Age Affects Over-Marking of Opposite-Sex Scent Marks in Meadow Voles, Microtus pennsylvanicus

Models of age-related effects on behavior predict that among short-lived species younger adults are more attractive and attracted to opposite-sex conspecifics than are older adults, whereas the converse is predicted for long-lived species. Although most studies of age-related effects on behavior support these predictions, they are not supported by many studies of scent marking, a behavior used in mate attraction. Over-marking, a form of scent marking, is a tactic used by many terrestrial mammals to convey information about themselves to opposite-sex conspecifics. The present study tested the hypothesis that the age of meadow voles, Microtus pennsylvanicus ; a microtine rodent, affects their over- and scent-marking behaviors when they encounter the marks of opposite-sex conspecifics. Sex differences existed in the over-marking behavior of adult voles among the three different age groups that were tested. Male voles that were 5–7 and 10–12 mo olds over-marked a higher proportion of the marks of females than did 2–3 mo old male voles. Female voles that were 2–3, 5–7, and 10–12 mo old over-marked a similar number of marks deposited by male voles. Overall, the data were not consistent with models predicting the behavior of short-lived animals such as rodents when they encounter the opposite sex. The differences in over-marking displayed by older and younger adult male voles may be associated with life history tradeoffs, the likelihood that they will encounter sexually receptive females, and being selected as mates.     

Response of Prairie Voles, Microtus ochrogaster (Rodentia, Arvicolidae), to Scent Over-marks of Two Same-sex Conspecifics: A Test of the Scent-masking Hypothesis

Previous work shows that after investigating a same-sex over-mark, two nonmonogamous species, meadow voles and golden hamsters, preferred the odor of the top-scent donor to that of the bottom-scent donor, and behaved as the odor of the bottom-scent donor was not familiar. This finding supported the scent-masking hypothesis; one of three hypotheses suggested previously to account for how an animal responds to the overlapping scent marks of two same-sex conspecifics. The present experiments tested whether one of these hypotheses, either scent-masking, scent-bulletin-board, or scent-blending, predicts how a monogamous species, the prairie vole, responds to such over-marks. Our data show that none of the three hypotheses adequately describes the way in which prairie voles respond to conspecific over-marks. Although prairie voles preferred the top scent to the bottom scent, they behaved as if the latter scent was familiar and less important than a novel scent (a scent not part of the over-mark). Overall, the data suggest that the manner in which males and females respond to same-sex over-marks reflects the different tactics they may use to attract and compete with conspecifics in monogamous and nonmonogamous species.     

Self-Discrimination in Meadow Voles, Microtus pennsylvanicus

Particular features of the signaling characteristics of the scent marks of temperate zone, seasonally breeding mammals may reflect differences in their reproductive state and, hence, be variable. Consequently, an individual's perception of self may depend more on the condition independent than on the condition-dependent signaling characteristics of the scent marks. Yet, we do not know whether an individual responds to changes in the signaling characteristics of its own scent marks, such as those associated with changes in an individual's reproductive state. Such changes may affect how and where an animal scent marks. Here we report on a series of experiments designed to test the hypothesis that individual meadow voles, Microtus pennsylvanicus , distinguish between scent marks they deposited when they were in different reproductive states. Results showed that voles discriminated their own scent marks from those of unfamiliar, same-sex conspecifics, and the scent marks of siblings. Voles did not behave as if they could distinguish between their own scent marks if the marks were deposited when the voles were in the same reproductive state, although the two scent marks used as stimuli differed in age by 30 d. However, they did so distinguish if they were exposed to scent marks taken when they were in different reproductive states. Overall, these findings suggest that voles behave as if their novel and familiar scent marks shared the similar signaling features. If, however, the reproductive condition of the voles differed when it provided the two scent marks, they behaved as if their own scent marks had different signal characteristics, which may have induced voles to treat the two scent marks as not being the same or having been deposited by two different donors. We speculate that the scent marks of individuals may have unique signaling characteristics that may be associated with that individual's 'current template for self'.     

Meadow Voles (Microtus pennsylvanicus) and Prairie Voles (M. ochrogaster) Differ in Their Responses to Over-Marks from Opposite- and Same-Sex Conspecifics

Over‐marking occurs when one individual deposits its scent mark on the scent mark of a conspecific. Previous studies have shown that meadow voles (Microtus pennsylvanicus) and prairie voles (M. ochrogaster) that were exposed to an over‐mark of two same‐sex conspecifics, later responded similarly to the top‐scent mark but differed in their response to the bottom‐scent mark. In the present study, we examined the responses of meadow voles and prairie voles to same‐sex and mixed‐sex over‐marks to ascertain whether their responses reflect the different tactics which males and females in promiscuous (meadow voles) and monogamous (prairie voles) species use to attract opposite‐sex conspecifics and to compete with same‐sex conspecifics. Males and females of both species spent more time investigating the mark of the top‐scent donor than that of the bottom‐scent donor of an over‐mark. Meadow voles exposed to a mixed‐sex over‐mark spent more time investigating the mark of the opposite‐sex conspecific independently of whether it was from the top‐ or bottom‐scent donor. In contrast, prairie voles spent more time investigating the mark of the opposite‐sex donor if it was from the top‐scent donor. These results suggest that: (i) over‐marking serves a competitive function; (ii) the scent marks of individuals attract multiple mates in promiscuous species such as the meadow vole; and (iii) the scent marks of individuals establish and maintain pair bonds between familiar opposite‐sex conspecifics in monogamous species such as the prairie vole.     

Persistence of the Attractiveness of Two Sex-specific Scents in Meadow Voles,Microtus pennsylvanicus

The function of an odour may be reflected in its fade-out time in the environment. In this study, we investigated fade-out times of two specific odours, the anogenital area scent and that of the posterolateral region. These two odours support opposite-sex preferences in male and female meadow voles, Microtus pennsylvanicus, but convey nonidentical information to conspecifics during the breeding season. The first experiment tested whether meadow voles respond preferentially to scents that were aged for 15 min (fresh) to 30 d. Males preferred female anogenital area scent to male anogenital area scent if both scents were ≤ 10 d old. By comparison, females preferred male anogenital area scent to female anogenital area scent if the scents were ≤ 25 d old. However, male and female voles preferred the posterolateral scent of males to that of females if the scents were ≤ 1 d old. Thus, fade-out times for these two scents differ for males and females, suggesting different functions. In the second experiment, male and female voles preferred fresh anogenital area scent and fresh posterolateral region scent compared with those same scents that were older. This result suggests that older scents may have lost information over time about the sex of the donor. Overall, data from both experiments indicate that voles may use specific scents for communication in different social contexts.     

Male and Female Meadow Voles,Microtus pennsylvanicus,Differ in Their Responses to Heterospecific/Conspecific Over‐Marks

Voles use runways, paths, and trails that may also be used by rabbits and mink. These shared areas could contain the scent marks of conspecifics and heterospecifics. Thus, it is likely that the scent marks of heterospecifics may overlap or be overlapped by those of voles, forming over‐marks. Much is known about how voles respond to over‐marks of two different conspecifics. However, we do not know how they would respond to an opposite‐sex conspecific whose scent marks are in an over‐mark with the scent marks of predator or the scent marks of a non‐predator heterospecifics. We tested the hypothesis that meadow voles, Microtus pennsylvanicus, differ in their response to the scent mark of the opposite‐sex conspecific if the scent mark was overlapped by that of a mink, a vole predator, or rabbit, a vole non‐predator. We found that female but not male voles showed a preference for the scent marks of the opposite‐sex conspecifics that were part of the mink‐vole over‐mark when compared to those of opposite‐sex conspecifics that were not part of the over‐mark. This preference by female voles was independent of whether the male vole was the top‐scent donor or bottom‐scent donor of the over‐mark. Male and female voles showed no preference between the scent marks of the opposite‐sex conspecifics whose marks were part of or not part of the rabbit‐vole over‐mark. Sex differences in the manner that meadow voles respond to rabbit‐vole and mink‐vole over‐marks are discussed.     

Seasonal Differences in Social Behavior among Adult and Juvenile Meadow Voles,Microtus pennsylvanicus

Seasonal differences in behavioral interactions between wild-caught adult and juvenile meadow voles, Microtus pennsylvanicus, were measured in paired encounters in a neutral arena. Seasonal differences existed in the adult-juvenile paired encounters. Early in the breeding season (May—Aug.) paired encounters between adult males and juvenile males were more agonistic than other adultjuvenile pairings. Late in the breeding season (Sep.—Dec.) adult female-juvenile male encounters were more agonistic than other adult-juvenile pairings. Odor preferences of juveniles were determined in a Y-maze olfactometer. Juveniles did not demonstrate seasonal differences in odor preference. Males preferred the odor of adult females to the odor of adult males or their own odor. Females preferred the scent of an adult male to the odor of an adult female, but did not show a preference between their own odor and the odor of an adult male or an adult female. The results are consistent with hypotheses that state in a promiscuous species, females should be philopatric, and males should disperse as a consequence of adult competition. The data suggest that agonistic behavior by adults may regulate juvenile dispersal and recruitment, and define the composition of overwintering groups.     

Clusters of Repetitive DNA Sequences in Chromosomes of Voles of the Subgenus Microtus (Microtus,Arvicolidae)

Russian Journal of Genetics - The study is focused on the analysis of the mechanisms underlying the formation and distribution of repeat clusters in mammalian chromosomes, as exemplified by a group...     

Food Deprivation Suppresses a Preference for the Top-Scent Mark of an Over-Mark in Meadow Voles (Microtus pennsylvanicus)

Food availability affects whether mammals communicate their interest in interacting with opposite‐sex conspecifics. This study examined the responses of voles to over‐marks, and factors that influence the formation and maintenance of a preference for the top‐scent in an over‐mark. Specifically, we investigated how food deprivation affected the amount of time male and female voles exposed to an over‐mark, later responded to the marks of the top‐ and bottom‐scent donors when subsequently presented with the two scents side by side. Males and females that were not food deprived and males that were food deprived 6 h before exposure to an over‐mark later maintained a preference for the donor of the top‐scent mark compared with the donor of the bottom‐scent mark of the over‐mark. Females that were food deprived for 6 h before or after exposure of the over‐mark and males food deprived 6 h after the exposure to the over‐mark showed no preference for the top‐scent mark donor. Re‐feeding females that were food deprived for 6 h before exposure to an over‐mark was sufficient to restore their preference for the mark of the top‐scent male over that of the bottom‐scent male. The different responses of food‐deprived male and female voles to over‐marks of opposite‐sex conspecifics may be associated with differences in their tactics for interacting with potential mates and the higher energetic costs of reproduction in female voles than in male voles.     

Female Meadow Voles, Microtus pennsylvanicus, Experience a Reduction in Copulatory Behavior During Postpartum Estrus

In many small mammalian species, females undergo postpartum estrus (PPE), mate, and become pregnant soon after delivering a litter. Mating during PPE implies a trade‐off between attending to the new litter and leaving that litter to mate. We tested the hypothesis that copulatory behavior is faster when a female mates during PPE than when it mates outside of PPE, during male‐induced estrus (MIE), a time when it is not lactating. We compared several variables of copulatory behavior in female meadow vole mating during both PPE and MIE. Females in PPE received significantly fewer intromissions, with shorter intervals between intromissions. Each ejaculatory series was also shorter for females in PPE. As a consequence, the total amount of time devoted to copulation was much shorter for females in PPE than for those in MIE. These data support the hypothesis that female meadow voles are able to reduce their copulatory behavior during PPE.     

Food Deprivation and Restriction During late Gestation Affect the Sexual Behavior of Postpartum Female Meadow Voles,Microtus pennsylvanicus

In many species of small mammals, including meadow voles, Microtus pennsylvanicus, females come into postpartum estrus (PPE) within 12–24 h of giving birth, allowing them to mate and become pregnant while raising the current litter. PPE females show increases in attractivity, proceptivity, and receptivity, the three components of sexual behavior, relative to females not in PPE. Several studies have shown that food deprivation and restriction reduce attractivity, proceptivity, and receptivity of females not in PPE. We tested the hypothesis that food deprivation and restriction during late gestation cause deficits and decrease the attractivity, proceptivity, and receptivity of females when they enter PPE. Our data support the hypothesis. On day 1 of lactation, females that were food deprived and food restricted produced scent marks that were significantly less attractive as those produced by control PPE females. Food deprivation but not food restriction caused females to no longer display significant preferences for the scent marks of males over those of females (proceptivity). Food deprivation and food restriction were sufficient to induce females to become significantly less sexually receptive than control females. Eleven of 12 control PPE females mated, 4 of 12 food‐restricted females mated, and 3 of 12 food‐deprived females mated. Dams facing food deprivation or restriction during late gestation may have to balance the benefits of mating during PPE with the increased costs associated with getting pregnant while they are lactating.     

Maternal Food Restriction During Lactation Affects Body Weight and Sexual Behavior of Male Offspring in Meadow Voles (Microtus pennsylvanicus)

Little is known about the occurrence of individual variation in sexual behavior and how maternal nutrition can affect this variation. We tested the hypothesis that male offspring of female meadow voles, Microtus pennsylvanicus, that were 30% food restricted (FR) during days 1–7 of lactation (FR 1–7), days 8–14 of lactation (FR 8–14), or late days 15–21 of lactation (FR 15–21) lactation show persistent, negative effects on their sexual behavior as adults relative to male offspring of females that were not food restricted. We measured three components of sexual behavior, attractivity, proceptivity, and receptivity, beginning when the males were 98 d of age. Food restriction during middle lactation (FR 8–14) but not during early (FR 1–7) and late lactation (FR 15–21) was sufficient to induce adult male voles to produce anogenital marks that were not as attractive as those produced by control males. Food restriction during lactation did not affect the proceptive behavior of male voles but did affect their receptivity. Only four of 12 FR 8–14 male voles mated compared to nine of 12 FR 1–7 males, eight of 12 FR 15–21 males, and eight of 11 control males. However, no differences existed in their copulatory behavior among the males that did mate. The body weight of FR 1–7 and FR 8–14 males was lower than that of FR 15–21 and control males when they were between 22 d of age (weaning) and 48 d of age (puberty) but was similar when the males were 98 d of age. Food intake was similar for the FR and control males between day 22 and day 98. It remains unclear, however, whether this type of maternal effect represents strategic programing of offspring behavior in response to the environment experienced by mothers or is a product of developmental processes of food restriction prior to weaning (Evolution 58 , 2004, 2574).     

The Reproductive State of Female Voles Affects their Scent Marking Behavior and the Responses of Male Conspecifics to Such Marks

During the breeding season, the reproductive condition of female mammals changes. Females may or may not be sexually receptive. We conducted a series of experiments to determine whether reproductive condition of female meadow voles affects their scent marking behavior as well as the scent marking behavior of male conspecifics. In expt 1, females in postpartum estrus (PPE females) deposited more scent marks than females that were neither pregnant nor lactating (REF females) or ovariectomized females (OVX females). In expt 2, male voles scent marked more and deposited more over‐marks in areas marked by PPE females than by REF and OVX females. In expt 3, PPE females deposited more scent marks and over‐marks in areas marked by males than did females in the other reproductive states. The results of these experiments showed that male and female voles may vary in the number, type and location of scent marks they deposit in areas scented by particular conspecifics.     

The Outcome of a Previous Social Interaction with a Same‐sex Conspecific Affects the Behavior of Meadow Voles,Microtus pennsylvanicus

In species where conspecifics compete for resources such as territories, remembering where a neighbor was previously encountered and the outcome of that interaction may give individuals advantages over nearby conspecifics. We used a two‐phase experiment to test the hypothesis that female meadow voles, Microtus pennsylvanicus, who during the breeding season are territorial and agonistic toward one another, can use details of an encounter with another female in one location to later navigate nearby areas. During the encounter phase, pairs of females interacted for two minutes in one isolated section of a Y‐maze; control females were placed in alone. Females were scored as either winners or losers. Winners displayed twice as many agonistic acts against their opponent. The test phase took place after a retention interval of one hour, 1 d, or 1 wk. Single females were returned to a clean and empty Y‐maze and allowed to explore the entire apparatus for 15 min. We recorded the amount of time spent in each section of the maze. After retention intervals of one hour and 7 d winners, losers, and controls spent similar amounts of time in each section. However, after 24 h, winners spent more time in the encounter section; losers and control females spent similar amounts of time in each section. The results suggest that meadow voles' memory of the details of a single encounter is influenced by the emotional valence attached to that event. The duration of memory may be associated with the establishment of territories by female meadow voles.     

Scent Counter-marking by Male Meadow Voles: Females Prefer the Top-scent Male

Scent counter-marking, in which one individual deposits scent in close proximity to the scent of another individual, is a widespread but poorly understood aspect of olfactory communication. Recent work with golden hamsters suggests that animals may have specially evolved mechanisms for determining which individual has marked most recently, and this work emphasizes the need for studies with other species. In Experiment 1 it was shown for the first time that male meadow voles, Microtus pennsylvanicus, scent mark with urine and anogenital scents and probably also counter-mark with these scents. Female meadow voles, after investigation of an area marked by two males, preferred the whole-body odours of the male that had marked the arena most recently (Experiment 2). After females investigated a male's home cage that had just been marked by another male, they again preferred the whole-body odours of the male that had marked in the cage most recently (Experiment 3). These results demonstrate that female voles, like male hamsters, can distinguish the top or most recent individual's scent from the bottom or older scent in places marked by two males, and further indicate that female voles may prefer the individual that deposited the top scent. Taken together, the results suggest that counter-marking by male voles may be a type of competitive advertising and that females may base mate-choice decisions on information from the pattern of such counter-marks.     

Olfactory Experience Affects the Response of Meadow Voles to the Opposite‐Sex Scent Donor of Mixed‐Sex Over‐Marks

Scent marking and over‐marking are important forms of communication between the sexes for many terrestrial mammals. Over the course of three experiments, we determined whether the amount of time individuals investigate the scent marks of opposite‐sex conspecifics is affected by 4 d of olfactory experience with those conspecifics. In Experiment 1, female meadow voles, Microtus pennsylvanicus, spent more time investigating the scent mark of the novel male conspecific than that of the familiar male donor, whereas male voles spent similar amounts of time investigating the scent mark of the familiar female and a novel female conspecific. In Experiment 2, voles were exposed to a mixed‐sex over‐mark in which subjects did not have 4 d of olfactory experience with either the top‐scent donor or the bottom‐scent donor. During the test phase, male and female voles spent more time investigating the scent mark of the opposite‐sex conspecific that provided the top‐scent mark than that of a novel, opposite‐sex conspecific. Male and female voles spent similar amounts of time investigating the scent mark of the bottom‐scent donor and that of a novel opposite‐sex conspecific. In Experiment 3, voles were exposed to a mixed‐sex over‐mark that contained the scent mark of an opposite‐sex conspecific with which they had 4 d of olfactory experience. During the test phase, male voles spent more time investigating the mark of the familiar, top‐scent female than the scent mark of a novel female donor but spent similar amounts of time investigating the mark of the familiar, bottom‐scent female and that of a novel female donor. In contrast, female voles spent more time investigating the mark of a novel male donor than that of either the familiar, top‐scent male or that of the familiar, bottom‐scent male. The sex differences in the responses of voles to scent marks and mixed‐sex over‐marks are discussed in relation to the natural history and non‐monogamous mating system of meadow voles.     

Meadow voles,Microtus pennsylvanicus,can discriminate between scents of individual house cats,Felis catus

Individual discrimination provides animals the opportunity to adjust their exposure and behavior when interacting with other animals, both conspecifics and heterospecifics. Meadow voles, Microtus pennsylvanicus, were exposed to scents of house cats, Felis catus. Our first experiment tested whether meadow voles could discriminate between caudal, interdigital, and facial scents produced by a cat with a habituation–dishabituation paradigm. Upon exposure to the familiar scent and a novel one, meadow voles did not investigate either scent more than the other. Our second experiment tested whether meadow voles discriminate between the facial scents of different cats. When exposed to a familiar scent of one cat and the unfamiliar scent of another cat, the meadow voles did discriminate and investigated the unfamiliar scent more than the familiar scent. The results suggest that meadow voles will discriminate between cats using any scent that the cat may inadvertently leave within the environment, thereby reducing the vole's risk of predation by that individual.