The influence of social cues on the reproductive endocrinology of male brown-headed cowbirds: field and laboratory studies

Captive male brown-headed cowbirds exposed to long days exhibit gonadal growth and have elevated plasma testosterone (T) levels. This photoperiodic response is enhanced if males are housed with female cowbirds: Photostimulated males with females increase plasma testosterone levels sooner than do individually housed photostimulated males. Peak plasma T levels are similar in both groups, although peak levels are maintained longer in males housed with females. The gonadal cycle is similarly affected; males in the presence of females have earlier gonadal recrudescence and maintain mature gonads longer than do photostimulated males without females. Plasma corticosterone levels increase in the unpaired males, suggesting that removal of social cues is stressful for these birds. Free-living paired males have significantly higher plasma testosterone levels than do unpaired/unknown males early in the season, when social relationships are being established; the levels are similar thereafter. There is no difference between the two groups in testicular maturation rates; nor do they differ in plasma corticosterone levels at any time of the season. These results suggest that social stimuli are important in modulating the secretion of testosterone in males early in the season when pairing occurs, and possibly late in the season as well, probably to prevent termination of breeding prior to that of females.     

Paternity assurance before and after fertilization by male burying beetles (Nicrophorus quadripunctatus)

Parental care requires a large investment of time and energy. This can reduce future parental survival and opportunities for mating. Because males are usually more uncertain of their parentage with respect to the caring of offspring than are females, the reduction in reproductive success is thought to be greater in males. Therefore, males are under selection to ensure paternity of the offspring for which they care. Males can increase paternity before and after fertilization. Before fertilization, males can increase paternity by increasing their competitive ability for fertilization. After fertilization, males can increase paternity by cannibalizing unrelated offspring. Here, we investigated the stage at which male burying beetles, Nicrophorus quadripunctatus, increase their paternity by evaluating the number of offspring sired by a nursing male in asynchronously hatched broods in relation to hatching time. We found that nursing males assure a very high level of the paternity of hatching offspring. We also found that the paternity of non-nursing and nursing males remained constant across hatching time within a brood, indicating that it is unlikely that filial cannibalism plays a role in increasing the paternity of offspring. We concluded that ensuring paternity before fertilization is more important in increasing the paternity of offspring.     

Re‐Feeding Food‐Deprived Male Meadow Voles Affects the Sperm Allocation of Their Rival Males

An individual's nutritional status affects the manner in which same‐ and opposite‐sex conspecifics respond to that individual, which may affect their fitness. Male meadow voles, Microtus pennsylvanicus, increase their sperm allocation if they encounter the scent mark of an unfamiliar male that is not nutritionally challenged. If, however, the scent mark comes from a male that has been food deprived for 24 h, stud male voles do not increase their sperm allocation. Food‐deprived males may be viewed as being lower quality and a reduced risk of sperm competition by rival males. We hypothesized that stud males in promiscuous mating systems tailor their sperm allocations depending on whether rival males have been food deprived and then re‐fed. We predicted that newly re‐fed males will be considered a strong risk of sperm competition because of the potentially high fitness and survival costs associated with food deprivation in males, and that they will cause stud males to increase their sperm allocation. Our results, however, showed that the recovery period from 24 h of food deprivation was a relatively slow process. It took between 96 and 336 h of re‐feeding male scent donors that were food deprived for 24 h to induce stud males to increase their sperm allocation to levels comparable to when scent donors were not food deprived. Stud male voles may be conserving the amount of sperm allocated until the male scent donors have recovered from food deprivation and subsequent re‐feeding.     

Experimental induction of social instability during early breeding does not alter testosterone levels in male black redstarts,a socially monogamous songbird

Testosterone plays an important role in territorial behavior of many male vertebrates and the Challenge Hypothesis has been suggested to explain differences in testosterone concentrations between males. For socially monogamous birds, the challenge hypothesis predicts that testosterone should increase during male–male interactions. To test this, simulated territorial intrusion (STI) experiments have been conducted, but only about a third of all bird species investigated so far show the expected increase in testosterone. Previous studies have shown that male black redstarts (Phoenicurus ochruros) do not increase testosterone during STIs or short-term male–male challenges. The aim of this study was to evaluate whether black redstarts modulate testosterone in an experimentally induced longer-term unstable social situation. We created social instability by removing males from their territories and compared the behavior and testosterone concentrations of replacement males and neighbors with those of control areas. Testosterone levels did not differ among replacement males, neighbors and control males. Injections with GnRH resulted in elevation of testosterone in all groups, suggesting that all males were capable of increasing testosterone. We found no difference in the behavioral response to STIs between control and replacement males. Furthermore, there was no difference in testosterone levels between replacement males that had expanded their territory and new-coming males. In combination with prior work these data suggest that testosterone is not modulated by male–male interactions in black redstarts and that testosterone plays only a minor role in territorial behavior. We suggest that territorial behavior in species that are territorial throughout most of their annual life-cycle may be decoupled from testosterone.     

Brain and gonadal aromatase activity and steroid hormone levels in female and polymorphic males of the peacock blenny Salaria pavo

In the peacock blenny Salaria pavo large males with well-developed secondary sexual characters establish nests and attract females while small “sneaker” males mimic female sexual displays in order to approach the nests of larger males and parasitically fertilize eggs. These alternative reproductive tactics are sequential, as sneakers irreversibly switch into nesting males. This transition involves major morphologic and behavioral changes and is likely to be mediated by hormones. This study focuses on the role of aromatase, an enzyme that catalyses the conversion of androgens into estrogens, in the regulation of male sexual polymorphism in S. pavo. For this, sex steroid plasma levels and aromatase activity (AA) in gonads, whole brain and brain macroareas were determined in sneakers, transitional males (i.e. sneakers undergoing the transition into nesting males), nesting males and females collected in the field. AA was much higher in ovarian tissue than in testicular tissue and accordingly circulating estradiol levels were highest in females. This supports the view that elevated AA and estradiol levels are associated with the development of a functional ovary. Transitional males are in a non-reproductive phase and had underdeveloped testes when compared with sneakers and nesting males. Testicular AA was approximately 10 times higher in transitional males when compared with sneakers and nesting males, suggesting high AA has a suppressive effect on testicular development. Nesting males had significantly higher plasma levels of both testosterone (T) and 11-ketotestosterone when compared with the other male morphs and previous studies demonstrated that these androgens suppress female-like displays in sneakers. In the brain, AA was highest in macroareas presumably containing hypothalamic nuclei traditionally associated with the regulation of reproductive behaviors. Overall, females presented the highest levels of brain AA. In male morphs AA increased from sneakers, to transitional males, to nesting males in all brain macroareas. These results suggest that the transition into the nesting male tactic is accompanied both by an increase in testicular androgen production and by a higher conversion of androgens into estrogens in the brain. The increase in androgen production is likely to mediate the development of male secondary sexual characters while the increase in brain AA may be related to the behavioral changes associated with tactic transition.     

The role of auditory stimuli in the courtship of Drosophila melanogaster

Simulated courtship song of male Drosophila melanogaster was played to males or females of this species. Upon receiving the song males increase their locomotor activity and start courting each other, whereas females reduce their locomotor activity. In wingless males the locomotor activity difference between the silent control and the experimental sound situation is much larger than in winged males, due to the inactivity of wingless males in the control situation. Males which had been kept singly up to the time of the experiment exhibit higher locomotor and sexual activity than group housed males. A second component of the male courtship song ‘sine song’ is described, together with experiments which investigate the sensory basis of the effect male courtship song has on males.     

Sperm: seminal fluid interactions and the adjustment of sperm quality in relation to female attractiveness

An important predictor of male fitness is the fertilizing efficiency of their ejaculates. Ejaculates are costly to produce and males are predicted to devote greater resources to copulations with reproductively superior females. It is well established that males allocate different numbers of sperm to ejaculates. However, less is known about how males adjust their sperm quality, which has important implications for our understanding of fertilization and the evolution of sexual strategies. Here we test in the fowl, Gallus gallus, whether males adjust their sperm velocity by differentially allocating seminal fluid to copulations with attractive and unattractive females. To disentangle the contributions of sperm and seminal fluid to sperm velocity, we separated and remixed sperm and seminal fluid from ejaculates allocated to females of different attractiveness. We show that dominant males increase the velocity of the sperm they invest in more attractive females by allocating larger ejaculates that contain seminal fluid that increases sperm velocity. Furthermore, we find weak evidence that males also allocate sperm with higher velocity, irrespective of seminal fluid, to more attractive females.     

Courtship Activity of Wandering and Burrow-holding Male Uca arcuata

The male of Uca (Deltuca) arcuata, a vertical claw-waving fiddler in the Indo-Pacific, is used to court females by approaching them from his burrow or while wandering. Differences in the rate of encounters with females between burrow-holding males and burrowless wandering males are found not to be significant. Burrow-holding males less often cause displacement of wandering by females than wandering males do. Wandering of females caused by wandering males occurs as often as wandering of males caused by other males. Thus, burrow-holding males tend not to reduce the number of their potential mates in the neighborhood. Burrow dwelling males are apt to start wandering after decreased encounters with females. Wandering males experience more interactions with other males than burrow-holding males do. Most wandering males that displace burrow owners descend the burrows one or more times after displacing the owners. The extra fighting for temporary burrows is responsible for no increase of the encounter rate with females during wandering.     

To be or not to be? Mating success and survival trade‐offs when switching between alternative reproductive tactics

Hormones underlie the decision of assuming a territorial or a nonterritorial role, with territorial individuals usually having higher hormonal levels than nonterritorial individuals. As a territorial status is linked to higher mating opportunities, it is unclear why animals do not keep high hormonal levels and one explanation is that this would imply survival costs. We have tested this using males of the territorial damselfly Argia emma in the field. We increased juvenile hormone (JH) levels using methoprene in both territorial and nonterritorial males and predicted that: (i) males will keep (the case of territorial males) or become (the case of nonterritorial males) territorial after hormonal increase, and (ii) there will be an increase in mating success for nonterritorial males only and an impaired survival for both male tactics. Hormonally treated males remained or became territorial but had their survival impaired compared with control groups. Also, hormonally treated, ex‐nonterritorial males increased their mating success compared with the other control, nonterritorial males. The reduced survival can be explained proximally by the energy devoted either to the enhanced aggression showed during territory defence or immune function (as detected previously in damselflies). Although nonterritorial males may increase their mating success by switching to a territorial tactic, they are possibly unable to do it naturally as JH is dietary dependent and usually nonterritorial animals are in poorer condition than territorial animals.     

Multiple traumatic insemination events reduce the ability of bed bug females to maintain water balance

To examine how traumatic insemination, a wounding process to females inflicted by males during copulation, reduces the longevity of females of the bedbug, Cimex lectularius, we assessed if multiple bouts of mating impact water relations of females by measuring net transpiration water loss rates. Our studies show that net transpiration rate of females correlates with frequency of mating (small increase after exposure to low numbers of males; large increase after exposure to large numbers of males), and this is reflected by reduced female survivorship for as much as 22 days at 75% RH, 25°C. Water loss occurs up to 28% more rapidly in females after being held with large groups of males. Females that were exposed to males having their paramere removed, females exposed only to other females, and females kept in isolation (unmated) exhibited no reduction in ability to retain water, indicating that traumatic insemination was responsible for the net transpiration rate increase. Mechanical piercing of the female’s abdominal wall leads to increased net transpiration rates for longer periods than puncturing the ectospermalege (regular mating site), implying that inaccurate copulation by males is extremely detrimental to the water balance of females and that the ectospermalege is uniquely modified to seal off more quickly to prevent excess water loss. Mating frequency and the associated increased water loss is considerably reduced by the addition of bed bug alarm pheromone components. Thus, females experience elevated water stress due to traumatic insemination, especially at high levels and when males fail to pierce the ectospermalege, and water loss prevention, likely by more rapid sealing of the wound, is a novel function of the ectopsermalege.     

Rôle of male accessory gland substance in the regulation of blood intake by mosquitoes

The presence of male accessory gland substance during mating is shown to increase the blood intake in the adult female Aedes aegypti and Culex pipiens fatigans. Females mated with males whose accessory glands were surgically removed took significantly less blood than females mated with normal males. However, females mated with males whose seminal vesicles were surgically removed took as much blood as the controls.     

Contrasting phenotypic correlations in food provision of male Tengmalm's owls (Aegolius funereus) in a temporally heterogenous environment

Summary We examined the food provision rate of male Tengmalm's owls,Aegolius funereus, during one 3 year vole cycle consisting of consecutive low, increase and peake vole years. The data were collected in the midnestling period when males provisioned the whole family. In the low vole year, males with a low loading index (g/cm²) of flying area fed their offspring more often than did males with a high loading index, whereas in the peak vole year the opposite trend was evident. Similar relationships were found in the food mass provisioned to the nest. In the increase vole year, male body size had no effect on feeding efficiency. In the peak vole year, when large voles are abundant, heavy males preyed on larger voles than were generally available in their territories, indicating that largeness may increase strike power in hunting attempts. In the low vole year, when breeding is costly due to food scarcity and extensive hunting area, small males are more economical fliers and efficient hunters than large males. The contrasting trends in correlations between male size and feeding efficiency in years of vole abundance versus scarcity suggest that no fixed phenotype may most efficiently cope with variable food supply.     

Changes in the activity budget of cycling female chimpanzees

This study is a preliminary report on the time allocated to various activities by female wild chimpanzees (Pan troglodytes schweinfurthii) during their sexual cycle. Cycling females with maximal tumescence (estrous females) tended to spend more time moving than cycling females with quiescent sexual skin (anestrous females). Although there was no statistically significant decrease in any specific activity that corresponded to the increase in time spent moving, feeding time did decrease in four of the five females. The frequency of approach by females toward males and the frequency of approach by males toward females significantly increased when females were in estrus. Direct aggression by males occurred more frequently toward estrous females than toward anestrous females. The copulation frequency and the frequency of approach to males was not significantly correlated with the increase in time spent moving. There was a high but not significant correlation between the time spent moving and the frequency of direct aggression by males toward females. Mating effort, feeding competition, male aggression, and other possible reasons that might explain the increase in moving time are discussed. Am. J. Primatol. 46:157–166, 1998. © 1998 Wiley-Liss, Inc.     

The Influence of Age on Male Mate‐Searching Behaviour in Thornbug Treehoppers

One prediction from life‐history theory is that males should increase investment in reproductive effort as they age because the opportunity for future reproductive events declines. However, older males may not be able to increase their reproductive effort if condition declines with age. The effect of age‐related changes in condition may be especially important for energetically costly activities such as moving within and between habitat patches while searching for mates. Although such searching is a component of many mating systems, the relationship between age and active mate searching has not been investigated. We investigated whether mate‐searching effort increased with age in the thornbug treehopper, Umbonia crassicornis (Hemiptera: Membracidae). In this species, males search for females using a ‘fly‐call‐walk’ strategy consisting of three phases: (1) flying from one plant to another; (2) walking and signalling while on a plant; and (3) close‐range courtship of encountered females. We measured several aspects of mate‐searching behaviour over the month‐long period of a male’s reproductive lifetime. Over the relevant period of male sexual activity (19–33 d), male condition remained stable. However, older males (25–33 d) did not search more actively than younger males as predicted; instead, younger males (19 d) had greater plant‐to‐plant flight activity and found females faster. Within‐plant walking rates and courtship duration did not differ among age classes. These results suggest that thornbug males may be investing so heavily in mate searching at younger ages that they are unable to increase investment in searching effort when they get older. As a result, older males are likely to be at a competitive disadvantage when active searching is required to locate sparsely distributed females.     

Agonistic Displays and the Benefits of Fighting in the Field Cricket, Gryllus bimaculatus

Fighting is often composed of discrete agonistic displays. Few studies have partitioned fighting behavior into its component agonistic displays and evaluated the relationships between the frequency of the displays and the potential benefits of fighting, particularly mating success. In this study, we quantified the frequency of male field cricket, Gryllus bimaculatus, agonistic displays. The displays were quantified under three social environments which varied in the potential benefits of fighting: males with other males only, males with other males and female scents, and males with other males and females. We found that (1) the presence of females elicited an increase in agonistic displays characteristic of intermediate levels of escalation, (2) female scents did not produce a similar increase in the frequency of agonistic displays, and (3) in the presence of females, the frequency of agonistic displays was positively correlated with mating success. Aggressive stridulation, an energetically low-cost display, was more strongly associated with mating success than were more costly displays. The results are discussed in the context of the evolutionary theory of aggression and in the context of cricket mating systems.     

Adaptive significance of environmental sex determination in an amphipod

Environmental sex determination (ESD) permits adaptive sex choice under patchy environmental conditions, where the environment affects sex-specific fitness and where offspring can predict their likely adult status by monitoring an appropriate environmental cue. For Gammarus duebeni, an amphipod with ESD, it has been proposed that this flexible sex determination system is adaptive because males gain more from large size. Under ESD, young which are born earlier in the season become mostly males and, experiencing longer to grow, are therefore larger at breeding than females which are born later in the season. In order to test the hypothesis that ESD is adaptive for this species we investigated the relationship between size and fitness for both males and females, in a population of G. duebeni known to have ESD. We measured size related pairing success and fecundity, and used these two measures to calculate the relative fitness gains achieved through an increase in size for either sex. The fitness of both males and females increased with size, but males gained more from an increase in size than did females, throughout the breeding season. The data support the adaptive explanation for the evolution and maintenance of ESD in this species.     

The state of gonads of resident sockeye salmon <Emphasis Type="Italic">Oncorhynchus nerka</Emphasis> from the Tolmachev Reservoir

Morphology of gonads and structure of sex cells in females and males of the resident sockeye salmon Oncorhynchus nerka—kokanee—from Tolmachevskoe Reservoir are investigated. Anomalies in structure of sex cells and structure of oocytes are found. Partial or mass resorpbion of cells in the ovaries of males and testes of males is found, leading to sterility of gonads. The relationship between anomalies found in structure of the sex system of kokanee, changes in biological parameters of fish and the increase of the population abundance is discussed.     

Testosterone supplementation in juvenile Psammodromus algirus lizards: consequences for aggressiveness and body growth

In some species, more aggressive individuals are more successful in resource competition. High testosterone level is associated with increased activity and aggressive behavior, and this may have a direct effect on metabolic rate and cause an increase in energy expenditure. Here, I examined the influence of exogenously administered testosterone on aggressiveness and body growth in juvenile Psammodromus algirus male lizards. Juvenile males were given testosterone-filled (experimental) or empty (control) implants. Testosterone produced an increase in aggressiveness and activity in the experimental males. However, despite being more aggressive, experimental males did not acquire larger home ranges than control males. Experimental males also experienced a significant reduction in growth rate over the 2-month period following implantation. Experimental males also were in poorer condition at the completion of the experiment, compared to control males. These results suggest that although an elevated testosterone level may have positive effects on aggressiveness and activity, it also may have negative effects manifested as reduced growth rate and body condition. Received in revised form: 15 June 2001 Electronic Publication     

Observations of the mating behavior and dentition of the round stingray,Urolophus halleri

Synopsis The mating behavior and dentition ofUrolophus halleri, the round stingray was examined. Males frequently bite females during the mating period but most male biting does not result in copulation. In bites that do not lead to copulation, males bite the posterior (or occasionally the medial) portion of the females' disc but females often free themselves from these bites. In bites that precede copulation, males bite the anterior portion of the females' disc and females do not struggle to free themselves. Thus, females may exert some form of choice when they are bitten. Mature males have sexually dimorphic dentition that may aid in holding females. A principal component analysis (PCA) showed that in juvenile males, the relative size of the teeth decrease while the relative thickness of the disc increases as body size enlarges; adult males displayed no clear pattern. In adult females, there is a relative decrease in tooth size and increase in relative disc thickness as body size enlarges. The relative increase in females disc thickness in areas where they are bitten may function to minimize the amount of damage due to non-copulatory biting. There is no indication that biting functions to induce female receptivity but it may allow females and males to acquire information about potential mates. Thus, copulatory biting functions to maintain contact during copulation while the function of non-copulatory biting is less clear.     

Aromatherapy on a large scale: exposing entire adult holding rooms to ginger root oil increases the mating competitiveness of sterile males of the Mediterranean fruit fly in field cage trials

The sterile insect technique (SIT) is widely used in integrated programs against fruit fly pests, particularly the Mediterranean fruit fly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae). Unfortunately, the mass-rearing procedures inherent to the SIT often lead to a reduction in male mating competitiveness. One potential solution involves the pre-release exposure of males to specific attractants. In particular, male exposure to ginger root oil [Zingiber officinale Roscoe (Zingiberaceae); hereafter GRO] has been shown to increase mating success dramatically in field cage trials. Initial studies exposed small groups of males (25 individuals), but more recent work has demonstrated that GRO exposure involving standard storage boxes (containing ≈ 36 000 males) also results in enhanced mating performance. The objective of the present study was to determine whether aromatization of entire trailers, holding ≈ 14 million sterile males from a genetic sexing [temperature sensitive lethal (tsl)] strain, increases male mating success. Independent of the total dose, spatial distribution, or type of dispenser used, sterile males exposed to GRO for a 24-h period displayed greater mating success than non-exposed males in mating cage trials (in which tsl males competed against males from a standard, bisexual strain for females from this same standard strain). Averaged over all experiments, tsl males exposed to GRO obtained 54% of all matings compared to 38% for non-exposed tsl males, an increase of 42%. The implications of these findings for SIT programs against C. capitata are discussed.