies

Captive male brown-headed cowbirds exposed to long days exhibit gonadal growth and have elevated plasma testosterone (T) levels. This photoperiodic response is enhanced if males are housed with female cowbirds: Photostimulated males with females increase plasma testosterone levels sooner than do individually housed photostimulated males. Peak plasma T levels are similar in both groups, although peak levels are maintained longer in males housed with females. The gonadal cycle is similarly affected; males in the presence of females have earlier gonadal recrudescence and maintain mature gonads longer than do photostimulated males without females. Plasma corticosterone levels increase in the unpaired males, suggesting that removal of social cues is stressful for these birds. Free-living paired males have significantly higher plasma testosterone levels than do unpaired/unknown males early in the season, when social relationships are being established; the levels are similar thereafter. There is no difference between the two groups in testicular maturation rates; nor do they differ in plasma corticosterone levels at any time of the season. These results suggest that social stimuli are important in modulating the secretion of testosterone in males early in the season when pairing occurs, and possibly late in the season as well, probably to prevent termination of breeding prior to that of females.     

Social networks in fish

Social network theory underwent rapid development in the 1970s by sociologists and was recently further refined by physicists. Network theory has been applied to systems consisting of interconnected components as different as that of actors in Hollywood, the neural network of the nematode worm Caenorhabditis elegans, and the US national power grid. Network theory has great potential as a conceptual tool, for example, it can be used to provide quantitative predictions regarding information transmission, spreading of diseases, and the potential for establishing reciprocal altruistic relationships. Conventionally behavioural studies have been restricted to the analysis of social interactions between isolated pairs or small groups of individuals. Here we built up, however, from pair‐wise interactions, the social network of an entire population of guppies (Poecilia reticulata) in Trinidad. We show that guppies have highly structured social networks with significant repeated interactions between male–male, female–female and male–female pairs. Furthermore, sexual segregation within the network is positively correlated with the strength of the social interactions. Finally, individuals were observed to be more variable in their inter‐sexual interactions than their intra‐sexual ones.     

Trust transitivity in social networks

Non-centralized recommendation-based decision making is a central feature of several social and technological processes, such as market dynamics, peer-to-peer file-sharing and the web of trust of digital certification. We investigate the properties of trust propagation on networks, based on a simple metric of trust transitivity. We investigate analytically the percolation properties of trust transitivity in random networks with arbitrary in/out-degree distributions, and compare with numerical realizations. We find that the existence of a non-zero fraction of absolute trust (i.e. entirely confident trust) is a requirement for the viability of global trust propagation in large systems: The average pair-wise trust is marked by a discontinuous transition at a specific fraction of absolute trust, below which it vanishes. Furthermore, we perform an extensive analysis of the Pretty Good Privacy (PGP) web of trust, in view of the concepts introduced. We compare different scenarios of trust distribution: community- and authority-centered. We find that these scenarios lead to sharply different patterns of trust propagation, due to the segregation of authority hubs and densely-connected communities. While the authority-centered scenario is more efficient, and leads to higher average trust values, it favours weakly-connected "fringe" nodes, which are directly trusted by authorities. The community-centered scheme, on the other hand, favours nodes with intermediate in/out-degrees, in detriment of the authorities and its "fringe" peers.     

Detecting social transmission in networks

In recent years researchers have drawn attention to a need for new methods with which to identify the spread of behavioural innovations through social transmission in animal populations. Network-based analyses seek to recognise diffusions mediated by social learning by detecting a correspondence between patterns of association and the flow of information through groups. Here we introduce a new order of acquisition diffusion analysis (OADA) and develop established time of acquisition diffusion analysis (TADA) methods further. Through simulation we compare the merits of these and other approaches, demonstrating that OADA and TADA have greater power and lower Type I error rates than available alternatives, and specifying when each approach should be deployed. We illustrate the new methods by applying them to reanalyse an established dataset corresponding to the diffusion of foraging innovations in starlings, where OADA and TADA detect social transmission that hitherto had been missed. The methods are potentially widely applicable by researchers wishing to detect social learning in natural and captive populations of animals, and to facilitate this we provide code to implement OADA and TADA in the statistical package R.     

Cultures, genes, and neurons in the development of song and singing in brown-headed cowbirds (Molothrus ater)

In brown-headed cowbirds, Molothrus ater, as in many songbird species, vocalizations are fundamental to reproduction. In our studies, experiments utilizing different social housing regimes and geographic comparisons have indicated the social learning of males' vocalizations and associated abilities to use vocalizations effectively during the breeding season. Here, we describe studies indicating roles of cultural and genetic background, and of social influences from females, on male vocal development. These influences can interact with neural regions, including song learning and song control nuclei, but also visual-processing nuclei, in the development of signaling. We argue that a developmental systems approach to the study of vocal behavior provides a structure to organize these different influences and how they may interact with one another over development. A systems approach requires that researchers study the social context in which signals and signalers develop - both the ontogenetic arena in which young animals learn their signals from older animals, and the functional arena in which young and older animals socially interact with one another.