Spatial distribution of communal nests in a colonial breeding bird: benefits without costs?

Abstract The spatial organization of individuals, or groups of individuals, within a population can provide valuable information about social organization and population dynamics. We analysed the spatial distribution of nests of the sociable weaver (Philetairus socius) on two farms in the Kalahari. Sociable weavers build large communal nests on big savannah trees, forming a pattern of trees with and without nests. We used two spatial statistics, Ripley's K and the pair correlation function, to describe characteristics of the point patterns over a range of distances. (i) At distances of 200 and 300 m, communal nests were clustered. (ii) At distances greater than 1000 m, communal nests were regularly distributed. These findings are independent of the spatial distribution of trees. Furthermore, we used Moran's I to analyse spatial autocorrelation of nest sizes. We expected negative autocorrelation because of competition between nests. But on both farms there was no significant autocorrelation of nest sizes for any distance class. The regular distribution observed at larger distances may indicate competition and/or territoriality among different nests, but the lack of spatial autocorrelation between nest sizes suggests that these interactions may happen between nest clusters rather than between single nests. This was confirmed by significant clustering of nests on small scales. We thus suggest, that colonies of P. socius consist of several nests on adjacent trees forming a cluster of subcolonies. The question why sociable weavers establish subcolonies instead of adding more chambers to the natal nest, could not simply be answered by limitation of nesting space. We suggest a strategy to avoid costs due to increasing colony size.     

Grooming as a reward? Social function of grooming between females in cooperatively breeding marmosets

Classical models of grooming predict that subordinate primates will direct grooming towards dominants to receive coalitionary support from them. In contrast, recent reviews suggest that grooming asymmetries can change with social system and ecological conditions and should reflect asymmetries in services provided by different members of the dyad. We studied grooming patterns between females in six wild groups of common marmosets, Callithrix jacchus, to investigate the relation between social structure and grooming between females in a cooperatively breeding species. We observed grooming frequently and consistently in all study groups. Breeding females groomed nonbreeding females more than vice versa, and grooming between breeding and nonbreeding females was not related to agonistic behaviour. Our results provide some support to the hypothesis that grooming asymmetries are related to differences in services provided by different group members. We suggest that, in cooperatively breeding systems, breeding females may use grooming as an incentive for helper females to stay in the group.     

Changes in biosynthesis and degradation of juvenile hormone during breeding by burying beetles: a reproductive or social role?

Burying beetles, Nicrophorus orbicollis, depend on the location of an unpredictable resource, a small vertebrate carcass, for reproduction. When they discover a carcass, they undergo a correlated rapid rise in titers of juvenile hormone (JH) in the hemolymph and ovarian development. This study investigates the regulation of the changes in JH during breeding in both male and female burying beetles and the role of JH in ovarian development. JH biosynthesis by the corpora allata (CA), measured in vitro, increased in females within an hour of their discovery of a carcass and increased later in males. After returning to low rates as oviposition began, JH biosynthesis rose again 3 days later in females but not in males. Neither the ovaries nor testes synthesized JH. There was a concomitant fall in JH esterase activity within 12 h of discovery of the carcass in both males and females. Although the rise in JH titers and biosynthesis and the fall in JH esterase is correlated with ovarian development, application of methoprene or JH III in the absence of a carcass did not result in vitellogenin uptake by the oocytes. Therefore, we conclude that, in spite of the rapid rise in JH before oviposition, it is not sufficient to regulate vitellogenin synthesis and/or its uptake by the ovaries. We suggest that its role has been preempted to organize social behavior and coordinate parental behavior between mates.     

What affects mRNA levels in leaves of field-grown aspen? A study of developmental and environmental influences

We have analyzed the abundance of mRNAs expressed from 11 nuclear genes in leaves of a free-growing aspen (Populus tremula) tree throughout the growing season. We used multivariate statistics to determine the influence of environmental factors (i.e. the weather before sampling) and developmental responses to seasonal changes at the mRNA level for each of these genes. The gene encoding a germin-like protein was only expressed early in the season, whereas the other tested genes were expressed throughout the season and showed mRNA variations on a day-to-day basis. For six of the genes, reliable models were found that described the mRNA level as a function of weather, but the leaf age was also important for all genes except one encoding an early light-inducible protein (which appeared to be regulated purely by environmental factors under these conditions). The results confirmed the importance of several environmental factors previously shown to regulate the genes, but we also detected a number of less obvious factors (such as the variation in weather parameters and the weather of the previous day) that correlated with the mRNA levels of individual genes. The study shows the power of multivariate statistical methods in analyzing gene regulation under field conditions.     

Stress hormones in relation to breeding status and territory location in colonial king penguin: a role for social density?

Because glucocorticoid (stress) hormones fundamentally affect various aspects of the behaviour, life history and fitness of free-living vertebrates, there is a need to understand the environmental factors shaping their variation in natural populations. Here, we examined whether spatial heterogeneity in breeding territory quality affected the stress of colonial king penguin (Aptenodytes patagonicus). We assessed the effects of local climate (wind, sun and ambient temperature) and social conditions (number of neighbours, distance to neighbours) on the baseline levels of plasma total corticosterone (CORT) in 77 incubating and 42 chick-brooding birds, breeding on territories of central or peripheral colony location. We also assessed the oxidative stress status of a sub-sample of central vs. peripheral chick-brooders to determine whether chronic stress arose from breeding on specific territories. On average, we found that brooders had 55 % higher CORT levels than incubators. Regardless of breeding status, central birds experienced greater social density (higher number of neighbours, shorter distance between territories) and had higher CORT levels than peripheral birds. Increasing social density positively explained 40 % of the variation in CORT levels of both incubators and brooders, but the effect was more pronounced in brooders. In contrast, climate was similar among breeding territories and did not significantly affect the CORT levels of breeding birds. In brooders, oxidative stress status was not affected by local density or weather conditions. These results highlight that local heterogeneity in breeding (including social) conditions may strongly affect the stress levels of breeding seabirds. The fitness consequences of such variation remain to be investigated.     

How does timing,duration and severity of heat stress influence pollen-pistil interactions in angiosperms?

Reproductive development in sexual plants is substantially more sensitive to high temperature stress than vegetative development, resulting in negative implications for food and fiber production under the moderate temperature increases projected to result from global climate change. High temperature exposure either during early pollen development or during the progamic phase of pollen development will negatively impact pollen performance and reproductive output; both phases of pollen development are considered exceptionally sensitive to moderate heat stress. However, moderately elevated temperatures either before or during the progamic phase can limit fertilization by negatively impacting important pollen pistil interactions required for successful pollen tube growth toward the ovules. This mini-review identifies the impacts of heat stress on pollen-pistil interactions and sexual reproduction in angiosperms. A special emphasis is placed on the biochemical response of the pistil to moderately high temperature and the resultant influence on in vivo pollen performance and fertilization.Key words: pollen-pistil interaction, carbohydrates, heat stress, fertilization, pollen tube growth, climate changeSexual reproduction is substantially more sensitive to moderately high temperature stress than vegetative processes.¹ Consequently, the yield of crops with valuable reproductive structures used for food (i.e., grain crops and horticultural crops) and fiber (i.e., cotton) would be especially sensitive to moderately elevated temperatures projected to result from global climate change. Sexual reproduction in angiosperms occurs in essentially three stages: gametophyte development (from meiosis to pollination), the progamic phase (from pollination to zygote formation) and embryo development (from zygote to seed).² During the pro-gamic phase, a number of reproductive processes must occur in a highly concerted fashion for successful fertilization to occur. (1) Anther dehiscence allows mature pollen grains to be transferred to a receptive stigmatic surface; (2) pollen grains germinate and pollen tubes penetrate the stigmatic surface of the pistil; (3) pollen tubes grow through the transmitting tissue of the style and towards a sexually competent ovule; finally, (4) double fertilization produces a zygote and its associated endosperm. Inhibition of any one of the aforementioned processes during the progamic phase, will necessarily limit seed development.³Although the timing and precise coordination of events during the progamic phase are strongly determined by genotype and occur in a unique and well-defined manner for a given species,⁴ the environment encountered either before or during the pro-gamic phase also exerts considerable control over the fertilization process, and can strongly influence reproductive success.⁵ Consequently, high temperature has been shown to substantially limit fertilization in vivo.⁵ Depending upon the timing, duration and severity, heat stress can limit fertilization⁵ by (1) inhibiting male⁶ and female^5,7 gametophyte development, (2) inhibiting pollen germination,^6,8,9 (3) limiting pollen tube growth,^8–11 or (4) by altering the development of tissues required to carry out reproductive processes (i.e., anther and pistil tissues).¹ Although the existing literature concerning heat stress and reproductive development in sexual plants is exhaustive (reviewed in ref. ¹ and ²), the approaches used by various investigators to elucidate plant reproductive responses to high temperature vary substantially from study to study. Consequently, it is the aim of this review to characterize the impact of timing, duration and severity of heat stress on sexual processes occurring during the progamic phase. A special emphasis is placed on the biochemical response of the pistil to moderately high temperature and the resultant influence on in vivo pollen performance.     

Plasma thyroxine-binding proteins and thyroid hormone levels in primate species; is callithricidae thyroid hormone resistant?

Thyroxine(T4)-binding to serum proteins in primates; catarrhini, prosimiae, and platyrrhini were studied by polyacrylamide gel electrophoresis T4 binding analysis. From the electrophoretic analysis, it was shown that thyroxine-binding proteins similar to human thyroxine-binding globulin (TBG) and thyroxine-binding prealbumin (TBPA) were present in catarrhini and prosimiae species, but not in platyrrhini (callithricidae and cebidae). T4-binding analysis also revealed that catarrhini and prosimiae have a high affinity T4-binding protein similar to human TBG. The association constant (Ka) for T4 of the plasma proteins in these species was approximately 2.0 X 10(10) M-1. On the other hand, it was unable to demonstrate a high affinity binding site for T4 in the plasma of platyrrhini species. Both the total and free thyroid hormone concentrations in catarrhini and prosimiae were similar to those in human. Total T4 in cebidae, one of the platyrrhini species, was extremely low. Among 8 animals examined, T4 in 6 was undetectable by radioimmunoassay and the mean T4 of the other two was 2.8 micrograms/dl. However, free thyroid hormone concentrations were similar to those in human. In callithricidae, another platyrrhini species, T4 in plasma was 6.90 +/- 2.11, which is comparable to the level in normal human subjects. However, in this species, high-affinity T4-binding protein was lacking and free thyroid hormone concentrations were extremely high (most were higher than the assay limit). Although the thyroid function of callithricidae remains to be studied, it will be interesting if callithricidae is resistant to thyroid hormone action.     

Captive breeding—a useful tool in the preservation of biodiversity?

Within the next decades species extinction may eliminate between 20 and 50% of the Earth's species. Captive breeding has often been claimed to be a useful tool in preservation of biodiversity. The role of zoos in conservation work and the value of captive breeding are discussed; the latter exemplified by the Peregrine Falcon (Falco peregrinus) Programme and the Arabian oryx (Oryx leucoryx) Programme. Captive breeding programmes are very resource demanding and can only be afforded for a very small number of species, which limits their value significantly. Zoos deal mainly with vertebrates, but these comprise less than 3% of the described species, and although the 878 zoos considered hold more than 20 000 specimens of 140 threatened mammal species, they probably only contribute to the conservation of 20 full species. The situation for birds, reptiles and amphibians is even worse. Zoos face serious problems with minimum viable population sizes and hybridization. However, zoos can make a major contribution to preservation of biodiversity through educating and informing the public. Today, where the crisis of extinction of species has reached such daunting dimensions, captive breeding and otherex situ conservation tools should be the last resort for preserving biodiversity, and captive breeding must not become an excuse to avoid dealing with preservation of habitats.     

Priming of Social Distance? Failure to Replicate Effects on Social and Food Judgments

Williams and Bargh (2008) reported an experiment in which participants were simply asked to plot a single pair of points on a piece of graph paper, with the coordinates provided by the experimenter specifying a pair of points that lay at one of three different distances (close, intermediate, or far, relative to the range available on the graph paper). The participants who had graphed a more distant pair reported themselves as being significantly less close to members of their own family than did those who had plotted a more closely-situated pair. In another experiment, people's estimates of the caloric content of different foods were reportedly altered by the same type of spatial distance priming. Direct replications of both results were attempted, with precautions to ensure that the experimenter did not know what condition the participant was assigned to. The results showed no hint of the priming effects reported by Williams and Bargh (2008).     

Is allantoin in serum and urine a useful indicator of exercise-induced oxidative stress in humans?

To assess whether allantoin levels in serum and urine are influenced by exhaustive and moderate exercise and whether allantoin is a useful indicator of exercise-induced oxidative stress in humans, we made subjects perform exhaustive and moderate (100% and 40% VO₂max) cycling exercise and examined the levels of allantoin, thiobarbituric acid reactive substances (TBARS) and urate in serum and urine. Immediately after exercise at 100% VO₂max, the serum allantoin/urate ratio was significantly elevated compared with the resting levels while the serum urate levels was significantly elevated 30 min after exercise. The serum TBARS levels did not increase significantly compared with the resting levels. Urinary allantoin excretion significantly increased during 60 min of recovery after exercise, however, urinary urate excretion decreased significantly during the same period. The urinary allantoin/urate ratio also rapidly increased during 60 min of recovery after exercise. Urinary TBARS excretion decreased during the first 60 min of the recovery period and thereafter significantly increased during the latter half of the recovery period. On the contrary, after 40% VO₂max of exercise, no significant changes in the levels of urate, allantoin and TBARS in serum or urine were observed. These findings suggest that allantoin levels in serum and urine may reflect the extent of oxidative stress in vivo and that the allantoin which appeared following exercise may have originated not from urate formed as a result of exercise but from urate that previously existed in the body. Furthermore, these findings support the view that allantoin in serum and urine is a more sensitive and reliable indicator of in vivo oxidative stress than lipid peroxidation products measured as TBARS.     

Memory in the grip of emotions and stress: a necessarily harmful impact?

While intense negative events are vividly recalled, information learned during stressful situations is poorly remembered. These differential effects of emotions and stress on memory have been attributed to the physiological manifestations generated during those affective states. Intense emotional and stressful events trigger the secretion of catecholamines and of glucocorticoids, in particular. These hormones would be modulatory agents of memory functions. In the first part of this paper, we review the specific effects emotions and stress have on memory. We then summarize the psychological and biological determinants responsible for these effects. Finally, we discuss different methodological issues that could explain the discrepancy found between the impact of emotions and stress on memory. Defining more precisely the effects emotion and stress have on memory will lead to a better comprehension of the cognitive problems that characterize patients dealing with emotional turmoil, such as patients suffering from depression or post-traumatic stress disorder.     

Does wintering north or south of the Sahara correlate with timing and breeding performance in black‐tailed godwits?

Migrating long distances requires time and energy, and may interact with an individual's performance during breeding. These seasonal interactions in migratory animals are best described in populations with disjunct nonbreeding distributions. The black‐tailed godwit (Limosa limosa limosa), which breeds in agricultural grasslands in Western Europe, has such a disjunct nonbreeding distribution: The majority spend the nonbreeding season in West Africa, while a growing number winters north of the Sahara on the Iberian Peninsula. To test whether crossing the Sahara has an effect on breeding season phenology and reproductive parameters, we examined differences in the timing of arrival, breeding habitat quality, lay date, egg volume, and daily nest survival among godwits (154 females and 157 males), individually marked in a breeding area in the Netherlands for which wintering destination was known on the basis of resightings. We also examined whether individual repeatability in arrival date differed between birds wintering north or south of the Sahara. Contrary to expectation, godwits wintering south of the Sahara arrived two days earlier and initiated their clutch six days earlier than godwits wintering north of the Sahara. Arrival date was equally repeatable for both groups, and egg volume larger in birds wintering north of the Sahara. Despite these differences, we found no association between wintering location and the quality of breeding habitat or nest survival. This suggests that the crossing of an important ecological barrier and doubling of the migration distance, twice a year, do not have clear negative reproductive consequences for some long‐distance migrants.     

Degradation of negative regulators: a common theme in hormone and light signaling networks?

Signal transduction pathways often modulate both positively and negatively acting components to optimize the efficiency of a signal. Recent results have shown that plants make extensive use of regulated proteolysis to modulate signal transduction pathways. An emerging theme from hormone (e.g. auxin and gibberellin) and light signaling pathways is signal or stimulus-induced degradation of negative regulators to optimize plant growth and development.     

Linking Social and Vocal Brains: Could Social Segregation Prevent a Proper Development of a Central Auditory Area in a Female Songbird?

Direct social contact and social interaction affect speech development in human infants and are required in order to maintain perceptual abilities; however the processes involved are still poorly known. In the present study, we tested the hypothesis that social segregation during development would prevent the proper development of a central auditory area, using a "classical" animal model of vocal development, a songbird. Based on our knowledge of European starling, we raised young female starlings with peers and only adult male tutors. This ensured that female would show neither social bond with nor vocal copying from males. Electrophysiological recordings performed when these females were adult revealed perceptual abnormalities: they presented a larger auditory area, a lower proportion of specialized neurons and a larger proportion of generalist sites than wild-caught females, whereas these characteristics were similar to those observed in socially deprived (physically separated) females. These results confirmed and added to earlier results for males, suggesting that the degree of perceptual deficiency reflects the degree of social separation. To our knowledge, this report constitutes the first evidence that social segregation can, as much as physical separation, alter the development of a central auditory area.     

Are corticosterone levels a good indicator of food availability and reproductive performance in a kittiwake colony?

We evaluated the use of corticosterone to gauge forage availability and predict reproductive performance in black-legged kittiwakes (Rissa tridactyla) breeding in Alaska during 1999 and 2000. We modeled the relationship between baseline levels of corticosterone and a suite of individual and temporal characteristics of the sampled birds. We also provided supplemental food to a sample of pairs and compared their corticosterone levels with that of pairs that were not fed. Corticosterone levels were a good predictor of forage availability in some situations, although inconsistencies between corticosterone levels and reproductive performance of fed and unfed kittiwakes suggested that this was not always the case. In general, higher corticosterone levels were found in birds that lacked breeding experience and in birds sampled shortly after arriving from their wintering grounds. All parameters investigated, however, explained only a small proportion of the variance in corticosterone levels. We also investigated whether corticosterone, supplemental feeding, year of the study, breeding experience, body weight, and sex of a bird were able to predict laying, hatching, and fledging success in kittiwakes. Here, breeding experience, year of the study, and body weight were the best predictors of a bird's performance. Corticosterone level and supplemental feeding were good predictors of kittiwake reproductive performance in some cases. For example, corticosterone levels of birds sampled during the arrival stage reliably predicted laying success, but were less reliable at predicting hatching and fledging success. Counts of active nests with eggs or chicks may be more reliable estimates of the actual productivity of the colony. Supplemental feeding had strong effects on kittiwake productivity when natural forage was poor, but had little effect when natural forage was plentiful.     

Pollen donor identity affects timing of stigma receptivity in Collinsia heterophylla (Plantaginaceae): a sexual conflict during pollen competition?

Theory predicts that, during pollen competition, selection may favor a pollen trait that increases donor competitive ability at the expense of the female reproductive function. One such pollen trait could be manipulation of the onset of stigma receptivity. We evaluated the potential occurrence of this kind of sexual conflict by testing female control of the timing of stigma receptivity in the self-compatible annual Collinsia heterophylla. By performing one-donor crosses in the greenhouse, we found that differences in both recipients and pollen donors influenced when stigmas became receptive. Because we did not detect an interaction effect, our result suggests that some donors were consistently better than others at germinating pollen and siring seeds earlier. Unexpectedly, self-pollen was able to fertilize seeds earlier during floral development compared with outcross pollen. These results suggest that female control on timing of stigma receptivity is not complete in this species. In addition, fertilizations that occurred early during floral development resulted in fewer seeds than later fertilizations, possibly indicating a cost of lost control over the onset of receptivity. The ability of pollen donors to influence the timing of stigma receptivity might reflect a conflict between the sexual functions in C. heterophylla.     

Divorce in cooperatively breeding long-tailed tits: a consequence of inbreeding avoidance?

The decision of whether to divorce a breeding partner between reproductive attempts can significantly affect individual fitness. In this paper, we report that 63% of surviving pairs of long-tailed tits Aegithalos caudatus divorced between years. We examine three likely explanations for the high divorce rate in this cooperative breeder. The 'better option' hypothesis predicts that divorce and re-pairing increases an individual's reproductive success. However, divorcees did not secure better partners or more helpers and there was no improvement in their reproductive success following divorce. The 'inbreeding avoidance' hypothesis predicts that females should disperse from their family group to avoid breeding with philopatric sons. The observed pattern of divorce was consistent with this hypothesis because, in contrast to the usual avian pattern, divorce was typical for successful pairs (81%) and less frequent in unsuccessful pairs (36-43%). The 'forced divorce' hypothesis predicts that divorce increases as the number of competitors increases. The pattern of divorce among failed breeders was consistent with this hypothesis, but it fails to explain the overall occurrence of divorce because divorcees rarely re-paired with their partners' closest competitors. We discuss long-tailed tits' unique association between divorce and reproductive success in the context of dispersal strategies for inbreeding avoidance.