Intracranial prolactin perfusion induces incubation behavior in turkey hens

Intracranial perfusion of ovine prolactin (oPrl) via osmotic pump in laying turkey hens caused a sudden onset in incubation behavior, defined as an increase in nest visits. The hens also displayed a gradual decrease in egg laying during the time they were receiving oPrl, another indicator of the onset of incubation. Circulating immunoreactive turkey Prl levels fell during the perfusion period, even though the hens were displaying persistent nesting activity and reduced egg laying. No effects on serum LH were noted. Perfusion of oPrl during the first 14 days of photostimulation delayed the onset of egg laying by several days. No effects on serum Prl or serum LH were noted. It is suggested that incubation behavior is facilitated by central levels of Prl.     

The role of prolactin in reproductive failure associated with heat stress in the domestic turkey

Reproductive failure associated with heat stress is a well-known phenomenon in avian species. Increased prolactin (PRL) levels in response to heat stress have been suggested as a mechanism involved in this reproductive malfunction. To test this hypothesis, laying female turkeys were subjected to 40 degrees C for 12 h during the photo-phase daily or maintained at 24-26 degrees C. Birds in each group received oral treatment with parachlorophenyalanine (PCPA; 50 mg/kg BW/day for 3 days), an inhibitor of serotonin (5-HT) biosynthesis, or immunized against vasoactive intestinal peptide (VIP). Both treatments are known to reduce circulating PRL levels. Nontreated birds were included as controls. In the control group, high ambient temperature terminated egg laying, induced ovarian regression, reduced plasma luteinizing hormone (LH) and ovarian steroids (progesterone, testosterone, estradiol) levels, and increased plasma PRL levels and the incidence of incubation behavior. Pretreatment with PCPA reduced (P < 0.05) heat stress-induced decline in egg production, increase in PRL levels, and expression of incubation behavior. Plasma LH and ovarian steroid levels of heat stressed birds were restored to that of controls by PCPA treatment. As in PCPA-treated birds, VIP immunoneutralization of heat-stressed turkeys reduced (P < 0.05) circulating PRL levels and prevented the expression of incubation behavior. But it did not restore the decline in LH, ovarian steroids, and egg production (P > 0.05). The present findings indicate that the detrimental effect of high temperature on reproductive performance may not be related to the elevated PRL levels in heat-stressed birds but to mechanism(s) that involve 5-HT neurotransmission and the induction of hyperthermia.     

Exogenous prolactin delays photo-induced sexual maturity and suppresses ovariectomy-induced luteinizing hormone secretion in the turkey (Meleagris gallopavo)

A direct effect of prolactin (Prl) on gonadotropin secretion has been suggested but not convincingly demonstrated. The secretion of LH in response to photostimulation (phs) and ovariectomy (ovx) was evaluated in adult female turkeys that had received injections of ovine Prl (124 IU/bird/day for 14 days). In experiment 1, oPrl administration initiated on the day of ovx and phs in reproductively quiescent birds suppressed (p less than 0.05) the elevated LH from a peak level of 11.7 +/- 3.5 ng/ml to 5.1 +/- 0.8 ng/ml in ovx hens. The photo-induced LH increase was unaffected by the oPrl treatment in intact birds. In experiment 2, the oPrl treatment was initiated 7 days before ovx and/or phs. This treatment blunted the peak increase in LH level that follows phs in intact (p less than 0.05) and in ovx (p less than 0.05) hens. In both experiments, onset of lay following phs was delayed (p less than 0.05) in the oPrl-treated groups (29.4 +/- 0.9 days vs. 22.3 +/- 0.9 days; 34.8 +/- 0.5 days vs. 25.0 +/- 0.9 days). In experiment 3, administration of oPrl after ovx of laying hens suppressed the LH rise at essentially all sampling times tested. At the end of the experimental period, 6 of the 7 sham-operated, oPrl-treated laying hens, but none of the sham controls, displayed incubation behavior and had Prl levels of 1020 +/- 370 ng/ml compared to 34 +/- 7 ng/ml in vehicle-treated controls. The results suggest a role for Prl in incubation behavior and LH secretion in the turkey.     

Hormonal induction of incubation behavior in ovariectomized female turkeys (Meleagris gallopavo)

Two experiments were conducted to evaluate the effect of estradiol benzoate (EB), progesterone (P), ovine prolactin (oPrl), or their combinations on temporal patterns of serum luteinizing hormone (LH) and Prl and on nesting behavior in adult ovariectomized female turkeys. Levels of serum LH were initially reduced (p greater than 0.05) by the steroid treatments, while continuation of treatments induced surges of LH to levels comparable to pretreatment levels. Administration of steroid increased (p less than 0.05) levels of serum Prl, which persisted until termination of treatments. Administration of oPrl had no effect on levels of serum LH but blunted the steroid-induced release of Prl. Neither EB, P, nor oPrl treatments alone nor a combination of EB + P elicited nest occupation. Nest occupation was observed after administration of P only in turkeys pretreated with EB. Administration of oPrl maintained and advanced the P-induced nesting to persistent nesting behavior (incubation behavior). Once persistent nesting behavior was established, hormonal treatments were terminated, yet nesting behavior was maintained and serum samples showed increasing levels of Prl and decreasing levels of LH. It is suggested that incubation behavior in the female turkey is facilitated by the combined action of estradiol, P, and Prl.     

Reinitiation of ovulatory cycles in incubating female turkeys by an inhibitor of serotonin synthesis, P-chlorophenylalanine

Nest deprivation of incubating turkeys caused a decrease in serum prolactin (Prl) levels from 1184.5 +/- 116.4 ng/ml to 896.8 +/- 83.0 ng/ml 1 day after initiation of deprivation, with a further decline to 156.5 +/- 111.7 ng/ml at the end of the 22-day experimental period. Serum luteinizing hormone (LH), progesterone and estradiol levels following nest deprivation were similar to those in birds allowed to incubate (controls). Oral administration of p-chlorophenylalanine (PCPA, 50 mg/kg) to incubating turkeys for 3 consecutive days reduced nesting frequency (P less than 0.05) on the 4th day after initiation of treatment and the nesting virtually ceased by the 9th day. Pretreatment Prl was 1655 +/- 210 ng/ml and declined (P less than 0.05) after PCPA administration to a low of 28.6 +/- 2.8 ng/ml. In addition, PCPA caused a sustained rise in serum LH peaking (5.59 +/- 1.09 ng/ml) 3 days after treatment initiation. Contrary to nest deprivation, serum levels of progesterone and estradiol increased (P less than 0.05) as a consequence of PCPA treatment. Seven of 8 PCPA-treated birds later came into lay when their Prl levels and nesting frequency increased again. The results suggest a role for serotonin (5-HT) in incubation behavior, and Prl and LH secretion in turkeys.     

Temperature and life history: experimental heating leads female tree swallows to modulate egg temperature and incubation behaviour

1. Life-history decisions are strongly affected by environmental conditions. In birds, incubation is energetically expensive and affected significantly by ambient temperature. We reduced energetic constraints for female tree swallows (Tachycineta bicolor) by experimentally heating nests during incubation by an average of 6.9 degrees C to test for changes in incubation behaviour. 2. Females in heated boxes (hereafter 'heated females') increased time spent incubating and maintained higher on-bout and off-bout egg temperatures. This indicates that female energetic constraints, not maximizing developmental conditions of offspring, determine incubation investment. Furthermore, this result suggests that embryonic developmental conditions in unmanipulated nests are suboptimal. 3. We found individual variation in how females responded to experimental heating. Early-laying (i.e. higher phenotypic quality) females with heated nests increased egg temperatures and maintained incubation constancy, while later-laying (lower quality) heated females increased incubation constancy. Changes in egg temperature were due to changes in female behaviour and not due directly to increases in internal nest-box temperatures. 4. Behaviour during the incubation period affected hatching asynchrony. Decreased variation in egg temperature led to lower levels of hatching asynchrony, which was also generally lower in heated nests. 5. Our study finds strong support for the prediction that intermittent incubators set their incubation investment at levels dictated by energetic constraints. Furthermore, females incubating in heated boxes allocated conserved energy primarily to increased egg temperature and increased incubation attentiveness. These results indicate that studies investigating the role of energetics in driving reproductive investment in intermittent incubators should consider egg temperature and individual variation more explicitly.     

Plasma prolactin levels in incubating turkey hens during pipping of the eggs and after introduction of poults into the nest

Plasma levels of prolactin (Prl) associated with incubation and maternal behavior were compared in turkey hens allowed to incubate 10 fertile eggs (Group I, n = 9) or 10 infertile eggs (Group II, n = 7) in open nest boxes. At the end of the day that the first egg hatched, all unhatched eggs were removed from Group I hens and each hen was given 10 poults. At the end of the following day, infertile eggs were removed from Group II hens and each hen was given 10 poults. Although pipping of the eggs changed the incubation behavior of Group I hens, it had no effect on plasma Prl. Subsequent hatch of the eggs and/or presence of poults resulted, within 24 h, in a sharp fall in Prl levels, abandonment of the nests, and a shift to maternal behavior. Visual and auditory exposure to Group I poults had no effect on plasma Prl or incubation behavior of Group II hens incubating infertile eggs in adjacent pens. However, within 24 h after the infertile eggs were exchanged for newly hatched poults, Prl levels in Group II hens declined sharply and the hens abandoned the nests and showed maternal behavior similar to that observed in Group I hens. No significant relationships were found in either group between plasma Prl levels and quality of incubation or maternal behavior.     

Heating nest-boxes reveals an energetic contraint on incubation behaviour in great tits,Parus major

According to the ''energetic-bottleneck'' hypothesis, incubation in birds is constrained by the availability of energy. Hence, uniparental incubators are predicted to respond to a change in energy supply by adjusting, positively or negatively, the time spent warming the clutch. Energetic constraints on incubation in the great tit (Parus major) were demonstrated by heating nests, so that the night-time thermostatic component of daily energy expenditure in females was reduced by comparison with a control group. Birds in heated nests increased the time allocated to incubation during the day by 55 minutes, consistent with the predictions of the energetic-bottleneck hypothesis. Daily energy expenditure of all birds was inversely related to night-time ambient temperature, and did not differ between warmed and control birds on mild nights. When temperatures were low, however, escalation of daily costs was less for birds in heated nests. It is suggested that the balance of the energy budget may effect a proximate control on the constancy of incubation, with likely implications for reproductive success.     

Phenotypic responses of hatchlings to constant versus fluctuating incubation temperatures in the multi-banded krait, Bungarus multicintus (Elapidae)

Most studies on egg incubation in reptiles have relied on constant temperature incubation in the laboratory rather than on simulations of thermal regimes in natural nests. The thermal effects on embryos in constant-temperature studies often do not realistically reflect what occurs in nature. Recent studies have increasingly recognized the importance of simulating natural nest temperatures rather than applying constant-temperature regimes. We incubated Bungarus multicintus eggs under three constant and one fluctuating-temperature regimes to evaluate the effects of constant versus fluctuating incubation temperatures on hatching success and hatchling phenotypes. Hatching success did not differ among the four treatments, and incubation temperature did not affect the sexual phenotype of hatchlings. Incubation length decreased as incubation temperature increased, but eggs incubated at fluctuating temperatures did not differ from eggs incubated at constant temperatures with approximately the same mean in incubation length. Of the hatchling phenotypes examined, residual yolk, fat bodies and locomotor performance were more likely affected by incubation temperature. The maximal locomotor speed was fastest in the fluctuating-temperature and 30 degrees C treatments and slowest in the 24 degrees C treatment, with the 27 degrees C treatment in between. The maximal locomotor length was longest in the fluctuating-temperature treatment and shortest in the 24 degrees C and 27 degrees C treatments, with the 30 degrees C treatment in between. Our results show that fluctuating incubation temperatures do not influence hatching success and hatchling size and morphology any differently than constant temperatures with approximately the same mean, but have a positive effect on locomotor performance of hatchlings.     

The evolution of sexual size dimorphism in the house finch. V. Maternal effects

The phenotype of a mother and the environment that she provides might differentially affect the phenotypes of her sons and daughters, leading to change in sexual size dimorphism. Whereas these maternal effects should evolve to accommodate the adaptations of both the maternal and offspring generations, the mechanisms by which this is accomplished are rarely known. In birds, females adjust the onset of incubation (coincident with the first egg or after all eggs are laid) in response to the environment during breeding, and thus, indirectly, determine the duration of offspring growth. In the two house finch (Carpodacus mexicanus) populations that breed at the extremes of the species' distribution (Montana and Alabama), females experience highly distinct climatic conditions during nesting. We show that in close association with these conditions, females adjusted jointly the onset of incubation and the sequence in which they produced male and female eggs and consequently modified the growth of sons and daughters. The onset of incubation in newly breeding females closely tracked ambient temperature in a pattern consistent with the maintenance of egg viability. Because of the very different climates in Montana and Alabama, females in these populations showed the opposite patterns of seasonal change in incubation onset and the opposite sex bias in egg-laying order. In females with breeding experience, incubation onset and sex bias in laying order were closely linked regardless of the climatic variation. In nests in which incubation began with the onset of egg laying, the first-laid eggs were mostly females in Montana, but mostly males in Alabama. Because in both populations, male, but not female, embryos grew faster when exposed to longer incubation, the sex-bias produced highly divergent sizes of male and female juveniles between the populations. Overall, the compensatory interaction between the onset of incubation and the sex-biased laying order achieved a compromise between maternal and offspring adaptations and contributed to rapid morphological divergence in sexual dimorphism between populations of the house finch breeding at the climatic extremes of the species range.     

The wild turkey as a host for Heterakis gallinarum and Histomonas meleagridis.

Freshly embryonated eggs of Heterakis gallinarum gathered from naturally infected domestic turkeys and chickens developed the first 4 weeks essentially as well in young wild turkeys as in domestic poults, but then became progressively retarded and failed in most birds to result in females with fertile eggs. There was no significant difference in the prevalence or progress of infections with Histomonas meleagridis in the two kinds of turkeys, both of which differed from chickens only in that the latter had neither liver involvement nor mortality. In a second test, heterakids hatched from eggs stored 5-6 months at 4 C (comparable to overwintering) sustained very heavy losses in all birds, with greatly accelerated liberations of H. meleagridis, Few worms reached maturity and still fewer produced fertile eggs. In turkeys, and especially in wild turkeys, replacement of infective stages was so poor, that these birds were of no importance in contaminating the soil.     

Metabolic heating and the prediction of sex ratios for green turtles (Chelonia mydas)

We compared incubation temperatures in nests (n=32) of the green turtle (Chelonia mydas) on Ascension Island in relation to sand temperatures of control sites at nest depth. Intrabeach thermal variation was low, whereas interbeach thermal variation was high in both control and nest sites. A marked rise in temperature was recorded in nests from 30% to 40% of the way through the incubation period and attributed to metabolic heating. Over the entire incubation period, metabolic heating accounted for a mean rise in temperature of between 0.07 degrees and 2.86 degrees C within nests. During the middle third of incubation, when sex is thought to be determined, this rise in temperature ranged between 0.07 degrees and 2.61 degrees C. Metabolic heating was related to both the number of eggs laid and the total number of hatchlings/embryos produced in a clutch. For 32 clutches in which temperature was recorded, we estimate that metabolic heating accounted for a rise of up to 30% in the proportion of females produced within different clutches. Previous studies have dismissed any effect of metabolic heating on the sex ratio of marine turtle hatchlings. Our results imply that metabolic heating needs to be considered when estimating green turtle hatchling sex ratios.     

Photoperiod and temperature affect reproductive and nonreproductive functions in male prairie voles (Microtus ochrogaster)

Adult male prairie voles (Microtus ochrogaster) were housed for 10 wk and exposed to long (16L:8D) or short (8L:16D) photoperiods at 21 degrees or 5 degrees C. Maintenance in short day lengths reduced testicular, epididymal, and seminal vesicle mass and also significantly depressed spermatogenic activity. Cold ambient temperature further suppressed gonadal size in voles exposed to short days. Several pelage characteristics were affected by photoperiod, but not by temperature. Increased fur density, fur depth, and length of guard hair and underhair were observed in voles exposed to short days. Intrascapular brown fat and gonadal fat pad mass as well as body mass were significantly less in voles housed in cold temperatures than in voles exposed to warm ambient temperatures; photoperiod did not affect these parameters. Approximately 30% of the male voles exposed to short days maintained their reproductive systems, yet they clearly processed photoperiodic information; all short-day males, regardless of reproductive condition, had comparable winter pelage development. Our results suggest that in prairie voles, photoperiod may be a predictive cue for reproductive function in nature; however, it appears that pelage development is a more obligatory response to photoperiod than is reproduction.     

桂西南褐翅鸦鹃的孵卵行为与节律

2016年3~6月,在广西西南部龙州县弄岗村(22°26′35.20′′~22°30′46.90′′N,106°57′46.35′′~107°03′32.99′′E),通过野外观察和自动温度记录仪相结合的方法对褐翅鸦鹃(Centropus sinensis)的孵卵行为与节律进行了研究。结果表明,1)褐翅鸦鹃边筑巢边产卵,每2 d产1枚卵,卵长径和短径分别为(36.11±0.42)mm和(28.46±0.38)mm,卵重(16.35±0.51)g(n=44枚)。窝卵数3~5枚,孵卵期为(16.75±1.65)d(n=4巢),孵化率为45.45%(n=44枚)。孵卵期与窝卵数之间无显著相关性(r=0.865,P0.05);2)白天双亲共同参与孵卵,夜晚则由其中1只负责。夜间亲鸟的在巢时间从19时左右持续至翌日晨6时左右;3)亲鸟采取离巢次数少和离巢时间长的孵卵策略。亲鸟日活动时间在700 min以上(n=45 d),日离巢次数为(8.82±0.34)次(n=45 d),平均每次离巢持续时间为(52.91±2.35)min(n=397次),每次离巢持续时间与环境温度呈显著负相关关系(r=﹣0.113,P0.05);4)巢内平均孵卵温度为(31.7±0.3)℃(n=4巢),随孵卵天数增加而增加,并与环境温度(最高温r=0.566,最低温r=0.537,平均温r=0.706,P0.01)和日活动时间正相关(r=0.506,P0.01);5)有延迟孵卵行为。延迟孵卵期间夜晚巢内最低温是22.1℃。在桂西南北热带气候环境中,高的环境温度是保障褐翅鸦鹃孵卵成功的主要因素之一。     

Evolution of passerine incubation behavior: influence of food, temperature, and nest predation

Abstract.— Incubation behavior is one component of reproductive effort and thus influences the evolution of life-history strategies. We examined the relative importance of body mass, frequency of mate feeding, food, nest predation, and ambient temperature to explain interspecific variation in incubation behavior (nest attentiveness, on- and off-bout durations, and nest trips per hour) using comparative analyses for North American passerines in which only females incubate. Body mass and frequency of mate feeding explained little variation in incubation behavior. We were also unable to detect any influence of food; diet and foraging strategy explained little interspecific variation in incubation behavior. However, the typical temperature encountered during reproduction explained significant variation in incubation behavior: Species breeding in colder environments take shorter bouts off the nest, which prevents eggs from cooling to temperatures below the physiological zero temperature. These species must compensate for shorter off-bouts by taking more of them (thus shorter on-bouts) to obtain needed energy for incubation. Nest predation also explains significant variation in incubation behavior among passerines: Species that endure high nest predation have evolved an incubation strategy (long on- and off-bouts) that minimizes activity that could attract predators. Nest substrate explained additional variation in incubation behavior (cavity-nesting birds have shorter on-bouts and make more frequent nest trips), presumably because nest predation and/or temperature varies among nest substrates. Thus, nest predation can influence reproductive effort in a way previously not demonstrated–by placing a constraint on parental activity at the nest. Incubating birds face an ecological cost associated with reproductive effort (predation of entire brood) that should be considered in future attempts to explain avian life-history evolution.     

Endocrine changes in the incubating and brooding turkey hen

Turkey hens were allowed to incubate eggs and to hatch and rear young. Plasma prolactin (Prl) levels increased prior to the start of continuous incubation and rose sharply as incubation progressed to reach a peak of 1178.2 +/- 221.8 ng/ml (mean +/- SEM) just before hatching. Prl levels then fell precipitously before the hens left the nest, and returned to preincubation levels (36.8 +/- 3.4 ng/ml) by the time the poults were 2 weeks old. These results show that the high plasma concentrations of Prl found during incubation are not initiated or maintained only by the stimulus of nesting. We suggest that the decline in Prl levels at the end of incubation could be related to the pipping and hatching of eggs, and the consequent shift to maternal behavior. Plasma growth hormone (GH) levels were significantly increased in hens which were brooding poults, but not in hens incubating eggs. An elevenfold, 1-day increase in plasma GH was observed immediately after the hens left the nests. Mean plasma GH levels rose from 12.0 +/- 4.7 ng/ml on the day that the hens left the nests to 133.0 +/- 32.0 ng/ml on the following day, and then declined to 23.1 +/- 9.6 ng/ml after an additional day. There were no significant changes in plasma thyroxine levels during laying, incubation and brooding. Plasma glucose concentration was significantly depressed during incubation.     

光周期和光刺激对雄性灰文鸟行为及生理状况的影响

光周期变化是导致动物行为发生变化的重要因素之一。为揭示不同光周期条件下鸟类对光刺激的生理和行为反应,本文通过比较雄性灰文鸟(Lonchura oryzivora)在经长(16L︰8D)、中(12L︰12D)、短(8L︰16D)3个光周期条件驯化后,再给予72 h的光刺激,研究其行为、血清皮质酮含量和血清蛋白的变化特征。结果表明,光周期改变直接影响雄性灰文鸟的行为。雄性灰文鸟的血清皮质酮含量受光周期和光刺激的影响,中光照周期条件下雄性灰文鸟皮质酮激素对光刺激的响应与长、短光照周期条件下明显不同。不同光周期条件下灰文鸟α-球蛋白区、β-球蛋白区和γ-球蛋白区血清蛋白的成分存在差异。上述结果表明,光周期显著影响雄性灰文鸟对光刺激的生理应激反应方式。     

Effects of ambient temperature on avian incubation behavior

Ambient temperature is commonly thought to influence avian incubation behavior. However, results of empirical studies examining correlationsbetween ambient temperature and bout duration are equivocal.We propose that these equivocal results can be partly explainedby developing a conceptual understanding of how we should expecttemperature to influence incubation. We demonstrate why linearcorrelation analyses across a wide range of temperatures canbe inappropriate based on development of an incubation model for small birds that incorporates how ambient temperature influencesboth embryonic development and adult metabolism. We found supportfor predictions of the model using incubation data from orange-crownedwarblers (Vermivora celata) in Arizona. Both off- and on-boutduration were positively correlated with ambient temperaturebetween 9° and 26°C, but unrelated to ambient temperature<9° and 26-40°C. Bout durations declined as ambienttemperature approached or exceeded 40°C. Incubating orange-crowned warblers appeared to avoid bouts off the nest <7 min andbouts on the nest <20 min. Time of day, duration of theprevious bout, and variation among nests all explained variationin both on- and off-bout duration. Although we found supportfor the general shape of the incubation model, temperature still explained only a small portion of the overall variation in on-and off-bout duration. Results of previous studies were generallyconsistent with the model for off-bout duration; most studiesin colder environments reported positive correlations withtemperature, and the one negative correlation reported was from a hot environment. However, the relationships between on-boutduration and temperature reported in previous studies wereless consistent with our model and our data. Although somediscrepancies could be explained by considering our model,some studies reported negative correlations in cold environments.The effect of ambient temperature on duration of on-bouts probablydiffers among species based on the amount of fat reserves females typically carry during incubation and the extent of male incubationfeeding. Additional studies of the effects of temperature onavian incubation will help improve the general model and ultimatelyaid our understanding of energetic and ecological constraintson avian incubation.     

Pair Formation in California Condors

SYNOPSIS. Pair formation is a prerequisite for reproductionin California condors and exploring the process has been centralto the captive breeding program for this endangered species.Observations show that once birds reach sexual maturity theycan form pairs within a matter of months; even birds who havelost or been separated from their mates will successfully bondwith others. However, some adults who are housed together showlittle indication of pairing. When such birds are placed withother potential mates they frequently successfully pair andproduce young, suggesting that individual differences play arole in pair formation. We recommend that adults who have notbonded within two years be housed with different potential mates. Condors raised together may or may not form pair bonds uponreaching sexual maturity; unfamiliar condors introduced at fouryears of age or older will form pair bonds. In the four caseswhere pairings have been successful the male initiates moreintrapair interactions with aggressive behavior than does thefemale. Unpaired condors housed in groups do show courtshipand the most dominant birds, usually males, are the most activein giving wings-out/head-down displays. Displays may be directedto males or females and may signal dominance and/or interest.Displays may be followed by mounts and one female may crowdin front of the bird to whom a male is displaying, increasingthe chances that she will receive the display and be mounted.Displaying and the response to displaying birds may be mechanismswhich influence mate selection.