The structure of the glands in the angulus oris of the Bank vole Clethrionomys glareolus

The presence of glands opening to the angulus oris region of the bank vole Clethrionomys glareolus has been confirmed. The glands are similar in general to the apocrine sweat glands of man and other animals although histological differences were noted. Also there was a marked sex difference in the diameters of the lumens of the secretory tubules in this study (undertaken in the late autumn and early winter) The hairs into which the glands open are larger than other hairs in the region, and the surface pattern of the scales is inverted.     

The spread of the Bank vole Clethrionomys glareolus in Ireland

The discovery of the Bank vole Clethrionomys glareolus in Ireland and the published information on its spread are reviewed, and the results of a complete resurvey of its distribution, in 1 982 , presented. The rate of spread is extremely variable and somewhat unpredictable. Over suitable habitat—agricultural land with hedges and other cover (and sometimes mountain valleys)—spread may average 2–4.5 kmyr^-1 but can sometimes be very much slower. This might be influenced by the type and extent of hedges and of permanent pasture. High ground and moorland usually prevent spread but there have been dramatic exceptions. In 1982 Bank voles occupied probably the whole of Co. Limerick; most of Co. Kerry, with the exception of substantial parts of the mountainous peninsulas in the southwest and contiguous interior; a large part of Co. Cork, though not the south-east and again excepting portions of the mountainous south-west, together with a substantial strip of east Co. Tipperary, and the south-east quarter of Co. Clare.     

Haemoglobin polymorphism in the Bank vole, Clethrionomys glareolus, in Britain

Three hundred and twenty-two Clethrionomys glareolus from 17 localities in England, Wales and southern Scotland were typed for haemoglobin and two phenotypes were discovered. They have been designated HbS and HbF. The latter is more electronegative at physiological pH. The nature of the molecular difference between the phenotypes was examined and it was concluded that HbF polymerises more readily than does HbS. Haematological investigations showed that although the red cells of animals with HbF contain more potassium, thereby ensuring electrostatic neutrality, there has not been concomitant dilution of the cell contents. It is suggested that the osmotic stress imposed by this high concentration of particles might be reduced by in vivo polymerisation of haemoglobin. HbS was found to predominate in the north of Britain and HbF in the south. As none of the populations sampled contained animals of both types, it is not yet known whether there is a north-south cline in haemoglobin type, or a sharp disjunction.     

Plasma esterase polymorphism in the Bank vole, Clethrionomys glareolus, in Britain

Three hundred and eighty-three Clethrionomys glareolus from 20 localities in England, Wales and Scotland were typed for plasma esterase and a genetic polymorphism was discovered. The esterase was named Es-1. Breeding tests suggested that three alleles were segregating: Es-1^o when homozygous results in complete absence of enzyme activity. The active alleles Es-1^f and Es-1^s code for enzyme variants which migrate more rapidly and less rapidly, respectively, under starch gel electrophoresis. Of these active alleles, Es-1^f is morc common in the north of Britain and Es-1⁸ in the south. A 23-month field study on two areas at Wicken Fen, Cambridgeshire, suggested that animals possessing Es-1^s survived less well at high population densities, perhaps through their being more likely to emigrate.     

Effects of diet on reproductive performance of the Bank vole (Clethrionomys glareolus)

Laboratory experiments were carried out to test the effect of dietary supplements of fresh sprouted wheat on reproduction of the Bank vole, Clethrionomys glareolus. One hundred and twenty females were divided into three groups at weaning: (a) group NSS (no sprout supplement), (b) group LSS (low sprout supplement) and (c) group HSS (high sprout supplement). Six- to seven-week-old females were paired with fertile males. Both groups on sprout supplement showed a significantly higher incidence of pregnancy in primiparous females (76%) compared with controls (NSS) (46%). Diet had no significant effect on the incidence of pregnancy in multiparous females. Control females had consistently higher mean litter sizes up to and including the fifth litter, compared with experimental females. Primiparous NSS females had the highest mean litter size and the experimental group HSS had the lowest. Differences in mean litter interval between groups were related to litter size (number of sucklings) rather than diet. The number of foetuses in utero had no effect on litter interval. The incidence of post-partum conception was higher in group LSS as compared with groups HSS and NSS. Diet did not affect the mean number of young produced per female in a given time, or the percentage of young surviving to weaning. The results are discussed with reference to their implications regarding habitat suitability of C. glareolus.     

Fertility of the post-partum bank vole (Clethrionomys glareolus)

Analysis of records of a bank vole breeding colony suggests that fertility is high immediately post partum, declines during established lactation and rises after weaning of young. Mating tests with lactating females and females whose young had been removed at birth showed that receptivity is reduced during lactation, although amongst the females which did mate there was no difference between lactating and non-lactating animals in the proportion which produced litters. However, average size of litters at birth was significantly larger for the lactating than for the non-lactating females. There is some evidence suggesting that this difference may arise after ovulation has occurred. Virgin females were no more receptive or fertile than lactating females.     

Social overwintering and food distribution in the bank vole Clethrionomys glareolus

We studied overwintering in the bank vole Clethrionomys glareolus in four 0.5 ha enclosures in an abandoned field in central Finland in the winter 1987/88. In two of the enclosures food was offered evenly distributed over the whole enclosed area (Even Enclosures = EE), in the two others food was offered in one feeding patch with four feeding chambers 2 m apart (Patchy Enclosures = PE). Food was provided in about the same amount in both enclosures. The experiment commenced in early October, with 13 females and 11 males in EEs and 12 + 13 voles in PEs. After two months the voles in the PEs were concentrated around the feeding patches. Territoriality was not observed in EEs, instead the voles formed small exclusive overwintering groups consisting of 2-3 females and at least one male. The size of the home range of the females and males was identical during mid-winter as the voles were non-breeding. By the onset of breeding, range size increased in both sexes, but significantly more in males, however. The survival was about the same in all populations. Every population showed a mid-winter decline suggesting the effect of the mustelid predators observed in and around the enclosures. In the PEs the overwintering aggregations lasted until the maturation of the first litters. Food distribution affected the spatial distribution of the populations. We conclude that the patchiness of the habitat and especially the availability of food are the most important factors determining the social structure of overwintering populations.     

Offspring Defence in Relation to Litter Size and Age: Experiment in the Bank vole Clethrionomys glareolus

According to parental investment theory, nest defence activity should be related to the reproductive value of the offspring. Alternative hypotheses suggest that defence activity may, for example, depend upon the conspicuousness of the young. Studies concerning this topic have been carried out almost exclusively on birds and experimental data on the diversity of organisms is lacking. Bank voles Clethrionomys glareolus were used to study the effects of the number and age of offspring upon the pup defence activity of mothers. Male bank voles are infanticidal and thus an adult male was used as a predator. Defence trials were conducted in the laboratory and filmed for subsequent analysis. Litter sizes were divided into three treatment groups: reduced (−2 pups), control (±0 pups) and enlarged (+2 pups). In order to study the effect of offspring age upon maternal defence activity the trials were conducted twice: when the pups were 3 and 8 days old. Defence activity increased with the number of offspring and enlarged litters were most actively defended. This result supports parental investment theory and conclusions drawn by earlier studies of birds. However, in contrast to the conclusions of earlier studies, older offspring were defended less than the younger ones. Whilst new-born pups are totally defenceless against predators their vulnerability decreases as they age. Therefore, we suggest that maternal aggression in female bank voles is related to the value as well as to the vulnerability of the offspring. The validity of this explanation and the determinants of parental investment decisions in small mammals in general deserve further study. This revised version was published online in July 2006 with corrections to the Cover Date.     

The reproductive cycle of the Skomer vole (Clethrionomys glareolus skomerensis)

A histological examination has been made of the reproductive organs of the Skomer vole to study the reproductive cycle and fertility of this race of the Bank vole ( Clethrionomys glareolus ) in its insular habitat. The main aim of the work was to compare the cycle with that of Clethrionomys on the mainland of Britain which was similarly investigated by Brambell & Rowlands (1936).     

Highly polymorphic microsatellite loci in the bank vole (Clethrionomys glareolus)

We describe seven polymorphic, dinucleotide microsatellite loci isolated from bank voles (Clethrionomys glareolus, Rodentia: Muridae) collected from the Wirral Peninsula, United Kingdom. Microsatellites were isolated as part of a long‐term study on the wider effects of host–pathogen interactions of an endemic viral disease. These microsatellites showed between five and 13 alleles per locus in these populations. Observed and expected heterozygosities varied between 0.275 to 0.777 and 0.487 to 0.794, respectively. These markers will allow us to investigate the structure of this bank vole population.     

Some population dynamics of the Bank vole, Clethrionomys glareolus and the Wood mouse, Apodemus sylvaticus in mixed woodland

The population dynamics of Clethrionomys glareolus Schr.and Apodemus sylvaticus (L.) in mixed woodland in County Durham, were studied from March 1963 to January 1965. Two areas of 0.9 ha (2.25 acres) each were trapped monthly and information was obtained on population size, reproduction, survival and growth from marked animals.
Clethrionomys populations increased from June to an autumn peak; then declined, at first rapidly, but then more slowly in winter, before reaching a spring trough. The breeding season of Clethrionomys was from May to December; juveniles were caught from mid-June to December. Survival was in general poor during the breeding season but good at other times. Survival of young born early in the summer was particularly good on one of the areas and some individuals lived long enough to breed in two successive years. On both areas young born early in the year matured rapidly and bred in the year of their birth. Young born in late summer and early autumn ceased growing at a weight of about 14.5 g, remained immature, and formed the bulk of the overwintering population. Growth was completed at the time of sexual maturation the following spring.
Trapping failed to provide adequate samples of juvenile Apodemus in summer to account for subsequent recruitment. The possibility that a substantial proportion of the adult population of this species was also either trap shy or had emigrated temporarily is discussed. Breeding occurred from April until the following January, but the numbers trapped remained very low throughout the early months of the breeding season. Large scale recruitment of young fecund animals into the trap-revealed population occurred during the autumn. Apodemus males continued to grow rapidly during the winter.     

Maternal effort and male quality in the bank vole, Clethrionomys glareolus.

Parental investment in reproduction is adjusted according to potential benefits in terms of offspring survival and/or mating success. If male quality affects the reproductive success of a female, then females mating with high-quality males should invest more in reproduction. Although the subject has been of general interest, further experimental verification of the hypothesis is needed. We studied whether female bank voles (Clethrionomys glareolus) adjusted their maternal effort according to male quality, measured as mating success. To enable the measurement of maternal effort during nursing separately from male genetic effects the litters were cross-fostered. Further, the genetic background of male quality was examined. Male quality did not correlate with litter size or offspring size at birth. Offspring growth was positively related to food consumption and milk production of mothers. However, these direct measurements of maternal effort were independent of male quality. Male mating success appeared to be significantly heritable indicating that there are genetic benefits. Still, females did not adjust maternal effort according to the genetic quality of their offspring. We suggest that female bank voles gain significant genetic benefits from mating with high-quality males whereas they cannot improve their reproductive success by increasing maternal effort.     

Individual variation and repeatability of basal metabolism in the bank vole, Clethrionomys glareolus

Basal metabolic rate (BMR) is a fundamental energetic trait and has been measured in hundreds of birds and mammals. Nevertheless, little is known about the consistency of the population-average BMR or its repeatability at the level of individual variation. Here, we report that average mass-independent BMR did not differ between two generations of bank voles or between two trials separated by one month. Individual differences in BMR were highly repeatable across the one month interval: the coefficient of intraclass correlation was 0.70 for absolute log-transformed values and 0.56 for mass-independent values. Thus, BMR can be a meaningful measure of an individual physiological characteristic and can be used to test hypotheses concerning relationships between BMR and other traits. On the other hand, mass-independent BMR did not differ significantly across families, and the coefficient of intraclass correlation for full sibs did not differ from zero, which suggests that heritability of BMR in voles is not high.     

Avoidance of used endometrial sites by blastocysts of the bank vole, Clethrionomys glareolus

In bank voles blastocysts arising from mating immediately after parturition do not implant in used zones of the preceding pregnancy but in the regions between them. However, if mating is postponed until 12 or more days after parturition blastocysts do implant in used zones of the previous pregnancy. Implantations also occur at zones used two pregnancies previously. It seems likely that the absence of implantations at recently used zones of the uterus is determined by the uterus rather than blastocysts.     

Effect of photoperiod on cultured granulosa cells of the bank vole, Clethrionomys glareolus

Gonadal function of the bank vole females depends on the photoperiod. This experiment was to show whether photoperiod applied on the whole animal in vivo would affect the function of ovarian cells in vitro. Granulosa cells from large ovarian follicles of bank vole reared in long or short photoperiod were cultured as monolayers in control or luteinizing hormone supplemented media. Formation of cell colonies, activity of delta5, 3beta-hydroxy steroid dehydrogenase and progesterone secretion were investigated. First colonies of long day cells were formed already on day 1. On day 2 they enlarged and became abundant. Short day cells formed colonies only on day 2. Colonies of similar size to 2 day colonies of long day cells appeared only on day 6. There were also differences in steroid dehydrogenase activity and in progesterone secretion between long and short day control and hormone treated cultures. We conclude that photoperiod applied in vivo affects ovarian cell function in vitro.     

The occurrence of erythropoiesis in the thymus of the bank vole (Clethrionomys glareolus)

Summary The thymus of wild young and adult bank voles (Clethrionomys glareolus) was examined by histological methods for the presence of developing erythroid cells. Nucleated erythroid cells were observed in 26% of the glands examined by light microscopy and in 69% of the glands examined by electron microscopy.The largest number of developing erythroid cells was observed in the thymus of pregnant females, also showing raised reticulocyte counts (3.1–10.2%). However, erythropoiesis could also be found in breeding and non-breeding, first year and older animals.Erythroid cells were mainly located in the cortex, sometimes in small groups interspersed between lymphoid cells, and also randomly scattered in the cortex. Occasionally, macrocytic erythroid cells were also present. Pyknotic cells were commonly present, and granulopoiesis was frequently observed.