Variable cultural acquisition costs constrain cumulative cultural evolution

One of the hallmarks of the human species is our capacity for cumulative culture, in which beneficial knowledge and technology is accumulated over successive generations. Yet previous analyses of cumulative cultural change have failed to consider the possibility that as cultural complexity accumulates, it becomes increasingly costly for each new generation to acquire from the previous generation. In principle this may result in an upper limit on the cultural complexity that can be accumulated, at which point accumulated knowledge is so costly and time-consuming to acquire that further innovation is not possible. In this paper I first review existing empirical analyses of the history of science and technology that support the possibility that cultural acquisition costs may constrain cumulative cultural evolution. I then present macroscopic and individual-based models of cumulative cultural evolution that explore the consequences of this assumption of variable cultural acquisition costs, showing that making acquisition costs vary with cultural complexity causes the latter to reach an upper limit above which no further innovation can occur. These models further explore the consequences of different cultural transmission rules (directly biased, indirectly biased and unbiased transmission), population size, and cultural innovations that themselves reduce innovation or acquisition costs.     

Why does human culture increase exponentially?

Historical records show that culture can increase exponentially in time, e.g., in number of poems, musical works, scientific discoveries. We model how human capacities for creativity and cultural transmission may make such an increase possible, suggesting that: (1) creativity played a major role at the origin of human culture and for its accumulation throughout history, because cultural transmission cannot, on its own, generate exponentially increasing amounts of culture; (2) exponential increase in amount of culture can only occur if creativity is positively influenced by culture. The evolution of cultural transmission is often considered the main genetic bottleneck for the origin of culture, because natural selection cannot favor cultural transmission without any culture to transmit. Our models suggest that an increase in individual creativity may have been the first step toward human culture, because in a population of creative individuals there may be enough non-genetic information to favor the evolution of cultural transmission.     

Stone toolmaking and the evolution of human culture and cognition

Although many species display behavioural traditions, human culture is unique in the complexity of its technological, symbolic and social contents. Is this extraordinary complexity a product of cognitive evolution, cultural evolution or some interaction of the two? Answering this question will require a much better understanding of patterns of increasing cultural diversity, complexity and rates of change in human evolution. Palaeolithic stone tools provide a relatively abundant and continuous record of such change, but a systematic method for describing the complexity and diversity of these early technologies has yet to be developed. Here, an initial attempt at such a system is presented. Results suggest that rates of Palaeolithic culture change may have been underestimated and that there is a direct relationship between increasing technological complexity and diversity. Cognitive evolution and the greater latitude for cultural variation afforded by increasingly complex technologies may play complementary roles in explaining this pattern.     

Cumulative cultural dynamics and the coevolution of cultural innovation and transmission: an ESS model for panmictic and structured populations

When individuals in a population can acquire traits through learning, each individual may express a certain number of distinct cultural traits. These traits may have been either invented by the individual himself or acquired from others in the population. Here, we develop a game theoretic model for the accumulation of cultural traits through individual and social learning. We explore how the rates of innovation, decay, and transmission of cultural traits affect the evolutionary stable (ES) levels of individual and social learning and the number of cultural traits expressed by an individual when cultural dynamics are at a steady‐state. We explore the evolution of these phenotypes in both panmictic and structured population settings. Our results suggest that in panmictic populations, the ES level of learning and number of traits tend to be independent of the social transmission rate of cultural traits and is mainly affected by the innovation and decay rates. By contrast, in structured populations, where interactions occur between relatives, the ES level of learning and the number of traits per individual can be increased (relative to the panmictic case) and may then markedly depend on the transmission rate of cultural traits. This suggests that kin selection may be one additional solution to Rogers's paradox of nonadaptive culture.     

Evolutionary tracking is determined by differential selection on demographic rates and density dependence

Recent ecological forecasts predict that ~25% of species worldwide will go extinct by 2050. However, these estimates are primarily based on environmental changes alone and fail to incorporate important biological mechanisms such as genetic adaptation via evolution. Thus, environmental change can affect population dynamics in ways that classical frameworks can neither describe nor predict. Furthermore, often due to a lack of data, forecasting models commonly describe changes in population demography by summarizing changes in fecundity and survival concurrently with the intrinsic growth rate (r). This has been shown to be an oversimplification as the environment may impose selective pressure on specific demographic rates (birth and death) rather than directly on r (the difference between the birth and death rates). This differential pressure may alter population response to density, in each demographic rate, further diluting the information combined to produce r. Thus, when we consider the potential for persistence via adaptive evolution, populations with the same r can have different abilities to persist amidst environmental change. Therefore, we cannot adequately forecast population response to climate change without accounting for demography and selection on density dependence. Using a continuous‐time Markov chain model to describe the stochastic dynamics of the logistic model of population growth and allow for trait evolution via mutations arising during birth events, we find persistence via evolutionary tracking more likely when environmental change alters birth rather than the death rate. Furthermore, species that evolve responses to changes in the strength of density dependence due to environmental change are less vulnerable to extinction than species that undergo selection independent of population density. By incorporating these key demographic considerations into our predictive models, we can better understand how species will respond to climate change.     

Gene-culture coevolutionary theory

Gene-culture coevolutionary theory is a branch of theoretical population genetics that models the transmission of genes and cultural traits from one generation to the next, exploring how they interact. These models have been employed to examine the adaptive advantages of learning and culture, to investigate the forces of cultural change, to partition the variance in complex human behavioral and personality traits, and to address specific cases in human evolution in which there is an interaction between genes and culture.     

Genome‐wide signature of local adaptation linked to variable CpG methylation in oak populations

It has long been known that adaptive evolution can occur through genetic mutations in DNA sequence, but it is unclear whether adaptive evolution can occur through analogous epigenetic mechanisms, such as through DNA methylation. If epigenetic variation contributes directly to evolution, species under threat of disease, invasive competition, climate change or other stresses would have greater stores of variation from which to draw. We looked for evidence of natural selection acting on variably methylated DNA sites using population genomic analysis across three climatologically distinct populations of valley oaks. We found patterns of genetic and epigenetic differentiations that indicate local adaptation is operating on large portions of the oak genome. While CHG methyl polymorphisms are not playing a significant role and would make poor targets for natural selection, our findings suggest that CpG methyl polymorphisms as a whole are involved in local adaptation, either directly or through linkage to regions under selection.     

Adaptive responses and disruptive effects: how major wildfire influences kinship-based social interactions in a forest marsupial

Environmental disturbance is predicted to play a key role in the evolution of animal social behaviour. This is because disturbance affects key factors underlying social systems, such as demography, resource availability and genetic structure. However, because natural disturbances are unpredictable there is little information on their effects on social behaviour in wild populations. Here, we investigated how a major wildfire affected cooperation (sharing of hollow trees) by a hollow-dependent marsupial. We based two alternative social predictions on the impacts of fire on population density, genetic structure and resources. We predicted an adaptive social response from previous work showing that kin selection in den-sharing develops as competition for den resources increases. Thus, kin selection should occur in burnt areas because the fire caused loss of the majority of hollow-bearing trees, but no detectable mortality. Alternatively, fire may have a disruptive social effect, whereby postfire home range-shifts 'neutralize' fine-scale genetic structure, thereby removing opportunities for kin selection between neighbours. Both predictions occurred: the disruptive social effect in burnt habitat and the adaptive social response in adjacent unburnt habitat. The latter followed a massive demographic influx to unburnt 'refuge' habitat that increased competition for dens, leading to a density-related kin selection response. Our results show remarkable short-term plasticity of animal social behaviour and demonstrate how the social effects of disturbance extend into undisturbed habitat owing to landscape-scale demographic shifts. We predicted long-term changes in kinship-based cooperative behaviour resulting from the genetic and resource impacts of forecast changes to fire regimes in these forests.     

Cultural transmission and the evolution of human behaviour: a general approach based on the Price equation

Transmitted culture can be viewed as an inheritance system somewhat independent of genes that is subject to processes of descent with modification in its own right. Although many authors have conceptualized cultural change as a Darwinian process, there is no generally agreed formal framework for defining key concepts such as natural selection, fitness, relatedness and altruism for the cultural case. Here, we present and explore such a framework using the Price equation. Assuming an isolated, independently measurable culturally transmitted trait, we show that cultural natural selection maximizes cultural fitness, a distinct quantity from genetic fitness, and also that cultural relatedness and cultural altruism are not reducible to or necessarily related to their genetic counterparts. We show that antagonistic coevolution will occur between genes and culture whenever cultural fitness is not perfectly aligned with genetic fitness, as genetic selection will shape psychological mechanisms to avoid susceptibility to cultural traits that bear a genetic fitness cost. We discuss the difficulties with conceptualizing cultural change using the framework of evolutionary theory, the degree to which cultural evolution is autonomous from genetic evolution, and the extent to which cultural change should be seen as a Darwinian process. We argue that the nonselection components of evolutionary change are much more important for culture than for genes, and that this and other important differences from the genetic case mean that different approaches and emphases are needed for cultural than genetic processes.     

Innovativeness, population size and cumulative cultural evolution

Henrich [Henrich, J., 2004. Demography and cultural evolution: how adaptive cultural processes can produce maladaptive losses—the Tasmanian case. Am. Antiquity 69, 197-214] proposed a model designed to show that larger population size facilitates cumulative cultural evolution toward higher skill levels. In this model, each newborn attempts to imitate the most highly skilled individual of the parental generation by directly-biased social learning, but the skill level he/she acquires deviates probabilistically from that of the exemplar (cultural parent). The probability that the skill level of the imitator exceeds that of the exemplar can be regarded as the innovation rate. After reformulating Henrich’s model rigorously, we introduce an overlapping-generations analog based on the Moran model and derive an approximate formula for the expected change per generation of the highest skill level in the population. For large population size, our overlapping-generations model predicts a much larger effect of population size than Henrich’s discrete-generations model. We then investigate by way of Monte Carlo simulations the case where each newborn chooses as his/her exemplar the most highly skilled individual from among a limited number of acquaintances. When the number of acquaintances is small relative to the population size, we find that a change in the innovation rate contributes more than a proportional change in population size to the cumulative cultural evolution of skill level.     

Rapid evolution of an adaptive cyanogenesis cline in introduced North American white clover (Trifolium repens L.)

White clover is polymorphic for cyanogenesis (HCN production after tissue damage), and this herbivore defence polymorphism has served as a classic model for studying adaptive variation. The cyanogenic phenotype requires two interacting biochemical components; the presence/absence of each component is controlled by a simple Mendelian gene (Ac/ac and Li/li). Climate-associated cyanogenesis clines occur in both native (Eurasian) and introduced populations worldwide, with cyanogenic plants predominating in warmer locations. Moreover, previous studies have suggested that epistatic selection may act within populations to maintain cyanogenic (AcLi) plants and acyanogenic plants that lack both components (acli plants) at the expense of plants possessing a single component (Acli and acLi plants). Here, we examine the roles of selection, gene flow and demography in the evolution of a latitudinal cyanogenesis cline in introduced North American populations. Using 1145 plants sampled across a 1650 km transect, we determine the distribution of cyanogenesis variation across the central United States and investigate whether clinal variation is adaptive or an artefact of population introduction history. We also test for the evidence of epistatic selection. We detect a clear latitudinal cline, with cyanogenesis frequencies increasing from 11% to 86% across the transect. Population structure analysis using nine microsatellite loci indicates that the cline is adaptive and not a by-product of demographic history. However, we find no evidence for epistatic selection within populations. Our results provide strong evidence for rapid adaptive evolution in these introduced populations, and they further suggest that the mechanisms maintaining adaptive variation may vary among populations of a species.     

How Darwinian is cultural evolution?

Darwin-inspired population thinking suggests approaching culture as a population of items of different types, whose relative frequencies may change over time. Three nested subtypes of populational models can be distinguished: evolutionary, selectional and replicative. Substantial progress has been made in the study of cultural evolution by modelling it within the selectional frame. This progress has involved idealizing away from phenomena that may be critical to an adequate understanding of culture and cultural evolution, particularly the constructive aspect of the mechanisms of cultural transmission. Taking these aspects into account, we describe cultural evolution in terms of cultural attraction, which is populational and evolutionary, but only selectional under certain circumstances. As such, in order to model cultural evolution, we must not simply adjust existing replicative or selectional models but we should rather generalize them, so that, just as replicator-based selection is one form that Darwinian selection can take, selection itself is one of several different forms that attraction can take. We present an elementary formalization of the idea of cultural attraction.     

Identifying innovation in laboratory studies of cultural evolution: rates of retention and measures of adaptation

In recent years, laboratory studies of cultural evolution have become increasingly prevalent as a means of identifying and understanding the effects of cultural transmission on the form and functionality of transmitted material. The datasets generated by these studies may provide insights into the conditions encouraging, or inhibiting, high rates of innovation, as well as the effect that this has on measures of adaptive cultural change. Here we review recent experimental studies of cultural evolution with a view to elucidating the role of innovation in generating observed trends. We first consider how tasks are presented to participants, and how the corresponding conceptualization of task success is likely to influence the degree of intent underlying any deviations from perfect reproduction. We then consider the measures of interest used by the researchers to track the changes that occur as a result of transmission, and how these are likely to be affected by differing rates of retention. We conclude that considering studies of cultural evolution from the perspective of innovation provides us with valuable insights that help to clarify important differences in research designs, which have implications for the likely effects of variation in retention rates on measures of cultural adaptation.     

Origins of altruism diversity ii: runaway coevolution of altruistic strategies via "reciprocal niche construction"

Understanding the evolution of altruism requires knowledge of both its constraints and its drivers. Here we show that, paradoxically, ecological constraints on altruism may ultimately be its strongest driver. We construct a two-trait, coevolutionary adaptive dynamics model of social evolution in a genetically structured population with local resource competition. The intensity of local resource competition, which influences the direction and strength of social selection and which is typically treated as a static parameter, is here allowed to be an evolvable trait. Evolution of survival/fecundity altruism, which requires weak local competition, increases local competition as it evolves, creating negative environmental feedback that ultimately inhibits its further evolutionary advance. Alternatively, evolution of resource-based altruism, which requires strong local competition, weakens local competition as it evolves, also ultimately causing its own evolution to stall. When evolving independently, these altruistic strategies are intrinsically self-limiting. However, the coexistence of these two altruism types transforms the negative ecoevolutionary feedback generated by each strategy on itself into positive feedback on the other, allowing the presence of one trait to drive the evolution of the other. We call this feedback conversion "reciprocal niche construction." In the absence of constraints, this process leads to runaway coevolution of altruism types. We discuss applications to the origins and evolution of eusociality, division of labor, the inordinate ecological success of eusocial species, and the interaction between technology and demography in human evolution. Our theory suggests that the evolution of extreme sociality may often be an autocatalytic process.     

DIRECTIONAL CULTURAL CHANGE BY MODIFICATION AND REPLACEMENT OF MEMES

Evolutionary approaches to culture remain contentious. A source of contention is that cultural mutation may be substantial and, if it drives cultural change, then current evolutionary models are not adequate. But we lack studies quantifying the contribution of mutations to directional cultural change. We estimated the contribution of one type of cultural mutations—modification of memes—to directional cultural change using an amenable study system: learned birdsongs in a species that recently entered an urban habitat. Many songbirds have higher minimum song frequency in cities, to alleviate masking by low‐frequency noise. We estimated that the input of meme modifications in an urban songbird population explains about half the extent of the population divergence in song frequency. This contribution of cultural mutations is large, but insufficient to explain the entire population divergence. The remaining divergence is due to selection of memes or creation of new memes. We conclude that the input of cultural mutations can be quantitatively important, unlike in genetic evolution, and that it operates together with other mechanisms of cultural evolution. For this and other traits, in which the input of cultural mutations might be important, quantitative studies of cultural mutation are necessary to calibrate realistic models of cultural evolution.     

The reality and importance of founder speciation in evolution

A founder event occurs when a new population is established from a small number of individuals drawn from a large ancestral population. Mayr proposed that genetic drift in an isolated founder population could alter the selective forces in an epistatic system, an observation supported by recent studies. Carson argued that a period of relaxed selection could occur when a founder population is in an open ecological niche, allowing rapid population growth after the founder event. Selectable genetic variation can actually increase during this founder-flush phase due to recombination, enhanced survival of advantageous mutations, and the conversion of non-additive genetic variance into additive variance in an epistatic system, another empirically confirmed prediction. Templeton combined the theories of Mayr and Carson with population genetic models to predict the conditions under which founder events can contribute to speciation, and these predictions are strongly confirmed by the empirical literature. Much of the criticism of founder speciation is based upon equating founder speciation to an adaptive peak shift opposed by selection. However, Mayr, Carson and Templeton all modeled a positive interaction of selection and drift, and Templeton showed that founder speciation is incompatible with peak-shift conditions. Although rare, founder speciation can have a disproportionate importance in adaptive innovation and radiation, and examples are given to show that "rare" does not mean "unimportant" in evolution. Founder speciation also interacts with other speciation mechanisms such that a speciation event is not a one-dimensional process due to either selection alone or drift alone.     

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).     

Complexity and Demographic Explanations of Cumulative Culture

Formal models have linked prehistoric and historical instances of technological change (e.g., the Upper Paleolithic transition, cultural loss in Holocene Tasmania, scientific progress since the late nineteenth century) to demographic change. According to these models, cumulation of technological complexity is inhibited by decreasing— while favoured by increasing—population levels. Here we show that these findings are contingent on how complexity is defined: demography plays a much more limited role in sustaining cumulative culture in case formal models deploy Herbert Simon''s definition of complexity rather than the particular definitions of complexity hitherto assumed. Given that currently available empirical evidence doesn''t afford discriminating proper from improper definitions of complexity, our robustness analyses put into question the force of recent demographic explanations of particular episodes of cultural change.     

Social complexity and linguistic diversity in the Austronesian and Bantu population expansions

Reconstructing the rise and fall of social complexity in human societies through time is fundamental for understanding some of the most important transformations in human history. Phylogenetic methods based on language diversity provide a means to reconstruct pre-historic events and model the transition rates of cultural change through time. We model and compare the evolution of social complexity in Austronesian (n = 88) and Bantu (n = 89) societies, two of the world's largest language families with societies representing a wide spectrum of social complexity. Our results show that in both language families, social complexity tends to build and decline in an incremental fashion, while the Austronesian phylogeny provides evidence for additional severe demographic bottlenecks. We suggest that the greater linguistic diversity of the Austronesian language family than Bantu likely follows the different biogeographic structure of the two regions. Cultural evolution in both the Bantu and Austronesian cases was not a simple linear process, but more of a wave-like process closely tied to the demography of expanding populations and the spatial structure of the colonized regions.     

Invasibility as an emergent property of native metapopulation structure

Biological invasions constitute major threats to global biodiversity. Eco‐evolutionary considerations highlight the importance of contemporary evolution in community responses to bioinvasions. However, effects of metapopulation structure on invasion success have been mostly overlooked even though metapopulation structure determines gene flow and is likely to affect evolutionary processes. Here, we investigate a stepping‐stone model with evolving alien native interaction strengths. We demonstrate analytically that the site of invasion can determine the success of an invading consumer because gene flow and demography of a local resource species interact to obstruct local resource adaptation. Our main results are 1) that invasion success is more likely in genetic sink populations of the native species and 2) that invasion is more likely to occur against the migrational flow of native species. These findings suggest that invasibility is best regarded as an emergent property not only of communities but of entire metapopulations. Since migration networks of aliens and natives are often mismatched due to anthropogenic interference, our results indicate how population structure eases the spread of invasives against the migrational flow of natives.