I analyze here biological regression equations known in the literature as allometries and scaling laws. My focus is on the alleged lawlike status of these equations. In particular I argue against recent views that regard allometries and scaling laws as representing universal, non-continent, and/or strict biological laws. Although allometries and scaling laws appear to be generalizations applying to many taxa, they are neither universal nor exceptionless. In fact there appear to be exceptions to all of them. Nor are the constants (...) in allometries and scaling laws truly constant, stable, or universal in character, but vary in value across different taxa and background conditions. Moreover, these equations represent evolutionary, strongly contingent generalizations, which threatens their lawlike status. Lastly, allometries and scaling laws do not offer stable probabilities to which they hold in different backgrounds. I further suggest that many allometries and scaling laws function to elucidate explananda rather than explanantia or covering laws. (shrink)

The current literature that attempts to bridge between geometric morphometrics (GMM) and finite element analyses (FEA) of CT-derived data from bones of living animals and fossils appears to lack a sound biotheoretical foundation. To supply the missing rigor, the present article demonstrates a new rhetoric of quantitative inference across the GMM–FEA bridge—a rhetoric bridging form to function when both have been quantified so stringently. The suggested approach is founded on diverse standard textbook examples of the relation between forms and the (...) way strains in them are produced by stresses imposed upon them. One potentially cogent approach to the explanatory purposes driving studies of this class arises from a close scrutiny of the way in which computations in both domains, shape and strain, can be couched as minimizations of a scalar quantity. For GMM, this is ordinary Procrustes shape distance; in FEA, it is the potential energy that is stored in the deformed configuration of the solid form. A hybrid statistical method is introduced requiring that all forms be subjected to the same detailed loading designs (the same “probes”) in a manner careful to accommodate the variations of those same forms before they were stressed. The proper role of GMM is argued to be the construction of regressions for strain energy density on the largest-scale relative warps in order that biological explanations may proceed in terms of the residuals from those regressions: the local residual features of strain energy density. The method, evidently a hierarchical one, might be intuitively apprehended as a geometrical approach to a formal allometric analysis of strain. The essay closes with an exhortation. (shrink)

The authors constructed an algorithm that relates leaf contour to the ratio between the two parts of the leaf using the function θ’=nθm, wherem is the allometric exponent. Using this model, it is possible to simulate the contour of symmetrical or asymmetrical leaves. The authors hypothesize that the portion of the leaf contour that agrees with the simulation is linked to a constraint imposed by the initial asymmetry of the leaf primordium. The final shape of the leaf results more from (...) a summation of cumulative processes through time than from a single process. In the analysis of foliar allometry, it is important to take into consideration the different phases of leaf development, since the nature of the allometric constants may change from one phase to the next, even though their numerical value may remain the same. (shrink)

Sexual selection drives the evolution of some of the most exaggerated traits in nature. Studies on sexual selection often focus on the size of these traits relative to body size, but few focus on energetic maintenance costs of the tissues that compose them, and the ways in which these costs vary with body size. The relationships between energy use and body size have consequences that may allow large individuals to invest disproportionally more in sexually selected structures, or lead to the (...) reduced per‐gram maintenance cost of enlarged structures. Although sexually selected traits can incur energetic maintenance costs, these costs are not universally high; they are dependent on the relative mass and metabolic activity of tissues associated with them. Energetic costs of maintenance may play a pervasive yet little‐explored role in shaping the relative scaling of sexually selected traits across diverse taxa. Also see the video abstract here: https://youtu.be/JyuoQIeA33Q. (shrink)

Despite the ontogenetic allometric size effects that explain much of phyletic variation in brain components, the residuals of some structures indicates that mosaic brain evolution was an important factor in hominid evolution, and that reorganization of the hominid brain may have occurred as early as 3+ MY. Finlay et al.'s allometric technique masks residual variation around allometric trends, and the patterns of residuals relevant to species-specific departures from strict allometric trends.

How micro- and macroevolutionary evolutionary processes produce phenotypic change is without question one of the most intriguing and perplexing issues facing evolutionary biologists. We believe that roadblocks to progress lie A) in the underestimation of the role of the environment, and in particular, that of the interaction of genotypes with environmental factors, and B) in the continuing lack of incorporation of development into the evolutionary synthesis. We propose the integration of genetic, environmental and developmental perspectives on the evolution of the (...) phenotype in the form of the concept of the developmental reaction norm (DRN) The DRN represents the set of multivariate ontogenies that can be produced by a single genotype when it is exposed to environmental variation. It encompasses: 1) the processes that alter the phenotype throughout the ontogenetic trajectory, 2) the recognition that different aspects of the phenotype are (and must be) correlated and 3) the ability of a genotype to produce phenotypes in different environments. This perspective necessitates the explicit study of character expression during development, the evaluation of associations between pairs or groups of characters (e.g., multivariate allometries), and the exploration of reaction norms and phenotypic plasticity. We explicitly extend the concept of the DRN to encompass adjustments made in response to changes in the internal environment as well. Thus, ‘typical’ developmental sequences (e.g., cell fate determination) and plastic responses are simply manifestations of different scales of ‘environmental’ effects along a continuum. We present: (1) a brief conceptual review of three fundamental aspects of the generation and evolution of phenotypes: the changes in the trajectories describing growth and differentiation (ontogeny), the multivariate relationships among characters (allometry), and the effect of the environment (plasticity); (2) a discussion of how these components are merged in the concept of the developmental reaction norm; and (3) a reaction norm perspective of major determinants of phenotypes: epigenesis, selection and constraint. (shrink)

Among all organisms, the size of each body part or organ scales with overall body size, a phenomenon called allometry. The study of shape and form has attracted enormous interest from biologists, but the genetic, developmental and physiological mechanisms that control allometry and the proportional growth of parts have remained elusive. Recent progress in our understanding of body‐size regulation provides a new synthetic framework for thinking about the mechanisms and the evolution of allometric scaling. In particular, insulin/IGF signaling, (...) which plays major roles in longevity, diabetes and the regulation of cell, organ and body size, might also be centrally involved in regulating organismal shape. Here we review recent advances in the fields of growth regulation and endocrinology and use them to construct a developmental model of static allometry expression in insects. This model serves as the foundation for a research program that will result in a deeper understanding of the relationship between growth and form, a question that has fascinated biologists for centuries. BioEssays 29:536–548, 2007. © 2007 Wiley Periodicals, Inc. (shrink)

1 Adaptationism is a research strategy that seeks to identify adaptations and the specific selective forces that drove their evolution in past environments. Since the mid-1970s, paleontologist Stephen J. Gould and geneticist Richard Lewontin have been critical of adaptationism, especially as applied toward understanding human behavior and cognition. Perhaps the most prominent criticism they made was that adaptationist explanations were analogous to Rudyard Kipling's Just So Stories. Since storytelling is an inherent part of science, the criticism refers to the acceptance (...) of stories without sufficient empirical evidence. In particular, Gould, Lewontin, and their colleagues argue that adaptationists often use inappropriate evidentiary standards for identifying adaptations and their functions, and that they often fail to consider alternative hypotheses to adaptation. Playing prominently in both of these criticisms are the concepts of constraint, spandrel, and exaptation. In this article we discuss the standards of evidence that could be used to identify adaptations and when and how they may be appropriately used. Moreover, building an empirical case that certain features of a trait are best explained by exaptation, spandrel, or constraint requires demonstrating that the trait's features cannot be better accounted for by adaptationist hypotheses. Thus, we argue that the testing of alternatives requires the consideration, testing, and systematic rejection of adaptationist hypotheses. Where possible, we illustrate our points with examples taken from human behavior and cognition. Key Words: adaptation; ADHD; brain allometry; constraint; epistemology; evolutionary psychology; exaptation; female orgasm ; optimization; special design; waist-hip ratio. Footnotes1 The authors contributed equally to this paper. Order of authorship was determined alphabetically. Correspondence may be addressed to any of the authors. (shrink)

The physical singularity of life phenomena is analyzed by means of comparison with the driving concepts of theories of the inert. We outline conceptual analogies, transferals of methodologies and theoretical instruments between physics and biology, in addition to indicating significant differences and sometimes logical dualities. In order to make biological phenomenalities intelligible, we introduce theoretical extensions to certain physical theories. In this synthetic paper, we summarize and propose a unified conceptual framework for the main conclusions drawn from work spanning a (...) book and several articles, quoted throughout. (shrink)

How does evolution grow bigger brains? It has been widely assumed that growth of individual structures and functional systems in response to niche-specific cognitive challenges is the most plausible mechanism for brain expansion in mammals. Comparison of multiple regressions on allometric data for 131 mammalian species, however, suggests that for 9 of 11 brain structures taxonomic and body size factors are less important than covariance of these major structures with each other. Which structure grows biggest is largely predicted by a (...) conserved order of neurogenesis that can be derived from the basic axial structure of the developing brain. This conserved order of neurogenesis predicts the relative scaling not only of gross brain regions like the isocortex or mesencephalon, but also the level of detail of individual thalamic nuclei. Special selection of particular areas for specific functions does occur, but it is a minor factor compared to the large-scale covariance of the whole brain. The idea that enlarged isocortex could be a a by-product of structural constraints later adapted for various behaviors, contrasts with approaches to selection of particular brain regions for cognitively advanced uses, as is commonly assumed in the case of hominid brain evolution. (shrink)

This paper proposes an abstract mathematical frame for describing some features of biological time. The key point is that usual physical (linear) representation of time is insufficient, in our view, for the understanding key phenomena of life, such as rhythms, both physical (circadian, seasonal …) and properly biological (heart beating, respiration, metabolic …). In particular, the role of biological rhythms do not seem to have any counterpart in mathematical formalization of physical clocks, which are based on frequencies along the usual (...) (possibly thermodynamical, thus oriented) time. We then suggest a functional representation of biological time by a 2-dimensional manifold as a mathematical frame for accommodating autonomous biological rhythms. The “visual” representation of rhythms so obtained, in particular heart beatings, will provide, by a few examples, hints towards possible applications of our approach to the understanding of interspecific differences or intraspecific pathologies. The 3- dimensional embedding space, needed for purely mathematical reasons, allows to introduce a suitable extra-dimension for “representation time”, with a cognitive significance. (shrink)

The scaling of body parts is central to the evolution of morphology and shape. Most traits scale proportionally with each other and body size such that larger adults are essentially magnified versions of smaller ones. This pattern is so ubiquitous that departures from it – disproportionate scaling between trait and body size – pique interest because it can generate dramatically exaggerated traits. These extreme morphologies are frequently hypothesized to result from sexual selection and their study has a long history, with (...) several hypotheses seeking to explain their evolution. Despite this effort, surprisingly little progress has been made in demonstrating the forms of selection that produce different scaling patterns or in identifying the mechanisms that underlie the expression and evolution of scaling relationships. Here we review recent insights regarding the proximate mechanisms that regulate and integrate trait growth and that offer a new framework for studying the evolution of morphological scaling. (shrink)

Researchers studying primate brain allometry often focus on departures from allometry more than the allometric relationships themselves because only the former reveal what brain regions and behavioral-cognitive abilities were the focus of selection. Allometric departures for the human brain provide insights into hominid brain evolution and cast doubt on the suggestion that the large human cerebral cortex is a.

Phenotypic Evolution explicitly recognizes organisms as complex genetic-epigenetic systems developing in response to changing internal and external environments. As a key to a better understanding of how phenotypes evolve, the authors have developed a framework that centers on the concept of the Developmental Reaction Norm. This encompasses their views: (1) that organisms are better considered as integrated units than as disconnected parts (allometry and phenotypic integration); (2) that an understanding of ontogeny is vital for evaluating evolution of adult forms (...) (ontogenetic trajectories, epigenetics, and constraints); and (3) that environmental heterogeneity is ubiquitous and must be acknowledged for its pervasive role in phenotypic expression. (shrink)

The growing interest and major advances of the last decades in evolutionary developmental biology (EvoDevo) have led to the recognition of the incompleteness of the Modern Synthesis of evolutionary theory. Here we discuss how paleontology makes significant contributions to integrate evolution and development. First, extinct organisms often inform us about developmental processes by showing a combination of features unrecorded in living species. We illustrate this point using the vertebrate fossil record and studies relating bone ossification to life history traits. Second, (...) we discuss exceptionally preserved fossils that document rare ontogenetic sequences and illustrate this case with the patterns of heterochrony observed in Cambrian crustacean larvae preserved three-dimensionally. Third, most fossils potentially document the evolutionary patterns of allometry and modularity, as well as some of the (paleo)ecological factors that had influenced them. The temporal persistence of adaptive patterns in rodent evolution serves to address the importance of ecological constraints in evolution. Fourth, we discuss how the macroevolutionary patterns observed in the tetrapod limb, in the mammal molar proportions, and in the molluscan shell provide independent tests of the validity of morphogenetic models proposed on living species. Reciprocally, these macroevolutionary patterns often act as a source of inspiration to investigate the underlying rules of development, because, at the end, they are the patterns that the neo-Darwinian theory was unable to account for. (shrink)

The human ability to live according to learned, shared rules of behavior requires cortical functions. Is the limbic system also necessary for culture or are its functions opposed to it, requiring cortical inhibition? The sizes of monkey and ape neocortical and major limbic structures scale with brain weight, but the neocortex expands more (has a steeper exponent) than limbic structures. As the human brain evolved it did not deviate from the scaling relationships found in nonhuman anthropoids. This evidence for conservation (...) in scaling supports the idea that limbic functions are necessary for human symbolism and culture. (shrink)