A mixed strategy model for the emergence and intensification of social learning in a periodically changing natural environment

Based on a population genetic model of mixed strategies determined by alleles of small effect, we derive conditions for the evolution of social learning in an infinite-state environment that changes periodically over time. Each mixed strategy is defined by the probabilities that an organism will commit itself to individual learning, social learning, or innate behavior. We identify the convergent stable strategies (CSS) by a numerical adaptive dynamics method and then check the evolutionary stability (ESS) of these strategies. A strategy that is simultaneously a CSS and an ESS is called an attractive ESS (AESS). For certain parameter sets, a bifurcation diagram shows that the pure individual learning strategy is the unique AESS for short periods of environmental change, a mixed learning strategy is the unique AESS for intermediate periods, and a mixed learning strategy (with a relatively large social learning component) and the pure innate strategy are both AESS's for long periods. This result entails that, once social learning emerges during a transient era of intermediate environmental periodicity, a subsequent elongation of the period may result in the intensification of social learning, rather than a return to innate behavior.     

Evolution of learning capacities and learning levels

Humans strongly depend on individual and social learning, both of which are highly effective and accurate. I study the effects of environmental change on the evolution of the effectiveness and accuracy of individual and social learning (individual and social learning levels) and the number of pieces of information learned individually and socially (individual and social learning capacities) by analyzing a mathematical model. I show that individual learning capacity decreases and social learning capacity increases when the environment becomes more stable; both decrease when the environment becomes milder. I also show that individual learning capacity increases when individual learning level increases or social learning level decreases, while social learning capacity increases when individual or social learning level increases. The evolution of high learning levels can be triggered when the environment becomes severe, but a high social learning level can evolve only when a high individual learning level can simultaneously evolve with it.     

The evolution of social learning rules: Payoff-biased and frequency-dependent biased transmission

Humans and other animals do not use social learning indiscriminately, rather, natural selection has favoured the evolution of social learning rules that make selective use of social learning to acquire relevant information in a changing environment. We present a gene-culture coevolutionary analysis of a small selection of such rules (unbiased social learning, payoff-biased social learning and frequency-dependent biased social learning, including conformism and anti-conformism) in a population of asocial learners where the environment is subject to a constant probability of change to a novel state. We define conditions under which each rule evolves to a genetically polymorphic equilibrium. We find that payoff-biased social learning may evolve under high levels of environmental variation if the fitness benefit associated with the acquired behaviour is either high or low but not of intermediate value. In contrast, both conformist and anti-conformist biases can become fixed when environment variation is low, whereupon the mean fitness in the population is higher than for a population of asocial learners. Our examination of the population dynamics reveals stable limit cycles under conformist and anti-conformist biases and some highly complex dynamics including chaos. Anti-conformists can out-compete conformists when conditions favour a low equilibrium frequency of the learned behaviour. We conclude that evolution, punctuated by the repeated successful invasion of different social learning rules, should continuously favour a reduction in the equilibrium frequency of asocial learning, and propose that, among competing social learning rules, the dominant rule will be the one that can persist with the lowest frequency of asocial learning.     

The evolution of fitness in life-history theory

Theory concerning the evolution of life history (the schedule of reproduction and survival) focuses on describing the life history which maximises fitness. Although there is an intuitive link between life history and fitness, there are in fact several measures of the 'black box' concept of fitness. There has been a debate in the bio-mathematical literature on the predictive difference between the two most commonly used measures; intrinsic rate of increase r and net reproductive ratio R0. Although both measures aim to describe fitness, models using one of the measures may predict the opposite of similar models using the other measure, which is clearly undesirable. Here, I review the evolution of these fitness measures over the last four decades, the predictive differences between these measures and the resulting shift of the fitness concept. I focus in particular on some recent developments, which have solved the dilemma of predictive differences between these fitness measures by explicitly acknowledging the game-theoretical nature of life-history evolution.     

Higher frequency of social learning in China than in the West shows cultural variation in the dynamics of cultural evolution

Cultural evolutionary models have identified a range of conditions under which social learning (copying others) is predicted to be adaptive relative to asocial learning (learning on one''s own), particularly in humans where socially learned information can accumulate over successive generations. However, cultural evolution and behavioural economics experiments have consistently shown apparently maladaptive under-utilization of social information in Western populations. Here we provide experimental evidence of cultural variation in people''s use of social learning, potentially explaining this mismatch. People in mainland China showed significantly more social learning than British people in an artefact-design task designed to assess the adaptiveness of social information use. People in Hong Kong, and Chinese immigrants in the UK, resembled British people in their social information use, suggesting a recent shift in these groups from social to asocial learning due to exposure to Western culture. Finally, Chinese mainland participants responded less than other participants to increased environmental change within the task. Our results suggest that learning strategies in humans are culturally variable and not genetically fixed, necessitating the study of the ‘social learning of social learning strategies'' whereby the dynamics of cultural evolution are responsive to social processes, such as migration, education and globalization.     

The genetic mating system of spotted sunfish (Lepomis punctatus): mate numbers and the influence of male reproductive parasites

In nest-building fish species, mature males often exhibit one of two alternative reproductive behaviours. Bourgeois males build nests, court females, and guard their eggs. Parasitic cuckolders attempt to steal fertilizations from bourgeois males and do not invest in parental care. Previous evidence from the bluegill sunfish (Lepomis macrochirus) suggests that adult males are morphologically specialized for these two tactics. Here, we used microsatellite markers to determine genetic parentage in a natural population of the spotted sunfish (L. punctatus) that also displayed both bourgeois and parasitic male morphs. As gauged by relative investments in gonadal vs. somatic tissues, between 5 and 15% of the mature adult males were parasites. Multi-locus genotypes were generated for more than 1400 embryos in 30 nests, their nest-guardian males, and for other adults in the population. Progeny in approximately 57% of the nests were sired exclusively by the guardian male, but the remaining nests contained embryos resulting from cuckoldry as well. Overall, the frequency of offspring resulting from stolen fertilizations was only 1.3%, indicating that the great majority of paternity is by bourgeois nesting males. With regard to maternity, 87% of the nests had at least three dams, and computer simulations estimate that about 7.2 dams spawned per nest.     

The sky is falling: evidence of a negativity bias in the social transmission of information

The method of serial reproduction has revealed that the social transmission of information is characterized by the gradual transformation of the original message. This transformation results from the preferential survival of certain types of information and the resolution of ambiguity. Here we present evidence of a bias favoring the social transmission of negatively-valenced information across multiple transmission episodes. Ninety-two, four-person chains transmitted a story containing unambiguously positive and unambiguously negative story events, along with ambiguous story events that could be interpreted positively or negatively. Analysis using mixed-effects modeling revealed the preferential survival of unambiguously negative events over positive events, and the increasingly negative resolution of ambiguous events across successive transmission episodes. Contrary to predictions, elevated state anxiety did not enhance the social transmission of negatively-valenced information. We also found that the survival of unambiguously negative story events was positively correlated with the negative resolution of ambiguous story events, reflecting a general negativity-bias in the social transmission of information.     

成年斑胸草雀在体HVC-RA突触传递的电生理特性

鸣禽高级发声中枢（high vocal center,HVC）至弓状皮质栎核（robust nucleus ofthe arcopallium,RA）的突触传递是鸣唱运动通路中的关键部分.本文运用在体场电位电生理记录的方法,研究了成年雄性斑胸草雀（Taeniopygia guttata）HVC-RA突触的电生理特性.实验结果显示,刺激HVC,在RA内所记录到的诱发场电位幅度较小.配对脉冲检测发现,HVC-RA突触传递具有明显的配对脉冲易化特性.当以强直刺激作用于HVC,RA内诱发场电位随即显著减小,并在15 min内逐渐恢复,表明HVC-RA突触传递在强直刺激过后出现了短时抑制.该通路的突触传递特性可能与其在发声控制中的作用有关.以上的实验结果为进一步研究发声运动过程中的突触可塑性提供了资料.