Costly steroids: egg testosterone modulates nestling metabolic rate in the zebra finch

The transfer of non-genetic resources from mother to the offspring often has considerable consequences for offspring performance. In birds, maternally derived hormones are known to influence a variety of morphological, physiological and behavioural traits in the chick. So far, the range of these hormonal effects involves benefits in terms of enhanced growth and competitive ability as well as costs in terms of immunosuppression. However, since yolk hormones can enhance growth and begging activity, high levels of these hormones may also involve energetic costs. Here, we show experimentally that elevated levels of prenatal testosterone increase resting metabolic rate in nestling zebra finches (Taeniopygia guttata). Surprisingly, however, elevation of prenatal testosterone did not result in higher growth rates and, thus, differences in resting metabolism do not seem to be linked to nestling growth. We conclude that apart from immunosuppressive effects, high levels of egg steroids may also entail costs in terms of increased energy expenditure.     

Negative effects of elevated testosterone on female fecundity in zebra finches

Although factors influencing androgen deposition in the avian egg and its effects on nestling fitness are recently receiving considerable attention, little is known about the potential costs of high testosterone levels in the females. Our study aimed at determining the effect of injections of testosterone (T) in female zebra finches (Taeniopygia guttata), on clutch size, egg mass, yolk mass, and yolk androgen content. Females were given a single bolus injection of T in a range of doses after laying the first egg. Results show that administration of T negatively affected clutch size; the strength of this effect increased with increasing doses of T. Females injected with the highest testosterone dose showed suppressed oviposition of the third and the fourth eggs. Interestingly, testosterone administration made females produce eggs with relatively large yolks, suggesting that T may mediate the trade-off between number and size of eggs. Testosterone injection resulted in elevated levels of androgen in the eggs, in contrast to control clutches, which showed a decreasing pattern of androgen concentration along the laying sequence. We conclude that high androgen investment in eggs may be limited by physiological requirements of the ovulatory process.     

The effects of dietary carotenoid intake on carotenoid accumulation in the retina of a wild bird, the house finch (Carpodacus mexicanus)

Carotenoid pigments accumulate in the retinas of many animals, including humans, where they play an important role in visual health and performance. Recently, birds have emerged as a model system for studying the mechanisms and functions of carotenoid accumulation in the retina. However, these studies have been limited to a small number of domesticated species, and the effects of dietary carotenoid access on retinal carotenoid accumulation have not been investigated in any wild animal species. The purpose of our studies was to examine how variation in dietary carotenoid types and levels affect retinal accumulation in house finches (Carpodacus mexicanus), a common and colorful North American songbird. We carried out three 8-week studies with wild-caught captive birds: (1) we tracked the rate of retinal carotenoid depletion, compared to other body tissues, on a very low-carotenoid diet, (2) we supplemented birds with two common dietary carotenoids (lutein + zeaxanthin) and measured the effect on retinal accumulation, and (3) we separately supplemented birds with high levels of zeaxanthin - an important dietary precursor for retinal carotenoids - or astaxanthin - a dominant retinal carotenoid not commonly found in the diet (i.e. a metabolic derivative). We found that carotenoids depleted slowly from the retina compared to other tissues, with a significant (∼50%) decline observed only after 8 weeks on a very low-carotenoid diet. Supplementation with lutein + zeaxanthin or zeaxanthin alone significantly increased only retinal galloxanthin and ε-carotene levels, while other carotenoid types in the retina remained unaffected. Concentrations of retinal astaxanthin were unaffected by direct dietary supplementation with astaxanthin. These results suggest highly specific mechanisms of retinal carotenoid metabolism and accumulation, as well as differential rates of turnover among retinal carotenoid types, all of which have important implications for visual health maintenance and interventions.     

Carotenoid discrimination by the avian embryo: a lesson from wild birds

The concentrations (μg/g wet yolk) of total carotenoids in eggs of the common moorhen (Gallinula chloropus), American coot (Fulica americana) and lesser black-backed gull (Larus fuscus), collected in the wild, were 47.5, 131.0 and 71.6, respectively. In contrast to data for eggs of the domestic chicken, β-carotene was a significant component in the yolks of these three wild species, forming 25–29% by wt. of the total carotenoids present. The concentration of total carotenoids in the livers of the newly-hatched chicks was 5–10 times higher than in the other tissues and β-carotene was again a major component, forming 37–58% of the hepatic carotenoids. In the newly-hatched gull, the proportions of both lutein and zeaxanthin were very low in the liver but high in the heart and muscle when compared with the yolk. By contrast canthaxanthin, echinenone and β-carotene were very minor constituents of heart and muscle when compared with their proportions in the yolk of the gull. The proportions of lutein and zeaxanthin in the liver of the newly-hatched coot and moorhen were also far lower than in the yolk whereas the liver was relatively enriched with β-cryptoxanthin, β-carotene and (in the moorhen) echinenone. The results indicate that avian embryos discriminate between different carotenoids during their distribution from the yolk to the various tissues.     

Influence of dietary vitamins A and E on serum alpha- and gamma-tocopherols, retinol, retinyl palmitate and carotenoid concentrations in Humboldt penguins (Spheniscus humboldti)

Serum retinol, retinyl palmitate, beta-carotene, cryptoxanthin, lutein, alpha-tocopherol and gamma-tocopherol were measured in 18 captive Humboldt penguins (Spheniscus humboldti) prior to and following the removal of Columbia River (CR) smelt (Thaleichthys pacificus) from the diet. Dietary vitamin A was reduced from 59.8 to 13.5 IU g-1 (dry matter basis) when CR smelt was removed from the diet. Minimal changes were noted in dietary vitamin E. Serum samples Without-CR smelt had significantly lower circulating retinol (1.19 +/- 0.09 vs. 1.94 +/- 0.08 micrograms ml-1) and retinyl palmitate (0.033 +/- 0.012 vs. 0.105 +/- 0.004 microgram ml-1) compared to samples With-CR. The Without-CR smelt diet resulted in increased serum alpha-tocopherol from 26.4 +/- 0.94 to 39.1 +/- 3.72 micrograms ml-1. More serum samples taken Without-CR smelt had detectable levels of gamma-tocopherol than those With-CR smelt. Serum lutein was higher for the samples taken Without versus With-CR smelt. Serum cryptoxanthin did not differ. beta-Carotene was not detected. Data indicate that high levels of dietary vitamin A can affect circulating levels of retinol, retinyl palmitate and vitamin E. Thus, dietary vitamin A and the interrelationship between vitamins A and E should be considered when assessing captive penguins.     

The response of the hypothalamo-gonadal system to environmental factors in the zebra finch,Poephila guttata castanotis

Summary Studies of the hypothalamo-gonadal response of the zebra finch (Poephila guttata castanotis) to environmental factors were conducted by the use of adult birds captured in arid regions of South Australia. In these birds breeding activity is not photoperiodically induced but follows rainfall. The hypothalamo-hypophysial system participating in this response involves centres of the rostral and mediobasal (tuberal) hypothalamus. Parvocellular neurones in both hypothalamic portions are arranged in clusters. In the rostral hypothalamus magnocellular neurosecretory perikarya are spatially closely associated with the clusters which contain different types of parvocellular peptidergic neurones. Extended, complex neuropile formations, rich in synapses, encompass the neuronal clusters and provide important loci of neural-neuroendocrine interaction.Experiments were undertaken to correlate indicators of breeding activity, such as increased size and mass of testes, spermatogenesis, and plasma levels of androgens and luteinizing hormone, with a greater availability of drinking water, increased air humidity, and green vegetation, all of which follow rainfall. Birds maintained in male/female pairs in controlled-environment rooms at 27° C on 14L10D received a diet of seeds (containing 10% water). Thirty-two birds were given drinking water at a basal level of 1 ml/week and one of eight different treatments, consisting of different combinations of drinking water, relative air humidity, and green grass, for two weeks (4 males/treatment). The size of the left testis was estimated at laparotomy and blood samples were taken from a wing vein before and after treatment. The observations suggest that, in moisture-deprived (1 ml water/week) adult zebra finches, an increase in testicular size is associated with a free supply of water; synergistic effects of supplying both water and green grass, or water and high humidity, were also observed. However, in six other groups (8–16 males/treatment), receiving a higher basal level of water (access for 3 min/day), an increase in testicular mass was associated with high humidity, while smaller non-significant increases followed the provision of free water or green grass. Spermatogenic activity was not significantly correlated with testicular size or mass. Leydig cells were sparse in all groups. Plasma androgen levels and luteinizing hormone levels were not obviously correlated with the treatments and were lower than values in zebra finches breeding in outdoor aviaries.According to K.C. Parkes; see Note on Taxonomy, p. XVII, Avian Biology, Vol. IV (D.S. Farner, J.R. King, eds.), Academic Press, New York/London, 1974. Taeniopygia guttata castanotis is widely used for this subspecies in non-taxonomic literature.     

Phototactic responses in Haematococcus lacustris and its modification by light intensity and the carotenoid biosynthesis inhibitor Norflurazon

At fluence rates below 45 W· m^-2 cells of the flagellate stage of Haematococcus lacustris react only positively phototactically with a rather high degree of orientation (indicated by r values up to 0.66 with the Rayleigh test). The directedness of orientation decreases with decreasing irradiance. The degree of directedness of the phototactic response depends on the intensity of preirradiation: Low light intensity applied after strong light application results in a dark reaction (low r values), low light given after darkness stimulates a rather high degree of directedness of positive phototaxis. Weak blue light (=483 nm; 0.4 W · m^-2) stimulates positive phototactic response, whereas comparable red light (=658 nm; 0.5 W · m^-2) does not.Cells which were grown in a medium containing 10^-4 M Norflurazon (effective in inhibition of carotenoid biosynthesis) although maintaining motility completely lose the ability to react positively phototactically. The possible role of carotenoids in the phototactic orientation is discussed.     

Inverse relationship between carotenoid and lipid formation in Rhodotorula gracilis according to the C/N ratio of the growth medium

Production of carotenoid by Rhodotorula gracilis was highest at 26 mg/g dry weight with a carbon/nitrogen (C/N) ratio in the medium of 10:1. This was 15 times higher than when the C/N ratio was 160:1. With this high C/N ratio, the yeast produced up to 55% lipid compared only 20% lipid with the low C/N ratio. Both carotenoid and lipid production were non-growth associated.     

Adrenocortical responses in zebra finches (Taeniopygia guttata): individual variation, repeatability, and relationship to phenotypic quality

Although individual variation is a key requirement for natural selection, little is known about the magnitude and patterns of individual variation in endocrine systems or the functional significance of that variation. Here we describe (1) the extent and repeatability of inter-individual variation in adrenocortical responses and (2) its relationship to sex-specific phenotypic quality, such as song duration and frequency and timing of egg laying. We measured adrenocortical responses to a standardized stressor in zebra finches (Taeniopygia guttata) at two life history stages: approximately day 16 (nestlings) and 3 months of age (sexually mature adults). Subsequently, we assessed phenotypic (reproductive) quality of all individuals as adults. Marked inter-individual variation in the adrenocortical response was seen in both sexes and ages, e.g., stress-induced corticosterone ranged from 2.2 to 62.5 ng/mL in nestlings and 5.0-64.0 ng/mL in adults. We found sex differences in (a) inter-individual variation in the adrenocortical response, (b) repeatability, and (c) relationships between corticosterone levels and phenotypic quality. In males, variation in nestling corticosterone was weakly but positively correlated with brood size and negatively correlated with nestling mass (though this relationship was dependent on one individual). There was no significant correlation of adrenocortical responses between two stages in males and adult phenotypic quality was significantly correlated only with adult corticosterone levels. In contrast, in females there was no relationship between nestling corticosterone and brood size or mass but adrenocortical response was repeatable between two stages (r2=0.413). Phenotypic quality of adult females was correlated with nestling baseline and adrenocortical response.     

Adaptations to fasting in a terrestrial mustelid, the sable (Martes zibellina)

The aim of this study was to investigate whether the actively wintering Palearctic sable Martes zibellina has evolved physiological adaptations to tolerate nutritional scarcity. Sixteen farm-bred male sables were divided into a fed control group and an experimental group fasted for 4 days. The rate of weight loss in the sable was similar to other medium-sized mustelids. Fasting led to hypoglycaemia and to a decreased lymphocyte percentage. The sable derived metabolic energy from both subcutaneous and intraabdominal white adipose tissues and the relative decrease in fat mass was the largest for the retroperitoneal and subcutaneous depots. Metabolic energy derived partly from body proteins indicated by the increased plasma levels of urea, uric acid and total essential amino acids. Triacylglycerols accumulated in the livers of the fasted sables and the increased plasma aminotransferase activities suggested hepatic dysfunction. The decreased plasma insulin concentrations and the elevated cortisol levels probably contributed to stimulated lipolysis and protein catabolism. Moreover, fasting increased the plasma ghrelin concentrations of the sables and down-regulated the thyroid activity.     

Super-normal length song preferences of female zebra finches (Taeniopygia guttata) and a theory of the evolution of bird song

Two way choice tests show a preference of female zebra finches for male songs four standard deviations longer than normal song. Further tests show the ontogeny of this preference to parallel song learning in general as well as a preference for songs with entirely heterogeneous notes compared to songs with four note repeats. These findings are discussed in relation to a theory of the evolution of bird song from bird calls due to female preferences for longer, more complex vocalizations. This revised version was published online in July 2006 with corrections to the Cover Date.