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Scientific activities take place within the structured sets of ideas and assumptions that define a field and its practices. The conceptual framework of evolutionary biology emerged with the Modern Synthesis in the early twentieth century and has since expanded into a highly successful research program to explore the processes of diversification and adaptation. Nonetheless, the ability of that framework satisfactorily to accommodate the rapid advances in developmental biology, genomics and ecology has been questioned. We review some of these arguments, focusing on literatures (evo-devo, developmental plasticity, inclusive inheritance and niche construction) whose implications for evolution can be interpreted in two ways—one that preserves the internal structure of contemporary evolutionary theory and one that points towards an alternative conceptual framework. The latter, which we label the ‘extended evolutionary synthesis' (EES), retains the fundaments of evolutionary theory, but differs in its emphasis on the role of constructive processes in development and evolution, and reciprocal portrayals of causation. In the EES, developmental processes, operating through developmental bias, inclusive inheritance and niche construction, share responsibility for the direction and rate of evolution, the origin of character variation and organism–environment complementarity. We spell out the structure, core assumptions and novel predictions of the EES, and show how it can be deployed to stimulate and advance research in those fields that study or use evolutionary biology.

The role of natural selection in the evolution of adaptive phenotypes has undergone constant probing by evolutionary biologists, employing both theoretical and empirical approaches. As Darwin noted, natural selection can act together with other processes, including random changes in the frequencies of phenotypic differences that are not under strong selection, and changes in the environment, which may reflect evolutionary changes in the organisms themselves. As understanding of genetics developed after 1900, the new genetic discoveries were incorporated into evolutionary biology. The resulting general principles were summarized by Julian Huxley in his 1942 book Evolution: the modern synthesis. Here, we examine how recent advances in genetics, developmental biology and molecular biology, including epigenetics, relate to today's understanding of the evolution of adaptations. We illustrate how careful genetic studies have repeatedly shown that apparently puzzling results in a wide diversity of organisms involve processes that are consistent with neo-Darwinism. They do not support important roles in adaptation for processes such as directed mutation or the inheritance of acquired characters, and therefore no radical revision of our understanding of the mechanism of adaptive evolution is needed.

Humans, unlike any other multicellular species in Earth's history, have emerged as a global force that is transforming the ecology of an entire planet. It is no longer possible to understand, predict, or successfully manage ecological pattern, process, or change without understanding why and how humans reshape these over the long term. Here, a general causal theory is presented to explain why human societies gained the capacity to globally alter the patterns, processes, and dynamics of ecology and how these anthropogenic alterations unfold over time and space as societies themselves change over human generational time. Building on existing theories of ecosystem engineering, niche construction, inclusive inheritance, cultural evolution, ultrasociality, and social change, this theory of anthroecological change holds that sociocultural evolution of subsistence regimes based on ecosystem engineering, social specialization, and non-kin exchange, or \"sociocultural niche construction,\" is the main cause of both the long-term upscaling of human societies and their unprecedented transformation of the biosphere. Human sociocultural niche construction can explain, where classic ecological theory cannot, the sustained transformative effects of human societies on biogeography, ecological succession, ecosystem processes, and the ecological patterns and processes of landscapes, biomes, and the biosphere. Anthroecology theory generates empirically testable hypotheses on the forms and trajectories of long-term anthropogenic ecological change that have significant theoretical and practical implications across the subdisciplines of ecology and conservation. Though still at an early stage of development, anthroecology theory aligns with and integrates established theoretical frameworks including social-ecological systems, social metabolism, countryside biogeography, novel ecosystems, and anthromes. The \"fluxes of nature\" are fast becoming \"cultures of nature.\" To investigate, understand, and address the ultimate causes of anthropogenic ecological change, not just the consequences, human sociocultural processes must become as much a part of ecological theory and practice as biological and geophysical processes are now. Strategies for achieving this goal and for advancing ecological science and conservation in an increasingly anthropogenic biosphere are presented.

The study of cultural evolution now includes multiple theoretical frameworks. Despite common influence from Darwinian evolutionary theory, there is considerable diversity. Thus, we recognize those most influenced by the tenets of the Modern Synthesis (evolutionary archaeology, cultural transmission theory, and human behavioral ecology) and those most aligned more closely with concepts emerging in the Extended Evolutionary Synthesis (cultural macroevolution and evolutionary cognitive archaeology). There has been substantial debate between adherents of these schools of thought as to their appropriateness and priority for addressing the fundamentals of cultural evolution. I argue that theoretical diversity is necessary to address research questions arising from a complex archaeological record. Concepts associated with the Extended Evolutionary Synthesis may offer unique insights into the cultural evolutionary process.

The Modern Synthesis (MS) is the current paradigm in evolutionary biology. It was actually built by expanding on the conceptual foundations laid out by its predecessors, Darwinism and neo-Darwinism. For sometime now there has been talk of a new Extended Evolutionary Synthesis (EES), and this article begins to outline why we may need such an extension, and how it may come about. As philosopher Karl Popper has noticed, the current evolutionary theory is a theory of genes, and we still lack a theory of forms. The field began, in fact, as a theory of forms in Darwin's days, and the major goal that an EES will aim for is a unification of our theories of genes and of forms. This may be achieved through an organic grafting of novel concepts onto the foundational structure of the MS, particularly evolvability, phenotypic plasticity, epigenetic inheritance, complexity theory, and the theory of evolution in highly dimensional adaptive landscapes.

In recent years, a number of researchers have advocated extending the modern synthesis in evolutionary biology. One of the core arguments made in favour of an extension comes from work on soft inheritance systems, including transgenerational epigenetic effects, cultural transmission and niche construction. In this study, we outline this claim and then take issue with it. We argue that the focus on soft inheritance has led to a conflation of proximate and ultimate causation, which has in turn obscured key questions about biological organization and calibration across the life span to maximize average lifetime inclusive fitness. We illustrate this by presenting hypotheses that we believe incorporate the core phenomena of soft inheritance and will aid in understanding them.

Niche Construction Theory (NCT) provides a powerful conceptual framework for understanding how and why humans and target species entered into domesticatory relationships that have transformed Earth’s biota, landforms, and atmosphere, and shaped the trajectory of human cultural development. NCT provides fresh perspective on how niche-constructing behaviors of humans and plants and animals promote co-evolutionary interactions that alter selection pressures and foster genetic responses in domesticates. It illuminates the role of niche-altering activities in bequeathing an ecological inheritance that perpetuates the co-evolutionary relationships leading to domestication, especially as it pertains to traditional ecological knowledge and the transmission of learned behaviors aimed at enhancing returns from local environments. NCT also provides insights into the contexts and mechanisms that promote cooperative interactions in both humans and target species needed to sustain niche-constructing activities, ensuring that these activities produce an ecological inheritance in which domesticates play an increasing role. A NCT perspective contributes to on-going debates in the social sciences over explanatory frameworks for domestication, in particular as they pertain to issues of reciprocal causation, co-evolution, and the role of human intentionality. Reciprocally, domestication provides a model system for evaluating on-going debates in evolutionary biology concerning the impact of niche construction, phenotypic plasticity, extra-genetic inheritance, and developmental bias in shaping the direction and tempo of evolutionary change.

Biologists and philosophers of science have recently called for an extension of evolutionary theory. This so-called ‘extended evolutionary synthesis’ (EES) seeks to integrate developmental processes, extra-genetic forms of inheritance, and niche construction into evolutionary theory in a central way. While there is often agreement in evolutionary biology over the existence of these phenomena, their explanatory relevance is questioned. Advocates of EES posit that their perspective offers better explanations than those provided by ‘standard evolutionary theory’ (SET). Still, why this would be the case is unclear. Usually, such claims assume that EES’s superior explanatory status arises from the pluralist structure of EES, its different problem agenda, and a growing body of evidence for the evolutionary relevance of developmental phenomena (including developmental bias, inclusive inheritance, and niche construction). However, what is usually neglected in this debate is a discussion of what the explanatory standards of EES actually are, and how they differ from prevailing standards in SET. In other words, what is considered to be a good explanation in EES versus SET? To answer this question, we present a theoretical framework that evaluates the explanatory power of different evolutionary explanations of the same phenomena. This account is able to identify criteria for why and when evolutionary explanations of EES are better than those of SET. Such evaluations will enable evolutionary biology to find potential grounds for theoretical integration.

A niche is the structural, temporal, and social context in which a species exists. Over the last two million years, the human lineage underwent clear morphological changes alongside less easily measurable, but significant, behavioral and cognitive shifts as it forged, and was shaped by, new niches. During this time period, core human patterns emerged, including the following: hypercooperation; lengthy childhood and complex parenting; intricate and diverse foraging and hunting patterns; novel and dynamic material and symbolic cultures; and complex communication and information sharing, eventually resulting in language. Approaches to human evolution grounded in paleoanthropology and archaeology offer fundamental insights into our past, and traditional evolutionary theory offers a strong grounding for explaining them. However, given the centrality of distinctive physiological, social, semiotic, and cognitive processes in human evolutionary histories, a broader anthropological approach can facilitate additional understanding of the human story. An integrative anthropology, reaching across subfields and foci, combined with contemporary evolutionary theory is an approach that can enhance our abilities to model and understand human evolution. Un nicho es el contexto estructural, temporal y social en el que una especie existe. Durante los últimos 2 millones de años, el linaje humano pasó por claros cambios morfológicos junto a cambios conductuales y cognitivos menos fácilmente medibles, pero significativos, en la medida, que forjó, y fue moldeado por, nuevos nichos. Durante este período de tiempo, patrones humanos centrales emergieron incluyendo los siguientes: hiper-cooperación; prolongada niñez y crianza compleja; patrones de caza y recolección intrincados y diversos; culturas simbólicas y materiales novedosas y dinámicas, y el compartir complejo de información y comunicación que finalmente resultó en lenguaje. Aproximaciones a la evolución humana basadas en paleo-antropología y arqueología ofrecen conocimiento fundamental de nuestro pasado, y la teoría evolucionaria tradicional ofrece un conocimiento básico sólido para explicarlo. Sin embargo, dada la centralidad de procesos fisiológicos, sociales, semióticos, y cognitivos distintivos en historias evolucionarias humanas, una aproximación antropológica más amplia puede facilitar un entendimiento adicional de la historia humana. Una antropología integrativa, extendiéndose a través de sub-campos y focos, combinada con teoría evolucionaria contemporánea es una aproximación que puede enriquecer nuestras habilidades para modelar y entender la evolución humana.

The development of agricultural societies, one of the most transformative events in human and ecological history, was made possible by plant and animal domestication. Plant domestication began 12,000–10,000 y ago in a number of major world areas, including the New World tropics, Southwest Asia, and China, during a period of profound global environmental perturbations as the Pleistocene epoch ended and transitioned into the Holocene. Domestication is at its heart an evolutionary process, and for many prehistorians evolutionary theory has been foundational in investigating agricultural origins. Similarly, geneticists working largely with modern crops and their living wild progenitors have documented some of the mechanisms that underwrote phenotypic transformations from wild to domesticated species. Ever-improving analytic methods for retrieval of empirical data from archaeological sites, together with advances in genetic, genomic, epigenetic, and experimental research on living crop plants and wild progenitors, suggest that three fields of study currently little applied to plant domestication processes may be necessary to understand these transformations across a range of species important in early prehistoric agriculture. These fields are phenotypic (developmental) plasticity, niche construction theory, and epigenetics with transgenerational epigenetic inheritance. All are central in a controversy about whether an Extended Evolutionary Synthesis is needed to reconceptualize how evolutionary change occurs. An exploration of their present and potential utility in domestication study shows that all three fields have considerable promise in elucidating important issues in plant domestication and in agricultural origin and dispersal research and should be increasingly applied to these issues.