Aggressive behavior of adult meadow voles (Microtus pennsylvanicus) towards young

Summary Field evidence indicates that adult microtines, especially females, may be a major cause of poor juvenile survival and this may be instrumental in their population regulation. This suggests that males and females behave differently towards young animals. To examine how adult males, nonlactating females, and lactating females behave towards strange young, I introduced young animals into the home cage of the adults. Lactating females were most aggressive towards young; most males investigated them; most nonlactating females ignored them. However, discriminant function analysis indicated a great deal of overlap in the behavior of the classes. None of the classes behaved differently towards young of either sex. Females did not vary their behavior during lactation. Behavior varied among lactating females, with 55% showing little aggression and 20% showing a great deal. Most lactating females showed similar behavior in each of their bouts: docile females remained docile, aggressive females remained aggressive. I conclude that lactating females are most aggressive towards strange young. These results are consistent with the field evidence, which suggests that adult females depress juvenile survival and recruitment and that the main culprits are breeding females.     

Anemia at the onset of winter in the meadow vole (Microtus pennsylvanicus)

1. From 1981 to 1986, 6120 meadow voles (Microtus pennsylvanicus) were sampled for hematological indices in southeastern Manitoba, Canada. This survey revealed the sporadic occurrence of anemia in early winter at mean temperatures below about -5 degrees C. 2. Anemia was associated with leukocytosis and circulating normoblasts, suggesting a sudden, large blood loss. Individuals became anemic quickly, with no obvious predisposing factors. 3. Attempts were made to induce anemia by exposing voles in traps to various temperatures. Temperatures characteristic of most trapping sessions failed to induce anemia in both wild and laboratory-born voles. 4. Short-term exposure to more extreme temperatures (-20 to -30 degrees C) induced anemia. Voles lost blood through erosions of the epithelium of the glandular stomach, and developed other pathological lesions characteristic of hypothermia. 5. Although there was a strong association between cold weather and anemia, we could find no clear relationship between winter survival and winter weather. However, in 1984, extraordinarily cold temperatures were associated with anemia and a subsequent population decline. These events suggest a threshold mean daily temperature of about -15 degrees C, below which vole survival is grossly affected. 6. Deteriorating protein levels and energy reserves of small mammals in winter may make them particularly susceptible to cold stress. Hence, sporadic bouts of sustained cold may be responsible for some of the enigmatic winter declines seen in northern small mammals.     

Reliability of transferrin and leucine aminopeptidase phenotyping in wild meadow voles (Microtus pennsylvanicus)

Replicate plasma samples from wild Microtus pennsylvanicus were typed by gradient polyacrylamide gel electrophoresis for the transferrin and leucine aminopeptidase polymorphisms. Phenotypes remained the same through major seasonal changes in reproductive activity and environmental conditions. Possible explanations for the anomalous phenotypic variation seen in Microtus ochrogaster [McGovern M., and Tracy, C. R. (1981). Oecologia 51:276] are discussed.     

The effects of mate removal on pregnancy success in prairie voles (Microtus ochrogaster) and meadow voles (Microtus pennsylvanicus).

The effects of removing the stud male have not been controlled in many studies relating pregnancy block to the presence of an unfamiliar male. We examined the effects of removing the male on pregnancy success in prairie voles and meadow voles, two species that differ in degree of paternal investment. Whereas prairie vole males provide extensive care to offspring and accelerate pup development, meadow vole males display little or no care and delay development of pups. We predicted that removal of the stud male would decrease pregnancy success in prairie voles and either have no effect or increase success in meadow voles. In experiment 1, females were in male-induced estrus, and their mates were either left with them or were removed 4 h, 1 day, 2 days, or 8 days after mating. In experiment 2, females were in postpartum estrus, and their mates were either left with them or were removed 1 day, 2 days, or 8 days after birth of their first litter. Removal of the male soon after mating in postpartum estrus decreased pregnancy success in prairie voles and increased success in meadow voles. Thus, although removal of the stud male influenced litter production, the direction of the effect varied with species.     

Seasonal control of odour preferences of meadow voles (Microtus pennsylvanicus) by photoperiod and ovarian hormones

During the spring-summer breeding season, female meadow voles prefer odours of males over those of females, but in the autumn-winter season of reproductive quiescence this preference is reversed. Females housed in long (14 h light/day) and short (10 h light/day) photoperiods, respectively, had odour preferences comparable to those of spring and autumn voles, respectively. The preference of long-photoperiod voles for male over female odours was reversed by ovariectomy and restored by treatment with oestradiol. By contrast, neither ovariectomy nor oestradiol affected odour preferences of short-photoperiod voles. Long days appear to influence olfactory preferences by altering ovarian hormone secretion. The failure of oestradiol to affect odour preferences in short photoperiods suggests that the neural substrates mediating this behavioural response are refractory to oestrogens during the nonbreeding season.     

Serologic study on the prevalence of murine viruses in a population of wild meadow voles (Microtus pennsylvanicus)

The demography and serology of a population of wild meadow voles (Microtus pennsylvanicus) were monitored from 1982 to 1984 near Pinawa, Manitoba, Canada. Serologic tests were performed on 486 samples to detect the presence of viral antibodies to 11 common murine viruses. Meadow voles showed evidence of infection with Theiler's encephalomyelitis, reovirus-type-3, ectromelia, lymphocytic choriomeningitis, adenovirus, and mouse hepatitis viruses. At times of good survival and breeding performance the population was nearly free of evidence of viral infection. During a period of severe mortality in the winter of 1982-1983, evidence of infection by Theiler's encephalomyelitis virus and reovirus-type-3 was obtained. A high prevalence of antibodies and high titers to these two viruses were characteristic of voles that were captured late in the decline in density in the spring of 1983. This association of mortality with a viral outbreak is consistent with the hypothesis that vole population declines are sometimes related to opportunistic pathogens present in the voles' biotic and social environment.     

Influence of daylength experienced by dams on post-natal development of young meadow voles (Microtus pennsylvanicus)

Meadow voles born to dams kept in short days (SD) beginning 5 or 11 weeks before parturition (SD-5, SD-11) had less developed testes, gained weight more slowly and built larger nests than did young gestated and maintained in long days (LD). Pelage development was greater in SD than in LD young at 21 and 45 days of age. At weaning, SD-5 young had less dense fur and shorter guard hairs than did SD-11 young, indicating that the photoperiodic history of dams before insemination affects the post-natal pelage development of their progeny. Short daylengths experienced by dams before mating may facilitate winter preparedness of their offspring.     

Influence of gonadal hormones on odours emitted by male meadow voles (Microtus pennsylvanicus).

Free-living male meadow voles (Microtus pennsylvanicus) emit odours that are attractive to females at the beginning, but not at the end, of the breeding season. The effect of gonadal hormones on female-attractant cues was examined in males born and reared in long (14 h light day-1) and short (10 h light day-1) photoperiods that simulate daylengths in the breeding and nonbreeding seasons, respectively. Gonadectomy affected the attractant properties of odours emitted by long photoperiod, but not short photoperiod, males. Long photoperiod females preferred odours of intact rather than those of gonadectomized long photoperiod males, and odours of gonadectomized long photoperiod males rather than those of intact short photoperiod males. Females did not show a preference between the odours of intact and castrated short photoperiod males. Gonadal hormone replacement in males affected female responses to the odours emitted by long photoperiod, but not short photoperiod, gonadectomized males. Long photoperiod females did not display a preference between odours of intact long photoperiod males and gonadectomized long photoperiod males treated with testosterone or oestradiol. We conclude that in spring and summer gonadal hormones increase attractiveness of male odours; this effect may require aromatization of testosterone to oestradiol. Substrates that control attractiveness of odour cues in male voles appear to be unresponsive to androgens during the nonbreeding season.     

Interaction of daylength and lactation in the control of pelage development and nest-building in female meadow voles (Microtus pennsylvanicus)

Pregnancy and lactation inhibited moult into winter pelage in voles maintained in short daylengths; development of a winter pelage was, however, greatly accelerated once the short-day dams weaned their litters. The presumed elevation of prolactin titres during lactation appears to mask full development and expression of pelage changes induced by short daylengths. Nest-building behaviour, by contrast, was increased in response to short photoperiods and was further augmented during lactation and may thereby facilitate thermoregulation in short-day dams that do not develop a winter pelage.     

Seasonal and interindividual variation in field water metabolism of female meadow voles Microtus pennsylvanicus.

We analyzed variations in water flux rates on a large sample of meadow voles (Microtus pennsylvanicus) to quantify the effect of season on water metabolism of individuals and to examine patterns of intra- and interindividual variability. Voles were nonreproductive females maintained in outdoor enclosures where they fed on natural vegetation. They were injected one to three times with doubly labeled water, which resulted in one to six measures of daily water flux rate per individual. Summer water flux rates of voles were 258% of the predicted values for herbivorous eutherian mammals of similar size. To date, very few studies have focused on mammals with such high water flux rates. Body water volume of individuals was higher in summer than in winter (75.6% vs. 72.5%), and water flux rate of animals was 12.5% higher in the winter season (0.99 vs. 0.88 mL H2O g-1 d-1). Between-season differences in water fluxes were proportional to differences in energy expenditures, hence the water economy index remained constant across seasons (0.30 mL H2O kJ-1). Intraindividual variability of water flux rate was high compared to interindividual variability (repeatability, r<0.30), which will make it difficult to study natural selection of water metabolism in a microevolutionary framework, at least in meadow voles.     

Gonadal hormones modulate sex differences in judgments of relative numerousness in meadow voles, Microtus pennsylvanicus

Animals in a variety of taxa discriminate between a greater quantity and a lesser quantity of the same object, an ability that is referred to as relative numerousness judgment. For example, meadow voles can distinguish between areas containing more over-marks by one opposite-sex scent donor and fewer over-marks by another opposite-sex scent donor. Females appear to be able to make better discriminations between more or less over-marks than do males. In that gonadal hormones have been implicated in modulating cognitive function associated with spatial tasks, we tested the hypothesis that high titers of testosterone and estradiol are necessary for male and female voles, respectively, to distinguish between the top- and bottom-scent donors in an area containing mixed over-marks. We gonadectomized voles, giving them either gonadal hormone replacement (testosterone for males and estradiol for females) or no hormone replacement, and tested their spontaneous judgments of distinguishing between the top- and bottom-scent donors in an area containing mixed over-marks; a task involving judgments of relative numerousness. Female voles given replacement estradiol performed better than did female voles not given replacement estradiol in determining the top-scent and bottom-scent males in areas containing mixed over-marks. In contrast, males not treated with replacement testosterone performed better than did males treated with testosterone in determining the top-scent and bottom-scent males in areas containing mixed over-marks. Thus, high titers of estradiol and low titers of testosterone are associated with better performance on tasks involving relative numerousness in female and male voles, respectively. The results of this task on relative numerousness judgments are discussed in relation to the effects of gonadal steroid hormone on spatial ability, a closely related cognitive domain, and the social biology of meadow voles.     

Meiosis in gray voles of the subgenus microtus (rodentia, arvicolinae) and in their hybrids

The results of light and electron microscopic (EM) studies of meiosis in Microtus arvalis males of the karyoform "arvalis" (2n = 46, NFa = 80), in hybrids between the chromosomal forms arvalis and obscurus (2n = 46, NFa = 68), in M. rossiaemeridionalis voles (2n = 54, NFa = 54), and in a hybrid between the species M. rossiaemeridionalis and M. kermanensis (2n = 54, NFa = 54) are presented. SC (synaptonemal complex) karyotypes of the parental forms and the hybrids were constructed on the basis of measurements of the length ofautosomal SCs revealed by the EM analysis in spermatocytes at the stage of middle pachytene. The SC karyotypes of M. arvalis and the hybrids female obscurus x male arvalis consist of 22 synaptonemal complexes of autosomal bivalents and the axial elements of the synaptonemal complexes of the sex chromosomes X and Y. The SC karyotypes of M. rossiaemeridionalis and the hybrid M. rossiaemeridionalis x M. kermanensis consist of 26 synaptonemal complexes of autosomal bivalents and a sex bivalent; they differ only in the length of the Y chromosome axis (Y chromosome in the hybrid was inherited from M. kermanensis). Asynaptic configurations of the autosomal SCs were not observed in the hybrids. The SC axial elements of the X and Y chromosomes in the parental forms and in the hybrids were located close to each other throughout pachytene, but they did not form a synaptic region. The normal synapsis in sterile hybrids (M. rossiaemeridionalis x M. kermanensis) and the behavior of the sex chromosomes in meiosis in fertile and sterile hybrids are discussed in the context of specific features of meiosis and reproductive isolation.     

A blond coat color variation in meadow vole (Microtus pennsylvanicus)

Color mutations occur frequently among rodents. Here we describe a blond coat color mutation in the meadow vole (Microtus pennsylvanicus) that arose in a captive breeding colony established from wild-caught animals from southern Illinois. The blond coat coloration results from changes in the color and distribution of pigments in the hair. The mutation is monogenic autosomal recessive.     

Experimental inoculation of meadow voles (Microtus pennsylvanicus), house mice (Mus musculus), and Norway rats (Rattus norvegicus) with Mycobacterium bovis

Mycobacterium bovis has a wide host range that includes several wildlife species, and this can hamper attempts to eradicate bovine tuberculosis from livestock. The purpose of this study was to determine if common rodent species, namely meadow voles (Microtus pennsylvanicus), house mice (Mus musculus), and Norway rats (Rattus norvegicus), that inhabit the bovine tuberculosis endemic area of Michigan, can be experimentally infected with M. bovis. The objectives of the study were: 1) to determine if these rodent species can be infected, and if so, to document attendant pathologic processes/pathogenesis; 2) to detect any fecal shedding of M. bovis; and 3) to evaluate the relative susceptibility of the three species to M. bovis infection. For each species (n=36) there were two treatment (n=12/group) and one or two control groups depending on species (n=6-12/group); the maximum study duration was 60 days. The meadow vole treatments consisted of high dose inocula that were given by oral or intranasal routes, whereas the house mice and Norway rats were given only oral inocula at either a high or low dose. Of the three species, meadow voles were most susceptible to M. bovis infection. Upon intranasal inoculation, all 12 voles were infected as determined by gross and microscopic lesions and culture of M. bovis from tissue and feces. Seven of the 12 meadow voles inoculated orally were infected. House mice also were susceptible; M. bovis was isolated from 14 of 24 animals. Only one Norway rat in the high dose treatment group was positive by culture and this was the only animal from which minimal attendant lesions were observed. Results of this study indicate that meadow voles and house mice can be infected with M. bovis and might serve as spillover hosts. Concerted efforts should, therefore, be made to reduce or eliminate these rodents on premises where M. bovis-infected livestock are present.     

Time course of androgenic modulation of odor preferences and odor cues in male meadow voles, Microtus pennsylvanicus.

During the breeding season, male meadow voles prefer female over male odors and females prefer male over female odors. Testosterone control of males' odor preferences and production of odors attractive to females differ. A male meadow vole's preference for female versus male odor was still evident 1 week after castration, but not 1 week later. This preference was reinstated in testosterone-treated male voles 2 weeks after the onset of hormone replacement. The attractiveness of male odors to females did not disappear until 3 weeks after castration. The attractiveness of male odors was reinstated 1 week after castrated males were treated with testosterone. The time course for the androgenic modulation of production of odors attractive to females may facilitate breeding. For example, at the end of the breeding season males may emit an odor that is still attractive to females. Similarly, at the beginning of the breeding season males may emit an odor that is attractive to females.     

The influence of multiple photoperiods and pinealectomy on gonads, pelage and body weight in male meadow voles, Microtus pennsylvanicus

1. Chronic exposure of male Microtus pennsylvanicus to photoperiods with 8, 10 and 12 hr of light per day results in gonadal regression, molt to winter pelage and significant reductions in body weight and food consumption relative to voles kept on photoperiods with 13 and 14 hr of light per day. 2. A precise critical daylength is observed in this species as exposure to photoperiods with 12 or fewer hours of light per day results in complete gonadal involution, seasonal molt and loss of body weight. 3. Pinealectomy abolishes all short-photoperiod induced responses in this species.     

The effects of habitat manipulation on population distribution and foraging behavior in meadow voles

Individuals, free to choose between different habitat patches, should settle among them such that fitness is equalized. Alternatives to this ideal free distribution result into fitness differences among the patches. The concordance between fitnesses and foraging costs among inhabitants of different quality patches, demonstrated in recent studies, suggests that the mode of habitat selection and the resulting fitness patterns may have important implications to the resource use of a forager and to the survival of its prey. We studied how coarse scale selection between habitat patches of different quality and quitting harvest rate in these patches are related to each other and to fine scale patch use in meadow voles (Microtus pennsylvanicus). To demonstrate these relationships, we manipulated habitat patches within large field enclosures by mowing vegetative cover and adding supplemental food according to a 2×2 factorial design. We tracked vole population densities, collected giving‐up densities (GUDs, a measure of patch quitting harvest rate), and monitored the removal of seeds from lattice grids with 1.5 m intervals (an index of fine‐scale space use) in the manipulated habitat patches. Changes in habitat quality induced changes in habitat use at different spatial scales. In preferred habitats with intact cover, voles were despotic and GUDs were low, but increased with the addition of food. In contrast, voles in less‐preferred mowed habitats settled into an ideal free distribution, GUDs were high and uninfluenced by the addition of food. Seed removal was enhanced by the presence of cover but inhibited by supplemental food. Across all treatments, vole densities and GUDs were strongly correlated making it impossible to separate their effects on seed removal rates. However, this relationship broke down in unmowed habitats, where GUDs rather than vole density primarily influenced seed removal by voles. GUDs and seed removal correlated with predation on tree seedlings formerly planted into the enclosures, demonstrating the mechanisms between coarse‐scale habitat manipulations and community level consequences on a forager's prey.