Patterns of anogenital swelling size and their endocrine correlates during ovulatory cycles and early pregnancy in free-ranging barbary macaques (Macaca sylvanus) of Gibraltar.

In macaques and other cercopithecoid primates, large anogenital swellings (AS) are generally found only in those species in which reproduction is not seasonally restricted. In this respect, the Barbary macaque is unusual because while it shows a marked degree of reproductive seasonality, it also exhibits a striking, exaggerated swelling of the circumanal region and labia. Information on the characteristics of AS in female Barbary macaques is limited in that it is largely based on semiquantitative assessments of swelling size, and there are no data on endocrine parameters associated with AS during ovulatory cycles or early pregnancy. In the present study, we combined quantitative measurements of four swelling size parameters (AS width, height, and depth, and labial width) using a video-imaging technique with fecal estrogen and progestagen determinations in free-ranging females of the Gibraltar Barbary macaque population to 1) characterize the pattern of AS throughout the mating season and early gestation, and 2) examine the relationships among changes in swelling size and endocrine parameters. The patterns of all four swelling parameters correlated significantly with one another, although measures of AS depth and labial width were difficult to obtain. Using the product of AS height and width, the data demonstrate that the occurrence of AS is highly seasonal, with pronounced cyclical changes during the mating season and early pregnancy. Furthermore, the swelling cycles are characterized by progressive size increases from the early to the late follicular phase, in association with an elevated estrogen:progestagen (E:P) ratio, with ovulation occurring during the maximum swelling phase. The results also demonstrated a conspicuous postconception increase in swelling between days 18-30 of gestation. The postconception swellings were on average 80% of the size of that of the conception cycles, and were preceded by a large increase in fecal estrogen levels and the E:P ratio. This is the first study to characterize swelling patterns and their endocrine correlates during ovarian cycles and early pregnancy in naturally reproducing female Barbary macaques. The data provide a solid basis for further studies to explore sociosexual behavioral patterns and the functional significance of AS in this species.

Hibernation effects on memory in European ground squirrels (Spermophilus citellus).

Effects of hibernation on memory were tested in European ground squirrels (Spermophilus citellus). The animals were trained in summer to successfully accomplish two tasks: a spatial memory task in a maze and an operant task on a feeding machine. One group hibernated normally, and the other was prevented from hibernation by maintaining ambient temperature at 22 degrees C. In spring, the same tasks were repeated for both groups and their individual performances compared to the initial training phase. The experimental groups differed significantly in both tasks. The nonhibernating animals had higher levels of retention and needed significantly fewer trials to relearn the tasks than the group that had hibernated. In addition to testing the retention of conditioned tasks, social memory was also studied. The ground squirrels were given a social recognition test in spring with one familiar and one unfamiliar conspecific. In contrast to the conditioned tasks, social memory did not seem to be affected by hibernation. The results indicate negative effects of hibernation on the retention of conditioned tasks, which could produce important constraints on animals. A potential explanation for this memory loss might be changes in neuronal activity, which occur during hibernation.

Water wave communication in the genus Bombina (amphibia).

Amphibians were phylogenetically the first vertebrates to leave the aquatic environment and cope with terrestrial conditions including effects of gravity and substrate on movement and communication. Studies of extant primitive amphibians, which have conserved ancestral morphology and behavior, may help us to understand how gravitational adaptation from aquatic to terrestrial environments occurred. The anuran genus Bombina is a candidate for this type of investigation. In particular, a member of this genus, B. orientalis, is known for its low reaction threshold to minor changes of angular acceleration. We hypothesize that a heightened sensitivity to angular and mechanical accelerations evolved with wave communication. Comparisons of such behavior among B. variegata, B. bombina and B. orientalis may shed light on the evolution of reproductive systems based on water wave communication and relevant vestibular sensitivity. This may represent a transition to derived vocalization modes, which is seen in B. bombina to a certain degree.

Seasonal relationships between plasma and fecal testosterone in response to GnRH in domestic ganders.

In two groups (eight individuals each) of socially acquainted, outdoor-kept, domestic ganders (male Hungarian white: Anser domesticus), basal and GnRH-stimulated plasma testosterone (T) concentrations were compared with fecal testosterone metabolites (TM) in and between three seasons, spring peak of reproductive activity, summer photorefractoriness, and fall sexual reactivation. Plasma was sampled 90 min following the challenge and T was analyzed by radioimmunoassay following the GnRH challenge. Fecal TM were measured by enzyme immunoassay using two group-specific antibodies against 17beta-OH-androgens or a novel antibody against 17-oxo groups, which was found to react with major testosterone metabolites without prior hydrolytic deconjugation. Baseline plasma T and systemic levels were high in spring and fall but low in summer. Plasma T increases in response to GnRH were followed by significantly elevated fecal TM levels 2 to 6 h following the challenge in spring and fall. In fall, at high plasma T levels, fecal TM levels were disproportionally lower than in spring. Variability of TM levels was two to five times higher in feces than in plasma, which explains why correlations between individual plasma T and fecal TM levels generally remained nonsignificant. This points to a low-level short-term relationship between the excreted TM and the plasma T levels. However, the reliability of the method was demonstrated by standard inter- and intraassay variabilities and by a high correspondence between results obtained by the two assays. It is suggested that, with appropriate sample size, fecal TM reflects plasma T increase. However, fecal TM was more variable than the plasma T, and fecal TM responses to GnRH did not always parallel the plasma T response. In addition, seasonal changes in androgen excretion regimes must be taken into account.

Estrus and estrogen changes in mated and unmated free-living European ground squirrels.

The course of behavioral and vaginal estrus and patterns of circulating estrogens were followed in free-living European ground squirrels (Spermophilus citellus) after their emergence from hibernation. Normally mating females were compared to a second group in an area where males had been removed from the population before female emergence. Both groups showed vaginal estrus, but the patterns differed. Mating shortened vaginal estrus to a 3-day period compared to 8 days in unmated females. The extent (cell number) of cell cornification during estrus and the cellular components (percentage distribution) of metestrus did not differ between the two groups. Females in the area without males had significantly higher estrogen levels during estrus and metestrus compared to those in the control area. European ground squirrels were found to be monestrous, as none of the unmated females reentered estrus after metor diestrus was detected. The prolongation of vaginal estrus in unmated females can be viewed as either a physiological inevitability or an adaptation to low mate availability. The extension is still relatively short compared to other sciurid species and perhaps a product of constraints producing a strict time frame for reproduction.

The autumnal fattening of the long-distance migratory garden warbler (Sylvia borin) is stimulated by intermittent fasting.

To investigate the proximate influence of a changing food availability on the seasonal fattening of migratory birds, garden warblers (Sylvia borin) following postnuptial moult were food restricted once a week. Body mass, food intake, plasma hormone and metabolite levels were measured and compared to birds which always had ad libitum food access. The food-restricted birds increased their body mass significantly earlier than the controls. The accelerated fattening was initially not accompanied by hyperphagia and may be due to either an increased food utilisation efficiency or a reduced metabolic rate. An increase of basal glucagon and corticosterone and a decrease of insulin levels prior to fattening were not significant, however, they resulted in a significant decrease of the insulin:glucagon ratio. This ratio was also lower in food-restricted birds than in control birds and may account for the difference in the fattening progress. We conclude that seasonal fattening may be stimulated by a catabolic impulse which could be imposed in free-living birds by a decrease of food availability and/or by an increase of energy expenditure. A negative energy balance is hypothesised to be a common proximate factor affecting migratory as well as winter fattening.

Fecal glucocorticoids document stress in female Barbary macaques (Macaca sylvanus).

Patterns of received aggression and the endocrine response were related to an increase in fecal glucocorticoid metabolites in an intact semifree-ranging group of Barbary macaque females in order to quantify the social stress incurred over a 20-week observation period. The novel result showed that immunoreactive cortisol and 11-oxoetiocholanolone found in the feces can indeed determine the endocrine response of the adrenal gland after a social stressor. After HPLC separation of pooled fecal samples, EIA analyses using three different assays (corticosterone, cortisol, and 11-oxoetiocholanolone) to quantify immunoreactive steroids showed that the corticosterone EIA had no distinctive immunoreactive peaks. Cortisol and 11-oxoetiocholanolone immunoassays showed respectively four and two immunoreactive substances. Time series analyses revealed a behaviorally initiated increase in concentrations of cortisol and 11-oxoetiocholanolone equivalents. Furthermore, both hormone curves exhibit comparable time functions. Either antibody is very suitable for determining glucocorticoid secretion after periods of stress.

Twenty-four hour melatonin profiles in a nocturnally migrating bird during and between migratory seasons.

The garden warbler Sylvia borin is a long-distance migrating bird that travels exclusively at night. During the migratory seasons caged warblers develop intense nocturnal activity which may become even more intense than that during the day. It is demonstrated that in spite of dramatic seasonal changes in the 24-h pattern of locomotor activity measured in caged garden warblers, the corresponding pattern of plasma melatonin changed only very little. As in other species melatonin levels were generally low during the day (below 50 pg/ml) and high at night (350 to 650 pg/ml). A slight reduction in the nocturnal melatonin peak (from 650 pg/ml to between 350 and 400 pg/ml) observed during the autumn and spring migratory seasons, was possibly due to an increased light perception of the birds moving around while being exposed to dim night lights. In general the results show that the 24-h plasma melatonin profiles of these birds are rather robust and that locomotor activity does not depend in a simple and direct way on plasma melatonin levels.

The effect of pinealectomy on circadian plasma melatonin levels in house sparrows and European starlings.

We determined 24-hr plasma melatonin profiles in intact, sham-pinealectomized, and pinealectomized European starlings (Sturnus vulgaris) and house sparrows (Passer domesticus) in a light-dark (LD) cycle and in constant darkness (DD). In the intact and sham-pinealectomized birds of both species, a melatonin rhythm was found, with low levels during the day and high levels during the night. Pinealectomy abolished the nighttime peak of melatonin in both species; hence, levels were low at all times sampled. This uniform response of plasma melatonin to pinealectomy contrasts with the differential response of circadian activity rhythms to pinealectomy for these two species. In DD, locomotor activity in pinealectomized house sparrows is usually arrhythmic, whereas in starlings a rhythm usually persists. This suggests that in the latter species free-running circadian rhythms are not necessarily dependent on a rhythm in plasma melatonin. The same is true for the synchronized activity rhythm observed in pinealectomized birds of both species in LD, as well as for the damped rhythm that persists in pinealectomized house sparrows following an LD-to-DD transfer. The results are consistent with the hypothesis that the pineal and its periodic output of melatonin constitute only one component in a system of at least two coupled pacemakers. They also suggest that there are species differences in the relative role played by the pineal and other pacemakers in controlling circadian rhythms in behavior.

Endogenous reproductive rhythms in a tropical bird.

Like other tropical birds, African stonechats breeding at the equator reproduce once a year during the main rainy season and subsequently carry out a complete molt. These two activities are controlled by endogenous circannual mechanisms: in stonechats held in constant conditions of photoperiod and temperature, a rhythm of gonadal size and molt persisted for up to 7.5 years, which would be a lifetime for free-living conspecifics. The fact that the period of these rhythms deviated from 12 months demonstrates their true endogenous nature. The results represent the longest circannual reproductive rhythm yet documented and suggest that circannual mechanisms may be of particular importance for reproductive timing in tropical organisms.

Circannual variations in plasma luteinizing hormone levels in castrated male European starlings (Sturnus vulgaris).

Intact and castrated male European starlings were held for about 2 years in a constant 12-hr photoperiod and constant temperature conditions. At 1- to 2-month intervals, testicular width was measured by laparotomy, and blood samples were taken for analysis of plasma luteinizing hormone (LH). Most of the control birds went through at least one circannual cycle of testicular width and plasma LH concentration. In the castrates, a similar proportion of birds went through circannual LH cycles with periods indistinguishable from those of the controls. It is concluded that the testes and their hormones are not essential components of the mechanism that generates circannual gonadal cycles in male European starlings.

Seasonal changes in follicle-stimulating hormone in a breeding population of barheaded geese, Anser indicus.

Changes in the plasma titers of follicle-stimulating hormone were investigated in a free-living population of barheaded geese over the whole year. Adult males had pronounced seasonal changes from prebreeding levels of approximately equal to 20 ng/ml to a peak in the breeding season of about 300 ng/ml. Changes in females were only slight. An increase from nondetectable, approximately equal to 20 ng/ml to a mean of 32.2 ng/ml before egg-laying was found. The data are discussed with regard to the literature on free-living species and the previously published data on LH and testosterone levels in the same birds.

Postjuvenile molt in east African and Central European stonechats (Saxicola torquata axillaris, S.t. rubicula) and its modification by photoperiod.

Twenty-eight stonechats of the European race (Saxicola torquata rubicula) from Austria and thirty-one stonechats of the Central African race (S.t. axillaris) from equatorial Kenya were handraised and subsequently investigated with regard to the temporal pattern of their postjuvenile molt. About one half of the birds of each race were held under their own native photoperiod and the other half under the photoperiodic conditions of the other race. The results demonstrated clear differences in the postjuvenile molt between the two races when birds were kept in the photoperiod under which they normally live. The African birds began to molt earlier and molted longer than their European conspecifics. The time course of postjuvenile molt was affected by photoperiod in both races as molt began and ended earlier under the equatorial photoperiod than under the European photoperiodic simulation. The question why the African birds showed strong photoperiodic reactions under these experimental conditions although in their natural environment they experience only minute photoperiodic variations, is critically evaluated: Three possible explanations are discussed: (1) the photoperiodic reaction may represent a relict from a time when ancestors of the tropical populations still lived in more temperate zones; (2) it may be due to the occasional immigration of conspecifics from populations living further north or south; (3) it may result from effects on a (possibly circadian) submechanism of the system controlling annual cycles which is normally affected by other environmental cues but can also be influenced by photoperiodic variations.

Seasonal changes in the behavior and plasma titers of various hormones in barheaded geese, Anser indicus.

A number of behavioral and physiological parameters including the circulating levels of 7 hormones were investigated in a free ranging population of barheaded geese in Seewiesen, W. Germany. Behavioral data were collected on the entire flock from which seasonal changes in the frequencies were determined. In addition, data on the behavior of individuals were obtained to aid in the comparison of seasonal changes in behavior and hormones. The resulting curves were used to examine whether the annual changes in the frequencies of various behaviors were related to changes in circulating hormonal concentrations. Lastly, the physiological data were discussed with regard to reproductive biology in birds.