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      Variation, mosaicism and degeneracy in the hominin foot

      review-article
      1 , 2 , 1 , * ,
      Evolutionary Human Sciences
      Cambridge University Press
      Foot, degeneracy, variability, biomechanics, locomotion, plasticity

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          Abstract

          Abstract

          The fossil record is scarce and incomplete by nature. Animals and ecological processes devour soft tissue and important bony details over time and, when the dust settles, we are faced with a patchy record full of variation. Fossil taxa are usually defined by craniodental characteristics, so unless postcranial bones are found associated with a skull, assignment to taxon is unstable. Naming a locomotor category based on fossil bone morphology by analogy to living hominoids is not uncommon, and when no single locomotor label fits, postcrania are often described as exhibiting a ‘mosaic’ of traits. Here, we contend that the unavoidable variation that characterises the fossil record can be described far more rigorously based on extensive work in human neurobiology and neuroanatomy, movement sciences and motor control and biomechanics research. In neurobiology, degeneracy is a natural mechanism of adaptation allowing system elements that are structurally different to perform the same function. This concept differs from redundancy as understood in engineering, where the same function is performed by identical elements. Assuming degeneracy, structurally different elements are able to produce different outputs in a range of environmental contexts, favouring ecological robusticity by enabling adaptations. Furthermore, as degeneracy extends to genome level, genetic variation is sustained, so that genes which might benefit an organism in a different environment remain part of the genome, favouring species’ evolvability.

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          Noise in the nervous system.

          Noise--random disturbances of signals--poses a fundamental problem for information processing and affects all aspects of nervous-system function. However, the nature, amount and impact of noise in the nervous system have only recently been addressed in a quantitative manner. Experimental and computational methods have shown that multiple noise sources contribute to cellular and behavioural trial-to-trial variability. We review the sources of noise in the nervous system, from the molecular to the behavioural level, and show how noise contributes to trial-to-trial variability. We highlight how noise affects neuronal networks and the principles the nervous system applies to counter detrimental effects of noise, and briefly discuss noise's potential benefits.
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            Adaptation to an extraordinary environment by evolution of phenotypic plasticity and genetic assimilation.

            Adaptation to a sudden extreme change in environment, beyond the usual range of background environmental fluctuations, is analysed using a quantitative genetic model of phenotypic plasticity. Generations are discrete, with time lag tau between a critical period for environmental influence on individual development and natural selection on adult phenotypes. The optimum phenotype, and genotypic norms of reaction, are linear functions of the environment. Reaction norm elevation and slope (plasticity) vary among genotypes. Initially, in the average background environment, the character is canalized with minimum genetic and phenotypic variance, and no correlation between reaction norm elevation and slope. The optimal plasticity is proportional to the predictability of environmental fluctuations over time lag tau. During the first generation in the new environment the mean fitness suddenly drops and the mean phenotype jumps towards the new optimum phenotype by plasticity. Subsequent adaptation occurs in two phases. Rapid evolution of increased plasticity allows the mean phenotype to closely approach the new optimum. The new phenotype then undergoes slow genetic assimilation, with reduction in plasticity compensated by genetic evolution of reaction norm elevation in the original environment.
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              The extended evolutionary synthesis: its structure, assumptions and predictions.

              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.
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                Author and article information

                Journal
                Evol Hum Sci
                Evol Hum Sci
                EHS
                Evolutionary Human Sciences
                Cambridge University Press (Cambridge, UK )
                2513-843X
                2022
                27 December 2021
                : 4
                : e2
                Affiliations
                [1 ]University of Liverpool , Liverpool, UK
                [2 ]Sheffield Hallam University , Sheffield, UK
                Author notes
                [* ]Corresponding author. E-mail: robinhuwcrompton@ 123456gmail.com
                Author information
                https://orcid.org/0000-0002-7441-1748
                Article
                S2513843X21000505
                10.1017/ehs.2021.50
                10426032
                37588898
                296ec1a9-690c-4d9f-b186-73c3133e71b8
                © The Author(s) 2022

                This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence ( http://creativecommons.org/licenses/by/4.0/), which permits unrestricted re-use, distribution and reproduction, provided the original article is properly cited.

                History
                Page count
                Figures: 1, References: 178, Pages: 19
                Funding
                Funded by: The Leverhulme Trust
                Award ID: EM-2017-010\2
                Categories
                Review

                foot,degeneracy,variability,biomechanics,locomotion,plasticity
                foot, degeneracy, variability, biomechanics, locomotion, plasticity

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