Structuralism and Adaptationism: Friends? Or foes?

Historically, the empirical study of phenotypic diversification has fallen into two rough camps; (1) "structuralist approaches" focusing on developmental constraint, bias, and innovation (with evo-devo at the core); and (2) "adaptationist approaches" focusing on adaptation, and natural selection. Whilst debates, such as that surrounding the proposed "Extended" Evolutionary Synthesis, often juxtapose these two positions, this review focuses on the grey space in between. Specifically, here I present a novel analysis of structuralism which enables us to take a more nuanced look at the motivations behind the structuralist and adaptationist positions. This makes clear how the two approaches can conflict, and points of potential commensurability. The review clarifies (a) the value of the evo-devo approach to phenotypic diversity, but also (b) how it properly relates to other predominant approaches to the same issues in evolutionary biology more broadly.

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.

Adaptation and its analogues: Biological categories for biosemantics.

"Teleosemantic" or "biosemantic" theories form a strong naturalistic programme in the philosophy of mind and language. They seek to explain the nature of mind and language by recourse to a natural history of "proper functions" as selected-for effects of language- and thought-producing mechanisms. However, they remain vague with respect to the nature of the proposed analogy between selected-for effects on the biological level and phenomena that are not strictly biological, such as reproducible linguistic and cultural forms. This essay critically explores various interpretations of this analogy. It suggests that these interpretations can be explicated by contrasting adaptationist with pluralist readings of the evolutionary concept of adaptation. Among the possible interpretations of the relations between biological adaptations and their analogues in language and culture, the two most relevant are a linear, hierarchical, signalling-based model that takes its cues from the evolution of co-operation and joint intentionality and a mutualistic, pluralist model that takes its cues from mimesis and symbolism in the evolution of human communication. Arguing for the merits of the mutualistic model, the present analysis indicates a path towards an evolutionary pluralist version of biosemantics that will align with theories of cognition as being environmentally "scaffolded". Language and other cultural forms are partly independent reproducible structures that acquire proper functions of their own while being integrated with organism-based cognitive traits in co-evolutionary fashion.

The evolution of altruism in humans.

Humans are an intensely social species, frequently performing costly behaviors that benefit others. Efforts to solve the evolutionary puzzle of altruism have a lengthy history, and recent years have seen many important advances across a range of disciplines. Here we bring together this interdisciplinary body of research and review the main theories that have been proposed to explain human prosociality, with an emphasis on kinship, reciprocity, indirect reciprocity, punishment, and morality. We highlight recent methodological advances that are stimulating research and point to some areas that either remain controversial or merit more attention.

Diversity and its causes: Lewontin on racism, biological determinism and the adaptationist programme.

Lewontin's 1972 paper (RC Lewontin, 1972 The apportionment of human diversity, in Evolutionary biology, vol. 6 (eds T Dobzhansky, MK Hecht, WC Steere), pp. 381-398) can be viewed as one foray in his battle against biological determinism. Our paper shows where Lewontin, The apportionment of human diversity, fits in the debate over human classification that it stimulated. We outline three assumptions inherent in the biological deterministic view of human phenotypic diversity and show how the 1972 paper, as well as Lewontin's papers in 1970 and 1974 on the problems with the heritability statistic and his 1979 criticism of naive pan-selectionism, invalidate these assumptions. These papers were crucial components of his campaign against biological determinism and the racism with which it was associated. In the current climate of widespread racism and the rise of sociogenomics, it is important to revisit Lewontin's writings and to disseminate the messages they contain. This article is part of the theme issue 'Celebrating 50 years since Lewontin's apportionment of human diversity'.

The Road Not Taken: What Developmental Psychology Might Learn From Darwin's Insights Concerning Sexual Selection.

Developmental Psychology is the branch of psychology that studies, not only human behavior, but how and why human behavior changes over time. This essay seeks to review to what extent Developmental Psychology has failed to perceive human behavior through the lens of evolutionary theory in general, and in particular sexual selection as first described by Darwin and later elaborated on by many, including Robert Trivers and Geoffrey Miller; the essay asserts that this failure has resulted in many wrong turns and missed opportunities. In some cases, major developmental theorists (e.g., Freud, Erikson) were bedeviled by sex-based differences which they saw but could not explain and which compromised the parsimony of their stage theories. In the case of stage theories of moral development, some major theorists (e.g., Piaget, Kohlberg) were able to offer simpler explanations of moral development only by limiting their studies to male subjects. And, while Developmental Psychology textbooks thoroughly describe sex differences in the timing of morphological changes in puberty, writers seldom discuss why the timing is different in the two sexes, universally, and functionally. On the other hand, several domains of developmental focus, including play, mate choice, parenting, and spatial cognition, have seen successful research efforts that utilized sexually selected predispositions as foundational assumptions. The essay concludes with a discussion of how a more evolutionary and functional view of human behavior might move the field of Developmental Psychology to an even more robust and accurate understanding of how humans change over the course of a lifetime.

Are Emotions Natural Kinds After All? Rethinking the Issue of Response Coherence.

The synchronized co-activation of multiple responses-motivational, behavioral, and physiological-has been taken as a defining feature of emotion. Such response coherence has been observed inconsistently however, and this has led some to view emotion programs as lacking biological reality. Yet, response coherence is not always expected or desirable if an emotion program is to carry out its adaptive function. Rather, the hallmark of emotion is the capacity to orchestrate multiple mechanisms adaptively-responses will co-activate in stereotypical fashion or not depending on how the emotion orchestrator interacts with the situation. Nevertheless, might responses cohere in the general case where input variables are specified minimally? Here we focus on shame as a case study. We measure participants' responses regarding each of 27 socially devalued actions and personal characteristics. We observe internal and external coherence: The intensities of felt shame and of various motivations of shame (hiding, lying, destroying evidence, and threatening witnesses) vary in proportion (i) to one another, and (ii) to the degree to which audiences devalue the disgraced individual-the threat shame defends against. These responses cohere both within and between the United States and India. Further, alternative explanations involving the low-level variable of arousal do not seem to account for these results, suggesting that coherence is imparted by a shame system. These findings indicate that coherence can be observed at multiple levels and raise the possibility that emotion programs orchestrate responses, even in those situations where coherence is low.

Differences in evolutionary accessibility determine which equally effective regulatory motif evolves to generate pulses.

Transcriptional regulatory networks (TRNs) are enriched for certain "motifs." Motif usage is commonly interpreted in adaptationist terms, i.e., that the optimal motif evolves. But certain motifs can also evolve more easily than others. Here, we computationally evolved TRNs to produce a pulse of an effector protein. Two well-known motifs, type 1 incoherent feed-forward loops (I1FFLs) and negative feedback loops (NFBLs), evolved as the primary solutions. The relative rates at which these two motifs evolve depend on selection conditions, but under all conditions, either motif achieves similar performance. I1FFLs generally evolve more often than NFBLs. Selection for a tall pulse favors NFBLs, while selection for a fast response favors I1FFLs. I1FFLs are more evolutionarily accessible early on, before the effector protein evolves high expression; when NFBLs subsequently evolve, they tend to do so from a conjugated I1FFL-NFBL genotype. In the empirical S. cerevisiae TRN, output genes of NFBLs had higher expression levels than those of I1FFLs. These results suggest that evolutionary accessibility, and not relative functionality, shapes which motifs evolve in TRNs, and does so as a function of the expression levels of particular genes.

What we (should) talk about when we talk about fruitfulness.

What are the relevant values to the appraisal of research programs? This question remains hotly debated, as philosophers have recently proposed many lists of values potentially relevant to scientific appraisal. Surprisingly, despite being mentioned in many lists, little attention has been paid to fruitfulness. It is unclear how fruitfulness should be explicated, and whether it has any substantial role in scientific appraisal. In this paper, I argue we should explicate fruitfulness as the capacity to develop of research programs. Moreover, I provide a novel strategy to assess and compare the fruitfulness of programs focused on their research questions and heuristics. To illustrate how this strategy would work, I will discuss a case study, namely the adaptationist program in evolutionary psychology.

Gender as a historical kind: a tale of two genders?

Is there anything that members of each binary category of gender have in common? Even many non-essentialists find the lack of unity within a gender worrying as it undermines the basis for a common political agenda for women. One promising proposal for achieving unity is by means of a shared historical lineage of cultural reproduction with past binary models of gender (e.g. Bach in Ethics 122:231-272, 2012). I demonstrate how such an account is likely to take on board different binary and also non-binary systems of gender. This implies that all individuals construed as members of the category, "women" are in fact not members of the same historical kind after all! I then consider different possible means of modifying the account but conclude negatively: the problem runs deeper than has been appreciated thus far.

Testing the Controversy : An Empirical Examination of Adaptationists' Attitudes Toward Politics and Science.

Critics of evolutionary psychology and sociobiology have advanced an adaptationists-as-right-wing-conspirators (ARC) hypothesis, suggesting that adaptationists use their research to support a right-wing political agenda. We report the first quantitative test of the ARC hypothesis based on an online survey of political and scientific attitudes among 168 US psychology Ph.D. students, 31 of whom self-identified as adaptationists and 137 others who identified with another non-adaptationist meta-theory. Results indicate that adaptationists are much less politically conservative than typical US citizens and no more politically conservative than non-adaptationist graduate students. Also, contrary to the "adaptationists-as-pseudo-scientists" stereotype, adaptationists endorse more rigorous, progressive, quantitative scientific methods in the study of human behavior than non-adaptationists.

A General Statistical Method for Identifying Adaptations by Parameterizing Trait Space.

It is obviously useful to think of evolved individuals in terms of their adaptations, yet the task of empirically classifying traits as adaptations has been claimed by some to be impossible in principle. I reject that claim by construction, introducing a formal method to empirically test whether a trait is an adaptation. The method presented is general, intuitive, and effective at identifying adaptations while remaining agnostic about their adaptive function. The test follows directly from the notion that adaptations arise from variation, heritability, and differential fitness in an evolving population: I operationalize these three concepts at the trait level, formally defining measures of individual traits. To test whether a trait is an adaptation, these measures are evaluated, locating the trait within a three-dimensional parameterized trait space. Within this space, I identify a region containing all adaptations; a trait's position relative to this adaptive region of trait space describes its status as an adaptation. The test can be applied in any evolving system where a few domain-specific statistical measures can be constructed; I demonstrate the construction of these measures, most notably a measure of an individual's hypothetical fitness if it were born with a different trait, in Packard's Bugs ALife model. The test is applied in Bugs, and shown to conform with our intuitive classification of adaptations.

Evolutionary perspective of cancer: myth, metaphors, and reality.

The evolutionary perspective of cancer (which origins and dynamics result from evolutionary processes) has gained significant international recognition over the past decade and generated a wave of enthusiasm among researchers. In this context, several authors proposed that insights into evolutionary and adaptation dynamics of cancers can be gained by studying the evolutionary strategies of organisms. Although this reasoning is fundamentally correct, in our opinion, it contains a potential risk of excessive adaptationism, potentially leading to the suggestion of complex adaptations that are unlikely to evolve among cancerous cells. For example, the ability of recognizing related conspecifics and adjusting accordingly behaviors as in certain free-living species appears unlikely in cancer. Indeed, despite their rapid evolutionary rate, malignant cells are under selective pressures for their altered lifestyle for only few decades. In addition, even though cancer cells can theoretically display highly sophisticated adaptive responses, it would be crucial to determine the frequency of their occurrence in patients with cancer, before therapeutic applications can be considered. Scientists who try to explain oncogenesis will need in the future to critically evaluate the metaphorical comparison of selective processes affecting cancerous cells with those affecting organisms. This approach seems essential for the applications of evolutionary biology to understand the origin of cancers, with prophylactic and therapeutic applications.

Engineering and Biology: Counsel for a Continued Relationship.

Biologists frequently draw on ideas and terminology from engineering. Evolutionary systems biology-with its circuits, switches, and signal processing-is no exception. In parallel with the frequent links drawn between biology and engineering, there is ongoing criticism against this cross-fertilization, using the argument that over-simplistic metaphors from engineering are likely to mislead us as engineering is fundamentally different from biology. In this article, we clarify and reconfigure the link between biology and engineering, presenting it in a more favorable light. We do so by, first, arguing that critics operate with a narrow and incorrect notion of how engineering actually works, and of what the reliance on ideas from engineering entails. Second, we diagnose and diffuse one significant source of concern about appeals to engineering, namely that they are inherently and problematically metaphorical. We suggest that there is plenty of fertile ground left for a continued, healthy relationship between engineering and biology.

Darwin and Spencer on the origin of music: is music the food of love?

Finding an evolutionary explanation for the origins of music serves as a rich test of broader ideas on the emergence of mind and the evolution of mental processes. Charles Darwin and Herbert Spencer both offered evolutionary explanations for the origins of music, indicating the importance of the question for these two leading nineteenth-century students of "descent with modification." Their discussion unfolded between the publication of Spencer's "The origin and function of music" in 1857 and Darwin's commentaries on music in The Descent of Man in 1871 with an addendum Spencer offered to his original article in light of Darwin's views. They had conflicting views on the lines of causation, asked differing questions, and had fundamentally different approaches. Their exchange laid the foundation for the discussion among contemporary adaptationists and nonadaptationists and contributed to the thinking of those who argue for Mixed Origins of Music or that it is a Transformative Technology of Mind.

The niche construction perspective: a critical appraisal.

Niche construction refers to the activities of organisms that bring about changes in their environments, many of which are evolutionarily and ecologically consequential. Advocates of niche construction theory (NCT) believe that standard evolutionary theory fails to recognize the full importance of niche construction, and consequently propose a novel view of evolution, in which niche construction and its legacy over time (ecological inheritance) are described as evolutionary processes, equivalent in importance to natural selection. Here, we subject NCT to critical evaluation, in the form of a collaboration between one prominent advocate of NCT, and a team of skeptics. We discuss whether niche construction is an evolutionary process, whether NCT obscures or clarifies how natural selection leads to organismal adaptation, and whether niche construction and natural selection are of equivalent explanatory importance. We also consider whether the literature that promotes NCT overstates the significance of niche construction, whether it is internally coherent, and whether it accurately portrays standard evolutionary theory. Our disagreements reflect a wider dispute within evolutionary theory over whether the neo-Darwinian synthesis is in need of reformulation, as well as different usages of some key terms (e.g., evolutionary process).

The mismeasure of machine: Synthetic biology and the trouble with engineering metaphors.

The scientific study of living organisms is permeated by machine and design metaphors. Genes are thought of as the "blueprint" of an organism, organisms are "reverse engineered" to discover their functionality, and living cells are compared to biochemical factories, complete with assembly lines, transport systems, messenger circuits, etc. Although the notion of design is indispensable to think about adaptations, and engineering analogies have considerable heuristic value (e.g., optimality assumptions), we argue they are limited in several important respects. In particular, the analogy with human-made machines falters when we move down to the level of molecular biology and genetics. Living organisms are far more messy and less transparent than human-made machines. Notoriously, evolution is an opportunistic tinkerer, blindly stumbling on "designs" that no sensible engineer would come up with. Despite impressive technological innovation, the prospect of artificially designing new life forms from scratch has proven more difficult than the superficial analogy with "programming" the right "software" would suggest. The idea of applying straightforward engineering approaches to living systems and their genomes-isolating functional components, designing new parts from scratch, recombining and assembling them into novel life forms-pushes the analogy with human artifacts beyond its limits. In the absence of a one-to-one correspondence between genotype and phenotype, there is no straightforward way to implement novel biological functions and design new life forms. Both the developmental complexity of gene expression and the multifarious interactions of genes and environments are serious obstacles for "engineering" a particular phenotype. The problem of reverse-engineering a desired phenotype to its genetic "instructions" is probably intractable for any but the most simple phenotypes. Recent developments in the field of bio-engineering and synthetic biology reflect these limitations. Instead of genetically engineering a desired trait from scratch, as the machine/engineering metaphor promises, researchers are making greater strides by co-opting natural selection to "search" for a suitable genotype, or by borrowing and recombining genetic material from extant life forms.

Los nuevos pilaees de la teoría evolutiva a la luz de la genómica / New Pillars of Evolutionary Theory in the Light of Genomics

La teoría de evolución de Darwin se ha constituido en uno de los pilares fundamentales de la biología. Algunos supuestos de dicha teoría han sido complementados con las bases genéticas de la herencia dando origen a la teoría moderna o sintética de la evolución. En los últimos años se han logrado grandes avances en el área de secuenciación y análisis de genomas completos que han aportado elementos para debatir varios de los postulados darwinistas de la evolución. La demostración de procesos de duplicación de genes y de genomas completos, la transferencia horizontal de genes y la teoría endosimbiótica cuestionan la idea de cambio evolutivo como un proceso de acumulación de pequeños cambios a través del tiempo geológico. La evidencia de la selección neutral en el contexto genómico pone de manifiesto otros mecanismos de evolución, que no necesariamente van acompañados de un programa adaptacionista ni con la idea de progreso. En este artículo se presentan estos y otros conceptos adicionales tales como la regulación génica y los mecanismos moleculares del desarrollo y del ambiente como puntos de partida para el planteamiento de una teoría evolutiva incluyente y acorde a los conocimientos actuales.

The evolutionist theory proposed by Darwin is one of the fundamental pillars in biology. Darwin’s theory was solidified with the Modern synthesis of evolutionary biology thanks to the rediscovery of Mendel’s work, which laid the genetic basis of heredity. In recent years, great progress has been acquired in the sequencing and analyses of complete genomes, which have provided several elements to discuss some Darwinists tenets of evolution. The evidence of gene duplication and whole-genome duplication, the horizontal gene transfer and the endosymbiosis process question the idea that evolution proceeds through the gradual accumulation of infinitesimally small random changes. The new evidence of neutral selection on the genomics context reveals other mechanisms of evolution not necessarily related with the idea of progress or with an adaptationist program as was originally stated by the Darwin’s theory. In this paper, I present these and other concepts such as gene regulation, molecular mechanisms of development and some environmental aspects (epigenesis and phenotypic plasticity) as starting points to think in the necessity to update the evolutionary theory which in my opinion should be more inclusive, pluralistic and consistent with our current knowledge.