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Wilson_1975_Wilson_1975_Sociobiology The Abridged Edition, p. 305-325 Adaptation In evolutionary biology, a particular structure, physiological process, or behavior that equips an organism to survive and reproduce. Also, the evolutionary process that leads to the formation of such a trait. (See also Natural selection.) Allele A particular form of a gene, distinguishable in its nucleotide sequence from other forms or alleles of the same gene, hence occupying the same locus in the chromosomes. Assimilation function The relation between the probability of adopting a culturgen or switching from one culturgen to another (or the rate at which such changes are made) and the proportion of other members of the society who possess the culturgen. For example, the probability may rise linearly as others adopt the culturgen, or not change at all. Coevolution An evolutionary change "in a trait of the individuals in one population in response to a trait of the individuals of a second population, followed by an evolutionary response by the second population to the change in the first" (Janzen, 1980). We have extended this biological term to include the reciprocal effects of genetic and cultural evolution (see Gene-culture coevolution). More generally, diachronic changes in two or more interacting objects or systems. Culture The sum of all of the artifacts, behavior, institutions, and mental concepts transmitted by learning among members of a society, and the holistic patterns they form. In human beings the culture of each society is characterized by some traits (culturgens) that are general to the whole species and others that are idiosyncratic. The transmission also entails cognition, which loads the traits with meaning and typically but not invariably labels them with words and other symbols that are then manipulated in language to create complex new messages. (See also Culturgen, Euculture, Gene-culture coevolution, Protoculture.) Culturgen (pronounced "kul' tur jen") The basic unit of culture. A relatively homogeneous set of artifacts, behaviors, or mentifacts (mental constructs having little or no direct correspondence to reality) that either shares without exception one or more attribute states selected for their functional importance or at least shares a consistently recurrent range of such attribute states within a given polythetic set. Culturgens can be mapped into node-link structures in long term memory and in many instances can be treated as identical with them. Culturgen assimilation The process whereby culturgens are invented and adopted in behavioral categories whose epigenetic rules are permissive toward the culturgens. (Compare with Genetic assimilation.) Differentiation In biology, the process whereby originally similar cells follow different developmental pathways. Dominance In behavioral biology, the physical domination of some members of a group by other members, usually in relatively orderly and long-lasting patterns (called dominance hierarchies). In genetics, the ability of one allelic form of a gene to determine the phenotype of a heterozygous individual, in which the homologous chromosome carries a different allele. For example, if A and a are two allelic forms of a gene, A is said to be dominant to a if AA diploids and Aa diploids are phenotypically identical (or nearly so) and are distinguishable from aa diploids. The a allele then is said to be recessive. Enculturation The transmission of a particular culture, especially to young members of the society. Some authors make a distinction between socialization, regarded as the development of social behavior basic to every normal human being, and enculturation, the act of learning one culture in all its uniqueness and particularity (see for example Mead, 1963). However, in this work we have used the two terms interchangeably. (See also Socialization.) Epigene A gene affecting a particular developmental process under study. Epigenesis The processes of interaction between genes and the environment that ultimately result in the distinctive anatomical, physiological, cognitive, and behavioral traits of the organism. Epigenetic events occur from the moment that RNA is transcribed from DNA, then forward through all phases of development to the final assembly of tissues and cognition itself; the interacting environment first is composed entirely of the cell medium but then expands until—in the case of human beings especially—it includes all aspects of culture. Epigenetic rule Any regularity during epigenesis that channels the development of an anatomical, physiological, cognitive, or behavioral trait in a particular direction. Epigenetic rules are ultimately genetic in basis, in the sense that their particular nature depends on the DNA developmental blueprint. They occur at all stages of development, from protein assembly through the complex events of organ construction to learning. Some epigenetic rules are inflexible, with the final phenotype being buffered from all but the most drastic environmental changes. Others permit a flexible response to the environment; yet even these may be invariant, in the sense that each possible response in the array is matched to one environmental cue or a set of cues through the operation of special control mechanisms. In cognitive development, the epigenetic rules are expressed in any one of the many processes of perception and cognition to influence the form of learning and the transmission of culturgens. The primary epigenetic rules of mental development are based upon the more automatic processes that lead from sensory filtering to perception (Chapter 2). The secondary epigenetic rules affect information displayed in the perceptual fields and include the channeling of memory, emotional response, decision making, and ultimately the usage bias curves (see also Chapter 3). Epigenotype The set of genes affecting a particular developmental process under study. Epistasis The nonadditive interaction of genes on different loci, especially the suppression of one set by another. Ethnographic curve The array of probabilities that a society will possess particular cultural patterns. In the present analysis of gene-culture coevolution, cultural patterns are defined as the various proportions of individuals in the society that possess one active culturgen as opposed to another. The curve can be based on many societies observed concurrently or on one or a few societies observed repetitively through time. Ethology The biological study of whole patterns of animal behavior in natural environments, stressing the analysis of adaptation and the evolution of the patterns. It is distinguished from sociobiology, which is the study of the biological basis of social behavior in particular, and especially of the properties of entire societies. (See also Sociobiology.) Euculture The most advanced form of culture, in which individuals not only teach and learn information but also conceptualize much of it into concrete entities that can be more readily labeled by symbols and handled by language. (See also Protoculture, Reification.) Evolution Any gradual change. Genetic or organic evolution, often referred to as evolution for short, is any genetic change in organisms from generation to generation; or, more strictly, a change in gene frequencies within populations from generation to generation. Cultural evolution is an alteration in culturally transmitted artifacts, behavior, institutions, or concepts within or across generations. As purely conceived, cultural evolution does not necessarily entail any form of genetic change, but when the two modes of change are coupled in a reciprocally interacting manner, the total process is referred to as gene-culture coevolution. Evolutionary biology The collective disciplines of biology that treat the evolutionary process and the characteristics of populations of organisms, as well as ecology, behavior, and systematics. (See also Population biology.) Fitness In evolutionary biology, differential representation in later generations. (See Gene-culture fitness, Genetic fitness.) Gene-culture adaptive landscape The array of gene-culture fitnesses of all of the combinations of genotypes and culturgens that can be assembled by a population. Gene-culture coevolution The coupled evolution of genes and culture. More precisely, any change in gene frequencies that alters culturgen frequencies in such a way that the culturgen changes alter the gene frequencies as well. (See also Coevolution.) Gene-culture coevolutionary theory The theory treating the full circuit of the coevolution process; sometimes referred to as gene-culture theory (GCT) for brevity. Gene-culture fitness The contribution to the next generation of one particular combination of genotype and culturgens relative to the contribution of other such combinations.By definition, the differential contribution leads eventually to the prevalence of the genotypes with the highest fitnesses. (Compare with Genetic fitness.) Gene-culture theory (GCT) See Gene-culture coevolutionary theory. Gene-culture translation The effect of epigenetic rules of individual development on social patterns. Gene-culture transmission The transmission of culturgens in which the choices are not all equiprobable. (Compare with Pure cultural transmission and Pure genetic transmission.) Gene flow The exchange of genes between different species (an extreme case referred to as hybridization) or between different populations, caused by the migration of individuals or the long-distance dispersal of gametes. Gene pool All the genes (hence, hereditary material) in a population. Genetic assimilation A change in the frequency of genes making it more likely that individuals will develop a previously rare trait. In a typical sequence the phenotype appears as a result of exposure to an unusual environment; the bearers of the trait prove superior at survival and reproduction; as a consequence, there is an increase in frequency of genes that promote the appearance of the trait even in normal environments. (Compare with Culturgen assimilation.) Genetic drift Evolution (change in gene proportions) by chance processes alone. Genetic evolution See Evolution. Genetic fitness The contribution to the next generation of one genotype relative to the contribution of other genotypes, averaged over all the culturgens possessed by individuals of various genotypes (hence in effect without reference to culturgens). By definition, the differential contribution leads eventually to the prevalence of the genotypes with the highest fitnesses. (Compare with Gene-culture fitness.) Group Any set of organisms, belonging to the same species, that remain together for a period of time while interacting with one another to a distinctly greater degree than with other conspecific organisms. Group selection Selection that operates on two or more members of a lineage group as a unit. Defined broadly, group selection includes both kin selection and selection of entire populations. (See also Kin selection.) Habituation The simplest form of learning, in which an animal presented with a stimulus without reward or punishment eventually ceases to respond. (See also Dishabituation.) Heritability The fraction of variation of a trait within a population—more precisely, the fraction of its variance, which is the statistical measure—due to heredity as opposed to environmental influences. A heritability score of one means that all the variation is genetic in basis; a heritability score of zero means that all the variation is the result of the environment. Heterarchy A mixed-level hierarchy: a hierarchy-like system of two or more levels of units, with the properties of the higher levels affecting the lower in some degree, but with additional activity in the lower units feeding back to some extent to influence the higher level. The units at the higher level are composed of the lower-level units. For example, individual behavior generates institutions and other macrocultural patterns, which then influence individual decisions. Hierarchy A system whose components are organized in such a way that two or more natural levels of system-into-subsystem decompositions can be distinguished. (See also Heterarchy.) Holism The method of explanation of complex systems, and the philosophy motivating it, that includes not just the properties of the components of the system but also the patterns and even the history of their relationships and the manner in which they are integrated to cause the system to function as a distinct, superordinate entity. Extreme holism eschews any meaningful connections between superordinate entities and their components. (Contrast with Reductionism.) Imprinting A rigid form of learning, usually induced during a brief, "sensitive" period of the life cycle, in which an animal comes to make a particular response to only one other animal or object. Inbreeding The mating of kin. The degree of inbreeding is measured by the fraction of genes that will be identical owing to common descent. Inclusive fitness The sum of an individual's own fitness plus all its influence on fitness in its relatives other than direct descendants; hence the total effect of kin selection with reference to an individual. Kin selection The change in gene frequencies due to one or more individuals favoring or disfavoring the survival and reproduction of relatives (other than offspring) who possess the same genes by common descent. Knowledge structure Any set of linked nodes in long term memory, from a single node to a schema to the entire contents of long term memory. Some knowledge structures can be mapped onto culturgens and in many cases are identical with them. Lamarckism The theory of evolution by acquired characteristics, as propounded by Jean Baptiste de Lamarck (1744-1829). The belief that traits acquired by the activities of individuals are passed directly to the offspring—in contradiction to Darwinism, which holds that offspring receive only genetic traits unaltered by the environment and that evolution occurs by the greater success of individuals inheriting these traits, in other words by natural selection. (See also Darwinism.) Learning Behavior that is modified according to an organism's experience. Learning rule An epigenetic rule that directly affects the learning of particular culturgens. Given appropriate methods, the learning rule can be evaluated by means of a usage bias curve. Leash principle The principle deduced from natural selection theory that epigenetic rules will tend to evolve in such a way as to make individuals choose certain culturgens over others; in other words, "the genes hold culture on a leash." (See also Epigenetic rule.) Logistic A particular form of change, such as growth of a population or the spread of a culturgen, that first accelerates and then slows steadily as the process approaches its upper limit. Major gene A gene that individually accounts for a large part of the variation of a trait under consideration, as opposed to a polygene. The effects of a major gene can be altered to some degree by modifier genes. (See also Polygenes.) Microevolution A small amount of evolutionary change, consisting in minor alterations in gene proportions, chromosome structure, or chromosome numbers. (A larger amount of change would be referred to as macroevolution or simply as evolution.) Multifactorial inheritance The control of variation in a trait by genes situated on more than one locus; the term usually implies further that the genes are polygenes. Mutant An allele that has recently been created by mutation or at least is rare enough in the population to be at the frequency level expected from recent mutation activity. Also, an individual organism carrying such an allele. Mutation In the broad sense, any discontinuous change in the genetic constitution of an organism. In the narrow sense, the word refers usually to a "point mutation," a change along a very narrow portion of the nucleic acid sequence. Natural selection The differential contribution to the next generation by individuals of different genetic types but belonging to the same population. This is the basic mechanism proposed by Darwin and is generally regarded today as the main guiding force in evolution. Penetrance In gene-culture theory, the number of culturgens of a particular category that various members of the population are able to incorporate. (See also Selectivity.) Perception The psychological process of receiving and recognizing cues from the environment. The environment includes both the outside world and the internal physiological state of the organism. (See also Cognition.) Perceptual field The configurations formed within the associative centers of the brain that correspond to information received through the sense organs. Thus the organization of light frequencies in the form of discrete hues represents a basic portion of the brain's perceptual field. Population A set of organisms belonging to the same species and occupying a clearly delimited space at the same time. Often a particular population is coterminous with a society, especially if the society forms a closed or partially closed breeding system. Population biology The scientific study of the biological properties peculiar to populations, including their distribution, ecology, growth, age characteristics, and genetic structure. (See also Demography, Population ecology, and Population genetics—all of which are subdisciplines of population biology.) Population ecology The scientific study of the distinctive relationships between whole populations of organisms and their environments, covering such topics as the determinants of mortality and fertility, the rate of population growth, and the processes of competition and symbiosis. Population genetics The scientific study of the distinctive properties of heredity viewed at the level of populations, including genetic diversity and the changes in gene frequencies that comprise evolution. Principle of parsimony The conjecture that epigenetic rules tend to evolve to the lowest level of genetic specification that will suffice; it follows that special physiological devices in cognition will decline when the need for specificity is reduced or eliminated. (See also Transparency principle.) Protoculture A form of culture, found in a few higher animals, in which information is transmitted by imitation and even by teaching, but in which reification and symbolization do not occur. (Contrast with Euculture.) Pure cultural transmission The transmission of culturgens in which all of the choices are equiprobable. (Compare with Pure genetic transmission and Geneculture transmission.) Pure genetic transmission In the theory of gene-culture coevolution, the extreme case in which the transmission of culturgens is limited to a single choice. (Compare with Pure cultural transmission and Gene-culture transmission.) Recessive In genetics, referring to an allele the phenotype of which is suppressed when the allele occurs in combination with a dominant allele. Reductionism Oversimplification in the explanation of a complex system owing to the attempt to account for the system solely on the basis of the properties of its components. Usually ascribed by social scientists to biologists, by biologists to chemists, and by chemists to physicists. (Contrast with Holism.) Reification The mental activity in which hazily perceived and relatively intangible phenomena, such as complex arrays of objects or activities, are given a factitiously concrete form, simplified, and labeled with words or other symbols. Selection pressure Any process in the environment that results in natural selection. For example, food shortage, the activity of a predator, or competition for a mate from other members of the same sex provide selection pressures by causing individuals of various genetic types to survive to different average ages, to reproduce at different rates, or both. Selectivity In gene-culture theory, the strength of the tendency for individuals to use one or a small set of culturgens in preference to others that are available. Sensory receptor A cell, tissue, or organ specialized for detecting physical signals (such as light or sound) and translating them into coded responses that can then be "read" by interneurons and transferred to the central nervous system. Social contagion The spreading of a successful culturgen through a society by means of personal contact. Socialization The total modification of behavior in an individual because of the interaction of that individual with other members of the society, including its parents. Some authors make a distinction between socialization, regarded as the development of social behavior basic to every normal human being, and enculturation, the act of learning one culture in all its uniqueness and particularity (see for example Mead, 1963). In this work we have used the two terms interchangeably. Society A group of individuals belonging to the same species and organized in a cooperative manner. The diagnostic criterion of a society is reciprocal communication of a cooperative nature, extending beyond sexual activity alone. (See Wilson, 1975.) Sociobiology The systematic study of the biological basis of all forms of social behavior, including sexual behavior and parent-offspring interaction, in all kinds of organisms. Teaching rule An epigenetic rule that directly affects the teaching of particular culturgens. Transparency principle The conjecture that the more the relation between a category of behavioral response and genetic fitness depends on environmental circumstances, the more clearly the conscious mind perceives the relation and the more flexible the response. Usage See Usage of culturgen. Usage bias curve A curve that displays the probabilities that an organism will use one or the other of various culturgens within a culturgen category, given that it possesses a particular genotype and lives in a particular environment. Usage of culturgen In the broad sense we have employed throughout, any response to a culturgen: either the initial perception and learning of the culturgen, or a relative preference for it during reflection and valuation, or a decision to employ it. Usage pattern The proportion of individuals in a population that use one culturgen as opposed to another.