Search

In dealing with the spatial and temporal variability of prey species, predators may be able to forage optimally if they have flexible and rapid behavioral plasticity rather than predetermined responses. For predators that learn to focus attention on the cryptic prey type most frequently encountered during recent searching (termed a “search image”), rare prey types may be overlooked because of a focus on more common prey. Search imaging reflects biased searching for one of a number of available prey types, and has been studied widely in birds and mammals. Here we discuss the significant implications of this phenomenon for insect predator–prey systems, particularly with respect to parasitic wasps searching for host species using learned olfactory cues. We (1) review studies about perceptual development through individual ontogeny, (2) define the term “search image” and discuss the cognitive mechanisms involved in search-image formation, (3) discuss the role of search images and frequency-dependent predation as a proximate mechanism in the maintenance of prey diversity, (4) examine data on host–parasitoid olfactory search imaging, and (5) conclude by identifying important research areas for future studies in the field of olfactory search images.

Although the study of adaptation is central to biology, two types of adaptation are recognized in the biological field: physiological adaptation (accommodation or acclimation; an individual organism’s phenotype is adjusted to its environment) and evolutionary–biological adaptation (adaptation is shaped by natural selection acting on genetic variation). The history of the former concept dates to the late nineteenth and early twentieth centuries, and has more recently been systemized in the twenty-first century. Approaches to the understanding of phenotypic plasticity and learning behavior have only recently been developed, based on cellular–histological and behavioral–neurobiological techniques as well as traditional molecular biology. New developments of the former concepts in phenotypic plasticity are discussed in bacterial persistence, wing di-/polymorphism with transgenerational effects, polyphenism in social insects, and defense traits for predator avoidance, including molecular biology analyses. We also discuss new studies on the concept of genetic accommodation resulting in evolution of phenotypic plasticity through a transgenerational change in the reaction norm based on a threshold model. Learning behavior can also be understood as physiological phenotypic plasticity, associating with the brain–nervous system, and it drives the accelerated evolutionary change in behavioral response (the Baldwin effect) with memory stock. Furthermore, choice behaviors are widely seen in decision-making of animal foragers. Incorporating flexible phenotypic plasticity and learning behavior into modeling can drastically change dynamical behavior of the system. Unification of biological sciences will be facilitated and integrated, such as behavioral ecology and behavioral neurobiology in the area of learning, and evolutionary ecology and molecular developmental biology in the theme of phenotypic plasticity.

Interspecific competition between an intermediate contest strain of Callosobruchus maculatus and a scramble strain of C. chinensis was investigated on two types of resource beans: the azuki (Vigna anguralis, small seed) and the black-eye bean (Vigna unguiculata, large seed). We conducted both single-generation competition experiments and multiple-generation experiments and then analyzed the system with a discrete version of the Lotka–Volterra competition model. The estimated competition coefficients showed the competitive dominance of the C. maculatus strain over the C. chinensis strain in both types of bean. Zero-growth isocline analysis based on the predicted model showed that an overall dominance of the contest species, C. maculatus, in azuki beans, whereas the outcome of competition was dependent on the initial population size of each species in the black-eye beans. In the multiple-generation experiments, C. maculatus was the successful competitor irrespective of the ratio of the two beans, whereas C. chinensis overcame C. maculatus in one of five replicates with the black-eye beans system. When we incorporated the cost of scramble competition, or density-dependent reduction in body mass into the predicted model, the model alteration decreased the survival region of C. chinensis in phase space on the black-eye bean. Thus, the competitive equations, which do not consider the density-dependent body size reduction, tend to overestimate the advantages of scramble strategists.