We discuss recent findings suggesting that non-human animals lack memory for stimulus sequences, and therefore do not represent the order of stimuli faithfully. These observations have far-reaching consequences for animal cognition, neuroscience, and studies of the evolution of language and culture. This is because, if non-human animals do not remember or process information about order faithfully, then it is unlikely that non-human animals perform mental simulations, construct mental world models, have episodic memory, or transmit culture faithfully. If this suggested sequence bottleneck proves to be a prevalent characteristic of animal memory systems, as suggested by recent work, it would require a re-examination of some influential concepts and ideas.

Developmental psychology and cultural evolution are concerned with the same research questions but rarely interact. Collaboration between these fields could lead to substantial progress. Developmental psychology and related fields such as educational science and linguistics explore how behavior and cognition develop through combinations of social and individual experiences and efforts. Human developmental processes display remarkable plasticity, allowing children to master complex tasks, many which are of recent origin and not part of our biological history, such as mental arithmetic or pottery. It is this potency of human developmental mechanisms that allow humans to have culture on a grand scale. Biological evolution would only establish such plasticity if the combinatorial problems associated with flexibility could be solved, biological goals be reasonably safeguarded, and cultural transmission faithful. We suggest that cultural information can guide development in similar way as genes, provided that cultural evolution can establish productive transmission/teaching trajectories that allow for incremental acquisition of complex tasks. We construct a principle model of development that fulfills the needs of both subjects that we refer to as Incremental Functional Development. This process is driven by an error-correcting mechanism that attempts to fulfill combinations of cultural and inborn goals, using cultural information about structure. It supports the acquisition of complex skills. Over generations, it maintains function rather than structure, and this may solve outstanding issues about cultural transmission. The presence of cultural goals gives the mechanisms an open architecture that become an engine for cultural evolution.

Identifying cognitive capacities underlying the human evolutionary transition is challenging, and many hypotheses exist for what makes humans capable of, for example, producing and understanding language, preparing meals, and having culture on a grand scale. Instead of describing processes whereby information is processed, recent studies have suggested that there are key differences between humans and other animals in how information is recognized and remembered. Such constraints may act as a bottleneck for subsequent information processing and behavior, proving important for understanding differences between humans and other animals. We briefly discuss different sequential aspects of cognition and behavior and the importance of distinguishing between simultaneous and sequential input, and conclude that explicit tests on non-human great apes have been lacking. Here, we test the memory for stimulus sequences-hypothesis by carrying out three tests on bonobos and one test on humans. Our results show that bonobos’ general working memory decays rapidly and that they fail to learn the difference between the order of two stimuli even after more than 2,000 trials, corroborating earlier findings in other animals. However, as expected, humans solve the same sequence discrimination almost immediately. The explicit test on whether bonobos represent stimulus sequences as an unstructured collection of memory traces was not informative as no differences were found between responses to the different probe tests. However, overall, this first empirical study of sequence discrimination on non-human great apes supports the idea that non-human animals, including the closest relatives to humans, lack a memory for stimulus sequences. This may be an ability that sets humans apart from other animals and could be one reason behind the origin of human culture.

In a recent study, Wasserman, Kain, and O'Donoghue (Current Biology, 33(6), 1112–1116, 2023) set out to resolve the associative learning paradox by showing that pigeons can solve a complex category learning task through associative learning. The present Outlook paper presents their findings, expands on this paradox, and discusses implications of their results.

Abnormal behaviours are common in captive animals, and despite a lot of research, the development, maintenance and alleviation of these behaviours are not fully understood. Here, we suggest that conditioned reinforcement can induce sequential dependencies in behaviour that are difficult to infer from direct observation. We develop this hypothesis using recent models of associative learning that include conditioned reinforcement and inborn facets of behaviour, such as predisposed responses and motivational systems. We explore three scenarios in which abnormal behaviour emerges from a combination of associative learning and a mismatch between the captive environment and inborn predispositions. The first model considers how abnormal behaviours, such as locomotor stereotypies, may arise from certain spatial locations acquiring conditioned reinforcement value. The second model shows that conditioned reinforcement can give rise to abnormal behaviour in response to stimuli that regularly precede food or other reinforcers. The third model shows that abnormal behaviour can result from motivational systems being adapted to natural environments that have different temporal structures than the captive environment. We conclude that models including conditioned reinforcement offer an important theoretical insight regarding the complex relationships between captive environments, inborn predispositions, and learning. In the future, this general framework could allow us to further understand and possibly alleviate abnormal behaviours.

Human language is unique in its compositional, open-ended, and sequential form, and its evolution is often solely explained by advantages of communication. However, it has proven challenging to identify an evolutionary trajectory from a world without language to a world with language, especially while at the same time explaining why such an advantageous phenomenon has not evolved in other animals. Decoding sequential information is necessary for language, making domain-general sequence representation a tentative basic requirement for the evolution of language and other uniquely human phenomena. Here, using formal evolutionary analyses of the utility of sequence representation we show that sequence representation is exceedingly costly and that current memory systems found in animals may prevent abilities necessary for language to emerge. For sequence representation to evolve, flexibility allowing for ignoring irrelevant information is necessary. Furthermore, an abundance of useful sequential information and extensive learning opportunities are required, two conditions that were likely fulfilled early in human evolution. Our results provide a novel, logically plausible trajectory for the evolution of uniquely human cognition and language, and support the hypothesis that human culture is rooted in sequential representational and processing abilities.

The Human Evolutionary Transition offers a unified view of the evolution of intelligence, presenting a bold and provocative new account of how animals and humans have followed two powerful yet very different evolutionary paths to intelligence. This incisive book shows how animals rely on robust associative mechanisms that are guided by genetic information, which enable animals to sidestep complex problems in learning and decision making but ultimately limit what they can learn. Humans embody an evolutionary transition to a different kind of intelligence, one that relies on behavioral and mental flexibility. The book argues that flexibility is useless to most animals because they lack sufficient opportunities to learn new behavioral and mental skills. Humans find these opportunities in lengthy childhoods and through culture. Blending the latest findings in fields ranging from psychology to evolutionary anthropology, The Human Evolutionary Transition draws on computational analyses of the problems organisms face, extensive overviews of empirical data on animal and human learning, and mathematical modeling and computer simulations of hypotheses about intelligence. This compelling book demonstrates that animal and human intelligence evolved from similar selection pressures while identifying bottlenecks in evolution that may explain why human-like intelligence is so rare.

We present a new mathematical formulation of associative learning focused on non-human animals, which we call A-learning. Building on current animal learning theory and machine learning, A-learning is composed of two learning equations, one for stimulus-response values and one for stimulus values (conditioned reinforcement). A third equation implements decision-making by mapping stimulus-response values to response probabilities. We show that A-learning can reproduce the main features of: instrumental acquisition, including the effects of signaled and unsignaled non-contingent reinforcement; Pavlovian acquisition, including higher-order conditioning, omission training, autoshaping, and differences in form between conditioned and unconditioned responses; acquisition of avoidance responses; acquisition and extinction of instrumental chains and Pavlovian higher-order conditioning; Pavlovian-to-instrumental transfer; Pavlovian and instrumental outcome revaluation effects, including insight into why these effects vary greatly with training procedures and with the proximity of a response to the reinforcer. We discuss the differences between current theory and A-learning, such as its lack of stimulus-stimulus and response-stimulus associations, and compare A-learning with other temporal-difference models from machine learning, such as Q-learning, SARSA, and the actor-critic model. We conclude that A-learning may offer a more convenient view of associative learning than current mathematical models, and point out areas that need further development.

Social transmission of information is a key phenomenon in the evolution of behaviour and in the establishment of traditions and culture. The diversity of social learning phenomena has engendered a diverse terminology and numerous ideas about underlying learning mechanisms, at the same time that some researchers have called for a unitary analysis of social learning in terms of associative processes. Leveraging previous attempts and a recent computational formulation of associative learning, we analyse the following learning scenarios in some generality: learning responses to social stimuli, including learning to imitate; learning responses to non-social stimuli; learning sequences of actions; learning to avoid danger. We conceptualize social learning as situations in which stimuli that arise from other individuals have an important role in learning. This role is supported by genetic predispositions that either cause responses to social stimuli or enable social stimuli to reinforce specific responses. Simulations were performed using a new learning simulator program. The simulator is publicly available and can be used for further theoretical investigations and to guide empirical research of learning and behaviour. Our explorations show that, when guided by genetic predispositions, associative processes can give rise to a wide variety of social learning phenomena, such as stimulus and local enhancement, contextual imitation and simple production imitation, observational conditioning, and social and response facilitation. In addition, we clarify how associative mechanisms can result in transfer of information and behaviour from experienced to naive individuals.