Effect of nutrition, temperature and photoperiod on the rate of sexual maturation of the field vole (Microtus agrestis)

Weanling male and female field voles were placed in long or short photoperiods, kept at 18 degrees C or 4 degrees C, and fed (ad libitum) diets containing 24%, 16%, 8% and 4% protein, for 6 weeks. Animals in the long photoperiod were more sexually mature than were animals in the short photoperiod. Temperatures had no effect on females, but did affect males: those kept at 4 degrees C had heavier testes and wider seminiferous tubules than those kept at 18 degrees C. There was little difference between the animals on 24%, 16% and 8% protein diets. Animals on 4% protein diets had retarded growth rates and were significantly less sexually mature than those on the other 3 diets, males having smaller testes and seminal vesicles and narrower seminiferous tubules and females having smaller ovaries and uteri.     

Gonadal development and gonadotrophin secretion in the male vole (Microtus agrestis) after an abrupt change in photoperiod

Male voles were reared from birth to age 28 days in 6L:18D. Pairs of animals showing similar sexual development were assigned at random to 16L:8D or 6L:18D. Treatments continued for a further 56 days. Increase in the activity of the hypothalamo-hypophysial system occurred within 4 days of exposure to 16L:8D, as shown by significant elevation of plasma LH and FSH. Pituitary LH did not increase until Day 7, and pituitary FSH did not increase until Day 21. After exposure to 16L:8D for 4 days, pituitary FSH was lower than in corresponding animals in 6L:18D. These discrepancies between pituitary and plasma values of gonadotrophins indicate that increase in hormone release occurs before synthesis is fully stimulated. Enhanced output of testicular hormones probably began between Day 7 and Day 14, as indicated by an increase in seminal vesicle weight, yet plasma and pituitary concentrations of LH and FSH remained elevated. This suggests that long photoperiods may cause direct stimulation of the hypothalamo-hypophysial system which increasing values of testicular hormones are initially unable to inhibit. The response of this system in voles to an abrupt change from a non-stimulating to a stimulating photoperiod has a time course resembling that for the Soay ram but appreciably slower than for the Japanese quail.     

Phytosterols affect endocrinology and metabolism of the field vole (Microtus agrestis)

Phytosterols or plant sterols (PS) enter the ecosystem via pulp mill effluents. They are also consumed by the general population of developed countries in natural remedies and margarines to lower elevated serum cholesterol levels. This study screened the endocrine and enzymatic parameters of the field vole (Microtus agrestis) for the effects of subchronic PS exposure at three doses (0, 5, or 50 mg of PS kg(-1) day(-1)). PS at 5 or 50 mg kg(-1) day(-1) decreased the relative liver weight of the voles. The kidney glycogen phosphorylase activity decreased at 5 or 50 mg kg(-1) day(-1), but the liver glycogen phosphorylase activity increased at 5 mg kg(-1) day(-1). The plasma estradiol and testosterone concentrations of males were higher due to PS supplement at 5 mg kg(-1) day(-1). This can be due to increased sex steroid synthesis from PS precursors. Biotransformation enzyme activities were not affected. PS caused multiple, previously unreported effects that were more pronounced at a low dose. As 5 mg PS kg(-1) day(-1) is the recommended dose for various health products, a thorough risk assessment of the effects and interactions of PS is warranted.     

The development of social behaviour in the vole (Microtus agrestis)

Juvenile voles of the species Microtus agrestis investigated the bodies of other juveniles with the nose, by touching the dorsal surface from snout to tail. Play movements often followed such nosing contacts and augmented further contacts. The most important scent stimulating play was produced by the skin at the back of the head. The secretion of the nose and mouth region may convey sexual identity, the rump region and feet individual identity.
Young born in the spring and early summer often interacted and played, the autumn young seldom. The behaviour of overwintering sexually immature adults was similar to that of the autumn juveniles. It is suggested that this seasonal difference in the behaviour of juveniles has consequences for subsequent mature males, and that this developmental factor may have an important effect on the annual population cycle.     

Apoptosis distribution in the first molar tooth germ of the field vole (Microtus agrestis)

Apoptosis represents an important process in organ and tissue morphogenesis and remodeling during embryonic development. A role for apoptosis in shape formation of developing teeth has been suggested. The field vole is a useful model for comparative studies in odontogenesis, particularly because of its contrasting molar morphogenesis when compared to the mouse. However, little is known concerning apoptosis in tooth development of this species. Morphological (cellular and nuclear alterations) and biochemical (specific DNA breaks--TUNEL staining) characteristics of apoptotic cells were used to evaluate the temporal and spatial occurrence of apoptosis in epithelial and mesenchymal tissues of the developing first molar tooth germs of the field vole. Apoptotic cells were found in non-proliferating areas (identified previously) throughout bud to bell stages, particularly in the epithelium, however, scattered also in the mesenchyme. A high concentration of TUNEL positive cells was evident in primary enamel knots at late bud stage with increasing density of apoptotic cells until ED 16 when the primary enamel knot in the field vole disappears and mesenchyme becomes protruded in the middle axes of the bell forming two shallow areas with zig-zag located secondary enamel knots. Distribution of TUNEL positive cells corresponded with localisation of secondary enamel knots as shown using histological and 3D analysis. Apoptosis was shown to be involved in the first molar development of the field vole, however, exact mechanisms and roles of this process in tooth morphogenesis require further investigation.     

Effects of temporal variation in the risk of predation by least weasel (Mustela nivalis) on feeding behavior of field vole (Microtus agrestis)

Predation risk tends to vary in time. Thus prey animals face a problem of allocating feeding and antipredator effort across different risk situations. A recent model of Lima and Bednekoff (1999) predicts that a prey should allocate more feeding effort to low risk situations and more antipredator effort to high risk situations with increasing relative degree of risk in high risk situations (attack ratio). Furthermore when the proportion of time the prey spends in the high risk situation (p) increases, the prey have to eventually feed also in the high risk situations. However the increase in feeding effort in low risk situations should clearly exceed that in high risk situations as p increases. To test these predictions we measured feeding effort of field voles (Microtus agrestis) exposed to varying presence of least weasel (Mustela nivalis) and its feces in laboratory conditions. We generated quantitative predictions by estimating attack ratios from results of a pilot experiment. The model explained 15% of the observed variation in feeding effort of voles. Further analyses indicated that feeding effort was lower in high risk situations than in low risk situations at high attack ratio, but not at a lower one. Voles exposed to a presence of a weasel for extended periods showed signs of nutritional stress. Still we did not find any increase in feeding effort with increasing p. This was obviously due to the relatively low maximal p we used as we included only conditions likely to occur in nature.     

Suppressed breeding in the field vole (Microtus agrestis): an adaptation to cyclically fluctuating predation risk

The densities of microtine rodents and their main predators,small mustelids, fluctuate synchronously in 3–5-year cyclesin central and northern Fennoscandia. Predation by small mustelidshas been suggested as one of the driving forces in microtinecyclicity, causing deep synchronous declines of several volespecies. We studied experimentally the effects of small mustelidson mating behavior, foraging, and breeding in nonwintered fieldvoles (Microtus agrestis) originating from a cyclic population.By using mustelid odors, we simulated a crash phase environmentwith high predation risk for breeding pairs of voles. In ourexperiments, 87% of the female field voles suppressed breedingwhen exposed to mustelid odors. Both female and male behaviorchanged, and no mating behaviors were observed under the simulatedpredation risk. Weights of both sexes decreased when exposedto mustelid odor, probably due to decreased foraging; weightsof the control females increased due to pregnancy; and no weightchanges occurred in control males. Decreased breeding and foragingpossibilities under high predation pressure may form the basisfor the ultimate explanation for breeding suppression. Thereare at least two different mechanisms for breeding suppression:either mating does not take place or malnutrition in femalesdoes not' allow breeding to occur. Delayed breeding under highrisk of predation, for whatever reason, could increase the probabilityof individuals, especially that of the females, to survive overthe crash to the next, safer breeding season when their youngwould have better possibilities to survive.     

Pregnancy blockage and the memory of the stud male in the vole (Microtus agrestis).

When female voles were allowed contact with the stud male for only 1 h at the time of mating, 55% exhibited pregnancy failure when exposed to a strange male 48 h later. When females were made psuedopregnant by hormone treatment and vaginal stimulation (i.e. no stud male involved), 87% exhibited luteal failure when exposed to a strange male. It is suggested that the characteristics of the stud male are rapidly imprinted upon the female at the time of mating and that this imprinting is important in preventing the female showing a blocking response to this male upon any subsequent exposure.     

Evaluation of methods to estimate field vole Microtus agrestis abundance in upland habitats

Field vole abundance was assessed on a range of upland habitats in spring and autumn, using live—trapping, snap—trapping and vole sign index methods. Three estimates of vole numbers were derived from live—trapping, five from snap—trapping and one from vole signs. Estimates were compared using a correlation matrix in order to establish the most accurate and consistent method. Snap—trapping for five nights provided enough information to calculate vole density, though the method was time consuming. Vole abundance indices derived from two nights' snaptrapping were significantly correlated with indices from five nights' trapping. Live—trapping and vole sign index were discounted as methods for this study area, as both proved unreliable.     

Short-term and circadian rhythms in the behaviour of the vole,Microtus agrestis (L.)

Summary The activity behaviour of the vole, Microtus agrestis, has been recorded in order to investigate the relationship between short-term rhythm and circadian rhythm. A simple device was developed, allowing separate monitoring of the time spent in or outside the nest, wheel-running, eating and drinking. Under natural light conditions during summer, a distinct differentiation between a short term rhythm of eating and drinking during the day-time and a circadian rhythm of wheel-running during the night was observed. The short-term rhythm depends closely on metabolic demands (hunger, thirst, excretion). Control of these demands by an endogenous oscillation could not be substantiated. The circadian rhythm of wheel-running activity is, however, controlled by an endogenous oscillation, synchronized by light conditions. It is subjected to seasonal variations. a) The threshold of light intensity below which wheel-running occurs is lowest during summer (<0.5 lx) and is higher during spring and autum (> 5 lx). b) Wheel-running is controlled by a circadian oscillation during summer only whereas it is an integrated part of the short-term rhythm during spring and autumn (experiments during the winter have not yet been performed). Experiments gave evidence that the properties of the cage can deeply influence the amount and pattern of wheel-running activity. It is concluded that wheel-running reflects a certain level of excitation, which may be caused by different behavioural intentions. The seasonal changes of the control of wheel-running activity are discussed with respect to this assumption. The relevancy of locomotor activity patterns as usually recorded in the laboratory to reveal the physiological and ecological significance of endogenously controlled behavioural patterns is discussed.Supported by the Deutsche Forschungsgemeinschaft     

Meiotic association and segregation of the achiasmatic giant sex chromosomes in the male field vole (Microtus agrestis)

Controversy exists regarding the meiotic behaviour of the giant sex chromosomes during spermatogenesis in the field vole, Microtus agrestis. Both univalents and bivalents have been observed between diakinesis and metaphase I. These differences seem to be dependent on the technique used. The present study employs electron microscopy of serially sectioned testes tubules and light microscopy of microspread preparations to re-examine the behaviour of sex chromosomes during meiosis. In microspreads, about one-third of the early pachytene nuclei examined showed end joining of the X and Y axes. The longitudinal heterogeneity of the chromosomes in the form of axial thickenings allowed the detection of two different end-joining patterns. In the remaining early pachytene cells as well as in all mid to late pachytene cells seen, the X and Y axes had, though near to each other, no contact in the form of a synaptonemal complex. If a synaptonemal complex is a prerequisite for genetic exchange, the sex chromosomes in M. agrestis males must be achiasmatic. The analysis of serial sections through an early pachytene and a late prophase I nucleus with the electron microscope revealed that the sex chromosomes occupied a common area. By metaphase I, the centromeres of the X and Y were oriented towards opposite spindle poles while the chromosomes remained attached to one another by their distal segments at the level of the metaphase I plate. As a consequence of the large size of the sex chromosomes their centromeres lay close to the spindle poles. In anaphase I the sex chromosomes maintained their metaphase position until the autosomes approached the spindle poles. During autosomal migration a medial constriction developed where the sex chromosomes were mutually associated, the X and Y became separated, and joined the autosomes. In metaphase II the chromatids of the sex chromosomes lay side by side and exhibited a delayed separation in the subsequent anaphase. It is suggested that heterochromatin, which represents a major part of both sex chromosomes, plays a role in the association of the two achiasmatic sex chromosomes in metaphase I and in the delayed separation of the chromatids of the sex chromosomes in anaphase II.Dedicated to Prof. C.-G. Arnold (Erlangen) on the occasion of his 60th birthday