Learning set formation in a group of Guinea baboons (Papio papio): Importance of the behavioral variable measured

A recent report of one-trial learning in a group of saddle-back tamarins (Saguinus fuscicollis) conflicted with views of learning set formation based on traditional techniques employing isolated subjects. An experiment is reported in which a group of Guinea baboons (Papio papio) was given a series of discrimination and habit reversal tasks. Both one-trial learning and learning set formation occurred. Analysis of spatial behavior revealed that the group learned in a single trial to discriminate stocked from unstocked zones. Improvement in successes (reinforced “digging”) was progressive but not rapid, indicating learning set formation. It appears that outcomes depend on the behavioral variables chosen.     

The evolutionary basis of human social learning

Humans are characterized by an extreme dependence on culturally transmitted information. Such dependence requires the complex integration of social and asocial information to generate effective learning and decision making. Recent formal theory predicts that natural selection should favour adaptive learning strategies, but relevant empirical work is scarce and rarely examines multiple strategies or tasks. We tested nine hypotheses derived from theoretical models, running a series of experiments investigating factors affecting when and how humans use social information, and whether such behaviour is adaptive, across several computer-based tasks. The number of demonstrators, consensus among demonstrators, confidence of subjects, task difficulty, number of sessions, cost of asocial learning, subject performance and demonstrator performance all influenced subjects' use of social information, and did so adaptively. Our analysis provides strong support for the hypothesis that human social learning is regulated by adaptive learning rules.     

The Effect of Familiarity with Demonstrators on Social Learning in Three‐Spined Sticklebacks (Gasterosteus aculeatus)

Social learning is an important process in the spread of information, especially in changing environments where inherited behaviors may not remain relevant. In many species, the decision of whom to trust to have reliable information depends on the relationship between individuals. Many fish species, including three‐spined sticklebacks, preferentially associate with familiar individuals. Previous studies in three‐spined sticklebacks have provided mixed evidence about the effect of familiarity on social learning in this species. Therefore, this study further explores familiarity and social learning in sticklebacks, specifically from a demonstrator‐focused perspective. We found that in a food patch discrimination task, individuals with unfamiliar demonstrators performed significantly better than those with familiar demonstrators. In a problem‐solving task, we found that focal fish attended to the behavior of demonstrators, but we did not detect an effect of familiarity on performance, and indeed the proportion of individuals to solve the task after observing demonstrators was low. These results suggest that sticklebacks have a preference for unfamiliar demonstrators, but that the use of social information varies depending on context.     

Social learning strategies and predation risk: minnows copy only when using private information would be costly

Animals can acquire information from the environment privately, by sampling it directly, or socially, through learning from others. Generally, private information is more accurate, but expensive to acquire, while social information is cheaper but less reliable. Accordingly, the 'costly information hypothesis' predicts that individuals will use private information when the costs associated with doing so are low, but that they should increasingly use social information as the costs of using private information rise. While consistent with considerable data, this theory has yet to be directly tested in a satisfactory manner. We tested this hypothesis by giving minnows (Phoxinus phoxinus) a choice between socially demonstrated and non-demonstrated prey patches under conditions of low, indirect and high simulated predation risk. Subjects had no experience (experiment 1) or prior private information that conflicted with the social information provided by the demonstrators (experiment 2). In both experiments, subjects spent more time in the demonstrated patch than in the non-demonstrated patch, and in experiment 1 made fewer switches between patches, when risk was high compared with when it was low. These findings are consistent with the predictions of the costly information hypothesis, and imply that minnows adopt a 'copy-when-asocial-learning-is-costly' learning strategy.     

Familiarity affects social network structure and discovery of prey patch locations in foraging stickleback shoals

Numerous factors affect the fine-scale social structure of animal groups, but it is unclear how important such factors are in determining how individuals encounter resources. Familiarity affects shoal choice and structure in many social fishes. Here, we show that familiarity between shoal members of sticklebacks (Gasterosteus aculeatus) affects both fine-scale social organization and the discovery of resources. Social network analysis revealed that sticklebacks remained closer to familiar than to unfamiliar individuals within the same shoal. Network-based diffusion analysis revealed that there was a strong untransmitted social effect on patch discovery, with individuals tending to discover a task sooner if a familiar individual from their group had previously done so than if an unfamiliar fish had done so. However, in contrast to the effect of familiarity, the frequency with which individuals had previously associated with one another had no effect upon the likelihood of prey patch discovery. This may have been due to the influence of fish on one another''s movements; the effect of familiarity on discovery of an empty ‘control’ patch was as strong as for discovery of an actual prey patch. Our results demonstrate that factors affecting fine-scale social interactions can also influence how individuals encounter and exploit resources.     

Social foraging: individual learning and cultural transmission of innovations

We present two stochastic models of individual and social learningthat count the number of individuals exhibiting a learned, resource-producingtrait in a group of social foragers. The novelty of our modelingresults from incorporating the empirically based assumptionthat rates of both individual and social learning should dependon the frequency of the learned trait within the group. Whenresources occur as clumps shared by group members, a naive individual'sacquisition of the skill required for clump discovery/productionshould involve opposing processes of frequency dependence. Theopportunity to learn via cultural transmission should increasewith the trait's frequency, but the opportunity for learningindividually should decrease as the trait's frequency increases.The results of the model suggest that the evolution of the capacityfor cultural transmission may be promoted in environments wherescrounging at resource clumps inhibits rates of individual learning.     

Conformist learning in nine-spined sticklebacks' foraging decisions

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.     

Simulated Trust: A cheap social learning strategy

Animals use heuristic strategies to determine from which conspecifics to learn socially. This leads to directed social learning. Directed social learning protects them from copying non-adaptive information. So far, the strategies of animals, leading to directed social learning, are assumed to rely on (possibly indirect) inferences about the demonstrator’s success. As an alternative to this assumption, we propose a strategy that only uses self-established estimates of the pay-offs of behavior. We evaluate the strategy in a number of agent-based simulations. Critically, the strategy’s success is warranted by the inclusion of an incremental learning mechanism. Our findings point out new theoretical opportunities to regulate social learning for animals. More broadly, our simulations emphasize the need to include a realistic learning mechanism in game-theoretic studies of social learning strategies, and call for re-evaluation of previous findings.     

Selectivity in social and asocial learning: investigating the prevalence,effect and development of young children's learning preferences

Culture evolution requires both modification and faithful replication of behaviour, thus it is essential to understand how individuals choose between social and asocial learning. In a quasi-experimental design, 3- and 5-year-olds (176), and adults (52) were presented individually with two novel artificial fruits, and told of the apparatus'' relative difficulty (easy versus hard). Participants were asked if they wanted to attempt the task themselves or watch an experimenter attempt it first; and then had their preference either met or violated. A significant proportion of children and adults (74%) chose to learn socially. For children, this request was efficient, as observing a demonstration made them significantly quicker at the task than learning asocially. However, for 5-year-olds, children who selected asocial learning were also found to be highly efficient at the task, showing that by 5 years children are selective in choosing a learning strategy that is effective for them. Adults further evidenced this trend, and also showed selectivity based on task difficulty. This is the first study to examine the rates, performance outcomes and developmental trajectory of preferences in asocial and social learning, ultimately informing our understanding of innovation.     

Training vervet monkeys to avoid electric wires: Is there evidence for social learning?

Before being released into a large park enclosed by an electric fence, a wild‐caught vervet group (Chlorocebus aethiops) had to learn to avoid electrified wires in a smaller cage. During this training, we observed the group continuously for 12 consecutive days to investigate if social learning was involved in the learning process. Results showed that all monkeys received an electric shock (average=2.5 shocks/individual). Most contacts with the wires occurred during the first few days of training and the vervets were never observed to come into contact with the electric fence in the 18 months after their release into the large park. This suggests that the vervets learned to avoid the electrified wires by trial‐and‐error learning. It is possible that local and stimulus enhancement may have played a role, but we could not carry out the necessary control experiments to quantify the role of these components. Observational conditioning of fear can be ruled out, however, because the vervets did not show fearful behavior toward the wires. Zoo Biol 24:145–151, 2005. © 2005 Wiley‐Liss, Inc.     

Success-biased social learning: Cultural and evolutionary dynamics

Success bias is a social learning strategy whereby learners tend to acquire the cultural variants of successful individuals. I develop a general model of success-biased social learning for discrete cultural traits with stochastic payoffs, and investigate its dynamics when only two variants are present. I find that success bias inherently favors rare variants, and consequently performs worse than unbiased imitation (i.e. random copying) when success payoffs are at least mildly stochastic and the optimal variant is common. Because of this weakness, success bias fails to replace unbiased imitation in an evolutionary model when selection is fairly weak or when the environment is relatively stable, and sometimes fails to invade at all. I briefly discuss the optimal strength of success bias, the complicated nature of defining success in social learning contexts, and the value of variant frequency as an important source of information to social learners. I conclude with predictions regarding the prevalence of success bias in different behavioral domains.     

Social Learning and Primate Reintroduction

Research on social learning may be of use in the conservation of primates, especially great apes, through (a) promoting their public image, (b) identifying specific adaptations, and (c) devising reintroduction training programs. We surveyed all the instigated social learning studies in primates published since 1950 in order to assess their usefulness to devise reintroduction training programs. We identified 99 publications containing 130 data sets from 27 species of primates. Great apes produced significantly more positive social learning effects than either cercopithecines or cebids. There was also an enhanced social learning effect when skilled demonstrators were used. Our survey indicates that the scientific understanding of many aspects of primate social learning relevant to conservation, including its function, learning spatial route plotting, food and sleeping site location, predator avoidance and detection, and the effect of model and tutee status, would benefit from greater research. Future instigated studies on primate social learning would be most informative for reintroduction if they included ecologically valid tasks presented to 2 similarly composed social groups, one of which functioned as a control, i.e., without being exposed to a model.     

Foraging zebra finches (Taeniopygia guttata) are public information users rather than conformists

Social learning enables adaptive information acquisition provided that it is not random but selective. To understand species typical decision-making and to trace the evolutionary origins of social learning, the heuristics social learners use need to be identified. Here, we experimentally tested the nature of majority influence in the zebra finch. Subjects simultaneously observed two demonstrator groups differing in relative and absolute numbers (ratios 1 : 2/2 : 4/3 : 3/1 : 5) foraging from two novel food sources (black and white feeders). We find that demonstrator groups influenced observers'' feeder choices (social learning), but that zebra finches did not copy the majority of individuals. Instead, observers were influenced by the foraging activity (pecks) of the demonstrators and in an anti-conformist fashion. These results indicate that zebra finches are not conformist, but are public information users.     

Social learning of fear and safety is determined by the demonstrator's racial group

Social learning offers an efficient route through which humans and other animals learn about potential dangers in the environment. Such learning inherently relies on the transmission of social information and should imply selectivity in what to learn from whom. Here, we conducted two observational learning experiments to assess how humans learn about danger and safety from members (‘demonstrators'') of an other social group than their own. We show that both fear and safety learning from a racial in-group demonstrator was more potent than learning from a racial out-group demonstrator.     

Age-dependent social learning in a lizard

Evidence of social learning, whereby the actions of an animal facilitate the acquisition of new information by another, is taxonomically biased towards mammals, especially primates, and birds. However, social learning need not be limited to group-living animals because species with less interaction can still benefit from learning about potential predators, food sources, rivals and mates. We trained male skinks (Eulamprus quoyii), a mostly solitary lizard from eastern Australia, in a two-step foraging task. Lizards belonging to ‘young’ and ‘old’ age classes were presented with a novel instrumental task (displacing a lid) and an association task (reward under blue lid). We did not find evidence for age-dependent learning of the instrumental task; however, young males in the presence of a demonstrator learnt the association task faster than young males without a demonstrator, whereas old males in both treatments had similar success rates. We present the first evidence of age-dependent social learning in a lizard and suggest that the use of social information for learning may be more widespread than previously believed.     

Understanding Behavioral Traditions in Primates: Are Current Experimental Approaches Too Focused on Food?

Recently, several researchers have highlighted the neglect of social behaviors relative to food-related behaviors in experimental research on social learning in primates, despite the significant number of apparent social traditions reported in the field. Here we aim to highlight the discrepancy between the relative number of nonfood-related behavioral traditions reported in the wild and food-related ones, and the almost exclusive investigation of food-related behaviors in an experimental context. First we discuss aspects of social and communicative customs that make them especially interesting. Then we consider reasons why experimental approaches are crucial to developing a full understanding of behavioral traditions observed in the wild. We report the results of a systematic literature survey in which we assessed the perceived discrepancy quantitatively. We also argue that the existing experimental literature, with its typical reliance on food as a motivator, may not be sufficient to elucidate the mechanisms underlying nonfood traditions, such as social conventions. Finally, we suggest new directions for the experimental investigation of social learning in primates, hoping to stimulate experimental research investigating social and communicative behavioral traditions.     

Cultural evolution of systematically structured behaviour in a non-human primate

Culture pervades human life and is at the origin of the success of our species. A wide range of other animals have culture too, but often in a limited form that does not complexify through the gradual accumulation of innovations. We developed a new paradigm to study cultural evolution in primates in order to better evaluate our closest relatives'' cultural capacities. Previous studies using transmission chain experimental paradigms, in which the behavioural output of one individual becomes the target behaviour for the next individual in the chain, show that cultural transmission can lead to the progressive emergence of systematically structured behaviours in humans. Inspired by this work, we combined a pattern reproduction task on touch screens with an iterated learning procedure to develop transmission chains of baboons (Papio papio). Using this procedure, we show that baboons can exhibit three fundamental aspects of human cultural evolution: a progressive increase in performance, the emergence of systematic structure and the presence of lineage specificity. Our results shed new light on human uniqueness: we share with our closest relatives essential capacities to produce human-like cultural evolution.     

Tolerance and Social Facilitation in the Foraging Behaviour of Free‐Ranging Crows (Corvus corone corone; C. c. cornix)

Social foraging provides animals with opportunities to gain knowledge about available food. Studies indicate that animals are influenced by social context during exploration and are able to learn socially. Carrion and hooded crows, which are opportunistic generalists with flexible social systems, have so far received little focus in this area. We combined observational and experimental approaches to investigate social interactions during foraging and social influences on crow behaviour within a free‐ranging population at Vienna Zoo, which included 115 individually marked crows. We expected the crows to be tolerant of conspecifics during foraging due to high food abundance. We predicted that social context would enhance familiar object exploration, as well as a specific foraging strategy: predation by crows on other species. We found that crows were highly tolerant of one another, as reflected by their high rates of cofeeding – where they fed directly beside conspecific(s) – relative to affiliative or agonistic interactions. Evidence for social facilitation – when the observer's behaviour is affected by the mere presence of a model – was found in both object exploration and predation behaviour. Specifically, crows touched the objects more frequently when others were present (whilst only approaching the objects when alone), and conspecifics were present more frequently during predation events involving the high‐risk target species. Evidence for enhancement during object exploration – where the observer's attention is drawn to a place or object by a model's actions – was not confirmed in this context. Our results highlight the role played by the presence of conspecifics across different contexts: natural foraging behaviour, familiar object exploration and a specific foraging strategy. To our knowledge, this is one of the first corvid studies aimed at teasing apart specific social influence and learning mechanisms in the field. These crows therefore make promising candidates for studying social learning and its consequences under naturalistic conditions.