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Is food availability a circannual zeitgeber in tropical birds? A field experiment on stonechats in tropical Africa.

Equatorial stonechats (Saxicola torquata axillaris) in Africa are seasonal breeders like their temperate-zone conspecifics (S.t. rubicola). Their annual cycle in gonadal size and function is controlled by an endogenous circannual rhythmicity that has been shown to run for up to 10 years in a constant equatorial photoperiod under laboratory conditions, with a period deviating from 12 months. In nature, however, this rhythm is synchronized with the actual year. Because photoperiod is essentially constant at the equator, it is likely that other environmental factors act as zeitgebers. The authors test whether food availability affects reproductive cycles of free-living East African stonechats. The authors offered supplemental food to the birds 2 months before the regular onset of the breeding season. Supplementally fed males started to sing and display earlier than males of control pairs that did not receive extra food. Although the supplemented food advanced the onset of the breeding season in the pairs that were fed, the onset of the postnuptial molt following the breeding season was not correspondingly shifted. Furthermore, in the year following the experiment, all pairs initiated breeding at the same time. The authors conclude that food availability does not act as a zeitgeber, but rather as a factor that modifies the timing of reproduction without affecting the underlying rhythmicity. The authors propose that this is adaptive under environmental conditions that are relatively constant within a given year but may vary considerably between years. The zeitgeber synchronizing the endogenous rhythmicity remains to be identified.

Animals↗

Winter adaptations of male deer mice (Peromyscus maniculatus) and prairie voles (Microtus ochrogaster) that vary in reproductive responsiveness to photoperiod.

Individuals of many nontropical rodent species restrict breeding to the spring and summer. Seasonal reproductive quiescence putatively reflects the energetic incompatibility of breeding and thermoregulatory activities. However, so-called "out-of-season" breeding occurs in virtually all rodent populations examined, suggesting that the incompatibility can be resolved. Both reproductive inhibition and development of energy-saving adaptations are mediated by environmental photoperiod, but some individuals do not inhibit reproduction in short days. In order to assess the costs and benefits of winter breeding, the present study examined the extent to which male prairie voles (Microtus ochrogaster) and deer mice (Peromyscus maniculatus) that maintained summer reproductive function in winter-simulated daylengths also maintained summer thermoregulatory adaptations. Circadian locomotor activity patterns, basal metabolic rate, capacity for nonshivering thermogenesis, nest building, body mass, and daily food consumption were compared among short-day (LD 8:16) regressed males, short-day (LD 8:16) nonregressed males, and long-day (LD 16:8) males. Short-day nonregressed deer mice resembled long-day conspecifics in terms of body mass and nest-building activities; however, the locomotor activity pattern of short-day nonregressed deer mice was similar to that of their short-day regressed conspecifics. Short-day nonregressed prairie voles had body masses similar to those of long-day conspecifics. Regardless of their reproductive response to photoperiod, short-day prairie voles reduced their daily food consumption and wheel-running activity, compared to long-day voles. These results suggest that winter breeding has energetic costs, most likely resulting from maintaining a "summer-like" body mass relative to that of reproductively regressed animals. These costs may be ameliorated to some extent by the reduction in locomotor activity and nest-building behavior emitted by short-day animals, regardless of reproductive response to short days. Thus, the occurrence of winter breeding may be the result of sufficient numbers of reproductively photoperiod-nonresponsive morphs in the population and sufficiently mild ambient conditions to permit survival of these larger animals.

Adaptation, Physiological↗

The evolution of activity breaks in the nest cycle of annual eusocial bees: a model of delayed exponential growth.

BACKGROUND: Social insects show considerable variability not only in social organisation but also in the temporal pattern of nest cycles. In annual eusocial sweat bees, nest cycles typically consist of a sequence of distinct phases of activity (queen or workers collect food, construct, and provision brood cells) and inactivity (nest is closed). Since the flight season is limited to the time of the year with sufficiently high temperatures and resource availability, every break reduces the potential for foraging and, thus, the productivity of a colony. This apparent waste of time has not gained much attention. RESULTS: We present a model that explains the evolution of activity breaks by assuming differential mortality during active and inactive phases and a limited rate of development of larvae, both reasonable assumptions. The model predicts a systematic temporal structure of breaks at certain times in the season which increase the fitness of a colony. The predicted pattern of these breaks is in excellent accordance with field data on the nest cycle of the halictid Lasioglossum malachurum. CONCLUSION: Activity breaks are a counter-intuitive outcome of varying mortality rates that maximise the reproductive output of primitively eusocial nests.

Animals↗

Courtship stimulates aromatase activity in preoptic area of brain in male ring doves.

Aromatase activity was quantified in vitro in different regions of the basal forebrain of male ring doves at various times after the initiation of courtship. By 24 h after the introduction of a female dove into the cage of a male, there was a significant, nearly two-fold, increase in aromatase activity which was confined to the ventromedial preoptic area (VM-POA) where enzyme activity was concentrated. This increase in aromatase activity, which was sustained through day 5 of courtship, presumably serves to activate nest-oriented behavior.

Animals↗

Alterations in hypothalamic vasoactive intestinal peptide-like immunoreactivity are associated with reproduction and prolactin release in the female turkey.

Vasoactive intestinal peptide (VIP) has potent PRL-releasing activity, but its physiological role in the regulation of PRL release during the avian reproductive cycle is not known. We used indirect immunofluorescence to determine if changes in hypothalamic VIP are associated with the shifts in circulating PRL during the reproductive cycle of the domestic turkey. In the naturally hyperprolactinemic incubating hen, the majority of VIP immunoreactivity (VIP-IR) existed within neurons of the infundibular nuclear complex (INF) and fibers in the external layer of the median eminence. Within the INF, the numbers of VIP-IR cells increased during the cycle, paralleling increases in serum PRL. In the reproductively inactive, nonphotostimulated hen with low serum PRL, essentially no positive cells were noted, whereas the incubating hen exhibited 32.1 +/- 2.2 cells/pair of adjacent sections in the anterior INF and 59.6 +/- 2.0 cells in the posterior INF. Exposure of inactive hens to a stimulatory photoperiod resulted in a 2.6-fold increase in serum PRL with the appearance of VIP-IR cells in the INF. During laying and incubation, further increases were observed in the number of positive cells in the INF and serum PRL as well as a greater fiber density in the median eminence. To further examine the association between changes in VIP-IR and serum PRL, circulating PRL was artificially lowered by depriving incubating hens of their nests for 0, 2, 5, and 10 days. On day 2 of nest deprivation, serum PRL declined markedly to 12% of day 0 levels, with VIP-IR cell numbers at 64% and 46% in the anterior and posterior INF, respectively. By day 10, birds exhibited cell numbers in the INF averaging 20% of those observed in the day 0 incubating hens, with serum PRL at 6% of day 0 levels. The results of these studies indicate a possible causal relationship between hypothalamic VIP and changes in PRL secretion during the avian reproductive cycle, providing a basis for further research on the importance of this peptide as well as factors responsible for the modulation of its expression in hypothalamic INF neurons.

Animals↗

Energetics of nestling growth and parental effort in Antarctic fulmarine petrels.

Antarctic fulmarine petrels breed in some of the coldest conditions encountered by any bird and their young grow twice as fast as predicted allometrically. To examine the energetic consequences of fast growth in a cold environment, we used the doubly labeled water technique to measure field metabolic rates of adults (three species) and different-aged nestlings (four species) of Antarctic fulmarine petrels in the Rauer Islands, East Antarctica: Antarctic fulmar Fulmarus glacialoides, Antarctic petrel Thalassoica antarctica, Cape petrel Daption capense and snow petrel Pagodroma nivea. We used our data to assess parental effort and, together with literature values on nestling growth and resting metabolic rate, to construct and partition nestling energy budgets. Nestling total energy expenditure and peak daily metabolic rate were significantly higher than predicted allometrically (33-73% and 17-66% higher, respectively), and the relative cost of growth in nestling petrels was among the highest reported for birds (54-72 kJ g(-1)). Parental effort during the nestling-feeding period was identical in adult Cape and Antarctic petrels (3.5 times basal metabolic rate, BMR), and was somewhat (but not significantly) higher in snow petrels (4.6 times BMR). These values are comparable to those of other high-latitude procellariiform birds. Thus, despite the constraints of a compressed breeding season, cold temperatures and fast-growing nestlings, adult Antarctic fulmarine petrels do not work harder than procellariid adults whose chicks grow much more slowly. Our findings suggest that obtaining sufficient food is generally not a constraint for adult fulmarine petrels and that factors operating at the tissue level limit nestling growth rate.

Adaptation, Physiological↗

Evidence for nest-odour recognition in two species of diving petrel.

In nearly every procellariiform species, the sense of smell appears to be highly adapted for foraging at sea, but the sense of smell among the diving petrels is enigmatic. These birds forage at considerable depth and are not attracted to odour cues at sea. However, several procellariiform species have recently been shown to relocate their nesting burrows by scent, suggesting that these birds use an olfactory signature to identify the home burrow. We wanted to know whether diving petrels use smell in this way. We tested the common diving petrel Pelecanoides urinatrix and the South-Georgian diving petrel Pelecanoides georgicus to determine whether diving petrels were able to recognise their burrow by scent alone. To verify the efficacy of the method, we also tested a bird that is known to use olfaction for foraging and nest recognition, the thin-billed prion Pachyptila belcheri. In two-choice T-maze trials, we found that, for all species, individuals significantly preferred the odour of their own nest material to that of a conspecific. Our findings strongly suggest that an individual-specific odour provides an olfactory signature that allows burrowing petrels to recognize their own burrow. Since this ability seems to be well developed in diving petrels, our data further implicate a novel adaptation for olfaction in these two species that have been presumed to lack a well-developed sense of smell.

Animals↗

Switch to diester preen waxes may reduce avian nest predation by mammalian predators using olfactory cues.

It has long been recognised that nest depredation by olfactory-searching mammals greatly influences the reproductive success of ground-nesting birds. Yet adaptations of birds to diminish smell during nesting have rarely been investigated. Recently, a remarkable shift in the composition of uropygial gland secretions (preen waxes) was discovered in many ground-nesting shorebirds and ducks that begin incubation, during which the usual mixtures of monoester preen waxes are replaced by mixtures of less volatile diester waxes. In this study we show experimentally that an olfactory-searching dog had greater difficulty detecting mixtures of the less volatile diesters than mixtures of monoesters. This is consistent with the hypothesis that diester preen waxes reduce birds' smell and thereby reduce predation risk.

Animals↗

Interspecific variation in beeswax as a biological construction material.

Beeswax is a multicomponent material used by bees in the genus Apis to house larvae and store honey and pollen. We characterized the mechanical properties of waxes from four honeybee species: Apis mellifera L., Apis andreniformis L., Apis dorsata L. and two subspecies of Apis cerana L. In order to isolate the material effects from the architectural properties of nest comb, we formed raw wax in to right, circular cylindrical samples, and compressed them in an electromechanical tensometer. From the resulting stress-strain curves, values for yield stress, yield strain, stress and strain at the proportional limit, stiffness, and resilience were obtained. Apis dorsata wax was stiffer and had a higher yield stress and stress at the proportional limit than all of the other waxes. The waxes of A. cerana and A. mellifera had intermediate strength and stiffness, and A. andreniformis wax was the least strong, stiff and resilient. All of the waxes had similar strain values at the proportional limit and yield point. The observed differences in wax mechanical properties correlate with the nesting ecology of these species. A. mellifera and A. cerana nest in cavities that protect the nest from environmental stresses, whereas the species with the strongest and stiffest wax, A. dorsata, constructs relatively heavy nests attached to branches of tall trees, exposing them to substantially greater mechanical forces. The wax of A. andreniformis was the least strong, stiff and resilient, and their nests have low masses relative to other species in the genus and, although not built in cavities, are constructed on lower, often shielded branches that can absorb the forces of wind and rain.

Animals↗

Corticosterone selectively decreases humoral immunity in female eiders during incubation.

Immunity is hypothesized to share limited resources with other physiological functions and this may partly account for the fitness costs of reproduction. Previous studies have shown that the acquired immunity of female common eider ducks (Somateria mollissima) is suppressed during their incubation, during which they entirely fast. Corticosterone was proposed to be an underlying physiological mechanism for such immunosuppression. Therefore, the current study aimed to assess the effects of exogenous corticosterone on acquired immunity in captive eiders. To this end, females were implanted with corticosterone pellets at different stages of their incubation fast. We measured total immunoglobulin levels, T-cell-mediated immune response, body mass and corticosterone levels in these females and compared them with those of control females prior to and after manipulation (i.e. corticosterone pellet implantation). To mimic corticosterone effects on body mass, we experimentally extended fasting duration in a group of females termed ;late fasters'. Implanted females had corticosterone levels 6 times higher and lost 35% more mass than control females. Corticosterone levels in ;late fasters' were similar to those in control females but body mass was 8% lower in the former. The decrease in the immunoglobulin levels of corticosterone implanted females was twice as high as in control females, while the T-cell-mediated immune response was not significantly affected by the treatment. We found a decrease in the T-cell-mediated immune response only in ;late fasters' (by 60%), while the immunoglobulin level was not lower in this group than in corticosterone implanted or control females. Our study shows that in incubating eiders, exogenous corticosterone only decreased humoral immunity. We suggest that the immunosuppressive effect of corticosterone could be mediated through its effects on body reserves. Further experiments are required to determine the relationship between body condition and immune system in fasting birds.

Animals↗

Embryonic and larval respiration in the arboreal foam nests of the African frog Chiromantis xerampelina.

In Zimbabwe, female Chiromantis xerampelina construct spherical foam nests that are suspended above temporary water. The nests average 624 ml in volume and contain 854 eggs. The 1.7 mm ova have exceptionally thin jelly capsules and are dispersed in the foamy core of the nest, which is surrounded by a layer of eggless foam. At 25 degrees C, each embryo requires 3.5 days to reach hatching at developmental stage 22, during which it consumes 30 microliters of oxygen. After hatching, each larva remains in the nest for 2 more days and consumes a further 123 microliters of oxygen. The fresh foam contains 77% air, which is sufficient to supply all of the oxygen requirements of the embryos until well after they hatch. Therefore, the size of the egg mass is not limited by oxygen availability as it is in many other anurans. Oxygen also diffuses into the nest from the atmosphere, but the rate is severely restricted by the wet foam, despite the presence of bubbles. Drying of the outer layer of foam greatly increases its oxygen conductance, but the larvae remain in the inner core of wet foam, where they compete for oxygen at the periphery. With further drying of the nest, the wet foam diminishes in volume and concentrates the larvae at a time when their oxygen demands are approaching the maximum. Oxygen pressures within the wet foam drop below 10 kPa and oxygen uptake by the larvae becomes progressively limited, possibly stimulating their emergence from the nest. The delay between hatching and escape from the nest permits the larvae to grow and mature to a stage at which all of the clutch can emerge simultaneously.

Animals↗

Improving housing conditions for laboratory mice: a review of "environmental enrichment".

Laboratory animal facilities have been designed to provide a standard environment where animals can be kept in good physical health at the same time as economic and ergonomic considerations are met. Recognizing the potential welfare problem associated with behavioural restriction in such housing systems, a number of attempts have been made to improve this environment, generally described under the term "environmental enrichment". Modifications of cages for mice usually consist of providing material for nest building and structures which can serve as hiding places and/or for climbing. We have reviewed 40 studies carried out between 1987 and 2000, in which preferences as well as the effect of housing modifications have been studied. Mice will work for access to nesting material and make use of this material to make nests in which they rest. They prefer a more complex cage to the standard cage and will also work for access to cages with shelter and raised platforms. On the basis of present knowledge, it is recommended that mice should have access to nesting material. Strategies for future research are outlined in the article.

Animal Husbandry↗

Rats' preferences for corn versus wood-based bedding and nesting materials.

Corn by-products can be used as bedding and nesting products. Corn-cob bedding resists ammonia build-up and corn-husk nesting material resists dampness. It is not clear whether these advantages are at the expense of animal comfort. Corn cob was compared to aspen chip bedding, and corn husk to paper strip nesting material. Data from 20 rats with differential early bedding experience suggested that they prefer aspen chip, but are also biased towards the bedding they were raised on. Data from 10 rats with no prior nesting material experience suggested that paper strip was preferred over cornhusk. Thus, corn-cob products are not recommended except in situations where air quality and/or flooding are significant problems.

Animal Husbandry↗

Observations on the prevalence of nest-building in non-breeding TO strain mice and their use of two nesting materials.

The spontaneous performance of nest-building behaviour by non-breeding laboratory mice suggests that routinely providing nesting material might be a suitable environmental enrichment. If nesting material is to be provided routinely, this should have characteristics which are preferred, or at least accepted by a considerable proportion of the animal population; it should also be inexpensive. The present study therefore examined the prevalence of nest-building behaviour in 39 individually-housed, non-breeding, female mice, and their preferences for a commercial nesting product and a less expensive source of material (paper towels). Within minutes of the materials being placed in the cages, the mice began manipulating the paper towels. Thirty-six of the mice subsequently constructed nests during the first dark phase after the materials had been placed in the cage--the remaining three mice constructed nests during the following 48 h. The nests were usually constructed from a mixture of the two materials, though observations indicated the mice might have preferred characteristics of the inexpensive paper towels. There was a strong tendency to build nests in the same location used for sleeping prior to the nesting material being provided, and similarly, the mice were conservative in the site chosen to build a second nest after the first was removed. The most frequently chosen site for nest-building was under the feeder. Other studies have reported a high motivation for nest-building behaviour, widespread performance amongst many strains, and nest-building as a thermoregulatory behaviour by animals housed in standard laboratory air temperature. In conjunction with these findings, the present results suggest that routinely providing paper towels is an inexpensive and practical means of environmental enrichment for non-breeding, laboratory mice.

Animal Husbandry↗

Preferences for nesting material as environmental enrichment for laboratory mice.

Behavioural and psychological needs of laboratory animals generally cannot adequately be met in standard laboratory cages. Environmental enrichment, which provides a more structured environment can enhance the well-being of laboratory animals. They may perform more of their species-specific behaviour and may control their environment in a better way. An easily applicable form of enrichment for laboratory mice is nesting material. Six different types of nesting materials were evaluated in a preference test with male and female animals of two strains [C57BL/6J or BALB/c, n = 48]. No significant differences in preference were found between the strains or between the sexes. All mice showed a clear preference for cages with tissues or towels as compared to paper strips or no nesting material, and for cages with cotton string or wood-wool as compared to wood shavings or no nesting material. Paper-derived materials were preferred over wood-derived materials, although the results also suggest that the nature (paper or wood) of the nesting material is less important than its structure, which determines the nestability of the material. Nesting material may be a relatively simple method to contribute to the well-being of laboratory mice.

Animal Husbandry↗

Blood parasites of some West African rainforest birds.

A total of 969 birds representing 121 species of 21 families from the West African nations of Cameroon, Equatorial Guinea and Ivory Coast were examined for haematozoa using thin blood smears; 277 individuals (28.6%) harbored blood parasites. The parasites identified included species of Haemoproteus (7.7% prevalence), Plasmodium (10.7%), Leucocytozoon (4.6%), and Trypanosoma (7.3%). In addition, microfilariae of filariid nematodes were present in 3.6% of the individuals examined. The birds were collected over a period of 12 years, from 1989-2001, from rainforest and ecotone habitats. We report a relatively high prevalence of parasites in colonial nesting birds, and two species of ground nesting birds. In addition, we compared data from bird species collected at a site identical to a previously published study, and did not find significant differences in parasite prevalence between the two years constituting two different seasons. Our results are also compared to other studies in Africa that implement similar and different methodologies.

Africa, Western↗