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Biomedical subjects

N Saino

Publications and source records attributed to N Saino.

17 recordsLinked to original sources

Increased egg estradiol concentration feminizes digit ratios of male pheasants (Phasianus colchicus).

The length ratio between individual digits differs between males and females in humans, other mammals, lizards, and one bird species. Sexual dimorphism in digit ratios and variation among individuals of the same sex may depend on differential exposure to androgens and estrogens during embryonic life. Organizational effects of sex hormones could cause the observed correlations between digit ratios and diverse phenotypic traits in humans. However, no study has investigated experimentally the effect of prenatal estrogens on digit ratios. We analyzed the effect of estradiol injection in ring-necked pheasant (Phasianus colchicus) eggs on digit ratios. Males from control eggs had higher ratios between the second or third and the fourth digit of the right foot compared to females. Estradiol-treated eggs produced males with lower (feminized) right foot second to fourth digit ratio. Thus, we provided the first experimental evidence that prenatal exposure to physiologically high estrogen levels affects bird digit ratios.

Animals↗

Differential effects of egg albumen content on barn swallow nestlings in relation to hatch order.

In diverse animal taxa, egg mass variation mediates maternal effects with long-term consequences for offspring ontogeny and fitness. Patterns of egg mass variation with laying order differ considerably among birds, but no study has experimentally investigated the function of variation in albumen or yolk egg content in the wild. In barn swallows (Hirundo rustica), absolute and relative albumen mass increased with egg laying order. Experimental albumen removal delayed hatching, had larger negative effects on growth of late-hatched nestlings, and reduced nestling survival. Laying order positively predicted hatch order. Because nestling competitive ability depends on size, and albumen egg content influences hatchling size, present results suggest that by increasing albumen content of late eggs mothers reduce hatching asynchrony and enhance growth particularly of late-hatched nestlings. Thus, variation in albumen mass with laying order may function to mitigate the negative phenotypic consequences of hatching late in species that adopt a 'brood-survival' strategy.

Animals↗

Early maternal, genetic and environmental components of antioxidant protection, morphology and immunity of yellow-legged gull (Larus michahellis) chicks.

Maternal effects mediated by egg quality are important sources of offspring phenotypic variation and can influence the course of evolutionary processes. Mothers allocate to the eggs diverse antioxidants that protect the embryo from oxidative stress. In the yellow-legged gull (Larus michahellis), yolk antioxidant capacity varied markedly among clutches and declined considerably with egg laying date. Analysis of bioptic yolk samples from clutches that were subsequently partially cross-fostered revealed a positive effect of yolk antioxidant capacity on embryonic development and chick growth, but not on immunity and begging behaviour, while controlling for parentage and common environment effects. Chick plasma antioxidant capacity varied according to rearing environment, after statistically partitioning out maternal influences mediated by egg quality. Thus, the results of this study indicate that egg antioxidants are important mediators of maternal effects also in wild bird populations, especially during the critical early post-hatching phase.

Animals↗

An analysis of continent-wide patterns of sexual selection in a passerine bird.

Patterns of selection are widely believed to differ geographically, causing adaptation to local environmental conditions. However, few studies have investigated patterns of phenotypic selection across large spatial scales. We quantified the intensity of selection on morphology in a monogamous passerine bird, the barn swallow Hirundo rustica, using 6495 adults from 22 populations distributed across Europe and North Africa. According to the classical Darwin-Fisher mechanism of sexual selection in monogamous species, two important components of fitness due to sexual selection are the advantages that the most attractive males acquire by starting to breed early and their high annual fecundity. We estimated directional selection differentials on tail length (a secondary sexual character) and directional selection gradients after controlling for correlated selection on wing length and tarsus length with respect to these two fitness components. Phenotype and fitness components differed significantly among populations for which estimates were available for more than a single year. Likewise, selection differentials and selection gradients differed significantly among populations for tail length, but not for the other two characters. Sexual selection differentials differed significantly from zero across populations for tail length, particularly in males. Controlling statistically for the effects of age reduced the intensity of selection by 60 to 81%, although corrected and uncorrected estimates were strongly positively correlated. Selection differentials and gradients for tail length were positively correlated between the sexes among populations for selection acting on breeding date, but not for fecundity selection. The intensity of selection with respect to breeding date and fecundity were significantly correlated for tail length across populations. Sexual size dimorphism in tail length was significantly correlated with selection differentials with respect to breeding date for tail length in male barn swallows across populations. These findings suggest that patterns of sexual selection are consistent across large geographical scales, but also that they vary among populations. In addition, geographical patterns of phenotypic selection predict current patterns of phenotypic variation among populations, suggesting that consistent patterns of selection have been present for considerable amounts of time.

Age Factors↗

Geographic variation in the G matrices of wild populations of the barn swallow.

In this paper, we present an analysis of genetic variation in three wild populations of the barn swallow, Hirundo rustica. We estimated the P, E, and G matrices for six linear morphological measurements and tested for variation among populations using the Flury hierarchical method and the jackknife followed by MANOVA method. Because of nonpositive-definite matrices, we had to employ 'bending' to analyse the G and E matrices with the Flury method. Both statistical methods agree in finding that the P and G matrices are significantly different but comparison between the analysis of the P matrices and pairwise analyses of the P, E, and G matrices suggests caution in interpreting the Flury results concerning differences in matrix structure. The significant variation among the populations in the G matrices appears to be due in large measure to the most geographically distant population.

Animals↗

Genetic variation in infestation with a directly transmitted ectoparasite.

Genetic variation in levels of parasitism of hosts is an underlying assumption of studies of coevolution, but few such estimates are available from the field. We studied genetic variation in the abundance of the chewing louse Hirundoecus malleus on its barn swallow host Hirundo rustica. These parasites are directly transmitted and a test of genetic variation of parasite abundance would thus provide a particularly strong test. The prevalence and the abundance of the chewing lice did not differ significantly between adult male and female hosts. The resemblance in parasite intensity of H. malleus of offspring and their parents was positive and highly significant, and an analysis of extra-pair paternity in the host allowed partitioning of this resemblance between genetic and common environment effects. There was no significant resemblance in parasite intensity between extra-pair offspring and their foster parents, although the resemblance remained for within-pair offspring. This provides evidence for the abundance of directly transmitted parasites having an additive genetic component. We found no evidence of common environment effects as parents did not resemble each other with respect to lice abundance.

Animals↗

Maternal antibodies but not carotenoids in barn swallow eggs covary with embryo sex.

Mothers influence their offspring phenotype by varying egg quality. Such maternal effects may be mediated by transmission of antibodies and antioxidants. Mothers should adjust allocation of maternal substances depending on embryonic sex because of differences in reproductive value, potentially dependent on paternal genetic effects as reflected by secondary sexual characters. We manipulated sexual attractiveness of male barn swallows (Hirundo rustica) and investigated maternal investment in eggs in relation to offspring sex. Mothers allocated more antibodies against a pathogen to eggs with a daughter than a son. However, concentration of antioxidants was independent of embryonic sex. Sex-dependent allocation was independent of paternal attractiveness. Thus, mothers adjusted allocation of substances to offspring in a complex manner, that may be part of a strategy of favouritism of daughters, which have larger mortality than sons. Such effects may have important consequences for secondary and tertiary sex ratios, but also for ontogeny of adult phenotype.

Animals↗

Humoral immune response in relation to senescence, sex and sexual ornamentation in the barn swallow (Hirundo rustica).

Performance of animals may decline with age. The effects of senescence, however, may differ between the sexes because of differences in physiology and behaviour. Acquired immunity provides hosts with efficient mechanisms of anti-parasite defence, but the effect of senescence on immunocompetence has never been studied in natural populations. In the barn swallow (Hirundo rustica), primary antibody response to an antigen during one breeding season declined with age in females, while secondary response during the following breeding season declined with age in both sexes. Parasite-mediated sexual selection theory posits that male secondary sexual characters reveal resistance to parasites. Males with large tail ornaments had stronger primary response, retained larger antibody levels until the following year, but did not differ in secondary response compared with short-tailed males, as predicted if ornamentation reflects resistance to parasites. This is the first study showing that immunocompetence declines with age in any vertebrate under natural conditions.

Aging↗

DDT homologues and PCBs in eggs of great crested grebe (Podiceps cristatus) and mallard (Anas platyrhynchos) from Lake Maggiore (Italy).

DDT homologues and PCBs were determined in eggs of two stationary species, great crested grebe (Podiceps cristatus) and mallard (Anas platyrhynchos), in three breeding areas of Lake Maggiore in 1998. As expected, much higher contamination levels of both DDTs and PCBs were found in the fish-eating great crested grebe than in the mallard. In many eggs these values exceeded the safety thresholds proposed for aquatic bird protection. However, no significant correlation could be found between the two classes of pollutants and shell thickness in great crested grebe eggs. While a DDT manufacturer was detected as the main source of DDT pollution in the Lake Maggiore environment, the reason for the surprisingly high PCB level remains unknown.

Animals↗

Offspring sexual dimorphism and sex-allocation in relation to parental age and paternal ornamentation in the barn swallow.

We analysed the morphology of nestling barn swallows (Hirundo rustica) in relation to their sex, and laying and hatching order. In addition, we studied sex-allocation in relation to parentage, parental age and expression of a secondary sexual character of fathers. Molecular sexing was conducted using the sex chromosome-linked avian CHD1 gene. Sex of the offspring was not associated with laying or hatching order. None of nine morphological, serological and immunological variables varied in relation to offspring sex. Sexual dimorphism did not vary in relation to parental age and expression of a paternal secondary sexual character. The proportion of sons declined with brood size. Individual males and females had a similar proportion of sons during consecutive breeding years. The proportion of sons of individual females declined with age, but increased with the expression of a secondary sexual character of their current mate. The generalized lack of variation in sexual dimorphism among nestlings may suggest that barn swallows do not differentially invest in sons vs. daughters. Alternatively, male offspring may require different parental effort compared to their female siblings in order to attain the same morphological state. The lack of variation in offspring sexual dimorphism with paternal ornamentation suggests no adjustment of overall parental effort in relation to reproductive value of the two sexes. However, male-biased sex ratio among offspring of highly ornamented males may represent an adaptive sex-allocation strategy because the expression of male ornaments is heritable and highly ornamented males are at a sexual selection advantage.

Age Factors↗

Better red than dead: carotenoid-based mouth coloration reveals infection in barn swallow nestlings.

Nestling birds solicit food from their parents by displaying their open brightly coloured gapes. Carotenoids affect gape colour, but also play a central role in immunostimulation. Therefore, we hypothesize that, by differentially allocating resources to nestlings with more brightly coloured gapes, parents favour healthy offspring which are able to allocate carotenoids to gape coloration without compromising their immune defence. We demonstrated that, in the barn swallow Hirundo rustica, (i) parents differentially allocate food to nestlings with an experimentally brighter red gape, (ii) nestlings challenged with a novel antigen (sheep red blood cells, SRBCs) have less bright gape colour than their control siblings, (iii) nestlings challenged with SRBCs but also provided with the principal circulating carotenoid (lutein) have more brightly coloured red gapes than their challenged but unsupplemented siblings and (iv) the gape colour of nestlings challenged with SRBCs and provisioned with lutein exceeds that of siblings that were unchallenged. This suggests that parents may favour nestlings with superior health by preferentially feeding offspring with the brightest gapes.

Animals↗

Genetic and environmental components of phenotypic variation in immune response and body size of a colonial bird, Delichon urbica (the house martin).

Directional selection for parasite resistance is often intense in highly social host species. Using a partial cross-fostering experiment we studied environmental and genetic variation in immune response and morphology in a highly colonial bird species, the house martin (Delichon urbica). We manipulated intensity of infestation of house martin nests by the haematophagous parasitic house martin bug Oeciacus hirundinis either by spraying nests with a weak pesticide or by inoculating them with 50 bugs. Parasitism significantly affected tarsus length, T cell response, immunoglobulin and leucocyte concentrations. We found evidence of strong environmental effects on nestling body mass, body condition, wing length and tarsus length, and evidence of significant additive genetic variance for wing length and haematocrit. We found significant environmental variance, but no significant additive genetic variance in immune response parameters such as T cell response to the antigenic phytohemagglutinin, immunoglobulins, and relative and absolute numbers of leucocytes. Environmental variances were generally greater than additive genetic variances, and the low heritabilities of phenotypic traits were mainly a consequence of large environmental variances and small additive genetic variances. Hence, highly social bird species such as the house martin, which are subject to intense selection by parasites, have a limited scope for immediate microevolutionary response to selection because of low heritabilities, but also a limited scope for long-term response to selection because evolvability as indicated by small additive genetic coefficients of variation is weak.

Analysis of Variance↗

Carotenoids, sexual signals and immune function in barn swallows from Chernobyl.

Carotenoids have been hypothesized to facilitate immune function and act as free-radical scavengers, thereby minimizing the frequency of mutations. Populations of animals exposed to higher levels of free radicals are thus expected to demonstrate reduced sexual coloration if use of carotenoids for free-radical scavenging is traded against use for sexual signals. The intensity of carotenoid-based sexual coloration was compared among three populations of barn swallows Hirundo rustica differing in exposure to radioactive contamination. Lymphocyte and immunoglobulin concentrations were depressed, whereas the heterophil:lymphocyte ratio, an index of stress, was enhanced in Chernobyl swallows compared to controls. Spleen size was reduced in Chernobyl compared to that of two control populations. Sexual coloration varied significantly among populations, with the size of a secondary sexual character (the length of the outermost tail feathers) being positively related to coloration in the two control populations, but not in the Chernobyl population. Thus the positive covariation between coloration and sexual signalling disappeared in the population subject to intense radioactive contamination. These findings suggest that the reliable signalling function of secondary sexual characters breaks down under extreme environmental conditions, no longer providing reliable information about the health status of males.

Animals↗

Immunocompetence, ornamentation, and viability of male barn swallows (Hirundo rustica).

Immunocompetence (i.e., the ability to produce an immune response to pathogens) can be predicted to influence the chances that organisms have to survive and reproduce. In this study we simulated a challenge to the immune systems of male barn swallows (Hirundo rustica) by injecting them intraperitoneally with a multigenic antigen, sheep red blood cells, and we analyzed long-term survival in relation to their immunocompetence. Males were assigned to four groups that differed for the treatment of the length of the outermost tail feathers, a sexually dimorphic ornamental character that is currently under directional sexual selection. Immunocompetence was measured as change of concentration of gamma globulins relative to plasma proteins. The intensity of the immune response was independent of age. Males that showed the highest short-term response to sheep red blood cells were more likely to survive until the breeding season following that in which they had been inoculated, a pattern consistently observed within each experimental group. Males with comparatively long tails were more likely to survive than those with short tails. To our knowledge, the results of this study are the first to demonstrate that immunocompetence can predict long-term survival in a free-ranging vertebrate. Moreover, they are compatible with current models of parasite-mediated sexual selection because long-tailed males are more immunocompetent than short-tailed ones, and females, by preferring to mate with the most ornamented males, may acquire the "good genes" for high immunocompetence and, hence, for high viability of their offspring.

Animals↗

Parasites, immunology of hosts, and host sexual selection.

Parasite-mediated sexual selection is reviewed with special emphasis on the bird literature. Choosy females may benefit from choosing parasite-free mates if such males provide better parental care, do not transmit contagious parasites, or provide resistance genes to offspring. There is evidence in support of each of these mechanisms. The immunocompetence handicap hypothesis posits that secondary sexual characters reliably reveal the ability of males to resist parasites due to the immunosuppressive effects of testosterone and other biochemicals. Several aspects of these negative feedback mechanisms are supported by laboratory studies, but evidence from free-living animals is almost completely absent. Corticosterone rather than testosterone may potentially mediate the immunocompetence handicap mechanism. A simple version of the immunocompetence handicap is developed suggesting that body condition of male hosts is a sufficient mediator of the handicap mechanism of reliable sexual signaling. Sexual selection appears to be more intense in sexually dichromatic bird species, and comparative studies using pairwise comparisons of closely related taxa reveal that sexually dichromatic bird species have larger spleens, larger bursa of Fabricius, and higher concentrations of leukocytes than monochromatic species. Parasite-mediated sexual selection is proposed to affect parasite biology by increasing (1) the variance-to-mean ratio in parasite abundance, (2) variance in the intensity of natural selection affecting hosts, and (3) speciation rates among parasites exploiting hosts subject to intense sexual selection as compared to those subject to less intense selection.

Animals↗