Aging alters properties of the circadian pacemaker controlling the locomotor activity rhythm in males of Drosophila nasuta

Effects of aging on the circadian rhythm of locomotor activity in males of Drosophila nasuta were investigated. The adult life of males was divided in 1-3 stages according to spontaneous changes in free-running period x in constant darkness (DD): stage 1, days 1-19; stage 2, days 20-36; stage 3, days 37-43. Stage 1 was characterized by a bimodal activity pattern with a short light-induced morning peak and a prolonged evening peak when the flies were entrained to light-dark cycles of 12 hours of light, 12 hours of darkness (LD 12:12). The morning peak had a phase angle difference Ψ_m (Ψ, the time from lights on in LD 12:12 cycles to the onset of morning peak) of about 0.1h, while Ψ_e (Ψ of evening peak) was about 9h at stage 1. The transient morning peak was curtailed at the end of stage 1. At stage 2, the Ψ_e was about 10h, and the activity end was delayed by an addition of about 3h of activity in the scotophase. The changes in W during DD free runs were determined in two groups of flies: flies reared in LD 12:12 and flies reared in DD. In both groups, W increased from about 23h at stage 1 to about 25h at stage 2. Stage 3 was characterized by arrhythmicity associated with highest mean activity level (total number of passes/fly/day) in the entrained and both free-running groups. The mean activity level increased significantly from stage 1 to stage 3 in all three groups of flies.     

Contrasting sexual selection on males and females in a role-reversed swarming dance fly, Rhamphomyia longicauda Loew (Diptera: Empididae)

Although there are several hypotheses for sex-specific ornamentation, few studies have measured selection in both sexes. We compare sexual selection in male and female dance flies, Rhamphomyia longicauda (Diptera: Empididae). Swarming females display size-enhancing abdominal sacs, enlarged wings and decorated tibiae, and compete for nuptial gifts provided by males. Males preferentially approach large females, but the nature of selection and whether it is sex-specific are unknown. We found contrasting sexual selection for mating success on structures shared by males and females. In females, long wings and short tibiae were favoured, whereas males with short wings and long tibiae had a mating advantage. There was no assortative mating. Females occupying potentially advantageous swarm positions were large and, in contrast to selection for mating success, tended to have larger tibiae than those of rivals. We discuss our findings in the context of both the mating biology of dance flies, and the evolution of sexual dimorphism in general.     

Can evolution of sexual dimorphism be triggered by developmental temperatures?

Genetic prerequisites for the evolution of sexual dimorphism, sex-specific heritabilities and low or negative genetic correlations between homologous traits in males and females are rarely found. However, sexual dimorphism is evolving rapidly following environmental change, suggesting that sexual dimorphism and its genetic background could be environmentally sensitive. Yet few studies have explored the sensitivity of the genetic background of sexual dimorphism on environmental variation. In this study, on Drosophila melanogaster, we used a large nested full-sib-half-sib breeding design where families were split into four different developmental temperatures: two constant temperature treatments of 25 and 30 °C and two cycling temperatures with means of 25 and 30 °C, respectively. After emergence, we tested heat shock tolerance of adult flies. We found that sexual dimorphism was strongly affected by temperature during development. Moreover, we found that female heritability was significantly lower in flies developing at hot temperature and more so under hot and cycling temperatures. Interestingly, most of the genetic variation for heat shock tolerance was orthogonal (i.e. noncorrelated) between sexes, allowing independent evolution of heat shock tolerance in males and females. These findings give support to the hypothesis that the evolution of sexual dimorphism can be influenced by the environments experienced during development.     

Circadian consequence of socio-sexual interactions in fruit flies Drosophila melanogaster

In fruit flies Drosophila melanogaster, courtship is an elaborate ritual comprising chasing, dancing and singing by males to lure females for mating. Courtship interactions peak in the night and heterosexual couples display enhanced nighttime activity. What we do not know is if such socio-sexual interactions (SSI) leave long-lasting after-effects on circadian clock(s). Here we report the results of our study aimed at examining the after-effects of SSI (as a result of co-habitation of males and females in groups) between males and females on their circadian locomotor activity rhythm. Males undergo reduction in the evening activity peak and lengthening of circadian period, while females show a decrease in overall activity. Such after-effects, at least in males, require functional circadian clocks during SSI as loss-of-function clock mutants and wild type flies interacting under continuous light (LL), do not display them. Interestingly, males with electrically silenced Pigment Dispersing Factor (PDF)-positive ventral lateral (LNv) clock neurons continue to show SSI mediated reduction in evening activity peak, suggesting that the LNv clock neurons are dispensable for SSI mediated after-effects on locomotor activity rhythm. Such after-effects in females may not be clock-dependent because clock manipulated females with prior exposure to males show decrease in overall activity, more or less similar to rhythmic wild type females. The expression of SSI mediated after-effects requires a functional olfactory system in males because males with compromised olfactory ability do not display them. These results suggest that SSI causes male-specific, long-lasting changes in the circadian clocks of Drosophila, which requires the presence of functional clocks and intact olfactory ability in males.     

Genetic constraints and the evolution of display trait sexual dimorphism by natural and sexual selection

The evolution of sexual dimorphism involves an interaction between sex-specific selection and a breakdown of genetic constraints that arise because the two sexes share a genome. We examined genetic constraints and the effect of sex-specific selection on a suite of sexually dimorphic display traits in Drosophila serrata. Sexual dimorphism varied among nine natural populations covering a substantial portion of the species range. Quantitative genetic analyses showed that intersexual genetic correlations were high because of autosomal genetic variance but that the inclusion of X-linked effects reduced genetic correlations substantially, indicating that sex linkage may be an important mechanism by which intersexual genetic constraints are reduced in this species. We then explored the potential for both natural and sexual selection to influence these traits, using a 12-generation laboratory experiment in which we altered the opportunities for each process as flies adapted to a novel environment. Sexual dimorphism evolved, with natural selection reducing sexual dimorphism, whereas sexual selection tended to increase it overall. To this extent, our results are consistent with the hypothesis that sexual selection favors evolutionary divergence of the sexes. However, sex-specific responses to natural and sexual selection contrasted with the classic model because sexual selection affected females rather than males.     

Parametric and nonparametric entrainment of circadian locomotor rhythm in the Japanese honeybee Apis cerana japonica

The circadian rhythm of locomotor activity in the Japanese honeybee Apis cerana japonica was studied to determine the involvement of parametric and/or nonparametric entrainment. The rhythm was entrained to a skeleton photoperiod in which a 1-h first light pulse was imposed in the morning along with a second light pulse in the evening, as well as to a complete photoperiodic regime (LD 12:12). However, the timing of peak activity relative to the lights-off in the evening in the skeleton photoperiod was earlier than that in the complete photoperiod. A single daily light pulse in the evening entrained the rhythm, whereas a daily light pulse in the morning allowed free-running as in constant darkness. The free-running period (τ) of locomotor activity in constant light became longer as the light intensity increased. A Winfree's type I phase response curve of the locomotor activity rhythm was obtained using a single 1-h light pulse. The results suggest that both parametric and nonparametric entrainment are involved in the circadian rhythm of individual locomotor activity in this honeybee.     

Two oscillators might control the locomotor activity rhythm of the high-altitude Himalayan strain of Drosophila helvetica

The properties of the pacemaker controlling the adult locomotor activity rhythm of the high-altitude Himalayan (haH) strain (Hemkund Sahib, 4121 m above sea level) of Drosophila helvetica are strikingly different from those of the low-altitude Himalayan (laH) strain (Birahi, 1132 m above sea level) of the same species. The haH strain has a unimodal activity pattern with a delayed peak occurring about 4.5 h after lights-on of the entraining light-dark (LD) cycle, while the laH strain has a bimodal activity pattern with the morning and evening peaks. It is rather unusual for a wild type strain of any Drosophila species to have a unimodal activity pattern during entrainment as observed in the haH strain. The single activity peak of the haH strain is regarded as a consequence of delayed morning peak merging with the evening one. Three experiments were performed to test this hypothesis. The first experiment examined whether the single activity peak could be dissociated into two components by LD cycles in which photoperiods varied from 10 to 16 h per 24 h. The haH strain again exhibited a unimodal activity pattern with a delayed peak in 10, 12, and 14 h photoperiods but a bimodal activity pattern in 16 h photoperiod. The laH strain had bimodality in 10 and 12 h photoperiods, unimodality in a 14 h photoperiod, but complete arrhythmicity in a 16 h photoperiod. In the second experiment, the haH flies were transferred from LD 16:8 to LL at 5 lux to confirm whether the bimodality of this strain in LD 16:8 cycles was not the result of masking by the long photoperiod of 16 h. Bimodality of the haH strain persisted in LL too; moreover, the morning component free-ran with period (tau) <24 h, while the evening component free-ran with tau>24 h. The third experiment examined the LL-induced splitting of activity peak of the haH strain. Flies were transferred from LD 12:12 cycles to LL at 0, 1, 5, and 15 lux. The haH strain was rhythmic in LL at 0 and 1 lux with a unimodal activity pattern. It was also rhythmic in LL at 5 lux, but the single activity peak was split into two discrete components; the morning component free-ran with tau<24 h, while the evening component free-ran with tau>24 h. This strain, however, was completely arrhythmic in LL at 15 lux. The laH strain was uniformly arrhythmic in LL at all levels of light intensity. These results suggest that the single but late activity component of the haH strain during entrainment appears to be the consequence of merging the delayed morning peak with the evening one as an adaptation to the environmental conditions at the altitude of origin of this strain, where these flies begin activity in the forenoon owing to non-permissible low temperature in the morning.     

Sex differences in insect immune function: a consequence of diet choice?

Males often have reduced immune function compared to females but the proximate mechanisms underlying this taxonomically widespread pattern are unclear. Because immune function is resource-dependent and sexes may have different nutritional requirements, we hypothesized that sexual dimorphism in immune function may arise from differential nutrient intake (acquisition hypothesis). To test this hypothesis, we examined patterns of phenoloxidase (PO) activity in relation to nutrient consumption in Queensland fruit flies (Q-flies). In the first experiment, flies were allowed to choose their preferred nutrient intake. Compared with males, female Q-flies had higher PO activity, consumed more calories, and preferred a higher protein:carbohydrate (P:C) diet, suggesting that differential acquisition could explain sex differences. In the second experiment, we restricted flies to one of 12 diets varying in protein and carbohydrate concentrations and mapped PO activity for each sex onto a nutritional landscape. Counter to our hypothesis, females had higher PO activity than males at any given level of nutrient intake. Both carbohydrate and protein intake affected PO activity in females but only protein affected PO activity in males. Our results indicate that sex differences in Q-fly immune function are not solely explained by sex differences in nutrient intake, although nutrition does contribute to the magnitude of these sex differences.     

The Role of Sex-specific Plasticity in Shaping Sexual Dimorphism in a Long-lived Vertebrate,the Snapping Turtle <Emphasis Type="Italic">Chelydra serpentina</Emphasis>

Sex-specific plasticity, the differential response that the genome of males and females may have to different environments, is a mechanism that can affect the degree of sexual dimorphism. Two adaptive hypotheses have been proposed to explain how sex-specific plasticity affects the evolution of sexual size dimorphism. The adaptive canalization hypothesis states that the larger sex exhibits lesser plasticity compared to the smaller sex due to strong directional selection for a large body size, which penalizes individuals attaining sub-optimal body sizes. The condition-dependence hypothesis states that the larger sex exhibits greater plasticity than the smaller sex due to strong directional selection for a large body size favoring a greater sensitivity as an opportunistic mechanism for growth enhancement under favorable conditions. While the relationship between sex-specific plasticity and sexual dimorphism has been studied mainly in invertebrates, its role in long-lived vertebrates has received little attention. In this study we tested the predictions derived from these two hypotheses by comparing the plastic responses of body size and shape of males and females of the snapping turtle (Chelydra serpentina) raised under common garden conditions. Body size was plastic, sexually dimorphic, and the plasticity was also sex-specific, with males exhibiting greater body size plasticity relative to females. Because snapping turtle males are larger than females, sexual size dimorphism in this species appears to be driven by an increased plasticity of the larger sex over the smaller sex as predicted by the condition-dependent hypothesis. However, male body size was enhanced under relatively limited resources, in contrast to expectations from this model. Body shape was also plastic and sexually dimorphic, however no sex by environment interaction was found in this case. Instead, plasticity of sexual shape dimorphism seems to evolve in parallel for males and females as both sexes responded similarly to different environments.     

Is sexual body shape dimorphism consistent in aquatic and terrestrial chelonians?

Comparisons between aquatic and terrestrial species provide an opportunity to examine how sex-specific adaptations interact with the environment to influence body shape. In terrestrial female tortoises, selection for fecundity favors the development of a large internal abdominal cavity to accommodate the clutch; in conspecific males, sexual selection favors mobility with large openings in the shell. To examine to what extent such trends apply in aquatic chelonians we compared the body shape of males and females of two aquatic turtles (Chelodina colliei and Mauremys leprosa). In both species, females were larger than males. When controlled for body size, females exhibited a greater relative internal volume and a higher body condition index than males; both traits potentially correlate positively with fecundity. Males were more streamlined (hydrodynamic), and exhibited larger openings in the shell providing more space to move their longer limbs; such traits probably improve mobility and copulation ability (the males chase and grab the female for copulation). Overall, although the specific constraints imposed by terrestrial and aquatic locomotion shape the morphology of chelonians differently (aquatic turtles were flatter, hence more hydrodynamic than terrestrial tortoises), the direction for sexual shape dimorphism remained unaffected. Our main conclusion is that the direction of sexual shape dimorphism is probably more consistent than sexual size dimorphism in the animal kingdom.     

Genetic sexing through the use of Y-linked transgenes

Sterile insect technique (SIT)-based pest control programs rely on the mass release of sterile insects to reduce the wild target population. In many cases, it is desirable to release only males. Sterile females may cause damage, e.g., disease transmission by mosquitoes or crop damage via oviposition by the Mediterranean fruit fly (Medfly). Also, sterile females may decrease the effectiveness of released males by distracting them from seeking out wild females. To eliminate females from the release population, a suitable sexual dimorphism is required. For several pest species, genetic sexing strains have been constructed in which such a dimorphism has been induced by genetics. Classical strains were based on the translocation to the Y chromosome of a selectable marker, which is therefore expressed only in males. Recently, several prototype strains have been constructed using sex-specific expression of markers or conditional lethal genes from autosomal insertions of transgenes. Here, we describe a novel genetic sexing strategy based on the use of Y-linked transgenes expressing fluorescent proteins. We demonstrate the feasibility of this strategy in a major pest species, Ceratitis capitata (Wiedemann), and discuss the advantages and disadvantages relative to other genetic sexing methods and potential applicability to other species.     

The evolution of sexual size dimorphism in the house finch. IV. Population divergence in ontogeny

Differences among taxa in sexual size dimorphism of adults can be produced by changes in distinct developmental processes and thus may reflect different evolutionary histories. Here we examine whether divergence in sexual dimorphism of adults between recently established Montana and Alabama populations of the house finch (Carpodacus mexicanus) can be attributed to population differences in growth of males and females. In both populations, males and females were similar at hatching, but as a result of sex-specific growth attained sexual size dimorphism by the time of independence. Timing and extent of growth varied between the sexes: Females maintained maximum rates of growth for a longer time than males, whereas males had higher initial growth rates and achieved maximum growth earlier and at smaller sizes than females. Ontogeny of sexual dimorphism differed between populations, but in each population, sexual dimorphism in growth parameters and sexual dimorphism at the time of nest leaving were similar to sexual dimorphism of adults. Variation in growth of females contributed more to population divergence than did growth of males. In each population, we found close correspondence between patterns of sexual dimorphism in growth and population divergence in morphology of adults: Traits that were the most sexually dimorphic in growth in each population contributed the most to population divergence in both sexes. We suggest that sex-specific expression of phenotypic and genetic variation throughout the ontogeny of house finches can result in different responses to selection between males and females of the same age, and thus produce fast population divergence in the sexual size dimorphism.     

Sex differences in lizard escape decisions vary with latitude,but not sexual dimorphism

Sexual selection is a powerful evolutionary mechanism that has shaped the physiology, behaviour and morphology of the sexes to the extent that it can reduce viability while promoting traits that enhance reproductive success. Predation is one of the underlying mechanisms accounting for viability costs of sexual displays. Therefore, we should expect that individuals of the two sexes adjust their anti-predator behaviour in response to changes in predation risk. We conducted a meta-analysis of 28 studies (42 species) of sex differences in risk-taking behaviour in lizards and tested whether these differences could be explained by sexual dichromatism, by sexual size dimorphism or by latitude. Latitude was the best predictor of the interspecific heterogeneity in sex-specific behaviour. Males did not change their escape behaviour with latitude, whereas females had increasingly reduced wariness at higher latitudes. We hypothesize that this sex difference in risk-taking behaviour is linked to sex-specific environmental constraints that more strongly affect the reproductive effort of females than males. This novel latitudinal effect on sex-specific anti-predator behaviour has important implications for responses to climate change and for the relative roles of natural and sexual selection in different species.     

Male pioneering as a mating strategy: the case of the beetle Maladera matrida

1. The phenomenon of male pioneering, whereby males are active (daily) before females, is compared with protandry, a phenomenon whereby males are sexually matured before females. The effects of natural selection and sexual selection on both phenomena are discussed. 2. In Maladera matrida Argaman (Coleoptera: Scarabaeidae), males emerged from the soil a few minutes before females, every evening, to feed and mate. 3. Aggregations of males and females are formed at sites of plant damage where males initiated feeding. 4. All copulations were initiated at the beginning of the activity period so males that emerge earlier may increase their probability of finding a mate. 5. Females mate only once in an evening. 6. Males that emerge later may not find receptive females with which to mate. 7. The aggregation of beetles per se did not contribute to the individual mating success because it occurred during and after the copulating had begun. 8. Male pioneering may evolve through sexual selection by means of male–male competition in time to achieve receptive females.     

奇台沙蜥生长过程中的两性异形

研究奇台沙蜥Phrynocephalus grumgrzimailoi头、尾、腋胯距大小在个体发育过程中的变化.成体体长(SVL)无显著的两性差异,两性异形主要表现为雄性个体有较大的头部(头长和头宽)和尾部,雌性具有较大的腋胯距.头、尾、腋胯距大小的两性异形在幼体就已存在,并随个体发育的进行变得更加显著.不同年龄组两性个体头部、尾部、腋胯距随SVL呈异速增长,表现为两性头部的增长速率在个体发育过程中逐渐增大,尾部的增长速率逐渐减慢,腋胯距在雌性蜥蜴中增长速率逐渐增大,在雄性中逐渐变小.     

Sexual Dimorphism of the Jilin Clawed Salamander,Onychodactylus zhangyapingi,(Urodela:Hynobiidae:Onychodactylinae) from Jilin Province,China

Sexual dimorphism in size and shape is common in many organisms, and is a key evolutionary feature. In this study, we analyzed morphometric data of the Jilin clawed salamander Onychodactylus zhangyapingi, an endemic Chinese salamander, to examine sexual size and shape dimorphism. The morphometric data included 14 characteristics of 13 females and 11 males and was analyzed using univariate and multivariate methods. Our results showed that sexual dimorphism occurs not only in body size, but also in body shape. Males have a longer snout-vent length than females, a rarely reported pattern of male-biased sexual size dimorphism. Females have a larger space between the axilla and groin than males, while males have longer and larger tails compared to females. The sexual dimorphism in body size and shape can be explained by existing theories, but there is little data for the mating system, behavior, reproduction, or ecology of O. zhangyapingi, so further studies are required.     

Consequences of sexual size dimorphism on energetics and locomotor performance of Grammostola rosea (Araneae; Teraphosidae)

Most male spiders are smaller than females; during sexual maturity, males change their behaviour, abandoning their web or nest to seek out receptive females actively, whereas females stalk prey near their web or nest and tend not to move away from it. Considering this behavioural difference to be associated with increased locomotor activity at maturity, it may be hypothesized that males will have traits that increase locomotor performance. The present study examines the kinetics and energetics of the movements of the mygalomorph spider Grammostola rosea Walckenaer, a large spider with sexual size dimorphism. It is found that males have a higher maximum aerobic speed, average speed, distance travelled and critical angle of climbing than females, indicating better performance. Males also have lower costs of transport than females. These results support the hypothesis that sexual dimorphism in wandering spiders with active males, which are characterized by smaller body size and longer legs than the larger and more static females, is associated with low transport cost, high velocity and better locomotor performance.     

Sex Ratio and Sexual Size Dimorphism in a Toad-headed Lizard,Phrynocephalus guinanensis

Phrynocephalus guinanensis has sexual dimorphism in abdominal coloration, but its ontogenetic development of sexual size dimorphism(SSD) is unknown. Using mark-recapture data during four days each year from August from 2014 to 2016, we investigated the development of sex ratios, SSD, sex-specific survivorship and growth rates in a population of P. guinanensis. Our results indicated that the sex ratio of males to females was 1:2.8. Males had a lower survival rate(6%) than females(14%) across the age range from hatchling to adult, which supported the discovered female-biased sex ratio potentially associated with the low survival rate of males between hatchlings and juveniles. Male-biased SSD in tail length and head width existed in adults rather than in hatchling or juvenile lizards. The growth rates in body dimensions were undistinguishable between the sexes during the age from hatchling to juvenile, but the growth rate in head length from juvenile to adult was significantly larger in males than females. Average growth rate of all morphological measurements from hatchling to juvenile were larger compared with corresponding measurements from juvenile to adult, but only being significant in tail length, head width, abdomen length in females and snout-vent length in males. We provided a case study to strengthen our understanding of the important life history traits on how a viviparous lizard population can survive and develop their morphology in cold climates.     

PROXIMATE MECHANISMS OF SEXUAL SELECTION IN WOOD FROGS

Observations and several types of field experiments on the mating behavior of wood frogs have revealed the proximate mechanisms for a size-related reproductive advantage in both males and females. For females, larger individuals produce larger clutches; for males, larger individuals can better remain clasped to females when contested by rival males and can better depose males clasped to other females. No results obtained support of the existence of mate choice in either males or females. Males were estimated to be 4.74 times as variable as females in the number of zygotes produced per individual per season; however, much of the variation in male RS resulted from a male-biased sex ratio at the breeding site rather than from sexual selection. After taking sex ratio effects into consideration, males were estimated to be only 1.63 times as variable as females. Patterns of variation in RS in males and females are associated with numerous sex-specific differences in life history and morphology. Life history differences include differential growth rates, ages at sexual maturity, and rates of mortality. Interpretation of how the body size dimorphism (females larger than males) in this species relates to sexual selection is consistent with information on how similar variations in body size influence RS for each sex, and how males and females differ in the functional relationship between body size and RS. Average RS increases more with body size in females than in males. Although body size directly influences RS for females, the possibility exists that, for males, other anatomical features correlated with body size more directly affect RS. Preliminary evidence suggests that sexual selection influences male arm length and that the male body size : RS relationship results as an incidental correlation.