Yolk Antioxidant Research Articles

Biliverdin- and protoporphyrin-based eggshell pigmentation in relation to antioxidant supplementation, female characteristics and egg traits in the canary (Serinus canaria)

Avian eggs exhibit a large variability in coloration and patterns, which are produced by blue-green biliverdin and red-brown protoporphyrin pigments. Several hypotheses have been proposed to explain the function of eggshell coloration. In this experimental study, we tested two hypotheses (signalling-function hypothesis and structural-function hypothesis) on both eggshell pigment types in an open-nesting songbird, the canary (Serinus canaria). Also, we aimed to examine whether deposition of pigments into the eggshell has any cost in terms of the plasma oxidative status of the female. We found that eggshell average blue-green chroma was increased by antioxidant supplementation, although we note that there had already been a pre-existing bias in plasma antioxidant capacity between the supplemented and control groups. Eggshell average blue-green chroma was positively related to female body condition during egg laying. However, blue-green eggshell colour was not related to female oxidative status during or after the laying period, and blue-green chroma increased with laying order. Accordingly, we found some support for that eggshell blue-green colour could reflect maternal antioxidant availability and body condition, but did not find evidence that it has a cost for the female’s oxidative status. By contrast, eggshell spot brightness was positively related to body condition, suggesting that darker spotting reflected poorer nutritional condition in the canary. Eggshell blue-green pigmentation was not significantly connected to the egg volume or yolk antioxidant level, but we found that eggs with lower yolk antioxidant concentration had higher average eggshell brown chroma. In sum, our results suggest that eggshell colour reflected female antioxidant and nutrient availability. Finally, we found that eggs with thinner eggshells had a more aggregated spot distribution, supporting the view that aggregated spots may help to strengthen eggshells. Avian eggs have a large variability in colours and patterns, which are due to two pigments: the blue-green biliverdin and the red-brown protoporphyrin. In this study, we tested whether eggshell pigmentation, measured by a spectrophotometer, could reflect female and egg quality in the canary, an open nesting songbird. We found that females supplemented with antioxidants before and during egg-laying laid more intense blue-green eggs. Females in better body condition laid eggs that had a more intense blue-green coloration, but lower intensity of brown coloration. Egg yolk antioxidant level was lower in eggs with more intense brown eggshell coloration. These results suggest that eggshell coloration could reflect the antioxidant and nutrient availability of females. Moreover, we found that eggs with thinner eggshells had a more aggregated spot distribution on the eggshell, supporting the view that aggregated brown pigment spots may help to strengthen eggshells.

Effects of environmental conditions on the egg mass, yolk antioxidant level, eggshell thickness and eggshell spotting patterns of Great Tits (Parus major)

Avian mothers can influence the fitness of their offspring by resource investment into the egg. Allocation of macro- and micronutrients into the eggs may be costly for the female, therefore, we expect that resource investment may be affected by the environmental and social conditions the mother experiences during egg formation. Here, we investigated whether environmental circumstances experienced by the reproducing female exert an influence on egg mass, yolk antioxidant (lutein and tocopherol) concentration, eggshell thickness, and the eggshell spotting patterns of Great Tits (Parus major). Our study showed that when caterpillars were less abundant, in colder weather and when breeding in an area of higher local breeding density, female Great Tits laid eggs of lower mass, suggesting that adverse environmental circumstances constrain the macronutrient investment into the eggs. However, we found no evidence that yolk antioxidant concentration and eggshell thickness were affected by environmental factors. Female Great Tits may use their endogenous stores or spend more time and energy in finding sufficient amounts of dietary antioxidants and calcium under unfavourable environmental conditions, which may have a cost effect on their own conditions. We found that birds that bred in the beginning of the season laid eggs with darker eggshell spots. Moreover, females laid more spotted eggs in colder weather and when breeding in higher density areas. Our results suggest that Great Tits deposit more of the potentially harmful pro-oxidant protoporphyrin pigment into the eggshell under unfavourable environmental conditions.

Darker eggshell spotting indicates lower yolk antioxidant level and poorer female quality in the Eurasian Great Tit (Parus major)

ABSTRACT Protoporphyrin pigment causes the red-brown eggshell colors; however, for many species, the function of this pigment is unknown. It has been proposed that eggshell pigmentation may strengthen the shell or that it may be a sexually selected signal, which advertises the quality of the female and that of her offspring to the male parent. In this study, we aimed to discover whether protoporphyrin-based eggshell pigmentation patterns of Eurasian Great Tits (Parus major) were related to female or egg quality. Additionally, we tested whether different methods of eggshell pigmentation estimation could be reliable predictors of eggshell protoporphyrin levels. We found that spot intensity, spot size, spotting coverage, and brown spot chroma indicated the protoporphyrin pigment concentration of the eggshell. Our results revealed that Eurasian Great Tit females that laid eggs with darker pigmentation had more lymphocytes in their circulation and had paler yellow breast and lower UV plumage reflectance, possi...

The effect of maternal immunization on female oxidative status, yolk antioxidants and offspring survival in a songbird

Immune defense involves inflammatory reactions in which immune cells produce reactive oxygen species (ROS) to fight pathogens. ROS may however cause damage to the host if they are not balanced by antioxidant defenses. Therefore, one should expect individuals undergoing an immune reaction to use antioxidants to prevent oxidative stress. Antioxidants are vital compounds that provide important protection against oxidative damage of embryos and newly hatched chicks. Thus, during egg laying a female that contracted an infection may face a trade-off between the allocation of antioxidants into self-maintenance and into her offspring via the eggs. In our study we investigated whether immunized females face this trade-off and consequently modify the antioxidant allocation into the eggs and whether this allocation affects offspring performance. We injected female zebra finches (Taeniopygia guttata) with lipopolysaccharide prior to egg laying while some females were left unimmunized. We removed the second egg of each clutch, while we allowed the other eggs to hatch. We assessed oxidative stress in females 24h after immunization, yolk antioxidant capacity of the second egg of the clutch and survival success of the offspring until adulthood. Compared to controls, immunized females had higher oxidative damage, but similar plasma non-enzymatic antioxidant levels. The treatment did not affect yolk antioxidants, clutch size, laying date and offspring survival. However, we found a positive correlation between yolk antioxidant capacity and offspring survival, irrespective of the treatment. Our study suggests that our immune challenge may not have changed female strategy of antioxidant allocation between self-maintenance and offspring survival.

Effects of breeding habitat (woodland versus urban) and metal pollution on the egg characteristics of great tits (Parus major)

In an urban environment, birds are exposed to metals, which may accumulate in their tissues and cause oxidative stress. Female birds may eliminate these pollutants through depositing them into eggs, thus eggs become suitable bioindicators of pollution. In this study, we aimed to analyse whether eggshell spotting pattern, egg volume, eggshell thickness and egg yolk antioxidant (lutein, tocopherol, retinol and selenium) levels were related to the breeding area (woodland versus urban) and the metal levels in the eggshell of a small passerine species, the great tit (Parus major). In the urban habitat, soil and eggshells contained higher concentrations of metals, and soil calcium level was also higher than that in the woodland. Eggshell spotting intensity and egg volume did not differ between eggs laid in the woodland and the urban park, and these traits were not related to the metal levels of the eggshell, suggesting that these egg characteristics are not sensitive indicators of metal pollution. A more aggregated eggshell spotting distribution indicated a higher Cu concentration of the eggshell. We found that eggshells were thinner in the less polluted woodland habitat, which is likely due to the limited Ca availability of the woodland area. Great tit eggs laid in the urban environment had lower yolk lutein, retinol and selenium concentrations, however, as a possible compensation for these lower antioxidant levels, urban females deposited more tocopherol into the egg yolk. It appears that females from different breeding habitats may provide similar antioxidant protection for their offspring against oxidative damage by depositing different specific dietary antioxidants. Egg yolk lutein and retinol levels showed a negative relationship with lead concentration of the eggshell, which may suggest that lead had a negative impact on the amount of antioxidants available for embryos during development in great tits.

Effect of high stocking density on performance, egg quality, and plasma and yolk antioxidant capacity in laying hens supplemented with organic chromium and vitamin C

This study was conducted to investigate the effects of dietary supplementation of chromium–methionine (CrMet) and vitamin C on performance, egg quality measurements, and plasma antioxidant capacity in laying hens subjected to high stocking density. A total of 360 Hy-Line W-36 laying hens, 26wk of age, were used in a 2×2×3 factorial arrangement of treatments consisting of 2 cage densities [5 or 7 hens/cage (40×45cm2)], 2 supplemental vitamin C levels (0 and 500mg/kg as l-ascorbic acid), and 3 supplemental Cr levels (0, 500, and 1000μg/kg as CrMet) with 5 replicate cages per treatment. The hens were kept in wire-floor cages in an environmentally-controlled room. After a 2-wk adaptation, the 70-d study was initiated and it was divided into two 35-d periods (28–33 and 33–38wk of age). Results showed that although subjecting hens to high stocking density had no effect on egg production percentage during 28–33wk of age, it decreased (P=0.031) during the 33–38wk of age, resulting in a marked reduction (P=0.038) in egg mass. Dietary Cr supplementation (500 and 1000μg/kg) increased (P<0.01) egg production and egg mass during both 35-d periods, and improved (P=0.005) feed conversion ratio (FCR) during the first 35-d period. There was a stocking density×vitamin C interaction (P=0.018) for FCR values during the first 35-d period, so that supplemental vitamin C improved FCR to a greater extent in high stocking density-challenged hens. Dietary CrMet supplementation (500 and 1000μg/kg) increased (P=0.001) feed intake during the second 35-d period. Shell thickness was greater (P=0.015) in hens under high stocking density challenge during the second 35-d period. Subjecting hens to high stocking density decreased (P=0.007) plasma vitamin C concentration, while plasma vitamin C level was increased (P<0.05) as the result of dietary supplementation of vitamin C. Dietary supplementation of CrMet (500 and 1000μg/kg) increased plasma vitamin C level only in high stocking density-challenged hens, resulting in a stocking density×Cr interaction (P=0.039). Although high stocking density increased (P<0.05) plasma and yolk concentrations of malondialdehyde (MDA), supplemental vitamin C decreased (P<0.05) plasma and yolk MDA contents. Similarly, dietary inclusion of CrMet (500 and 1000μg/kg) decreased (P=0.046) plasma MDA level. The present findings indicate that dietary CrMet supplementation could increase antioxidant capacity in stressed hens, which improve production performance of laying hens.

Female oxidative status, egg antioxidant protection and eggshell pigmentation: a supplemental feeding experiment in great tits

Oxidative stress has been suggested as a mechanism underlying the costs of reproduction and life history trade-offs. Reproductive activities may lead to high production of pro-oxidants, whose activity can generate oxidative damage when not countered by adequate antioxidant defenses. Because inter-individual differences in the efficiency of the antioxidant system are influenced by an individual’s diet, food availability experienced during reproduction may affect the females’ antioxidant status and, in birds, their ability to transfer antioxidants into their eggs. Moreover, a female’s ability to cope with oxidative stress has been suggested to influence pigment deposition in the eggshell, suggesting a possible signaling function of eggshell maculation. Here we performed a food supplementation experiment in a natural population of great tits (Parus major) in order to investigate how nutritional conditions experienced during the egg laying period affect the female’s oxidative status and egg investment and how maternal oxidative status and egg antioxidant protection relate to eggshell pigmentation. We show that food-supplemented females had lower oxidative damage levels (ROMs) than non-food-supplemented females. Furthermore, a female’s ROMs levels were negatively associated with the levels of yolk antioxidant protection in her eggs, but this negative association was only significant in non-food-supplemented females. This suggests that oxidative stress experienced during reproduction influences the allocation of antioxidants into the eggs. Moreover, we observed a positive relationship between eggshell pigment distribution and maternal and yolk antioxidant protection, suggesting that eggshell pigmentation is a cue of female (and offspring) quality.

A comparative assessment of the fatty acid profiles and antioxidant status of supermarket eggs

SummaryNutrient deposition in eggs is largely dictated by the dietary composition of laying hen feed, particularly in terms of specific fatty acids and antioxidants. In the present study, the nutritional quality of a range of commercially available egg varieties, marketed as omega-3 enriched; corn-fed; free range and standard caged, were assessed by determining fatty acid profiles and antioxidant status. Across each egg variety, significant differences were observed in key fatty acids such as palmitic, oleic, linoleic, alpha-linolenic and docosahexaenoic acid (DHA) (P ≤ 0.05). Egg yolks enriched with a stated dietary source of omega-3 fatty acid DHA were shown to have significantly improved levels of DHA (P ≤ 0.05), approximately 4.5-fold higher than standard caged eggs. Compared with free range, corn fed and caged, eggs from diets enriched with a source of omega-3 were shown to have considerably altered omega-6: omega-3 ratios, amounting to 1.5–2.1 fold reductions. Yolk antioxidant activity was improved for omega-3 enriched eggs, particularly in hexane fractionated samples. The inclusion of omega-3 fatty acids to the diet resulted in eggs with improved DHA contents and antioxidant status, highlighting the importance of poultry diet composition for egg nutritional quality.

Embryonic growth and antioxidant provision in avian eggs

Avian embryos undergo extremely rapid development over a relatively short period of time, and so are likely to suffer high levels of oxidative damage unless this is mitigated by sufficient maternal allocation of appropriate antioxidants. At a species level, it is therefore predicted that antioxidants should be allocated to eggs according to the rate of embryonic growth, such that eggs containing embryos that grow faster are furnished with higher antioxidant levels, independent of egg size. We tested this prediction for three potentially important classes of dietary-derived yolk antioxidants: carotenoids, vitamin E and vitamin A. Across species, we found positive relationships between embryonic growth rate and total yolk levels of each of the three antioxidant classes. Moreover, there were consistent differences in antioxidant provision between pairs of species that share a common initial egg mass yet have differing rates of embryonic growth, such that the eggs of the faster-developing species have higher levels of carotenoids and vitamin E. These results may explain the marked interspecific variation in antioxidant provision and provide evidence for the role that these antioxidants play during embryonic development.

Effects of Astragalus membranaceus on laying performance and antioxidant status of laying hens

SUMMARY Interest in the use of natural feed additives in the animal and poultry industries is growing. In the present study, we assessed the effects of dietary supplementation of Astragalus membranaceus root powder (AMP) on the laying performance and serum and egg yolk antioxidant status of laying hens. Four groups (135 each) of laying hens fed a corn- and soybean meal-based diet were supplemented with 0, 5, 10, or 15 g/kg of diet of AMP for 10 wk, and their laying performance and serum and egg yolk antioxidant status were measured. Supplementation of AMP did not affect ADFI or average egg weight but did linearly increase egg mass of the laying hens. Antioxidant status of the serum and egg yolk was improved, as judged by an increase in antioxidant enzymatic activities and a decrease in concentrations of oxidized products, with supplementation of AMP in a dose-dependent manner. In conclusion, supplementing a cornand soybean meal-based laying hen diet with AMP at levels of 5 to 10 g/kg of diet had the potential to improve the antioxidant status of laying hens and improve laying performance.

Mind the gap: the ratio of yolk androgens and antioxidants varies between sons and daughters dependent on paternal attractiveness

Females are expected to partition resources between offspring in a context-dependent way to maximise total fitness returns from a reproductive attempt. Female zebra finches (Taeniopygia guttata) vary the allocation of yolk androgens and antioxidants among offspring. Importantly, the balance between androgens and antioxidants in yolks may be more important than their independent absolute amounts in terms of fitness consequences for developing young. Therefore, we tested whether the relative allocation of these two resources in yolks varies according to either the Trivers–Willard, positive or compensatory maternal investment hypothesis. We manipulated male attractiveness using coloured leg bands (red-banded males appear attractive; green-banded males, unattractive) and measured yolk androgens and antioxidants in each egg, egg sex, clutch sex ratio and female condition. While female zebra finches manipulated the balance of androgens and antioxidants within and between clutches in response to mate attractiveness, offspring sex and their own condition, they did not do so in a way that consistently followed any of the hypotheses. Mothers paired with unattractive males allocated a larger antioxidant/androgen ratio to daughters than sons. This pattern was reversed when paired to an attractive male; sons received a larger antioxidant/androgen ratio than daughters. We also found offspring sex ratio decreased with increasing female condition for unattractive males, but not for attractive males. However, without knowing the fitness consequences of the balance of different egg constituents, it is difficult to interpret the patterns consistently in terms of the Trivers–Willard, compensatory and positive investment hypotheses.

Blue-green eggshell coloration reflects yolk antioxidant content in spotless starlings Sturnus unicolor

The hypothesis that eggshell colouration is a sexually selected trait of female birds is based on the fact that biliverdin, the pigment responsible for blue-green colours of the eggshell, is a potent antioxidant and that only females with high antioxidant capacity can deposit higher concentrations of biliverdin as eggshell pigment. Antioxidants (e.g. carotenoids, vitamins) are also abundant in the egg yolk, which serve as nutrient reserves for the developing embryo, and eggshell colour intensity may reflect maternal investment in yolk antioxidants. Here, we test the relationship between blue-green eggshell colour intensity and concentration and amount of carotenoids, vitamin A, and vitamin E in the egg yolk of spotless starling Sturnus unicolor, a species for which we have previously shown good evidence of sexual selection driving egg coloration. As could be extrapolated from the hypothesis of sexual selection driving the evolution of blue-green eggshells, we found that eggshell colour intensity was positively related to the concentration and amount of carotenoids and vitamin E in the yolk. Thus, mothers may use egg colour intensity to signal to fathers the antioxidant status of their offspring. Moreover, we provide evidence suggesting that maternal yolk investment in more coloured eggs can also explain the detected association between feeding decisions of males and egg colour intensity that we have found previously in this species.

Egg yolk antioxidant deposition as a function of parental ornamentation, age, and environment in great tits Parus major

Females can modify phenotype of their offspring through the deposition of biologically active compounds into eggs, including carotenoids, vitamins and other antioxidants. Understanding patterns of deposition is critical for better insight into the significance of maternal effects. Here we investigated how egg yolk antioxidants (lutein, zeaxanthin, β-carotene, vitamin A and E) related to environmental conditions and parental characteristics in great tits Parus major using data from three breeding seasons. Male and female traits included condition, age and multiple feather ornaments, both carotenoid- and melanin-based (carotenoid and UV chroma of yellow breast feathers, area of black breast band, white cheek immaculateness). Yolk mass increased with ambient temperature during laying, laying date, and the area of male black breast band. Lutein, zeaxanthin, and vitamin E increased with laying date. Total antioxidants increased with female age, immaculateness of female white cheek patch, and UV chroma of carotenoid-based yellow breast feathers of the social mate. These patterns were thus consistent with 1) environmental effects on yolk mass and composition, 2) higher quality females depositing more antioxidants, and 3) differential allocation of resources in females in relation to male ornamentation. Overall, environmental factors, female traits, and male traits all had an influence on egg yolk characteristics in this socially monogamous songbird.

Effects of ginger root (Zingiber officinale) on laying performance and antioxidant status of laying hens and on dietary oxidation stability

To investigate the effects of ginger root (Zingiber officinale) on laying performance and antioxidant status of laying hens and on dietary oxidation stability, 675 Hy-Line brown laying hens at 27 wk of age were randomly allocated to 5 treatments with 5 replicates in a complete randomized design. Laying hens were fed corn-soybean meal based diets and supplemented with 0, 5, 10, 15, or 20 g of ginger powder/kg of diet for 10 wk. The ADFI, laying rate, average egg weight, egg mass, and feed conversion of each replicate were measured weekly. Blood samples from 10 laying hens and 15 egg samples/treatment were obtained on d 35 and 70 of the experiment to determine the serum or egg yolk antioxidant enzymatic activities. Three separate batches of each treatment diet were also made at the beginning of the experiment and stored at 20°C. Subsamples were taken at 10-d intervals to determine the dietary oxidation stability over a 60-d period. All laying hens had similar ADFI, laying rate, average egg weight, and feed conversion. However, laying hens supplemented with ginger powder had higher (P = 0.024) egg mass than the control group. Supplementation of ginger powder increased (P < 0.05) the serum and egg yolk antioxidant enzymatic activities of laying hens on d 35 and 70. Increasing ginger powder from 0 to 20 g/kg of diet linearly increased (P < 0.05) the peroxide value but linearly reduced (P < 0.05) the acid value of lipid extracted from stored diets for the first 30 d, but not for the second 30 d. Dietary supplementation of ginger powder improved laying performance and serum and egg yolk antioxidant status and enhanced dietary oxidation stability in a dose-dependent manner. The optimum supplementation rate of ginger powder in the diet of laying hens appeared to be between 10 and 15 g/kg of diet.

Pterin-based ornamental coloration predicts yolk antioxidant levels in female striped plateau lizards (Sceloporus virgatus)

1. Maternal investment in egg quality can have important consequences for offspring fitness. For example, yolk antioxidants can affect embryonic development as well as juvenile and adult phenotype. Thus, females may be selected to advertise their yolk antioxidant deposition to discriminatory males via ornamental signals, perhaps depending on the reproductive costs associated with signal production. 2. Female striped plateau lizards (Sceloporus virgatus) develop pterin-based orange colour patches during the reproductive season that influence male behaviour and that are positively associated with the phenotypic quality of the female and her offspring. Here, we assessed one potential developmental mechanism underlying the relationship between offspring quality and female ornamentation in S. virgatus, by examining the relationship between ornament expression and yolk antioxidant levels. 3. As expected, concentrations of the yolk antioxidants vitamin A, vitamin E and carotenoids (lutein and zeaxanthin) were strongly positively intercorrelated. Eggs from larger clutches had fewer antioxidants than eggs from smaller clutches, suggesting that females may be limited in antioxidant availability or use. Fertilized and unfertilized eggs did not differ in yolk antioxidant levels. 4. The size of a female's ornament was positively related to both the concentration and total amount of yolk antioxidants, and ornament colour was positively related to yolk antioxidant concentration. Thus, in S. virgatus, female ornaments may advertise egg quality. In addition, these data suggest that more ornamented females may produce higher-quality offspring, in part because their eggs contain more antioxidants. As the colour ornament of interest is derived from pterins, not carotenoids, direct resource trade-offs between ornaments and eggs may be eliminated, reducing reproductive costs associated with signalling. 5. This is the first example of a positive relationship between female ornamentation and yolk antioxidants in reptiles and may indicate the general importance of these patterns in oviparous vertebrates.

Patterns of egg yolk antioxidant co-variation in an avian brood parasite–host system

Despite their importance to offspring fitness outcomes, there has been little previous work on egg-mediated maternal effects in avian brood parasites which lay their eggs in the nests of other species. Here, we examine patterns of egg yolk antioxidant deposition in an avian host (red-winged blackbird, Agelaius phoeniceus) and their brood parasite (brown-headed cowbird, Molothrus ater). Cowbird nestlings compete directly for food with host blackbird chicks and succeed, despite hatching from a smaller egg, by begging more intensely and growing as or more rapidly than their hosts. Dietary-derived antioxidants, such as carotenoids and vitamins E and A, deposited in egg yolk, may enhance growth rate and reduce oxidative stress, and therefore provide a potential mechanism by which female cowbirds could enhance the competitiveness of their young in host nests. However, provision of antioxidants to eggs is costly so we predicted that female cowbirds should adjust the amount of antioxidants in response to variation in host quality. We found that whilst red-winged blackbird clutches parasitised by cowbirds did not differ in the pattern of antioxidant deposition compared to non-parasitised clutches, levels of vitamin E deposited in cowbird eggs were closely matched to that of the individual host clutch in which cowbirds laid their egg. This supports the prediction that female cowbirds adjust the antioxidant concentration of their eggs to increase the likely competitiveness of their offspring in the host nest into which they are laid and may help explain the success of cowbirds in competing with larger host young.

Effects of dietary resveratrol supplementation on egg production and antioxidant status

Resveratrol, a polyphenol derived from red grapes, berries, and peanuts, exerts antiinflammatory, antioxidant, and immunomodulatory effects. The objective of this study was to investigate the effects of dietary resveratrol supplementation on performance and serum and egg yolk antioxidant status in quail (Coturnix coturnix japonica). A total of 150 five-week-old quails were allocated randomly to 1 of 3 dietary treatments: basal diet and basal diet supplemented with 200 or 400 mg of resveratrol/kg of diet. Each diet was offered to 10 cages of 5 birds in each from 4 to 16 wk of age. Serum and egg samples were collected at the beginning and end the experimental period to be evaluated for malondialdehyde (MDA), vitamin A, and vitamin E. Data were subjected to analysis of covariance using the MIXED procedure. There was no treatment effect on feed intake, egg production, or egg quality parameters related to shell, yolk, and albumen. There were no effects of resveratrol supplementation on serum and egg yolk vitamin A concentrations. The quails supplemented with resveratrol had a lower serum MDA concentration (0.56 vs. 0.88 mg/L, P<0.03) and a higher serum vitamin E concentration (5.72 vs. 3.56 mg/L, P<0.008) than those not supplemented with resveratrol. Moreover, there was a linear decrease in serum MDA concentration (P<0.02) and a linear increase in serum vitamin E concentration (P<0.01) as supplemental resveratrol level increased. The treatment groups had less egg yolk MDA concentration than the control group (0.21 vs. 0.15 microg/g, P<0.002). Egg yolk MDA concentration decreased linearly in response to increasing dietary resveratrol level (P<0.0003). In conclusion, inclusion of resveratrol up to 400 mg/kg into quail diets enhanced antioxidant status of birds and eggs. Further studies should investigate the carryover effects of dietary resveratrol supplementation on product quality with respect to shelf life, antioxidant stability, and its nutritive value for human consumption.

Complex trade-offs in the pigeon (Columba livia): egg antioxidant capacity and female serum oxidative status in relation to diet quality

Despite the great deal of ecological research interest in modulators of offspring quality and consequences of reproduction on female status, we still know little about the relationships among diet quality, antioxidant capacity of egg components (yolk and albumen) and oxidative status of female birds. In this study, I compared the egg quality (egg size, albumen and yolk antioxidant capacity) and serum oxidative status (oxidative damage, total serum antioxidant capacity, and serum thiols) of female pigeons (Columba livia) fed with foods of different quality (standard quality and decreased quality). I also analysed the patterns of covariation among egg and female traits. The study focussed on the first clutch laid by the female in the breeding season and on the short-term effects of a decrease in diet quality. The treatment did not affect the egg volume, the lipophilic and hydrophilic components of antioxidant capacity (lipOXY and hydrOXY, respectively) or the antioxidant capacity of the albumen (albumOXY). However, females fed a higher quality diet were fatter and had marginally higher values of serum hydroperoxides (oxidative damage) than females fed a lower quality diet. Moreover, females that showed an increase in yolk hydrOXY and serum hydroperoxides also showed a decrease in yolk lipOXY, albumOXY and serum antioxidant capacity. These results show that the female's oxidative status can be correlated with the antioxidant content of her eggs, but the nature of these correlations is complex, depending on the molecular component measured. The results also suggest that in the pigeon the deposition of hydrophilic and lipophilic antioxidants in the egg may trade off against each other.

Eggshell characteristics and yolk composition in the common cuckoo Cuculus canorus: are they adapted to brood parasitism?

The developmental rate of cuckoo embryos and their hatching size is greater than that of host species, which may require more nutrient resources in the egg and more intensive gas exchange during development. In the present study, we compared various egg characteristics of a brood parasite, the common cuckoo Cuculus canorus, and its frequent host, the great reed warbler Acrocephalus arundinaceus. As maternally‐derived testosterone is known to enhance growth rate of embryos and hatchlings, cuckoo eggs are expected to contain higher concentration of testosterone than host eggs. In addition, we expected higher concentration of antioxidants in cuckoo eggs to protect embryos from oxidative stress associated with accelerated growth. Our results showed that cuckoo eggs had thicker shells and higher pore density than great reed warbler eggs. Yolk was significantly heavier in cuckoo eggs and contained higher concentrations of carotenoids and vitamin E, however, yolk androgen and immunoglobulin concentrations were lower in cuckoo eggs as compared to great reed warbler eggs. We also examined whether eggshell colour was associated to egg quality, and detected a positive association between blue‐green chroma and yolk antioxidant concentration in both species, suggesting that eggshell colour reflects the antioxidant investment of the female into the eggs. Our results suggest that cuckoo females increase the size, growth rate and competitive ability of their young by providing them with more nutrients and more dietary antioxidants for embryonic development, and not through elevated yolk testosterone or antibody levels. In addition, increased porosity of cuckoo eggshells may allow embryos to develop more rapidly because of a greater capacity of gas exchange.

Egg composition in relation to social environment and maternal physiological condition in the collared flycatcher

Offspring survival can be influenced by resources allocated to eggs, which in turn may be affected by the environmental factors the mother experiences during egg formation. In this study, we investigated whether experimentally elevated social interactions and number of neighbouring pairs influence yolk composition of collared flycatchers (Ficedula albicollis). Social challenge was simulated by presentation of a conspecific female. Experimental females spent more time near the cage and produced eggs with higher androgen concentration, but local breeding density did not affect yolk androgen level. Moreover, we found that females exposed to more intra-specific interactions and those that bred at higher density produced eggs with smaller yolk. These females may be more constrained in foraging time due to more frequent social encounters, and there might be increased competition for food at areas of higher density. In contrast, the present study did not reveal any evidence for the effect of social environment on yolk antioxidant and immunoglobulin levels. However, we found that yolk lutein and immunoglobulin concentrations were related to the female’s H/L ratio. Also, yolk lutein and α-tocopherol levels showed a seasonal increase and were positively related to the female’s plasma carotenoid level. Mothers may incur significant costs by transferring these compounds into the eggs, thus only females in good physiological condition and those that lay eggs later, when food is probably more abundant, could allocate higher amounts to the eggs without compromising their defence mechanisms. Our results suggest that environmental circumstances during egg formation can influence conditions for embryonic development.