Social organization of the bank vole (Clethrionomys glareolus,Schreber 1780) and its demographic consequences: a model

Summary A unified model of social organization, spatial distribution, and demographic parameters in the bank vole was developed. It is based on social relations among females, among males and also between reproductive females and males. In the model, social status and reproductive condition of an individual depend exclusively on interactions with its nearest neighbours. A result of interactions between two neighbours remains local, i.e., it cannot affect other, more distant individuals. The simulated variables show similar trend and scatter as those found in a growing real population of the bank vole. The relevance of the model for theories of population dynamics is discussed.     

Patterns of host-parasite interactions between the nematode Heligmosomum mixtum (Schulz, 1952) and the bank vole (Clethrionomys glareolus Schreber, 1780)

Some aspects of the host-parasite interaction of the nematode Heligmosomum mixtum and the bank vole have been studied. The dependence of infestation on the host sex, age and weight, the seasonal and annual abundance dynamics are investigated. It has been found that the bank vole is the main host of H. mixtum; the distribution of H. mixtum abundances in the host population is described by a negative binomial distribution model. The greatest infestation is characteristic for mature bank voles; males are infected more heavily than females. Infestation with the nematode increases alongside with the host weight. Seasonal dynamics of the nematode abundance in a bank vole population appears as a curve with a maximum in January and a minimum in August; the infestation curve for 1-month-old animals captured from June through October has two peaks (in July and October), with a minimum in June. The number of parasites in the ecosystem over a long-term period changed synchronously with the host abundance. The synchronism in the host-parasite system dynamics is possible when the parasite has a short life span, and does not influence the host abundance.     

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.     

Influence of low-chitin krill meal on reproduction of Clethrionomys glareolus (Schreber, 1780)

1. Clethrionomys glareolus fed on a diet containing krill meal assimilated excessive amounts of fluoride.2. Excess of fluoride caused disturbances in the reproduction of these animals: reduction of the number of litters, higher mortality of offspring and degenerative changes in seminiferous tubules.3. The disturbances in the reproduction occurred still more distinctly in the successive generation.4. Among the bank vole offspring which received the diet with the greater amount of krill meal, both in the parent generation and in the generation F₁, a changed sex ratio was found, which was evidenced by a significantly higher number of males.     

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.     

Immunolocalization of androgen receptors in testicular cells during postnatal development of the bank vole (Clethrionomys glareolus, S.)

Determination of the cellular distribution of the androgen receptors within the testis is of great importance for an understanding of their essential role in mediating of androgen action in the male gonad. In bank voles, which are seasonally breeding rodents, photoperiod is one of the most important factors inducing profound changes in the morphology and hormonal activity of the testes. Immunolocalization of androgen receptors was found in all somatic cells such as Sertoli cells, Leydig cells, and peritubular-myoid cells, however, distribution of the androgen receptors in various cell types depended on age of animals. Intensity of immunoreactivity was noticed as age and photoperiod-dependent. Males reared under different light regimes showed a significant correlation between the length of light and sexual maturation. Therefore, morphology of the testis from young and adult bank voles was also presented.     

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.     

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.     

Low ambient temperature decreases cadmium accumulation in the liver and kidneys of the bank vole (Clethrionomys glareolus)

The importance of photoperiod and ambient temperature on the accumulation of cadmium in the liver and kidneys of bank voles was determined in the present study. Males and females, aged 1 month, were given 3.0 g Cd ml^–1 drinking water and divided into four groups according to photoperiod (16 h light/8 h dark and 8 h light/16 h dark) and ambient temperature (20 or 5°C); liver and kidneys were removed for cadmium as well as copper, iron and zinc analyses at the end of 6 weeks. Bank voles exposed to 5°C in both photoperiods consumed approximately 30% less water containing cadmium than those kept at 20°C. However, the total accumulation of cadmium in the liver and kidneys of males and females exposed to the low temperatures was 4.3–4.8 and 2.2–3.3 times less than that in animals maintained at room temperature in the long and short photoperiod, respectively. Simultaneously, the low temperature brought about an increase in the copper concentrations in the liver (12–43%) and kidneys (47–78%), giving rise to an inverse correlation between the cadmium accumulation and the tissue copper concentration. In contrast to cadmium and copper, the concentrations of iron and zinc were affected primarily by photoperiod. These findings indicate that ambient temperature is an important determinant of cadmium retention in the bank vole. It appears that low temperature decreases tissue cadmium accumulation not only by reducing cadmium intake but also through changes in copper metabolism.     

Photoperiodic modulation of testicular LH receptors in the bank vole (Clethrionomys glareolus)

The temporal changes in testicular binding of 125I-labelled hCG in juvenile bank voles (18 days of age, born and reared in a 18L:6D photoperiod) exposed to a long (18L:6D, Group L) or short (6L:18D, Group S) photoperiod for 0, 3, 7, 14 and 42-56 days were investigated. During testicular maturation, in Group L, there was a slight initial decrease in LH receptor numbers per testis followed by a marked prepubertal rise during the initial phase of rapid testicular growth after which a decrease took place. In Group S, during testicular regression, the temporal changes in LH receptor numbers per testis resembled those of Group L except that the corresponding increase in hCG binding during the initial week was considerably less marked and the receptor numbers remained thereafter at a significantly lower level than in Group L. Leydig cell count indicated that the observed changes in LH receptors per testis were due to changes in the number of Leydig cells as well as in LH receptors per Leydig cell. The present results indicate, that (1) photoperiod is an important modulator of testicular LH receptor numbers in this species, (2) photoperiod or age has no significant effect on the binding affinity of LH receptors, (3) short photoperiods arrest the induction of LH receptors as well as the increase in Leydig cell numbers associated with normal testicular maturation, and (4) changes in LH receptor numbers per testis correlate well with the photoperiod-induced changes in androgen biosynthesis, spermatogenesis and Leydig cell morphology observed in our previous studies.     

Dietary cadmium decreases lipid peroxidation in the liver and kidneys of the bank vole (Clethrionomys glareolus).

The effect of elevated levels of dietary cadmium on lipid peroxidation in the liver and kidneys of a small rodent, the bank vole, was determined in the present study. Males and females, aged 1 month, were given diets containing 0.40 and 80 mg Cd per kg; liver and kidneys were removed for TBA-RS as well as iron, copper, zinc, cadmium and metallothionein analyses at the end of 6 weeks. Dietary Cd significantly decreased the TBA-RS level in the liver and kidneys of both sexes; however, this effect appeared to be dose-dependent only for the male liver. The changes in hepatic and renal TBA-RS paralleled closely those of tissue iron. Copper concentration decreased significantly only in the male liver, while hepatic and renal zinc were not influenced by dietary Cd. The concentrations of Cd and metallothionein in the liver and kidneys increased significantly in a dose-dependent fashion. Regression analysis confirmed that TBA-RS in both organs correlated closely with iron. The data suggest that dietary Cd decreases hepatic and renal lipid peroxidation indirectly, through lowering the tissue iron concentration.     

Adaptation of the bank vole (Clethrionomys glareolus Schreber) to conditions of biogeochemical province with abnormally high content of nickel, cobalt and chromium

Morphophysiological characteristics and peculiarities of adrenal gland of bank vole (Clethrionomys glareolus) were studied in the area of natural biogeochemical province with abnormally high content of nickel, cobalt and chromium. The control population inhabited area with usual content of these elements. We used 4-factor analysis of variance to estimate the influence of geochemical conditions, phase of population cycle, sex and reproductive state on the morphophysiological characteristics of animals and functional activity of adrenal gland. Animals from area with high concentration of Ni, Co and Cr show an increase in relative mass of adrenal glands, fascicular zone of adrenal cortex, size of cells and their nuclei. All these changes can be considered as an evidence of increased secretion of glucocorticoids. It is shown that phase of population cycle influences fatness of animals, size of nuclei, cells and adrenal cortex. Females in comparison with males are characterized with higher indexes of liver and adrenal gland, as well as morphometric indexes of adrenal cortex. The maturation of animals is accompanied with increase in body mass, fatness and relative mass of adrenal glands, the size of cortex zone, nuclei and cells themselves. It is supposed that the effect of "geochemical factor" results in intensification of glucocorticoid secretion of adrenal costex, thus increasing non-specific resistance of animals inhabiting area with high concentration of heavy metals. Such factors as "phase of population cycle", "sex" and "reproductive state", influence mineralocorticoid activity, glucocorticoid and androgenic functions of adrenal cortex. Some factors show synergetic effect.     

Limitation of reproductive success by food availability and litter size in the bank vole,Clethrionomys glareolus

Food limitation has been suggested as one of the most important factors affecting life history evolution in terrestrial vertebrates. However, this inference is based mainly on evidence from birds, and reproductive trade-offs may differ among groups with different forms of parental care. To study whether the costs of enlarged litters (decreased mass of offspring) would appear when food is not limiting, we performed outdoor enclosure experiments in which we manipulated simultaneously the litter size (control versus litter plus two pups) and food availability (control versus food-supplemented) of female bank voles, Clethrionomys glareolus. The weaning success of females increased significantly in response to supplementary food. When females were provided with extra food, no differences were observed in body masses of weanlings of control and enlarged litters. Further, food-supplemented females grew to larger sizes during nursing than unsupplemented females. Our experiment suggests that energetic requirements during nursing constrain the number of offspring that can be raised successfully in a particular breeding attempt. The results also indicate that unlimited food resources may increase future reproductive potential of females because they can use more energy for somatic growth.     

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.     

Blood system peculiarities in the bank vole (Clethrionomys glareolus) under chronic environmental pollution

The parameters of peripheral blood and hemopoietic organs in mature and immature bank voles inhabiting a chemically polluted area were studied. Variability of the blood system parameters depending on the level of toxic load and the animals’ reproductive status was determined. Alteration of the cell composition of erythrocytes and leucocytes, the structure of erythrocytes, and the hemoglobin fractions and leucocyte functions describe the adaptive response to the factors of a changed environment more than the concentration of leucocytes, erythrocytes, and blood hemoglobin.     

Hepatic and renal cadmium accumulation is associated with mass-specific daily metabolic rate in the bank vole (Clethrionomys glareolus)

Previous study has shown that photoperiod and age affect tissue accumulation of cadmium (Cd) in a small rodent, the bank vole. Since the body mass is also influenced by these factors, the present study was designed to determine whether mass-specific daily metabolic rate might be responsible for differential accumulation of Cd in the liver and kidneys of the short- and long-photoperiod bank voles as well as of the young and old animals. One- and five-month old male bank voles were held under short (8 h light/16 h dark) or long (16 h light/8 h dark) photoperiods and exposed to dietary Cd (100 microg/g) for 6 weeks. The bank voles raised under the short photoperiod and those injected subcutaneously with melatonin (7 micromol/kg/day) under the long photoperiod showed significantly higher concentrations of Cd in the liver (43-60%) and kidneys (40-47%) than the age-matched long-photoperiod animals. The old bank voles accumulated significantly less Cd in both organs than the young animals. These differences in Cd accumulation appeared not to be associated with the relative Cd intake. However, the hepatic and renal Cd levels followed a pattern similar to that of the mass-specific daily metabolic rate (or energy expenditure) and energy assimilation efficiency. These data indicate that mass-specific daily metabolic rate and energy assimilation efficiency (an indicative of digestive and absorptive processes) may be responsible for differential tissue Cd accumulation in the bank vole.     

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.