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The concepts of social learning and exploration have been central to debates in comparative cognition research. While their roles in the origins of human cumulative culture on the one hand and creativity on the other have been highlighted, the two concepts have mostly been studied separately. In this article, we examine the relationship between adopting similar or different behaviours within a group, focusing on how exploration and exploitation shape primate communication systems. Using a comparative approach, we discuss how similarity and differentiation of communicative behaviour can be viewed as two endpoints on a continuum, impacting both individual- and group-level behavioural variation. While group-level variation is evident in some ape behaviours (e.g. foraging traditions), individual variation in communicative behaviour appears to outweigh group-level differences, making a widespread communicative culture in apes unlikely. Drawing parallels to language acquisition in human infants, we propose that ape communication follows an exploration–exploitation trajectory, with initial exploration gradually giving way to focused exploitation of genetically predisposed and/or individually developed communicative repertoires. By integrating the individual and social learning processes underlying communicative behaviour, we can gain a deeper understanding of how exploration–exploitation tensions shape communication systems across species. [ABSTRACT FROM AUTHOR]

Ocean deoxygenation and standing levels of hypoxia are shrinking fundamental niches, particularly in coastal areas, yet documented repercussions on species development and behaviour are limited. Here, we tackled the impacts of deoxygenation (7 mg O2 l−1), mild hypoxia (nocturnal 5 mg O2 l−1) and severe hypoxia (nocturnal 2 mg O2 l−1) on cuttlefish (Sepia officinalis) development (hatching success, development time, mantle length), cognition (ability to learn individually and socially) and behaviour (ability to camouflage and to explore its surroundings spatially). We found that hypoxia yielded lower survival rates, smaller body sizes and inhibited predatory (increased latency to attack the prey) and anti-predator (camouflage) behaviours. Acute and chronic exposure to low oxygen produced similar effects on cognition (inability to socially learn, increased open-field activity levels, no changes in thigmotaxis). It is thus expected that, although cuttlefish can withstand oxygen limitation to a certain degree, expanding hypoxic zones will diminish current habitat suitability. [ABSTRACT FROM AUTHOR]

Mate-choice copying is a type of social learning in which females can change their mate preference after observing the choice of others. This behaviour can potentially affect population evolution and ecology, namely through increased dispersal and reduced local adaptation. Here, we simulated the effects of mate-choice copying in populations expanding across an environmental gradient to understand whether it can accelerate or retard the expansion process. Two mate-choice copying strategies were used: (i) when females target a single individual and (ii) when females target similar individuals. We also simulated cases where the male trait singled out by females with mate choice maps perfectly onto his genotype or is influenced by genotype-by-environment interactions. These rules have different effects on the results. When a trait is determined by genotype alone, populations where copier females target all similar males expand faster and the number of potential copiers increased. However, when preference is determined by genotype-by-environment interactions, populations where copier females target a single male had higher dispersal and also expand faster, but the potential number of copiers decreases. The results show that mate-choice copying can accelerate the expansion process, although its adaptiveness depends on the information animals use in different contexts. [ABSTRACT FROM AUTHOR]

Direct reciprocity is a mechanism for the evolution of cooperation in repeated social interactions. According to the literature, individuals naturally learn to adopt conditionally cooperative strategies if they have multiple encounters with their partner. Corresponding models have greatly facilitated our understanding of cooperation, yet they often make strong assumptions on how individuals remember and process payoff information. For example, when strategies are updated through social learning, it is commonly assumed that individuals compare their average payoffs. This would require them to compute (or remember) their payoffs against everyone else in the population. To understand how more realistic constraints influence direct reciprocity, we consider the evolution of conditional behaviours when individuals learn based on more recent experiences. Even in the most extreme case that they only take into account their very last interaction, we find that cooperation can still evolve. However, such individuals adopt less generous strategies, and they cooperate less often than in the classical setup with average payoffs. Interestingly, once individuals remember the payoffs of two or three recent interactions, cooperation rates quickly approach the classical limit. These findings contribute to a literature that explores which kind of cognitive capabilities are required for reciprocal cooperation. While our results suggest that some rudimentary form of payoff memory is necessary, it suffices to remember a few interactions. [ABSTRACT FROM AUTHOR]

Evolutionary biologists have long been interested in parsing out the roles of genetics, plasticity and their interaction on adaptive trait divergence. Since males and females often have different ecological and reproductive roles, separating how their traits are shaped by interactions between their genes and environment is necessary and important. Here, we disentangle the sex-specific effects of genetic divergence, developmental plasticity, social learning and contextual plasticity on foraging behaviour in Trinidadian guppies (Poecilia reticulata) adapted to high- or low-predation habitats. We reared second-generation siblings from both predation regimes with or without predator chemical cues, and with adult conspecifics from either high- or low-predation habitats. We then quantified their foraging behaviour in water with and without predator chemical cues. We found that high-predation guppies forage more efficiently than low-predation guppies, but this behavioural difference is context-dependent and shaped by different mechanisms in males and females. Higher foraging efficiency in high-predation females is largely genetically determined, and to a smaller extent socially learned from conspecifics. However, in high-predation males, higher foraging efficiency is plastically induced by predator cues during development. Our study demonstrates sex-specific differences in genetic versus plastic responses in foraging behaviour, a trait of significance in organismal fitness and ecosystem dynamics. [ABSTRACT FROM AUTHOR]

The waggle dances of honeybees are a strikingly complex form of animal communication that underlie the collective foraging behaviour of colonies. The mechanisms by which bees assess the locations of forage sites that they have visited for representation on the dancefloor are now well-understood, but few studies have considered the remarkable backward translation of such information into flight vectors by dance-followers. Here, we explore whether the gene expression patterns that are induced through individual learning about foraging locations are mirrored when bees learn about those same locations from their nest-mates. We first confirmed that the mushroom bodies of honeybee dancers show a specific transcriptomic response to learning about distance, and then showed that approximately 5% of those genes were also differentially expressed by bees that follow dances for the same foraging sites, but had never visited them. A subset of these genes were also differentially expressed when we manipulated distance perception through an optic flow paradigm, and responses to learning about target direction were also in part mirrored in the brains of dance followers. Our findings show a molecular footprint of the transfer of learnt information from one animal to another through this extraordinary communication system, highlighting the dynamic role of the genome in mediating even very short-term behavioural changes. [ABSTRACT FROM AUTHOR]

The study of cultural evolution benefits from detailed analysis of cultural transmission in specific human domains. Chess provides a platform for understanding the transmission of knowledge due to its active community of players, precise behaviours and long-term records of high-quality data. In this paper, we perform an analysis of chess in the context of cultural evolution, describing multiple cultural factors that affect move choice. We then build a population-level statistical model of move choice in chess, based on the Dirichlet-multinomial likelihood, to analyse cultural transmission over decades of recorded games played by leading players. For moves made in specific positions, we evaluate the relative effects of frequency-dependent bias, success bias and prestige bias on the dynamics of move frequencies. We observe that negative frequency-dependent bias plays a role in the dynamics of certain moves, and that other moves are compatible with transmission under prestige bias or success bias. These apparent biases may reflect recent changes, namely the introduction of computer chess engines and online tournament broadcasts. Our analysis of chess provides insights into broader questions concerning how social learning biases affect cultural evolution. [ABSTRACT FROM AUTHOR]

Animal cultures have now been demonstrated experimentally in diverse taxa from flies to great apes. However, experiments commonly use tasks with unrestricted access to equal pay-offs and innovations seeded by demonstrators who are trained to exhibit strong preferences. Such conditions may not reflect those typically found in nature. For example, the learned preferences of natural innovators may be weaker, while competition for depleting resources can favour switching between strategies and generalizing from past experience. Here we show that in experiments where wild jackdaws (Corvus monedula) can freely discover depleting supplies of novel foods, generalization has a powerful effect on learning, allowing individuals to exploit multiple new opportunities through both social and individual learning. Further, in contrast to studies with trained demonstrators, individuals that were first to innovate showed weak preferences. As a consequence, many individuals ate all available novel foods, displaying no strong preference and no group-level culture emerged. Individuals followed a 'learn from adults' strategy, but other demographic factors played a minimal role in shaping social transmission. These results demonstrate the importance of generalization in allowing animals to exploit new opportunities and highlight how natural competitive dynamics may impede the formation of culture. [ABSTRACT FROM AUTHOR]

Are human cultures distinctively cumulative because they are uniquely compositional? We addressed this question using a summative learning paradigm where participants saw different models build different tower elements, consisting of discrete actions and objects: stacking cubes (tower base) and linking squares (tower apex). These elements could be combined to form a tower that was optimal in terms of height and structural soundness. In addition to measuring copying fidelity, we explored whether children and adults (i) extended the knowledge demonstrated to additional tower elements and (ii) productively combined them. Results showed that children and adults copied observed demonstrations and applied them to novel exemplars. However, only adults in the imitation condition combined the two newly derived base and apex, relative to adults in a control group. Nonetheless, there were remarkable similarities between children's and adults' performance across measures. Composite measures capturing errors and overall generativity in children's and adults' performance produced few population by condition interactions. Results suggest that early in development, humans possess a suite of cognitive skills—compositionality and generativity—that transforms phylogenetically widespread social learning competencies into something that may be unique to our species, cultural learning; allowing human cultures to evolve towards greater complexity. [ABSTRACT FROM AUTHOR]

The pathways through which primates acquire skills are a central focus of cultural evolution studies. The roles of social and genetic inheritance processes in skill acquisition are often confounded by environmental factors. Hybrid macaques from Koram Island (Thailand) provide an opportunity to examine the roles of inheritance and social learning to skill acquisition within a single ecological setting. These hybrids are a cross between tool-using Burmese long-tailed (Macaca fascicularis aurea) and non-tool-using common long-tailed macaques (Macaca fascicularis fascicularis). This population provides an opportunity to explore the roles of social learning and inheritance processes while being able to exclude underlying ecological factors. Here, we investigate the roles of social learning and inheritance in tool use prevalence within this population using social network analysis and simulation. Agent-based modelling (ABM) is used to generate expectations for how social/asocial learning and inheritance structure the patterning in a social network. The results of the simulation show that various transmission mechanisms can be differentiated based on associations between individuals in a social network. The results provide an investigative framework for discussing tool use transmission pathways in the Koram social network. By combining ABM, network analysis, and behavioural data from the field we can investigate the roles social learning and inheritance play in tool acquisition of wild primates. [ABSTRACT FROM AUTHOR]

The past 2 Myr have seen both unprecedented environmental instability and the evolution of the human capacity for complex culture. This, along with the observation that cultural evolution occurs faster than genetic evolution, has led to the suggestion that culture is an adaptation to an unstable environment. We test this hypothesis by examining the ability of human social learning to respond to environmental changes. We do this by inserting human participants (n = 4800) into evolutionary simulations with a changing environment while varying the social information available to individuals across five conditions. We find that human social learning shows some signs of adaptation to environmental instability, including critical social learning, the adoption of up-and-coming traits and, unexpectedly, contrariness. However, these are insufficient to avoid significant fitness declines when the environment changes, and many individuals are highly conformist, which exacerbates the fitness effects of environmental change. We conclude that human social learning reflects a compromise between the competing needs for flexibility to accommodate environmental change and fidelity to accurately transmit valuable cultural information. [ABSTRACT FROM AUTHOR]

Learning to respond appropriately to novel dangers is often essential to survival and success, but carries risks. Learning about novel threats from others (social learning) can reduce these risks. Many species, including the Trinidadian guppy (Poecilia reticulata), respond defensively to both conspecific chemical alarm cues and conspecific anti-predator behaviours, and in other fish such social information can lead to a learned aversion to novel threats. However, relatively little is known about the neural substrates underlying social learning and the degree to which different forms of learning share similar neural mechanisms. Here, we explored the neural substrates mediating social learning of novel threats from two different conspecific cues (i.e. social cue-based threat learning). We first demonstrated that guppies rapidly learn about threats paired with either alarm cues or with conspecific threat responses (demonstration). Then, focusing on acquisition rather than recall, we discovered that phospho-S6 expression, a marker of neural activity, was elevated in guppies during learning from alarm cues in the putative homologue of the mammalian lateral septum and the preoptic area. Surprisingly, these changes in neural activity were not observed in fish learning from conspecific demonstration. Together, these results implicate forebrain areas in social learning about threat but raise the possibility that circuits contribute to such learning in a stimulus-specific manner. [ABSTRACT FROM AUTHOR]

Culture is an outcome of both the acquisition of knowledge about behaviour through social transmission, and its subsequent production by individuals. Acquisition and production are often discussed or modelled interchangeably, yet to date no study has explored the consequences of their interaction for cultural diffusions. We present a generative model that integrates the two, and ask how variation in production rules might influence diffusion dynamics. Agents make behavioural choices that change as they learn from their productions. Their repertoires may also change, and the acquisition of behaviour is conditioned on its frequency. We analyse the diffusion of a novel behaviour through social networks, yielding generalizable predictions of how individual-level behavioural production rules influence population-level diffusion dynamics. We then investigate how linking acquisition and production might affect the performance of two commonly used inferential models for social learning; network-based diffusion analysis, and experience-weighted attraction models. We find that the influence that production rules have on diffusion dynamics has consequences for how inferential methods are applied to empirical data. Our model illuminates the differences between social learning and social influence, demonstrates the overlooked role of reinforcement learning in cultural diffusions, and allows for clearer discussions about social learning strategies. [ABSTRACT FROM AUTHOR]

There is growing evidence that offspring receive information about their environment vertically, i.e. from their parents (environmental parental effects or transgenerational plasticity). For example, parents exposed to predation risk may produce offspring with heightened antipredator defences. At the same time, organisms can gain information about the environment horizontally, from conspecifics. In this study, we provide some of the first evidence that horizontally acquired social information can be transmitted vertically across generations. Three-spined stickleback (Gasterosteus aculeatus) fathers produced larval offspring with altered antipredator behaviour when fathers received visual and olfactory cues from predator-chased neighbours. Although fathers did not personally witness their neighbours being chased (i.e. they never saw the predator), changes in offspring traits were similar to those induced by direct paternal exposure to predation risk. These findings suggest that two different non-genetic pathways (horizontal transfer of social information, vertical transfer via sperm-mediated paternal effects) can combine to affect offspring phenotypes. The implications of simultaneous horizontal and vertical transmission are widely appreciated in the context of disease and culture; our results suggest that they could be equally important for the maintenance of phenotypic variation and could have profound consequences for the rate at which information flows within and across generations. [ABSTRACT FROM AUTHOR]

High levels of within-population behavioural variation can have drastic demographic consequences, thus changing the evolutionary fate of populations. A major source of within-population heterogeneity is personality. Nonetheless, it is still relatively rarely accounted for in social learning studies that constitute the most basic process of cultural transmission. Here, we performed in female mosquitofish (Gambusia holbrooki) a social learning experiment in the context of mate choice, a situation called mate copying (MC), and for which there is strong evidence that it can lead to the emergence of persistent traditions of preferring a given male phenotype. When accounting for the global tendency of females to prefer larger males but ignoring differences in personality, we detected no evidence for MC. However, when accounting for the bold–shy dichotomy, we found that bold females did not show any evidence for MC, while shy females showed significant amounts of MC. This illustrates how the presence of variation in personality can hamper our capacity to detect MC. We conclude that MC may be more widespread than we thought because many studies ignored the presence of within-population heterogeneities. [ABSTRACT FROM AUTHOR]

Social learning is fundamental to human development, helping individuals adapt to changing circumstances and cooperate in groups. During the formative years of adolescence, the social environment shapes people's socio-cognitive skills needed in adulthood. Although peer influence among adolescents is traditionally associated with risky and unruly conduct, with long-term negative effects on educational, economic and health outcomes, recent findings suggest that peers may also have a positive impact. Here, we present a series of experiments with 10–20-year-olds (n = 146) showing that positive and negative peer effects reflect a domain-general factor of social information use which declines during adolescence. Exposure to disobedient peers provoked rule breaking, and selfish peers reduced prosocial behaviour, particularly in early adolescence. However, compliant peers also promoted rule compliance and fair peers increased prosociality. A belief formation task further revealed that younger adolescents tend to assimilate social information, while older adolescents prioritize personal views. Our results highlight early adolescence as a key window for peer-based interventions to improve developmental trajectories. [ABSTRACT FROM AUTHOR]

Learning and decision-making are greatly influenced by context. When navigating a complex social world, individuals must quickly ascertain where to gain important resources and which group members are useful sources of such information. Such dynamic behavioural processes require neural mechanisms that are flexible across contexts. Here we examine how the social context influences the learning response during a cue discrimination task and the neural activity patterns that underlie acquisition of this novel information. Using the cichlid fish, Astatotilapia burtoni, we show that learning of the task is faster in social groups than in a non-social context. We quantify the neural activity patterns by examining the expression of Fos, an immediate-early gene, across brain regions known to play a role in social behaviour and learning (such as the putative teleost homologues of the mammalian hippocampus, basolateral amygdala and medial amygdala/BNST complex). We find that neural activity patterns differ between social and non-social contexts. Taken together, our results suggest that while the same brain regions may be involved in the learning of a cue association, the activity in each region reflects an individual's social context. [ABSTRACT FROM AUTHOR]

Slow-fast differences in cognition among individuals have been proposed to be an outcome of the speed-accuracy trade-off in decision-making. Based on the different costs associated with acquiring information via individual and social learning, we hypothesized that slow-fast cognitive differences would also be tied to the adoption of these different learning modes. Since foragers in honeybee colonies likely have both these information acquisition modes available to them, we chose to test them for interindividual differences in individual and social learning. By measuring performance on a learning task with and without a social cue and quantifying learning rate and maximum accuracy in these two tasks, our results show the existence of a speed-accuracy trade-off in both the individual and the social learning contexts. However, the trade-off is steeper during individual learning, which was slower than social learning but led to higher accuracy. Most importantly, our results also show that bees that attained high accuracy on the individual learning task had low accuracy on the social learning task and vice versa. We discuss how these two information acquisition strategies tie to slow-fast differences in cognitive phenotypes and how they might contribute to division of labour and social behaviour. [ABSTRACT FROM AUTHOR]

Human cooperation is often claimed to be special and requiring explanations based on gene–culture coevolution favouring a desire to copy common social behaviours. If this is true, then individuals should be motivated to both observe and copy common social behaviours. Previous economic experiments, using the public goods game, have suggested individuals' desire to sacrifice for the common good and to copy common social behaviours. However, previous experiments have often not shown examples of success. Here we test, on 489 participants, whether individuals are more motivated to learn about, and more likely to copy, either common or successful behaviours. Using the same social dilemma and standard instructions, we find that individuals were primarily motivated to learn from successful rather than common behaviours. Consequently, social learning disfavoured costly cooperation, even when individuals could observe a stable, pro-social level of cooperation. Our results call into question explanations for human cooperation based on cultural evolution and/or a desire to conform with common social behaviours. Instead, our results indicate that participants were motivated by personal gain, but initially confused, despite receiving standard instructions. When individuals could learn from success, they learned to cooperate less, suggesting that human cooperation is maybe not so special after all. [ABSTRACT FROM AUTHOR]

Social learning underpins our species's extraordinary success. Learning through observation has been investigated in several species, but learning from advice—where information is intentionally broadcast—is less understood. We used a pre-registered, online experiment (n = 1492) combined with computational modelling to examine learning through observation and advice. Participants were more likely to immediately follow advice than to copy an observed choice, but this was dependent upon trust in the adviser: highly paranoid participants were less likely to follow advice in the short term. Reinforcement learning modelling revealed two distinct patterns regarding the long-term effects of social information: some individuals relied fully on social information, whereas others reverted to trial-and-error learning. This variation may affect the prevalence and fidelity of socially transmitted information. Our results highlight the privileged status of advice relative to observation and how the assimilation of intentionally broadcast information is affected by trust in others. [ABSTRACT FROM AUTHOR]

As infants develop, caregivers adjust their behaviour to scaffold their infant's emerging skills, such that changes in infants' social abilities are expected to elicit changes in caregiver behaviour. We examined whether changes in the probability of infant-directed caregiving behaviour—specifically, greeting, a ubiquitous signal used by caregivers to initiate reciprocal interactions—differ between infant–caregiver dyads with an infant later diagnosed with autism and dyads with a neurotypically developing infant during infants' first 6 months. Using longitudinal data from 163 dyads, we found that caregivers in autism dyads (n = 40) used greeting less and at later infant ages than caregivers with a neurotypically developing infant (neurotypical dyads, n = 83). Caregivers in dyads with infants at elevated familial genetic likelihood for autism who did not receive an autism diagnosis (EL-non-autism dyads, n = 40) showed no differences in greeting compared with neurotypical dyads. Socioeconomic status partially mediated the difference between autism and neurotypical dyads. These findings show that autism and socioeconomic status were associated with the mutually adapted dynamics of dyadic interaction beginning in the first postnatal weeks. Importantly, differences in caregiver greeting observed in autism dyads are not interpreted as suboptimal behaviour from caregivers but rather indicate how early emerging social differences related to autism, years before overt features are present, may alter social learning opportunities elicited by the infant. [ABSTRACT FROM AUTHOR]

Changes to social settings caused by migration, cultural change or pandemics force us to adapt to new social norms. Social norms provide groups of individuals with behavioural prescriptions and therefore can be inferred by observing their behaviour. This work aims to examine how cognitive learning processes affect adaptation and learning of new social norms. Using a multiplayer game, I found that participants initially complied with various social norms exhibited by the behaviour of bot-players. After gaining experience with one norm, adaptation to a new norm was observed in all cases but one, where an active-harm norm was resistant to adaptation. Using computational learning models, I found that active behaviours were learned faster than omissions, and harmful behaviours were more readily attributed to all group members than beneficial behaviours. These results provide a cognitive foundation for learning and adaptation to descriptive norms and can inform future investigations of group-level learning and cross-cultural adaptation. [ABSTRACT FROM AUTHOR]

Social learning is a primary mechanism for information acquisition in social species. Despite many benefits, social learning may be disadvantageous when independent learning is more efficient. For example, searching independently may be more advantageous when food sources are ephemeral and unpredictable. Individual differences in cognitive abilities can also be expected to influence social information use. Specifically, better spatial memory can make a given environment more predictable for an individual by allowing it to better track food sources. We investigated how resident food-caching chickadees discovered multiple novel food sources in both harsher, less predictable high elevation and milder, more predictable low elevation winter environments. Chickadees at high elevation were faster at discovering multiple novel food sources and discovered more food sources than birds at low elevation. While birds at both elevations used social information, the contribution of social learning to food discovery was significantly lower at high elevation. At both elevations, chickadees with better spatial cognitive flexibility were slower at discovering food sources, likely because birds with lower spatial cognitive flexibility are worse at tracking natural resources and therefore spend more time exploring. Overall, our study supported the prediction that harsh environments should favour less reliance on social learning. [ABSTRACT FROM AUTHOR]

A key goal of conservation is to protect biodiversity by supporting the longterm persistence of viable, natural populations of wild species. Conservation practice has long been guided by genetic, ecological and demographic indicators of risk. Emerging evidence of animal culture across diverse taxa and its role as a driver of evolutionary diversification, population structure and demographic processes may be essential for augmenting these conventional conservation approaches and decision-making. Animal culture was the focus of a ground-breaking resolution under the Convention on the Conservation of Migratory Species of Wild Animals (CMS), an international treaty operating under the UN Environment Programme. Here, we synthesize existing evidence to demonstrate how social learning and animal culture interact with processes important to conservation management. Specifically, we explore how social learning might influence population viability and be an important resource in response to anthropogenic change, and provide examples of how it can result in phenotypically distinct units with different, socially learnt behavioural strategies. While identifying culture and social learning can be challenging, indirect identification and parsimonious inferences may be informative. Finally, we identify relevant methodologies and provide a framework for viewing behavioural data through a cultural lens which might provide new insights for conservation management. [ABSTRACT FROM AUTHOR]

Noise pollution is commonly associated with human environments and mounting evidence indicates that noise has a variety of negative effects on wildlife. Noise has also been linked to cognitive impairment in humans and because many animals use cognitively intensive processes to overcome environmental challenges, noise pollution has the potential to interfere with cognitive function in animals living in urban areas or near roads. We experimentally examined how road traffic noise impacts avian cognitive performance by testing adult zebra finches (Taeniopygia guttata) on a battery of foraging tasks in the presence or absence of traffic noise playback. Here, we show that traffic noise reduces cognitive performance, including inhibitory control, motor learning, spatial memory and social learning, but not associative colour learning. This study demonstrates a novel mechanism through which anthropogenic noise can impact animals, namely through cognitive interference, and suggests that noise pollution may have previously unconsidered consequences for animals. [ABSTRACT FROM AUTHOR]

Human cumulative cultural evolution (CCE) is recognized as a powerful ecological and evolutionary force, but its origins are poorly understood. The long-standing view that CCE requires specialized social learning processes such as teaching has recently come under question, and cannot explain why such processes evolved in the first place. An alternative, but largely untested, hypothesis is that these processes gradually coevolved with an increasing reliance on complex tools. To address this, we used large-scale transmission chain experiments (624 participants), to examine the role of different learning processes in generating cumulative improvements in two tool types of differing complexity. Both tool types increased in efficacy across experimental generations, but teaching only provided an advantage for the more complex tools. Moreover, while the simple tools tended to converge on a common design, the more complex tools maintained a diversity of designs. These findings indicate that the emergence of cumulative culture is not strictly dependent on, but may generate selection for, teaching. As reliance on increasingly complex tools grew, so too would selection for teaching, facilitating the increasingly open-ended evolution of cultural artefacts. [ABSTRACT FROM AUTHOR]

Honeybees can be directed to profitable food sources by following waggle dances performed by other bees. Followers can often choose between using this social information or relying on memories about food sources they have visited in the past, so-called private information. While the circumstances that favour the use of either social or private information have received considerable attention, still little is known about the neurophysiological basis of information use. We hypothesized that octopamine and dopamine, two biogenic amines with important functions in reward signalling and learning, affect dance use in honeybees. We orally administered octopamine and dopamine when bees collected food at artificial feeders and tested if this affected interest in dance information about a new food source. We predicted that octopamine reduces interest in dances and strengthens private information use via an increase in the perceived value of the previously exploited resource. Since dopamine has been shown to lower reward perception, we expected it to act in the opposite direction. Octopamine-treated foragers indeed followed 32% fewer dances than control bees and increased the use of private information. Conversely, dopamine-treated bees followed dances 15% longer than control bees, but surprisingly did not use social information more. Overall, our results suggest that biogenic amine signalling affects interactions among dancers and dance followers and, thus, information flow about high-quality food sources. [ABSTRACT FROM AUTHOR]

In shaping how individuals explore their environment and interact with others, personality may mediate both individual and social learning. Yet increasing evidence indicates that personality expression is contingent on social context, suggesting that group personality composition may be key in determining how individuals learn about their environment. Here, we used recovery latency following simulated predator attacks to identify Trinidadian guppies (Poecilia reticulata) that acted in a consistently bold or shy manner. We then employed network-based diffusion analysis to track the spread of a novel foraging behaviour through groups containing different proportions of bold and shy fish. Informed associates promoted learning to a greater extent in bold individuals, but only within groups composed predominately of bold fish. As the proportion of shy fish within groups increased, bold individuals instead emerged as especially effective demonstrators that facilitated learning in others. Individuals were also more likely to learn overall within shy-dominated groups than in bold-dominated ones. We demonstrate that whether and how individuals learn is conditional on group personality composition, indicating that selection may favour traits enabling individuals to better match their behavioural phenotype to their social environment. [ABSTRACT FROM AUTHOR]

Recent studies have emphasized the role of social learning and cultural transmission in promoting conformity and uniformity in animal groups, but little attention has been given to the role of negative frequencydependent learning in impeding conformity and promoting diversity instead. Here, we show experimentally that under competitive conditions that are common in nature, social foragers (although capable of social learning) are likely to develop diversity in foraging specialization rather than uniformity. Naive house sparrows that were introduced into groups of foraging specialists did not conform to the behaviour of the specialists, but rather learned to use the alternative food-related cues, thus forming groups of complementary specialists. We further show that individuals in such groups may forage more effectively in diverse environments. Our results suggest that when the benefit from socially acquired skills diminishes through competition in a negative frequency-dependent manner, animal societies will become behaviourally diverse rather than uniform. [ABSTRACT FROM AUTHOR]

Flight is a unique adaptation at the core of many behaviours in most bird species, whether it be foraging, migration or breeding. Birds have developed a wide diversity of flight modes (e.g. flapping, gliding, soaring, hovering) which involves very specialized behaviours. A key issue when studying flight behaviours is to understand how they develop through all the ontogenetic stages of birds, from the embryo to the flying adult. This question typically involves classical debates on animal behaviour about the importance of maturation and experience. Here, we review the literature available on the development of flight behaviours in birds. First, we focus on the early period when young birds are not yet capable of flight. We discuss examples and show how endogenous processes (e.g. wing flapping in the nest, flight development timing) and environmental factors (e.g. maternal stress, nutritional stress) can influence the development of flight behaviours. Then, we review several examples showing the different processes involved in the development of flight in flight-capable juveniles (e.g. practice, trial and error learning, social learning). Despite the lack of experimental studies investigating this specific question at different developmental stages, we show that several patterns can be identified, and we anticipate that the development of new tracking techniques will allow us to study this question more thoroughly in more bird species. [ABSTRACT FROM AUTHOR]

A defining feature of human culture is that knowledge and technology continually improve over time. Such cumulative cultural evolution (CCE) probably depends far more heavily on how reliably information is preserved than on how efficiently it is refined. Therefore, one possible reason that CCE appears diminished or absent in other species is that it requires accurate but specialized forms of social learning at which humans are uniquely adept. Here, we develop a Bayesian model to contrast the evolution of high-fidelity social learning, which supports CCE, against low-fidelity social learning, which does not. We find that high-fidelity transmission evolves when (1) social and (2) individual learning are inexpensive, (3) traits are complex, (4) individual learning is abundant, (5) adaptive problems are difficult and (6) behaviour is flexible. Low-fidelity transmission differs in many respects. It not only evolves when (2) individual learning is costly and (4) infrequent but also proves more robust when (3) traits are simple and (5) adaptive problems are easy. If conditions favouring the evolution of high-fidelity transmission are stricter (3 and 5) or harder to meet (2 and 4), this could explain why social learning is common, but CCE is rare. [ABSTRACT FROM AUTHOR]

Understanding how information about threats in the environment is shared and transmitted between individuals is crucial for explaining adaptive, survival-related behaviour in humans and other animals, and for developing treatments for phobias and other anxiety disorders. Research across species has shown that observing a conspecific’s, a ‘demonstrator’s,’ threat responses causes strong and persistent threat memories in the ‘observer’. Here, we examined if physiological synchrony between demonstrator and observer can serve to predict the strength of observationally acquired conditioned responses. We measured synchrony between demonstrators’ and observers’ phasic electrodermal signals during learning, which directly reflects autonomic nervous system activity. Prior interpersonal synchrony predicted the strength of the observer’s later skin conductance responses to threat predicting stimuli, in the absence of the demonstrator. Dynamic coupling between an observer’s and a demonstrator’s autonomic nervous system activity may reflect experience sharing processes facilitating the formation of observational threat associations. [ABSTRACT FROM AUTHOR]

Social learning occurs when animals acquire knowledge or skills by observing or interacting with others and is the fundamental building block of culture. Within populations, some individuals use social learning more frequently than others, but why social learning phenotypes differ among individuals is poorly understood. We modelled the evolution of social learning frequency in a system where foragers compete for resources, and there are many different foraging options to learn about. Social learning phenotypes diverged when some options offered much better rewards than others and expected rewards changed moderately quickly over time. When options offered similar rewards or when rewards changed slowly, a single social learning phenotype evolved. This held for fixed and simple conditional social learning rules. Sufficiently complex conditional social learning rules prevented the divergence of social learning phenotypes under all conditions. Our results explain how competition can promote the divergence of social learning phenotypes. [ABSTRACT FROM AUTHOR]

In many species, individuals gather information about their environment both through direct experience and through information obtained from others. Social learning, or the acquisition of information from others, can occur both within and between species and may facilitate the rapid spread of antipredator behaviour. Within birds, acoustic signals are frequently used to alert others to the presence of predators, and individuals can quickly learn to associate novel acoustic cues with predation risk. However, few studies have addressed whether such learning occurs only though direct experience or whether it has a social component, nor whether such learning can occur between species. We investigate these questions in two sympatric species of Parids: blue tits (Cyanistes caeruleus) and great tits (Parus major). Using playbacks of unfamiliar bird vocalizations paired with a predator model in a controlled aviary setting, we find that blue tits can learn to associate a novel sound with predation risk via direct experience, and that antipredator response to the sound can be socially transmitted to hetero-specific observers, despite lack of first-hand experience. Our results suggest that social learning of acoustic cues can occur between species. Such interspecific social information transmission may help to mediate the formation of mixed-species aggregations. [ABSTRACT FROM AUTHOR]

The unique cumulative nature of human culture has often been explained by high-fidelity copying mechanisms found only in human social learning. However, transmission chain experiments in human and non-human primates suggest that cumulative cultural evolution (CCE) might not necessarily depend on high-fidelity copying after all. In this study, we test whether defining properties of CCE can emerge in a non-copying task. We performed transmission chain experiments in Guinea baboons and human children where individuals observed and produced visual patterns composed of four squares on touchscreen devices. In order to be rewarded, participants had to avoid touching squares that were touched by a previous participant. In other words, they were rewarded for innovation rather than copying. Results nevertheless exhibited fundamental properties of CCE: an increase over generations in task performance and the emergence of systematic structure. However, these properties arose from different mechanisms across species: children, unlike baboons, converged in behaviour over generations by copying specific patterns in a different location, thus introducing alternative copying mechanisms into the non-copying task. In children, prior biases towards specific shapes led to convergence in behaviour across chains, while baboon chains showed signs of lineage specificity. We conclude that CCE can result from mechanisms with varying degrees of fidelity in transmission and thus that high-fidelity copying is not necessarily the key to CCE. [ABSTRACT FROM AUTHOR]

Much evidence for non-human culture comes from vocally learned displays, such as the vocal dialects and song displays of birds and cetaceans. While many oscine birds use song complexity to assess male fitness, the role of complexity in humpback whale (Megaptera novaeangliae) song is uncertain owing to population-wide conformity to one song pattern. Although songs change gradually each year, the eastern Australian population also completely replaces their song every few years in cultural 'revolutions'. Revolutions involve learning large amounts of novel material introduced from the Western Australian population. We examined two measures of song structure, complexity and entropy, in the eastern Australian population over 13 consecutive years. These measures aimed to identify the role of complexity and information content in the vocal learning processes of humpback whales. Complexity was quantified at two hierarchical levels: the entire sequence of individual sound 'units' and the stereotyped arrangements of units which comprise a 'theme'. Complexity increased as songs evolved over time but decreased when revolutions occurred. No correlation between complexity and entropy estimates suggests that changes to complexity may represent embellishment to the song which could allow males to stand out amidst population-wide conformity. The consistent reduction in complexity during song revolutions suggests a potential limit to the social learning capacity of novel material in humpback whales. [ABSTRACT FROM AUTHOR]

The relative contributions of genetic and social factors in shaping the living world are a crucial question in ecology. The annual migration of birds to their wintering grounds and back provides significant knowledge in this field of research. Migratory movements are predominantly genetically determined in passerine birds, while in large soaring birds, it is presumed that social (cultural) factors play the largest role. In this study, we show that genetic factors in soaring birds are more important than previously assumed. We used global positioning system (GPS)-telemetry to compare the autumn journeys and wintering ranges of two closely related large raptorial bird species, the greater spotted eagle Clanga clanga and the lesser spotted eagle Clanga pomarina, and hybrids between them. The timing of migration in hybrids was similar to that of one parental species, but the wintering distributions and home range sizes were similar to those of the other. Tracking data were supported by habitat suitability modelling, based on GPS fixes and ring recoveries. These results suggest a strong genetic influence on migration strategy via a trait-dependent dominance effect, although we cannot rule out the contribution of social interactions. [ABSTRACT FROM AUTHOR]

In recent years, the phenomenon of cumulative cultural evolution (CCE) has become the focus of major research interest in biology, psychology and anthropology. Some researchers argue that CCE is unique to humans and underlies our extraordinary evolutionary success as a species. Others claim to have found CCE in non-human species. Yet others remain sceptical that CCE is even important for explaining human behavioural diversity and complexity. These debates are hampered by multiple and often ambiguous definitions of CCE. Here, we review how researchers define, use and test CCE. We identify a core set of criteria for CCE which are both necessary and sufficient, and may be found in non-human species. We also identify a set of extended criteria that are observed in human CCE but not, to date, in other species. Different socio-cognitive mechanisms may underlie these different criteria. We reinterpret previous theoretical models and observational and experimental studies of both human and non-human species in light of these more fine-grained criteria. Finally, we discuss key issues surrounding information, fitness and cognition. We recommend that researchers are more explicit about what components of CCE they are testing and claiming to demonstrate. [ABSTRACT FROM AUTHOR]

The importance of culture for human social evolution hinges largely on the extent to which culture supports outcomes that would not otherwise occur. An especially controversial claim is that social learning leads groups to coalesce around group-typical behaviours and associated social norms that spill over to shape choices in asocial settings. To test this, we conducted an experiment with 878 groups of participants in 116 communities in Sudan. Participants watched a short film and evaluated the appropriate way to behave in the situation dramatized in the film. Each session consisted of an asocial condition in which participants provided private evaluations and a social condition in which they provided public evaluations. Public evaluations allowed for social learning. Across sessions, we randomized the order of the two conditions. Public choices dramatically increased the homogeneity of normative evaluations. When the social condition was first, this homogenizing effect spilled over to subsequent asocial conditions. The asocial condition when first was thus alone in producing distinctly heterogeneous groups. Altogether, information about the choices of others led participants to converge rapidly on similar normative evaluations that continued to hold sway in subsequent asocial settings. These spillovers were at least partly owing to the combined effects of conformity and self-consistency. Conformity dominated self-consistency when the two mechanisms were in conflict, but self-consistency otherwise produced choices that persisted through time. Additionally, the tendency to conform was heterogeneous. Females conformed more than males, and conformity increased with the number of other people a decision-maker observed before making her own choice. [ABSTRACT FROM AUTHOR]

Information is a crucial currency for living organisms as it allows them to adjust their behaviour to environmental fluctuations. Thus, natural selection should have favoured the capacity of collecting information from different sources, including social interactions whereby individuals could quickly gain reliable information. However, such conditions may also favour the gathering of potentially detrimental information, such as false or misinterpreted accounts of environmental and social phenomena such as rumours, which may spread via informational cascades.We applied ecological and evolutionary principles to investigate howthe propagation of social information at a populational level affects the propensity to assimilate it, here defined as the gullibilty. Our results show that the evolution of an individual's susceptibility to assimilate information strongly depends on eco-evolutionary feedbacks, in particular when both useful and detrimental information circulate.We discuss our results regarding the different information transmission mechanisms involved with particular attention to specific cases of social learning. [ABSTRACT FROM AUTHOR]

Various non-human animal species have been shown to exhibit behavioural traditions. Importantly, this research has been guided by what we know of human culture, and the question of whether animal cultures may be homologous or analogous to our own culture. In this paper, we assess whether models of human cultural transmission are relevant to understanding biological fundamentals by investigating whether accounts of human payoff-biased social learning are relevant to chimpanzees (Pan troglodytes). We submitted 4- and 5-year-old children (N = 90) and captive chimpanzees (N = 69) to a token-reward exchange task. The results revealed different forms of payoff-biased learning across species and contexts. Specifically, following personal and social exposure to different tokens, children's exchange behaviour was consistent with proportional imitation, where choice is affected by both prior personally acquired and socially demonstrated token-reward information. However, when the socially derived information regarding token value was novel, children's behaviour was consistent with proportional observation; paying attention to socially derived information and ignoring their prior personal experience. By contrast, chimpanzees' token choice was governed by their own prior experience only, with no effect of social demonstration on token choice, conforming to proportional reservation. We also find evidence for individual- and group-level differences in behaviour in both species. Despite the difference in payoff strategies used, both chimpanzees and children adopted beneficial traits when available. However, the strategies of the children are expected to be the most beneficial in promoting flexible behaviour by enabling existing behaviours to be updated or replaced with new and often superior ones. [ABSTRACT FROM AUTHOR]

Partner discrimination is crucial in mutualistic interactions between organisms to counteract cheating by the partner. Trophobiosis between ants and aphids is a model system of such mutualistic interaction. To establish and maintain the mutualistic association, ants need to correctly discriminate mutualistic aphids. However, the mechanism by which ants recognize aphids as their partners is poorly understood, despite its ecological and evolutionary importance. Here, we show for the first time the evidence that interaction with nest-mates that have tended aphids (Aphis craccivora) allows ants (Tetramorium tsushimae) to learn to recognize the aphid species as their partner. When ants had previously tended aphids, they moderated their aggressiveness towards aphids. More importantly, ants that had interacted with aphid-experienced nest-mates also reduced their aggressiveness towards aphids, even though they had never directly experienced them, indicating that aphid information was transmitted from aphid-experienced ants to inexperienced ants. Furthermore, inhibition of mouth-to-mouth contact (trophallaxis) from aphid-experienced ants to inexperienced ants by providing the inexperienced ants with artificial honeydew solution caused the inexperienced ants to become aggressive towards aphids. These results, with further supporting data, strongly suggest that ants transfer information on their mutualists during trophallactic interactions. [ABSTRACT FROM AUTHOR]

Brood parasites face considerable cognitive challenges in locating and selecting host nests for their young. Here, we test whether female brown-headed cowbirds, Molothrus ater, could use information acquired from observing the nest prospecting patterns of conspecifics to influence their own patterns of nest selection. In laboratory-based experiments, we created a disparity in the amount of personal information females had about the quality of nests. Females with less personal information about the quality of two nests spent more time investigating the nest that more knowledgeable females investigated. Furthermore, there was a strong negative relationship between individual's ability to track nest quality using personal information and their tendency to copy others. These two contrasting strategies for selecting nests are equally effective, but lead to different patterns of parasitism. [ABSTRACT FROM AUTHOR]

Language acquisition in humans and song learning in songbirds naturally happen as a social learning experience, providing an excellent opportunity to reveal social motivation and reward mechanisms that boost sensorimotor learning. Our knowledge about the molecules and circuits that control these social mechanisms for vocal learning and language is limited. Here we propose a hypothesis of a role for oxytocin (OT) in the social motivation and evolution of vocal learning and language. Building upon existing evidence, we suggest specific neural pathways and mechanisms through which OT might modulate vocal learning circuits in specific developmental stages. [ABSTRACT FROM AUTHOR]

The type and variety of learning strategies used by individuals to acquire behaviours in the wild are poorly understood, despite the presence of behavioural traditions in diverse taxa. Social learning strategies such as conformity can be broadly adaptive, but may also retard the spread of adaptive innovations. Strategies like pay-off-biased learning, by contrast, are effective at diffusing new behaviour but may perform poorly when adaptive behaviour is common.We present a field experiment in a wild primate, Cebus capucinus, that introduced a novel food item and documented the innovation and diffusion of successful extraction techniques.We develop a multilevel, Bayesian statistical analysis that allows us to quantify individual-level evidence for different social and individual learning strategies. We find that pay-offbiased and age-biased social learning are primarily responsible for the diffusion of new techniques. We find no evidence of conformity; instead rare techniques receive slightly increased attention. We also find substantial and important variation in individual learning strategies that is patterned by age, with younger individuals being more influenced by both social information and their own individual experience. The aggregate cultural dynamics in turn depend upon the variation in learning strategies and the age structure of the wild population. [ABSTRACT FROM AUTHOR]

Humans have a sophisticated ability to learn from others, termed social learning, which has allowed us to spread over the planet, construct complex societies, and travel to the moon. It has been hypothesized that social learning has played a pivotal role in making human societies cooperative, by favouring cooperation even when it is not favoured by genetical selection. However, this hypothesis lacks direct experimental testing, and the opposite prediction has also been made, that social learning disfavours cooperation. We experimentally tested how different aspects of social learning affect the level of cooperation in public-goods games. We found that: (i) social information never increased cooperation and usually led to decreased cooperation; (ii) cooperation was lowest when individuals could observe how successful individuals behaved; and (iii) cooperation declined because individuals preferred to copy successful individuals, who cooperated less, rather than copy common behaviours. Overall, these results suggest that individuals use social information to try and improve their own success, and that this can lead to lower levels of cooperation. [ABSTRACT FROM AUTHOR]

There has been extensive game-theoretic modelling of conditions leading to equilibria of producer--scrounger dichotomies in groups. However there is a surprising paucity of experimental evidence in wild populations. Here, we examine producer--scrounger games in five subpopulations of birds feeding at a socially learnt foraging task. Over four weeks, a bimodal distribution of producers and scroungers emerged in all areas, with pronounced and con- sistent individual tactic specialization persisting over 3 years. Tactics were unrelated to exploratory personality, but correlated with latency to contact and learn the foraging task, with the late arrivers and slower learners more likely to adopt the scrounging role. Additionally, the social environment was also important: at the broad scale, larger subpopulations with a higher social density contained proportionally more scroungers, while within subpopulations scroungers tended to be central in the social network and be observed in larger foraging flocks. This study thus provides a rare example of a stable, dimorphic distribution of producer--scrounger tactics in a wild population. It further gives support across multiple scales for a major prediction of social foraging theory; that the frequency of scroungers increases with group size. [ABSTRACT FROM AUTHOR]

Understanding how humans and other animals learn to perform an act from seeing it done has been a major challenge in the study of social learning. To determine whether this ability is based on 'true imitation', many studies have applied the two-action experimental paradigm, examining whether subjects learn to perform the specific action demonstrated to them. Here, we show that the insights gained from animals' success in two-action experiments may be limited, and that a better understanding is achieved by monitoring subjects' entire behavioural repertoire. Hand-reared house sparrows that followed a model of a mother demonstrator were successful in learning to find seeds hidden under a leaf, using the action demonstrated by the mother (either pushing the leaf or pecking it). However, they also produced behaviours that had not been demonstrated but were nevertheless related to the demonstrated act. This finding suggests that while the learners were clearly influenced by the demonstrator, they did not accurately imitate her. Rather, they used their own behavioural repertoire, gradually fitting it to the demonstrated task solution through trial and error. This process is consistent with recent views on how animals learn to imitate, and may contribute to a unified process-level analysis of social learning mechanisms. [ABSTRACT FROM AUTHOR]

Innovative behaviour in animals, ranging from invertebrates to humans, is increasingly recognized as an important topic for investigation by behavioural researchers. However, what constitutes an innovation remains controversial, and difficult to quantify. Drawing on a broad definition whereby any behaviour with a new component to it is an innovation, we propose a quantitative measure, which we call the magnitude of innovation, to describe the extent to which an innovative behaviour is novel. This allows us to distinguish between innovations that are a slight change to existing behaviours (low magnitude), and innovations that are substantially different (high magnitude). Using mathematical modelling and evolutionary computer simulations, we explored how aspects of social interaction, cognition and natural selection affect the frequency and magnitude of innovation. We show that high-magnitude innovations are likely to arise regularly even if the frequency of innovation is low, as long as this frequency is relatively constant, and that the selectivity of social learning and the existence of social rewards, such as prestige and royalties, are crucial for innovative behaviour to evolve. We suggest that consideration of the magnitude of innovation may prove a useful tool in the study of the evolution of cognition and of culture. [ABSTRACT FROM AUTHOR]

Socially transmitted wildlife behaviours that create human-wildlife conflict are an emerging problem for conservation efforts, but also provide a unique opportunity to apply principles of infectious disease control to wildlife management. As an example, California sea lions (Zalophus californianus) have learned to exploit concentrations of migratory adult salmonids below the fish ladders at Bonneville Dam, impeding endangered salmonid recovery. Proliferation of this foraging behaviour in the sea lion population has resulted in a controversial culling programme of individual sea lions at the dam, but the impact of such culling remains unclear. To evaluate the effectiveness of current and alternative culling strategies, we used network-based diffusion analysis on a long-term dataset to demonstrate that social transmission is implicated in the increase in dam-foraging behaviour and then studied different culling strategies within an epidemiological model of the behavioural transmission data. We show that current levels of lethal control have substantially reduced the rate of social transmission, but failed to effectively reduce overall sea lion recruitment. Earlier implementation of culling could have substantially reduced the extent of behavioural transmission and, ultimately, resulted in fewer animals being culled. Epidemiological analyses offer a promising tool to understand and control socially transmissible behaviours. [ABSTRACT FROM AUTHOR]