Nestling diet,secondary sexual traits and fitness in the zebra finch

We examined the effect of nestling diet quality on a suite of physiological, morphological and life-history traits in adult male zebra finches, Taeniopygia guttata. Compared with birds reared on a supplemented diet, nestlings reared on a seed-only diet showed a reduced rate of growth and reduced cell-mediated immune function as measured by an in vivo response to a T lymphocyte-dependent mitogen. There were no differences between birds reared on the two diets in any of the following adult traits: body size, primary sexual traits (testes mass, numbers of stored sperm, sperm function, velocity and morphology), secondary sexual traits (beak colour and song rate), serological traits or immunological traits. The only differences we detected were a lower body mass and a greater proportion of individuals with plumage abnormalities among those reared on a seed-only diet (this latter effect was transient). The fact that male zebra finches reared on a seed-only diet were, as adults, virtually indistinguishable from those reared on a supplemented diet, despite having reduced growth and immune function as nestlings, demonstrates that they subsequently compensated through the differential allocation of resources. Our results indicate that differential allocation is costly in terms of fitness since birds reared on a seed-only diet experienced a significantly greater mortality rate than those reared on a supplemented diet. This in turn suggests the existence of a trade-off between the development of traits important for reproduction, such as primary and secondary sexual traits and longevity.     

Sex-specific differences in compensation for poor neonatal nutrition in the zebra finch Taeniopygia guttata

Individuals can compensate for poor early nutrition by accelerating their growth rates once diet improves, but if malnutrition occurs at a key stage of development only certain body structures may compensate fully. This degrees of compensation is predicted to differ between the sexes and also between species with different life history strategies. In this paper we determine how males and females in a short-lived bird species, with only slight size dimorphism, differ in their abilities to compensate for a poor start in life. Here, zebra finch Taeniopygia guttata chicks from 93 broods were reared on either a standard quality (SQ), or low quality (LQ) diet for the first 15 d of life, followed by a standard diet for the rest of their lives. Thus, the period of nutritional manipulation was relatively short compared with previous studies. Nestlings on a LQ diet showed slower body mass and tarsus growth rates than those on SQ diet but this was reversed after the diet of the LQ birds was improved. LQ birds of both sexes were able to fully compensate in terms of body mass and beak colour, but not tarsus length. Body size and beak colour are sexually selected traits in male zebra finches. By adulthood LQ females had significantly shorter wings than other birds, apparently directing resources into sex-specific structures instead of feather and skeletal growth. Thus, our experiment showed that the sexes differed in how they phenotypically compensated for a poor start in life. Males in particular invested sparse resources into structures associated with mate acquisition, as proposed by life history theory for a species that 'lives fast and dies young.' We predict that this relatively short period of malnutrition during early development will have long term, sex-specific, fitness consequences for these birds.     

State-dependent incubation behaviour in the zebra finch

State dependence is thought to be an important factor in resource allocation decisions, particularly in those decisions relating to parental care. Incubation behaviour is a costly part of avian parental care, and therefore likely to depend on the parent's body condition. We manipulated the body condition of zebra finches, Taeniopygia guttata, during incubation by using prelaying diets of different protein content, and induced birds to lay a similar number of eggs. We tested the hypothesis that birds in better body condition would invest more in incubation than birds in poorer condition when incubating on the same diet. Females that had received a high-protein prelaying diet lost more body mass than those that had received a low-protein diet. They also increased the length of their incubation bouts between early and middle incubation, whereas females that had received a low-protein prelaying diet increased bout length only between middle and late incubation. There were no differences between males from the two diet groups, and males were responsible for a lesser proportion of incubation than females. These results indicate that incubation behaviour is state dependent in female zebra finches, and that birds of different body condition adopt different incubation strategies. We found no differences in incubation duration and hatching success between the two incubation strategies in captivity, but the potential risk of nest predation in the wild may differ. We suggest that only females in good condition can afford to adopt a strategy of increasing bout length early in incubation; females in poorer condition first have to recover their body condition after having produced a clutch. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour       

Patterns and mechanisms of growth of fifth-instar Manduca sexta caterpillars following exposure to low- or high-protein food during early instars.

For many insect herbivores, variation in protein availability is a pervasive part of the environment. I explore how variable protein availability affects growth rates of fifth-instar Manduca sexta caterpillars and how growth is related to behavior and physiology. Groups of larvae were reared on low- or high-protein artificial diets (5.9% and 17.7% casein by dry weight, respectively) and then transferred in the fifth instar to the same or opposite diet. During or after the 24-h period following transfer, I measured growth rate, consumption rate, growth efficiency, midgut proteolytic activity, and masses of midgut contents and tissues. Fifth-instar caterpillars reared in earlier instars on high-protein diet grew about 20% more rapidly over 24 h than did caterpillars reared on low-protein diet. This growth pattern appears to be caused by differences in consumption and growth efficiency: caterpillars reared on high protein consumed more food, and used it more efficiently, than did caterpillars reared on low-protein diet. Over the short term (24 h), in contrast, fifth instars that received low-protein diet grew as rapidly as caterpillars that received high-protein diet. Increased (compensatory) consumption appears to be the primary mechanism by which caterpillars consuming low-protein food maintained growth rates.     

Experimental dissociation of the effects of diet, age and breeding experience on primary reproductive effort in zebra finches Taeniopygia guttata

Reproductive performance varies with age in a wide range of organisms, and increasingly such patterns are interpreted in terms of state-dependent models. We sought to characterise 'state' with regards to age-related variation in clutch size, egg mass and timing of breeding in captive zebra finches Taeniopygia guttata , focusing on the roles of diet quality, age and breeding experience. Females on a high-quality diet laid larger clutches of larger eggs than did females on a low-quality diet. The effect of age on reproductive performance was examined by comparing females breeding (i.e. paired) for the first time at either 3- and/or 6-months of age. Clutch size increased with age but on the low-quality diet only, not on the high-quality diet. Furthermore, clutch size decreased between 3- and 6-months of age in birds bred first on the high-quality diet and then on the low-quality diet. Age did not affect egg mass but older birds had shorter laying intervals. Reproductive performance did not differ between females breeding at 6-months of age for the first or second time: the effects of age were not due to 'training' effects or experience specific to breeding (e.g. undergoing the physiological process of egg formation). In conclusion, nutritional condition (diet) emerged as a central component of state that could strongly influence, and even reverse, any age-dependent increase in primary reproductive performance.     

Basal metabolic rate: heritability and genetic correlations with morphological traits in the zebra finch

Studies of genetic variation in metabolic traits have so far not focused on birds. In our study population of captive zebra finches we found evidence for a significant heritable genetic component in basal metabolic rate (BMR). Heritability of all morphological traits investigated (body mass, head length, tars length and wing length) was significantly larger than zero. All traits were positively phenotypically correlated. Eight of 10 genetic correlations presented in this study differed significantly from zero, all being positive, suggesting the possibility of correlated responses to any selection acting on the traits. When conditioned on the genetic variance in body mass, the heritability of BMR was reduced from 25% to 4%. Hence, our results indicate that genetic changes in BMR through directional selection are possible, but the potential for adaptation independent of body mass may be limited.     

Is clutch size in birds affected by environmental conditions during growth?

Only environmental conditions occurring at the time of breeding have been shown to affect clutch size in birds, even though conditions experienced during growth are known to affect body size or egg size. We show here that environmental conditions experienced during early life can affect clutch size in captive zebra finches (Taeniopygia guttata) and wild great tits (Parus major). Not only do factors outside the immediate breeding season affect clutch size, but clutch size control mechanism is permanently influenced by conditions experienced during ontogeny.     

Female mate choice as a condition-dependent life-history trait

The acquisition of resources is an important determinant of patterns of variation in and covariation among traits that are costly to produce and are dependent on condition for their expression. However, the extent to which variation in female mate choice behavior is condition dependent, and how this is related to other life-history traits, remains largely unknown. We manipulated the acquisition of dietary protein in the black field cricket, Teleogryllus commodus, and measured the effects of this on several important life-history traits and on female mate choice behavior. Females reared on a high-protein diet developed faster, were heavier at eclosion, and lived longer than females reared on a low-protein diet. Two lines of evidence suggest that female mate choice behavior in T. commodus is condition dependent. First, females reared on the high-protein diet were more sexually responsive and expressed stronger linear and quadratic preference functions for call rate and dominant frequency, respectively. Second, within treatments, females that developed faster were lighter, generally less sexually responsive, and, in the high-protein-diet treatment, expressed weaker preferences than slower-developing females. Collectively, our findings suggest an important role for resource acquisition in generating variation in mate choice behavior.     

Experimental (antiestrogen-mediated) reduction in egg size negatively affects offspring growth and survival

The relationship between egg size and offspring phenotype is critical to our understanding of the selective pressures acting on the key reproductive life-history traits of egg size and number. Yet there is surprisingly little empirical evidence to support a strong, positive relationship between egg size and offspring quality (i.e., offspring growth, condition, and survival) in birds, in part because of confounding effects of parental quality and the lack of experimental techniques for directly manipulating avian egg size independently of maternal condition. Previously, we showed that treatment of laying female zebra finches (Taeniopygia guttata) with the antiestrogen tamoxifen can decrease egg size by ca. 8% but that this reduction in egg size had few effects on offspring mass and size at fledging. Here, we extend the use of this technique to induce larger decreases in egg size (up to 50% in individual females) and show that a reduction in egg size of ca. 18% is associated with decreased embryo viability, increased hatchling mortality, and lower posthatching offspring survival. Furthermore, we show that although hatchlings from eggs reduced in size by ca. 9% can survive to fledging, these chicks show slower initial growth during the linear growth phase (5-10 d of age), fledge at lower masses than chicks from control eggs, and show postfledging compensatory growth. Our results provide empirical support for significant effects of egg size on offspring quality and further suggest that among individual females there is a minimum egg size required to maintain embryo viability and offspring quality.     

Song learning with audiovisual compound stimuli in zebra finches

We investigated the effects of audiovisual compound training on song learning in zebra finches, Taeniopygia guttata. In the first experiment, presentation of a stuffed adult zebra finch male was found to be reinforcing to zebra finch males in an operant task. In a separate experiment, zebra finch males were reared without their father from day 7 after hatching onwards. Between 35 and 76 days, they were placed in isolation and exposed to taped songs of a zebra finch male, according to a random schedule (20 presentations/h). For half of the birds, presentation of the song coincided with presentation of a stuffed zebra finch male. For the remaining birds, each presentation of the song was followed by presentation of a stuffed male. The birds were subsequently isolated until day 142, when their own songs were recorded and analysed. Birds in both groups shared significantly more song elements with their tutor songs than with an unfamiliar song. There was no significant difference in song learning between the groups. These results confirm that zebra finches can learn part of their songs from taped tutor songs. Furthermore, simultaneous presentation of the tutor song and a relevant, salient visual stimulus is not superior to sequential presentation. Copyright 1999 The Association for the Study of Animal Behaviour.     

No heightened condition dependence of zebra finch ornaments – a quantitative genetic approach

The developmental stress hypothesis offers a mechanism to maintain honesty of sexually selected ornaments, because only high quality individuals will be able to develop full ornamentation in the face of stress during early development. Experimental tests of this hypothesis have traditionally involved the manipulation of one aspect of the rearing conditions and an examination of effects on adult traits. Here, we instead use a statistically powerful quantitative genetic approach to detect condition dependence. We use animal models to estimate environmental correlations between a measure of early growth and adult traits. This way, we could make use of the sometimes dramatic differences in early growth of more than 800 individually cross‐fostered birds and measure the effect on a total of 23 different traits after birds reached maturity. We find strong effects of environmental growth conditions on adult body size, body mass and fat deposition, moderate effects on beak colour in both sexes, but no effect on song and plumage characters. Rather surprisingly, there was no effect on male attractiveness, both measured in mate choice trials and under socially complex conditions in aviaries. There was a trend for a positive effect of good growth conditions on the success at fertilizing eggs in males breeding in aviaries whereas longevity was not affected in either sex. We conclude that zebra finches are remarkably resilient to food shortage during growth and can compensate for poor growth conditions without much apparent life‐history trade‐offs. Our results do not support the hypothesis that sexually selected traits show heightened condition dependence compared to nonsexually selected traits.     

中国鸟类的生活史和生态学特征数据集

中国现有1,445种鸟类, 是世界上鸟类物种数最多的国家之一。物种特征反映了生物有机体的功能和适合度, 在生态学、进化生物学和保护生物学研究中具有重要作用。但是, 目前还没有关于我国鸟类生活史、生态学和地理分布等物种特征的完整数据库。通过系统查阅文献和各种数据资料, 本文共收集整理出了中国1,445种鸟类17个功能特征数据: 体重、体长、嘴峰长、翅长、尾长、跗蹠长、食性、窝卵数、卵大小、卵体积、巢址、巢的类型、集群状况、迁徙状况、是否特有种、地理分布范围和分布省份等。在这些特征中, 除迁徙状况、是否特有种、地理分布范围和分布省份外, 其余特征数据均存在不同程度的缺失, 数据的完整度为60.83%‒100%。本数据库是目前关于中国鸟类最新和最全的物种特征数据库, 期望能为我国鸟类生态学、进化生物学、生物地理学、保护生物学等研究提供支持。     

Anhydrolutein in the zebra finch: a new,metabolically derived carotenoid in birds

Many birds acquire carotenoid pigments from the diet that they deposit into feathers and bare parts to develop extravagant sexual coloration. Although biologists have shown interest in both the mechanisms and function of these colorful displays, the carotenoids ingested and processed by these birds are poorly described. Here we document the carotenoid-pigment profile in the diet, blood and tissue of captive male and female zebra finches (Taeniopygia guttata). Dietary carotenoids including: lutein; zeaxanthin; and β-cryptoxanthin were also present in the plasma, liver, adipose tissue and egg-yolk. These were accompanied in the blood and tissues by a fourth pigment, 2′,3′-anhydrolutein, that was absent from the diet. To our knowledge, this is the first reported documentation of anhydrolutein in any avian species; among animals, it has been previously described only in human skin and serum and in fish liver. We also identified anhydrolutein in the plasma of two closely related estrildid finch species (Estrilda astrild and Sporaeginthus subflavus). Anhydrolutein was the major carotenoid found in zebra finch serum and liver, but did not exceed the concentration of lutein and zeaxanthin in adipose tissue or egg yolk. Whereas the percent composition of zeaxanthin and β-cryptoxanthin were similar between diet and plasma, lutein was comparatively less abundant in plasma than in the diet. Lutein also was proportionally deficient in plasma from birds that circulated a higher percentage of anhydrolutein. These results suggest that zebra finches metabolically derive anhydrolutein from dietary sources of lutein. The production site and physiological function of anhydrolutein have yet to be determined.     

Prenatal developmental conditions have long-term effects on offspring fecundity

Maternal effects, in which differences in parental state cause differences in offspring fitness, are important in trade-offs influencing an individual's optimal reproductive strategy. In zebra finches (Taeniopygia guttata) we manipulated the nutritional state for four weeks before the start of breeding through protein supplementation. Zebra finches were kept on identical diets during the rest of the experiment. We then tested the effects of maternal state on offspring size, survival and fecundity. In order to separate the effects of maternal state occurring through egg production, incubation and chick-rearing, we used a cross-fostering experiment. We show that a protein-rich diet prior to laying improved maternal body weight prior to breeding compared with birds on a protein-poor diet. Poorer maternal state prior to breeding gave rise to offspring with lower fecundity than offspring from birds in a better nutritional state. Maternal state is thought to affect the conditions developing offspring experience through the bird's ability to produce and incubate eggs. Male and female embryos differed in their responses to conditions at different developmental stages. This shows that embryonic developmental conditions and sex differences in vulnerability to these conditions need to be incorporated into future models of selection, life-history evolution and sex-ratio theory.     

Compensatory growth impairs adult cognitive performance

Several studies have demonstrated that poor early nutrition, followed by growth compensation, can have negative consequences later in life. However, it remains unclear whether this is attributable to the nutritional deficit itself or a cost of compensatory growth. This distinction is important to our understanding both of the proximate and ultimate factors that shape growth trajectories and of how best to manage growth in our own and other species following low birth weight. We reared sibling pairs of zebra finches on different quality nutrition for the first 20 d of life only and examined their learning performance in adulthood. Final body size was not affected. However, the speed of learning a simple task in adulthood, which involved associating a screen colour with the presence of a food reward, was negatively related to the amount of growth compensation that had occurred. Learning speed was not related to the early diet itself or the amount of early growth depression. These results show that the level of compensatory growth that occurs following a period of poor nutrition is associated with long-term negative consequences for cognitive function and suggest that a growth-performance trade-off may determine optimal growth trajectories.     

Inheritance of body size in the Barnacle Goose under different environmental conditions

Heritabilities, genetic variances and covariances for body size traits, i.e. tarsus length, head length and body mass, were estimated under different environmental conditions in a Barnacle Goose (Branta leucopsis) population. Under poor growth conditions, that is, when average body size of fully grown offspring in a given cohort was small, the offspring-parent regressions and full-sib analyses yielded heritability estimates not significantly different from zero. By contrast, when growth conditions were normal or good the heritability estimates were generally significantly positive. Comparisons of genetic covariance estimates indicated that they also differed across the analysed environmental conditions. This result, together with similar results obtained in studies of passerine birds, suggests that genotype-environment interactions might be frequent within the range of environments normally encountered by birds in natural populations. If general, such results might question the validity of assuming approximate constancy of additive genetic variances and covariances over time and environments in evolutionary models.     

Use of labelled methionine to investigate the contribution of muscle proteins to egg production in zebra finches

Many bird species show a loss of female muscle mass at the time of egg formation. In this study we investigated whether there was a causal link between the loss in muscle condition and the formation of egg proteins by feeding ³⁵S-methionine to female zebra finches to label muscle proteins. When these birds subsequently bred the isotope was transferred to the egg proteins: isotope loss from female muscle tissue was significantly greater in birds which had bred than in control groups which had not.     

Nest destruction elicits indiscriminate con‐ versus heterospecific brood parasitism in a captive bird

Following nest destruction, the laying of physiologically committed eggs (eggs that are ovulated, yolked, and making their way through the oviduct) in the nests of other birds is considered a viable pathway for the evolution of obligate interspecific brood parasitism. While intraspecific brood parasitism in response to nest predation has been experimentally demonstrated, this pathway has yet to be evaluated in an interspecific context. We studied patterns of egg laying following experimental nest destruction in captive zebra finches, Taeniopygia guttata, a frequent intraspecific brood parasite. We found that zebra finches laid physiologically committed eggs indiscriminately between nests containing conspecific eggs and nests containing heterospecific eggs (of Bengalese finches, Lonchura striata vars. domestica), despite the con‐ and heterospecific eggs differing in both size and coloration. This is the first experimental evidence that nest destruction may provide a pathway for the evolution of interspecific brood parasitism in birds.     

Renal response to dietary protein in the house sparrow Passer domesticus

Many birds switch seasonally or during ontogeny between diets of varying protein content. In mammals, high-protein diets induce hypertrophy of the kidney in general and of the thick ascending limbs (TAL) in particular, along with increases in glomerular filtration rate (GFR) and urine flow. A hypothesis to explain these phenomena is that the TAL become increasingly sensitive to peptide hormones (glucagon and antidiuretic hormone [ADH]) released in response to protein feeding; the consequent enhancement of ion reabsorption dilutes urine reaching the macula densa, thereby suppressing tubulo-glomerular feedback (TGF) and causing a rise in GFR. Avian kidneys possess most of the elements involved in this mechanism, including loops of Henle with TAL, sensitivity of TAL to ADH (arginine vasotocin [AVT] in birds), and the elements of TGF. We therefore hypothesized that switching from a low-protein to a high-protein diet would induce responses in birds similar to those found in mammals. We tested this hypothesis by feeding house sparrows, Passer domesticus, isocaloric diets containing either 8% or 30% protein. Birds on high-protein food had larger renal medullae, both in mass and in TAL diameter, but no increase in whole-kidney mass. Urine flow was approximately doubled on high-protein food, but there was no change in GFR. We were not able to detect an increased sensitivity of AVT-induced adenylyl cyclase activity in TAL from high-protein animals, and responsiveness to glucagon was higher in TAL from birds eating low-protein food. We are unable to conclude that a suppression of TGF is responsible for the rise in urine flow in birds eating high-protein foods, and the mechanisms behind the medullary hypertrophy and the diuresis remain to be fully explored.     

Haematocrit is weakly related to condition in nestling Barn Swallows Hirundo rustica

Currently, there is no agreement about the suitability of haematocrit (the proportion of blood volume occupied by packed red blood cells) as a predictor of condition in birds. In order to clarify this point, genetic and environmental components of phenotypic variation for a number of traits were estimated in nestling Barn Swallows Hirundo rustica by means of a partial cross-fostering experiment. The studied traits were haematocrit, two morphological traits used as estimates of condition in birds (body mass and body mass relative to body size) and a morphological trait that presumably is not associated with condition (tarsus length). First, we found that body mass relative to body size was related to haematocrit, supporting the suggestion that haematocrit reflects condition in this species. Secondly, we found that the coefficient of residual (i.e. environmental) variation of haematocrit was larger than that of tarsus length, but smaller than those of body mass or body mass relative to body size. Under the hypothesis that traits closely related to condition (and, ultimately, to fitness) are characterized by large residual variance, these results also suggest that haematocrit is related to condition, but that this relationship must be weak. Therefore, the use of haematocrit as an estimate of condition is not recommended. Finally, heritabilities of the studied traits were calculated, differing significantly from zero only for tarsus length, the trait with the smallest residual variation. However, a consistent pattern in the relationship between heritability and genetic variation was not found.