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Whalen, A., Cownden, D., & Laland, K. (2015). The learning of action sequences through social transmission. Anim. Cogn., 18(5), 1093–1103.
Abstract: Previous empirical work on animal social learning has found that many species lack the ability to learn entire action sequences solely through reliance on social information. Conversely, acquiring action sequences through asocial learning can be difficult due to the large number of potential sequences arising from even a small number of base actions. In spite of this, several studies report that some primates use action sequences in the wild. We investigate how social information can be integrated with asocial learning to facilitate the learning of action sequences. We formalize this problem by examining how learners using temporal difference learning, a widely applicable model of reinforcement learning, can combine social cues with their own experiences to acquire action sequences. The learning problem is modeled as a Markov decision process. The learning of nettle processing by mountain gorillas serves as a focal example. Through simulations, we find that the social facilitation of component actions can combine with individual learning to facilitate the acquisition of action sequences. Our analysis illustrates that how even simple forms of social learning, combined with asocial learning, generate substantially faster learning of action sequences compared to asocial processes alone, and that the benefits of social information increase with the length of the action sequence and the number of base actions.
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Webster, M. M., & Laland, K. N. (2008). Social learning strategies and predation risk: minnows copy only when using private information would be costly. Proc. R. Soc. Lond. B, 275(1653), 2869–2876.
Abstract: 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.
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Reader, S. M., & Laland, K. N. (2002). Social intelligence, innovation, and enhanced brain size in primates. Proc. Natl. Acad. Sci. U.S.A., 99(7), 4436–4441.
Abstract: Despite considerable current interest in the evolution of intelligence, the intuitively appealing notion that brain volume and “intelligence” are linked remains untested. Here, we use ecologically relevant measures of cognitive ability, the reported incidence of behavioral innovation, social learning, and tool use, to show that brain size and cognitive capacity are indeed correlated. A comparative analysis of 533 instances of innovation, 445 observations of social learning, and 607 episodes of tool use established that social learning, innovation, and tool use frequencies are positively correlated with species' relative and absolute “executive” brain volumes, after controlling for phylogeny and research effort. Moreover, innovation and social learning frequencies covary across species, in conflict with the view that there is an evolutionary tradeoff between reliance on individual experience and social cues. These findings provide an empirical link between behavioral innovation, social learning capacities, and brain size in mammals. The ability to learn from others, invent new behaviors, and use tools may have played pivotal roles in primate brain evolution.
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Reader, S. M., & Laland, K. N. (2000). Diffusion of foraging innovations in the guppy. Anim. Behav., 60(2), 175–180.
Abstract: The way in which novel learned behaviour patterns spread through animal populations remains poorly understood, despite extensive field research and the recognition that such processes play an important role in the behavioural development, social interactions and evolution of many animal species. We conducted a series of controlled diffusions of foraging information in replicate experimental populations of the guppy, Poecilia reticulata. We presented novel foraging tasks over 15 trials to mixed-sex groups, made up of food-deprived and nonfood-deprived adults (experiment 1) or small, young fish and old, large adults (experiment 2). In these diffusions, knowledge of a route to a feeder could spread through the group by subjects learning from others, discovering the route for themselves, or, most likely, by some combination of these social and asocial learning processes. We found a striking sex difference, with novel foraging information spreading at a significantly faster rate through subgroups of females than of males. Females both discovered the goal and learned the route more quickly than males. Food-deprived individuals were faster at completing the tasks over the 15 trials than nonfood-deprived guppies, and there was a significant interaction between sex and size, with a sex difference in adults but not young individuals. There was also an interaction between sex and hunger level, with food deprivation having a stronger effect on male than female performance. We suggest that information may diffuse in a similar nonrandom or 'directed' manner through many natural populations of animals. Copyright 2000 The Association for the Study of Animal Behaviour.
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Reader, S. M., & Laland, K. N. (2009). Animal Innovation. Oxford: Oxford University Press.
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Reader, S. M., Kendal, J. R., & Laland, K. N. (2003). Social learning of foraging sites and escape routes in wild Trinidadian guppies. Anim. Behav., 66(4), 729–739.
Abstract: We describe two field experiments with wild guppies, Poecilia reticulata, in Trinidad that demonstrated that guppies can acquire foraging and predator escape-response information from conspecifics. In the foraging experiment, subjects were presented with two distinctly marked feeders in their home rivers. One feeder contained a conspecific shoal in a transparent container. Guppies preferred to enter the feeder containing this artificial shoal over the other feeder. In a test phase, the artificial shoal was removed and the feeders replaced at the testing site after a 5-min delay. More guppies entered the feeder that had contained the artificial shoal over the other feeder, a difference that can be explained only by the fish learning the characteristics or location of the feeder during the training phase. We suggest that subjects acquired a foraging patch preference through a propensity to approach feeding conspecifics, a local enhancement process. In the predator escape-response experiment, naive 'observer' guppies could avoid an approaching trawl net by escaping through either a hole to which 'demonstrator' guppies had been trained or through an alternative hole. When the demonstrators were present, the naive observers escaped more often and more rapidly by the demonstrated route than the alternative route. When the demonstrators were removed, observers maintained a route preference according to the training of their demonstrators, which suggests that the observers had learned an escape route through following or observing their more knowledgeable conspecifics. Thus, both experiments reveal that guppies can socially learn in the wild. Copyright 2003 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.
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Reader, S., & Laland, K. (2001). Primate Innovation: Sex, Age and Social Rank Differences. Int. J. Primatol., 22(5), 787–805.
Abstract: Analysis of an exhaustive survey of primate behavior collated from the published literature revealed significant variation in rates of innovation among individuals of different sex, age and social rank. We searched approximately 1,000 articles in four primatology journals, together with other relevant databases, for examples of innovation. The reported incidence of innovation is higher in males and adults, and lower in females and nonadults, than would be expected by chance given the estimated relative proportions of these groups. Amongst chimpanzees, the only species for which there are sufficient data to consider alone, there is a similar sex difference in the propensity to innovate, but no effect of age. Chimpanzees of low social rank are reported as innovators more frequently than high-ranking chimpanzees are. Male chimpanzees innovate more often than females in sexual, courtship, mating and display contexts; that is, in contexts likely to increase access to mates. The largest number of recorded observations are in the foraging context, wherein contrary to expectations, there is no evidence for female chimpanzees exhibiting more innovation than males. The study is the first extensive investigation of behavioral innovation in primates and provides evidence that much individual variation in the propensity to innovate can be explained in terms of sex, age, and social rank.
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Reader, S. M., & Laland, K. N. (2003). Animal Innovation. Oxford: Oxford University Press.
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Laland, K. N., & Reader, S. M. (1999). Foraging innovation in the guppy. Anim. Behav., 57(2), 331–340.
Abstract: When novel behaviour patterns spread through animal populations, typically one animal will initiate the diffusion. It is not known whether such 'innovators' are particularly creative individuals, individuals exposed to the appropriate environmental contingencies, or individuals in a particular motivational state. We describe three experiments that investigated the factors influencing foraging innovation in the guppy Poecilia reticulata. We exposed small laboratory populations of fish to novel foraging tasks, which involved exploration and problem solving to locate a novel food source. Experiments 1 and 2 found that (1) females were more likely to innovate than males, (2) food-deprived fish were more likely to innovate than nonfood-deprived subjects, and (3) smaller fish were more likely to innovate than larger fish. We suggest that the sex difference may reflect parental investment asymmetries in males and females. Experiment 3 found that past innovators were more likely to innovate than past noninnovators. Collectively, the results suggest that differences in foraging innovation in guppies are best accounted for by differences in motivational state, but, in addition, guppies may vary in their predisposition to innovate. Copyright 1999 The Association for the Study of Animal Behaviour.
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Laland, K. N., & van Bergen, Y. (2003). Experimental studies of innovation in the guppy. Animal Innovation, , 155–174.
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