All Publications

  • Conditions that favour cumulative cultural evolution. Philosophical transactions of the Royal Society of London. Series B, Biological sciences Denton, K. K., Ram, Y., Feldman, M. W. 2023; 378 (1872): 20210400


    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'.

    View details for DOI 10.1098/rstb.2021.0400

    View details for PubMedID 36688392

  • Conformity and anti-conformity in a finite population. Journal of theoretical biology Lappo, E., Denton, K. K., Feldman, M. W. 2023: 111429


    Conformist and anti-conformist cultural transmission have been studied both empirically, in several species, and theoretically, with population genetic models. Building upon standard, infinite-population models (IPMs) of conformity, we introduce finite-population models (FPMs) and study them via simulation and a diffusion approximation. In previous IPMs of conformity, offspring observe the variants of n adult role models, where n is often three. Numerical simulations show that while the short-term behavior of the FPM with n=3 role models is well approximated by the IPM, stable polymorphic equilibria of the IPM become effective equilibria of the FPM at which the variation persists prior to fixation or loss, and which produce plateaus in curves for fixation probabilities and expected times to absorption. In the FPM with n=5 role models, the population may switch between two effective equilibria, which is not possible in the IPM, or may cycle between frequencies that are not effective equilibria, which is possible in the IPM. In all observed cases of 'equilibrium switching' and 'cycling' in the FPM, model parameters exceed O(1/N), required for the diffusion approximation, resulting in an over-estimation of the actual times to absorption. However, in those cases with n=5 role models that have one effective equilibrium and stable fixation states, even if conformity coefficients exceed O(1/N), the diffusion approximation matches closely the numerical simulations of the FPM. This suggests that the robustness of the diffusion approximation depends not only on the magnitudes of coefficients, but also on the qualitative behavior of the conformity model.

    View details for DOI 10.1016/j.jtbi.2023.111429

    View details for PubMedID 36746297

  • Social evolution and the what, when, why and how of the major evolutionary transitions in the history of life FRONTIERS IN ECOLOGY AND EVOLUTION Nonacs, P., Denton, K. K., Robin, A. N., Helantera, H., Kapheim, K. M. 2022; 10
  • Polychotomous traits and evolution under conformity. Proceedings of the National Academy of Sciences of the United States of America Denton, K. K., Liberman, U., Feldman, M. W. 2022; 119 (39): e2205914119


    Conformist and anticonformist transmission of dichotomous cultural traits (i.e., traits with two variants) have been studied both experimentally, in many species, and theoretically, with mathematical models. Signatures of types of conformity to polychotomous traits (with more than two variants; e.g., baby names and syllables in bird song) have been inferred from population-level data, but there are few models that include individual-level biases among more than two discrete variants. We generalize the standard dichotomous trait conformity model by Boyd and Richerson to incorporate [Formula: see text] role models and [Formula: see text] variants. Our analysis shows that in the case of [Formula: see text] role models, under anticonformity, the central polymorphic equilibrium [Formula: see text] is globally stable, whereas under conformity, if initially the frequencies of [Formula: see text] variants are all equal to the maximum variant frequency in the population, there is global convergence to an equilibrium in which the frequencies of these variants are all [Formula: see text] and all other variants are absent. With a general number n of role models, the same result holds with conformity, whereas under anticonformity, global convergence is not guaranteed, and there may be stable frequency cycles or chaos. If both conformity and anticonformity occur for different configurations of variants among the n role models, a variety of novel polymorphic equilibria may exist and be stable. Future empirical studies may use this formulation to directly quantify an individual's level of (anti)conformist bias to a polychotomous trait.

    View details for DOI 10.1073/pnas.2205914119

    View details for PubMedID 36122242

  • Major Evolutionary Transitions and the Roles of Facilitation and Information in Ecosystem Transformations FRONTIERS IN ECOLOGY AND EVOLUTION Robin, A. N., Denton, K. K., Horna Lowell, E. S., Dulay, T., Ebrahimi, S., Johnson, G. C., Mai, D., O'Fallon, S., Philson, C. S., Speck, H. P., Zhang, X., Nonacs, P. 2021; 09
  • Conformity and content-biased cultural transmission in the evolution of altruism. Theoretical population biology Denton, K. K., Ram, Y., Feldman, M. W. 2021


    The evolution of altruism has been extensively modeled under the assumption of genetic transmission, whereas the dynamics under cultural transmission are less well understood. Previous research has shown that cultural transmission can facilitate the evolution of altruism by increasing 1) the probability of adopting the altruistic phenotype, and 2) assortment between altruists. We incorporate vertical and oblique transmission, which can be conformist or anti-conformist, into models of parental care, sibling altruism, and altruism between individuals that meet assortatively. If oblique transmission is conformist, it becomes easier for altruism to invade a population of non-altruists as the probability of vertical transmission increases. If oblique transmission is anti-conformist, decreasing vertical transmission facilitates invasion by altruism in the assortative meeting model, whereas in other models, there is a trade-off: greater vertical transmission produces greater assortment among genetically related altruists, but lowers the probability of adopting altruism via anti-conformity. Compared to conditions for invasion under genetic transmission, e.g., Hamilton's rule, we show that invasion can be easier with sufficiently strong anti-conformity, and in some models, with sufficiently high assortment even if oblique transmission is conformist. We also explore invasion by an allele A that increases individuals' content bias for altruism, in the absence of other forms of cultural transmission. If costs and benefits combine additively, A invades under previously known conditions. If costs and benefits combine multiplicatively, invasion by A and by altruism become more difficult than in the corresponding additive models.

    View details for DOI 10.1016/j.tpb.2021.10.004

    View details for PubMedID 34793823

  • On randomly changing conformity bias in cultural transmission. Proceedings of the National Academy of Sciences of the United States of America Denton, K. K., Liberman, U., Feldman, M. W. 2021; 118 (34)


    Humans and nonhuman animals display conformist as well as anticonformist biases in cultural transmission. Whereas many previous mathematical models have incorporated constant conformity coefficients, empirical research suggests that the extent of (anti)conformity in populations can change over time. We incorporate stochastic time-varying conformity coefficients into a widely used conformity model, which assumes a fixed number n of "role models" sampled by each individual. We also allow the number of role models to vary over time ([Formula: see text]). Under anticonformity, nonconvergence can occur in deterministic and stochastic models with different parameter values. Even if strong anticonformity may occur, if conformity or random copying (i.e., neither conformity nor anticonformity) is expected, there is convergence to one of the three equilibria seen in previous deterministic models of conformity. Moreover, this result is robust to stochastic variation in [Formula: see text] However, dynamic properties of these equilibria may be different from those in deterministic models. For example, with random conformity coefficients, all equilibria can be stochastically locally stable simultaneously. Finally, we study the effect of randomly changing weak selection. Allowing the level of conformity, the number of role models, and selection to vary stochastically may produce a more realistic representation of the wide range of group-level properties that can emerge under (anti)conformist biases. This promises to make interpretation of the effect of conformity on differences between populations, for example those connected by migration, rather difficult. Future research incorporating finite population sizes and migration would contribute added realism to these models.

    View details for DOI 10.1073/pnas.2107204118

    View details for PubMedID 34417299

  • Cultural evolution of conformity and anticonformity. Proceedings of the National Academy of Sciences of the United States of America Denton, K. K., Ram, Y. n., Liberman, U. n., Feldman, M. W. 2020


    Conformist bias occurs when the probability of adopting a more common cultural variant in a population exceeds its frequency, and anticonformist bias occurs when the reverse is true. Conformist and anticonformist bias have been widely documented in humans, and conformist bias has also been observed in many nonhuman animals. Boyd and Richerson used models of conformist and anticonformist bias to explain the evolution of large-scale cooperation, and subsequent research has extended these models. We revisit Boyd and Richerson's original analysis and show that, with conformity based on more than three role models, the evolutionary dynamics can be more complex than previously assumed. For example, we show the presence of stable cycles and chaos under strong anticonformity and the presence of new equilibria when both conformity and anticonformity act at different variant frequencies, with and without selection. We also investigate the case of population subdivision with migration and find that the common claim that conformity can maintain between-group differences is not always true. Therefore, the effect of conformity on the evolution of cooperation by group selection may be more complicated than previously stated. Finally, using Feldman and Liberman's modifier approach, we investigate the conditions under which a rare modifier of the extent of conformity or the number of role models can invade a population. Understanding the dynamics of conformist- and anticonformist-biased transmission may have implications for research on human and nonhuman animal behavior, the evolution of cooperation, and frequency-dependent transmission in general.

    View details for DOI 10.1073/pnas.2004102117

    View details for PubMedID 32461360