3.3 million years of stone tool complexity suggests that cumulative culture began during the Middle Pleistocene.

Cumulative culture, the accumulation of modifications, innovations, and improvements over generations through social learning, is a key determinant of the behavioral diversity across Homo sapiens populations and their ability to adapt to varied ecological habitats. Generations of improvements, modifications, and lucky errors allow humans to use technologies and know-how well beyond what a single naive individual could invent independently within their lifetime. The human dependence on cumulative culture may have shaped the evolution of biological and behavioral traits in the hominin lineage, including brain size, body size, life history, sociality, subsistence, and ecological niche expansion. Yet, we do not know when, in the human career, our ancestors began to depend on cumulative culture. Here, we show that hominins likely relied on a derived form of cumulative culture by at least ~600 kya, a result in line with a growing body of existing evidence. We analyzed the complexity of stone tool manufacturing sequences over the last 3.3 My of the archaeological record. We then compare these to the achievable complexity without cumulative culture, which we estimate using nonhuman primate technologies and stone tool manufacturing experiments. We find that archaeological technologies become significantly more complex than expected in the absence of cumulative culture only after ~600 kya.

The Evolution of the Optimization of Cognitive and Social Functions in the Cerebellum and Thereby the Rise of Homo sapiens Through Cumulative Culture.

The evolution of the prominent role of the cerebellum in the development of composite tools, and cumulative culture, leading to the rise of Homo sapiens is examined. Following Stout and Hecht's (2017) detailed description of stone-tool making, eight key repetitive involvements of the cerebellum are highlighted. These key cerebellar learning involvements include the following: (1) optimization of cognitive-social control, (2) prediction (3) focus of attention, (4) automaticity of smoothness, appropriateness, and speed of movement and cognition, (5) refined movement and social cognition, (6) learns models of extended practice, (7) learns models of Theory of Mind (ToM) of teachers, (8) is predominant in acquisition of novel behavior and cognition that accrues from the blending of cerebellar models sent to conscious working memory in the cerebral cortex. Within this context, the evolution of generalization and blending of cerebellar internal models toward optimization of social-cognitive learning is described. It is concluded that (1) repetition of movement and social cognition involving the optimization of internal models in the cerebellum during stone-tool making was the key selection factor toward social-cognitive and technological advancement, (2) observational learning during stone-tool making was the basis for both technological and social-cognitive evolution and, through an optimizing positive feedback loop between the cerebellum and cerebral cortex, the development of cumulative culture occurred, and (3) the generalization and blending of cerebellar internal models related to the unconscious forward control of the optimization of imagined future states in working memory was the most important brain adaptation leading to intertwined advances in stone-tool technology, cognitive-social processes behind cumulative culture (including the emergence of language and art) and, thereby, with the rise of Homo sapiens.

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.

Evolved Open-Endedness in Cultural Evolution: A New Dimension in Open-Ended Evolution Research.

The goal of Artificial Life research, as articulated by Chris Langton, is "to contribute to theoretical biology by locating life-as-we-know-it within the larger picture of life-as-it-could-be." The study and pursuit of open-ended evolution in artificial evolutionary systems exemplify this goal. However, open-ended evolution research is hampered by two fundamental issues: the struggle to replicate open-endedness in an artificial evolutionary system and our assumption that we only have one system (genetic evolution) from which to draw inspiration. We argue not only that cultural evolution should be seen as another real-world example of an open-ended evolutionary system but that the unique qualities seen in cultural evolution provide us with a new perspective from which we can assess the fundamental properties of, and ask new questions about, open-ended evolutionary systems, especially with regard to evolved open-endedness and transitions from bounded to unbounded evolution. Here we provide an overview of culture as an evolutionary system, highlight the interesting case of human cultural evolution as an open-ended evolutionary system, and contextualize cultural evolution by developing a new framework of (evolved) open-ended evolution. We go on to provide a set of new questions that can be asked once we consider cultural evolution within the framework of open-ended evolution and introduce new insights that we may be able to gain about evolved open-endedness as a result of asking these questions.

To copy or not to copy? That is the question! From chimpanzees to the foundation of human technological culture.

A prerequisite for copying innovative behaviour faithfully is the capacity of observers' brains, regarded as 'hierarchically mechanistic minds', to overcome cognitive 'surprisal' (see 2.), by maximising the evidence for their internal models, through active inference. Unlike modern humans, chimpanzees and other great apes show considerable limitations in their ability, or 'Zone of Bounded Surprisal', to overcome cognitive surprisal induced by innovative or unorthodox behaviour that rarely, therefore, is copied precisely or accurately. Most can copy adequately what is within their phenotypically habitual behavioural repertoire, in which technology plays scant part. Widespread intra- and intergenerational social transmission of complex technological innovations is not a hall-mark of great-ape taxa. 3 Ma, precursors of the genus Homo made stone artefacts, and stone-flaking likely was habitual before 2 Ma. After that time, early Homo erectus has left traces of technological innovations, though faithful copying of these and their intra- and intergenerational social transmission were rare before 1 Ma. This likely owed to a cerebral infrastructure of interconnected neuronal systems more limited than ours. Brains were smaller in size than ours, and cerebral neuronal systems ceased to develop when early Homo erectus attained full adult maturity by the mid-teen years, whereas its development continues until our mid-twenties nowadays. Pleistocene Homo underwent remarkable evolutionary adaptation of neurobiological propensities, and cerebral aspects are discussed that, it is proposed here, plausibly, were fundamental for faithful copying, which underpinned social transmission of technologies, cumulative learning, and culture. Here, observers' responses to an innovation are more important for ensuring its transmission than is an innovator's production of it, because, by themselves, the minimal cognitive prerequisites that are needed for encoding and assimilating innovations are insufficient for practical outcomes to accumulate and spread intra- and intergenerationally.

Corrigendum: Small steps for mankind: Modeling the emergence of cumulative culture from joint active inference communication.

[This corrects the article DOI: 10.3389/fnbot.2022.944986.].

Conditions that favour cumulative cultural evolution.

The emergence of human societies with complex language and cumulative culture is considered a major evolutionary transition. Why such a high degree of cumulative culture is unique to humans is perplexing given the potential fitness advantages of cultural accumulation. Here, Boyd & Richerson's (1996 Why culture is common, but cultural evolution is rare. Proc. Br. Acad. 88, 77-93) discrete-cultural-trait model is extended to incorporate arbitrarily strong selection; conformist, anti-conformist and unbiased frequency-dependent transmission; random and periodic environmental variation; finite population size; and multiple 'skill levels.' From their infinite-population-size model with success bias and a single skill level, Boyd and Richerson concluded that social learning is favoured over individual learning under a wider range of conditions when social learning is initially common than initially rare. We find that this holds only if the number n of individuals observed by a social learner is sufficiently small, but with a finite population and/or a combination of success-biased and conformist or unbiased transmission, this result holds with larger n. Assuming social learning has reached fixation, the increase in a population's mean skill level is lower if cumulative culture is initially absent than initially present, if population size is finite, or if cultural transmission has a frequency-dependent component. Hence, multiple barriers to cultural accumulation may explain its rarity. This article is part of the theme issue 'Human socio-cultural evolution in light of evolutionary transitions'.

Human cooperation and evolutionary transitions in individuality.

A major evolutionary transition in individuality involves the formation of a cooperative group and the transformation of that group into an evolutionary entity. Human cooperation shares principles with those of multicellular organisms that have undergone transitions in individuality: division of labour, communication, and fitness interdependence. After the split from the last common ancestor of hominoids, early hominins adapted to an increasingly terrestrial niche for several million years. We posit that new challenges in this niche set in motion a positive feedback loop in selection pressure for cooperation that ratcheted coevolutionary changes in sociality, communication, brains, cognition, kin relations and technology, eventually resulting in egalitarian societies with suppressed competition and rapid cumulative culture. The increasing pace of information innovation and transmission became a key aspect of the evolutionary niche that enabled humans to become formidable cooperators with explosive population growth, the ability to cooperate and compete in groups of millions, and emergent social norms, e.g. private property. Despite considerable fitness interdependence, the rise of private property, in concert with population explosion and socioeconomic inequality, subverts potential transition of human groups into evolutionary entities due to resurgence of latent competition and conflict. This article is part of the theme issue 'Human socio-cultural evolution in light of evolutionary transitions'.

Capuchin monkeys learn to use information equally well from individual exploration and social demonstration.

The limited evidence of complex culture in non-human primates contrasts strikingly with human behaviour. This may be because non-human primates fail to use information acquired socially as effectively as they use information acquired individually. Here, monkeys were trained on a stimulus discrimination task with a win-stay, lose-shift (WSLS) reward structure. In a social learning condition, the experimenter performed an information trial by choosing between the available stimuli; in an individual condition, monkeys made this choice themselves. The monkeys' subsequent test trials displayed the same stimulus array. They were rewarded for repetition of rewarded ('win-stay') and avoidance of unrewarded ('lose-shift') information trial selections. Nine monkeys reached our pre-determined performance criterion on the initial two-stimulus stage. Their ability to generalise the WSLS strategy was then evaluated by transfer to a three-stimulus stage. Minimal differences were found in information use between the social and individual conditions on two-stimuli. However, a bias was found towards repetition of the information trial, regardless of information source condition or whether the information trial selection was rewarded. Proficient subjects were found to generalise the strategy to three-stimuli following rewarded information trials, but performed at chance on unrewarded. Again, this was not found to vary by source condition. Overall, results suggest no fundamental barrier to non-human primates' use of information from a social source. However, the apparent struggle to learn from the absence of rewards hints at a difficulty with using information acquired from unsuccessful attempts; this could be linked to the limited evidence for cumulative culture in non-human primates.

Bringing cumulative technological culture beyond copying versus reasoning.

The dominant view of cumulative technological culture suggests that high-fidelity transmission rests upon a high-fidelity copying ability, which allows individuals to reproduce the tool-use actions performed by others without needing to understand them (i.e., without causal reasoning). The opposition between copying versus reasoning is well accepted but with little supporting evidence. In this article, we investigate this distinction by examining the cognitive science literature on tool use. Evidence indicates that the ability to reproduce others' tool-use actions requires causal understanding, which questions the copying versus reasoning distinction and the cognitive reality of the so-called copying ability. We conclude that new insights might be gained by considering causal understanding as a key driver of cumulative technological culture.

Blind alleys and fruitful pathways in the comparative study of cultural cognition.

A mere few decades ago, culture was thought a unique human attribute. Evidence to the contrary accumulated through the latter part of the twentieth century and has exploded in the present one, demonstrating the transmission of traditions through social learning across all principal vertebrate taxa and even invertebrates, notably insects. The scope of human culture is nevertheless highly distinctive. What makes our cultural capacities and their cognitive underpinnings so different? In this article I argue that in behavioural scientists' endeavours to answer this question, fruitful research pathways and their ensuing discoveries have come to exist alongside popular, yet in the light of current empirical evidence, highly questionable scenarios and even scientific blind alleys. I particularly re-evaluate theories that rely on the centrality of a supposed uniquely human capacity for imitative copying in explaining the distinctive capacity for massive cumulative cultural evolution (CCE) in our species. The most extreme versions of this perspective suffer logical incoherence and severe limits on scientific testability. By contrast the field has generated a range of rigorous observational and experimental methodologies that have revealed both long-term cultural fidelity and limited forms of CCE in non-human species. Attention now turns to directly investigating the scope, limits and underlying cognition of non-human versus human CCE, with a broader approach to factors additional to cultural transmission, notably the role of invention, innovation and evolved motivational biases underlying the scope of CCE in the species studied.

Domestic spaces as crucibles of Paleolithic culture: An archaeological perspective.

The places in which people live, sleep, prepare food, and undertake other activities-known variably as homes, residential sites, living sites, and domestic spaces-play a key role in the emergence and evolution of modern human culture. The dynamic influence of domestic spaces began early in human evolutionary history, during the Paleolithic/Stone Age. Drawing on examples from Africa and western Eurasia, this article explores aspects of the changing social and cultural significance of domestic spaces throughout this time using several lines of evidence: repeated site visitation, behavioral structuring of living spaces, and information gained by dissecting palimpsest records. With the development of pyrotechnology, living sites become hearth-centered domestic spaces that provided a common hub for activities. Through time the activities around hearths increased in their complexity and diversity. The parsing of palimpsest records by archaeologists also reveals changes in the nature, variety, and intensity of on-site activities through time, indicating shifts in site function and the spatial expression of cultural norms. Archaeological evidence shows that the entwined development of domestic spaces and human cultural activities was gradual, albeit nonlinear from the Lower Paleolithic through the Upper Paleolithic/later Middle Stone Age. In this process, domestic spaces emerged as common arenas of opportunity for social interaction and knowledge transmission, qualities that may have contributed to and enhanced the development of cumulative culture in Paleolithic society.

When the "satisficing" is the new "fittest": how a proscriptive definition of adaptation can change our view of cognition and culture.

Since Darwin's theory of evolution, adaptationism is frequently invoked to explain cognition and cultural processes. Adaptationism can be described as a prescriptive view, as phenotypes that do not optimize fitness should not be selected by natural selection. From an epistemological perspective, the principle of a prescriptive definition of adaptation seems incompatible with recent advances in epigenetics, evolutionary developmental biology, ethology, and genomics. From these challenges, a proscriptive view of adaptation has emerged, postulating that phenotypes that are not deleterious will be evolutionary maintained. In this epistemological investigation, we examine how the shift from adaptationism to a proscriptive view changes our view of cognition and culture. We argue that, while adaptationism leads to cognitivism and a view of culture as strategies to optimize overall fitness, the proscriptive definition favors embodied theories of cognition and a view of culture as the cumulative diffusion of behaviors allowed by the constraints of reproduction.

Belief correlations with parental vaccine hesitancy: Results from a national survey.

We conducted a nationally representative survey of parents' beliefs and self-reported behaviors regarding childhood vaccinations. Using Bayesian selection among multivariate models, we found that beliefs, even those without any vaccine or health content, predicted vaccine-hesitant behaviors better than demographics, social network effects, or scientific reasoning. The multivariate structure of beliefs combined many types of ideation that included concerns about both conspiracies and side effects. Although they are not strongly related to vaccine-hesitant behavior, demographics were key predictors of beliefs. Our results support some of the previously proposed pro-vaccination messaging strategies and suggest some new strategies not previously considered.

Realizamos una encuesta nacionalmente representativa sobre las creencias y comportamientos autodeclarados por los padres con relación a la vacunación infantil. Usando selección bayesiana entre modelos multivariados, encontramos que las creencias, aun aquellas sin ningún contenido sobre vacunas o salud, predijeron comportamientos indecisos sobre la vacuna mejor que las características demográficas, los efectos de las redes sociales o el razonamiento científico. La estructura multivariada de las creencias combinó muchos tipos de ideación que incluyó preocupaciones tanto sobre conspiraciones como efectos secundarios. Aunque no están relacionados fuertemente con los comportamientos de indecisión, las características demográficas fueron predictores centrales de las creencias. Nuestros resultados apoyan algunas de las estrategias de mensajes pro­vacunación propuestas previamente y sugieren algunas nuevas estrategias no consideradas anteriormente. [vacunación, encuesta, cultura acumulativa, bayesiana].

Costly teaching contributes to the acquisition of spear hunting skill among BaYaka forager adolescents.

Teaching likely evolved in humans to facilitate the faithful transmission of complex tasks. As the oldest evidenced hunting technology, spear hunting requires acquiring several complex physical and cognitive competencies. In this study, we used observational and interview data collected among BaYaka foragers (Republic of the Congo) to test the predictions that costlier teaching types would be observed at a greater frequency than less costly teaching in the domain of spear hunting and that teachers would calibrate their teaching to pupil skill level. To observe naturalistic teaching during spear hunting, we invited teacher-pupil groupings to spear hunt while wearing GoPro cameras. We analysed 68 h of footage totalling 519 teaching episodes. Most observed teaching events were costly. Direct instruction was the most frequently observed teaching type. Older pupils received less teaching and more opportunities to lead the spear hunt than their younger counterparts. Teachers did not appear to adjust their teaching to pupil experience, potentially because age was a more easily accessible heuristic for pupil skill than experience. Our study shows that costly teaching is frequently used to transmit complex tasks and that instruction may play a privileged role in the transmission of spear hunting knowledge.

Restricted Access to Working Memory Does Not Prevent Cumulative Score Improvement in a Cultural Evolution Task.

Some theories propose that human cumulative culture is dependent on explicit, system-2, metacognitive processes. To test this, we investigated whether access to working memory is required for cumulative cultural evolution. We restricted access to adults' working-memory (WM) via a dual-task paradigm, to assess whether this reduced performance in a cultural evolution task, and a metacognitive monitoring task. In total, 247 participants completed either a grid search task or a metacognitive monitoring task in conjunction with a WM task and a matched control. Participants' behaviour in the grid search task was then used to simulate the outcome of iterating the task over multiple generations. Participants in the grid search task scored higher after observing higher-scoring examples, but could only beat the scores of low-scoring example trials. Scores did not differ significantly between the control and WM distractor blocks, although more errors were made when under WM load. The simulation showed similar levels of cumulative score improvement across conditions. However, scores plateaued without reaching the maximum. Metacognitive efficiency was low in both blocks, with no indication of dual-task interference. Overall, we found that taxing working-memory resources did not prevent cumulative score improvement on this task, but impeded it slightly relative to a control distractor task. However, we found no evidence that the dual-task manipulation impacted participants' ability to use explicit metacognition. Although we found minimal evidence in support of the explicit metacognition theory of cumulative culture, our results provide valuable insights into empirical approaches that could be used to further test predictions arising from this account.

Tradition and invention: The bifocal stance theory of cultural evolution.

Cultural evolution depends on both innovation (the creation of new cultural variants by accident or design) and high-fidelity transmission (which preserves our accumulated knowledge and allows the storage of normative conventions). What is required is an overarching theory encompassing both dimensions, specifying the psychological motivations and mechanisms involved. The bifocal stance theory (BST) of cultural evolution proposes that the co-existence of innovative change and stable tradition results from our ability to adopt different motivational stances flexibly during social learning and transmission. We argue that the ways in which instrumental and ritual stances are adopted in cultural transmission influence the nature and degree of copying fidelity and thus also patterns of cultural spread and stability at a population level over time. BST creates a unifying framework for interpreting the findings of otherwise seemingly disparate areas of enquiry, including social learning, cumulative culture, overimitation, and ritual performance. We discuss the implications of BST for competing by-product accounts which assume that faithful copying is merely a side-effect of instrumental learning and action parsing. We also set out a novel "cultural action framework" bringing to light aspects of social learning that have been relatively neglected by behavioural ecologists and evolutionary psychologists and establishing a roadmap for future research on this topic. The BST framework sheds new light on the cognitive underpinnings of cumulative cultural change, selection, and spread within an encompassing evolutionary framework.

Human cumulative culture and the exploitation of natural phenomena.

Cumulative cultural evolution (CCE)-defined as the process by which beneficial modifications are culturally transmitted and progressively accumulated over time-has long been argued to underlie the unparalleled diversity and complexity of human culture. In this paper, I argue that not just any kind of cultural accumulation will give rise to human-like culture. Rather, I suggest that human CCE depends on the gradual exploitation of natural phenomena, which are features of our environment that, through the laws of physics, chemistry or biology, generate reliable effects which can be exploited for a purpose. I argue that CCE comprises two distinct processes: optimizing cultural traits that exploit a given set of natural phenomena (Type I CCE) and expanding the set of natural phenomena we exploit (Type II CCE). I argue that the most critical features of human CCE, including its open-ended dynamic, stems from Type II CCE. Throughout the paper, I contrast the two processes and discuss their respective socio-cognitive requirements. This article is part of a discussion meeting issue 'The emergence of collective knowledge and cumulative culture in animals, humans and machines'.

The origins of human cumulative culture: from the foraging niche to collective intelligence.

Various studies have investigated cognitive mechanisms underlying culture in humans and other great apes. However, the adaptive reasons for the evolution of uniquely sophisticated cumulative culture in our species remain unclear. We propose that the cultural capabilities of humans are the evolutionary result of a stepwise transition from the ape-like lifestyle of earlier hominins to the foraging niche still observed in extant hunter-gatherers. Recent ethnographic, archaeological and genetic studies have provided compelling evidence that the components of the foraging niche (social egalitarianism, sexual and social division of labour, extensive co-residence and cooperation with unrelated individuals, multilocality, fluid sociality and high between-camp mobility) engendered a unique multilevel social structure where the cognitive mechanisms underlying cultural evolution (high-fidelity transmission, innovation, teaching, recombination, ratcheting) evolved as adaptations. Therefore, multilevel sociality underlies a 'social ratchet' or irreversible task specialization splitting the burden of cultural knowledge across individuals, which may explain why human collective intelligence is uniquely able to produce sophisticated cumulative culture. The foraging niche perspective may explain why a complex gene-culture dual inheritance system evolved uniquely in humans and interprets the cultural, morphological and genetic origins of Homo sapiens as a process of recombination of innovations appearing in differentiated but interconnected populations. This article is part of a discussion meeting issue 'The emergence of collective knowledge and cumulative culture in animals, humans and machines'.