Built for speed, not for comfort. Darwinian theory and human culture.

Darwin believed that his theory of evolution would stand or fall on its ability to account for human behavior. No species could be an exception to his theory without imperiling the whole edifice. The ideas in the Descent of Man were widely discussed by his contemporaries although they were far from being the only evolutionary theories current in the late nineteenth century. Darwin's specific evolutionary ideas and those of his main followers had very little impact on the social sciences as they emerged as separate disciplines in the early Twentieth Century. Not until the late twentieth century were concerted, sophisticated efforts made to apply Darwinian theory to human behavior. Why such a long delay? We argue that Darwin's theory was rather modern in respects that conflicted with Victorian sensibilities and that he and his few close followers failed to influence any of the social sciences. The late Twentieth Century work takes up almost exactly where James Baldwin left off at the turn of the century.

Complex societies : The evolutionary origins of a crude superorganism.

The complexity of human societies of the past few thousand years rivals that of social insect societies. We hypothesize that two sets of social "instincts" underpin and constrain the evolution of complex societies. One set is ancient and shared with other social primate species, and one is derived and unique to our lineage. The latter evolved by the late Pleistocene, and led to the evolution of institutions of intermediate complexity in acephalous societies. The institutions of complex societies often conflict with our social instincts. The complex societies of the past few thousand years can function only because cultural evolution has created effective "work-arounds" to manage such instincts. We describe a series of work-arounds and use the data on the relative effectiveness of WWII armies to test the work-around hypothesis.

Group selection among alternative evolutionarily stable strategies.

Many important models of the evolution of social behavior have more than one evolutionarily stable strategy (ESS). Examples include co-ordination games, contests, mutualism, reciprocity, and sexual selection. Here we show that when there are multiple evolutionarily stable strategies, selection among groups can cause the spread of the strategy that has the lowest extinction rate or highest probability of contributing to the colonization of empty habitats, and that this may occur even when groups are usually very large, migration rates are substantial, and "extinction" entails only the disruption of the group and the dispersal of its members. The main requirements are: (1) individuals drawn from a single surviving group make up a sufficiently large fraction newly formed groups, and (2) the processes increasing the frequency of successful strategies within groups are strong compared to rate of migration among groups. The latter condition suggests that this form of group selection will be particularly important when behavioral variation is culturally acquired.

The evolution of reciprocity in sizable groups.

Recently, several authors have investigated the evolution of reciprocal altruism using the repeated prisoner's dilemma game. These models suggest that natural selection is likely to favor behavioral strategies leading to reciprocal cooperation when pairs of individuals interact repeatedly in potentially cooperative situations. Using the repeated n-person prisoner's dilemma game, we consider whether reciprocal altruism is also likely to evolve when social interactions involve more individuals. We show that the conditions that allow the evolution of reciprocal cooperation become extremely restrictive as group size increases.

The cultural transmission of acquired variation: effects on genetic fitness.

In a cultural species individuals acquire some aspects of their phenotypes by imitating conspecifics. Presumably, cultural transmission has some adaptive function in such species. Some authors have suggested that one such function may be the ability to transmit phenotypes that have been modified by learning or some other form of phenotypic plasticity. To examine this hypothesis we have compared the genetic fitness of cultural and noncultural individuals in three different models. In each model the environment is assumed to be variable, either in space or time, and learning modifies each individual's phenotype so as to increase the individual's fitness in the local environment. Cultural individuals transmit the modified phenotype to their offspring, while noncultural individuals do not. The models are different in the following ways: two models assume a dichotomous phenotype, one in a temporally varying environment and one in a spatially varying environment; the third model assumes a quantitative phenotype in a temporally variable environment. The two models which assume a temporally varying environment yield similar results. Genetic transmission is favored in highly autocorrelated environments while cultural transmission in environments with intermediate autocorrelation. In the spatially varying model cultural transmission is always favored.

Effect of phenotypic variation on kin selection.

An expression for the equilibrium of the mean phenotypic value of a quantitative character is derived for a model in which the fitness of an individual depends on its own phenotype and the mean phenotypic value of a group of related individuals. When selection is weak the equilibrium mean is well predicted by Hamilton's k > 1/r rule (k is the ratio of mean fitness gained by recipient of altruistic behavior to mean fitness lost by donor; r is mean coefficient of relationship between donor and recipient). When selection is strong, however, the equilibrium mean depends on the heritability of the character. Low heritability can lead to substantially more "altruism" than predicted by the k > 1/r rule.

Spatial scales of current speed and phytoplankton biomass fluctuations in lake tahoe.

Spectral analysis of current speed and chlorophyll a measurements in Lake Tahoe, California and Nevada, indicates that considerably more variance exists at longer length scales in chlorophyll than in the current speeds. Increasingly, above scales of approximately 100 meters, chlorophyll does not behave as a simple passive contaminant distributed by turbulence, which indicates that biological processes contribute significantly to the observed variance at these large length scales.