Chemical and visual communication during mate searching in rock shrimp

Mate searching in crustaceans depends on different communicational cues, of which chemical and visual cues are most important. Herein we examined the role of chemical and visual communication during mate searching and assessment in the rock shrimp Rhynchocinetes typus. Adult male rock shrimp experience major ontogenetic changes. The terminal molt stages (named "robustus") are dominant and capable of monopolizing females during the mating process. Previous studies had shown that most females preferably mate with robustus males, but how these dominant males and receptive females find each other is uncertain, and is the question we examined herein. In a Y-maze designed to test for the importance of waterborne chemical cues, we observed that females approached the robustus male significantly more often than the typus male. Robustus males, however, were unable to locate receptive females via chemical signals. Using an experimental set-up that allowed testing for the importance of visual cues, we demonstrated that receptive females do not use visual cues to select robustus males, but robustus males use visual cues to find receptive females. Visual cues used by the robustus males were the tumults created by agitated aggregations of subordinate typus males around the receptive females. These results indicate a strong link between sexual communication and the mating system of rock shrimp in which dominant males monopolize receptive females. We found that females and males use different (sex-specific) communicational cues during mate searching and assessment, and that the sexual communication of rock shrimp is similar to that of the American lobster, where females are first attracted to the dominant males by chemical cues emitted by these males. A brief comparison between these two species shows that female behaviors during sexual communication contribute strongly to the outcome of mate searching and assessment.     

Economy of mate attraction in the Cassin's finch

Mate attraction can be costly. Thus, individuals should modulate it according to its probable benefits. Specifically, individuals should modulate mate-attraction efforts based on their need for, the probability of attracting, and the reproductive competence of prospective mates. We tested these predictions by monitoring song output in laboratory-housed male Cassin's finches (Carpodacus cassinii) before, during and after brief female exposure following variable periods of isolation from females. We inferred individual reproductive competence from the product of season and reproductive schedule, the latter estimated from moult progress. Males produced little song in the presence of a female but robustly elevated song output in response to female loss. However, mere absence of a female did not elevate song output in males unaccustomed to female proximity. Furthermore, song output in response to female loss increased with her reproductive competence. We suggest that individuals modulate mate-attraction effort based on the benefits such efforts are likely to yield.     

The importance of mate attraction for intrasexual competition in men and women

Buss (1988a), using an evolutionary perspective, tested hypotheses about intrasexual competition within the context of behavior to attract mates. He found sex differences in intrasexual competition predicted by the goal of mate attraction. We used Buss's methodology while asking subjects about the acts they performed to compete with members of the same sex, rather than about acts for attracting potential mates. Competitive acts were nominated and placed into 26 tactics in Study 1. Study 2 obtained self-reports of male and female act performance frequency. Study 3 replicated Study 2 using observer-reports. Study 4 obtained act effectiveness judgments for men and women. We largely replicated Buss's results despite the focus on intrasexual competition instead of mate attraction. Thus, we provide strong evidence for the importance of mate attraction in intrasexual competition.     

Trimedlure affects mating success and mate attraction in male Mediterranean fruit flies

Male Mediterranean fruit flies, Ceratitis capitata (Wiedemann), are attracted to the synthetic compound trimedlure. Despite the common use of trimedlure in control programs, the underlying basis of male attraction remains unknown. In a series of laboratory trials, we compared the mating success of (treated) males exposed to trimedlure and (control) males not given access to trimedlure. When tested immediately after exposure, treated males mated more frequently than control males. This mating advantage was short-lived, however, and males tested as little as 24 h after exposure had no advantage over control males. When tested immediately after exposure, treated males exhibited higher levels of pheromone calling than control males. Consequently, more females were attracted to treated males than to control males in field tests.     

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.     

Green nesting material has a function in mate attraction in the European starling

The function of fresh green nest material has long been debated. It has been suggested that it reduces the number of ectoparasites in nests and on nestlings (nest protection hypothesis), or is used by males to signal condition and paternal quality (male quality hypothesis) or is used as a sexually selected ornament to attract females (courtship hypothesis). We simultaneously tested these three hypotheses in the European starling, Sturnus vulgaris, in the field. Green material was carried by male starlings only, and mainly during nest building. It was not used to reduce ectoparasites. Males nesting in nestboxes that were experimentally contaminated with ectoparasites did not carry more green nest material than males nesting in control boxes, and experimental removal and addition of green material had no effect on the number of ectoparasites on the nestlings or on their body mass. Furthermore, the amount of green material carried into a nestbox was not associated with male body mass, paternal incubation attendance or nestling food provisioning. There is two-fold experimental evidence that males use green plant material to attract mates. First, removal of greenery resulted in a significantly lower percentage of nestboxes containing a clutch than the control or addition treatment. Second, unpaired male starlings sang more and carried more greenery into a nestbox when a caged female was positioned adjacent to the nestbox than when a caged male or an empty cage was present. Paired males, when subjected to the same experimental design, did not respond.     

Temperature coupling of mate attraction signals and female mate preferences in four populations of Enchenopa treehopper (Hemiptera: Membracidae)

Variation in temperature can affect the expression of a variety of important fitness‐related behaviours, including those involved with mate attraction and selection, with consequences for the coordination of mating across variable environments. We examined how temperature influences the expression of male mating signals and female mate preferences—as well as the relationship between how male signals and female mate preferences change across temperatures (signal–preference temperature coupling)—in Enchenopa binotata treehoppers. These small plant‐feeding insects communicate using plantborne vibrations, and our field surveys indicate they experience significant natural variation in temperature during the mating season. We tested for signal–preference temperature coupling in four populations of E. binotata by manipulating temperature in a controlled laboratory environment. We measured the frequency of male signals—the trait for which females show strongest preference—and female peak preference—the signal frequency most preferred by females—across a range of biologically relevant temperatures (18°C–36°C). We found a strong effect of temperature on both male signals and female preferences, which generated signal–preference temperature coupling within each population. Even in a population in which male signals mismatched female preferences, the temperature coupling reinforces predicted directional selection across all temperatures. Additionally, we found similar thermal sensitivity in signals and preferences across populations even though populations varied in the mean frequency of male signals and female peak preference. Together, these results suggest that temperature variation should not affect the action of sexual selection via female choice, but rather should reinforce stabilizing selection in populations with signal–preference matches, and directional selection in those with signal–preference mismatches. Finally, we do not predict that thermal variation will disrupt the coordination of mating in this species by generating signal–preference mismatches at thermal extremes.     

Attracting mates: Effects of paternal investment on mate attraction strategies

It is proposed here that mate-attraction strategies among females and among males vary facultatively with expectations about paternal investment. The following four hypotheses are argued and tested: 1. Females who expect to find investing mates will be more likely than other females to emphasize their need for investment by suppressing their resources and appearance of competence. 2. Females who expect to find investing mates will try to attract them by acting chaste and emphasizing their fidelity, while females who expect non-investing mates will flaunt their sexuality in order to get pre-reproductive investment from as many males as possible. 3. Males who think it is appropriate to invest heavily in offspring will be more likely than other males to attract mates by emphasizing their ability and willingness to invest. 4. Investing males will be more likely than non-investing males to emphasize chastity and fidelity, whereas non-investing males will flaunt their sexuality and sexual attractiveness to females.These hypotheses were evaluated among undergraduates by administering questionnaires measuring (a) attitudes toward paternal investment and (b) reported tactics used to attract mates. Hypotheses 2–4 were supported by the data, while hypothesis 1 received only partial support. The mixed results for hypothesis 1 indicate that self-deprecating acts and deferential acts may be signaling different things.     

A conserved heptapeptide sequence in the waterborne attractin pheromone stimulates mate attraction in Aplysia

Mate attraction in the marine mollusk Aplysia involves long-distance waterborne chemical signaling via the release of the peptide pheromone attractin during egg laying. Aplysia californica attractin attracts conspecifics, reduces the latency to mating, and stimulates hermaphroditic mating. Four additional members of the Aplysia attractin family have recently been characterized from Aplysia brasiliana, Aplysia fasciata, Aplysia depilans, and Aplysia vaccaria. The five sequences differ significantly, but share six cysteine residues and the strictly conserved sequence Ile30-Glu-Glu-Cys-Lys-Thr-Ser36. Attractin is attractive to geographically and evolutionarily distant species, suggesting that the conserved heptapeptide region may be important for mate attraction. Consistent with this prediction, a synthetic constrained cyclic peptide that contains the conserved heptapeptide sequence is significantly attractive in T-maze bioassays. The attractins are the first family of waterborne peptide pheromones characterized in invertebrates and are unique in that family members are not species-specific pheromonal attractants.     

Male sexual attractiveness predicts differential ovulatory shifts in female extra-pair attraction and male mate retention

Because ancestral women faced trade-offs in choosing mates, they may have evolved to pursue a dual-mating strategy in which they secured investment through one partner and obtained good genes through others. The dual-mating theory predicts that women will display greater interest in extra-pair sex near ovulation, especially if they are mated to a primary male partner who is low in sexual attractiveness. Forty-three normally ovulating women rated their partner's sexual attractiveness and separately reported their own desires and their partner's mate retention behaviors at high and low fertility (confirmed using luteinizing hormone tests). In the high-fertility session relative to the low, women who assessed their partners as being lower in sexual attractiveness reported greater extra-pair desires and more expressed love and attention from their male partners. Women's desire for their own partners did not differ significantly between high and low-fertility sessions.     

Structural aspects of sexual attraction and chemical communication in insects

In the animal kingdom, the relationship between individuals and the environment is often modulated by chemical communication. In the olfaction of insects, small antennary proteins--such as odorant- and pheromone-binding proteins (OBPs and PBPs, respectively) and chemosensory proteins (CSP)--have been proposed to carry their ligand from the air-fluid interface to the olfactory receptors. Binding experiments and recent structural studies of three PBPs, an OBP and a CSP have illustrated their versatility and ability to accommodate ligands of different shapes and chemical structures. The role of these proteins--as simple transporters or receptor triggers--is still a matter of debate, but some recent data seem to support the latter.     

Sex and age differences in preferences and tactics of mate attraction: Analysis of published advertisements

Using the theoretical perspective of evolutionary biology, 13 hypotheses were generated concerning sex and age differences in human mate preferences and tactics of mate attraction. Classified “lonely hearts” advertisements (N = 1,000) from a nationally circulated, fortnightly magazine were content-analyzed. Men, more than women, sought cues to reproductive value (i.e., physical appearance and youth), whereas women, more than men, sought cues revealing an ability to acquire resources (i.e., actual and potential financial security and older men). Women also sought to ascertain a man's willingness to provide resources (in the form of time, emotions, money, and status) in a relationship. Both sexes offered those traits sought by the opposite sex. Men were more promiscuously inclined than women, favoring casual relationships and being more likely to be married, whereas women sought long-term monogamous relationships. These differences support evolutionary predictions based on concepts of sexual selection, parental investment, and reproductive capacities and confirm the use of personal advertisements as a valuable method of research.     

Baffling: a condition-dependent alternative mate attraction strategy using self-made tools in tree crickets

Intense sexual selection in the form of mate choice can facilitate the evolution of different alternative reproductive strategies, which can be condition-dependent. Tree cricket males produce long-distance acoustic signals which are used by conspecific females for mate localization and mate choice. Our study shows that baffling, an acoustic call amplification strategy employed by male tree crickets using self-made tools, is a classic example of a condition-dependent alternative strategy. We show that though most males can baffle, less preferred males, such as smaller and lower-amplitude callers, predominantly use this alternative strategy. Baffling allows these males to increase their call amplitude and advertisement range, which attracts a higher number of females. Baffling also gives these males a mating benefit because females mate for longer durations with them. Our results suggest that the advantage of baffling in terms of gain in the number of sperm cells transferred while mating is primarily limited to less preferred males, thus maintaining the polymorphism of calling strategies in the population. We summarize that baffling is a condition-dependent strategy used by less preferred tree cricket males to obtain mating benefits.     

Behavioral,physiological, and morphological components of dominance and mate attraction in male green iguanas

This study investigated the morphological, physiological, and behavioral components of social dominance important for mate attraction in male green iguanas (Iguana iguana). A group of 9 male and 11 female adult green iguanas was studied in a large semi-natural enclosure during one reproductive season (October–January). Four of the nine males never initiated aggressive encounters; the other five were observed to display aggressively toward each other and were ranked in a linear dominance hierarchy. Head size was the most important factor influencing fighting success. Head size and display frequency were positively correlated with plasma testosterone levels. Dominance rank directly influenced ability to monopolize areas containing resources used by females. The quality of a male's home range, measured as his access to a large basking rock in the enclosure, was related to the proportion of potential mates found within his home range. One male greatly surpassed the others in his ability to defend a home range of high quality and attract potential mates. These data suggest that physiological and morphological factors, through their influence on social behavior, may ultimately affect male reproductive fitness. © 1992 Wiley-Liss, Inc.     

Newly identified water-borne protein pheromones interact with attractin to stimulate mate attraction in Aplysia

The water-borne protein attractin is a potent sex pheromone involved in forming and maintaining mating and egg-laying aggregations in the marine mollusk Aplysia. Binary blends of attractin and either enticin, temptin, or seductin, three other Aplysia protein pheromones, stimulate mate attraction. The four pheromones are thought to act in concert during egg-laying. The new data presented here show that: (1) the water-borne odor of non-laying Aplysia brasiliana further increases the attractiveness of attractin and of eggs in T-maze bioassays. This suggests that individual Aplysia release additional factors that enhance the effects of attractin, enticin, temptin, and seductin during egg-laying; (2) the N-terminal region of enticin aligns well with the conserved epidermal growth factor (EGF)-like domain of mammalian reproductive proteins known as fertilins, which may mediate intercellular adhesion interactions between eggs and sperm; (3) temptin, according to fold recognition servers, may also have an EGF-like fold. Enticin and temptin also have conserved metal binding sequences that may play a role in their signaling behavior. These results suggest that aspects of mammalian egg-sperm interactions (fertilins) may have evolved from pheromonal signaling mechanisms. We also review the structure, expression, localization, release, and behavioral actions of attractin, enticin, temptin, and seductin.     

Phenotypic covariance structure and its divergence for acoustic mate attraction signals among four cricket species

The phenotypic variance-covariance matrix (P) describes the multivariate distribution of a population in phenotypic space, providing direct insight into the appropriateness of measured traits within the context of multicollinearity (i.e., do they describe any significant variance that is independent of other traits), and whether trait covariances restrict the combinations of phenotypes available to selection. Given the importance of P, it is therefore surprising that phenotypic covariances are seldom jointly analyzed and that the dimensionality of P has rarely been investigated in a rigorous statistical framework. Here, we used a repeated measures approach to quantify P separately for populations of four cricket species using seven acoustic signaling traits thought to enhance mate attraction. P was of full or almost full dimensionality in all four species, indicating that all traits conveyed some information that was independent of the other traits, and that phenotypic trait covariances do not constrain the combinations of signaling traits available to selection. P also differed significantly among species, although the dominant axis of phenotypic variation (p(max)) was largely shared among three of the species (Acheta domesticus, Gryllus assimilis, G. texensis), but different in the fourth (G. veletis). In G. veletis and A. domesticus, but not G. assimilis and G. texensis, p(max) was correlated with body size, while p(max) was not correlated with residual mass (a condition measure) in any of the species. This study reveals the importance of jointly analyzing phenotypic traits.     

Social learning and human mate preferences: a potential mechanism for generating and maintaining between-population diversity in attraction

Inspired by studies demonstrating mate-choice copying effects in non-human species, recent studies of attractiveness judgements suggest that social learning also influences human preferences. In the first part of our article, we review evidence for social learning effects on preferences in humans and other animals. In the second part, we present new empirical evidence that social learning not only influences the attractiveness of specific individuals, but can also generalize to judgements of previously unseen individuals possessing similar physical traits. The different conditions represent different populations and, once a preference arises in a population, social learning can lead to the spread of preferences within that population. In the final part of our article, we discuss the theoretical basis for, and possible impact of, biases in social learning whereby individuals may preferentially copy the choices of those with high status or better access to critical information about potential mates. Such biases could mean that the choices of a select few individuals carry the greatest weight, rapidly generating agreement in preferences within a population. Collectively, these issues suggest that social learning mechanisms encourage the spread of preferences for certain traits once they arise within a population and so may explain certain cross-cultural differences.