Chemical communication and mate attraction in echinostomes

Mate attraction is widespread among animals and appears to facilitate mating and to prevent hybridisation between closely related species. In this study we investigated mate preference between two geographical isolates of Echinostoma caproni (Trematoda, Platyhelminth) and another species of the genus Echinostoma E. sp. Because previous experiments showed a partial reproductive isolation between echinostome isolates, we examined the possibility that such isolation resulted from differential mate attraction. We compared intra-isolate, inter-isolate and interspecific pairings using two in vitro experimental designs. In the first experiment we compared mate attraction of two individuals belonging to or not belonging to the same isolate, while in the second experiment we examined mate choice when individuals were in the presence of individuals from both the same isolate and from a different isolate or a different species. Distances between worms were measured over a period of 90 min. Results from both experiments suggested that mate attraction was similar for intra-isolate, inter-isolate or interspecific combinations. This lack of mate preference in vitro would therefore support an alternative hypothesis of a reproductive isolation through sperm selection.     

The evolution of sex roles in mate searching

Searching for mates is a critical stage in the life cycle of most internally, and many externally, fertilizing species. Males usually invest more in this costly activity than females, but the reasons for this are poorly understood. Previous models have shown that female‐biased parental investment, including anisogamy, does not by itself select for male‐biased mate searching, so it requires additional explanations. Here, we correct and expand upon earlier models, and present two novel hypotheses that might explain the evolution of male‐biased mate searching. The “carry‐over hypothesis” states that females benefit less from searching if the associated costs affect other stages of the life cycle, rather than arising only while searching. It is relevant to the evolution of morphological traits that improve searching efficiency but are also expressed in other contexts. The “mating window hypothesis” states that females benefit less from searching if their life cycle includes intervals during which the exact timing of mating does not matter for the appropriate timing of reproduction (e.g., due to sperm storage or delayed embryo implantation). Such intervals are more likely to exist for females given the general pattern of greater female parental investment. Our models shed new light on classic arguments about sex role evolution.     

Complex mate searching in the satin bowerbird Ptilonorhynchus violaceus

Mate-choice studies typically focus on male traits affecting female mating decisions, but few studies seek to identify the behavioral rules females use when searching for mates. Current models suggest that females may either directly compare a set of males ("pooled comparison") or compare each male to an internal standard ("sequential-search rule") when judging the suitability of potential mates. Models also differ in other specific aspects, such as the predicted number of sampling bouts initiated and the tendency of females to return to males after previous visits. We monitored 63 female satin bowerbirds, Ptilonorhynchus violaceus, during mate sampling to reconstruct their search patterns. We found that females typically sampled several males and returned to the most attractive male for mating: a behavior consistent with the pooled-comparison tactic. Females, however, varied in the number of males sampled; some visited only one male before mating. We found that this variation can be explained by differences among females in the number of mates, the date mate searching is initiated, and long-term experience with males. Further, females were observed to initiate two distinct sampling bouts, with the rejection of most of their potential mates occurring before the start of the second sampling bout. This suggests that the choices of potential mates are narrowed prior to the second sampling bout and that the later visits may function to reconsider preliminary decisions made during the first sampling bout or to resolve decisions concerning the remaining potential mates. Our results indicate that mate searching is a complex process in which females use multiple sampling bouts to find suitable mates and in which several different factors influence their search behavior.     

The role of chemical communication in mate choice

Chemical signals are omnipresent in sexual communication in the vast majority of living organisms. The traditional paradigm was that their main purpose in sexual behaviour was to coordinate mate and species recognition and thus pheromones were conserved in structure and function. In recent years, this view has been challenged by theoretical analyses on the evolution of pheromones and empirical reports of mate choice based on chemical signals. The ability to measure precisely the quantity and quality of chemicals emitted by single individuals has also revealed considerable individual variation in chemical composition and release rates, and there is mounting evidence that prospecting mates respond to this variation. Here, we review the evidence for pheromones as indicators of mate quality and examine the extent of their use in individual mate assessment. We begin by briefly defining the levels of mate choice--species recognition, mate recognition and mate assessment. We then explore the degree to which pheromones satisfy the key criteria necessary for their evolution and maintenance as cues in mate assessment; that is, they should exhibit variation across individuals within a sex and species; they should honestly reflect an individual's quality and thus be costly to produce and/or maintain; they should display relatively high levels of heritability. There is now substantial empirical evidence that pheromones can satisfy all these criteria and, while measurements of the actual metabolic cost of pheromone production remain to some degree lacking, trade-offs between pheromone production and various fitness-related characters such as growth rate, immunocompetence and longevity have been reported for a range of species. In the penultimate section, we outline the growing number of studies where the consequences of chemical-based mate assessment have been investigated, specifically focussing on the reported direct and genetic benefits accrued by the receiver. Finally, we highlight potential areas for future research and in particular emphasise the need for interdisciplinary research that combines exploration of chemical, physiological and behavioural processes to further our understanding of the role of chemical cues in mate assessment.     

Competitive mate searching in male common toads, Bufo bufo

The reproductive behaviour of a population of individually marked toads Bufo bufo was studied at a pond where males outnumbered females by between four and five to one. There was intense competition between the males for mates and only 20·5 % of them bred successfully. Of the successful males, 38·5 % got mates by fighting and displacing other males from the backs of females (takeovers). Larger males enjoyed greater reproductive success because they were stronger and better able to achieve takeovers. When competing for females, some males searched at the spawn site while others searched away from spawn. The numbers searching in the two areas can be predicted by a model which assumes that unpaired males distribute themselves so that there is a spatial ESS, where individuals have equal expectations of finding a female both at and away from the spawning ground.     

Male versus female mate searching in fiddler crabs: a comparative analysis

We present a comparative analysis of mate searching in fiddlercrabs, genus Uca. Several ecological factors determine whichsex will search for mates and how complex male signaling willbe. Female searching is most tightly correlated with matingin male burrows. Female searching is associated with high burrowdensity, small body size, and large soil size. These factors explain variation in a female's need for male-defended incubationsites. Female searching also is correlated with short eyestalks.In species in which females search for mates, males use a morecomplex mate attraction signal than in species in which malessearch.     

Sex roles and mutual mate choice matter during mate sampling

The roles of females and males in mating competition and mate choice have lately proven more variable, between and within species, than previously thought. In nature, mating competition occurs during mate search and is expected to be regulated by the numbers of potential mates and same-sex competitors. Here, we present the first study to test how a temporal change in sex roles affects mating competition and mate choice during mate sampling. Our model system (the marine fish Gobiusculus flavescens) is uniquely suitable because of its change in sex roles, from conventional to reversed, over the breeding season. As predicted from sex role theory, courtship was typically initiated by males and terminated by females early in the breeding season. The opposite pattern was observed late in the season, at which time several females often simultaneously courted the same male. Mate-searching females visited more males early than late in the breeding season. Our study shows that mutual mate choice and mating competition can have profound effects on female and male behavior. Future work needs to consider the dynamic nature of mating competition and mate choice if we aim to fully understand sexual selection in the wild.     

On the cost of searching for a mate in female pied flycatchers Ficedula hypoleuca

Female pied flycatchers, Ficedula hypoleuca, were trapped in one area and released in another a few kilometres away, where the positions of all available nest sites and males were known beforehand. Many of the females moved only a short distance before mating, settling after about 2 (0–6) days in early spring and about 1 day later on. The length of the premating period was negatively correlated with female body weight and prevailing air temperature, indicating that the females were subject to an energy constraint when searching. The premating period was longer for females that settled with an alreadymated male than for those mating with an unmated male. The significance of the results is discussed in relation to the evolution of avian mating systems, and of male behaviour and plumage colour. Competition for a mate, rather than for food, may explain the spacing of males (territoriality).     

Sex-specific timing of mate searching and territory prospecting in the nightingale: nocturnal life of females

Formal models have shown that diel variation in female mate searching is likely to have profound influence on daily signalling routines of males. In studies on acoustic communication, the temporal patterns of the receivers'' signal evaluation should thus be taken into account when investigating the functions of signalling. In bird species in which diel patterns of signalling differ between males singing to defend a territory or to attract a mate, the diel patterns of mate and territory prospecting are suggested to depend on the sex of the prospector. We simulated newly arriving female nightingales (Luscinia megarhynchos) by translocating radio-tagged females to our study site. The mate-searching females prospected the area mostly at night, visiting several singing males. The timing of female prospecting corresponded to the period of the night when the singing activity of unpaired males was higher than that of paired males. In contrast to females, territory searching males have been shown to prospect territories almost exclusively during the dawn chorus. At dawn, both paired and unpaired males sang at high rates, suggesting that in contrast to nocturnal singing, dawn singing is important to announce territory occupancy to prospecting males. In the nightingale, the sex-specific timing of prospecting corresponded to the differential signalling routines of paired and unpaired males. The temporal patterns in the behaviour of signallers and receivers thus appear to be mutually adapted.     

Egg coats of the rock shrimp Rhynchocinetes typus: ultrastructural characterization and their function during fertilization

The aim of the present work was to characterize structurally and ultrastructurally the egg coats of the rock shrimp, Rhynchocinetes typus, and to describe their functional roles during fertilization. Oocytes fixed directly from the ovary, have a total diameter of 549 microm and are covered by a 10- microm-thick transparent envelope. Electron microscope sections (dehydrated) of the egg envelope revealed an electron-dense external coat of 0.4 microm covered by filamentous processes, and a granular inner coat of 4- microm thickness. Oocytes placed for 5 min in seawater had a significantly larger diameter (573 microm), because of the increase in the thickness of the egg coats (32 microm) and the formation of a 16- microm perivitelline space. The diameter of the egg proper was reduced by the same extent as the size of the perivitelline space. All these changes were associated to the loss of the egg fertilizability. SDS-PAGE of isolated and solubilized egg coats with 20% beta-mercaptoethanol or 25 mM dithiothreitol (DTT) showed bands between 58 and 105 kDa and between 44 and 103 kDa, respectively. During normal fertilization, the sperm undergoes a drastic change in shape after first contact with the egg. We observed a similar change when solubilized egg coats were placed with vas deferens sperms. When the solubilized egg coat proteins were ultrafiltrated with a membrane of 10,000 MWCO (pore size) and then assayed for their effect on fertilization, an inhibitory effect of 30%, 41%, and 59% was found when oocytes were incubated with spermatozoa pre-treated with 30, 60, and 120 microg/ml of proteins solubilized with beta-mercaptoethanol. A similar inhibitory effect was found when egg coat proteins solubilized with 25 mM DTT were used. Our results suggest that, in the shrimp, the egg coats play an active role during the morphological changes of the sperm during their passage through them.     

Olfactory conditioning in mate searching by the parasitoid Aphidius ervi (Hymenoptera: Braconidae)

Despite the fact that insect learning capacity has been broadly demonstrated, the role that this process plays during mate searching has been scarcely explored. We studied whether the sexual behaviour of a male parasitic wasp can be conditioned to the odours from two alternative host plant complexes (HPCs) present during its first copulation. The experimental subjects were newly emerged males of the aphid parasitoid, Aphidius ervi, and two alternative HPCs (alfalfa or wheat). In the training protocol, copulation experience corresponded to an unconditioning stimulus and HPC odours to the conditioning stimuli. The initial (just after eclosion) and trained responses were assessed in a glass Y-olfactometer. The results showed that neither alfalfa HPC nor wheat HPC stimuli elicited sexual-related behaviours in initial male responses. Conversely, both HPCs triggered strong attraction and wing fanning courtship behaviour in trained responses when the male was exposed to a female plus HPC during training. In males trained with females plus a given HPC but tested with the alternative HPC in the olfactometer, trained response showed a similar trend to the non-associative treatments. Hence, through learning, the olfactory stimulus context present during copulation could become a predictive cue for further mate searching. These results are discussed in terms of parasitic wasp ecology and host fidelity.     

Chemical communication during foraging in the harvesting ants Messor pergandei and Messor andrei

We combined behavioral analyses in the laboratory and field to investigate chemical communication in the formation of foraging columns in two Nearctic seed harvesting ants, Messor pergandei and Messor andrei. We demonstrate that both species use poison gland secretions to lay recruitment trails. In M. pergandei, the recruitment effect of the poison gland is enhanced by adding pygidial gland secretions. The poison glands of both species contain 1-phenyl ethanol. Minute quantities (3 μl of a 0.1 ppm solution) of 1-phenyl ethanol drawn out along a 40 cm long trail released trail following behavior in M. pergandei, while M. andrei required higher concentrations (0.5–1 ppm). Messor pergandei workers showed weak trail following to 5 ppm trails of the pyrazines 2,5-dimethylpyrazine and 2,3,5-trimethylpyrazine, whereas M. andrei workers showed no behavioral response. Minute quantities of pyrazines were detected in M. pergandei but not in M. andrei poison glands using single ion monitoring gas chromatography–mass spectrometry.     

Natural selection and divergence in mate preference during speciation

Sexual isolation can evolve due to natural selection against hybrids (reinforcement). However, many different forms of hybrid dysfunction, and selective processes that do not involve hybrids, can contribute to the evolution of sexual isolation. Here we review how different selective processes affect the evolution of sexual isolation, describe approaches for distinguishing among them, and assess how they contribute to variation in sexual isolation among populations of Timema cristinae stick-insects. Pairs of allopatric populations of T. cristinae living on different host-plant species exhibit greater sexual isolation than those on the same host, indicating that some sexual isolation has evolved due to host adaptation. Sexual isolation is strongest in regions where populations on different hosts are in geographic contact, a pattern of reproductive character displacement that is indicative of reinforcement. Ecological costs to hybridization do occur but traits under ecological selection (predation) do not co-vary strongly with the probability of between-population mating such that selection on ecological traits is not predicted to produce a strong correlated evolutionary response in mate preference. Moreover, F1 hybrid egg inviability is lacking and the factors contributing to reproductive character displacement require further study. Finally, we show that sexual isolation involves, at least in part, olfactory communication. Our results illustrate how understanding of the evolution of sexual isolation can be enhanced by isolating the roles of diverse ecological and evolutionary processes.     

Characterizing background heterogeneity in visual communication

How is a visual signal effectively transmitted through an environment triggering a response by a perceiver? Experimental and theoretical works in disparate fields, such as sexual selection and plant–animal interactions, have demonstrated selection based on signal conspicuousness. However, to properly describe the conspicuousness of a signal, it is necessary to quantify the spatial and temporal heterogeneity of background coloration of a signaler. We intend to compare the effectiveness of four methods developed to characterize background heterogeneity. To describe the background in a seasonal vegetation, we collected reflectance data in dry and wet seasons of (i) target leaves (those leaves against which a signal is predominantly displayed); (ii) overall leaves (leaves from the most common plant species in the community); (iii) frames in a limited area in space, by measuring the reflectance of all material found in the background; and (iv) background items from the most common species, based on their abundance along transects. We analyzed seasonal color changes in terms of hue, chroma and brightness and described the conspicuousness of fruit coloration according to avian vision. All methods found significant differences in the background coloration between seasons, with higher reflectance values in the dry season. Hue values were higher in the dry season, and chroma values were higher in the wet season. Only the methods most restricted in space (target leaves and frames) recorded seasonal differences in fruit conspicuousness. The transects and frames methods are useful for describing the backgrounds of non-stationary signals (i.e., those of most animals). For plant signals, which are displayed against a fixed background in space, we recommend specific sampling of target leaves. Our results support the importance of measuring the seasonal heterogeneity of the background but also indicate that a monthly sampling design is not necessary to evaluate the conspicuousness of fruit signals.     

Context-dependent visual preferences in starlings and blue tits: mate choice and light environment

There is considerable interest in the role that ultraviolet (UV) cues play in the foraging and mate choice decisions of birds. However, with the exception of the zebra finch, Taeniopygia guttata, it is not yet clear whether ultraviolet preferences are context specific, or whether birds show a general preference for full-avian-spectrum environments (320-700 nm) irrespective of the activity in which they are engaged. We investigated whether European starlings, Sturnus vulgaris, and blue tits, Parus caeruleus, show general (nonresource based) or context-specific preferences for full-spectrum environments. We found that neither species showed a general preference for UV-present (UV+) over UV-deficient (UV−) environments, when those environments contained no resources (experiment 1). Furthermore, neither species showed a UV+ preference when cages contained food, water and perches (starlings; experiment 2) or food, perches and heterospecifics (blue tits; Hunt et al. 1999. Animal Behaviour, 58, 809-815). However, both species did show highly significant preferences for UV+ conditions when viewing potential mates. Such experiments are necessary before one can conclude that particular wavebands have specific relevance to mate choice. In fact, our results suggest that the importance of particular wavelength compositions do indeed vary with behavioural context.     

Chemical communication in heliothine moths

The chemical and temporal features of the sex-pheromone emitted by Heliothis virescens females are encoded by a diverse array of output pathways from the male-specific macroglomerular complex (MGC) in the antennal lobe. Most output neurons (29 out of 32) were activated by antennal stimulation with the principal component of the sex-pheromone blend of this species, (Z)-11-hexadecenal. Six neurons were excited solely by this component, 8 neurons also responded to the second essential blend component, (Z)-9-tetradecenal, and 14 neurons displayed equivalent responses to the two. Many neurons also effectively encoded the onset and duration of the stimulus. In one additional neuron, a prolonged excitatory response (synergism) was evoked only by the blend of the two components, indicating that some MGC neurons function as blend detectors.In contrast to the situation in Helicoverpa zea, none of the MGC neurons in H. virescens responded selectively to (Z)-9-tetradecenal, suggesting that these two noctuid species employ different neural strategies to encode information about their respective pheromone blends.Three MGC-output neurons responded selectively to (Z)-11-hexadecenyl acetate, an odorant released by some sympatric species that disrupts normal upwind flight to pheromones. Thus, changes in the attractant and deterrent chemical signals, as well as the physical features of these odor plumes, are encoded in the MGC across a diverse parallel array of output pathways to the protocerebrum.Abbreviations AL antennal lobe - AN antennal nerve - 16:AL hexadecanal - MGC macroglomerular complex - 14:AL tetradecanal - Z11-16:AL (Z) 11-hexadecenal - Z11-16:AC (Z) 11-hexadecenyl acetate - Z9-14:AL (Z) 9-tetradecenal - Z9-14:FO (Z) 9-tetradecenyl formate     

The evolution of sex differences in mate searching when females benefit: new theory and a comparative test

Sexual selection is thought to have led to searching as a profitable, but risky way of males obtaining mates. While there is great variation in which sex searches, previous theory has not considered search evolution when both males and females benefit from multiple mating. We present new theory and link it with data to bridge this gap. Two different search protocols exist between species in the bush-cricket genus Poecilimon (Orthoptera): females search for calling males, or males search for calling females. Poecilimon males also transfer a costly nuptial food gift to their mates during mating. We relate variations in searching protocols to variation in nuptial gift size among 32 Poecilimon taxa. As predicted, taxa where females search produce significantly larger nuptial gifts than those where males search. Our model and results show that search roles can reverse when multiple mating brings about sufficiently strong material benefits to females.     

Digital video playback and visual communication in lizards

Experimental analyses of dynamic visual signals have to overcome the technical obstacle of reproducing complex motor patterns such as those found in courtship and threat displays. Video playback offers a potential solution to this problem, but it has recently been criticized because of sensory differences between humans and nonhuman animals, which suggest that video stimuli might be perceived as deficient relative to live conspecifics. Quantitative comparisons are therefore necessary to determine whether video sequences reliably evoke natural responses. Male Jacky dragons, Amphibolurus muricatus, compete for territories using complex displays delivered in a rapid stereotyped sequence. We evaluated video playback as a technique for studying this visual signal. Digital video sequences depicting a life-sized displaying male were indistinguishable from live male conspecifics in the rate and structure of aggressive displays evoked. Other measures of social behaviour suggested that video stimuli were more effective in this context. Lizards produced significantly more appeasement displays and had higher rates of substrate licking and locomotor activity in response to video playback than to confined male opponents, which failed to produce aggressive displays. Lizards tracked temporal changes in the display rate of video stimuli and were also sensitive to individual differences in morphology and behaviour between video exemplars. These results show that video stimuli are appropriate for the experimental analysis of Jacky dragon aggressive displays. We compare the potential shortcomings of video playback with those of other techniques and conclude that no approach offers a panacea, but that several have complementary characteristics. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

金龟子化学通讯与信息化学物质

本文综述了金龟子的化学通讯与信息化学物质研究与应用的新进展,重点评述了丽金龟科和鳃金龟科性信息素的鉴定与化学结构。迄今,已有14种金龟子的性信息素被鉴定,还有一些金龟子的聚集信息素和两性引诱剂被报道。对金龟子性信息素、聚集信息素和引诱剂等信息化学物质的应用也作了概述。