Age-related patterns of reproductive success in house sparrows Passer domesticus

A common life history pattern in many organisms is that reproductive success increases with age. We report a similar pattern in house sparrows Passer domesticus , older individuals performed better than yearlings for most measures of reproductive success. Older males and females began breeding earlier in a given season and fledged more young than their yearling counterparts. Individual males also fledged more young in their second breeding season than they did in their first, but individual females did not show consistent improvement in reproductive success from year one to two. A path analysis indicated that age in both sexes acted primarily through the timing of breeding; earlier nesters laid more eggs and hence fledged more young but did not have more nesting attempts. We tested whether the increased reproductive success with age arose from high quality individuals surviving to be older (selection hypothesis). In contrast to the main prediction of this hypothesis that reproductive success and survival should be positively related, we found that survival from one year of age to two years of age was negatively related to reproductive success in the first year for males and females combined. Additionally, individuals that survived to breed as two-year-olds did not differ in total young fledged in their first year from those that did not survive to their second season of breeding. Our results indicate that fledgling production increases with age due to improvements in timing of breeding, particularly in females, and not because of the loss of poor breeders or increased output. Mechanisms producing age-related differences in timing of breeding warrant further study.     

Synergistic effects of food and predators on annual reproductive success in song sparrows

The behaviour literature is full of studies showing that animals in every taxon balance the probability of acquiring food with the risk of being preyed upon. While interactions between food and predators clearly operate at an individual scale, population-scale studies have tended to focus on only one factor at a time. Consequently, interactive (or 'synergistic') effects of food and predators on whole populations have only twice before been experimentally demonstrated in mammals. We conducted a 2 x 2 experiment to examine the joint effects of food supply and predator pressure on the annual reproductive success of song sparrows (Melospiza melodia). Our results show that these two factors do not operate in an additive way, but instead have a synergistic effect on reproduction. Relative to controls, sparrows reared 1.1 more young when food was added and 1.3 more when predator pressure was low. When these treatments were combined 4.0 extra young were produced, almost twice as many as expected from an additive model. These results are a cause for optimism for avian conservation because they demonstrate that remedial actions, aimed at simultaneously augmenting food and reducing predators, can produce dramatic increases in reproductive success.     

Influences of weather on reproductive function in female Song sparrows, Melospiza melodia

Prolonged and severe periods of inclement weather can retard gonadal development in early spring or disrupt the breeding cycle in summer. Previous investigations on the endocrine responses of free-living avian species to storms have been restricted almost entirely to males. This communication presents the endocrine responses of free-living female Song sparrows, Melospiza melodia, during gonadal recrudescence after a severe winter, and during storms in early spring and summer. As in males, gonadal recrudescence was delayed by the severe winter weather, and this effect was exacerbated by an aseasonal snow storm in early spring. Unlike males, female Song sparrows did appear to be stressed by this snow storm, since body mass and fat deposit decreased, and plasma levels of corticosterone increased. A second storm in early summer did not, however, prove to be stressful to females, although males were apparently stressed by the same storm. At the time the early summer storm occurred, young had been fledged and were being fed mostly by the male. Females begin preparing for a second clutch of eggs and do not feed fledglings to the same extent as males. Thus it is possible that males are stressed by storms at this time, because they are feeding several young as well as themselves during a period of reduced availability of food, whereas females are feeding themselves only. These data suggest that, not only do the endocrine responses to inclement weather vary in relation to stage in the reproductive cycle when storms strike, but also that there are sex differences in those responses.     

Influences of weather on reproductive function in male Song sparrows, Melospiza melodia

The avian reproductive cycle is integrated and timed by a variety of environmental factors that include day length, availability of food, nest sites, presence of a mate, etc. However, reproduction can be delayed or disrupted by inclement weather and, until recently, the endocrine responses of avian species to storms were entirely unknown. This communication describes changes in endocrine state in free-living male Song sparrows, Melospiza melodia, during gonadal recrudescence after a severe winter, and in response to storms in early spring and summer. Both the severe winter weather and an early spring snow storm delayed testicular development, and depressed circulating levels of testosterone. The sparrows did not appear to be stressed, however, since body mass and fat deposits were higher than in birds sampled during the previous spring after a less severe winter.
A second storm occurred in late May and early June when Song sparrows were feeding young. Curiously, plasma levels of testosterone were not affected, but the sparrows did appear to be stressed since there was a significant decline in body mass, and a marked increase in circulating levels of corticosterone. These data suggest that the effects of inclement weather on reproductive function can vary in relation to the stage in the reproductive cycle when the storm strikes, and that the endocrine mechanisms underlying acclimatization to severe meteorological conditions are also different.     

Organizational effects of maternal testosterone on reproductive behavior of adult house sparrows

Despite the well-known, long-term, organizational actions of sex steroids on phenotypic differences between the sexes, studies of maternal steroids in the vertebrate egg have mainly focused on effects seen in early life. Long-term organizational effects of yolk hormones on adult behavior and the underlying mechanisms that generate them have been largely ignored. Using an experiment in which hand-reared house sparrows (Passer domesticus) from testosterone- or control-treated eggs were kept under identical conditions, we show that testosterone treatment in the egg increased the frequency of aggressive, dominance, and sexual behavior of 1-year-old, reproductively competent house sparrows. We also show that circulating plasma levels of progesterone, testosterone, 5alpha-dihydrotestosterone, and 17beta-estradiol did not differ between treatment groups. Thus, a simple change in adult gonadal hormone secretion is not the primary physiological cause of long-term effects of maternal steroids on adult behavior. Rather, differences in adult behavior caused by exposure to yolk testosterone during embryonic development are likely generated by organizational modifications of brain function. Furthermore, our data provide evidence that hormone-mediated maternal effects are an epigenetic mechanism causing intra-sexual variation in adult behavioral phenotype.     

The period gene and allochronic reproductive isolation in Bactrocera cucurbitae

Clock genes that pleiotropically control circadian rhythm and the time of mating may cause allochronic reproductive isolation in the melon fly Bactrocera cucurbitae (Coquillett) (Diptera: Tephritidae). Flies with a shorter circadian period (ca. 22 h of locomotor activity rhythm) mated 5 h earlier in the day than those with a longer circadian period (ca. 30 h). Mate-choice tests demonstrated significant pre-mating isolation between populations with short and long circadian periods. Pre-mating isolation did not occur when the mating time was synchronized between the two populations by photoperiodic controls, indicating that reproductive isolation is due to variations in the time of mating and not any unidentified ethological difference between the two populations. We cloned the period (per) gene of B. cucurbitae that is homologous to the per gene in Drosophila. The relative level of per mRNA in the melon fly exhibited a robust daily fluctuation under light : dark conditions. The fluctuation of per expression under dark : dark conditions is closely correlated to the locomotor rhythm in B. cucurbitae. These results suggest that clock genes can cause reproductive isolation via the pleiotropic effect as a change of mating time.     

Plastid development in primary leaves of Phaseolus vulgaris: The light-induced development of the chloroplast cytochromes

Summary Etioplasts obtained from the primary leaves of dark-grown bean plants contained cytochromes f, b-559_LP and b-563 in a molar ratio of approximately 1.0:2.0:1.5. On illumination of the plants there was a lag of between 10 and 15 h before these cytochromes increased in amount, but after 48 h they had increased from 6- to 10-fold on a per plastid basis. The presence of cytochrome b-559_HP in the plastids was first detected after 15 h of illumination, which coincided with the commencement of grana formation and the onset of a number of photosynthetic reactions in the greening leaves. After 48 h of illumination the molar ratio for cytochromes f, b-559_HP, b-559_LP and b-563 was 1.0:1.2:2.8:2.6.Agranal chloroplasts formed by the exposure of dark-grown plants to intense light flashes contained high amounts of cytochromes f, b-559_LP and b-563 but cytochrome b-559_HP could not be detected.As the light-induced formation of cytochromes f, b-559_LP and b-563 was substantially inhibited by D-threo chloramphenicol, but not by the L-threo isomer, it seems likely that their formation was dependent on 70S ribosomes. Both chloramphenicol isomers gave plastids which lacked cytochrome b-559_HP.     

RNA metabolism during light-induced chloroplast development in euglena

Methods are described which provide good recoveries of non-degraded chloroplast and non-chloroplast RNAs from Euglena gracilis var. bacillaris. These have been characterized by comparing the RNA from W₃BUL (an aplastidic mutant of Euglena), with that of wild-type cells which have been resolved into chloroplast and non-chloroplast fractions. Using E. coli RNA as a standard, the RNAs from W₃BUL and from the non-chloroplast fraction of green cells exhibit optical density peaks, upon sucrose gradient centrifugation, at 4S, 10S, and 19S. The chloroplast fraction exhibits optical density peaks at 19S and 14S with the 19S component predominating. Application of various techniques for the separation of RNAs to the problem of separating the chloroplast and non-chloroplast RNAs, without prior separation of the organelle, have not proven successful.

³²P_i is readily incorporated into RNA by cells undergoing light-induced chloroplast development, and fractionation at the end of development reveals that although chloroplast RNAs have a higher specific activity, the other RNAs of the cells are significantly labeled as well. The succession of labeling patterns of total cellular RNA as light-induced chloroplast development proceeds are displayed and reveal that all RNA species mentioned above eventually become labeled. In contrast, cells kept in darkness during this period incorporate little ³²P_i into any RNA fraction. In addition, a heavy RNA component, designated as 28S, while representing a negligible fraction of the total RNA, becomes significantly labeled during the first 24 hours of illumination. While there is light stimulated uptake of ³²P_i into the cells, this uptake is never limiting in the light or dark, for RNA labeling.

On the basis of these findings, we suggest that extensive activation of non-chloroplast RNA labeling during chloroplast development is the result of the activation of the cellular synthetic machinery external to the chloroplast necessary to provide metabolic precursors for plastid development. Thus the plastid is viewed as an auxotrophic resident within the cell during development. Other possibilities for interaction at this and other levels are also discussed.

     

Male age and its association with reproductive traits in captive and wild house sparrows

Evolutionary theory predicts that females seek extra‐pair fertilizations from high‐quality males. In socially monogamous bird species, it is often old males that are most successful in extra‐pair fertilizations. Adaptive models of female extra‐pair mate choice suggest that old males may produce offspring of higher genetic quality than young males because they have proven their survivability. However, old males are also more likely to show signs of reproductive senescence, such as reduced sperm quality. To better understand why old males account for a disproportionally large number of extra‐pair offspring and what the consequences of mating with old males are, we compared several sperm traits of both captive and wild house sparrows, Passer domesticus. Sperm morphological traits and cloacal protuberance volume (a proxy for sperm load) of old and young males did not differ substantially. However, old males delivered almost three times more sperm to the female's egg than young males. We discuss the possibility of a post‐copulatory advantage for old over young males and the consequences for females mated with old males.     

Experimentally prolonged sexual activity in female sparrows delays termination of reproductive activity in their untreated mates

Oestradiol treatment of free-ranging female song sparrows delays the normal, late summer, termination of reproductive behaviour not only in treated females, but also in their untreated mates. Males mated to oestradiol-implanted females continued to respond to experimental playbacks of tape-recorded song after control males had ceased territorial defence. Experimental birds remained paired on their summer breeding territories for up to 3 months after the controls had terminated reproductive activity. Elevated levels of plasma testosterone were maintained well into the autumn in experimental males. The data suggest a major role for behavioural interactions in the temporal modulation of seasonal reproductive activity.     

Role of refractory period in homoclinic models of neural synchronization

We study the properties of a homoclinic model of neuron by introducing a suitable one-dimensional map. We show that the system is characterized by a response time to external signals which is a decreasing function of the signal strength, in contrast to excitable models whose response time is signal-independent. In a one-dimensional array of these systems with bidirectional coupling, we observe a sudden transition to a synchronized state at a certain value of the coupling strength. The transition occurs when the response time of a site to the signals of the adjacent sites is of the order of refractory time. Near the transition, we find an intermittent behavior due to the competition between a turbulent and a synchronized state. The observed behavior distinguishes homoclinic systems from excitable systems.