Internal magnesium, 2,3-diphosphoglycerate, and the regulation of the steady-state volume of human red blood cells by the Na/K/2Cl cotransport system.

This study is concerned with the relationship between the Na/K/Cl cotransport system and the steady-state volume (MCV) of red blood cells. Cotransport rate was determined in unfractionated and density-separated red cells of different MCV from different donors to see whether cotransport differences contribute to the difference in the distribution of MCVs. Cotransport, studied in cells at their original MCVs, was determined as the bumetanide (10 microM)-sensitive 22Na efflux in the presence of ouabain (50 microM) after adjusting cellular Na (Nai) and Ki to achieve near maximal transport rates. This condition was chosen to rule out MCV-related differences in Nai and Ki that might contribute to differences in the net chemical driving force for cotransport. We found that in both unfractionated and density-separated red cells the cotransport rate was inversely correlated with MCV. MCV was correlated directly with red cell 2,3-diphosphoglycerate (DPG), whereas total red cell Mg was only slightly elevated in cells with high MCV. Thus intracellular free Mg (Mgifree) is evidently lower in red cells with high 2,3-DPG (i.e., high MCV) and vice versa. Results from flux measurements at their original MCVs, after altering Mgifree with the ionophore A23187, indicated a high Mgi sensitivity of cotransport: depletion of Mgifree inhibited and an elevation of Mgifree increased the cotransport rate. The apparent K0.5 for Mgifree was approximately 0.4 mM. Maximizing Mgifree at optimum Nai and Ki minimized the differences in cotransport rates among the different donors. It is concluded that the relative cotransport rate is regulated for cells in the steady state at their original cell volume, not by the number of copies of the cotransporter but by differences in Mgifree. The interindividual differences in Mgifree, determined primarily by differences in the 2,3-DPG content, are responsible for the differences in the relative cotransport activity that results in an inverse relationship with in vivo differences in MCV. Indirect evidence indicates that the relative cotransport rate, as indexed by Mgifree, is determined by the phosphorylated level of the cotransport system.     

Sexual character intensity and its relationship to breeding timing, fecundity and mate choice in the great cormorant Phalacrocorax carbo lucidus

Reproductive success and the intensity of five separate seasonal sexual characters, expressed similarly in both sexes and likely to be dependent on age and/or individual condition, were documented at a colony of great cormorants at Lake Naivasha, Kenya (00°46' S, 36°22' E) in 1996. The characters assessed were head filoplumes, thigh patch feathers, colour of cheek, neck and upper breast plumage, colour of gular skin, and colour of suborbital skin. More intensely developed sexual characters at the time of pair formation were associated with significantly earlier breeding. Pairs that bred earlier fledged significantly more chicks than those breeding later. However, correlations between the number of chicks fledged and the intensity of the parents' sexual characters at pair formation were generally weak and not significant. An exception was the colour of the cheek, foreneck and upper breast plumage: males (but not females) with the darkest plumage in these areas fledged significantly more chicks than others. Correlations between paired male and female sexual character intensity were weak, and significant only in the case of suborbital skin colour. Other factors may have been more important than these characters in influencing mate selection.     

Haematocrit values of breeding Red-billed queleas Quelea quelea (Aves: Ploceidae) in relation to body condition and thymus activity

In a Red-billed quelea colony in north-eastern Nigeria that was later abandoned as a result of the poor nutritional condition of the adults, laying females showed low haematocrit values compared with normal pre-breeding haematocrit levels but nestbuilding males did not. In females the low haematocrits were correlated with extremely low levels of reserve lipid and labile protein but males in equally poor condition did not have such low haematocrits. This difference must result in some way from the additional nutritional burden for females of mobilizing sufficient protein to form eggs. Earlier observations of massive red blood cell production in the thymus of queleas during incubation suggest that low haematocrits, indicating anaemia, may be a normal occurrence in queleas during breeding, and were not peculiar to this abnormal abandoned colony.     

The growth bars on tail feathers in the male Styan's Grasshopper Warbler may indicate quality

We propose that growth bar size may indicate the quality of individuals in Styan's Grasshopper Warblers Locustella pleskei . Positive correlations of an individual's standardized growth bar width between years showed that some individuals of both sexes were usually in better condition than others. The survival rate of males with wider growth bars was higher than that of other males. These survivors might have survived better because of a better body condition. We also examined the relationships between growth bar width and reproductive parameters (arrival date, pairing date and reproductive success). Males with wider growth bars arrived earlier, acquired a mate earlier and achieved higher total reproductive success by breeding twice. In females, growth bar width was correlated with reproductive parameters (arrival date, pairing date and total reproductive success), but not with survival. These results suggest that males with wider growth bars were of higher quality than those with narrower bars. The contrasting results for the two sexes may be explained by differences in their breeding tactics.     

Sexual size dimorphism and positive assortative mating in red-backed shrike Lanius collurio: an adaptive value?

During field studies in 1997–1999 in South Bohemia (Czech Republic), we found significant differences in size between the sexes in a local breeding population of red-backed shrike Lanius collurio. Males were significantly larger than females for wing length and tarsus length, but had smaller body mass than females. However, there was considerable overlap in the ranges of these parameters between the sexes. Interestingly, pairs were formed at random with respect to wing length and tarsus length, but assortative mating was significant for body mass/body condition. Among tested variables, only male wing length correlated significantly with nestling body mass at day 7. However, clutch size and the number of fledglings strongly depended on differences in tarsus length between mates, but not on body size of mates. Individual improvements in foraging skills and/or courtship feeding rates are proposed as possible explanations for these findings.     

Correlations between body size, defensive behaviour and reproductive success in male Little Blue Penguins Eudyptula minor: implications for female choice

Field studies exploring relationships among body size, defensive behaviour (including vocalizations) and reproductive success in male Little Blue Penguins Eudyptula minor were conducted to identify behavioural and physical attributes that might provide females or opponents with information on male quality. Head length provided a simple estimator of body size in both sexes. Nests defended by large males produced eggs and chicks earlier in the breeding season, and contained chicks that grew more quickly than chicks in nests defended by small males. In contrast, female body size did not affect any of the reproductive parameters we measured. The pitch of male defensive calls varied significantly with male size, so females (or opponents) could potentially gauge male size by assessing the calls. Large males were also more likely to respond vocally to simulated nest intrusions. As a result, by 'keeping company' with prospective partners early in the breeding season, females can obtain information about males that would allow them to select better partners and improve their own reproductive potential.     

The effect of parental age,experience and historical reproductive success on wandering albatross (Diomedea exulans) chick growth and survival

Growth and survival of altricial young are influenced by their parents’ abilities to invest in a breeding attempt. As a result, chick growth and survival in one breeding season may be indicative of their parents’ long-term reproductive potential. To determine whether variation in long-term reproductive success is driven by differential breeding investment, parental care and chick growth in wandering albatrosses (Diomedea exulans) were correlated with parental historical reproductive success. Effects of age and breeding experience (determined from past breeding attempts) and pre-laying body condition (mass–size indices) on chick growth and survival also were tested. Longer brooding of chicks increased their survival, but length of chick brooding did not differ between historically unproductive and successful breeders. Past reproductive success also was not correlated with chick growth rates or fledging mass or size. Chick brooding period, chick growth rates, final mass and size were independent of parental body condition. Older and more experienced parents brooded chicks for longer and their chicks grew faster, supporting previous findings that breeding competence is a learnt skill. Chick care and growth characteristics differed more between than within pairs, suggesting that differences in these characteristics are driven by variation among pairs.     

Sex-Based Differences in Adélie Penguin (Pygoscelis adeliae) Chick Growth Rates and Diet

Sexually size-dimorphic species must show some difference between the sexes in growth rate and/or length of growing period. Such differences in growth parameters can cause the sexes to be impacted by environmental variability in different ways, and understanding these differences allows a better understanding of patterns in productivity between individuals and populations. We investigated differences in growth rate and diet between male and female Adélie Penguin (Pygoscelis adeliae) chicks during two breeding seasons at Cape Crozier, Ross Island, Antarctica. Adélie Penguins are a slightly dimorphic species, with adult males averaging larger than adult females in mass (~11%) as well as bill (~8%) and flipper length (~3%). We measured mass and length of flipper, bill, tibiotarsus, and foot at 5-day intervals for 45 male and 40 female individually-marked chicks. Chick sex was molecularly determined from feathers. We used linear mixed effects models to estimate daily growth rate as a function of chick sex, while controlling for hatching order, brood size, year, and potential variation in breeding quality between pairs of parents. Accounting for season and hatching order, male chicks gained mass an average of 15.6 g d^-1 faster than females. Similarly, growth in bill length was faster for males, and the calculated bill size difference at fledging was similar to that observed in adults. There was no evidence for sex-based differences in growth of other morphological features. Adélie diet at Ross Island is composed almost entirely of two species—one krill (Euphausia crystallorophias) and one fish (Pleuragramma antarctica), with fish having a higher caloric value. Using isotopic analyses of feather samples, we also determined that male chicks were fed a higher proportion of fish than female chicks. The related differences in provisioning and growth rates of male and female offspring provides a greater understanding of the ways in which ecological factors may impact the two sexes differently.     

The breeding season duration hypothesis: acute handling stress and total plasma concentrations of corticosterone and androgens in male and female striped plateau lizards (Sceloporus virgatus)

Acute glucocorticoid elevations can be adaptations to short-term stressors. The breeding season hypothesis predicts reduced glucocorticoid responsiveness to acute stressors in populations or species with short breeding seasons. The striped plateau lizard (Sceloporus virgatus) has a short breeding season in Arizona. We measured plasma corticosterone and total androgen levels (dihydrotestosterone and testosterone) following one of the four stress-handling treatments (0, 10, 60, or 180 min). In both sexes, longer handling stress yielded higher corticosterone; females had higher corticosterone than males at all time points. Androgens did not vary with handling duration, in either sex. Combining treatments, plasma androgens correlated positively with corticosterone (CORT) in females but not in males; plasma CORT and body mass residuals were negatively correlated in both sexes, suggesting lizards in poor body condition and/or not investing heavily in reproduction (follicle mass) have higher acute corticosterone. Total plasma androgens and body mass residuals were positively associated in males, but showed no association in females. The maximal CORT elevation after handling stress in this single-clutching species was of comparable magnitude to responses in related multi-clutching lizard species with longer breeding seasons. Using data from studies of multiple populations of three Sceloporus species, we found no relationship between the relative magnitude of the CORT increase and either latitude or elevation, two variables in the literature correlated with duration of the breeding season, and only weak relationships with geographic elevation and actual (not relative) stress-elevated CORT values in this multi-population comparison.     

Changes in hematological parameters in free-living pigeons (Columba livia f. urbana) during the breeding cycle

The blood parameters—red blood cell (RBC) count, hematocrit (Ht), hemoglobin concentration (Hb), mean corpuscular volume (MCV), mean corpuscular hemoglobin (MCH), and mean corpuscular hemoglobin concentration (MCHC)—were studied for the first time in free ranging pigeons Columba livia f. urbana. The aim of the study was to establish a set of reference values for this ubiquitous urban species and to examine a potential influence of the factors such as sex and phase of the breeding cycle on the hematological parameters. Blood was sampled from the adult individuals of both sexes during the breeding cycle. Significant variations in Ht and Hb were observed, while RBC count remained unchanged. The lower hematocrit value and hemoglobin concentration found in both males and females during molt probably resulted from the increase of plasma volume not accompanied by an increase in RBC. On the other hand, a significant physiological stress of nutritional deficiency during this energy-consuming stage may manifest itself in the decreased Ht and Hb values. The higher values of Ht and Hb during the laying phase could be related to the dehydration status. There were no overall differences in MCV, MCH, and MCHC values in either of the breeding stage. A lack of sex dimorphism in the measured and calculated blood parameters was apparent. Data collected herein contribute to the better understanding of general physiology of the ubiquitous urban species. The knowledge of variation in the blood indexes with respect to sex and breeding stage is crucial before blood parameters can provide an useful information on physical condition of the bird.     

Spawning colouration,parasites and habitat selection in Salvelinus alpinus: initiating speciation by sexual selection?

The relationship between parasite infection and breeding ornamentation in an Arctic charr Salvelinus alpinus population was examined. Interrelationships between the intensities of parasites with different dietary transmission routes suggested that hosts were segregated by habitat and diet preferences. That is, intensities of parasites transmitted through intermediate hosts that share habitat were, in both sexes, positively interrelated. Negative interrelationships in the intensities of parasites transmitted through different hosts were found only in females. The intensity of breeding colouration was positively correlated with intensities of amphipod‐transmitted parasites in both sexes. Intensities of parasites transmitted through copepods showed no relationship to morphological or ornamental characters. Seen in the context of morphology and ornaments of previously described sympatric populations of Arctic charr, these relationships suggested that individual variation in niche background may provide a source for the initial variation in morphology and secondary sexual traits leading to sympatric speciation through sexual selection.     

Effects of experimental tail shortening on the phenotypic condition of barn swallows Hirundo rustica: implications for tail‐length evolution

Some studies have suggested that tail streamers in the barn swallow Hirundo rustica may have been elongated 10–12 mm by sexual selection, but according to other studies, the length of these feathers is at the aerodynamic optimum or very close to it. To shed light on this issue, outermost tail feathers were experimentally shortened in male and female barn swallows by 1, 11 or 21 mm. Changes in four physiological parameters commonly used to estimate phenotypic condition in birds (weight, erythrocyte sedimentation rate, blood leukocyte concentration and heterophil/lymphocyte ratio) were checked one month later. Health improved (blood leukocyte concentration decreased) in the group of birds with tails shortened by 11 mm (both males and females), but body condition deteriorated (weight decreased) compared to the other two experimental groups. There was no significant effect of tail‐length manipulation on the other two physiological parameters. These contradictory results suggest trade‐offs between components of phenotypic condition. Possible negative relationships between condition‐related traits imply that using one or very few physiological parameters to estimate phenotypic condition might not be appropriate. The most plausible explanation for the turning point in phenotypic condition when streamers were shortened by 11 mm is that these feathers are 7–15 mm longer than the aerodynamic optimum in both sexes. Therefore, our results are consistent with the hypothesis that tail streamers have been elongated 10–12 mm by sexual selection. This conclusion disagrees with a previous study on the effect of experimental tail shortening on haematocrit, but the complexity of interpreting changes in haematocrit might account for this discrepancy.     

Temporal but not spatial environmental variation drives adaptive offspring sex allocation in a plural cooperative breeder

Cooperatively breeding birds have been used frequently to study sex allocation because the adaptive value of the sexes partly depends upon the costs and benefits for parents of receiving help. I examined patterns of directional sex allocation in relation to maternal condition (Trivers-Willard hypothesis), territory quality (helper competition hypothesis), and the number of available helpers (helper repayment hypothesis) in the superb starling, Lamprotornis superbus, a plural cooperative breeder with helpers of both sexes. Superb starlings biased their offspring sex ratio in relation to prebreeding rainfall, which was correlated with maternal condition. Mothers produced relatively more female offspring in wetter years, when they were in better condition, and more male offspring in drier years, when they were in poorer condition. There was no relationship between offspring sex ratio and territory quality or the number of available helpers. Although helping was male biased, females had a greater variance in reproductive success than males. These results are consistent with the Trivers-Willard hypothesis and suggest that although females in most cooperatively breeding species make sex allocation decisions to increase their future direct reproductive success, female superb starlings appear to base this decision on their current body condition to increase their own inclusive fitness.     

Chick growth and survival in northern lapwings Vanellus vanellus indicate that secondary females do the best of a bad job

The polygyny threshold model predicts that monogamous and secondary females on average settle at the same time and have similar reproductive success. This is not generally found. Incorporating varying female competitive strength into the model, changes the predictions to state that secondary females should breed later and show a reduced success compared to that of monogamous and primary females. We examined if this was the case by investigating growth and survival in chicks of northern lapwings Vanellus vanellus from mothers of monogamous, primary and secondary mating status. Chicks where monitored from hatching to the age of 15–18 d. Growth and survival in secondary chicks was lower than in monogamous and primary chicks. Primary chicks survived significantly better than secondary chicks. Survival of monogamous chicks was comparable to primary chicks and close to significantly higher than in secondary chicks (p = 0.086). Among surviving chicks, daily weight gain in monogamous chicks was significantly higher than in secondary chicks. Growth rates of primary chicks were comparable to monogamous chicks and tended to be higher than in secondary chicks (p = 0.11). Monogamous and primary females both bred significantly earlier than secondary females, and chick survival and body‐mass growth decreased significantly with hatching date. Given the premium on early breeding in lapwings, secondary females appeared to do the best of a bad job, and their later onset of breeding could have been caused by poorer condition and/or lower breeding experience. Additional costs might also have accrued from sharing breeding resources with primary females that presumably were stronger competitors.     

Haematocrit correlates with tail ornament size in three populations of the Barn Swallow (Hirundo rustica)

1. Handicap models of sexual selection propose that male ornaments are indicators of male quality and that honesty is enforced by the costs imposed by the exaggerated ornamental traits. In long-distance migratory birds that feed on the wing, the aerodynamic cost of exaggerated ornamental characters should be particularly high because the size of the ornaments deviates from the natural selection optimum. During migration, birds are expected to raise their oxygen consumption in relation to the energetic demands imposed by their morphology. An increase of haematocrit is an adaptive response to enhance oxygen uptake and efficiency of transfer to the muscular tissues during spells of intense muscular activity.
2. The change of haematocrit of Barn Swallows ( Hirundo rustica ) after their arrival to the breeding sites, and the relationships between haematocrit values recorded after migration and the size of ordinary and sexually selected morphological characters in three Barn Swallow populations were analysed.
3. Males had higher haematocrit values than females. Individual haematocrit values declined after arrival to the breeding sites. Haematocrit values of males were significantly and positively correlated with the size of their ornamental tail but not correlated with other characters, thus suggesting that well-ornamented males, in order to arrive early, have to raise their haematocrit above the level of short-tailed males.
4. Males and females of similar tail length did not differ in their haematocrit, thus suggesting that sexual dimorphism in haematocrit might be functionally related to dimorphism in tail length.
5. Our results are consistent with the handicap principle because long-tailed males experience lower mortality and larger seasonal reproductive success compared with short-tailed males.     

Annual productivity and individual female reproductive success in a Great Bustard Otis tarda population

The reproductive success of Great Bustards Otis tarda in north-western Spain was studied between 1987 and 1998, both at the population ( c . 700 adult females breeding in our study area) and the individual level (sample of 32 marked females). Overall productivity was low, with a population mean of only 0.14 chicks reared per adult female, and an average breeding success of 0.15 chicks per year in the sample of marked females, but interannual variability was high (0.04–0.29). Population productivity was positively correlated with winter (October–March) precipitation prior to each breeding season, and negatively correlated with the number of days of rain during the hatching period. High annual productivity resulted from a high proportion of females rearing two chicks. Reproductive success was higher in females older than 6 years than in younger birds. The proportion of females in the marked sample that failed in breeding after having bred successfully the previous season was significantly higher than the proportion of those that did not. Finally, females with a higher than average breeding success tended to breed successfully in years of both low and high population productivity, whereas those with lower than average breeding success did so only in years of high productivity.     

Size of ornament is negatively correlated with baseline corticosterone in males of a socially monogamous colonial seabird

The Goymann–Wingfield model predicts that glucocorticoid levels in social animals reflect the costs of acquiring and maintaining social status. The crested auklet is one of the few avian colonial species where a mutual ornament in males and females is used in both sexual and aggressive displays. Previous studies of the crested auklet support the notion that the crest ornament is a badge of status in this species. Here, we examined the relationship between the crest ornament size and the adrenocortical function in breeding crested auklets. Crest length was negatively correlated with corticosterone at baseline in males, but not in females. Baseline corticosterone in females (but not in males) was negatively correlated with body condition index. Although male and female crested auklets are monomorphic in their ornamental traits, our results suggest that the socially mediated physiological costs associated with status signaling may differ between the sexes.     

Notes on the behavioural ecology of the Magnificent Frigatebird Fregata magnificens

The Magnificent Frigatebird population breeding on Man-o-War Cay, Belize in 1980 laid their single eggs between mid-December and early April, and had a mean hatching date of mid- to late April. There were no differences between the sexes in the percent of time spent incubating eggs or brooding small chicks (males 48%, females 52%). However, males were not observed to feed larger (75–100 d) chicks. All 81 feedings of juvenile (approximately 1 year old) frigatebirds observed during 126 hr of observations were by females, and the majority of these feedings occurred at midday (46%) when approximately equal numbers of females and juveniles frequented the study area. Our data confirm Diamond's (1972, 1973) reports of unequal roles in chick rearing and post-fledging care between male and female F. magnificens , and support the hypothesis that males and successful females may breed at one- and two-year intervals, respectively.     

Corticosterone responses of grey-faced petrels (Pterodroma macroptera gouldi) are higher during incubation than during other breeding stages

We report the results of the first field study examining seasonal changes in corticosterone responses of typically long-lived birds of the order Procellariiformes. In particular, we examined whether grey-faced petrels Pterodroma macroptera gouldi showed changes in circulating baseline corticosterone concentrations and corticosterone responses to a standardized handling protocol across the breeding season. Such changes have been associated with changes in body condition and variations in energy demands on adult birds through the breeding season. During early incubation, males were in significantly better condition than females that had just completed laying, whereas during late incubation, males were in significantly poorer condition than females. In spite of these differences, there was no significant difference in baseline corticosterone concentrations between sexes or among birds at different reproductive stages. However, we detected significant differences in corticosterone responses associated with a standardized handling protocol at different stages through the breeding season. Responses were significantly greater during incubation compared with the prelay period and late chick rearing. Body condition was weakly and negatively correlated with maximum and total integrated corticosterone level, indicating that some of the individual variability in stress corticosterone responses could be explained by variation in body condition. However, the largest stress response occurred during late incubation and was independent of sex, although males were in relatively poor condition and females in relatively good condition. This period coincided with the breeding stage in which energy constraints on individual adults were higher than at other periods of the reproductive cycle and birds may be physiologically primed for extended fasts.     

The influence of habitat management on the breeding success of the Great Bittern Botaurus stellaris in Britain

Great Bitterns Botaurus stellaris have experienced a population decline in the UK, such that in 1997 the total number of breeding males was just 11. This study aimed to identify factors affecting productivity, and how management could be used to manipulate this. An intensive study of Great Bittern breeding success was conducted between 1997 and 2001. The date that males established their booming territories was closely correlated with when females started nesting. Wetter sites with greater fish densities had males that established their booming territories earlier in the season. However, only the date that males started booming determined when females started nesting. The mean clutch size of Great Bitterns was four and the only cause of nest failure was predation. Of eight nests suspected to be second attempts the mean interval between these first and second attempts was 12.25 ± 0.88 (sd) days (range 8–15). The fate of radiotagged chicks followed to fledging revealed that the overall probability of a chick surviving to fledge was 39.1%. Daily losses of Great Bittern chicks due to starvation/exposure accounted for 76.25% and predation 21.25%. The youngest chicks in poor condition were most likely to die, particularly in periods of high rainfall. A simulated renesting model allowed estimation of Great Bittern productivity as 1.24 chicks per female and 1.52 nesting attempts per female. Habitat management, or lack of water control that resulted in sites being drier in spring, delayed nesting, although statistically there was no difference in productivity compared with wetter sites.