Food, vigilance, and sperm: the role of male direct benefits in the evolution of female preference in a polygamous bird

The adaptive significance of female selection of copulationpartners remains unresolved, particularly in polygamous specieswhere males do not provide paternal care. In these species thepossibility that direct benefits other than paternal care mayplay an important role in the evolution of female choice hasreceived little attention. I tested whether direct benefitsare associated with female choice in the polygamous feral fowl,Gallus g. domesticus, where females prefer socially dominantcopulation partners and males do not care for the young butdo provide females with three commodities: food, vigilance,and sperm. I used a combination of empirical and experimentaldata to show that male propensity to offer food and vigilance,but not sperm, was positively associated with male social status,suggesting that the provision of these resources may be costlyand condition dependent in males. Copulation success was correlatedwith male status but not with the number of feedings a femalereceived from a male, indicating that a female preferred dominantpartners that in general provided any female with more food,rather than partners that provided only her with more food,consistent with the idea that females may use male resourceprovisioning as a proximate mechanism to assess male condition.Together, these results indicate that male resources provisioningis (1) tightly linked to male social status, (2) a potentialindicator of male condition and possibly genetic quality, and(3) a potential criterion for females to select dominant partners,thus playing an important role in the evolution of partner choiceeven in polygamous species lacking paternal care.     

Sequential mate encounters: female but not male body size influences female remating behavior

Whether and how individuals choose sequentially among matesis an important but largely neglected aspect in sexual selectionstudies. Here, we explore female remating behavior in the cellarspider Pholcus phalangioides. We focus on body size as one ofthe most important traits involved in mate choice. Large andsmall females (n = 216) were double mated with large or smallmales in all eight possible combinations. All females copulatedwhen virgin, but only 82% accepted a second male. The chanceof a female remating was not significantly predicted by thebody size of the second or first male or by the size differencebetween the two. In contrast, a previous study demonstrateda male size effect in that larger males monopolized femalesuntil egg laying when two males of different sizes were present.We suggest that sequential encounters are more common undernatural conditions than male monopolization of females becauseestimates of concurrent multiple paternity together with observationsin a natural population do not favor mate guarding as the predominantmating strategy in this species. It follows from our study thatthe intensity of sexual selection on male size may be greatlyoverestimated when using a competitive laboratory setting fora species in which females generally encounter mates in a sequentialfashion. Female remating probability was significantly predictedby female size, with large females remating with higher probabilitythan small females. Thus, when mating with large females, malesmay gain higher fertilization success through increased femalefecundity but also face a higher sperm competition risk.     

Hungry females show stronger mating preferences

Female mating decisions that are based on condition-dependenttraits, such as male nutritional state, may be associated witha female's own condition. In the swordtail fish, Xiphophorusbirchmanni, females prefer the chemical cues of well-fed malesto cues of food-deprived males. Here we show that this preferenceis significantly stronger in females when they were experimentallyfood deprived than when they were well fed. Our results suggestthat if females have limited access to food resources, and arethemselves food deprived, they will attend to cues indicatingmale nutritional condition more than when environmental conditionsallow for greater access to food. Furthermore, not only is theslope of the preference function condition dependent but also,in all trials, the latency to respond to the presented stimuliwas shorter in food-deprived females, suggesting that responsivenessto environmental cues is condition dependent as well. Undernatural conditions, females of many species likely experiencevariation in resource availability. Thus, we predict that covariancebetween the strength of female preferences and resource availabilitymay be widespread and may represent a common source of femalepreference variation within and between populations.     

Sex allocation and nestling survival in a dimorphic raptor: does size matter?

Fisher's theory predicts equal sex ratios at the end of parentalcare if the costs and benefits associated with raising eachsex of offspring are equal. In raptors, which display variousdegrees of reversed sexual size dimorphism (RSD; females thelarger sex), sex ratios biased in favor of smaller males areonly infrequently reported. This suggests that offspring ofeach sex may confer different fitness advantages to parents.We examined the relative returns associated with raising eachsex of offspring of the brown falcon Falco berigora, a medium-sizedfalcon exhibiting RSD (males approximately 75% of female bodymass) and subsequent sex ratios. Female nestlings hatched eitherfirst or second did not receive more food nor did they hatchfrom larger eggs or remain dependent on parents for longer periodsthan male offspring from these hatch orders. Together with previousstudies this result indicates that even in markedly dimorphicspecies, the required investment to raise the larger sex islikely to be less than that predicted by body size differencesalone. Moreover, among last-hatched nestlings, both sexes faceda reduced food allocation and suffered a slower growth rateand thus final body size, with a concurrent increased probabilityof mortality. For last-hatched females the reduction in foodallocation was more marked, with complete mortality of all last-hatchedfemale nestlings monitored in this study. Once independent,males of any size but only larger females are likely to be recruitedinto the breeding population. The sex-biased food allocationamong last-hatched offspring favoring males thus reflects therelative returns to parents in raising a small member of eachsex.     

Predation risk affects trade-off between nest guarding and foraging in Seychelles warblers

The fitness costs of egg loss for Seychelles warblers (Acrocephalus sechellensis)on Cousin Island are considerable because warblers have a single-eggclutch and no time to lay a successful replacement clutch. Onthe islands of Cousin and Cousine, with equal densities of Seychellesfodies (Foudia sechellarum), nearly 75% of artificial eggs placedin artificial nests were predated by fodies after 3 days. OnAride Island with no fodies present, loss of artificial eggswas not observed. Female warblers incubate the clutch, and malewarblers guard the clutch when females are absent. Deterrenceof fodies by male warblers is efficient: loss rate of eggs fromunattended warbler nests was seven times as high as from attendednests, and the more nest guarding, the lower the egg loss andthe higher the hatching success. Egg loss is independent ofthe amount of incubation by females. There is no trade-off betweenincubating and foraging by females. Nest guarding competes withforaging by males, and this trade-off has a more pronounced effecton egg loss when food availability is low. The transfer of breeding pairsfrom Cousin to either Cousine with egg-predating fodies or toAride without fodies allowed us to experimentally investigatethe presumed trade-off between nest guarding and foraging. OnCousine, individual males spent the same amount of time nestguarding and foraging as on Cousin, and egg loss was similarand inversely related to time spent nest guarding as on Cousin.Males that guarded their clutch on Cousin did not guard theclutch on Aride but allocated significantly more time to foragingand gained better body condition. Loss of warbler eggs on Aridewas not observed. Time allocation to incubating and foragingby individual females before and after both translocations remainedthe same.     

Mice with an autism-associated R451C mutation in neuroligin-3 show a cautious but accurate response style in touchscreen attention tasks

One of the earliest identifiable features of autism spectrum disorder (ASD) is altered attention. Mice expressing the ASD-associated R451C mutation in synaptic adhesion protein neuroligin-3 (NL3) exhibit impaired reciprocal social interactions and repetitive and restrictive behaviours. The role of this mutation in attentional abnormalities has not been established. We assessed attention in male NL3^R451C mice using two well-established tasks in touchscreen chambers. In the 5-choice serial reaction task, rodents were trained to attend to light stimuli that appear in any one of five locations. While no differences between NL3^R451C and WT mice were seen in accuracy or omissions, slower response times and quicker reward collection latencies were seen across all training and probe trials. In the rodent continuous-performance test, animals were required to discriminate, and identify a visual target pattern over multiple distractor stimuli. NL3^R451C mice displayed enhanced ability to attend to stimuli when task-load was low during training and baseline but lost this advantage when difficulty was increased by altering task parameters in probe trials. NL3^R451C mice made less responses to the distractor stimuli, exhibiting lower false alarm rates during all training stages and in probe trials. Slower response times and quicker reward latencies were consistently seen in NL3^R451C mice in the rCPT. Slower response times are a major cognitive phenotype reported in ASD patients and are indicative of slower processing speed. Enhanced attention has been shown in a subset of ASD patients and we have demonstrated this phenotype also exists in the NL3^R451C mouse model.     

Food affects the potential reproductive rates of sand goby females but not of males

The interspawning interval of female sand gobies, Pomatoschistusminutus, a batch-spawning fish with paternal care, was significantlyshorter when the fish were fed daily than when they were fedevery fourth day. The incubation time of males was not affectedby feeding, nor was the interbrood interval Males have an equalor higher potential reproductive rate than females. As femalesreproduce more slowly when food is scarce than when it is abundant,and males do not, the difference between the sexes in potentialreproductive rate increases when there is food shortage. Becauseof this difference, both male bias in operational sex ratioand intensity in male-male competition for mates are predictedto increase as food availability decreases. Furthermore, a tradeoffbetween current and future reproduction is demonstrated to operateonly when resources are limited, because the correlation betweenegg number of the first and second clutch was positive amonghigh-food females but negative among low-food females. The numberof eggs per female clutch did, however, not differ between treatmentsin first or second dutch. I conclude that operational sex ratioand sexual selection are expected to vary within and betweensand goby populations in accordance with prey availability     

Food availability and parasite infection influence mating tactics in guppies (Poecilia reticulata)

Despite the important effects of diet and parasite infectionon male reproductive behavior, few studies have simultaneouslyaddressed their influence on intrasexual selection (male–malecompetition). We examined the synergistic effects of 2 naturallyvarying environmental factors, lifetime food intake and infection,with the monogenean parasite Gyrodactylus turnbulli on the matingtactics and foraging behavior of male guppies (Poecilia reticulata).We allowed fish to interact directly with each other duringobservations and found that unparasitized males won more intermalecontests, courted females more frequently, and received positiveresponses to courtship displays more frequently than males thathad been infected. Infected males devoted more time to foragingand less time to courtship and competition than uninfected males,suggesting that they were energetically limited and could notincrease reproductive effort despite their reduced expectedlifespan. This interpretation was supported by the observationthat greater food intake ameliorated the negative effects ofparasite infection on courtship effort. Our results have bearingon how natural variation in food availability and parasite prevalenceinfluence geographic variation in reproductive behavior.     

Condition-dependent signaling affects male sexual attractiveness in field crickets, Gryllus campestris

The condition dependence of sexually selected traits is an importantassumption of sexual selection theory. Several laboratory studieshave documented a positive relationship between food availability,body condition, and sexual display. However, these studies mightnot reflect the resource allocation between body maintenance,reserves, and the sexually selected trait under natural conditions.Further, the effect of condition-dependent signaling on femalemate choice has hardly been investigated experimentally in thefield. We therefore investigated the effect of food availabilityon body condition, calling behavior, and sexual attractivenessof male field crickets, Gryllus campestris, under field conditions.Food availability was manipulated for individual males by supplementingfood in a confined area close to the burrow. Food-supplementedmales showed a significant increase in body condition, whereasthe opposite was found in the control males. Males receivingextra food called more frequently, whereas the calling-songcharacteristics were not affected by the treatment. Further,food-supplemented males attracted more females than did controlmales, and their higher attractiveness was partly explainedby their superior calling rate. Our study thus indicates condition-dependentsignaling as an important determinant of the sexual attractivenessof males to females under natural condition.     

Male fruit flies learn to avoid interspecific courtship

Experimental data suggest, and theoretical models typicallyassume, that males of many fruit flies (Drosophila spp) areat least partially indiscriminate while searching for mates,and that it is mostly the females who exert selective mate choice,which can lead to incipient speciation. Evidence on learningby male D. melanogaster in the context of courtship, however,raises the possibility that the initially indiscriminate malesbecome more selective with experience. I tested this possibilityby comparing the courtship behavior of male D. melanogasterexperienced at courting females of the closely related species,D. simulans, and inexperienced males. I found that comparedwith the inexperienced males, the males experienced with courtingD. simulans females showed significantly lower courtship towardfemale D. simulans. Both male treatments, however, showed virtuallyidentical courtship durations with female D. melanogaster. Theseresults indicate that male fruit flies adaptively refine theircourtship behavior with experience and suggest that the malescontribute more to assortative mating and incipient speciationthan is commonly assumed.     

Adaptations of Polygynous Breeding in the Bobolink, Dolichonyx oryzivorus

In some species of polygynous songbirds, males assist in feedingnestlings of their first (primary) female but ignore offspringof subsequent (secondary) mates. A number of adaptations importantin minimizing the disadvantages of a secondary status in femaleBobolinks (Dolichonyx oryzivorus) have been discovered. In Bobolinks, clutches of secondary females are significantlysmaller than those of primary females, thereby adjusting broodsize to correspond better with food delivery capabilities ofsecondary hens. Commencement of incubation with deposition ofthe penultimate egg allows additional brood size adjustment,for the resulting hatching asynchronism promotes efficient cullingof the late-hatching chick in times of food shortage, withoutendangering the entire brood. Secondary females differ fromprimary pairs in food exploitation techniques by capitalizingon insects found close to the nest and by showing less discriminationin food selection. Such behavior permits compensation in therate of food delivery trips to secondary nestlings so as toequal the tempo maintained by both members of the primary pair,yet results in quantitative disparities in food brought to primaryand secondary nestlings. Flexibility in male feeding responses,resulting in partial male assistance at secondary nests in situationswhere secondary brood size is exceptionally large, is also importantin helping to maximize the reproductive performance of birdsengaging in this mating strategy.     

Perseverance and food sharing among closely affiliated female chimpanzees

Chimpanzees (Pan troglodytes) have been frequently observed to share food with one another, with numerous hypotheses proposed to explain why. These often focus on reciprocity exchanges for social benefits (e.g., food for grooming, food for sex, affiliation, kinship, and dominance rank) as well as sharing based on begging and deterring harassment. Although previous studies have shown that each of these hypotheses has a viable basis, they have only examined situations in which males have preferential access to food whereby females are required to obtain the food from males. For example, studies on male chimpanzee food sharing take advantage of successful crop-raids and/or acquisitions of meat from hunting, situations that only leave females access to food controlled by male food possessors. This begs the question how and with whom might a female chimpanzee in sole possession of a high-quality food item choose to share? In two large captive groups of chimpanzees, we examined each of the hypotheses with female food possessors of a high-quality food item and compared these data to a previous study examining food transfers from male chimpanzees. Our results show that alpha females shared significantly more with closely affiliated females displaying perseverance, while kinship and dominance rank had no effect. This positive interaction between long-term affiliation and perseverance shows that individuals with whom the female possessor was significantly affiliated received more food while persevering more than those with neutral or avoidant relationships towards her. Furthermore, females with avoidant relationships persevered far less than others, suggesting that this strategy is not equally available to all individuals. In comparison to the mixed-sex trials, females chose to co-feed with other females more than was observed when the alpha male was sharing food. This research indicates that male and female chimpanzees (as possessors of a desired food item) share food in ways influenced by different factors and strategies.     

Reaction norms of size and age at maturity of Pomacea canaliculata (Gastropoda: Ampullariidae) under a gradient of food deprivation

Pomacea canaliculata, an apple snail native to South America,has become a serious pest of aquatic crops and a promoter ofecosystem changes in natural wetlands worldwide. Its successas an invader has been attributed to its great phenotypic plasticityin life-history traits. Our aims were to determine the reactionnorms of size and age at maturity under a gradient of food deprivation.Full sibling experimental snails were reared in isolation fromhatching and maintained until maturity under seven differentlevels of relative food deprivation based on size-specific ingestionrates. To detect the onset of sexual activity of experimentalsnails, fully mature virgin snails reared in the laboratorywere used as consorts. The reaction norms for age and size atmaturity of P. canaliculata showed marked sexual dimorphism.Shell length was the main component of variation in the malereaction norms for both copulation and egg-laying by femaleconsorts, whereas age was the main component of variation forfemales. Irrespective of the intensity of food deprivation,males mature at the same age at the expense of size, since sizeis apparently irrelevant in the access to females and male fitnesscan be maximized through fast maturation. In contrast, a minimumsize is required for females to reach maturity, perhaps as aresult of their higher reproductive costs. The highly dimorphicreaction norms lead to an increasing lag between male and femalematurity as deprivation increases; in temperate regions, malesborn early in the reproductive season would mature in the sameseason irrespective of food availability, while most femaleswould have to overwinter before attaining sexual maturity inunproductive habitats or those dominated by unpalatable macrophytes.The great life-history plasticity reported in invaded areascould be a heritage from populations in the native range. (Received 10 August 2007; accepted 30 July 2008)     

棉铃虫成虫对性信息素的电生理和行为反应研究

通过EAG和风洞实验,研究了棉铃虫雌雄成虫对性信息素组分和诱芯(Z-11-16Ald∶Z-9-16Ald=97∶3)的电生理反应。其中棉铃虫雌、雄蛾对诱芯的平均EAG反应测定值分别为1.06mV和4.32mV,分别高出对照(无性信息素空白诱芯)0.67mV和0.366mV,差异均达到极显著水平(雌蛾：t＝25.020, P≤0.01;雄蛾：t＝44.269,P≤0.01);棉铃虫雌蛾对性信息素组分(Z-11-16-Ald和Z-9.16Ald)的EAG反应值随浓度增加而增加;雄蛾在被剪除触角后与雌蛾不能正常交配,而雌蛾在被剪除触角后仍有40%的交配率,比正常雌雄蛾的交配率(70%)有所下降;在风洞实验中,雄蛾没有顺风远离诱源的飞行行为,趋向诱源的比率为81.8%,与对照有显著差异。研究表明性信息素组分对棉铃虫的交配活动有明显的影响。     

Variation in courtship rate in the fiddler crab Uca annulipes: is it related to male attractiveness?

We investigated among-male variation in courtship waving inthe fiddler crab Uca annulipes. Wave rate is positively correlatedwith both male carapace size and relative claw size (controlledfor body size), and relative claw size is positively correlatedwith an index of body condition. An experimental reduction inthe availability of food decreased male wave rate. These datasuggest that some of the variation in wave rate among malesis due to variation in male condition combined with energeticcosts to waving (differential costs). However, we also foundthat the correlation between male size and wave rate decreasedover the semilunar cycle. Later in the cycle, smaller malesincrease their wave rate relative to that of larger males. Previouswork has shown that females are more likely to accept a smallermale as a mate later in the cycle. We suggest that smaller malesinvest disproportionately more in courtship later in the cyclebecause the potential benefits are greater due to their increasedattractiveness to females (differential benefits). Alternativeexplanations for the observed temporal trend are also discussed.     

Sexual selection for large male size in a polyandrous butterfly: the effect of body size on male versus female reproductive success in Pieris napi

Sexual selection theory predicts that the larger sex shouldbe that for which fitness increases at the faster rate withsize. In butterflies, as in most invertebrates, females areusually the larger sex, but previous comparative analysis hasshown that relative male size increases with female polyandryamong butterflies. In agreement with this pattern, males arelarger than females in the strongly polyandrous green-veinedwhite butterfly, Pieris napi L., and in this article we assessthe size dependence of reproductive success in both sexes. Inan experiment where virgin males and females were released inthe field, we found no strong association between size and malemating success. However, laboratory experiments showed thatthere was a strong correlation between size and the ejaculatethat the male delivered to the female at mating and that largeejaculates delayed female remating for a longer time comparedto small ejaculates. Moreover, female P. napi utilize male-derivednutrients received at mating to increase their fecundity. Hence,large males sire more offspring both by way of donating morenutrients to female egg production and by way of delaying femaleremating (given that the last male to mate with the female willfather most of the offspring). Laboratory experiments showedthat the association between size and fecundity was low, ornonexistent, among P. napi females allowed to mate only once.However, weak size dependence was found for polyandrous females.We hypothesize that size dependence of female fecundity maybe especially weak among polyandrous butterflies because a fundamentalsource of variation in fecundity relates to their ability tofind nutrient giving males, an ability which may be unrelatedto female size. According to this hypothesis there is a causalassociation between weak size dependence of female fecundityand polyandry, and a strong size dependence of male reproductivesuccess that may underlie the comparative pattern of positivecorrelation between relative male size and polyandry.     

Paternal Care in Canids

Paternal care has never been reported as absent in any canidspecies, and some form of care has been seen in 18 of the 36species in the family. Food provisioning, active defense ofthe young, and protecting young by remaining at the den as thefemale forages appear to be the commonest forms of male care.In addition males may groom, retrieve, play and rest with young.Male canids are rarely involved in den selection or construction.The effect on the fitness of the young of indirect forms ofmale care such as provisioning the female and territory defenseare hard to assess. Quantitative studies of male provisioningin seven species offer few generalizations. In two species (Canisaureus, C mesomelas) females provided more food to the youngthan males; in one species (Alopex lagopus) the pair contributedequally to feeding young, and in four species (Canis lupus,Vulpes vulpes, Chrysocyon brachyurus, and Lycaon pictus), malesprovided more food than females. Much more data are required,particularly from field studies, before patterns of variationcan be interpreted.     

Mate choice based on static versus dynamic secondary sexual traits in the dark-eyed junco

Some secondary sexual traits (SSTs) such as structural characteristicsare semi-permanent or static, while others, such as courtshipdisplay, are more labile or dynamic. In this paper we reportresults from two experiments designed to test the relative attractivenessto female dark-eyed juncos (Junco hyemalis, Passeriformes, Aves)of a relatively static plumage trait, the amount of white inthe tail, and a relatively dynamic behavioral trait, courtshipintensity. The experiments derived from a study showing that femalejuncos prefer males that court more vigorously. We asked whether femalesalso base their preferences on plumage traits and how they respond whenpresented with a choice between attractive traits that are eitherstatic (plumage) or dynamic (courtship) in nature. In the firstexperiment we presented males to females in paired mate-choicetrials and found that males enhanced with more white in theirtails were more attractive to females than controls with unenhancedtails. Females spent more time with enhanced males and directedmore sexual displays toward them. In the second experiment we testedwhether females preferred males with enhanced tails (a staticSST) or males with enhanced hormone-mediated courtship behavior(a dynamic SST). In this experiment females did not demonstratea consensus preference for either the static or the dynamictrait. Instead, some females preferred the male whose courtshipperformance was enhanced with testosterone, while others preferredthe male with an enhanced tail. We conclude that both kindsof traits are important in junco mate choice, but that somefemales apparently weigh static traits more heavily than dynamicones, while other females use opposite weightings.     

Reduced Inhibition of Return to Food Images in Obese Individuals

Previous research has shown that obese individuals may be biased towards attending to food over non-food information, and this bias may contribute to the development and/or maintenance of obesity. The present study sought to extend our understanding of maladaptive attentional processing in this population by investigating whether obese individuals have difficulty in disengaging attention from food compared with non-food images, relative to normal-weight controls. To address this question, we measured inhibition of return (IOR) in an attentional cueing task. The participants were 29 obese and 35 normal-weight satiated females without eating disorders. The obese group displayed less IOR to food images than the normal-weight group, while there was no difference in IOR between the groups for non-food images. This suggests that obese females have greater difficulty disengaging attention from food than normal-weight females. Our findings provide a new focus for studies investigating maintenance factors in obesity and are discussed in relation to a theory of incentive-sensitisation.     

Sexual selection in a moth: effect of symmetry on male mating success in the wild

Sexual selection is generally caused by female choice and male–malecompetition. In female choice process, female preference isfavored indirectly and/or directly by sexual selection. In indirectselection, females expressing the preference might gain indirectgenetic benefits. In direct selection, females expressing thepreference might gain direct benefits or avoid male-imposedcosts. The white-tailed zygaenid moth Elcysma westwoodii ismonandrous, and males often gather around a female to mate withher, suggesting a high opportunity for sexual selection on maletraits. We quantified phenotypic selection on male morphologyin this species in the field. The morphological characters analyzedincluded body weight, antenna length, forewing length, hindwing length, hind wing tail length, genital clasper length,and the fluctuating asymmetry (FA) of these bilateral traits.In E. westwoodii, selection favored males with more symmetricgenital claspers, as well as longer and more symmetrical hindwings and antennae. Negative correlations between FA and sizewere also detected in the clasper and the antenna. Our resultssuggest that FAs of male traits, in particular the genital clasper,may have indirect and direct influences on mating success. Duringa copulatory attempt, an E. westwoodii male will try to graspthe female's abdominal tip with his claspers but often failto do so because of the female's reluctance to mate. The femaleabdominal tips are smooth and strongly sclerotized and couldthus be difficult for males to grasp. We hypothesize that moresymmetrical male claspers are more efficient in overcoming femalereluctance.