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The role of androgens in the trade-off between territorial and parental behavior in the Azorean rock-pool blenny, Parablennius parvicornis.

Androgen hormones have been shown to facilitate competitive ability in courtship and territorial behavior, while suppressing paternal behavior. The rock-pool blenny, Parablennius parvicornis, provides an excellent model to study the proximate regulation of such a trade-off between territorial and parental behavior, because nest-holder males of this species display these behaviors simultaneously. A field study was carried out in which territorial nest holder males were either treated with long-lasting implants filled with 11-ketotestosterone (11-KT) or with control implants. Males treated with 11-KT showed a higher frequency of aggressive behavior, were more responsive to aggressive challenges, and were more persistent in aggressive behavior than control males. In addition, territories were larger in males treated with 11-KT than in controls. We found evidence for incompatibility between defense of a large territory and high levels of parental behavior. However, contrary to expectation, 11-KT did not suppress parental behavior. We suggest that trade-offs between territorial and parental behavior may not be regulated by androgen hormones but may result from a time constraint in the individual's activity budget.

Absorbable Implants↗

Vocal production in different social contexts relates to variation in immediate early gene immunoreactivity within and outside of the song control system.

In songbirds, a major function of song during the breeding season is mate attraction, and song in this context can be highly sexually motivated. Vocal learning, perception, and production are regulated by the song control system, but there is no evidence that this system participates in the motivation to sing. Instead, brain regions involved in sexual motivation and arousal, including the medial preoptic nucleus (POM), bed nucleus of the stria terminalis (BST), nucleus taeniae (Tn), and area ventralis of Tsai (AVT) might regulate the motivation to sing, at least in a sexual context. The role of these nuclei and song control nuclei (area X and HVC) in vocal production within a breeding context, and other courtship behaviors, was investigated using immunocytochemistry for protein products of immediate early genes (IEGs), ZENK and c-fos (Fos), in flocks of male house sparrows (Passer domesticus) presented with females. Compared to vocalizations from other perches, vocal behavior from a nest box is more likely directed toward females, and sexually motivated. The numbers of ZENK and Fos labeled cells within rostral, but not caudal POM related positively only to vocalizations produced from a nest box. In contrast, the number of ZENK-labeled cells within area X related negatively to vocalizations from a nest box. Additionally, numbers of IEG-labeled cells within rPOM, Tn and AVT related positively to mount attempts. The results support the hypothesis that the POM interacts with the song control system to regulate sexually motivated vocal expression, and are consistent with work indicating that (a) rostral and caudal POM play distinct roles in sexual behavior, and (b) involvement of area X in song is context specific.

Amygdala↗

Prolactin and parental behavior in Adélie penguins: effects of absence from nest, incubation length, and nest failure.

Adélie penguin (Pygoscelis adeliae) males and females, nesting in Antarctica, alternate attendance at the nest with absences of many days to forage at sea. We investigated the importance of tactile input from egg and chicks on prolactin levels by observing nest attendance patterns and obtaining blood samples (1) during the first nest exchange of the incubation stage, (2) from birds whose incubation period was artificially increased or decreased by about 10 days, and (3) from birds whose nests had failed. Prolactin levels in females after 8 to 11 days of absence from the breeding colony did not differ from those in incubating males and did not change after females resumed incubation. Moving eggs between nests resulted in nests in which chicks hatched after about 26, 36 (normal), or 46 days. Duration of incubation did not affect prolactin levels in the parents measured during incubation, at the pip stage, hatch stage, or early brood stage. Adults first left their chicks unguarded on about the same calendar date, regardless of chick age. However, chicks from long incubation nests averaged 8 days younger when they were left unguarded than chicks from control or short-incubation nests. In females, there was no effect of nest failure on prolactin levels. In males, prolactin levels were slightly lower after nest failure than in males tending nests. Testosterone was significantly higher in males after nest failure than in males still tending nests. Prolactin is elevated in Adélie penguins as part of the program of cyclical hormonal changes that accompany the lengthy reproductive season and is relatively independent of tactile input. Sustained prolactin secretion is probably required for the maintenance of parental behavior in offshore feeding species that must be absent from the nest for many days at a time.

Animals↗

The interaction of testosterone and breeding phase on the reproductive behavior and use of space of male zebra finches.

It is well known that androgens play a critical role in mediating the reproductive behavior of males. However, many laboratory experiments that examined the effects of testosterone in male songbirds typically limited their investigations to the early phase of breeding. We sought to determine the influence of testosterone on social behavior, pair bonding, nesting, and use of space in captive zebra finch (Taeniopygia guttata) males as a function of breeding stage (pre-laying, incubation, and nestling phases). Fourteen males were released into an aviary with 14 females and allowed to breed for 7 weeks. Half of the males were given testosterone implants and half were given control implants. During the pre-laying phase, testosterone-implanted males spent significantly more time in nesting activities than control birds and more time elapsed from starting to build a nest to when their mates initiated egg-laying. During the incubation phase, testosterone-implanted subjects spent significantly more time in female-directed and undirected singing. Use of space varied between hormone conditions depending upon breeding phase: there was no difference during pre-laying, but during the incubation and nestling phases, testosterone-implanted subjects used significantly more space. This significant increase in "home range" during the latter phase of the breeding cycle coincides with results from field studies on other species. These results underscore the importance of considering breeding phase in assessing the behavioral sensitivity to hormones.

Animals↗

Hormonal influences on the duration of postpartum maternal responsiveness in the rat.

These studies investigated the role of gestational hormones in controlling the duration of the postpartum period of elevated maternal responsiveness in rats. In the first study, females had pregnancies terminated by Caesarean section, and had ovaries removed on day 16 or 22 of pregnancy. Removal of the ovaries did not prevent the sustaining of an elevated maternal responsiveness for 7 days after pregnancy termination on either day. In the second study, changes in concentrations of estrogen and progesterone during pregnancy, and following parturition, were determined by radioimmunoassay. Levels of these hormones postpartum were found to decline to virgin levels by 7 days postpartum and could not therefore explain the elevated maternal behavior seen at this time. In the third study, ovariectomized virgins were given a pregnancy-mimicking schedule of estradiol and progesterone, with and without prolactin, and were tested for maternal behavior 2 and 7 days later. A 22-day estradiol-progesterone schedule induced rapid maternal behavior within 2 days, but did not sustain it for 7 days. Adding prolactin to the 22-day estradiol-progesterone schedule sustained some components of maternal behavior, notably nest-building and retrieval, but not maternal behavior as a whole.

Animals↗

Effects of maternal state on the responsiveness to nest odors of hooded rats.

Two experiments were designed to determine if pregnant and lactating rats are differentially attracted to their own bedding at various stages of the reproductive cycle and whether they will discriminate between their own nest materials and those of virgins and other lactating animals. We found that from just before birth until the second week postpartum, rats are attracted to their own bedding but do not differentiate between their own and that of other maternal animals. It is suggested that the responsiveness to nest odors in prepartum pregnant animals is a result of hormonal events prior to parturition. In addition, early maternal activities are thought to be facilitated by changes in responsiveness to olfactory stimuli and also by the presence of an attractant which contributes to nest odors.

Animals↗

Maternal behavior following rehabilitation of rats with intergenerational malnutrition. 1. Persistent changes in lactation-related behaviors.

Female rats rehabilitated for one or two generations on an adequate level of dietary protein following a history of intergenerational malnutrition were studied during the postnatal suckling period. No recovery occurred after two generations of dietary rehabilitation for most measures of maternal behavior including active nursing, passive nursing, pup-oriented behavior, time spent in the nest or time spent in contact with the young. In contrast, nest quality improved to normal levels after rehabilitation. In addition, growth of pups born to females rehabilitated for one generation was similar to growth of control pups, and in the case of mothers rehabilitated for two generations, growth of offspring exceeded that of the control pups. These results demonstrate variable sensitivity of different maternal behaviors to rehabilitation following a history of intergenerational malnutrition, with lack of normalization of a considerable number of maternal behaviors. Consequently, some of these abnormalities in maternal behavior may result in persistent abnormalities in the behavior of offspring over several generations.

Age Factors↗

Inter-seasonal maintenance of individual nest site preferences in hawksbill sea turtles.

Within a single population of hawksbill sea turtles (Eretmochelys imbricata), we found a behavioral polymorphism for maternal nest site choice with respect to beach microhabitat characteristics. Some females preferred to nest in littoral forest and in places with overstory vegetation cover, and others preferred to nest in more open, deforested areas. Nest site choice was consistent within and between nesting seasons two years apart. This was not a result of females simply returning to the same location along the shoreline; beach sections used by individual turtles varied between seasons. Nest site choice was not influenced by changes in beach environment (e.g., beach width and foliage cover) or by changes in females' reproductive output (e.g., clutch size), suggesting that fidelity to particular microhabitats is a major determinant of the observed nesting patterns. Because hawksbills exhibit temperature-dependent sex determination, if the behavioral polymorphism in nest site choice has a genetic basis, as is plausible, then this would have implications for sex ratio evolution and offspring survival. By taking an individual-based approach to the study of maternal behavior we reveal previously overlooked individual variation and hope to provide some impetus for more detailed studies of nest site choice.

Animals↗

Use of partial mammectomy to produce malnutrition in the rat.

Partial mammectomy in the Sprague-Dawley rat was studied as a method for producing malnutrition without accompanying differences in litter size. Throughout the litter period, experimental litters weighed less than controls. No difference in the rate of development of various anatomical landmarks was noted with this degree of malnutrition. Maternal behavior in the nursing dam was measured with respect to nursing, nesting, and retrieval behavior. Mammectomized subjects built and maintained better nests and spent more time in a nursing posture, but were less likely to retrieve their litter than were control animals. At weaning the intact mammae of the experimental animals were larger than those of control animals.

Animals↗

Intact neurons of the lateral habenular nucleus are necessary for the nonhormonal, pup-mediated display of maternal behavior in sensitized virgin female rats.

Our research has demonstrated that the lateral habenular nucleus (Lhb) is necessary for the hormonal onset but not the postpartum maintenance of maternal behavior in the rat (K. P. Corodimas, J. S. Rosenblatt, & J. I. Morrell, 1992; K. P. Corodimas, J. S. Rosenblatt, M. E. Canfield, & J. I. Morell, 1993; T. Matthews-Felton, K. P. Corodimas, J. S. Rosenblatt, & J. I. Morell, 1995). To test the role of the Lhb in the nonhormonal onset of maternal behavior, we used the sensitization model in which the continual exposure of females to pups induces maternal behavior. Ovariectomized females received bilateral cytotoxic lesions of neurons of either the Lhb or the dorsal medial cingulate cortex-hippocampus, or they were unoperated. Maternal behavior, activity, and oromotor carrying capability were tested. Complete lesions of the neurons of the Lhb induced significant deficits in pup retrieval and nest building. Sniffing, licking, and crouching behaviors were unaltered. Activity and carrying ability were normal. These results indicate a role for the Lhb that extends to the nonhormonally dependent onset of maternal behavior, but they also indicate a more limited role than in the mediation of the hormonal onset of the behavior.

Animals↗

Behavioral strategies in the cold: effects of feeding and nesting costs.

Animals can meet energetic challenges by acquiring or conserving energy. In the present experiment, we pitted these strategies against each other by housing rats in a cold environment and requiring them to bar press for food pellets and for access to a heated nest. Our question was how rats would exploit these resources to meet the added energetic demand of the cold when food was abundant or scarce. Results showed that rats' allocation of time and effort between competing activities was a function of demand. Feeding and nesting were partially substitutable in the cold: rats traded food for warmth and vice versa, economizing on both time and energy by adjusting their feeding and nesting patterns. Depending upon food abundance and the cost of nesting, rats acquired calories from food, conserved calories by nesting, and borrowed calories from the defense of body weight.

Acclimatization↗

Applications of behavior to poultry management.

The application of poultry behavior to management is discussed with examples of behavior-management interactions relating to commercial poultry husbandry practices. Behaviors that are important for the adaptation of poultry to husbandry include: social behavior, aggression, sexual behavior, feeding, broodiness, cannibalism, nest site selection, and comfort behaviors.

Agonistic Behavior↗

Pineal-dependent and -independent effects of photoperiod on immune function in Siberian hamsters (Phodopus sungorus).

Siberian hamsters (Phodopus sungorus) exhibit reproductive and immunological responses to photoperiod. Short (<10-h light/day) days induce gonadal atrophy, increase leukocyte concentrations, and attenuate thermoregulatory and behavioral responses to infection. Whereas hamster reproductive responses to photoperiod are dependent on pineal melatonin secretion, the role of the pineal in short-day induced changes in immune function is not fully understood. To examine this, adult hamsters were pinealectomized (PINx) or sham-PINx, and transferred to short days (9-h light/day; SD) or kept in their natal long-day (15-h light/day; LD) photoperiod. Intact and PINx hamsters housed in LD maintained large testes over the next 12 weeks; sham-PINx hamsters exhibited gonadal regression in SD, and PINx abolished this effect. Among pineal-intact hamsters, blood samples revealed increases in leukocyte, lymphocyte, CD62L+ lymphocyte, and T cell counts in SD relative to LD; PINx did not affect leukocyte numbers in LD hamsters, but abolished the SD increase in these measures. Hamsters were then treated with bacterial lipopolysaccharide (LPS), which induced thermoregulatory (fever), behavioral (anorexia, reductions in nest building), and somatic (weight loss) sickness responses in all groups. Among pineal-intact hamsters, febrile and behavioral responses to LPS were attenuated in SD relative to LD. PINx did not affect sickness responses to LPS in LD hamsters, but abolished the ameliorating effects of SD on behavioral responses to LPS. Surprisingly, PINx failed to abolish the effect of SD on fever. In common with the reproductive system, PINx induces the LD phenotype in most aspects of the immune system. The pineal gland is required for photoperiodic regulation of circulating leukocytes and neural-immune interactions that mediate select aspects of sickness behaviors.

Animals↗

Changes in plasma androgen levels associated with changes in male reproductive behavior in a brood cycling marine fish.

Like many paternal-caring fishes, male garibaldi (Hypsypops rubicundus) exhibit changes in reproductive behavior during the course of a brood cycle. The purpose of this study was to compare plasma levels of testosterone and 11-ketotestosterone among males in different phases of the brood cycle. These phases are characterized by differences in male courtship, nest maintenance, and egg fanning behavior. Blood samples were taken from wild-caught males, and plasma androgen levels were measured using radioimmunoassay techniques. Androgen levels among males in different phases of the brood cycle were concordant with differences in courtship behavior and also showed a general, positive correspondence with nest maintenance behavior. Levels were nearly inversely related to egg fanning rates. These relationships were independent of seasonal or population level environmental effects. These data suggest a strong three-way relationship among androgen levels, reproductive behavior, and stimuli associated with the progressing brood cycle.

Androgens↗

Nest site attributes and temporal patterns of northern flicker nest loss: effects of predation and competition.

To date, most studies of nest site selection have failed to take into account more than one source of nest loss (or have combined all sources in one analysis) when examining nest site characteristics, leaving us with an incomplete understanding of the potential trade-offs that individuals may face when selecting a nest site. Our objectives were to determine whether northern flickers (Colaptes auratus) may experience a trade-off in nest site selection in response to mammalian nest predation and nest loss to a cavity nest competitor (European starling, Sturnus vulgaris). We also document within-season temporal patterns of these two sources of nest loss with the hypothesis that flickers may also be constrained in the timing of reproduction under both predatory and competitive influence. Mammalian predators frequently depredated flicker nests that were: lower to the ground, less concealed by vegetation around the cavity entrance and at the base of the nest tree, closer to coniferous forest edges and in forest clumps with a high percentage of conifer content. Proximity to coniferous edges or coniferous trees increased the probability of nest predation, but nests near conifers were less likely to be lost to starlings. Flickers may thus face a trade-off in nest site selection with respect to safety from predators or competitors. Models suggested that peaks of nest predation and nest loss to eviction occurred at the same time, although a competing model suggested that the peak of nest loss to starlings occurred 5 days earlier than the peak of mammalian predation. Differences in peaks of mammalian predation and loss to starlings may constrain any adjustment in clutch initiation date by flickers to avoid one source of nest loss.

Animals↗

Maturation of olfactory exploration in golden hamsters.

Following placement into a test cage filled with pine shavings, a litter of 7-8 golden hamster pups (aged 3-18 days postnatal: P3-18) initially displays a period of locomotion which ends reliably in huddling. The latency to establish a huddle (i.e., the duration of locomotion) is significantly longer in the presence of novel odors (fresh or lemon shavings) than more familiar odors (slightly soiled fresh or lemon shavings) but only in pups aged P12 or older. Pups aged P9 or younger do not locomote differentially in the presence of novel or familiar odors. This age difference represents the emergence of olfactory exploration in hamsters between P9 and P12. Exploration of novel odors interferes with initial attempts to establish a single huddle site by a litter, but does not preclude the ultimate aggregation of all pups at a single site as guided by conspecific odors and possibly thermotactile cues as well. Such shifts in the control of behavior by non-nest and nest-related, conspecific stimuli correspond well with the first occurrence of nest exits at P11-12 (e.g., Dieterlen, 1959) coupled with the persistent return of hamster pups to the maternal nest for as long as it is maintained (Rowell, 1961).

Age Factors↗