The Replicator in Retrospect

The history and theoretical role of the concept of a ``replicator'is discussed, starting with Dawkins' and Hull's classic treatmentsand working forward. I argue that the replicator concept is still auseful one for evolutionary theory, but it should be revised insome ways. The most important revision is the recognition that notall processes of evolution by natural selection require thatsomething play the role of a replicator.     

Evolutionary Epistemology, Social Epistemology, and the Demic Structure of Science

One of the principal difficulties in assessing Science as aProcess (Hull 1988) is determining the relationship between the various elements of Hull's theory. In particular, it is hard to understand precisely how conceptual selection is related to Hull's account of the social dynamics of science. This essay aims to clarify the relation between these aspects of his theory by examining his discussion of the``demic structure' of science. I conclude that the social account cando significant explanatory work independently of the selectionistaccount. Further, I maintain that Hull's treatment of the demicstructure of science points us toward an important set of issues insocial epistemology. If my reading of Science as a Process iscorrect, then most of Hull's critics (e.g., those who focus solelyon his account of conceptual selection) have ignored promisingaspects of his theory.     

黑莓的实生选优

南京中山植物园于1986年从美国引入一批黑莓品种赫尔（Hull）种子,培育出实生苗350株,其中无刺苗232株。经过8年实生选优,从中选出钟山17－13优株,并通过组培无性繁殖。其果实大,平均果重5．98g,最大9．7g,不仅远高于其他实生单株,也高于原品种赫尔。萌枝粗壮,具有较好的栽培性状,值得扩繁种植。     

Evolutionary Change and Epistemology

This paper is concerned with the debate in evolutionary epistemology about the nature of the evolutionary process at work in the development of science: whether it is Darwinian or Lamarckian. It is claimed that if we are to make progress through the many arguments that have grown up around this issue, we must return to an examination of the concepts of change and evolution, and examine the basic kinds of mechanism capable of bringing evolution about. This examination results in two kinds of processes being identified, dubbed direct and indirect, and these are claimed to exhaust all possibilities. These ideas are then applied to a selection of the debates within evolutionary epistemology. It is shown that while arguments about the pattern and rate of evolutionary change are necessarily inconclusive, those concerning the origin of novel variations and the mode of inheritance can be resolved by means of the distinctions made here. It is claimed that the process of selection in the evolution of science can also be clarified. The conclusion is that the main process producing the evolution of science is a direct or Lamarckian one although, if realism is correct, an indirect or Darwinian process plays a vital role.     

Intelligence and evolutionary epistemology

Intelligence is considered as one level in a multiple level model of evolution, each level being defined by its own units of selection and by the capacity to furnish adaptations to environmental events that occur within particular frequency bands. The specific implications for intelligence are that it is never an across-species generalist capacity but always focussed around species-typical adaptive behaviours; and that it interacts with evolution at other levels, notably at the sociocultural level in the case of Humans.     

Differential selection between the sexes and selection for sex

Anisogamy is known to generate an important cost for sexual reproduction (the famous "twofold cost of sex"). However, male-female differences may have other consequences on the evolution of sex, due to the fact that selective pressures may differ among the sexes. On the one hand, intralocus sexual conflict should favor asexual females, which can fix female-beneficial, male-detrimental alleles. On the other hand, it has been suggested repeatedly that sexual selection among males may help to purge the mutation load, providing an advantage to sexual females. However, no analytical model has computed the strength of selection acting on a modifier gene affecting the frequency of sexual reproduction when selection differs between the sexes. In this article, we analyze a two-locus model using two approaches: a quasi-linkage-equilibrium (QLE) analysis and a local stability analysis, whose predictions are verified using a multilocus simulation. We find that costly sex can be maintained when selection is stronger in males than in females, but acts in the same direction in both. Complete asexuality, however, evolves under any other form of selection. Finally, we discuss how experimental measurements of fitness variances and covariances between sexes could be used to determine the overall direction and strength on selection for sex arising from differences in selection between males and females.     

Anisogamy,sexual selection,and the evolution and maintenance of sex

Summary In the present paper we distinguish between two aspects of sexual reproduction. Genetic recombination is a universal features of the sexual process. It is a primitive condition found in simple, single-celled organisms, as well as in higher plants and animals. Its function is primarily to repair genetic damage and eliminate deleterious mutations. Recombination also produces new variation, however, and this can provide the basis for adaptive evolutionary change in spatially and temporally variable environments.The other feature usually associated with sexual reproduction, differentiated male and female roles, is a derived condition, largely restricted to complex, diploid, multicellular organisms. The evolution of anisogamous gametes (small, mobile male gametes containing only genetic material, and large, relatively immobile female gametes containing both genetic material and resources for the developing offspring) not only established the fundamental basis for maleness and femaleness, it also led to an asymmetry between the sexes in the allocation of resources to mating and offspring. Whereas females allocate their resources primarily to offspring, the existence of many male gametes for each female one results in sexual selection on males to allocate their resources to traits that enhance success in competition for fertilizations. A consequence of this reproductive competition, higher variance in male than female reproductive success, results in more intense selection on males.The greater response of males to both stabilizing and directional selection constitutes an evolutionary advantage of males that partially compensates for the cost of producing them. The increased fitness contributed by sexual selection on males will complement the advantages of genetic recombination for DNA repair and elimination of deleterious mutations in any outcrossing breeding system in which males contribute only genetic material to their offspring. Higher plants and animals tend to maintain sexual reproduction in part because of the enhanced fitness of offspring resulting from sexual selection at the level of individual organisms, and in part because of the superiority of sexual populations in competition with asexual clones.     

Emergence of cooperative linkages by random intensity of selection on a network

By assuming the random intensity of selection, the emergence of cooperation on a network is studied. We constructed an evolutionary model in which an individual plays the prisoner's dilemma game, and updates both its strategy and neighbor connections in response to its relative success in the game. The constant (strong or weak) and random intensities of selection are compared. The random intensities of selection are introduced to realize complex environmental effects on the fitness of each individual. Breaking the links on the network is realized according to fixed global parameters. We found that cooperative clusters emerged when cooperators unilaterally broke the link with defectors. The emergent networks under these conditions had a high clustering coefficient and shared some properties with a scale-free network. In addition, after a cooperator with high fitness emerged circumstantially under the random intensity of selection, we observed that the cooperative linkages emerged and spread rapidly through the network. This situation frequently occurred because of the stochastic effect on the fitness of cooperators. Thus, the origin of such phenomena is qualitatively different from the Lotka-Volterra system in which deterministic processes control the system. Cooperative linkages spread more when defectors maintained many links with cooperators.     

Sexual selection favours male parental care, when females can choose

Explaining the evolution of male care has proved difficult. Recent theory predicts that female promiscuity and sexual selection on males inherently disfavour male care. In sharp contrast to these expectations, male-only care is often found in species with high extra-pair paternity and striking variation in mating success, where current theory predicts female-only care. Using a model that examines the coevolution of male care, female care and female choice; I show that inter-sexual selection can drive the evolution of male care when females are able to bias mating or paternity towards parental males. Surprisingly, female choice for parental males allows male care to evolve despite low relatedness between the male and the offspring in his care. These results imply that predicting how sexual selection affects parental care evolution will require further understanding of why females, in many species, either do not prefer or cannot favour males that provide care.     

The extended replicator

This paper evaluates and criticises the developmental systems conception of evolution and develops instead an extension of the gene's eye conception of evolution. We argue (i) Dawkin's attempt to segregate developmental and evolutionary issues about genes is unsatisfactory. On plausible views of development it is arbitrary to single out genes as the units of selection. (ii) The genotype does not carry information about the phenotype in any way that distinguishes the role of the genes in development from that other factors. (iii) There is no simple and general causal criterion which distinguishes the role of genes in development and evolution. (iv) There is, however, an important sense in which genes but not every other developmental factor represent the phenotype. (v) The idea that genes represent features of the phenotype forces us to recognise that genes are not the only, or almost the only, replicators. Many mechanisms of replication are involved in both development and evolution. (vi) A conception of evolutionary history which recognises both genetic and non-genetic replicators, lineages of replicators and interactors has advantages over both the radical rejection of the replicator/interactor distinction and the conservative restriction of replication to genetic replication.     

Population dynamics and evolutionary processes: the manifold roles of habitat selection

Summary Any character that has a substantial effect on a species' distribution and abundance can exert a variety of indirect effects on evolutionary processes. It is suggested that an organism's capacity for habitat selection is just such a character. Habitat selection can constrain the selective environment experienced by a population. Habitat selection can also indirectly influence the relative importance of natural selection, drift, and gene flow, through its effect on population size and growth rate. In many circumstances (but not all), habitat selection increases population size and growth rate, and thereby makes selection in a local environment more effective than drift and gene flow.     

The Philosophy of Donald T. Campbell: A Short Review and Critical Appraisal

Aside from his remarkable studies in psychology and the social sciences, Donald Thomas Campbell (1916–1996) made significant contributions to philosophy, particularly philosophy of science,epistemology, and ethics. His name and his work are inseparably linked with the evolutionary approach to explaining human knowledge (evolutionary epistemology). He was an indefatigable supporter of the naturalistic turn in philosophy and has strongly influenced the discussion of moral issues (evolutionary ethics). The aim of this paper is to briefly characterize Campbells work and to discuss its philosophical implications. In particular, I show its relevance to some current debates in the intersection of biology and philosophy. In fact, philosophy of biology would look poorer without Campbells influence. The present paper is not a hagiography but an attempt to evaluate and critically discuss the meaning of Campbells work for philosophy of biology and to encourage scholars working in this field to read and re-read this work which is both challenging and inspiring.     

Assessing evolutionary epistemology

There are two interrelated but distinct programs which go by the name evolutionary epistemology. One attempts to account for the characteristics of cognitive mechanisms in animals and humans by a straightforward extension of the biological theory of evolution to those aspects or traits of animals which are the biological substrates of cognitive activity, e.g., their brains, sensory systems, motor systems, etc. (EEM program). The other program attempts to account for the evaluation of ideas, scientific theories and culture in general by using models and metaphors drawn from evolutionary biology (EET program). The paper begins by distinguishing the two programs and discussing the relationship between them. The next section addresses the metaphorical and analogical relationship between evolutionary epistemology and evolutionary biology. Section IV treats the question of the locus of the epistemological problem in the light of an evolutionary analysis. The key questions here involve the relationship between evolutionary epistemology and traditional epistemology and the legitimacy of evolutionary epistemology as epistemology. Section V examines the underlying ontological presuppositions and implications of evolutionary epistemology. Finally, section VI, which is merely the sketch of a problem, addresses the parallel between evolutionary epistemology and evolutionary ethics.This research was supported, in part, by a grant from the National Science Foundation # SES—8308720. I want to thank Richard Lewontin and the Museum of Comparative Zoology, Harvard University for their hospitality and support during Spring 1984. I also want to thank David Hull, Charles Dyke and Michael Ruse for helpful comments on an earlier draft, and Andy Altman for pointing out to me the passage from Inherit the Wind.     

Kin selection may inhibit the evolution of reciprocation

Kin selection and reciprocal cooperation provide two candidate explanations for the evolution of cooperation. Models of the evolution of cooperation have typically focussed on one or the other mechanism, despite claims that kin selection could pave the way for the evolution of reciprocal cooperation. We describe a computer simulation model that explicitly supports both kin selection and reciprocal cooperation. The model simulates a viscous population of discrete individuals with social interaction taking the form of the Prisoner's Dilemma and selection acting on performance in these interactions. We recount how the analytical and empirical study of this model led to the conclusion that kin selection may actually inhibit the evolution of effective strategies for establishing reciprocal cooperation.     

Multilevel selection: the evolution of cooperation in non-kin groups

Hamiltons (1964a, 1964b) landmark papers are rightly recognized as the formal basis for our understanding of the evolution of altruistic traits. However, Hamiltons equation as he originally expressed it is simplistic. A genetically oriented approach to studying multilevel selection can provide insights into how the terminology and assumptions used by Hamilton can be generalized. Using contextual analysis I demonstrated that Hamiltons rule actually embodies three distinct processes, group selection, individual selection, and transmission genetics or heritability. Whether an altruistic trait will evolve depends the balance of all of these factors. The genetical approach, and particularly, contextual analysis provides a means of separating these factors and examining them one at a time. Perhaps the greatest issue with Hamiltons equation is the interpretation of r. Hamilton (1964a) interpreted this as relatedness. In this paper I show that what Hamilton called relatedness is more generally interpreted as the proportion for variance among groups, and that many processes in addition to relatedness can increase the variance among groups. I also show that the evolution of an altruistic trait is driven by the ratio of the heritability at the group level to the heritability at the individual level. Under some circumstances this ratio can be greater than 1. In this situation altruism can evolve even if selection favoring selfish behavior is stronger than selection favoring altruism.     

Natural selection and veridical perceptions

Does natural selection favor veridical perceptions, those that more accurately depict the objective environment? Students of perception often claim that it does. But this claim, though influential, has not been adequately tested. Here we formalize the claim and a few alternatives. To test them, we introduce “interface games,” a class of evolutionary games in which perceptual strategies compete. We explore, in closed-form solutions and Monte Carlo simulations, some simpler games that assume frequency-dependent selection and complete mixing in infinite populations. We find that veridical perceptions can be driven to extinction by non-veridical strategies that are tuned to utility rather than objective reality. This suggests that natural selection need not favor veridical perceptions, and that the effects of selection on sensory perception deserve further study.     

Synergistic selection and graded traits

Fitness interactions where benefits are shared only between individuals with similar traits are often referred to as synergistic. Examples include defence characters, like insect warning colouration and plant unpalatability, and joint activities needing the active participation of all group members, such as cooperative hunting. Previous analyses, assuming discrete variation in the trait, have shown that synergistic selection can be a sufficient explanation for the evolutionary stability of such traits. Here, we investigate the consequences of graded variation in the trait responsible for synergistic effects. Classifying the synergism as unbiased when an individual receives maximum associational benefit by having the same trait value as its neighbours, and letting a positive (negative) bias represent the maximum above (below) this value, we show that only positively biased synergistic selection can enhance a graded trait. Thus for graded traits, a synergistic benefit is not in itself sufficient for evolutionary stability. We study possible reasons for synergistic bias in a simple model of plant defences against herbivores, and suggest that the processes of herbivore avoidance learning and diet selection are probable causes of positive bias. We propose that mammalian herbivores exposed to a given level of toxicity will show stronger feeding aversion to higher toxicity, resulting in positively biased synergistic selection of plant defence traits. Positive bias produced by avoidance learning may, in a similar way, also select for defence signals.     

Michael Ruse and His Fifteen Years of Booknotes – For Better or for Worse

In this paper I trace Michael Ruse's Booknotes from the first volumeof Biology and Philosophy in 1986 to the present. I deal withboth the style and the content of these booknotes. Ruse paid specialattention to authors outside of the traditional English axis as wellas to feminist writers. He complained that too much attention wasbeing paid to certain topics (e.g., evolutionary ethics, evolutionaryepistemology, the species problem and reduction) while other, moreimportant topics were all but ignored (e.g., natural selection,population genetics, levels of selection and extraterrestrial life).He also dealt with the Darwin Industry. Creationism, his love-haterelationships with several authors and his undiluted love of CharlesDickens.     

Unconscious selection and the evolution of domesticated Plants

Two types of selection operate (and complement each other) in plants under domestication: (a) conscious or intentional selection applied by the growers for traits of interest to them; (b) unconscious or automatic selection brought about by the fact that the plants concerned were taken from their original wild habitats and placed in new (and usually very different) human-made or human-managed environments. The shift in the ecology led automatically to drastic changes in selection pressures. Numerous adaptations vital for survival in the wild environments lost their fitness under the new sets of conditions. New traits were automatically selected, resulting in the build-up of characteristic “domestication syndromes,” each fitting the specific agricultural environment provided by the farmer. The present paper assesses the evolutionary consequences of the introduction of the wild plants into several sets of contrasting farming situations. These include: (a) the type of maintenance applied, whether seed planting or vegetative propagation; (b) the plant organs for which the crop has been grown, whether they are reproductive parts or vegetative parts; (c) the impact of the system of tilling, sowing, and reaping on the evolution of grain crops; (d) the impact of the horticultural environment on fruit crops.     

Assortment and the analysis of natural selection on social traits

A central problem in evolutionary biology is to determine whether and how social interactions contribute to natural selection. A key method for phenotypic data is social selection analysis, in which fitness effects from social partners contribute to selection only when there is a correlation between the traits of individuals and their social partners (nonrandom phenotypic assortment). However, there are inconsistencies in the use of social selection that center around the measurement of phenotypic assortment. Here, we use data analysis and simulations to resolve these inconsistencies, showing that: (i) not all measures of assortment are suitable for social selection analysis; and (ii) the interpretation of assortment, and how to detect nonrandom assortment, will depend on the scale at which it is measured. We discuss links to kin selection theory and provide a practical guide for the social selection approach.