Natural selection without survival of the fittest

Susan Mills and John Beatty proposed a propensity interpretation of fitness (1979) to show that Darwinian explanations are not circular, but they did not address the critics' chief complaint that the principle of the survival of the fittest is either tautological or untestable. I show that the propensity interpretation cannot rescue the principle from the critics' charges. The critics, however, incorrectly assume that there is nothing more to Darwin's theory than the survival of the fittest. While Darwinians all scoff at this assumption, they do not agree about what role, if any, this principle plays in Darwin's theory of natural selection. I argue that the principle has no place in Darwin's theory. His theory does include the idea that some organisms are fitter than others. But greater reproductive success is simply inferred from higher fitness. There is no reason to embody this inference in the form of a special principle of the survival of the fittest.I would like to thank John Beatty, Ron Giere, Philip Kitcher and John Winnie for detailed and helpful criticisms of an earlier draft of this paper.     

Eskimo hunting groups,social kinship,and the possibility of kin selection in humans

Contemporary Eskimo whaling crews show very high coefficients of kinship, with the consequent possibility of kin selection, even though the spoken rules of crew membership are in terms of social kinship.     

Natural selection, fitness entropy, and the dynamics of coevolution

The coevolutionary dynamics of interacting populations were studied by combining continuous time Lotka-Volterra models of population growth with single-locus genetic models of weak selection. The effects of natural selection on population growth were evaluated using Ginzburg's fitness entropy function as a measure of the deviation of a population's initial allele frequencies from their polymorphic equilibrium values. This entropy measure was used to relate the dynamics of a community composed of evolving populations to the dynamics of a "reference community" whose populations are initially in genetic equilibrium. Specifically, a quantity called the "selective difference area" was defined as the total difference between the population size trajectories of a reference and evolving population. The selective difference area represents the amount of extra life a species would realize if the entire community were at genetic equilibrium. It was shown that this selective difference area is a simple linear function of the initial fitness entropies of each species. This prediction is independent of the strength of selection and holds for any arbitrary set of initial population densities. Numerical examples were presented to illustrate the results. Under the assumption of weak selection, a generalization for arbitrary population growth models was outlined.     

Sexual selection,redundancy and survival of the most beautiful

A model is described of a highly redundant complex organism that has overlapping banks of genes such that each vital function is specified by several different genetic systems. This generates a synergistic profile linking probability of survival to the number of deleterious mutations in the genome. Computer models show that there is a dynamic interaction between the mean number of new deleterious mutations per generation (X), the mean number of deleterious mutations in the genome of the population (Y) and percentage zygote survival (Zs). IncreasedX leads to increasedY and a fall in Zs but it takes several generations before a new equilibrium is reached. If sexual attraction is influenced by the number of deleterious mutations in the genome of individuals thenY is reduced and Zs increased for any given value ofX. This fall inY and rise in Zs is more marked in polygamous than monogamous mating systems. The model is specified such that deleterious mutations can occur without any observable or measurable effect on function. Thus sexual selection, in this organism, for low levels of deleterious mutations cannot be based on assessment of performance. Instead it is based on a simple symmetrical surface pattern that is flawlessly reproduced by organisms with no deleterious mutations, but is less than perfect, and therefore less attractive, if genetic systems have been deleted. A complex vital task requires a system with a high level of redundancy that acts so that the loss of one component has no observable effect and therefore cannot be used for sexual selection. The reproduction of a beautiful surface pattern also requires a low error, high redundancy genetic system; however, in this case there is advantage if a single deleterious mutation produces a recognisable change. This leads to the conclusion that sexual selection and sexual attraction should be based on beauty rather than utility, and explains the common observation in nature that it is the most beautiful that survive.     

Natural selection at the MJD locus: phenotypic diversity, survival and fertility among Machado-Joseph Disease patients from the Azores.

Machado-Joseph Disease (MJD) is an autosomal dominant neurodegenerative disorder of adult onset, associated with the expansion of a (CAG)n tract in the coding region of the causative gene, localized on 14q32.1. Machado-Joseph Disease shows non-Mendelian features typical of other triplet repeat disorders, including clinical heterogeneity, variable age at onset and anticipation. Three phenotypes have been proposed (clinical types 1, 2 and 3). Type 1 is associated with early age at onset and a high repeat number of the CAG sequence, and Types 2 and 3 have later onset and lower numbers of CAG repeats. This paper investigates whether there is selection against the MJD gene, acting through differential survival. nuptiality and fertility associated with clinical type and age at onset. The study sample comprised 40 MJD patients from the Azores (Portugal) having fully documented reproductive histories and known dates of death. The proportion of married patients of each clinical type increased from 0.22 among Type 1 patients, to 0.40 in Type 2 and 0.95 in Type 3. Age at onset and length of survival were also associated with marital status, with the married cases having later mean age at onset and longer mean survival time. In the whole sample, clinical type was associated with fertility, with significantly fewer children born to Type 1 patients. Among married patients clinical type was not associated with age at marriage, reproductive span or number of children. No reduction of fertility was detected among married patients in whom the onset of MJD was below the age of 50. The authors' interpretation of these results is that the high-repeat CAG haplotypes associated with early age at onset and clinical Type 1 are selected against through reduced survival and fertility. The fertility component of selection is mediated by nuptiality rather than marital fertility.     

Natural selection and the Michaelis constant

Natural selection operating on a biological process with Michaelis-Menten kinetics adjusts the Michaelis (half-saturation) constant relative to ambient substrate concentration. From the perspective developed here, the relative fitnesses of alternative evolutionary “strategies” determine the trajectory of the Michaelis constant (K_m) over evolutionary time. If substrate concentration is held relatively constant or fluctuates randomly about mean concentration over evolutionary time, while processing rate increases, K_m tends toward a value greater than or equal to substrate concentration; if processing rate is held relatively constant over evolutionary time, K_m tends to become large relative to substrate concentration. The diversity of the supportive evidence cited suggests a broad applicability of this argument across taxonomic groups and levels of biological organization.     

Natural selection and history

In “Spandrels,” Gould and Lewontin criticized what they took to be an all-too-common conviction, namely, that adaptation to current environments determines organic form. They stressed instead the importance of history. In this paper, we elaborate upon their concerns by appealing to other writings in which those issues are treated in greater detail. Gould and Lewontin’s combined emphasis on history was three-fold. First, evolution by natural selection does not start from scratch, but always refashions preexisting forms. Second, preexisting forms are refashioned by the selection of whatever mutational variations happen to arise: the historical order of mutations needs to be taken into account. Third, the order of environments and selection pressures also needs to be taken into account.     

Natural selection and self-organization

The Darwinian concept of natural selection was conceived within a set of Newtonian background assumptions about systems dynamics. Mendelian genetics at first did not sit well with the gradualist assumptions of the Darwinian theory. Eventually, however, Mendelism and Darwinism were fused by reformulating natural selection in statistical terms. This reflected a shift to a more probabilistic set of background assumptions based upon Boltzmannian systems dynamics. Recent developments in molecular genetics and paleontology have put pressure on Darwinism once again. Current work on self-organizing systems may provide a stimulus not only for increased problem solving within the Darwinian tradition, especially with respect to origins of life, developmental genetics, phylogenetic pattern, and energy-flow ecology, but for deeper understanding of the very phenomenon of natural selection itself. Since self-organizational phenomena depend deeply on stochastic processes, self-organizational systems dynamics advance the probability revolution. In our view, natural selection is an emergent phenomenon of physical and chemical selection. These developments suggest that natural selection may be grounded in physical law more deeply than is allowed by advocates of the autonomy of biology, while still making it possible to deny, with autonomists, that evolutionary explanations can be modeled in terms of a deductive relationship between laws and cases. We explore the relationship between, chance, self-organization, and selection as sources of order in biological systems in order to make these points.     

Natural selection and neoteny

Summary Even today, a century after the publication of the Origin of Species, current zoological literature often reveals an insufficient grasp of the implications of the now generally accepted view that it is natural selection that confers direction on the evolutionary process.This is, in part, due to a reaction against oversimplified teleology and against Lamarckism. In rejecting Lamarck's thesis that the activities of an animal directly affect its hereditary characters it is frequently assumed that this implies that such activities are irrelevant to the study of evolution. This is a non-sequitur, for activities may affect evolution not directly, through heredity, but indirectly, by influencing the direction of the selective forces impinging on the organism. Reasons are given for concluding that changes in habits and behaviour frequently precede structural change and, in fact, determine the direction of the latter.As an example of a concept which deserves a more evolutionary treatment than it commonly receives, the subject of neoteny is considered. It appears that failure to think in terms of the selective forces involved has frequently led to error. An analysis in functional terms is made of the theory that the chordates represent secondarily glorified sea-squirts and it is concluded that this idea is unlikely to be correct. The characteristics of man that are commonly considered to be neotenous are also discussed. It is concluded that a facile and unanalytical application of the label neoteny to many of the evolutionary changes involved has tended to obscure rather than to clarify their significance: the neotenous features of man are not those that have been of primary importance in human evolution, but are secondary to the acquisition of new characters which are not to be found in either the adult or the young stages of any other primate.

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Zusammenfassung Sogar heutzutage, ein Jahrhundert nach dem Erscheinen der Origin of Species, zeigt die jetzige zoologische Literatur oft eine unzulängliche Einsicht in die Folgerungen, zu denen die jetzt allgemein akzeptierte Annahme leitet, dass die natürliche Selektion die Richtung des Evolutionsprozesses bedingt.Dies ist teilweise zu sehen als eine Reaktion gegen eine einseitige Teleologie und gegen das Lamarckismus. Bei der Ablehnung der TheseLamarck's, dass die Aktivitäten eines Tieres einen direkten Einfluss ausüben auf seine erbliche Konstitution, nimmt man oft an, dass dies bedeutet, dass derartige Aktivitäten demnach nichts zur Sache tun bei dem Studium der Evolution. Zu einer derartigen Konklusion ist man jedoch nicht berechtigt, denn, obwohl die Aktivitäten die Evolution nicht direkt beeinflussen können mittels der Erblichkeit, so können sie dies jedoch indirekt, weil sie die Richtung der selektiven Kräfte, welche auf das Organismus einwirken, bestimmen. Es werden Argumente gegeben, welche die Konklusion rechtfertigten, dass Änderungen in Gewohnheit und Verhalten oft Strukturänderungen vorhergehen und tatsächlich die Richtung dieser letztgenannten bestimmen. Die Neotenie wirdt als ein Beispiel eines Begriff betrachtet, welcher eine mehr evolutionäre Behandlung verdient als bisher geschehen ist. Es ergibt sich dass man viele Irrtümer hat gemacht, indem man den betreffenden selektiven Kräften keine Rechnung getragen hat. Eine Analyse der Theorie, welche die Chordaten als neotenische Ascidien betrachtet, wird von einem funktionellen Gesichtspunkt aus durchgeführt. Daraus scheint der Schluss berechtigt, dass diese Theorie falsch ist. Auch werden die menschlichen Kennzeichen, welche gewöhnlich als neotenische betrachtet werden, besprochen. Die Konklusion lautet, dass, wenn man leichtfertig und ohne genügende Analyse den Zettel Neotenie für mancherlei evolutive Änderungen anwendet, man die richtige Bedeutung dieser Erscheinung vielmehr verdunkelt als aufklärt: nicht die neotenische Kennzeichen des Menschen sind am wichtigsten in seiner Evolution; in Gegenteil sie erscheinen nur sekundär, nachdem die neuen Kennzeichen, welche sich weder in den Erwachsenen noch in den jüngeren Stadien von den anderen Primaten finden lassen, erschienen sind.

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Résumé Aujourd'hui, un siècle après l'apparition de l'Originedes Espèces, la conviction est presque générale que la sélection naturelle dirige le procès de l'évolution. Néanmoins, les publications zoologiques actuelles révèlent souvent une compréhension insuffisante de ce qui en résulte.Ceci doit en partie être considéré comme réaction contre une téléologie simpliste et contre le Lamarckisme. En rejetant la thèse de Lamarck, que les activités d'un animal ont une influence directe sur ses caractéristiques héréditaires, on admet fréquemment que de pareilles activités n'ont rien à voir avec l'étude le l'évolution. Cependant, une telle conclusion n'est pas justifiée, car les activités peuvent influer sur l'évolution non pas directement, par la voie de l'hérédité, mais indirectement en agissant sur la direction des forces sélectives que l'organisme rencontre. Des raisons ont été données qui mènent à conclure que des changements dans les habitudes et dans le comportement précèdent maintes fois le changement structural et même déterminent la direction de celui-ci.Comme exemple d'un concept qui mérite un traitement plus évolutionniste qu'il ne reçoive en général, la néoténie a été examinée. Il paraît que l'on a fait beaucoup d'erreurs en ne tenant pas compte des forces sélectives engagées. Une analyse au pointde-vue de la fonction a été appliquée à la théorie que les chordées ne sont que des ascidies néoténiques transfigurées par après, et il s'ensuit que cette idée n'est probablement pas correcte. Aussi les caractéristiques de l'homme généralement regardées comme neoténiques ont été discutées. Il en résulte que, en désignant légèrement et sans analyse plusieurs changements évolutionnaires comme néoténie, on a plutôt obsurci qu'éclairci leur signification. Ce ne sont pas les traits néoteniques de l'homme qui ont joué le rôle principal dans l'évolution humaine; au contraire, ils n'apparaissent qu'après l'acquisition de quelques nouvelles caractéristiques qui ne se trouvent ni dans l'adulte ni dans les phases plus jeunes des autres primates.

     

Natural selection and the evolution of neutralism

The efficiency of evolutionary search increases as the density of acceptable proteins in a protein space increases. Populations caught in regions whose density is too low to support evolution can be pulled into high density regions by hitchhiking selection. As they move into such regions, the action of natural selection becomes more effective, yet these populations will satisfy conditions which lead to predictions made by neutral, so-called non-Darwinian models.     

Natural selection and the elusiveness of happiness

The quest for happiness has expanded from a focus on relieving suffering to also considering how to promote happiness. However, both approaches have yet to be conducted in an evolutionary framework based on the situations that shaped the capacities for happiness and sadness. Because of this, the emphasis has almost all been on the disadvantages of negative states and the benefits of positive states, to the nearly total neglect of 'diagonal psychology', which also considers the dangers of unwarranted positive states and the benefits of negative emotions in certain situations. The situations that arise in goal pursuit contain adaptive challenges that have shaped domain-general positive and negative emotions that were partially differentiated by natural selection to cope with the more specific situations that arise in the pursuit of different kinds of goals. In cultures where large social groups give rise to specialized and competitive social roles, depression may be common because regulation systems are pushed far beyond the bounds for which they were designed. Research on the evolutionary origins of the capacities for positive and negative emotions is urgently needed to provide a foundation for sensible decisions about the use of new mood-manipulating technologies.     

Natural selection and the maximization of fitness

The notion that natural selection is a process of fitness maximization gets a bad press in population genetics, yet in other areas of biology the view that organisms behave as if attempting to maximize their fitness remains widespread. Here I critically appraise the prospects for reconciliation. I first distinguish four varieties of fitness maximization. I then examine two recent developments that may appear to vindicate at least one of these varieties. The first is the ‘new’ interpretation of Fisher's fundamental theorem of natural selection, on which the theorem is exactly true for any evolving population that satisfies some minimal assumptions. The second is the Formal Darwinism project, which forges links between gene frequency change and optimal strategy choice. In both cases, I argue that the results fail to establish a biologically significant maximization principle. I conclude that it may be a mistake to look for universal maximization principles justified by theory alone. A more promising approach may be to find maximization principles that apply conditionally and to show that the conditions were satisfied in the evolution of particular traits.     

Natural selection, larval dispersal, and the geography of phenotype in the sea

Populations evolve generalist, specialist, and plastic strategies in response to environmental heterogeneity. Describing such within-species variation in phenotype and how it arises is central to understanding a variety of ecological and evolutionary topics. The literature on phenotypic differences among populations is highly biased; for every one article published on a marine species, at least 10 articles are published on a terrestrial species and eight focus on terrestrial plants. Here, I outline what we know from the marine literature about geographic variation in phenotype in the sea, with a principal focus on local adaptation. The theory of environmental "grain" predicts that the most likely evolutionary response (e.g., local adaptation, phenotypic plasticity, generalism, and balanced polymorphism) depends on the spatial scale of environmental variation relative to the distance that an organism disperses. Consistent with these predictions, phenotypic plasticity is stronger among invertebrates with geographically broad dispersal versus restricted dispersal (i.e., planktonic-dispersers versus direct-developers). However, contrary to predictions, the relative frequency, and spatial scale of local adaptation is not consistently greater among direct-developers relative to planktonic disperers. This indicates that the likelihood of local adaptation depends on other organismal or environmental traits. Two of the most vexing issues that remain include (1) predicting the extent to which barriers to dispersal are a cause versus consequence of phenotypic differentiation and (2) delineating the relative importance of evolutionary forces that favor or impede local adaptation. Understanding the mechanistic basis of the geography of phenotypic differences, or phenogeography, has gained recent momentum because of a need to predict impacts of global climatic change, anthropogenic disturbances, and dispersal of organisms to non-native habitats.     

Sub-antarctic House mice: colonization, survival and selection

House mice have colonized and survived successfully on a number of Sub-Antarctic islands, where the mean annual temperature is only about 5°C, but where there is little seasonal fluctuation in climate. Surprisingly this allows almost continuous breeding. On at least two islands (Macquarie and Marion), there are significant changes in gene frequency in electro-phoretically detected enzymes between young (less than three months of age) and old animals from the same population. This indicates natural selection acting in opposite directions at different stages of the life cycle. However the genetical compositions of the Macquarie and Marion populations are more distinct from each other than either is from most British samples. This means that detailed studies of the Sub-Antarctic mouse populations are likely to reveal much about local adaptation, while comparison between the responses of different populations may lead to important generalisations about the possible reaction to evolutionary challenges of a species living close to its physiological limit.