Motivational and Heritable Determinants of Dispersal Latency in Wild Male House Mice (Mus musculus musculus)

Divergence of dispersal regimens has been suggested to be the selective basis for the evolutionary differentiation of agonistic phenotypes found in natural populations of house mice. Dispersal propensity may, therefore, be expected to exhibit heritable variation in wild house mice, ultimately related to motivational differences causing observable differences in agonistic behaviour. To test for heritable components in dispersal propensity in wild house mice, father–offspring regressions of dispersal latencies from residential social groups were determined in standardized seminatural social settings. To evaluate potential motivational causes of phenotypic variation in dispersal behaviour, all test animals (fathers, sons, and daughters) were scored prior to the dispersal experiment in a standardized behavioural test, at 60 d of age. Activities were monitored in a 1 m² square test arena during 10‐min observation periods. Test arenas exhibited four equidistant openings leading to cages containing fresh, own, sibling, or foreign bedding material. The apparatus allowed for scoring anxiety, exploratory activity, and kin preference. Subsequently, test animals were exposed to a resident population in a semi‐natural enclosure providing a dispersal opportunity. Father–son regressions of dispersal latencies were significantly positive, but no significant relationship was found for daughters. Dispersal latency decreased with increasing exploratory activity scores in males, but increased in females. Anxiety as well as kin preferences did not affect dispersal propensity. Hence sex‐linked, motivational components reflect heritable social behaviour variation in male house mice that may ultimately be caused by diverging dispersal regimens.     

Dispersal in house mice

This review evaluates direct (live-trapping) and indirect (genetic) methods to study dispersal in wild house mice ( Mus musculus ) and summarizes field and experimental data to examine the causes and consequences of dispersal. Commensal house mice (associated with human habitations, farms, food stores and other anthropogenic habitats) typically show lower rates of dispersal than feral house mice (living in crops, natural and semi-natural habitats). However, early claims of long-term fine-scale genetic structure in commensal house mice (due to low rates of dispersal) are not supported by recent data. Dispersal becomes obligatory when habitat conditions deteriorate, but most dispersal occurs below the local environmental carrying capacity and is due to social interactions with conspecifics. Excursions are relatively frequent and probably allow mice to assess the quality of habitats before dispersing. Young males have the greatest tendency to disperse, apparently prompted mainly by aggressive interactions with dominant males. If they do disperse, females integrate into new groups more easily than do males. Dispersing house mice risk loss of condition or death, but may gain reproductive opportunities on arrival in a new location. House mice can be transported passively as stowaways with humans; this contributes to population persistence and genetic structure at regional scales and has allowed house mice to spread world-wide.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 565–583.     

Effects of early weaning and housing conditions on the development of stereotypies in farmed mink

Female mink pups were weaned at 6, 8 or 10 weeks of age and subjected to two different housing conditions. They were either kept together with a single male sibling in traditional mink cages (30x45x90 cm) or housed socially with all litter-mates in an alternative system consisting of three adjoining traditional cages (90x45x90 cm). All cages were supplied with nest boxes. At 5 months of age, the siblings were removed leaving the females socially isolated in the two different cage systems. Females' stereotypies were quantified by repeated scanning observations under the social housing conditions immediately before removal of the siblings, and again at the age of 7 and 9 months, when the animals had stayed solitary in the two systems for 2 and 4 months. Solitary females showed significantly more stereotypies than females under social housing conditions in both cage systems. Stereotypies were more frequent in the smaller traditional cages. Stereotypies declined from 7 to 9 months of age among solitary animals in traditional cages but not in alternative cages. Early-weaned solitary females in traditional cages showed more stereotypies than later-weaned animals, but only when measured at the age of 7 months. It is suggested that early weaning, individual housing and small cages promote the development of stereotypies in farmed mink. The influence of early weaning on stereotypies seems to decline with age, while effects related to individual housing and small cages appear to be more persistent.     

Mate Choice for Non-siblings in Wild House Mice: Evidence from a Choice Test and a Reproductive Test

Two experiments were designed to test whether wild house mice discriminate between olfactory cues from different kin and, if so, whether given preferences would relate to actual reproductive decisions. Experimental animals were mice born to the offspring of wild-caught house mice. Litter-mates stayed together until 60 d of age and were then housed individually. In a choice test, animals were placed in the middle of an arena with 4 openings which led to small cages containing bedding material from opposite-sex animals of known kinship (full-sib, cousin, unrelated) or clean material. Test animals (11 oestrous females, 11 males tested with oestrous females' bedding, 8 males tested with material from non-oestrous females) preferred conspecific to control bedding. Males tested with oestrous females' bedding significantly preferred unrelated to full-sib odours. In a second experiment, 34 males were each mated simultaneously to 3 females (sister, cousin, unrelated) and these groups were then housed together for 5, 10, and 15 d. Females were checked for litters during the next 20 d. Reproductive rate increased significantly in the 15 d cohabitation group, and significantly more cousin and unrelated females than sisters gave birth to a litter.     

Intermittent stimulation and delay of puberty by urinary chemosignals and social contact in wild stock female house mice (Mus Domesticus)

A sequence of six experiments using wild stock house mouse (Mus domesticus) tested the effects of intermittent stimulation with either the urinary chemosignal released by grouped female mice or social contact from grouped females on the age of first vaginal oestrus in young females. Weanling female mice were exposed to bedding soiled by grouped females or cages containing grouped females for 15 min periods, then removed for a prescribed period, and placed again in a cage with soiled bedding or grouped females. The nature of the exposure to the puberty delaying effect, the number of total exposures each day, the total length of exposure to the stimulus, and the total time period over which the exposures occurred were varied. None of the treatment regimes employed here with soiled bedding from grouped females resulted in delays in the onset of first oestrus in test females. Young females exposed to grouped females for 6 or 8 exposures in a 4 h period, 6 or 8 exposures in an 8 h period, or 8 exposures in a 12 h period were significantly delayed in attaining puberty relative to control females that were exposed to cages containing clean bedding. These results are in contrast to earlier findings involving chemosignals that accelerate first oestrus wherein young females exhibited the capacity to accumulate the exposures to the urinary chemosignals from males, females in oestrus and pregnant or lactating females. Direct exposure to the grouped females on an intermittent basis can provide stimulation that is cummulative and results in delays in the onset of first oestrus.     

Response,use and habituation to a mouse house in C57BL/6J and BALB/c mice

Animal welfare depends on the possibility to express species-specific behaviours and can be strongly compromised in socially and environmentally deprived conditions. Nesting materials and refuges are very important resources to express these behaviours and should be considered as housing supplementation items. We evaluated the effects of one item of housing supplementation in standard settings in laboratory mice. C57BL/6JOlaHsd (B6) and BALB/cOlaHsd (BALB) young male and female mice, upon arrival, were housed in groups of four in standard laboratory cages and after 10 days of acclimatization, a red transparent plastic triangular-shaped Mouse House™ was introduced into half of the home cages. Animals with or without a mouse house were observed in various contexts for more than one month. Body weight gain and food intake, home cage behaviours, emotionality and response to standard cage changing procedures were evaluated. The presence of a mouse house in the home cage did not interfere with main developmental and behavioural parameters or emotionality of BALB and B6 male and female mice compared with controls. Both strains habituated to the mouse house in about a week, but made use of it differently, with BALB mice using the house more than the B6 strain. Our results suggest that mice habituated to the mouse house rather quickly without disrupting their home cage activities. Scientists can thus be encouraged to use mouse houses, also in view of the implementation of the EU Directive (2010/63/EU).     

Dispersal behaviour of African wild dogs in Kenya

Dispersal behaviour plays a key role in social organisation, demography and population genetics. We describe dispersal behaviour in a population of African wild dogs (Lycaon pictus) in Kenya. Almost all individuals, of both sexes, left their natal packs, with 45 of 46 reproductively active “alpha” individuals acquiring their status through dispersal. Dispersal age, group size and distance did not differ between males and females. However, only females embarked on secondary dispersal, probably reflecting stronger reproductive competition among females than males. When dispersing, GPS-collared wild dogs travelled further than when resident, both in daylight and by night, following routes an order of magnitude longer than the straight-line distance covered. Dispersers experienced a daily mortality risk three times that experienced by adults in resident packs. The detailed movement data provided by GPS-collars helped to reconcile differences between dispersal patterns reported previously from other wild dog populations. However, the dispersal patterns observed at this and other sites contrast with those assumed in published demographic models for this endangered species. Given the central role of dispersal in demography, models of wild dog population dynamics need to be updated to account for improved understanding of dispersal processes.     

Hunter feedback of individually marked wild boar <Emphasis Type="Italic">Sus scrofa</Emphasis> L.: dispersal and efficiency of hunting in northeastern Germany

Increasing wild boar (Sus scrofa L.) population densities all over Europe cause severe economic problems. For understanding mechanisms of epidemics, the knowledge of dispersal is required. Thus, we investigated dispersal rates and distances with regard to sex and age of wild boar in southwestern Mecklenburg-Western Pomerania. From 152 marked wild boar, 105 have been registered as dead, of which, 51% were males and 49% females. Forty-five percent were shot as piglets, 41% as yearlings, and 14% as adults. The distance between capture site and site of death ranged between 184 m and 41.5 km. Piglets were shot closer to their capture site (mean distance 1 km) than older animals (mean 4 km), although this difference was only significant for males. In general, males tended to disperse further before being shot (3.8 km) than females (1.6 km). Only 3.8% of all animals were shot at distances larger than 10 km. As most animals (84.6%) were shot inside their natal home range, only a small proportion (15.4%) did actually disperse (shot outside mothers home range), which is 32% of all animals surviving to the age of yearlings. Of those dispersed animals, 25% were females. The low dispersal rate is biased by female philopatry and allows actual dispersal only at very high population densities or in sparsely populated regions. In consideration for the low natural mortality proved by radio-tagged animals, the harvest rate is lower than the net reproduction. We did not detect any sex-biased hunting. The dominating hunting method was single hunt at bait, although drive hunts are highly effective. However, hunting rates on piglets and females were too low for regulating the population.     

Seasonal variation in fertility, fecundity and litter sex ratio in laboratory and wild stocks of house mice (Mus domesticus)

Data compilations were made for three parameters pertaining to reproduction in a domestic strain (14 years) and a wild stock (4 years) of commensal house mice: (a) the percentage of females mated that produced litters; (b) the average number of pups per litter; and (c) the sex ratio of the pups in the litters. Fecundity and fertility varied seasonally in both domestic and wild mice. More females become pregnant and litter sizes were larger in the spring, summer, and fall months than during the winter season. Sex ratios also varied seasonally with more male biased litters produced during the 4 winter months. There appear to be seasonal shifts in productivity for both stocks of mice and these seasonal trends have not been altered by domestication under laboratory conditions. In spite of the fact that house mice are generally opportunistic, it is possible that there has been selection in the mice for shifting rates of production in relation to the best seasons of the year in terms of climate and resource availability. The higher production of males during summer months may be geared toward greater success in dispersal at that season and a higher probability that the males can find a territory and mate successfully in summer rather than winter. These results have potential implications for animal breeders and for those who maintain mouse colonies to produce animals for scientific investigations.     

Cage-change interval preference in mice

Before animal research facilities began using individually ventilated cage (IVC) systems for mice, cages were often changed one or more times per week. When using IVC systems, however, it is standard practice to change cages only once every 2-3 weeks. When deciding how often to change cages, personnel may consider the cost of labor needed to change the cage, as well as the cage type and bedding type, rather than animal preference or concern for animal well-being. The authors carried out a simple preference test in groups of mice. Mice were allowed to choose between an unsoiled cage and cages that had not been changed for 1 d, 7 d or 14 d. When evaluating where mice positioned their nests and the amount of time mice spent in the various cages, the authors found that the mice preferred the unsoiled cage. Younger mice (<150 d old) showed a stronger preference for the unsoiled cage than did older mice (>150 d old). Further studies are warranted to evaluate mice's preferences for cages changed at different intervals and to determine whether prolonging the interval between cage changes has any negative effects on mice.     

Trypanosoma cruzi: effects of social stress in Calomys callosus a natural reservoir of infection

Social environment can represent a major source of stress affecting cortisol and/or corticosterone levels, thereby altering the immune response. We have investigated the effects of social isolation on the development of Trypanosoma cruzi infection in female Calomys callosus, a natural reservoir of this protozoan parasite. Animals were divided in groups of five animals each. The animals of one group were kept together in a single cage. In a second group, four females were kept together in a cage with one male. In the final group, five individuals were kept isolated in private cages. The isolated animals showed body weight reduction, decreased numbers of peritoneal macrophages, lower global leucocytes counts, smaller lytic antibody percentage and a significantly higher level of blood parasites compared to the other animals. Their behavior was also altered. They were more aggressive than grouped females, or females exposed to the presence of a male. These results suggest that isolation creates a distinct social behavior in which immunity is impaired and pathogenesis is enhanced.     

Aggressive behaviour in related and unrelated wild house mice (Mus musculus L.)

Aggressive behaviour was observed to be rare in small family groups of confined wild house mice, Mus musculus L. Unrelated mice were attacked when they were introduced to a family group and in their presence intra-family aggressive behaviour increased. When two family groups of mice were allowed to meet there were frequent aggressive encounters between unrelated animals and the two groups remained separate. Resident mice were found to be aggressive towards males and females individually isolated and returned to their own family after 2 or 3 weeks absence but not after 1 week. The possibility is discussed that in wild mice odour discrimination influences the dispersal and build-up of free-living populations.     

Impact of Social Ties on Dispersal,Reproduction and Dominance in Feral House Mice (Mus musculus domesticus)

The existence of a relationship between the social ties an individual has to other family members and its further role within the family was tested in feral house mice (Mus musculus domesticus), according to the ‘Social cohesions hypothesis’. It is predicted by the hypothesis that individuals not forming strong social ties are the first who emigrate. House mice were studied using a population cage system that allowed continuous observation of individually marked animals. Data on time staying with other animals (social ties), aggressive interactions, body weight, reproduction, and emigration were collected daily. The results may be summarized as follows:

• 1 Male emigrants were less integrated in cohorts of male littermates compared with their brothers of the same age. These male cohorts appeared to protect single males from attacks by the dominant male. No difference could be observed in social ties to other family members.
• 2 After weaning, there was no difference in social ties of male and female offspring. However, after sexual maturation social ties of males decreased significantly while those of females remained almost constant.
• 3 Female emigrants showed the same intensity of social ties as their resident sisters.
• 4 No difference could be found between social ties of females becoming pregnant and their nonreproductive sisters of the same age. Reproduction or reproductive suppression could not be explained by having more or less contact with other reproductive females.
• 5 Dominant males spent least of all time with other family members.

The social cohesions hypothesis has to be rejected in analysing proximate causes of emigration. In house mice, male emigration was caused by aggression of the dominant male in competition for the top rank within the group. This was enhanced by a lower integration in the group of same-aged brothers but is not related to a lack of integration into the family.     

The effect of preweaning and postweaning housing on the behaviour of the laboratory mouse (Mus musculus)

Standard housing for laboratory mice severely restricts natural behaviour and the control that the animal has over its environment. Providing the cage with objects is a method that has been used to both increase environmental complexity, promote the performance of natural behaviour and provide greater controllability for the animal. This method of furnishing cages has mostly been studied in adult animals, and little is known about the influence that the preweaning environment has on the behaviour of mice as adults. This study aimed to investigate the effects on mice behaviour of preweaning and postweaning housing environment. In this experiment, 64 pairs of animals of the strain C57BL/6J were used. Half of the animals were born and reared until weaning in standard cages and the other half in cages twice the size of the standard and furnished with nesting material, a cardboard tube, a PVC nest box and a wooden chewblock. After weaning, half the animals in each group were changed to the other type of cage, whereas the other half remained in the same environment; in both cases they were kept in single-sex pairs of littermates. Behaviour during the dark, active period was studied through video recordings. We found no main effects of preweaning environment on behaviour; however, mice moved from furnished to standard cages at weaning showed a decrease in inactive behaviour at four weeks of age. Mice housed after weaning in standard cages spent less time inactive, and more time engaging in activities like feeding and drinking, self-grooming and allogrooming. A sex difference was also found, in that females showed a greater performance of exploratory behaviour as well as a greater prevalence of stereotypies. The use of different objects and locations within the furnished cage was also analysed at both ages. Results show that at eight weeks of age mice spent more time at the top of the cage, and that the use of the nest box (although not for resting) increased between four and eight weeks. Mice were found to use the nest box as a nesting site/sleeping place only at age four weeks, whereas they always used the nesting material for sleeping.     

A qualitative investigation of major urinary proteins in relation to the onset of aggressive behavior and dispersive motivation in male wild house mice (<Emphasis Type="Italic">Mus musculus domesticus</Emphasis>)

The physiological basis for population differentiation of dispersal timing during individual development in male wild house mice is still unknown. As major urinary proteins (MUPs) are known to convey information about competitive ability in male mice, we examined individual MUP profiles defined by isoelectric-focusing (IEF) patterns in relation to developmental timing of dispersive motivation. As an experimental paradigm marking the development of the dispersal propensity, we used agonistic onset between litter mate brothers when kept in pairs under laboratory conditions. Agonistic onset is known to reflect the initiation of dispersive motivation. Hence, we compared individual MUP IEF patterns between fraternal pairs that did or did not develop agonistic relationships before the age of 2 months. Urine was collected on the day of weaning and at the beginning of adulthood. We investigated whether there was a significant co-occurrence of particular MUP IEF patterns with the agonistic onset in male mice. We assumed that, based on this co-occurrence, particular MUP IEF patterns and/or a particular dynamic of MUP IEF expression from weaning to adulthood may be considered a physiological predictor of a specific behavioral strategy in male mice (i.e. submissive-philopatric or agonistic-dispersive strategy). We found that agonistic males expressed more MUP IEF bands than amicable ones at weaning, but these differences disappeared later on. The presence of two particular IEF bands at weaning was significantly associated with early agonistic onset. Our study suggests that MUPs could have a predictive value for the onset of aggressive behavior and dispersal tendency in male wild house mice.     

Cage design reduces emotionality in mice

To see if a more natural cage design would alter the reactivity of laboratory mice, 192 mice were reared in cages with (1) no dividers, (2) five vertical dividers, (3) nine vertical dividers, or (4) nine vertical dividers and one horizontal platform. The mice preferred the most complex cages, and on almost all measures they were less emotional when reared in the more complex cages. Results suggest that a more natural housing environment would lead to healthier animals.     

Transient hypertension and sustained tachycardia in mice housed individually in metabolism cages

The novel environment of a metabolic cage can be stressful for rodents, but few studies have attempted to quantify this stress-response. Therefore, we determined the effects on mean arterial pressure (MAP) and heart rate (HR), of placing mice of both sexes in metabolism cages for 2 days. After surgical implantation of a carotid artery catheter mice recovered individually in standard cages for 5 days. Mice then spent 2 days in metabolism cages. MAP and HR were monitored in the standard cage on Day 5 and in metabolism cages on Days 6-7. MAP increased by 18+/-3 and 22+/-4 %, while HR increased by 27+/-4 and 27+/-6 %, in males and females, respectively, during the first hours after cage switch. MAP decreased to baseline in the fourth and eighth h following metabolism cage switch in males and females, respectively. However, HR remained significantly elevated in both sexes during the entire two-day period in metabolism cages. Females had lower MAP than males both pre- and post-metabolism cage switch, but there were no sex differences in HR. These results demonstrate sustained changes in cardiovascular function when mice are housed in metabolism cages, which could potentially affect renal function.     

Female house mice develop a unique ovarian lesion in colonies that are at maximum population density

Colonies of house mice reach maximum population density in 120-180 days, irrespective of cage size and initial number of colonizing animals. Reproduction ceases because the females become aggressive and unreceptive to mating. The aggressive behavior is correlated with elevated levels of testosterone (T) and corticosterone (B) (Chapman et al., Phys Behav 64:529-533, 1998). In two of seven strains of mice, females developed ovarian lesions. The occurrence of the lesion in one strain was correlated with the age of the animal and duration of the study. In the second strain, cage size was the determining factor. Lesioned ovaries weighed significantly more than nonlesioned ovaries. The lesion consisted of accumulations of luteal membrane and organelle fragments, and other cellular debris, suggestive of incomplete and prolonged luteolysis. Electron microscopic (EM) analyses revealed the presence of deposits of permanganate-resistant congophilic amyloid fibrils in the intima and smooth muscle cells of luteal thecal arteries. Population females had thymus glands and uteri that weighed significantly less than the same organs from females housed in the breeding colony, whereas the adrenal glands from the population females weighed significantly more. It is proposed that the female aggression is due to high levels of T. It is also proposed that the high levels of B suppress the immune cells involved in normal luteolysis and contribute to the incomplete and prolonged luteolysis.     

Genetic Basis of Mating Preferences in Wild House Mice

This paper reviews work conducted over the last several yearson the effect of genetic differences within the t-complex ofwild house mice on female mating preference. Wild mice are polymorphicfor a mutation within the t complex on chromosome 17. About25% of wild mice are heterozygous (+/t) for a t-haplotype andthe remainder are +/+. These t-haplotypes have a number of deleteriouseffects when homozygous and hence t/t individuals are rarelyfound in wild populations. We have examined preferences of +/+and +/t females for males of both genotypes. We have found that+/t, but not +/+ females have strong preferences for +/+ males.These preferences can be modified by a variety of factors includingestrous condition of the female (the preferences are strongeramong estrous than diestrous females) and the dominance statusof the male (when forced to choose, females give priority tomale dominance status over t complex genotype in choosing males).The restiction of preference to +/t females indicates that geneson t haplotypes modulate these preferences. Because t haplotypesinclude the major histocompatibility complex (MHC) of the mousewe designed a study to ascertain whether the preferences of+/t females were associated with the MHC. Results of the studyindicate that the preferences are independent of the MHC. Furtherwork testing females carrying a partial t-haplotype (t^w18) indicatesthat the genes for mating preference are localized in the regionof the t complex distal to the MHC. A large number of t haplotypesare found in wild mouse populations. Females that are themselves+/t when forced to choose between 2 +/t males (one carryinga haplotype that is the same as their own and one carrying ahaplotype that is different) prefer males carrying t-haplotypesthat differ from their own. Finally, we conclude that matingpreference may only be a weak force regulating the frequencyof t-mutations in wild mouse populations. The impact of matingpreference on population genetics of genes within this regionis muted because of the great importance of male dominance rankin determining mating patterns within interacting social groups.     

Characterizing spring emergence of adult Halyomorpha halys using experimental overwintering shelters and commercial pheromone traps

To improve our understanding of adult Halyomorpha halys (Stål) (Hemiptera: Pentatomidae) overwintering biology and to better inform models of its population dynamics, its temporal pattern of spring emergence was investigated using experimental overwintering shelters in screened cages within protective structures. In 2012, plastic shelters containing 100 adults were deployed in unheated, unlighted buildings, and adjacent woodlots in Virginia, USA. In 2013 and 2014, wooden shelters containing 300 paint‐marked adults were deployed in pairs in six woodlots across Virginia, West Virginia, and Maryland, USA; one in a closed cage and one in a cage with the top removed, enabling emerged adults to be counted or to disperse, respectively. In 2013 and 2014, pheromone‐baited and non‐baited pyramid traps encircled the shelters at each site. Regular counts of adults that emerged into the closed cage and of marked and ‘wild’ (unmarked) adults captured in traps were conducted from February or March through early July. In 2012, emergence patterns from shelters in buildings and woodlots were very similar and matched those recorded from woodlots in 2013 and 2014. In all years, a small peak of emergence occurred in about mid‐April, a larger and more prolonged peak was observed between mid‐May and early June, and emergence ended by early July. Of the 449 H. halys adults captured in traps between 2013 and 2014, only three were marked individuals from shelters in the open cage, suggesting that adults emerging from overwintering sites may require a dispersal flight before responding to pheromone‐baited traps. In 2013 and 2014, respectively, 98 and 93% of captures were in pheromone‐baited traps, but there was no correlation between the weekly number of adults that emerged from shelters in the closed cages and captures in traps.