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Observational learning is a form of social learning in which a demonstrator performs a target task in the company of an observer, who may as a consequence learn something about it. In this study, we approach social learning in terms of the dynamics of coordination rather than the more common perspective of transmission of information. We hypothesised that observers must continuously adjust their visual attention relative to the demonstrator's time-evolving behaviour to benefit from it. We eye-tracked observers repeatedly watching videos showing a demonstrator solving one of three manipulative puzzles before attempting at the task. The presence of the demonstrator's face and the availability of his verbal instruction in the videos were manipulated. We then used recurrence quantification analysis to measure the dynamics of coordination between the overt attention of the observers and the demonstrator's manipulative actions. Bayesian hierarchical logistic regression was applied to examine (1) whether the observers' performance was predicted by such indexes of coordination, (2) how performance changed as they accumulated experience, and (3) if the availability of speech and intentional gaze of the demonstrator mediated it. Results showed that learners better able to coordinate their eye movements with the manipulative actions of the demonstrator had an increasingly higher probability of success in solving the task. The availability of speech was beneficial to learning, whereas the presence of the demonstrator's face was not. We argue that focusing on the dynamics of coordination between individuals may greatly improve understanding of the cognitive processes underlying social learning., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier B.V. All rights reserved.)

As a form of adaptive plasticity that allows organisms to shift their phenotype toward the optimum, learning is inherently a source of developmental bias. Learning may be of particular significance to the evolutionary biology community because it allows animals to generate adaptively biased novel behavior tuned to the environment and, through social learning, to propagate behavioral traits to other individuals, also in an adaptively biased manner. We describe several types of developmental bias manifest in learning, including an adaptive bias, historical bias, origination bias, and transmission bias, stressing that these can influence evolutionary dynamics through generating nonrandom phenotypic variation and/or nonrandom environmental states. Theoretical models and empirical data have established that learning can impose direction on adaptive evolution, affect evolutionary rates (both speeding up and slowing down responses to selection under different conditions) and outcomes, influence the probability of populations reaching global optimum, and affect evolvability. Learning is characterized by highly specific, path-dependent interactions with the (social and physical) environment, often resulting in new phenotypic outcomes. Consequently, learning regularly introduces novelty into phenotype space. These considerations imply that learning may commonly generate plasticity first evolution., (© 2019 Wiley Periodicals, Inc.)

I introduce seven criteria for determining the validity of competing theories for the original function of language. I go on to present a novel explanation that meets all the criteria: language originally evolved to teach kin. I suggest that the use of symbols subsequently generated evolutionary feedback at two levels, in the form of self-modified selection pressures that favored structures in the mind that functioned to manipulate and use symbols with efficiency, and cultural selection on languages for learnability.

The directed social learning hypothesis suggests that information does not spread evenly through animal groups, but rather individual characteristics and patterns of physical proximity guide the social transmission of information along specific pathways. Network-based diffusion analysis (NBDA) allows researchers to test whether information spreads following a social network. However, the explanatory power of different social networks has rarely been compared, and current models do not easily accommodate random effects (e.g. allowing for individuals within groups to correlate in their asocial solving rates). We tested whether the spread of two novel foraging skills through captive starling groups was affected by individual- and group-level random and fixed effects (i.e. sex, age, body condition, dominance rank and demonstrator status) and perching or foraging networks. We extended NBDA to include random effects and conducted model discrimination in a Bayesian context. We found that social learning increased the rate at which birds acquired the novel foraging task solutions by 6.67 times, and acquiring one of the two novel foraging task solutions facilitated the asocial acquisition of the other. Surprisingly, the spread of task solutions followed the perching rather than the foraging social network. Upon acquiring a task solution, foraging performance was facilitated by the presence of group mates. Our results highlight the importance of considering more than one social network when predicting the spread of information through animal groups. This article is part of a Special Issue entitled: Cognition in the wild., (Copyright © 2014 Elsevier B.V. All rights reserved.)

Social learning mechanisms are widely thought to vary in their degree of complexity as well as in their prevalence in the natural world. While learning the properties of a stimulus that generalize to similar stimuli at other locations (stimulus enhancement) prima facie appears more useful to an animal than learning about a specific stimulus at a specific location (local enhancement), empirical evidence suggests that the latter is much more widespread in nature. Simulating populations engaged in a producer-scrounger game, we sought to deploy mathematical models to identify the adaptive benefits of reliance on local enhancement and/or stimulus enhancement, and the alternative conditions favoring their evolution. Surprisingly, we found that while stimulus enhancement readily evolves, local enhancement is advantageous only under highly restricted conditions: when generalization of information was made unreliable or when error in social learning was high. Our results generate a conundrum over how seemingly conflicting empirical and theoretical findings can be reconciled. Perhaps the prevalence of local enhancement in nature is due to stimulus enhancement costs independent of the learning task itself (e.g. predation risk), perhaps natural habitats are often characterized by unreliable yet highly rewarding payoffs, or perhaps local enhancement occurs less frequently, and stimulus enhancement more frequently, than widely believed., (Copyright © 2013 Elsevier Inc. All rights reserved.)

Organisms often respond to environmental change phenotypically, through learning strategies that enhance fitness in variable and changing conditions. But which strategies should we expect in population exposed to those conditions? We address this question by developing a mathematical model that specifies the consequences of different mixtures of individual and social learning strategies on the frequencies of different cultural variants in temporally and spatially changing environments. Assuming that alternative cultural variants are differently well-adapted to diverse environmental conditions, we are able to evaluate which mixture of learning strategies maximises the mean fitness of the population. We find that, even in rapidly changing environments, a high proportion of the population will always engage in social learning. In those environments, the highest adaptation levels are achieved through relatively high fractions of individual learning and a strong conformist bias. We establish a negative relationship between the proportion of the population learning socially and the strength of conformity operating in a population: strong conformity requires fewer conformists (i.e. larger proportion of individual learning), while many conformists can only be found when conformist transmission is weak. Investigations of cultural diversity show that in frequently changing environments high levels of adaptation require high level of cultural diversity. Finally, we demonstrate how the developed mathematical framework can be applied to time series of usage or occurrence data of cultural traits. Using Approximate Bayesian Computation we are able to infer information about the underlying learning processes that could have produced observed patterns of variation in the dataset., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

Many animals have socially transmitted behavioural traditions, but human culture appears unique in that it is cumulative, i.e. human cultural traits increase in diversity and complexity over time. It is often suggested that high-fidelity cultural transmission is necessary for cumulative culture to occur through refinement, a process known as 'ratcheting', but this hypothesis has never been formally evaluated. We discuss processes of information transmission and loss of traits from a cognitive viewpoint alongside other cultural processes of novel invention (generation of entirely new traits), modification (refinement of existing traits) and combination (bringing together two established traits to generate a new trait). We develop a simple cultural transmission model that does not assume major evolutionary changes (e.g. in brain architecture) and show that small changes in the fidelity with which information is passed between individuals can lead to cumulative culture. In comparison, modification and combination have a lesser influence on, and novel invention appears unimportant to, the ratcheting process. Our findings support the idea that high-fidelity transmission is the key driver of human cumulative culture, and that progress in cumulative culture depends more on trait combination than novel invention or trait modification.

The demographic and ecological success of our species is frequently attributed to our capacity for cumulative culture. However, it is not yet known how humans combine social and asocial learning to generate effective strategies for learning in a cumulative cultural context. Here we explore how cumulative culture influences the relative merits of various pure and conditional learning strategies, including pure asocial and social learning, critical social learning, conditional social learning and individual refiner strategies. We replicate the Rogers' paradox in the cumulative setting. However, our analysis suggests that strategies that resolved Rogers' paradox in a non-cumulative setting may not necessarily evolve in a cumulative setting, thus different strategies will optimize cumulative and non-cumulative cultural learning., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

Vigorous debates as to the evolutionary origins of culture remain unresolved due to an absence of methods for identifying learning mechanisms in natural populations. While laboratory experiments on captive animals have revealed evidence for a number of mechanisms, these may not necessarily reflect the processes typically operating in nature. We developed a novel method that allows social and asocial learning mechanisms to be determined in animal groups from the patterns of interaction with, and solving of, a task. We deployed it to analyse learning in groups of wild meerkats (Suricata suricatta) presented with a novel foraging apparatus. We identify nine separate learning processes underlying the meerkats' foraging behaviour, in each case precisely quantifying their strength and duration, including local enhancement, emulation, and a hitherto unrecognized form of social learning, which we term 'observational perseverance'. Our analysis suggests a key factor underlying the stability of behavioural traditions is a high ratio of specific to generalized social learning effects. The approach has widespread potential as an ecologically valid tool to investigate learning mechanisms in natural groups of animals, including humans.

Controversy over claims of cultures in nonhuman primates and other animals has led to a call for quantitative methods that are able to infer social learning from freely interacting groups of animals. Network-based diffusion analysis (NBDA) is such a method that infers social transmission of a behavioral trait when the pattern of acquisition follows the social network. As, relative to other animals, primates may be unusual in their heavy reliance on social learning, with learning frequently directed along pathways of association; in this study, we draw attention to the significance of this method for primatologists. We provide a "users guide" to NBDA methodology, discussing the choice of NBDA model and social network, and suggest model selection procedures. We also present the results of simulations that suggest that NBDA works well even when the assumptions of the underlying model are violated., (© 2011 Wiley-Liss, Inc.)

There are consistent individual differences in human intelligence, attributable to a single 'general intelligence' factor, g. The evolutionary basis of g and its links to social learning and culture remain controversial. Conflicting hypotheses regard primate cognition as divided into specialized, independently evolving modules versus a single general process. To assess how processes underlying culture relate to one another and other cognitive capacities, we compiled ecologically relevant cognitive measures from multiple domains, namely reported incidences of behavioural innovation, social learning, tool use, extractive foraging and tactical deception, in 62 primate species. All exhibited strong positive associations in principal component and factor analyses, after statistically controlling for multiple potential confounds. This highly correlated composite of cognitive traits suggests social, technical and ecological abilities have coevolved in primates, indicative of an across-species general intelligence that includes elements of cultural intelligence. Our composite species-level measure of general intelligence, 'primate g(S)', covaried with both brain volume and captive learning performance measures. Our findings question the independence of cognitive traits and do not support 'massive modularity' in primate cognition, nor an exclusively social model of primate intelligence. High general intelligence has independently evolved at least four times, with convergent evolution in capuchins, baboons, macaques and great apes.

Research into social learning (learning from others) has expanded significantly in recent years, not least because of productive interactions between theoretical and empirical approaches. This has been coupled with a new emphasis on learning strategies, which places social learning within a cognitive decision-making framework. Understanding when, how and why individuals learn from others is a significant challenge, but one that is critical to numerous fields in multiple academic disciplines, including the study of social cognition., (Copyright © 2010 Elsevier Ltd. All rights reserved.)

Theoretical analyses have reported that in most circumstances where natural selection favours reliance on social learning, conformity (positive frequency-dependent social learning) is also favoured. These findings suggest that much animal social learning should involve a copy-the-majority strategy, yet there is currently surprisingly little evidence for conformist learning among animals. Here, we investigate this possibility in the nine-spined stickleback (Pungitius pungitius) by manipulating the number of demonstrator fish at two feeders, one rich and one poor, during a demonstration phase and evaluating how this affects the likelihood that the focal fish copy the demonstrators' apparent choices. As predicted, we observed a significantly increased level of copying with increasing numbers of demonstrators at the richer of the two feeders, with copying increasing disproportionately, rather than linearly, with the proportion of demonstrators at the rich foraging patch. Control conditions with non-feeding demonstrators showed that this was not simply the result of a preference for shoaling with larger groups, implying that nine-spined sticklebacks copy in a conformist manner.

Theoretical analyses within the broad field of social learning research give mixed conclusions on whether the shape of a diffusion curve can be used to infer that a learned trait increases through social or asocial learning. Here we explore how factors such as task structure (e.g., multiple-step tasks), task abandonment, subgoal learning, and neophobia affect the shape of the diffusion curve for both asocially learned and socially learned behavior. We demonstrate that, whereas social learning increases the likelihood of S-shaped curves, sigmoidal patterns can be generated by entirely asocial processes, and cannot be reliably interpreted as indicators of social learning. Our findings reinforce the view that diffusion curve analysis is not a reliable way of detecting social transmission. We also draw attention to the fact that task structure can similarly confound interpretation of network-based diffusion analyses, and suggest resolutions to this problem. Supplemental materials for this article may be downloaded from http://lb.psychonomic-journals.org/content/supplemental.

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., ((c) 2010 Elsevier Ltd. All rights reserved.)

We explore the evolution of reliance on social and asocial learning using a spatially explicit stochastic model. Our analysis considers the relative merits of four evolved strategies, two pure strategies (asocial and social learning) and two conditional strategies (the "critical social learner," which learns asocially only when copying fails, and the "conditional social learner," which copies only when asocial learning fails). We find that spatial structure generates outcomes that do not always conform to the finding of earlier theoretical analyses that social learning does not enhance average individual fitness at equilibrium (Rogers' paradox). Although we describe circumstances under which the strategy of pure social learning increases the average fitness of individuals, we find that spatial structure introduces a new paradox, which is that social learning can spread even when it decreases the average fitness of individuals below that of asocial learners. We also show that the critical social learner and conditional social learner both provide solutions to the aforementioned paradoxes, although we find some conditions in which pure (random) social learning out-competes both conditional strategies. Finally, we consider the relative merits of critical and conditional social learning under various conditions.

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.

Background: Studies of natural animal populations reveal widespread evidence for the diffusion of novel behaviour patterns, and for intra- and inter-population variation in behaviour. However, claims that these are manifestations of animal 'culture' remain controversial because alternative explanations to social learning remain difficult to refute. This inability to identify social learning in social settings has also contributed to the failure to test evolutionary hypotheses concerning the social learning strategies that animals deploy., Methodology/principal Findings: We present a solution to this problem, in the form of a new means of identifying social learning in animal populations. The method is based on the well-established premise of social learning research, that--when ecological and genetic differences are accounted for--social learning will generate greater homogeneity in behaviour between animals than expected in its absence. Our procedure compares the observed level of homogeneity to a sampling distribution generated utilizing randomization and other procedures, allowing claims of social learning to be evaluated according to consensual standards. We illustrate the method on data from groups of monkeys provided with novel two-option extractive foraging tasks, demonstrating that social learning can indeed be distinguished from unlearned processes and a social learning, and revealing that the monkeys only employed social learning for the more difficult tasks. The method is further validated against published datasets and through simulation, and exhibits higher statistical power than conventional inferential statistics., Conclusions/significance: The method is potentially a significant technological development, which could prove of considerable value in assessing the validity of claims for culturally transmitted behaviour in animal groups. It will also be of value in enabling investigation of the social learning strategies deployed in captive and natural animal populations.

Many species are known to acquire valuable life skills and information from others, but until recently it was widely believed that animals did not actively facilitate learning in others. Teaching was regarded as a uniquely human faculty. However, recent studies suggest that teaching might be more common in animals than previously thought. Teaching is present in bees, ants, babblers, meerkats and other carnivores but is absent in chimpanzees, a bizarre taxonomic distribution that makes sense if teaching is treated as a form of altruism. Drawing on both mechanistic and functional arguments, we integrate teaching with the broader field of animal social learning, and show how this aids understanding of how and why teaching evolved, and the diversity of teaching mechanisms.

In most studies of social learning in animals, no attempt has been made to examine the nature of the strategy adopted by animals when they copy others. Researchers have expended considerable effort in exploring the psychological processes that underlie social learning and amassed extensive data banks recording purported social learning in the field, but the contexts under which animals copy others remain unexplored. Yet, theoretical models used to investigate the adaptive advantages of social learning lead to the conclusion that social learning cannot be indiscriminate and that individuals should adopt strategies that dictate the circumstances under which they copy others and from whom they learn. In this article, I discuss a number of possible strategies that are predicted by theoretical analyses, including copy when uncertain, copy the majority, and copy if better, and consider the empirical evidence in support of each, drawing from both the animal and human social learning literature. Reliance on social learning strategies may be organized hierarchically, their being employed by animals when unlearned and asocially learned strategies prove ineffective but before animals take recourse in innovation.

Although it is natural to expect that group-living animals will utilize social learning, the expectation for non-grouping species is less clear. Only a few studies have explored the relationship between sociality and social learning. Here we presented 4 non-grouping fish species, fifteenspine sticklebacks (Spinachia spinachia), bullhead sculpins (Cottus gobio), stone loach (Barbatula barbatula) and juvenile European flounders (Platichthys flesus) with social information provided by groups of a shoal-forming heterospecific, the threespine stickleback (Gasterosteus aculeatus). Using a binary choice procedure we allowed individual test subjects to select between simulated prey patches. Although the test subjects could not sample the patches directly they were able to use information generated by the heterospecific demonstrators to select the "richer" of the 2 patches. For comparison we also recorded social information use in 2 shoaling species, threespine, and ninespine sticklebacks (Pungitius pungitius). We saw evidence of social information use and social learning in all 6 species, with no differences seen between social and non-grouping species. We argue that social learning is not likely to be restricted to group-living species, since many solitary species too are regularly exposed to social stimuli from both conspecifics and heterospecifics, and can benefit from using social information. We suggest that researchers have much to learn about the sensory, perceptive, and cognitive mechanisms underlying social learning, and the extent to which these vary (if at all) between grouping and non-grouping species. [ABSTRACT FROM AUTHOR]

Background: Social learning is potentially advantageous, but evolutionary theory predicts that (i) its benefits may be self-limiting because social learning can lead to information parasitism, and (ii) these limitations can be mitigated via forms of selective copying. However, these findings arise from a functional approach in which learning mechanisms are not specified, and which assumes that social learning avoids the costs of asocial learning but does not produce information about the environment. Whether these findings generalize to all kinds of social learning remains to be established. Using a detailed multi-scale evolutionary model, we investigate the payoffs and information production processes of specific social learning mechanisms (including local enhancement, stimulus enhancement and observational learning) and their evolutionary consequences in the context of skill learning in foraging groups. Results: We find that local enhancement does not benefit foraging success, but could evolve as a side-effect of grouping. In contrast, stimulus enhancement and observational learning can be beneficial across a wide range of environmental conditions because they generate opportunities for new learning outcomes. Conclusions: In contrast to much existing theory, we find that the functional outcomes of social learning are mechanism specific. Social learning nearly always produces information about the environment, and does not always avoid the costs of asocial learning or support information parasitism. Our study supports work emphasizing the value of incorporating mechanistic detail in functional analyses. [ABSTRACT FROM AUTHOR]

Niche construction is the modification of components of the environment through an organism's activities. Humans modify their environments mainly through ontogenetic and cultural processes, and it is this reliance on learning, plasticity and culture that lends human niche construction a special potency. In this paper we aim to facilitate discussion between researchers interested in niche construction and those interested in human cognitive development by highlighting some of the related processes. We discuss the transmission of culturally relevant information, how the human mind is a symbol-generating and artefact-devising system, and how these processes are bi-directional, with infants and children both being directed, and directing, their own development. We reflect on these in the light of four approaches: natural pedagogy, activity theory, distributed cognition and situated learning. Throughout, we highlight pertinent examples in non-humans that parallel or further explicate the processes discussed. Finally we offer three future directions; two involving the use of new techniques in the realms of neuroscience and modelling, and the third suggesting exploration of changes in the effects of niche construction across the lifespan. [ABSTRACT FROM AUTHOR]

Putative traditions in animal populations remain unsubstantiated in the absence of methods to isolate the mechanisms of social transmission in social groups. Here we address this problem by quantifying the effect of two social learning processes, namely stimulus enhancement and observational learning, on the adoption of a novel extractive foraging behavior in groups of callitrichid monkeys. We simulate the effect of these processes in a model for the spread of a novel behaviour and also select which model-parameters provide the best fit to the monkey data. Our analysis reveals evidence for asocial processes and stimulus enhancement but not observational learning. The latency to solve the tasks co-varied with the strength of the asocial, but not social, processes. Further, only asocial parameters were required for the model to fit the data. Both model and monkey diffusion data exhibited deceleratory diffusion curves. We discuss the relationship between both the asocial and social processes and the diffusion dynamics. [ABSTRACT FROM AUTHOR]

Researchers with diverse interests in topics ranging from the formation of dominance hierarchies and social intelligence to animal personalities have predicted specific, and often conflicting, relations between social rank, neophobia and learning ability. We investigated the relations between these variables in captive groups of wild-caught starlings, Sturnus vulgaris, adopting a multidimensional approach to social rank and neophobia. Both agonistic and competitive rank orders were determined for each group and we tested individuals in the absence of their groupmates for object neophobia, latency to feed in a novel environment and performance on an extractive foraging task. In each starling group, the fastest learners occupied the highest competitive ranks, supporting the hypothesis that cognitive ability is positively correlated with social dominance. Competitive rank orders, however, did not correlate significantly with agonistic rank orders. Situation-specific foraging neophobia was suggested: individuals showed consistency in their latencies to feed near a variety of novel objects, but no significant correlation was found between this measure of object neophobia and latency to feed in a novel environment. Starlings fastest to feed in the novel environment were fastest in solving the foraging task. We discuss the implications of these findings for researchers studying hierarchy formation in animal groups, social intelligence and animal personalities. [Copyright &y& Elsevier]