The non-existence of a principle of natural selection

The theory of natural selection is a rich systematization of biological knowledge without a first principle. When formulations of a proposed principle of natural selection are examined carefully, each is seen to be exhaustively analyzable into a proposition about sources of fitness and a proposition about consequences of fitness. But whenever the fitness of an organic variety is well defined in a given biological situation, its sources are local contingencies together with the background of laws from disciplines other than the theory of natural selection; and the consequences of fitness for the long range fate of organic varieties are essentially applications of probability theory. Hence there is no role and no need for a principle of the theory of natural selection, and any generalities that may hold in that theory are derivative rather than fundamental.     

An assessment of Gallistel's (2012) rationalistic account of extinction phenomena

Gallistel (2012) asserts that animals use rationalistic reasoning (i.e., information theory and Bayesian inference) to make decisions that underlie select extinction phenomena. Rational processes are presumed to lead to evolutionarily optimal behavior. Thus, Gallistel's model is a type of optimality theory. But optimality theory is only a theory, a theory about an ideal organism, and its predictions frequently deviate appreciably from observed behavior of animals in the laboratory and the real world. That is, behavior of animals is often far from optimal, as is evident in many behavioral phenomena. Hence, appeals to optimality theory to explain, rather than illuminate, actual behavior are misguided.     

Are individuals really subordinated to genes? A theory of living entities

Modern evolutionary theory consists of an explicit theory of change and an implicit theory of the existence of living entities. A theory of the latter type, called the theory of the process of living (POL), is proposed. It contains a two-level definition of living entities, an analysis of possible and impossible similarity and differences between parents and offspring, and a logical division of the information governing the functioning of a living organism. The theory of POL emphasizes the proceeding of living entities in time while the modern evolutionary theory emphasizes the types of entities proceeding in time. The theory of POL produces a basis for reanalysing relations between individuals, their genes and the environment. It produces some predictions deviating from classic evolutionary theory. These conflicts occur in areas where analyses of genetic dynamics have made implicit assumptions of rules of POL. For example, the theory of POL states that the genetic relatedness of successive generations is a technical solution more than the reason of reproduction, that genetic relatedness as such does not imply altruism between individuals and that genes are the tools whereby organisms exist and not the contrary. Implications of the theory of POL on paradoxes of evolutionary theory are discussed.     

Salt-water coupling in leaky epithelia

Summary The theory of quasi-isotonic transport by cellular osmosis (the standing-gradient theory) has been challenged on the grounds that the osmotic permeabilities of the mucosal and interspace membranes are too low; if they were as high as the theory requires then the osmotic permeability of the whole epithelium would be 2–3 orders of magnitude higher than observed. This objection has basically been accepted for it is now claimed that these enormous permeabilities do exist, but are masked by unstirred-layer effects; I show that this is incorrect because unstirredlayer corrections are small and that the situation has not changed since 1975.The view that the route of fluid transport is junctional is replacing the cellular theory, and trans-junctional water flows seem to account for major fractions of the flow in various epithelia. I argue on grounds of general theory that these are unlikely to be osmotic flows because the junctional pores cannot satisfy both the osmotic and diffusive properties required of them, but the basic osmotic theory is also rather vague here.Non-osmotic theories, if junctional flow is accepted, must be either electro-kinetic or peristaltic.     

The validity and value of inclusive fitness theory

Social evolution is a central topic in evolutionary biology, with the evolution of eusociality (societies with altruistic, non-reproductive helpers) representing a long-standing evolutionary conundrum. Recent critiques have questioned the validity of the leading theory for explaining social evolution and eusociality, namely inclusive fitness (kin selection) theory. I review recent and past literature to argue that these critiques do not succeed. Inclusive fitness theory has added fundamental insights to natural selection theory. These are the realization that selection on a gene for social behaviour depends on its effects on co-bearers, the explanation of social behaviours as unalike as altruism and selfishness using the same underlying parameters, and the explanation of within-group conflict in terms of non-coinciding inclusive fitness optima. A proposed alternative theory for eusocial evolution assumes mistakenly that workers' interests are subordinate to the queen's, contains no new elements and fails to make novel predictions. The haplodiploidy hypothesis has yet to be rigorously tested and positive relatedness within diploid eusocial societies supports inclusive fitness theory. The theory has made unique, falsifiable predictions that have been confirmed, and its evidence base is extensive and robust. Hence, inclusive fitness theory deserves to keep its position as the leading theory for social evolution.     

The subtleties of error management

Error management theory is a theory of considerable scope and emerging influence. The theory claims that cognitive biases do not necessarily reflect flaws in evolutionary design, but that they may be best conceived as design features. Unfortunately, existing accounts are vague with respect to the key concept of bias. The result is that it is unclear that the cognitive biases that the theory seeks to defend are not simply a form of behavioral bias, in which case the theory reduces to a version of expected utility theory. We propose some clarifications and refinements of error management theory by emphasizing important distinctions between different forms of behavioral and cognitive bias. We also highlight a key assumption, that the capacity for Bayesian beliefs is subject to constraints. This assumption is necessary for what we see as error management theory's genuinely novel claim: that behavioral tendencies to avoid costly errors can rest on systematic departures from Bayesian beliefs and that the latter can be adaptive insofar as they generate the former.     

DNA and the neutral theory

The neutral theory claims that the great majority of evolutionary changes at the molecular (DNA) level are caused not by Darwinian selection but by random fixation of selectively neutral or nearly neutral mutants. The theory also asserts that the majority of protein and DNA polymorphisms are selectively neutral and that they are maintained in the species by mutational input balanced by random extinction. In conjunction with diffusion models (the stochastic theory) of gene frequencies in finite populations, it treats these phenomena in quantitative terms based on actual observations. Although the theory has been strongly criticized by the 'selectionists', supporting evidence has accumulated over the years. Particularly, the recent outburst of DNA sequence data lends strong support to the theory both with respect to evolutionary base substitutions and DNA polymorphism, including rapid evolutionary base substitutions in pseudogenes. In addition, the observed pattern of synonymous codon choice can now be readily explained in the framework of this theory. I review these recent findings in the light of the neutral theory.     

Analytical comparison between Nixon–Logvinenko’s and Jung–Brown’s theories of slow neurofilament transport in axons

This paper develops analytical solutions describing slow neurofilament (NF) transport in axons. The obtained solutions are based on two theories of NF transport: Nixon–Logvinenko’s theory that postulates that most NFs are incorporated into a stationary cross-linked network and only a small pool is slowly transported and Jung–Brown’s theory that postulates a single dynamic pool of NFs that are transported according to the stop-and-go hypothesis. The simplest two-kinetic state version of the model developed by Jung and Brown was compared with the theory developed by Nixon and Logvinenko. The model for Nixon–Logvinenko’s theory included stationary, pausing, and running NF populations while the model used for Jung–Brown’s theory only included pausing and running NF populations. Distributions of NF concentrations resulting from Nixon–Logvinenko’s and Jung-Brown’s theories were compared. In previous publications, Brown and colleagues successfully incorporated slowing of NF transport into their model by assuming that some kinetic constants depend on the distance from the axon hillock. In this paper we defined the average rate of NF transport as the rate of motion of the center of mass of radiolabeled NFs. We have shown that for this definition, if all kinetic rates are assumed constant, Jung–Brown’s theory predicts a constant average rate of NF transport. We also demonstrated that Nixon–Logvinenko’s theory predicts slowing of NF transport even if all kinetic rates are assumed constant, and the obtained slowing agrees well with published experimental data.     

Recent evidence for the exon theory of genes

I provide a history of the Exon Theory of Genes, a theory that holds that introns are extremely ancient characteristics of genes and that early genes were created through the intron-mediated shuffling of exons. The theory has existed since the late seventies. An uncompromising version of the theory, in which all introns were considered to be ancient, dominated the early work. However, in the late eighties and early nineties evidence arose that at least some introns are more recently acquired. Due to the previously overly polarized nature of the debate, many results that demonstrated the existence of recent introns have been interpreted by some as evidence that all introns are late, leading some to believe that the theory is no longer a viable theory. However, recent work emphasizes a mixed model, in which some introns are ancient and some new. This model has proven to provide impressive explanatory power for a broad array of observations. Thus, this moderated form of the Exon Theory of Genes still offers a coherent picture of the origin and evolutionary history of the intron–exon structure of eukaryotic genes.     

The neutral theory of molecular evolution and the world view of the neutralists

The main tenet of the neutral theory is that the great majority of evolutionary changes at the molecular level are caused not by Darwinian selection but by random fixation of selectively neutral (or very nearly neutral) alleles through random sampling drift under continued mutation pressure. The theory also asserts that the majority of protein and DNA polymorphisms are selectively neutral, and that they are maintained in the species by mutational input balanced by random extinction rather than by "balancing selection." The neutral theory is based on simple assumptions. This enabled us to develop mathematical theories (using the diffusion equation method) that can treat these phenomena in quantitative terms and that permit theory to be tested against actual observations. Although the neutral theory has been severely criticized by the neo-Darwinian establishment, supporting evidence has accumulated over the last 20 years. In particular, the recent burst of DNA sequence data helped to strengthen the theory a great deal. I believe that the neutral theory triggered reexamination of the traditional "synthetic theory of evolution." In this paper, I review the present status of the neutral theory, including discussions of such topics as "molecular evolutionary clock," very high evolutionary rates observed in RNA viruses, a deviant coding system found in Mycoplasm together with the concept of mutation-driven neutral evolution, and the origin of life. I also present a worldview based on the conception of what I call "survival of the luckiest."     

Physiological regulation-deregulation and psychiatric disorders

This article discusses regulation-deregulation theory, a theory that integrates both old and new concepts to explain behavior-physiology interactions and changes in psychiatric disorder probability. The theory postulates: that specific types and frequencies of social interactions are essential to maintain normal physiological function; when essential interactions fail to occur, atypical physiological states (deregulation) develop; and deregulation is associated with the presence of unpleasant feelings and an increase in the frequency of psychiatric disorders. Implications of the theory are that certain types of social interactions are critical proximate mechanisms in determining the onset of psychiatric disorders, and that social interaction type and frequency, through affecting different physiological systems, alter the course of psychiatric disorders.     

Mixed function oxidases and herbivore polyphagy: the devil's advocate position

Abstract. 1. Evidence used to support the theory that mixed function oxidases (MFOs) are important in the survival of polyphagous herbivores is briefly reviewed. This evidence includes data on patterns of variation in MFO activity among species and among developmental stages within species. It also includes data on induction of MFO activity by plant compounds and metabolism of plant cornpounds by MFOs.
2. It is argued that the evidence presently available does not offer strong support of the theory because key pieces of information are lacking. Evidence which tends to refute the theory is reviewed and discussed.
3. Experiments are proposed which could more rigorously test the theory.     

Dynamic decision-making in uncertain environments II. Allais paradox in human behavior

In both animal and human behavioral repertoires, classical expected utility theory is considered a fundamental element of decision making under conditions of uncertainty. This theory has been widely applied to problems of animal behavior and evolutionary game theory, as well as to human economic behavior. The Allais paradox hinges on the expression of avoidance of bankruptcy by humans, or death by starvation in animals. This paradox reveals that human behavioral patterns are often inconsistent with predictions under the classical expected utility theory as formulated by von Neumann and Morgenstern. None of the many attempts to reformulate utility theory has been entirely successful in resolving this paradox with rigorous logic. We present a simple, but novel approach to the theory of decision making, in which utility is dependent on current wealth, and in which losses are more heavily weighted than gains. Our approach resolves the Allais paradox in a manner that is consistent with how humans formulate decisions under uncertainty. Our results indicate that animals, including humans, are in principle risk-averse. Our restructuring of dynamic utility theory presents a basic optimization scheme for sequential or dynamic decisions in both animals and humans.     

Two neo-Darwinisms

There are two extant theories of evolution, each of which deserves the honourific "neo-Darwinism": Modern Synthesis Replicator theory and a theory I shall call Developmental Darwinism. The principal difference concerns the canonical unit of biological organization. Modern Synthesis replicator theory explains the process of evolution by appeal to the activities of genes or replicators. Developmental Darwinism explains the process of evolution by appeal to the capacities of organisms. In particular, it is the plasticity of organisms, manifested most distinctly during development, that causes adaptive evolution. Despite the fact that each, in its own way, traces its origin to the theory outlined by Darwin, they are radically different. The objectives of this essay are twofold: to underscore the differences between these theories, and to argue that Developmental Darwinism, though nascent, is a viable alternative to Modern Synthesis replicator theory.     

The derivation of D’Arcy Thompson’s theory of transformations from the theory of optimal design

It is shown informally that Cohn’s theory of Optimal Forms can be construed as a comparative theory, and that when this is done the celebrated theory of transformations of D’Arcy Thompson follows as a consequence. The implications of this type of theoretical foundation for the Thompson theory with regard to problems of comparative morphology are discussed, and some suggestions for the further implementation of the theory are described. This research was supported by the United States Air Force through the Air Force Office of Scientific Research of the Air Research and Development Command, under Contract No. AF 49(638)-917.     

Motivation and placebos: do different mechanisms occur in different contexts?

This paper challenges the common assumption that the mechanisms underlying short-term placebo paradigms (where there is no motivation for health improvement) and long-term placebo paradigms (where patients value improvement in their health) are the same. Three types of motivational theory are reviewed: (i) classical placebo motivation theory that the placebo response results from the desire for therapeutic improvement; (ii) goal activation model that expectancy-driven placebo responses are enhanced when the placebo response satisfies an activated goal; and (iii) motivational concordance model that the placebo response is the consequence of concordance between the placebo ritual and significant intrinsic motives. It is suggested that current data are consistent with the following theory: response expectancy, conditioning and goal activation are responsible for short-term placebo effects but long-term therapeutic change is achieved through the effects of goal satisfaction and affect on the inflammatory response system and hypothalamic-pituitary-adrenal axis. Empirical predictions of this new theory are outlined, including ways in which placebo effects can be combined with other psychologically mediated effects on short-term and long-term psychological and physiological state.     

Towards an integration of ecological stoichiometry and the metabolic theory of ecology to better understand nutrient cycling

Ecologists have long recognized that species are sustained by the flux, storage and turnover of two biological currencies: energy, which fuels biological metabolism and materials (i.e. chemical elements), which are used to construct biomass. Ecological theories often describe the dynamics of populations, communities and ecosystems in terms of either energy (e.g. population-dynamics theory) or materials (e.g. resource-competition theory). These two classes of theory have been formulated using different assumptions, and yield distinct, but often complementary predictions for the same or similar phenomena. For example, the energy-based equation of von Bertalanffy and the nutrient-based equation of Droop both describe growth. Yet, there is relatively little theoretical understanding of how these two distinct classes of theory, and the currencies they use, are interrelated. Here, we begin to address this issue by integrating models and concepts from two rapidly developing theories, the metabolic theory of ecology and ecological stoichiometry theory. We show how combining these theories, using recently published theory and data along with new theoretical formulations, leads to novel predictions on the flux, storage and turnover of energy and materials that apply to animals, plants and unicells. The theory and results presented here highlight the potential for developing a more general ecological theory that explicitly relates the energetics and stoichiometry of individuals, communities and ecosystems to subcellular structures and processes. We conclude by discussing the basic and applied implications of such a theory, and the prospects and challenges for further development.     

Play-like behaviour: An essay in speculative ethology

It is claimed that certain processes of individual behaviour and of interaction between individuals run parallel. Such parallels are seen along three axes: antagonism-coordination, constriction-expansion and neutral-play-like. Characteristics of ritualized behaviour and play are analysed and the two categories of behaviour are compared in detail. They are shown to differ largely in degree of expansion. They also differ along the antagonism-coordination axis. Both are play-like. A theory comprising both ritualized behaviour and play is attempted. The concept of field interaction is introduced. The physical theory of elect ro-magnetism is used as a conceptual model. Positive and problematic aspects of the theory are discussed: Is the behavioural field polarised or not? The relationship between aggression and play-like behaviour is considered, both as a motivational problem and in phylogenetic perspective. Some limits to the theory are indicated. Relationships of the theory to some other theories are briefly discussed.