So far in this course we have dealt entirely with the evolution of characters that are controlled by simple Mendelian inheritance at a single locus. There are notes on the course website about gametic disequilibrium and how allele frequencies change at two loci simultaneously, but we didn’t discuss them. In every example we’ve considered we’ve imagined that we could understand something about evolution by examining the evolution of a single gene. That’s the domain of classical population genetics. For the next few weeks we’re going to be exploring a field that’s actually older than classical population genetics, although the approach we’ll be taking to it involves the use of population genetic machinery. If you know a little about the history of evolutionary biology, you may know that after the rediscovery of Mendel’s work in 1900 there was a heated debate between the “biometricians” (e.g., Galton and Pearson) and the “Mendelians” (e.g., de Vries, Correns, Bateson, and Morgan). Biometricians asserted that the really important variation in evolution didn’t follow Mendelian rules. Height, weight, skin color, and similar traits seemed to

Human biochemical genetics

Genetics and analysis of quantitative traits

Recent studies suggest that populations and species often exhibit behavioral syndromes; that is, suites of correlated behaviors across situations. An example is an aggression syndrome where some individuals are more aggressive, whereas others are less aggressive across a range of situations and contexts. The existence of behavioral syndromes focuses the attention of behavioral ecologists on limited (less than optimal) behavioral plasticity and behavioral carryovers across situations, rather than on optimal plasticity in each isolated situation. Behavioral syndromes can explain behaviors that appear strikingly non-adaptive in an isolated context (e.g. inappropriately high activity when predators are present, or excessive sexual cannibalism). Behavioral syndromes can also help to explain the maintenance of individual variation in behavioral types, a phenomenon that is ubiquitous, but often ignored. Recent studies suggest that the behavioral type of an individual, population or species can have important ecological and evolutionary implications, including major effects on species distributions, on the relative tendencies of species to be invasive or to respond well to environmental change, and on speciation rates. Although most studies of behavioral syndromes to date have focused on a few organisms, mainly in the laboratory, further work on other species, particularly in the field, should yield numerous new insights.

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Behavioral syndromes: an ecological and evolutionary overview.

Temperament describes the idea that individual behavioural differences are repeatable over time and across situations. This common phenomenon covers numerous traits, such as aggressiveness, avoidance of novelty, willingness to take risks, exploration, and sociality. The study of temperament is central to animal psychology, behavioural genetics, pharmacology, and animal husbandry, but relatively few studies have examined the ecology and evolution of temperament traits. This situation is surprising, given that temperament is likely to exert an important influence on many aspects of animal ecology and evolution, and that individual variation in temperament appears to be pervasive amongst animal species. Possible explanations for this neglect of temperament include a perceived irrelevance, an insufficient understanding of the link between temperament traits and fitness, and a lack of coherence in terminology with similar traits often given different names, or different traits given the same name. We propose that temperament can and should be studied within an evolutionary ecology framework and provide a terminology that could be used as a working tool for ecological studies of temperament. Our terminology includes five major temperament trait categories: shyness-boldness, exploration-avoidance, activity, sociability and aggressiveness. This terminology does not make inferences regarding underlying dispositions or psychological processes, which may have restrained ecologists and evolutionary biologists from working on these traits. We present extensive literature reviews that demonstrate that temperament traits are heritable, and linked to fitness and to several other traits of importance to ecology and evolution. Furthermore, we describe ecologically relevant measurement methods and point to several ecological and evolutionary topics that would benefit from considering temperament, such as phenotypic plasticity, conservation biology, population sampling, and invasion biology.

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Integrating animal temperament within ecology and evolution.

Beginning with Emlen (1966) and MacArthur and Pianka (1966) and extending through the last ten years, several authors have sought to predict the foraging behavior of animals by means of mathematical models. These models are very similar,in that they all assume that the fitness of a foraging animal is a function of the efficiency of foraging measured in terms of some "currency" (Schoener, 1971) -usually energy- and that natural selection has resulted in animals that forage so as to maximize this fitness. As a result of these similarities, the models have become known as "optimal foraging models"; and the theory that embodies them, "optimal foraging theory." The situations to which optimal foraging theory has been applied, with the exception of a few recent studies, can be divided into the following four categories: (1) choice by an animal of which food types to eat (i.e., optimal diet); (2) choice of which patch type to feed in (i.e., optimal patch choice); (3) optimal allocation of time to different patches; and (4) optimal patterns and speed of movements. In this review we discuss each of these categories separately, dealing with both the theoretical developments and the data that permit tests of the predictions. The review is selective in the sense that we emphasize studies that either develop testable predictions or that attempt to test predictions in a precise quantitative manner. We also discuss what we see to be some of the future developments in the area of optimal foraging theory and how this theory can be related to other areas of biology. Our general conclusion is that the simple models so far formulated are supported are supported reasonably well by available data and that we are optimistic about the value both now and in the future of optimal foraging theory. We argue, however, that these simple models will requre much modification, espicially to deal with situations that either cannot easily be put into one or another of the above four categories or entail currencies more complicated that just energy.

Citation classic - optimal foraging - a selective review of theory and tests

Three major hypotheses, based upon mechanisms of sexual selection, intersexual food competition and reproductive role division, have been advanced to explain the evolution of sexual dimorphism in body size and morphology of animals Genetic models suggest that all of the hypotheses are plausible, and empirical studies demonstrate that each of the three mechanisms operates in natural populations However, problems arise in testing hypotheses for the evolution of sexual dimorphism: more than one mechanism may be operating simultaneously, and the demonstrated occurrence of a mechanism does not indicate that it actually results in selection for dimorphism A recent statistical technique offers a solution to these problems and provides a promising new approach to the study of sexual dimorphism, in which researchers can assess the relative importance of each mechanism in present-day selection for sexual dimorphism within a species

The evolution of sexual dimorphism in animals: Hypotheses and tests

Mate choice by female crickets is influenced by predation risk

A test for correlations among fitness-linked behavioural traits in the spider Agelenopsis aperta (Araneae, Agelenidae)

The calls of male field crickets attract sexually receptive females. In Gryllus integer, males differ from one another in their durations of uninterrupted calling (calling bout lengths). Tape recordings of the calls of 50 wild-caught males revealed that 14 males spent most of their calling time in short bouts (Fig. 1A), 18 in both short and long bouts (Fig. 1B), and 18 in long bouts (Fig. 1C). Re-recordings of 32 males after 3 weeks showed that calling bout lengths of individual males are stable with time (age) (Fig. 2). Three phonotaxis experiments investigated whether calling bout lengths of males affect female preferences. They demonstrated that (1) females can discriminate among conspecific males on the basis of calls alone; (2) females are preferentially attracted to males with long calling bout lengths; and (3) calling bout length is the specific factor responsible for preferential attraction. These results precisely identify a criterion that females use to discriminate among potential mates of their own species.

Female preferences for male calling bout duration in a field cricket

Modern models for the evolution of conspicuous male mating displays assume that males with conspicuous displays must bear the cost of enhanced predation risk. However, if males can compensate behav...

Ann V. Hedrick

Papers

The evolution of sexual dimorphism in animals: Hypotheses and tests

Mate choice by female crickets is influenced by predation risk

A test for correlations among fitness-linked behavioural traits in the spider Agelenopsis aperta (Araneae, Agelenidae)

Female preferences for male calling bout duration in a field cricket

Crickets with extravagant mating songs compensate for predation risk with extra caution