Cultural traits operating in senders are driving forces of cultural evolution.

We introduce a mathematical model of cultural evolution to study cultural traits that shape how individuals exchange information. Current theory focuses on traits that influence the reception of information (receiver traits), such as evaluating whether information represents the majority or stems from a trusted source. Our model shifts the focus from the receiver to the sender of cultural information and emphasizes the role of sender traits, such as communicability or persuasiveness. Here, we show that sender traits are probably a stronger driving force in cultural evolution than receiver traits. While receiver traits evolve to curb cultural transmission, sender traits can amplify it and fuel the self-organization of systems of mutually supporting cultural traits, including traits that cannot be maintained on their own. Such systems can reach arbitrary complexity, potentially explaining uniquely human practical and mental skills, goals, knowledge and creativity, independent of innate factors. Our model incorporates social and individual learning throughout the lifespan, thus connecting cultural evolutionary theory with developmental psychology. This approach provides fresh insights into the trait-individual duality, that is, how cultural transmission of single traits is influenced by individuals, who are each represented as an acquired system of cultural traits.

Sequence representation as an early step in the evolution of language.

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.

How associations become behavior.

The Rescorla and Wagner (1972) model is the first mathematical theory to explain associative learning in the presence of multiple stimuli. Its main theoretical construct is that of associative strength, but this is connected to behavior only loosely. We propose a model in which behavior is described by a collection of Poisson processes, each with a rate proportional to an associative strength. The model predicts that the time between behaviors follows an exponential or hypoexponential distribution. This prediction is supported by two data sets on autoshaped and instrumental behavior in rats.

Correlations in fertility across generations: can low fertility persist?

Correlations in family size across generations could have a major influence on human population size in the future. Empirical studies have shown that the associations between the fertility of parents and the fertility of children are substantial and growing over time. Despite their potential long-term consequences, intergenerational fertility correlations have largely been ignored by researchers. We present a model of the fertility transition as a cultural process acting on new lifestyles associated with fertility. Differences in parental and social influences on the acquisition of these lifestyles result in intergenerational correlations in fertility. We show different scenarios for future population size based on models that disregard intergenerational correlations in fertility, models with fertility correlations and a single lifestyle, and models with fertility correlations and multiple lifestyles. We show that intergenerational fertility correlations will result in an increase in fertility over time. However, present low-fertility levels may persist if the rapid introduction of new cultural lifestyles continues into the future.

Coevolution of intelligence, behavioral repertoire, and lifespan.

Across many taxa, intriguing positive correlations exist between intelligence (measured by proxy as encephalization), behavioral repertoire size, and lifespan. Here we argue, through a simple theoretical model, that such correlations arise from selection pressures for efficient learning of behavior sequences. We define intelligence operationally as the ability to disregard unrewarding behavior sequences, without trying them out, in the search for rewarding sequences. We show that increasing a species' behavioral repertoire increases the number of rewarding behavior sequences that can be performed, but also the time required to learn such sequences. This trade-off results in an optimal repertoire size that decreases rapidly with increasing sequence length. Behavioral repertoire size can be increased by increasing intelligence or lengthening the lifespan, giving rise to the observed correlations between these traits.

A test of memory for stimulus sequences in great apes.

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.

Models of conditioned reinforcement and abnormal behaviour in captive animals.

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.

Repeated learning makes cultural evolution unique.

Although genetic information is acquired only once, cultural information can be both abandoned and reacquired during an individual's lifetime. Therefore, cultural evolution will be determined not only by cultural traits' ability to spread but also by how good they are at sticking with an individual; however, the evolutionary consequences of this aspect of culture have not previously been explored. Here we show that repeated learning and multiple characteristics of cultural traits make cultural evolution unique, allowing dynamical phenomena we can recognize as specifically cultural, such as traits that both spread quickly and disappear quickly. Importantly, the analysis of our model also yields a theoretical objection to the popular suggestion that biological and cultural evolution can be understood in similar terms. We find that the possibility to predict long-term cultural evolution by some success index, analogous to biological fitness, depends on whether individuals have few or many opportunities to learn. If learning opportunities are few, we find that the existence of a success index may be logically impossible, rendering notions of "cultural fitness" meaningless. On the other hand, if individuals can learn many times, we find a success index that works, regardless of whether the transmission pattern is vertical, oblique, or horizontal.

Cultural evolution and individual development of openness and conservatism.

We present a model of cultural evolution in which an individual's propensity to engage in social learning is affected by social learning itself. We assume that individuals observe cultural traits displayed by others and decide whether to copy them based on their overall preference for the displayed traits. Preferences, too, can be transmitted between individuals. Our results show that such cultural dynamics tends to produce conservative individuals, i.e., individuals who are reluctant to copy new traits. Openness to new information, however, can be maintained when individuals need significant time to acquire the cultural traits that make them effective cultural models. We show that a gradual enculturation of young individuals by many models and a larger cultural repertoire to be acquired are favorable circumstances for the long-term maintenance of openness in individuals and groups. Our results agree with data about lifetime personality change, showing that openness to new information decreases with age. Our results show that cultural remodeling of cultural transmission is a powerful force in cultural evolution, i.e., that cultural evolution can change its own dynamics.

Exposure to mother's pregnancy and lactation in infancy is associated with sexual attraction to pregnancy and lactation in adulthood.

INTRODUCTION: Several theories, including psychodynamic theories, sexual imprinting and early conditioning have been formulated to explain sexual development. Empirical data, however, remain insufficient for a thorough evaluation of these theories. AIM: In this study, we test the hypothesis that a critical period exists for the acquisition of sexual preferences, as suggested by empirical findings in birds and mammals (sexual imprinting). METHODS: An Internet questionnaire was used. MAIN OUTCOME MEASURES: We gather data from individuals with a sexual preference for pregnant and/or lactating women, under the hypothesis that pregnancy or lactation may become sexually attractive in adulthood following an exposure to pregnant or lactating women in infancy. RESULTS: We find that these preferences are more common in older siblings, i.e., in individuals who have been exposed to more maternal pregnancy and lactation. This result is independent of respondent and sibling sex. In addition, only maternal pregnancies and lactations experienced between 1.5 and 5 years of age are associated with the preferences. CONCLUSIONS: We discuss our findings in relation to theories of sexual development and to earlier reports of birth order effects on sexual behavior. We suggest that this age range may constitute a sensitive period for the acquisition of sexual preferences.

Modelling cultural systems and selective filters.

A specific goal of the field of cultural evolution is to understand how processes of transmission and selection at the individual level lead to population-wide patterns of cultural diversity and change. Models of cultural evolution have typically assumed that traits are independent of one another and essentially exchangeable. But culture has a structure: traits bear relationships to one another that affect the transmission and selection process itself. Here, we introduce a modelling framework to explore the effect of interdependencies on the process of learning. Through simulations, we find that introducing a simple structure changes the cultural dynamics. Based on a basic filtering mechanism for parsing trait relationships, more elaborate cultural filters emerge. In a mostly incompatible cultural domain of traits, these filters organize culture into mostly (but not fully) consistent and stable systems. Incompatible domains produce small homogeneous cultures, while more compatibility increases size, diversity and group divergence. When individuals copy based on a trait's features (here, its compatibility relationships), they produce more homogeneous cultures than when they copy based on the agent carrying the cultural trait. We discuss the implications of considering cultural systems and filters in the dynamics of cultural change. This article is part of the theme issue 'Foundations of cultural evolution'.

Underappreciated features of cultural evolution.

Cultural evolution theory has long been inspired by evolutionary biology. Conceptual analogies between biological and cultural evolution have led to the adoption of a range of formal theoretical approaches from population dynamics and genetics. However, this has resulted in a research programme with a strong focus on cultural transmission. Here, we contrast biological with cultural evolution, and highlight aspects of cultural evolution that have not received sufficient attention previously. We outline possible implications for evolutionary dynamics and argue that not taking them into account will limit our understanding of cultural systems. We propose 12 key questions for future research, among which are calls to improve our understanding of the combinatorial properties of cultural innovation, and the role of development and life history in cultural dynamics. Finally, we discuss how this vibrant research field can make progress by embracing its multidisciplinary nature. This article is part of the theme issue 'Foundations of cultural evolution'.

Sustainability of culture-driven population dynamics.

We consider models of the interactions between human population dynamics and cultural evolution, asking whether they predict sustainable or unsustainable patterns of growth. Phenomenological models predict either unsustainable population growth or stabilization in the near future. The latter prediction, however, is based on extrapolation of current demographic trends and does not take into account causal processes of demographic and cultural dynamics. Most existing causal models assume (or derive from simplified models of the economy) a positive feedback between cultural evolution and demographic growth, and predict unlimited growth in both culture and population. We augment these models taking into account that: (1) cultural transmission is not perfect, i.e., culture can be lost; (2) culture does not always promote population growth. We show that taking these factors into account can cause radically different model behavior, such as population extinction rather than stability, and extinction rather than growth. We conclude that all models agree that a population capable of maintaining a large amount of culture, including a powerful technology, runs a high risk of being unsustainable. We suggest that future work must address more explicitly both the dynamics of resource consumption and the cultural evolution of beliefs implicated in reproductive behavior (e.g., ideas about the preferred family size) and in resource use (e.g., environmentalist stances).

A-learning: A new formulation of associative learning theory.

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.

Accumulation of independent cultural traits.

In a species capable of (imperfect) social learning, how much culture can a population of a given size carry? And what is the relationship between the individual and the population? In the first study of these novel questions, here we develop a mathematical model of the accumulation of independent cultural traits in a finite population with overlapping generations.

On the role of responses in Pavlovian acquisition.

A defining feature of Pavlovian conditioning is that the unconditioned stimulus (US) is delivered whether or not the animal performs a conditioned response (CR). This has lead to the question: Does CR performance play any role in conditioning? Between the 1930s and 1970s, a consensus emerged that CR acquisition is driven by CS-US (CS, conditioned stimulus) experiences, and that CRs play a minimal role, if any. Here we revisit the question and present 2 new quantitative methods to evaluate whether CRs influence the course of learning. Our results suggest that CRs play an important role in Pavlovian acquisition, in such paradigms as rabbit eye blink conditioning, pigeon autoshaped key pecking, and rat autoshaped lever pressing and magazine entry. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

Social learning through associative processes: a computational theory.

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.

Memory for stimulus sequences: a divide between humans and other animals?

Humans stand out among animals for their unique capacities in domains such as language, culture and imitation, yet it has been difficult to identify cognitive elements that are specifically human. Most research has focused on how information is processed after it is acquired, e.g. in problem solving or 'insight' tasks, but we may also look for species differences in the initial acquisition and coding of information. Here, we show that non-human species have only a limited capacity to discriminate ordered sequences of stimuli. Collating data from 108 experiments on stimulus sequence discrimination (1540 data points from 14 bird and mammal species), we demonstrate pervasive and systematic errors, such as confusing a red-green sequence of lights with green-red and green-green sequences. These errors can persist after thousands of learning trials in tasks that humans learn to near perfection within tens of trials. To elucidate the causes of such poor performance, we formulate and test a mathematical model of non-human sequence discrimination, assuming that animals represent sequences as unstructured collections of memory traces. This representation carries only approximate information about stimulus duration, recency, order and frequency, yet our model predicts non-human performance with a 5.9% mean absolute error across 68 datasets. Because human-level cognition requires more accurate encoding of sequential information than afforded by memory traces, we conclude that improved coding of sequential information is a key cognitive element that may set humans apart from other animals.

Time Does Not Help Orangutans Pongo abelii Solve Physical Problems.

Many questions in animal intelligence and cognition research are challenging. One challenge is to identify mechanisms underlying reasoning in experiments. Here, we provide a way to design such tests in non-human animals. We know from research in skill acquisition in humans that reasoning and thinking can take time because some problems are processed in multiple steps before a solution is reached (e.g., during mental arithmetics). If animals are able to learn through similar processes their decision making can be time consuming, and most importantly improve if more time to process information is allowed. We tested if performance of two Sumatran orangutans (Pongo abelii) increased in a two-choice experiment when they were allowed extra time before making their decisions, compared to when they were forced to decide immediately. We found that the performance of the orangutans did not depend on the time they were allowed to process the information before making their decisions. This methodology provides a potential avenue for empirical tests of mechanisms underlying reasoning in non-human animals.