The structure of a chromosomal hybrid zone of house mice (Mus domesticus) in central Italy: cytogenetic analysis

Parapatric hybridization between the chromosomal race “CD” (2n = 22) and standard karyotype populations (2n = 40) of Mus domesticus occurs extensively in central Italy. The present paper reports the results of a ctogenetic surve on a transect crossin the hybrid zone north of Rome. No clinal variation in eitier diloid nuders and chromosome frequencies was found to occur in this area, and drift seems to be responsible for the observed atchy pattern of variation. The previous assumption of a strong fertility reduction in structuray heterozyous hybrids contrasts sharply with the width (32 km) of the zone and the average structural aeterozygosity of the hybrid poulations. It is suggested that fitness of structural heterozygotes in nature is not strongly aPfected as has been inferred from laboratory experiments. The results of this study are discussed in context together with the role of hybrid zones in chromosomal speciation in Mus domesticus.     

WHERE ARE THE WORMY MICE? A REEXAMINATION OF HYBRID PARASITISM IN THE EUROPEAN HOUSE MOUSE HYBRID ZONE

Wormy mice in a hybrid zone have been interpreted as evidence of low hybrid fitness, such that parasites contribute to species separation. However, because of its natural heterogeneity, observations of parasite load must be numerous with good field area coverage. We sampled 689 mice from 107 localities across the Bavaria-Bohemia region of the European house mouse hybrid zone and calculated their hybrid indices using 1401 diagnostic single nucleotide polymorphisms (SNPs). We tested whether hybrids have greater or lesser diversity and load of parasite helminths than additive expectations, performing load analyses on the four most common taxa. We found hybrids have significantly reduced diversity and load of each of the commonest helminths; rarer helminths further support reduced load. Although within-locality comparisons have little power, randomization tests show the repeated pattern is unlikely to be due to local parasite heterogeneity, and simulations show a patch of low parasite diversity is unlikely to fall by chance just so in the field area, such that it produces the observed effects. Our data therefore contradict the idea that helminths reduce hybrid fitness through increased load. We discuss a vicariant Red Queen model that implies immune genes tracking parasites will escape Dobzhansky-Muller incompatibilities, generating hybrid variants untargeted by parasites.     

Adaptation and plasticity in insular evolution of the house mouse mandible

Morphometric methods allow the quantification of directions of phenotypic changes and their statistical comparison in a morphometric space. We applied this approach to investigate several candidate factors to explain changes in mandible shape occurring in house mice (Mus musculus domesticus, Mammalia, Rodentia) in Corsica and a nearby islet. The role of niche widening and of the concomitant change in diet was evaluated by comparing the micro‐evolutionary insular change to the macro‐evolutionary difference between omnivorous and herbivorous rodents. Phenotypic plasticity, which may contribute to rapid insular evolution, was assessed by breeding laboratory mice on hard versus soft food. The related change in mandible shape was compared with differences between continental and insular populations. The role of allometry was evaluated by assessing shape change related to size within the continental population and comparing this direction of change with differences on islands. Finally, evolution may be facilitated along the direction of the greatest phenotypic variance. This hypothesis was tested by computing in wild populations vectors corresponding to this direction and by comparing these vectors with those corresponding to estimates of shape changes related to plasticity, micro‐ and macro‐evolutionary processes. In Corsica, the congruence in directions of macro‐ and micro‐evolutionary phenotypic vectors (Corsican/continental mice versus omnivorous/herbivorous rodents) supports the hypothesis of adaptation in mandible shape evolution. By contrast, on the islet, phenotypic divergence follows directions of plastic response to food consistency as well as within‐population allometry. Thus, results suggest differences in the relative importance of processes which may influence rodent mandibular shape depending on the size of the islands they colonized. Faster evolution and plasticity may be more evident in small and often ephemeral populations living on small islands, whereas micro‐evolutionary processes may have enough time and genetic variability to progressively ‘align’ with macro‐evolutionary trends in large populations from big islands.     

The influence of sex and body size on nestling survival and recruitment in the house sparrow

Differences in the survival rates of males and females over the period from hatching to recruitment can have important impacts on individual fitness and population demographics. However, whilst the influence of an individual's sex on nestling growth and survival has been well studied, less is known about sex‐specific survival over the period between fledging and recruitment. Here, we analyse nestling survival and recruitment in an isolated, island population of house sparrows (Passer domesticus), using data collected over a 4‐year period. Nestlings that had a greater mass at 1 day old were more likely to fledge. Recruitment was also positively associated with day 11 mass. The positive influence of nestling mass on survival to fledging also increased as brood size increased. There was no difference in the survival of male and female individuals prior to fledging. In contrast, over the period from fledging to recruitment, females had significantly less mortality than males. Recruitment was also positively associated with 11‐day‐old mass. Neither the nestling sex ratio nor the fledging sex ratio deviated from 0.5, but the sex ratio amongst recruits was female biased. Our study shows that sex can influence juvenile survival, but also shows that its effect varies between different life‐history stages; therefore, these stages should be considered separately if we want to understand at what point sex‐specific differences in juvenile survival occur. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 680–688.     

Parental effort and response to nestling begging in the house sparrow: repeatability,heritability and parent–offspring co‐evolution

Parental effort has a direct impact on individual fitness. Theoretical models exploring how parental effort evolves to cope with offspring demand and sexual conflicts may differ in the assumptions they make in respect to the genetic heritability of parental behaviours. Only a few attempts, however, have been made to estimate the heritability of parental behaviours and their possible co‐evolution with offspring solicitation behaviour. Analysing parent and offspring behaviours in four generations of cross‐fostered broods of house sparrows, we found that parental effort (food delivery rate) was repeatable across consecutive broods and heritable across generations. In contrast, parental response to experimentally induced changes in nestling begging was neither repeatable across broods nor heritable across generations or correlated to nestling begging. Thus, the results give no indication for genetic covariance between begging intensity and parental response, but provide the first cross‐fostering‐based evidence for the heritability of parental investment levels across generations.     

Linkage-dependent gene flow in a house mouse chromosomal hybrid zone

In the alpine valley of Valtellina there are two Robertsonian chromosomal races of house mouse, the Poschiavo (POS: 2n = 24-26) characterized by metacentric 8.12 and acrocentrics 2 and 10 and the Upper Valtellina (UV: 2n = 22-24) characterized by metacentrics 2.8 and 10.12. The races inhabit separate villages in the valley except in Sommacologna and Sondalo, where they both occur together with hybrids. A total of 179 mice from 16 villages were typed at 13 microsatellite loci. Seven of these loci were localized close to the centromeres of chromosomes 10 and 12, with the prediction that these regions on the race-specific chromosomes would be the most likely to experience a barrier to gene flow. The remaining six loci were localized at the telomeres of chromosomes 10 and 12 and at the centromeres of chromosomes that do not differ between the races. Substantial differences in allelic frequencies were found between the villages with POS and UV races at five of the loci at the centromeres of chromosomes 10 and 12 but at none of the other loci. These differences were not found to distinguish the two races in Sommacologna and Sondalo. Therefore, the centromeric regions of race-specific chromosomes do appear to experience a barrier to gene flow, although this can break down under intense interbreeding between the races. These results are considered in the context of Harrison's (1990) concept of the semipermeability of hybrid zones to gene exchange and in relation to parapatric speciation.     

Sexual variation in heritability and genetic correlations of morphological traits in house sparrow (Passer domesticus)

Estimates of genetic components are important for our understanding of how individual characteristics are transferred between generations. We show that the level of heritability varies between 0.12 and 0.68 in six morphological traits in house sparrows (Passer domesticus L.) in northern Norway. Positive and negative genetic correlations were present among traits, suggesting evolutionary constraints on the evolution of some of these characters. A sexual difference in the amount of heritable genetic variation was found in tarsus length, wing length, bill depth and body condition index, with generally higher heritability in females. In addition, the structure of the genetic variance-covariance matrix for the traits differed between the sexes. Genetic correlations between males and females for the morphological traits were however large and not significantly different from one, indicating that sex-specific responses to selection will be influenced by intersexual differences in selection differentials. Despite this, some traits had heritability above 0.1 in females, even after conditioning on the additive genetic covariance between sexes and the additive genetic variances in males. Moreover, a meta-analysis indicated that higher heritability in females than in males may be common in birds. Thus, this indicates sexual differences in the genetic architecture of birds. Consequently, as in house sparrows, the evolutionary responses to selection will often be larger in females than males. Hence, our results suggest that sex-specific additive genetic variances and covariances, although ignored in most studies, should be included when making predictions of evolutionary changes from standard quantitative genetic models.     

Mucosal acidification and an acid microclimate in the hen colon in vitro

Experiments were performed on isolated, stripped colonic epithelia of low-salt-adapted hens (Gallus domesticus) in order to characterize acid secretion by this tissue. With symmetric, weak buffer solutions, colonic epithelia acidified both mucosal and serosal sides. Titration measurements of the mucosal acidification rate (pH-stat technique) averaged 1.63±0.25 Eq·cm^-2·h^-1. Mucosal acidification was also evident in colons from high-salt-adapted birds and in low-salt-adapted coprodeum, but was completely abolished in the high-salt coprodeum. Mucosal acidification by low-salt-adapted colonic epithelium was unaffected by sodium replacement, mucosal amiloride (10^-3 mol·l^-1), and serosal ouabain (5x10^-4 mol·l^-1), although all three treatments significantly reduced or reversed the short-circuit current. Acetazolamide (10^-3 mol·l^-1, serosal) reduced mucosal acidification by 15% and simultaneously increased short-circuit current by a similar amount. Colonic epithelia incubated in glucose-free solutions had significantly lower acidification rates (0.59±0.13 Eq·cm^-2·h^-1, P<0.002 versus controls) and addition of glucose (15 mmol·l^-1), but not galactose, partially restored acidification to control levels. Anoxia (N₂ gassing) completely inhibited short-circuit current, but reduced acidification by only 30%. A surface microclimate pH, nearly 2 pH units more acidic than the bath pH of 7.1–7.4 was measured in low-salt-adapted colon and coprodeum. The acid microclimate of both tissues was partially attenuated by adaptation to a high-salt diet. Colonic microclimate pH was dependent on the presence of glucose and sensitive to the bath pH. Histochemical staining for carbonic anhydrase localized this enzyme to cytoplasm and lateral margins of one subfraction of colonic cells, and to cytoplasm in a second subpopulation Intense staining was also evident in subepithelial capillaries. These results suggest that a large part of mucosal acidification and maintenance of the acid microclimate in hen colon may be dependent on glycolysis and metabolic acid production, although a smaller, electrogenic and acetazolamidesensitive component also appears to exist. This latter component may become more prominent under conditions of cellular acidification.Abbreviations CA carbonic anhydrase - I _SC short circuit current - NFM N-ethylmaleimide - PD transepithelial potential - SCFA short chain fatty acids     

Endogenous ecdysteroid levels and rates of ecdysone acylation by intact ovaries in vitro in relation to ovarian development in adult female house crickets,Acheta domesticus

Ecdysteroid titres have been determined in adult female house crickets (Acheta domesticus) in relation to reproductive maturation. Ecdysteroid levels in newly emerged adult females are low except in the gut and carcass, which probably reflect the remnants of the preecdysial ecdysteroid peak. Ecdysteroid levels in all compartments increase markedly once ovarian weight surpasses 10 mg. Apolar ecdysteroid conjugates (ecdysone 22-fatty acyl esters) predominate in ovarian tissue throughout ovarian maturation, but low levels of free ecdysteroid and polar conjugated ecdysteroids are also present. During this period, two peaks of ecdysteroids (mainly free and apolar conjugated ecdysteroids) are observed in the haemolymph, gut, and carcass compartments. The peaks in the haemolymph occur when the ovarian mass reaches 30 and 100 mg. The gut and carcass may be acting as sinks or sites of metabolism for the hormone released from the ovaries. The rate of ecdysone acylation by ovaries was found to be developmentally regulated, increasing from low levels in the immature ovaries of newly emerged females as the ovaries increase in size. A semiquantitative assay has been developed to identify compounds which inhibit the conversion of [³H] ecdysone into 22-fatty acyl [³H] ecdysone by ovaries in vitro. A number of ecdysteroids possessing a free hydroxyl group at C-22 as well as the side-chain stereochemistry of ecdysone effectively inhibit this conversion, probably by acting as competitive substrates. In the cases of 20-hydroxyecdysone and ponasterone A, it was clearly demonstrated that these compounds are converted to a mixture of C-22 fatty acyl esters. Several other compounds which have been sugested to affect ecdysteroid metabolism/mode of action in other systems were also tested for their effects on the acyltransferase activity of ovaries in vitro. Arch. Insect Biochem. Physiol. 35:279-299, 1997.© 1997 Wiley-Liss, Inc.     

Raciation and speciation in house mice from the Alps: the role of chromosomes

There are at least 24 different karyotypic races of house mouse in the central Alps, each characterized by a different complement of ancestral acrocentric and derived metacentric chromosomes; altogether 55 different metacentric chromosomes have been described from the region. We argue that this chromosome variation largely arose in situ. If these races were to make contact, in most cases they would produce F1 hybrids with substantial infertility (sometimes complete sterility), due to nondisjunction and germ cell death associated with the formation of long-chain and/or ring configurations at meiosis. We present fertility estimates to confirm this for two particular hybrid types, one of which demonstrates male-limited sterility (in accordance with Haldane's Rule). As well as a model for speciation in allopatry, the Alpine mouse populations are of interest with regards speciation in parapatry: we discuss a possible reinforcement event. Raciation of house mice appears to have happened on numerous occasions within the central Alps. To investigate one possible source of new karyotypic races, we use a two-dimensional stepping stone model to examine the generation of recombinant races within chromosomal hybrid zones. Using field-derived ecological data and laboratory-derived fertility estimates, we show that hybrid karyotypic races can be generated at a reasonable frequency in simulations. Our model complements others developed for flowering plants that also emphasize the potential of chromosomal hybrid zones in generating new stable karyotypic forms.