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.     

Life histories of Clethrionomys and Microtus (Microtinae)

Although there have been numerous life-history reviews of mammals at high taxonomic levels (e.g. among families within orders), there are far fewer studies at lower taxonomic levels (e.g. among species within genera). Data on adult weights, litter size, gestation length, neonate weight, age and weight at weaning, growth rate to weaning, maximum life span, and length of the breeding season were compiled from the literature on five species of Clethrionomys and 33 species of Microtus. Variability in litter size and male body weights was not significantly different when compared between cyclic and non-cyclic populations. Coefficients of variation were also calculated for the three species with the most data (C. gapperi, C. glareolus and M. pennsylvanicus ). These values showed that the amount of intraspecific variation differed among traits as well as among species. Gestation length was the most invariable of all traits and variation in adult weights, neonate weight, gestation length, and litter size had similar values to those reported for Peromyscus maniculatus. Five and eight traits differed among Clethrionomys and Microtus species, respectively. Differences in litter size, adult weights and length of the breeding season were common to both genera. Male weight, gestation length and neonate weight as well as length of the breeding season were different between genera. Very few traits covaried within C. gapperi, C. glareolus or M. pennsylvanicus. Similarly, few traits covaried among all Clethrionomys populations. However, among all Microtus populations and Microtus species, 11 and 12 correlations were significant. Many of the patterns found in Microtus involved positive relationships between female weight and some other trait. These patterns have also been found by broader surveys at higher taxonomic levels. Large species of Microtus had larger offspring, a greater litter size and occurred in short-season environments relative to small species of this genus.     

Predation by Tawny owls (Strix aluco) on Bank voles (Clethrionomys glareolus) and Wood mice (Apodemus sylvaticus)

During 1954–56 we made a study of the numerical relationships between a population of Tawny owls ( Strix aluco ) and populations of their main prey species, the Bank vole ( Clethrionomys glareolus ) and the Wood mouse ( Apodemus sylvaticus ), in a 48.6-ha area of mainly deciduous woodland near Oxford. The owls were censused by plotting their territorial challenges (hooting), the rodents by the capture-mark-recapture technique. Since the rodents were marked with numbered monel metal leg rings, we were able to recover a proportion of these in the pellets of undigested material cast by the owls.
The results for Bank voles, which were the more numerous of the two prey species, indicated that 20–30% of the standing crop was removed by owls in any two-month period. We also made an independent estimate of predation rate from the number of rings recovered from owl pellets in relation to the number of ringed rodents released into the population each two months. This coincided with the limits indicated by the first method for voles (20-30% removed of the standing crop each 2 months).
Wood mice were scarcer than Bank voles and were not amenable to satisfactory capturemark-recapture analysis but, when treated on the system of recovery of rings in owl pellets, we found that they were preyed upon relatively more heavily than were the voles. Of the latter 18–46% of the marked animals were recovered in the owl pellets compared with 28–70% of the marked mice. Either the mice were preferred prey or they were more vulnerable to owl predation by reason of their preference for more open habitats.     

棕背Ping昼夜活动节律的研究

The diurnal activity rhythm of Clethrionomys rufocanus was studied under three different food conditions in a laboratory. The activity occurred both in the daytime and at night, but mainly at night (19:00--4:00). The mice fed with Mouse food exhibited six small peaks in their daily activity, and those fed with grass or hay exhib-ited seven, but the peak of the out-nest night activity was 2 ～ 3 hours earlier. The activity amount of out-nest was 30 518.0 ± 3 694.9 s for those fed with grass, the next was 21 811.7 ± 2 288.0 s for those fed with hay,and the least was 15 038.0 ± 666.0 s for those fed with mouse food. The activity amount of feeding was 10867.3± 1 612.1 s for those fed with grass, much more than those fed with mouse food and hay. The activity amount for drinking was 988.5 ± 79.1 s (fed with hay), 568.9 ± 60.9 s (fed with mouse food) and 139.3 ±47.2 s (fed with grass).     

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).     

Laboratory breeding of the red-backed vole (Clethrionomys rufocanus bedfordiae)]

Laboratory matings were attempted to establish breeding colonies of red-backed voles (Clethrionomys rufocanus bedformidae) as experimental animals. For these mating, 10 pairs of red-backed voles which were captured in the Tohbetsu region of Hokkaido, Japan and their litters were used. In the results for two years, 1987 to 1988, the rates of pregnancy, birth and weaning were 35.4%, 94.5% and 79.5%, respectively. The mean litter size was 5:1 +/- 1.6 with a range of 1 to 9. The mean gestation period was 20.0 +/- 0.7 days with a range of 18 to 22. These results suggest that planned production of red-backed voles in the laboratory is possible. To determine intraregional variations of red-backed voles with a view to the establishment of a strain by inbreeding, restriction patterns of mitochondrial DNAs using seven restriction endonucleases were compared. Four different patterns were obtained from wild red-backed voles used in the present study.     

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.     

Hormonal requirements and the influence of age on the decidual cell reaction in the bank vole (Clethrionomys glareolus)

Laboratory bred bank voles of different age and reproductive status were bilaterally ovariectomized, treated with exogenous oestrogen or progesterone or both, and given an artificial decidual stimulus (crushing with artery forceps). An hormonal regimen of oestrogen (100 ng) followed by progesterone (1 mg) and small amounts of oestrogen (10 ng) was necessary for maximal development of the decidual cell reaction (DCR). A DCR did not develop if the animals were treated with only oestrogen or progesterone. Middle-aged virgin females 5-7 months old had larger responses than did young virgin females 2-3 months old, and parous females, 15-19 months old and having had at least 10 litters, had a response similar to that of virgin females of about the same age. The size of the DCR in old parous females was inversely related to the time elapsing since the last litter. It is concluded that in its hormonal requirements for a successful DCR the bank vole is similar to the mouse. The results also suggest that the increase in fertility of bank voles up to the age of about 100 days may in part be explained by increased response of the endometrium to blastocysts. The reduction in size of the DCR in old animals is a consequence of ageing rather than of repeated pregnancies.     

Influence of age and artificial vaginal stimulation on fertility in female bank voles (Clethrionomys glareolus)

Female bank voles mated when 30-50 days old showed only 20-25% fertility at the first mating, but fertility increased to about 80% in animals greater than 100 days of age at the first mating. Young females subjected to artificial cervical stimulation after the completion of a normal mating showed a much higher fertility than normally mated controls (76 compared with 29%). The reason for the low fertility therefore appears to be that young females require more stimulation to activate the corpora lutea than is normally received at a mating.     

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.     

Seasonal effects on sexual maturation of male bank voles (Clethrionomys glareolus)

The weights of the testes and accessory glands, and the degree of morphological and functional development of the seminiferous tubules, were used as indicators of sexual maturation in male bank voles. Males reared under constant laboratory conditions showed a significant relationship between the seasons and sexual maturation. Young males reached maturity most rapidly in the reproduction season (mid-April to mid-October), while adult males matured earlier, in the spring season (mid-January to mid-April). The influence of season on animals reared in outdoor cages was much more pronounced. One of the important factors was the photoperiod: animals reared from 3 to 12 weeks of age in a short photoperiod (8L:16D) matured less rapidly than did those reared in 16L:8D.     

Parental care in captive red-backed vole (Clethrionomys rutilus)

Parental behavior in ten pairs of male and female red-backed voles, Clethrionomus rutilus, was studied under laboratory conditions. Significant sex differences were found in total time spent in the nest, time spent by adults alone in the nest, grooming time, and nest-building activity; in half of the males under study, the level of paternal care was extremely low. These findings indicate different paternal contributions of female and male red-backed voles into the rearing of young, in agreement with the spatial-ethological structure of populations of this species. Parental behavior of C. rutilus is discussed in comparison with that of other species of the subfamily Microtinae.     

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.     

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.     

Postglacial expansion of the southern red-backed vole (Clethrionomys gapperi) in North America

Dynamic climatic oscillations of the Pleistocene dramatically changed the distributions of high latitude species. Molecular investigations of a variety of organisms show that processes of postglacial colonization of boreal regions were more complex than initially thought. Phylogeographical and coalescent analyses were conducted on partial sequences of the cytochrome b gene (600 bp) from 64 individuals of Clethrionomys gapperi from North Carolina, Pennsylvania, Minnesota, Idaho, Washington, British Columbia, Northwest Territories, and Alaska to test hypotheses relating to Pleistocene refugia and postglacial colonization routes. Three divergent clades (east, west, central) were identified with highest net divergence (dA = 5.2%) between the eastern and western clades. Populations from the recently deglaciated higher latitudes of Canada and Alaska are closely related to lower latitude populations of the central clade (dA = 1.2%) suggesting recent expansion from this midwestern region. No representatives from the east or west clade were found at latitudes higher than 50 degrees N, indicating that postglacial colonization occurred through a midcontinental route. The high latitude population from the Northwest Territories exhibited demographic patterns and genetic diversity consistent with a stable noncolonizing population. This population is found near the Mackenzie range, where the two continental ice sheets were believed to have coalesced. Molecular variation observed in this population may be the result of leading edge population diversifying in the continental corridor or may reflect the signal of a high latitude refugial population.     

The long-term effect of cadmium exposure through food on the postnatal development of the bank vole (Clethrionomys glareolus Schreber, 1780)

Cadmium is well known for its toxicity to the animal body. However, its effect on pregnancy and the development of young animals is still not well understood. This study examined such effects, using bank voles captured from the wild to make the results closer to those which could be expected in the natural environment. One group of animals was fed 7 microg g(-1) cadmium in the food, a second 35 microg g(-1), and a third no cadmium, as a control. The concentrations of cadmium in the whole bodies of young bank voles were determined on the 3rd, 5th, or 10th day of life. The cadmium level in the bodies of animals exposed to 35 microg g(-1) of cadmium was significantly higher than in those from either the control group or the group receiving 7 microg g(-1) of cadmium, which did not differ from each other. The cadmium level did not change with animal age in any of the study groups. Concentrations of Zn, Cu, and Fe were also determined in the whole body of young animals, as cadmium is known to disturb the metabolism of these essential metals through antagonistic activity. Both Cu and Fe levels were negatively correlated with cadmium concentrations, while a positive correlation was found between zinc and cadmium in the young animal bodies. Also found was higher offspring mortality in the group receiving 35 microg g(-1) of cadmium in food. There was no difference in young animal body weight between the study groups.     

Photoperiod-induced changes in the testicular metabolism of [4-14C]17 alpha-hydroxyprogesterone in the bank vole (Clethrionomys glareolus)

Minces of the testes of bank voles, born and reared in a long (18L:6D) photoperiod until weaning (18-22 days of age) and subjected thereafter to a short (6L:18D, Group S) or a long (18L:6D, Group L) photoperiod for 6-9 weeks, were incubated with [4-14C]17 alpha-hydroxyprogesterone in the presence of cofactors (NADP/NADPH, 1.3 mmol/1) for 1 h at 37 degrees C. The radioactive metabolites were characterized and identified by thin-layer chromatography with derivative formation and chromatography to constant specific activity and isotope ratio. In Group L virtually all of the substrate was utilized and it was readily converted to androgens (48% of the radioactivity recovered) such as androstenedione and testosterone. The only pregnane metabolite identified was 17 alpha-hydroxy,20 alpha-dihydroxyprogesterone (43.3%). In Group S there was a decreased production of 17 alpha-hydroxy,20 alpha-dihydroprogesterone and androgens (25.4% and 10.4% respectively) and a substantial portion of the substrate was not metabolized (38.8%). The main androgen metabolites identified, androst-4-ene-3 beta,17 beta-diol and 5 alpha-androstane-3,17-dione are hormonally quite inert steroids. No androstenedione or testosterone was found. The results indicate that exposure to short photoperiod induces a decrease in the testicular C17-C20 lyase and 20 alpha-hydroxysteroid dehydrogenase.     

DNA sequence variation in the mitochondrial control region of red-backed voles (Clethrionomys)

The complete mitochondrial DNA (mtDNA) control region was sequenced for 71 individuals from five species of the rodent genus Clethrionomys both to understand patterns of variation and to explore the existence of previously described domains and other elements. Among species, the control region ranged from 942 to 971 bp in length. Our data were compatible with the proposal of three domains (extended terminal associated sequences [ETAS], central, conserved sequence blocks [CSB]) within the control region. The most conserved region in the control region was the central domain (12% of nucleotide positions variable), whereas in the ETAS and CSB domains, 22% and 40% of nucleotide positions were variable, respectively. Tandem repeats were encountered only in the ETAS domain of Clethrionomys rufocanus. This tandem repeat found in C. rufocanus was 24 bp in length and was located at the 5' end of the control region. Only two of the proposed CSB and ETAS elements appeared to be supported by our data; however, a "CSB1-like" element was also documented in the ETAS domain.     

Circadian organization of a subarctic rodent, the northern red-backed vole (Clethrionomys rutilus)

Arctic and subarctic environments are exposed to extreme light: dark (LD) regimes, including periods of constant light (LL) and constant dark (DD) and large daily changes in day length, but very little is known about circadian rhythms of mammals at high latitudes. The authors investigated the circadian rhythms of a subarctic population of northern red-backed voles (Clethrionomys rutilus). Both wild-caught and third-generation laboratory-bred animals showed predominantly nocturnal patterns of wheel running when exposed to a 16:8 LD cycle. In LL and DD conditions, animals displayed large phenotypic variation in circadian rhythms. Compared to wheel-running rhythms under a 16:8 LD cycle, the robustness of circadian activity rhythms decreased among all animals tested in LL and DD (i.e., decreased chi-squared periodogram waveform amplitude). A large segment of the population became noncircadian (60% in DD, 72% in LL) within 8 weeks of exposure to constant lighting conditions, of which the majority became ultradian, with a few individuals becoming arrhythmic, indicating highly labile circadian organization. Wild-caught and laboratory-bred animals that remained circadian in wheel running displayed free-running periods between 23.3 and 24.8 h. A phase-response curve to light pulses in DD showed significant phase delays at circadian times 12 and 15, indicating the capacity to entrain to rapidly changing day lengths at high latitudes. Whether this phenotypic variation in circadian organization, with circadian, ultradian, and arrhythmic wheel-running activity patterns in constant lighting conditions, is a novel adaptation to life in the arctic remains to be elucidated.