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Trade up polygyny and breeding synchrony in avian populations.

The advent of the molecular techniques used to assign paternity has focused attention on the differences between the social and the genetic mating systems of sexual species. In particular, the interrelations between breeding synchrony-the degree to which the fertility periods of individual females in a population overlap and the degree of extra pair paternity (EPP) in that population, has became a subject of a lively debate. Investigation of the subject can be facilitated by examining the criteria that females use in choosing extra pair partners. These preferences constitute a continuum ranging between two extremes. At one end, there are situations wherein all the females in a population exhibit a preference for males with particular phenotypic markers, and females mated to males lacking such "quality" markers seek extra pair fertilizations from males that do -trade up polygyny. At the other extreme, there are situations wherein females seek to maximize the total number of male partners, rather than secure fertilization by males of particular type -indiscriminate polygyny. Previously, we used game theoretical methods to model the interrelations between breeding synchrony and EPP in the context of indiscriminate polygyny. Here we present an analogous investigation in the context of trade up polygyny. Our results for the two cases, which delimit the range of the possible behavior, are similar. That is, we see that it is the pursuit of extra pair fertilizations opportunities that determines breeding synchrony of populations, rather than the vice versa as has been previously suggested.

Animals↗

Prevention of incubation behavior expression in turkey hens by active immunization against prolactin.

The consequences of active immunization against prolactin on expression of incubation, reproductive performance and hormonal profiles were evaluated in turkey hens. Hens were injected weekly for 4 wk starting 8 wk before being submitted to a stimulatory photoperiod and 3 times thereafter at intervals of 4 to 5 wk. The hens were injected i.d. with 0.5 mL of a mixture diluted half in Freund's adjuvant. The mixture was prediluted in .9% saline and contained 100 micrograms of a fusion protein (GST-tPRL), GST, oPRL or vehicle. The results indicate that active immunizations with GST-tPRL or oPRL both induce production of specific prolactin antibodies. The onset of egg production was unaffected but higher egg production was observed for the GST-tPRL immunized hens. No GST-tPRL immunized hens expressed incubation behavior, whereas 20 to 30% of hens in the other experimental groups did so. Apparent hyperprolactinemia was detected by RIA for the GST-tPRL immunized groups starting before photostimulation and lasting until Week 10 of egg production but not in other groups. No significant differences were observed in either plasma LH or estradiol concentrations of immunized and nonimmunized turkey hens. In conclusion, both GST-tPRL or oPRL induced the production of antibodies against prolactin in turkey hens. However, only active immunization using GST-tPRL induced higher antibody titers as well as full prevention of incubation behavior expression. Such a pharmacological approach is of great practical interest, although its uses need to be carefully evaluated under commercial conditions.

Animals↗

The role of prolactin in the regulation of clutch size and onset of incubation behavior in the American kestrel.

In most bird species, the timing of incubation onset may influence the degree of hatching asynchrony, which, together with variation in clutch size, affects reproductive success. In some domesticated species that usually show no hatching asynchrony, plasma prolactin concentrations in females rise with the onset of incubation and the end of laying, and this rise enhances incubation behavior and may terminate laying. To investigate whether a rise in prolactin during laying is involved in the regulation of clutch size and incubation onset in a species with hatching asynchrony, we measured plasma concentrations of immunoreactive prolactin (ir-prolactin) in laying American kestrels, Falco sparverius, and quantified clutch size and incubation behavior. In a separate study, we administered one of three concentrations of ovine prolactin (o-prolactin) via osmotic pumps implanted in females when egg 2 of a clutch was laid. ir-Prolactin concentrations during laying were higher in small than in large clutches and increased in parallel with the development of incubation behavior. o-Prolactin treatment enhanced incubation behavior, but did not affect clutch size, possibly because the manipulation was performed after clutch size had already been determined. Consistent with studies on domesticated species that show synchronous hatching, our results indicate that rising prolactin during laying enhances the expression of incubation behavior in a species that shows hatching asynchrony. Further studies are necessary to determine whether the relationship between prolactin and clutch size in the American kestrel is one of causation or of mere association.

Animals↗

Timing of incubation bouts by ring doves (Streptopelia risoria).

Ring dove parents share in incubation and brooding such that the male sits for a block of time toward the middle of the day and the female sits the rest of the time. In order to evaluate the role of the sexes in determining nest exchange, approaches to the nest, the timing of incubation behavior, and the topography of interactions at the nest site were continuously monitored. Most nest exchanges (87%) were initiated by the nonsitting mate. There was less variability associated with the onset of sitting by the female than by the male. The male entered the nest area repeatedly from the time of lights-on until nest exchange several hours later. In contrast, the female rarely entered the nest area when the male was sitting, and when she did approach, nest exchange usually ensued. When the male's entrances to the nest area were prevented until the usual time of nest exchange, the timing of the subsequent sitting bout and nest exchange was not altered, which indicates that male visits are not a necessary prerequisite for nest exchange. Each pair developed a cooperative interaction as incubation progressed. When mates were exchanged among physiologically synchronized pairs, a serious disruption of sitting resulted. It is concluded that behavioral synchrony between the mates in the form of each pair's bond is as necessary as physiological synchrony in maintaining intermate cooperation in parental behavior of ring doves.

Animals↗

Hormonal mediation of reproductive behavior.

Reproductive capability requires synchronization of both endocrine and behavioral components of reproduction. The classic view of reproductive behavior was that there was simply direct stimulation of a behavioral response by the appropriate gonadal steroid. However, it has become clear with recent developments in the field of behavioral endocrinology that the relationship of hormonal and behavioral processes is complex. In addition to the complexity of the mechanisms involved, other factors, both social and environmental, influence both the endocrine and behavioral responses. This paper provides an overview of selected issues within the field of behavioral endocrinology which deal with mechanisms of hormonal induction of behavior at various stages of the life cycle and with factors that interact with these processes.

Aging↗

External factors affecting the duration of broody behavior in the ring dove (Streptopelia risoria).

The object of this study was to investigate factors which determine the duration of sitting in ring doves. This normally lasts ca. 19 days from laying, with very small variation. The period is made up of 15 days incubating eggs and 4-5 days brooding squabs. The duration of sitting is unaffected by substituting fresh or sterile eggs, and can only be slightly influenced by substituting foster squabs or new eggs just before or after hatching. The sitting period appears to be predetermined to run for 19 days following laying. The maintenance of sitting, however, requires the presence of the nest and eggs, and can be drastically varied by keeping subjects in continuous daylight or on a day of 6 hr light:6 hr dark (6 L:6 D). Birds kept on ahemeral cycles (11 L: 10 D; 13 L:14 D) displayed significantly different periods of incubation on infertile eggs and recycling, indicative of an endogenous circadian basis to the timing mechanism.

Animals↗

The significance of the nest-building of the nine-spined stickleback, Pungitius pungitius.

The reproductive behavior of the stickleback is unique and odd, occurring inevitably with nesting. However, upon investigation, it has been found that nesting is an obstacle to hatching. The author conducted a survey in the necessity of nesting behavior in the nine-spined stickleback, Pungitius pungitius, and reached following conclusion: For Pungitius pungitius, nesting is the means of communicating to descendants settlement at a fixed area where cold water flowed bountifully and the water temperature was constant. Pungitius pungitius has evolved a behavior pattern that involves settling where water temperature is uniform, without evolving physiologically or morphologically, thereby enabling Pungitius pungitius to adapt to the changes in environment.

Animals↗

Prior-residence effect in Betta splendens.

Male Siamese fighting fish were presented with the visual image of an aggressive male conspecific for a 150-min test, with attack behaviors monitored continually. A 10-day period of residency in either the test tank or its exact replica resulted in more persistent attack than 10 min of residency. Testing in water other than that in which subjects had resided for 10 days did not produce a reduction in attack. Finally, the extent of attack behavior occurring early in testing was highly and positively correlated with subsequent attack duration; that is, at the start of an encounter, and before severe physical damage has been caused, Bettas may communicate to opponents their "intention" to engage in persistent, injurious aggression.

Aggression↗

[Studies on the Sexual Behavior of Cyrtophora cicatrosa (Stolíczka)(Araneae, Araneidae)].

Courtship and copulation behaviour of the tropical spider Cyrtophora cicatrosa were observed in the laboratory. They are described and compared with the behaviour of C. citricola. Emphasis is placed on the roles of pheromones and web tapping, and on the species' different ways of overcoming the problems resulting from extreme sexual dimorphism.

Animal Communication↗

Effects of elevated egg corticosterone levels on behavior, growth, and immunity of yellow-legged gull (Larus michahellis) chicks.

Eggs of vertebrates contain steroid hormones of maternal origin that may influence offspring performance. Recently, it has been shown that glucocorticoids, which are the main hormones mediating the stress response in vertebrates, are transmitted from the mother to the egg in birds. In addition, mothers with experimentally elevated corticosterone levels lay eggs with larger concentrations of the hormone, which produce slow growing offspring with high activity of the hypothalamo-adrenal axis under acute stress. However, the effects and function of transfer of maternal corticosterone to the eggs are largely unknown. In the present study, we injected corticosterone in freshly laid eggs of yellow-legged gulls (Larus michahellis), thus increasing the concentration of the hormone within its natural range of variation, and analyzed the effect of manipulation on behavioral, morphological, and immune traits of the offspring in the wild. Eggs injected with corticosterone had similar hatching success to controls, but hatched later. Mass loss during incubation was greater for corticosterone-treated eggs, except for the last laid ones. Corticosterone injection reduced rate and loudness of late embryonic vocalizations and the intensity of chick begging display. Tonic immobility response, reflecting innate fearfulness, was unaffected by hormone treatment. Elevated egg corticosterone concentrations depressed T-cell-mediated immunity but had no detectable effects on humoral immune response to a novel antigen, viability at day 10, or growth. Present results suggest that egg corticosterone can affect the behavior and immunity of offspring in birds and disclose a mechanism mediating early maternal effects whereby stress experienced by females may negatively translate to offspring phenotypic quality.

Animals↗

Differential effects of 11-ketotestosterone on dimorphic traits in a teleost with alternative male reproductive morphs.

Species with dimorphic males typically have one morph (type I) associated with territoriality and courtship, and another morph (type II) associated with cuckoldry. In teleost fishes, type I males generally have higher levels of the androgen 11-ketotestosterone (KT) than type II males. Is KT causal to phenotypic differences between morphs? We investigated this question in the midshipman fish (Porichthys notatus) in which type I males have detectable levels of KT during the breeding season, whereas levels in type II males are usually undetectable. Type I midshipman will either cuckold or hold territories and court females, whereas type II males are only known to cuckold. Acoustic courtship by type I's is supported by the sonic motor nucleus, which innervates a sound-producing sonic muscle. Type I males have larger sonic motor nuclei and larger sonic muscles than type II males, consistent with the more dynamic vocal repertoire of type I's. Here, we tested whether intraperitoneal KT implants in adult type II males would induce type I male-like traits in brain, sonic muscle, and behavior. Type II's treated with KT did not differ from blank-implanted type II's in sonic motor nucleus volume. Sonic muscle mass increased in KT-implanted type II's, but did not reach the relative mass naturally observed in type I's. While neither territoriality nor courtship were induced, cuckoldry behavior intensified in KT-implanted type II's. Thus, for some but not all characters, KT exaggerated the expression of already existing type II male traits rather than inducing the type I male-like traits of territoriality, courtship, and an expansive vocal motor system.

Animals↗

The influence of testosterone on territorial defence and parental behavior in male free-living rufous whistlers, Pachycephala rufiventris.

We studied a population of rufous whistlers, Pachycephala rufiventris, throughout a single breeding season in central New South Wales, Australia. We evaluated the relation between plasma testosterone (T) and reproductive behaviors using both simulated territorial intrusions (STIs) and subcutaneous T implants. We compared circulating T values to aggression levels of males (using STI) during pair bond and territory establishment and again during incubation. Although plasma T levels were significantly lower in the latter period, male responsiveness to STI, in terms of proximity to decoy, call rate, and number of attacks on the decoy, was indistinguishable between the two breeding stages. T levels of males exposed to STI were not different from the levels of unexposed free-living males at the same breeding stage. The effect of exogenous T on parental behavior was examined by comparing duration of incubation bouts of males and their mates prior to and after T treatment. T males significantly reduced the amount of time they incubated following implantation, whereas Control males maintained their incubation effort. After cessation of breeding activities, T males displayed significantly higher call rates due to increased use of the primary intersexual advertisement call in this species. The reduction of incubation behavior following T implantation emphasises the functional significance of the rapid decline in T in free-living males during incubation. The results from both experiments suggest that intersexual advertisement, rather than territorial aggression, may be dependent on high T levels in this species.

Aggression↗

Home-orienting behavior in rat pups: the effect of 2 and 3 generations of rehabilitation following intergenerational malnutrition.

In an earlier study in this laboratory, we examined home-orienting ability in rats with histories of intergenerational malnutrition that were rehabilitated postnatally with an adequate protein diet (Galler, 1979). Despite increased size and activity, 1st-generation rehabilitated pups showed no improvement in homing ability when compared to malnourished pups. In the current study, offspring of rats with intergenerational malnutrition were rehabilitated for 2 additional generations and tested for homing ability from postnatal Day 4 through Day 12. The pups rehabilitated for 2 and 3 generations did not perform as well as control pups, even though growth and activity returned to normal levels. Furthermore, the rehabilitated pups failed to demonstrate any preference for the nest on all days of testing; in contrast, control pups favored the nest quadrant by Day 8 of life. No significant improvement was evident on any measure from the 2nd to the 3rd generation of rehabilitation.

Animals↗

Individually variable energy management strategies in relation to energetic costs of egg production.

Marked interindividual variation in metabolic rate suggests considerable complexity in energy management strategies, but attempts to further our understanding of the relationship between resting metabolic rate (RMR), daily energy expenditure (DEE), and reproductive effort have been hampered by the complexity of studying this system in the field. Here, we describe energy management strategies in a captive-breeding system, using Zebra Finches (Taeniopygia guttata), to demonstrate the high level of complexity and interindividual variability in energy expenditure, food intake, locomotor activity, and reproductive effort. In particular, we investigated whether the increase in RMR associated with egg production is additive, resulting in higher DEE and a need for elevated food intake, or whether this cost is compensated by reduced expenditure in nonreproductive components of the energy budget. We found high levels of intra-individual variation in energy expenditure associated with egg production in female Zebra Finches, e.g., comparing nonbreeding stage with the one-egg stage, change in RMR varied from 4.0% and 41.3%, and change in DEE varied from -33.3% to +46.4%. This variation was systematically related to aspects of locomotor activity and reproductive effort. Females with the largest increase in RMR during egg production decreased locomotor activity the most but still had increased DEE at the one-egg stage, and females with high DEE at the one-egg stage produced larger clutches. Our study suggests that females minimize increases in DEE during egg production through behavioral energy reallocation (reduced locomotor activity) but that individuals differ in their use of this strategy, which, in turn, is related to the absolute level of reproductive investment. This suggests a very complex, individually variable system of energy management to meet the demands of egg production.

Animals↗

Dispersed male networks in western gorillas.

Although kin-selection theory has been widely used to explain the tendency of individuals to bias beneficial behaviors towards relatives living within the same social group, less attention has focused on kin-biased interactions between groups. For animal societies in which females emigrate, as is the case for mountain gorillas (Gorilla beringei beringei), encounters between males in different groups often involve aggressive displays that can escalate to physical violence and fatal injuries. However, recent findings on the little-studied western gorilla (Gorilla gorilla) indicate that interactions between social groups occur more frequently than they do in mountain gorillas and are often, although not always, surprisingly nonaggressive. We investigated the pattern of genetic relationships between individuals of different groups and found evidence suggesting a previously unrecognized "dispersed male network" social structure in western gorillas in which the single males leading social groups were usually related to one or more nearby males. We propose that this provides a basis for extra-group, kin-biased behaviors and may explain the reported peaceful intergroup interactions. Furthermore, these results suggest that a patrilocal social structure, in which males remain in their natal region and potentially benefit from kin associations, is a feature unifying African apes and humans.

Animals↗

Limited changes of mouse maternal care after prenatal oxazepam: dissociation from pup-related stimulus perception.

There is evidence that activity at the GABA/benzodiazepine receptor complex in specific brain areas might be enhanced during rodent motherhood. We tested the hypothesis that the manipulation of this neurochemical system by prenatal benzodiazepine exposure affects typical behavioral responses of lactating mouse dams. Outbred CD-1 mouse fetuses were administered either oxazepam (OX, 15 mg/kg) or vehicle twice a day on embryonic days 12-16 and fostered at birth to untreated dams. Female offspring were subsequently mated at the young adult stage. In a first experiment, the behavioural repertoire of the two groups of lactating females was scored (single 10-min session) from postpartum days 3 to 18. When compared with VEH dams, OX females showed a shorter duration of pup-sniffing at 7-10 days and enhanced crouching behaviour when pups had reached the age of 14-18 days. In addition, OX-treated dams used more cotton for nest construction than the controls. The two female groups were differentiated only in the presence of their offspring. In a second experiment aimed at investigating possible OX-induced changes in pup-stimulus perception, the same lactating females were challenged in sequence on postpartum day 8 with three different patterns of pup-related cues consisting of: three 8-day-old live male pups (LP), three same-age dead pups DP, or three dead pups accompanied by pre-recorded ultrasounds (DPU). In the absence of carry-over effects of prenatal dam treatment, LP stimuli elicited a higher frequency of sniffing and digging than the others, whereas the level of licking, and grooming was reduced. In conclusion, the present results indicate that the slight alteration in maternal care resulting from prenatal OX treatment can be dissociated from changes in pup-related stimulus perception.

Animals↗

Exogenous testosterone inhibits several forms of male parental behavior and stimulates song in a monogamous songbird: the blue-headed vireo (Vireo solitarius).

Natural variation in circulating testosterone is thought to play a role in creating inter- and intraspecific variation in paternal care in birds. Experimental elevation of plasma testosterone in males has reduced rates of male care in many species. However, a small number of studies suggest that male care is not inhibited by testosterone when there appears to be strong selection for male care. Both genetic monogamy and male incubation are thought to be the result of selection for biparental care, yet the sensitivity of male care to inhibition by T had not been examined in a species with both characteristics. Male blue-headed vireos have been shown to be strictly monogamous and provide extensive parental care, including incubation. Males received subcutaneous implants of testosterone, flutamide (an anti-androgen), or empty controls. Consistent with previous studies, testosterone treatment decreased the relative contribution to incubation by males and increased the average duration that the nest was left unattended during incubation, relative to control and flutamide treatments. The relative and absolute rates of feeding offspring and nest sanitation by T-treated males also decreased relative to controls. Flutamide-treated males incubated eggs and brooded young more than testosterone-treated males, with control males performing at intermediate values. Testosterone-treated males sang more often and at higher rates, and flutamide-treated males sang less often, than controls. These results demonstrate that even in a strictly monogamous songbird with biparental incubation, all forms of paternal care remain sensitive to elevated plasma testosterone, suggesting a shift from parental effort to mating effort.

Animals↗

Long-term effects of acute perinatal asphyxia on rat maternal behavior.

In this study we used a rat model of graded perinatal asphyxia to study the long-term consequences of this manipulation on rat maternal behavior at adulthood. Rats were delivered by cesarean (C) section and the pups, still in the uterus horns, were placed into a water bath at 37 degrees C for periods of 0 (controls) or 20 min (asphyxia). Subsequently, female pups were given to surrogate mothers, weaned at 21 days postnatally and then left undisturbed until adulthood, when they were mated. Once they gave birth, on postnatal days (Pnds) 1, 3, 5, 7, 9, 11 and 13 they were observed in the home cage five times per day to assess their maternal behavior in an undisturbed condition. In addition, maternal behavior was observed for 30 min in a novel cage on Pnds 4 and 8. Perinatal asphyxia affected maternal behavior in the home cage, hypoxic females being more often found outside the nest area and performing more often behaviors such as self-grooming. Principal component analysis confirmed a more 'active' behavioral profile for hypoxic females. Hypoxic mothers were characterized by a longer latency to perform on-nest behavior and by a reduced frequency of pup retrieval and licking in the novel cage. No significant differences in corticosterone secretion in response to an acute stressor were found in dams belonging to the different treatments or in the body weights of the offspring. These results are suggestive of an arousal deficit due to perinatal hypoxia and point to the dopaminergic system as a potential neurochemical target for an early hypoxic insult.

Age Factors↗