Simple sequence repeats in zebra finch (Taeniopygia guttata) expressed sequence tags: a new resource for evolutionary genetic studies of passerines

Background Passerines (perching birds) are widely studied across many biological disciplines including ecology, population biology, neurobiology, behavioural ecology and evolutionary biology. However, understanding the molecular basis of relevant traits is hampered by the paucity of passerine genomics tools. Efforts to address this problem are underway, and the zebra finch (Taeniopygia guttata) will be the first passerine to have its genome sequenced. Here we describe a bioinformatic analysis of zebra finch expressed sequence tag (EST) Genbank entries. Results A total of 48,862 ESTs were downloaded from GenBank and assembled into contigs, representing an estimated 17,404 unique sequences. The unique sequence set contained 638 simple sequence repeats (SSRs) or microsatellites of length ≥20 bp and purity ≥90% and 144 simple sequence repeats of length ≥30 bp. A chromosomal location for the majority of SSRs was predicted by BLASTing against assembly 2.1 of the chicken genome sequence. The relative exonic location (5' untranslated region, coding region or 3' untranslated region) was predicted for 218 of the SSRs, by BLAST search against the ENSEMBL chicken peptide database. Ten loci were examined for polymorphism in two zebra finch populations and two populations of a distantly related passerine, the house sparrow Passer domesticus. Linkage was confirmed for four loci that were predicted to reside on the passerine homologue of chicken chromosome 7. Conclusion We show that SSRs are abundant within zebra finch ESTs, and that their genomic location can be predicted from sequence similarity with the assembled chicken genome sequence. We demonstrate that a useful proportion of zebra finch EST-SSRs are likely to be polymorphic, and that they can be used to build a linkage map. Finally, we show that many zebra finch EST-SSRs are likely to be useful in evolutionary genetic studies of other passerines

Testes asymmetry, condition and sexual selection in birds: an experimental test

The functional significance of the marked directional asymmetry in testes size observed in many bird species is obscure. Møller suggested that (i) the smaller of the two testes serves a compensatory role and increases in size (and hence reduces asymmetry) when the larger one is defective in some way, and (ii) as a consequence, the degree of directional asymmetry in testes size reflects male quality and covaries positively with the expression of secondary sexual traits.We conducted an experimental test of these two hypotheses in the zebra finch,Taeniopygia guttata. Neither hypothesis was supported. First, there was no significant relationship between the size of the left testis and relative testes asymmetry. Second, we obtained no support for the hypothesis that the degree of directional asymmetry in testes mass covaried with condition. On the contrary, directional asymmetry in testes mass was signifcantly greater in birds whose condition was experimentally reduced, compared with control birds. Moreover, we found no significant relationships between testes asymmetry and secondary sexual traits. We conclude that directional asymmetry in testes size does not reflect male condition in the zebra finch

Sperm competition and sperm midpiece size: no consistent pattern in passerine birds

Sperm competition is thought to be a major force driving the evolution of sperm shape and function. However, previous studies investigating the relationship between the risk of sperm competition and sperm morphometry revealed inconclusive results and marked differences between taxonomic groups. In a comparative study of two families of passerines (Fringillidae and Sylviidae) and also across species belonging to different passerine families, we investigated the relative importance of the phylogenetic background on the relationship between sperm morphometry and the risk of sperm competition. The risk of sperm competition was inferred from relative testis mass as an indicator of investment in sperm production. We found: (i) a significant positive association between both midpiece length and flagellum length and relative testis mass in the Fringillidae, (ii) a significant negative association between sperm trait dimensions and relative testis mass in the Sylviidae, and (iii) no association across all species. Despite the striking difference in the patterns shown by the Sylviidae and the Fringillidae, the relationship between midpiece length and flagellum length was positive in both families and across all species with positive allometry. Reasons for the differences and similarities between passerine families are discussed

The identity of the bird known locally in sixteenth- and seventeenth-century Norfolk, United Kingdom, as the Spowe

In the kitchen record books of the L'Estrange family in the sixteenth and seventeenth centuries, there are references to a bird, widely shot on the Norfolk coast, called a Spowe. On the basis of the similarity to the Icelandic name, J. H. Gurney (sen.) and Fisher (in their "An account of birds found in Norfolk" published in 1846) assumed this to be the Whimbrel (Numenius phaeopus) as have all ornithological texts ever since. Internal evidence from the kitchen records strongly suggest that the Spowe was a winter visitor, not a passage migrant, thus throwing considerable doubt on Gurney and Fisher's ascription. We suggest that it is much more likely that the Spowe was the Bar-tailed Godwit (Limosa lapponica)

Sophisticated sperm allocation in male fowl

When a female is sexually promiscuous, the ejaculates of different males compete for the fertilization of her eggs; the more sperm a male inseminates into a female, the more likely he is to fertilize her eggs. Because sperm production is limited and costly, theory predicts that males will strategically allocate sperm (1) according to female promiscuity, (2) saving some for copulations with new females, and (3) to females producing more and/or better offspring. Whether males allocate sperm in all of these ways is not known, particularly in birds where the collection of natural ejaculates only recently became possible. Here we demonstrate male sperm allocation of unprecedented sophistication in the fowl Gallus gallus. Males show status-dependent sperm investment in females according to the level of female promiscuity; they progressively reduce sperm investment in a particular female but, on encountering a new female, instantaneously increase their sperm investment; and they preferentially allocate sperm to females with large sexual ornaments signalling superior maternal investment. Our results indicate that female promiscuity leads to the evolution of sophisticated male sexual behaviour

Sophisticated sperm allocation in male fowl

When a female is sexually promiscuous, the ejaculates of different males compete for the fertilization of her eggs; the more sperm a male inseminates into a female, the more likely he is to fertilize her eggs. Because sperm production is limited and costly, theory predicts that males will strategically allocate sperm (1) according to female promiscuity, (2) saving some for copulations with new females, and (3) to females producing more and/or better offspring. Whether males allocate sperm in all of these ways is not known, particularly in birds where the collection of natural ejaculates only recently became possible. Here we demonstrate male sperm allocation of unprecedented sophistication in the fowl Gallus gallus. Males show status-dependent sperm investment in females according to the level of female promiscuity; they progressively reduce sperm investment in a particular female but, on encountering a new female, instantaneously increase their sperm investment; and they preferentially allocate sperm to females with large sexual ornaments signalling superior maternal investment. Our results indicate that female promiscuity leads to the evolution of sophisticated male sexual behaviour

The insemination window provides a distorted view of sperm competition in birds

The aim of this study is to identify and rectify a misunderstanding about the optimal timing of inseminations in birds. In species laying clutches of more that one egg, a copulation during the hour following egg-laying can result in sperm reaching the site of fertilization in time to fertilize the next egg to be laid. Cheng et al. (1983) referred to this period as the insemination window and proposed that it was an 'espcially favourable period' for males to obtain extra-pair copulations. As stated in their paper, this is true only in terms of the next ovum to be fertilized, but subsequent authors assumed that the insemination window represents a general peak in female fertility and have made predictions about the optimal timing of extra-pair behaviours and paternity guards relative to it. Far from being a general peak in female fertilty, we show by a re-analysis of Cheng et al.s data and by using published information on the domestic fowl Gallus domesticus, turkey Gallopavo meleagris and Muscovy duck Cairina Moschata, that inseminations either just after egg laying or just before it are much less likely overall to result in fertilization than inseminations made at other times. The reduced efficacy of inseminations made close to the time of egg-laying occurs because the retention of sperm by females inseminated at this time is low. The fact that inseminations made around the time of the egg laying in the domestic fowl, turkey and Muscovy duck have a reduced probability of fertilization is consistent with the fact that very few wild birds,even those in which sperm competition is intense, alter their copulation or mate guarding behaviour during the insemination window

Sperm precedence in the domestic fowl

The aim of this study was to examine last-male sperm precedence in the domestic fowl. We used sperm from two different genotypes to assign paternity: and in seven experiments females were artificially inseminated with either equal or unequal numbers of sperm at intervals of 4 or 24 h. We were unable to replicate the results of a previous study by Compton et al. (1978) in which a strong last-male precedence effect had been recorded when two equal sized inseminations were made 4 h apart. We observed no marked last-male sperm precedence and our results did not differ significantly from that predicted by the passive sperm loss model, in which a last-male effect is determined by the rate at which sperm are lost from the female tract and the interval between successive inseminations. The most likely explanation for the disparity between our result and Compton et al.'s is a difference in the timing of inseminations. The implications of this for studies of sperm competition in birds is discussed

A comparison of SNPs and microsatellites as linkage mapping markers: lessons from the zebra finch (Taeniopygia guttata)

Background: Genetic linkage maps are essential tools when searching for quantitative trait loci (QTL). To maximize genome coverage and provide an evenly spaced marker distribution a combination of different types of genetic marker are sometimes used. In this study we created linkage maps of four zebra finch (Taeniopygia guttata) chromosomes (1, 1A, 2 and 9) using two types of marker, Single Nucleotide Polymorphisms (SNPs) and microsatellites. To assess the effectiveness and accuracy of each kind of marker we compared maps built with each marker type separately and with both types of marker combined. Linkage map marker order was validated by making comparisons to the assembled zebra finch genome sequence. Results: We showed that marker order was less reliable and linkage map lengths were inflated for microsatellite maps relative to SNP maps, apparently due to differing error rates between the two types of marker. Guidelines on how to minimise the effects of error are provided. In particular, we show that when combining both types of marker the conventional process of building linkage maps, whereby the most informative markers are added to the map first, has to be modified in order to improve map accuracy. Conclusions: When using multiple types and large numbers of markers to create dense linkage maps, the least error prone loci (SNPs) rather than the most informative should be used to create framework maps before the addition of other potentially more error prone markers (microsatellites). This raises questions about the accuracy of marker order and predicted recombination rates in previous microsatellite linkage maps which were created using the conventional building process, however, provided suitable error detection strategies are followed microsatellite-based maps can continue to be regarded as reasonably reliable

The evolution of sperm morphometry in pheasants

Postcopulatory sexual selection is thought to be a potent evolutionary force driving the diversification of sperm shape and function across species. In birds, insemination and fertilisation are separated in time and sperm storage increases the duration of sperm female interaction and hence the opportunity for sperm competition and cryptic female choice. We performed a comparative study of 24 pheasant species (Phasianidae, Galliformes) to establish the relative importance of sperm competition and the duration of sperm storage for the evolution of sperm morphometry (i.e. size of different sperm traits). We found that sperm size traits were negatively associated with the duration of sperm storage but were independent of the risk of sperm competition estimated from relative testis mass. Our study emphasises the importance of female reproductive biology for the evolution of sperm morphometry particularly in sperm storing taxa