Computational mate choice: theory and empirical evidence

The present review is based on the thesis that mate choice results from information-processing mechanisms governed by computational rules and that, to understand how females choose their mates, we should identify which are the sources of information and how they are used to make decisions. We describe mate choice as a three-step computational process and for each step we present theories and review empirical evidence. The first step is a perceptual process. It describes the acquisition of evidence, that is, how females use multiple cues and signals to assign an attractiveness value to prospective mates (the preference function hypothesis). The second step is a decisional process. It describes the construction of the decision variable (DV), which integrates evidence (private information by direct assessment), priors (public information), and value (perceived utility) of prospective mates into a quantity that is used by a decision rule (DR) to produce a choice. We make the assumption that females are optimal Bayesian decision makers and we derive a formal model of DV that can explain the effects of preference functions, mate copying, social context, and females' state and condition on the patterns of mate choice. The third step of mating decision is a deliberative process that depends on the DRs. We identify two main categories of DRs (absolute and comparative rules), and review the normative models of mate sampling tactics associated to them. We highlight the limits of the normative approach and present a class of computational models (sequential-sampling models) that are based on the assumption that DVs accumulate noisy evidence over time until a decision threshold is reached. These models force us to rethink the dichotomy between comparative and absolute decision rules, between discrimination and recognition, and even between rational and irrational choice. Since they have a robust biological basis, we think they may represent a useful theoretical tool for behavioural ecologist interested in integrating proximate and ultimate causes of mate choice.     

Alternative models of conspecific attraction in flies and crabs

Animals are often attracted to one another when selecting habitats,but little is known about the rules governing conspecific attraction.We use Akaike Information Criterion to evaluate alternativemodels of the effects of conspecifics on individual choice inthe context of habitat selection. One set of models was testedusing data collected on virgin female flies, Drosophila melanogaster,selecting habitats in the laboratory; a second set of modelswas tested using data collected on crabs, Ocypode rotundata,selecting foraging patches in the field. Patterns of space usein the flies were most consistent with models indicating thatindividuals were attracted to other females that selected trapsduring the same hour, as well as to cues left by females thathad entered traps during the previous hour. Results for thecrabs were most consistent with a model which assumes that individualspreferred to join the patch with the most crabs but that theirability to assess the number of crabs in alternative patcheswas constrained by basic psychophysical principles (Weber'slaw). These results provide support for hypotheses about thefunctional significance of conspecific attraction in the fliesand the crabs and illustrate the richness of information aboutconspecific attraction that can be obtained when the same dataset is confronted with alternative models of the ways that animalsrespond to one another when selecting habitats.     

Active choice, passive attraction and the cognitive machinery of acoustic preferences

Female preferences for loud calls are widespread among animals that communicate acoustically, but their ‘strategic’ role is still controversial. According to the ‘passive attraction’ hypothesis, females are more likely to respond to loud calls simply because these calls are more effective at stimulating their acoustic receptors and primary neurons. The ‘active choice’ hypothesis, however, assumes that females use sound intensity as a cue to assess the power and location of sound sources and to respond accordingly. To test which hypothesis could best explain the preference for loud calls of female green toads, Bufo viridis, we carried out multitrial discrimination experiments, in which females were given a choice between two stimuli differing in either the power or the distance of their sources or both. In the laboratory, females could discern differences in intensity caused by differences in power or location of the acoustic sources. This ability increased in field experiments, when the speakers and receivers were further apart than in the laboratory experiments. Phonotactic behaviour did not vary significantly between females, but did within females: in their second trial, females were more likely to approach the closer speaker than in the first trial. These findings suggest that sound intensity plays a more complex role in female mate choice than that recognized by the passive attraction hypothesis. To understand female preferences and, ultimately, mate choice, attention should be focused directly on the computational processes of the female's central nervous system.     

Male and female Trybliographa rapae (Hymenoptera: Figitidae) behavioural responses to food plant, infested host plant and combined volatiles

Many parasitoids use volatiles produced by plants as important cues during their food and host search process. We investigated the attraction of the parasitic wasp Trybliographa rapae Westwood (Hymenoptera: Figitidae) to volatiles emitted from plants infested by the cabbage root fly Delia radicum L. (Diptera: Anthomyiidae), as well as to volatiles from a nectar food plant. Behavioural choice tests showed that male parasitoids were not attracted to any volatiles from plants infested by D. radicum or from nectar plants, while females showed clear attraction to both volatile sources. Young females were more attracted to combined volatiles of host and food plants over those from only the host plant, whereas older females showed no differences in attraction to the two odour sources. This suggests that intercropping attractive flowers with host plants could potentially be used to recruit newly emerged parasitoids from surrounding fields while older parasitoids invest more energy in host location than in additional food search. Volatiles from a whole infested plant were chosen over those emitted from separated above- and below-ground parts from infested plants. It is important to consider the availability of both energy and host resources for parasitoids when designing an eco-compatible management of a vegetable crop system.     

SEXUAL SELECTION AND FEMALE CHOICE IN MOLE CRICKETS (SCAPTERISCUS: GRYLLOTALPIDAE): MODELLING THE EFFECTS OF INTENSITY AND MALE SPACING

ABSTRACT

Attraction of flying mole crickets to individual males calling in an outdoor arena was influenced significantly by the relative intensity of the males calling in the arena. Louder males attracted more individuals than males whose calling songs were less intense. Making simple assumptions about the acoustic output from calling males and about the flight pattern and response of flying females, the differential attraction can be explained by a mathematical model. Computer simulation of the model was used to examine the importance of a male's intensity relative to others and the effect of distance between males on the attraction of females. The model and its relation to active female choice and passive attraction are discussed. The model makes predictions about differences in spacing behaviour of males that maximize attraction relative to other males.     

Female mate-choice behavior and sympatric speciation

Many models have investigated how the process of speciation may occur in sympatry. In these models, individuals are either asexual or mate choice is determined by very simple rules. Females, for example, may be assumed either to compare their phenotype to that of a potential mate, preferring to mate with similar males (phenotype matching), or to possess preference genes that determine which male phenotype they prefer. These rules often do not reflect the mate-choice rules found in empirical studies. In this paper, we compare these two modes of female choice with various types of sexual imprinting. We examine the efficacy of different mate-choice behavior in causing divergence in male traits under simple deterministic one-locus population genetic models as well as under polygenic, individual-based simulations based on the models of Dieckmann and Doebeli (1999). We find that the inheritance mechanism of mate choice can have a large effect on the ease of sympatric speciation. When females imprint on their mothers, the result of the model is similar to phenotype matching, where speciation can occur fairly easily. When females imprint on their fathers or imprint obliquely, speciation becomes considerably less likely. Finally, when females rely on preference genes, male trait evolution occurs easily, but the correlation between trait and preference can be weak, and interpreting these results as speciation may be suspect.     

A cognitive framework for mate choice and species recognition

Mating decisions contribute to both the fitness of individuals and the emergence of evolutionary diversity, yet little is known about their cognitive architecture. We propose a simple model that describes how preferences are translated into decisions and how seemingly disparate patterns of preference can emerge from a single perceptual process. The model proposes that females use error-prone estimates of attractiveness to select mates based on a simple decision rule: choose the most attractive available male that exceeds some minimal criterion. We test the model in the tungara frog, a well-characterized species with an apparent dissociation between mechanisms of mate choice and species recognition. As suggested by our model results, we find that a mate attraction feature alters assessments of species status. Next, we compare female preferences in one-choice and two-choice tests, contexts thought to emphasize species recognition and mate choice, respectively. To do so, we use the model to generate maximum-likelihood estimators of preference strengths from empirical data. We find that a single representation of preferences is sufficient to explain response probabilities in both contexts across a wide range of stimuli. In this species, mate choice and species recognition are accurately and simply summarized by our model. While the findings resolve long-standing anomalies, they also illustrate how models of choice can bridge theoretical and empirical treatments of animal decisions. The data demonstrate a remarkable congruity of perceptual processes across contexts, tasks, and taxa.     

Spectral composition of light sources and insect phototaxis,with an evaluation of existing spectral response models

Artificial illumination attracts insects, but to what extent light attracts insects, depends on the spectral composition of the light. Response models have been developed to predict the attractiveness of artificial light sources. In this study we compared attraction of insects by existing light sources used for streetlights as well as newly developed environment friendly alternatives, and used this data to test the predictive ability of the existing response models. Light sources differed in overall attractiveness to insects and relative attractiveness was dependent on insect order. The attraction patterns predicted by the two models correlated weakly with the number of insects attracted when the only light source rich in UV, a mercury vapour light, was included in the tested spectra. When the mercury vapour light, which is going to be banned in Europe, was not included in the test no correlation was found between predicted and observed attraction patterns. We conclude that currently existing attraction response models are insufficiently sensitive to evaluate new light sources.     

一种测定植物挥发物对棉铃虫引诱作用的无选择性风洞嗅觉仪(英文)

发展了一套测定棉铃虫Helicoverpaarmigera (H櫣ｂｎｅｒ)对植物挥发物的行为反应的生测系统。这套系统由嗅觉仪、风洞和风洞室组成。用这套系统测定了棉铃虫对萎蔫后的枫杨Pterocaryastenoptera叶、加拿大杨Populuscanadensis叶、香椿Toonasisnensis叶、新鲜的芹菜与胡萝卜花挥发物以及苯乙醛的行为反应。结果显示 ,萎蔫的枫杨叶、加拿大杨叶、香椿叶的挥发物均对棉铃虫处女雌蛾表现出显著的引诱作用 ,而对已交配过的雌蛾和雄蛾无明显的引诱作用 ;新鲜的芹菜与胡萝卜花挥发物对棉铃虫两性性成虫均不表现出显著的引诱作用 ;苯乙醛对棉铃虫处女雌、雄蛾有较强的引诱作用。讨论了杨树枝把引诱棉铃虫的原因。     

A NO‐CHOICE WIND TUNNEL OLFACTOMETER SYSTEM FOR EVALUATING THE ATTRACTIVENESS OF PLANT VOLATILES TO ADULT HELICOVERPA ARMIGERA (HÜBNER) (LEPIDOPTERA:NOCTUIDAE) *

Abstract A bioassay system to evaluate adult cotton bollworm (CBW), Helicoverpa armigera, behavioral responses to plant volatiles was described. The system was comprised of a no‐choice olfactometer, a wind tunnel, and a wind tunnel room. The olfactometer was designed to provide for the testing and evaluation of volatiles from a variety of natural and synthetic sources. The behavioral responses in this system demonstrated that the volatiles of wilted leaves of Pterocarya stenoptera, Toona sinensis and Populus canadensis, the main materials of which poplar bundles were made, only could attract virgin females but not mated females and males. The volatiles emitted from fresh carrot and celery flowers did not exhibit significant attractiveness to both sexes of CBW. 0.1% phenylacetaldehyde showed significant attraction for virgin females and males. The biological significance of attraction of volatiles from these plants for CBW was discussed.     

A six-arm olfactometer permitting simultaneous observation of insect attraction and odour trapping

Abstract.  Behavioural assays to study insect attraction to specific odours are tedious, time consuming and often require large numbers of replications. Olfactometer and flight tunnel tests can usually only be conducted with one or two odour sources at a time. Moreover, chemical information on the odour sources has to be obtained in separate analytical studies. An olfactometer was developed in which six odours can be tested simultaneously for their relative attractiveness while during the assays, part of each test odour can be trapped for further analyses. The effectiveness of this six-arm olfactometer was tested by observing the responses of the solitary endoparasitoid Cotesia marginiventris (Cresson) to host-induced odours from young maize plants. For statistical analyses, we used log-linear models were adapted to account for overdispersion and possible positional biases. Female wasps responded extremely well in tests where they were offered a single odour source, as well as in tests with multiple choices. The responses of wasps released in groups were the same as those released individually and it was found that females did not attract or repel each other, but males preferred arms in which females had been released. Dose–response tests with varying numbers of plants or host larvae on plants revealed that the wasps responded in a dose-related manner, thus showing that the system is well suited to measure relative preference. The clear choices of the insects amongst six possibilities provided substantial statistical power. Gas chromatographic analyses of sampled air revealed clean and effective odour trapping, which largely facilitates the comparison of results from behavioural assays with the actual blends of volatiles that were emitted by the various odour sources. Advantages and disadvantages compared to other methods are discussed.     

石斛篓象成虫对不同来源气味的行为反应

【目的】研究不同来源气味对石斛篓象Nassophasis sp.成虫嗅觉选择行为的影响,对于发展新型石斛篓象成虫防治技术具有重要意义。【方法】应用陷阱式嗅觉测定仪测试了石斛篓象成虫对同种个体气味、蛹壳气味、成虫粪便、球花石斛Dendrobium thyrsiflorum受害植株气味的嗅觉反应。【结果】雌成虫气味对雄成虫有引诱作用,对雌成虫没有引诱作用;雄虫气味对雌、雄成虫均无引诱效果。雌、雄蛹壳气味对雌、雄成虫均有显著引诱效果。两性成虫聚集时产生的粪便对雌、雄成虫均有显著引诱作用,雌虫或者雄虫单独放置时产生的粪便对雌、雄成虫均无引诱效果。被雌虫为害的球花石斛植株茎干释放的气味仅对雌虫有引诱作用, 而受雄虫为害后的茎干产生的气味仅对雄成虫有引诱作用。【结论】本研究为进一步分离、 鉴定对该虫有引诱作用的活性物质奠定了基础。     

Female mate choice tactics in a resource-based mating system: field tests of alternative models

Abstract In this study we test theoretical models of female mate choice tactics in natural populations of pine engravers, Ips pini (Say) (Coleoptera: Scolytidae), a species with a resource-based mating system and high search costs. We first develop distinguishing predictions for each of four models of mate choice: random, comparison tactics, and fixed and adjustable thresholds. These predictions relate to commonly collected field data that include the visiting behavior of females and the quality of accepted and rejected mates. Using these types of data, we conclude that pine engravers use an adjustable threshold mate choice tactic because females often accepted the first male encountered, rarely revisited males, visited similar numbers of males in patches of different quality, accepted higher-quality males than those they rejected even on their first encounter with a male in a patch, and had higher acceptance thresholds in high-quality patches than in low-quality patches. This adjustable threshold tactic is consistent with a one-step decision rule and is predicted to occur in species such as pine engravers in which search costs are high and females have information about patch quality before beginning a search in a patch.     

Choosing mates: good genes versus genes that are a good fit

Female choice for male ornamental traits is widely accepted as a mechanism by which females maximize their reproductive success and/or offspring quality. However, there is an increasing empirical literature that shows a fitness benefit of genetic diversity and a tendency for females to use genetic dissimilarity as a criterion for mate choice. This genetic compatibility hypothesis for female mate choice presents a paradox. How can females use both an absolute criterion, such as male ornamentation, and a relative criterion, such as genetic dissimilarity, to choose their mates? Here, we present potential solutions for this dilemma and the empirical evidence supporting them. The interplay between these two contrasting forms of female mate choice presents an exciting empirical and theoretical challenge for evolutionary ecologists.     

Effect of olfactory and visual stimuli on the orientation of the 4th instar larvae of the stem borer Chilo partellus swinhoe (Lepidoptera: Pyralidae).

The orientational responses of 4th instar larvae of Chilo partellus to different sources of stimuli being artificial diet, leaves and stems of maize and sorghum were tested, under free choice and no-choice situations. Larvae were attracted to maize and sorghum in a moderate to high degree dependent on what choice they were given. The orientational preference of the larvae, offered a choice between the visual and the odour sources, depended upon their stimulating capacities which were represented by the percentages of individuals responding to the sources of stimuli. Odour played a greater role than visual stimuli in this close range attraction when the two competed with each other.     

Gaze-dependent evidence accumulation predicts multi-alternative risky choice behaviour

Choices are influenced by gaze allocation during deliberation, so that fixating an alternative longer leads to increased probability of choosing it. Gaze-dependent evidence accumulation provides a parsimonious account of choices, response times and gaze-behaviour in many simple decision scenarios. Here, we test whether this framework can also predict more complex context-dependent patterns of choice in a three-alternative risky choice task, where choices and eye movements were subject to attraction and compromise effects. Choices were best described by a gaze-dependent evidence accumulation model, where subjective values of alternatives are discounted while not fixated. Finally, we performed a systematic search over a large model space, allowing us to evaluate the relative contribution of different forms of gaze-dependence and additional mechanisms previously not considered by gaze-dependent accumulation models. Gaze-dependence remained the most important mechanism, but participants with strong attraction effects employed an additional similarity-dependent inhibition mechanism found in other models of multi-alternative multi-attribute choice.     

Sequential mate choice and sexual isolation in threespine stickleback species

Sequential mate choice strategies predict how females should alter their choosiness based on the availability of attractive males. There are many studies on sequential mate choice within species, but few have asked whether females apply these strategies to interactions between species and how these strategies may affect hybridization. We tested how previous interactions with conspecific and heterospecific males affect mate preference and sexual isolation in two threespine stickleback species (benthics and limnetics: Gasterosteus spp.). Consistent with previous work, we found that within species, stickleback females gauge male attractiveness relative to previously encountered males. If females extend these decision rules between species, we predicted that previous interactions with conspecifics should make heterospecifics less attractive, whereas interactions with heterospecifics should make conspecifics more attractive. However, females found heterospecifics less attractive after prior experience, largely independent of the species of male first encountered. Thus, sequential mate choice strategies are used within but not between species in sticklebacks. Further, learning from prior courtship interactions acts to enhance existing sexual isolation between species.     

The empirical question of thresholds and mechanisms of mate choice

Summary Theoretical discussions concerning how animals might best sample and select mates have suggested that individuals could base decisions either on a sample of mates (sampled-based decisions) or on a threshold of comparison (threshold-based decisions). Recent theoretical work demonstrates that threshold-based mating decisions generate higher expected fitness than sample-based mating decisions when search costs exist. Empirical results from most unmanipulated systems, however, either conclude that females make sample-based decisions or are inconclusive. A few experimental studies designed to detect mating thresholds purport to demonstrate threshold-based choice but an examination of these studies indicates such conclusions were premature. We believe that few examples of threshold-based choice exist because protocols designed to identify mating thresholds were often inconsistent with models of threshold choice. We suggest that future empirical work strive not to document mating thresholdsper se. Rather, future work might best reveal decision rules by manipulating the distribution of quality among potential mates; such manipulations predict uniquely how females using sample-based and threshold-based decision rules should behave.     

Optimism and pessimism in optimised replay

The replay of task-relevant trajectories is known to contribute to memory consolidation and improved task performance. A wide variety of experimental data show that the content of replayed sequences is highly specific and can be modulated by reward as well as other prominent task variables. However, the rules governing the choice of sequences to be replayed still remain poorly understood. One recent theoretical suggestion is that the prioritization of replay experiences in decision-making problems is based on their effect on the choice of action. We show that this implies that subjects should replay sub-optimal actions that they dysfunctionally choose rather than optimal ones, when, by being forgetful, they experience large amounts of uncertainty in their internal models of the world. We use this to account for recent experimental data demonstrating exactly pessimal replay, fitting model parameters to the individual subjects’ choices.     

Is Self‐Grooming by Male Prairie Voles a Predictor of Mate Choice?

Self-grooming by mammals is a form of scent dissemination in which individuals anoint themselves with salivary, anogenital, and other body odours. Self-grooming has been proposed to be a sexually selected trait favoured in reproductive competition and sexual attraction. We tested the hypothesis that females would show a mating preference for males that self-groomed more than a reproductive competitor that groomed less. In mate-choice experiments in which females had a choice of two tethered males, non pair-bonded females did not choose males based on their frequency of self-grooming. In a second experiment in which pair-bonded females in postpartum oestrus had access to their current mate and two strange males, strange males groomed significantly more than pair-bonded mates, yet attained the fewest copulations. Non pair-bonded females and pair-bonded males and females groomed significantly less often than did non pair-bonded males. Self-grooming behaviour was consistent with the sexual attraction hypothesis, but the frequency of self-grooming did not increase a male's mating success. We conclude that the frequency and time spent self grooming are not good predictors of mating success.