Naloxone increases conditioned fear responses during social buffering in male rats.

Social buffering is the phenomenon in which the presence of an affiliative conspecific mitigates stress responses. We previously demonstrated that social buffering completely ameliorates conditioned fear responses in rats. However, the neuromodulators involved in social buffering are poorly understood. Given that opioids, dopamine, oxytocin and vasopressin play an important role in affiliative behaviour, here, we assessed the effects of the most well-known antagonists, naloxone (opioid receptor antagonist), haloperidol (dopamine D2 receptor antagonist), atosiban (oxytocin receptor antagonist) and SR49059 (vasopressin V1a receptor antagonist), on social buffering. In Experiment 1, fear-conditioned male subjects were intraperitoneally administered one of the four antagonists 25 min prior to exposure to a conditioned stimulus with an unfamiliar non-conditioned rat. Naloxone, but not the other three antagonists, increased freezing and decreased walking and investigation as compared with saline administration. In Experiment 2, identical naloxone administration did not affect locomotor activity, anxiety-like behaviour or freezing in an open-field test. In Experiment 3, after confirming that the same naloxone administration again increased conditioned fear responses, as done in Experiment 1, we measured Fos expression in 16 brain regions. Compared with saline, naloxone increased Fos expression in the paraventricular nucleus of the hypothalamus and decreased Fos expression in the nucleus accumbens shell, anterior cingulate cortex and insular cortex and tended to decrease Fos expression in the nucleus accumbens core. Based on these results, we suggest that naloxone blocks social buffering of conditioned fear responses in male rats.

Social buffering enhances extinction of conditioned fear responses by reducing corticosterone levels in male rats.

The presence of an affiliative conspecific reduces stress responses to a wide variety of stimuli, which is termed "social buffering." We previously reported that social buffering in male rats ameliorated behavioral responses, as well as hypothalamic-pituitary-adrenal axis activation, elicited by an auditory conditioned stimulus (CS). In addition, subjects that experienced social buffering did not show stress responses when re-exposed to the CS the next day in the absence of an accompanying rat. However, the mechanisms underlying this enhancement of between-session extinction are poorly understood. In Experiment 1, we compared corticosterone levels at 0, 10, and 15 min after extinction training. Subjects that experienced social buffering had lower corticosterone levels than subjects that trained alone at the end of extinction training. However, corticosterone levels at 10 and 15 min after training were not affected by the experience of social buffering. These results suggest that a lower level of corticosterone during extinction training had an important role in the enhancement of extinction. To directly assess this, in Experiment 2, we manipulated the corticosterone level during extinction training. We found that a subcutaneous injection of corticosterone before extinction training blocked the enhancement of extinction by social buffering. These results demonstrate that the enhancement is caused by a low level of corticosterone during the training. Taken together, we suggest that social buffering enhances extinction of conditioned fear responses by reducing corticosterone levels in male rats.

Identification of a pheromone that increases anxiety in rats.

Chemical communication plays an important role in the social lives of various mammalian species. Some of these chemicals are called pheromones. Rats release a specific odor into the air when stressed. This stress-related odor increases the anxiety levels of other rats; therefore, it is possible that the anxiety-causing molecules are present in the stress-related odorants. Here, we have tried to identify the responsible molecules by using the acoustic startle reflex as a bioassay system to detect anxiogenic activity. After successive fractionation of the stress-related odor, we detected 4-methylpentanal and hexanal in the final fraction that still possessed anxiogenic properties. Using synthetic molecules, we found that minute amounts of the binary mixture, but not either molecule separately, increased anxiety in rats. Furthermore, we determined that the mixture increased a specific type of anxiety and evoked anxiety-related behavioral responses in an experimental model that was different from the acoustic startle reflex. Analyses of neural mechanisms proposed that the neural circuit related to anxiety was only activated when the two molecules were simultaneously perceived by two olfactory systems. We concluded that the mixture is a pheromone that increases anxiety in rats. To our knowledge, this is the first study identifying a rat pheromone. Our results could aid further research on rat pheromones, which would enhance our understanding of chemical communication in mammals.

Social buffering ameliorates conditioned fear responses in female rats.

The stress experienced by an animal is ameliorated when the animal is exposed to distressing stimuli along with a conspecific animal(s). This is known as social buffering. Previously, we found that the presence of an unfamiliar male rat induced social buffering and ameliorated conditioned fear responses of a male rat subjected to an auditory conditioned stimulus (CS). However, because our knowledge of social buffering is highly biased towards findings in male subjects, analyses using female subjects are crucial for comprehensively understanding the social buffering phenomenon. In the present studies, we assessed social buffering of conditioned fear responses in female rats. We found that the estrus cycle did not affect the intensity of the rats' fear responses to the CS or their degree of vigilance due to the presence of a conspecific animal. Based on these findings, we then assessed whether social buffering ameliorated conditioned fear responses in female rats without taking into account their estrus cycles. When fear conditioned female rats were exposed to the CS without the presence of a conspecific, they exhibited behavioral responses, including freezing, and elevated corticosterone levels. By contrast, the presence of an unfamiliar female rat suppressed these responses. Based on these findings, we conclude that social buffering can ameliorate conditioned fear responses in female rats.

The strain of an accompanying conspecific affects the efficacy of social buffering in male rats.

Social buffering is a phenomenon in which stress in an animal is ameliorated when the subject is accompanied by a conspecific animal(s) during exposure to distressing stimuli. We previously reported that in male Wistar rats, the presence of another Wistar rat mitigates conditioned fear responses to an auditory conditioned stimulus (CS). Subsequent analyses revealed several characteristics of this social buffering of conditioned fear responses. However, information regarding the specificity of accompanying conspecifics is still limited. In the present study, we assessed whether rats of other strains could induce social buffering in Wistar rats. When a fear-conditioned Wistar subject was re-exposed to the CS alone, we observed increased freezing and decreased investigation and walking, as well as elevated corticosterone levels. The presence of a Wistar, Sprague-Dawley, or Long-Evans rat blocked these responses, suggesting that social buffering was induced by these strains of rats. In contrast, a Fischer 344 rat did not induce social buffering in the Wistar subject. We further found that an inbred Lewis rat induced social buffering whereas a Brown Norway rat, a strain that has been established independently from Wistar rats, did not. These results suggest that the difference in origin, rather than the inbred or outbred status of the associate rat, seemed to account for the lack of social buffering induced by the F344 rats. Based on these findings, we conclude that strains of an accompanying conspecific can affect the efficacy of social buffering in rats.

Social buffering reduces male rats' behavioral and corticosterone responses to a conditioned stimulus.

In social species, the presence of an affiliative same-sex conspecific ameliorates acute stress responses in threatening conditions. We previously found that the presence of an unfamiliar male rat separated by a wire mesh barrier blocks the behavioral responses and Fos expression in the paraventricular nucleus of the hypothalamus (PVN) in a male subject rat that had previously been exposed to an auditory conditioned stimulus (CS) paired with foot shocks. Based on the Fos expression in the PVN, we hypothesized that the presence of a conspecific ameliorated the hypothalamic-pituitary-adrenal (HPA) axis activation and induced social buffering of conditioned fear responses. The direct evidence for this hypothesis, however, is still lacking. To clarify this point, we exposed fear-conditioned and non-conditioned subjects to the CS either alone or with a conspecific separated by a wire mesh barrier. When the fear-conditioned subject alone was re-exposed to the CS, it exhibited increased freezing, decreased sniffing, and elevated corticosterone levels. In contrast, the presence of the conspecific suppressed these behavioral and HPA axis responses to a level similar to those observed in the non-conditioned subjects. These results suggest that the presence of a conspecific suppressed the behavioral responses and HPA axis activation to the CS. The present results provide direct evidence for the existence of social buffering of conditioned fear responses in male rats.

Coexistence of roof rats and carnivores in barns on a livestock farm in Japan.

Brown rats (Rattus norvegicus), roof rats (Rattus rattus), and house mice (Mus musculus) are considered to be important pests on livestock farms. Although the diel activity patterns of rodents are key to their control, information on this aspect of their ecology is limited. Furthermore, the effect of carnivores on rodent activity patterns as well as the carnivore species present on livestock farms is unclear. Here, we set camera traps in an open-type cow barn and in an enclosed pig barn on the same livestock farm in Japan from August through October 2021. The only rodents observed in both barns were roof rats, and the carnivore species observed were dogs (Canis familiaris), cats (Felis catus), and Japanese weasels (Mustela itatsi). Roof rats showed different patterns of activity and behavior between the barns. However, because the pattern in both barns was nocturnal, the activity patterns of roof rats and carnivores showed a moderate to high degree of overlap. Therefore, roof rats did not appear to shift their activity patterns to avoid nocturnal carnivores. Taken together, the present study provides valuable information for rodent control on livestock farms in Japan.

Approach/Avoidance Behavior to Novel Objects is Correlated with the Serotonergic and Dopaminergic Systems in the Brown Rat (Rattus norvegicus).

The brown rat (Rattus norvegicus) is known to show three types of behavioral responses to novel objects. Whereas some rats are indifferent to novel objects, neophobic and neophilic rats show avoidance and approach behavior, respectively. Here, we compared the dopaminergic, serotonergic, and noradrenergic systems immunohistochemically among these rats. Trapped wild rats and laboratory rats were first individually exposed to the novel objects in their home cage. Wild rats were divided into neophobic and indifferent rats depending on their behavioral responses. Similarly, laboratory rats were divided into neophilic and indifferent rats. Consistent with the behavioral differences, in the paraventricular nucleus of the hypothalamus, Fos expression in corticotropin-releasing hormone-containing neurons was higher in the neophobic rats than in the indifferent rats. In the anterior basal amygdala, the neophobic rats showed higher Fos expression than the indifferent rats. In the posterior basal amygdala, the neophobic and neophilic rats showed lower and higher Fos expressions than the indifferent rats, respectively. When we compared the neuromodulatory systems, in the dorsal raphe, the number of serotonergic neurons and Fos expression in serotonergic neurons increased linearly from neophobic to indifferent to neophilic rats. In the ventral tegmental area, Fos expression in dopaminergic neurons was higher in the neophilic rats than in the indifferent rats. These results demonstrate that approach/avoidance behavior to novel objects is correlated with the serotonergic and dopaminergic systems in the brown rat. We propose that the serotonergic system suppresses avoidance behavior while the dopaminergic system enhances approach behavior to novel objects.

Alarm pheromone is detected by the vomeronasal organ in male rats.

It is widely known that a stressed animal releases specific pheromones, possibly for alarming nearby conspecifics. We previously investigated an alarm pheromone in male rats and found that this alarm pheromone evokes several responses, including increases in the defensive and risk assessment behaviors in a modified open-field test, and enhancement of the acoustic startle reflex. However, the role of the vomeronasal organ in these pheromone effects remains unclear. To clarify this point, vomeronasal organ-excising or sham surgeries were performed in male rats for use in 2 experimental models, after which they were exposed to alarm pheromone. We found that the vomeronasal organ-excising surgery blocked the effects of this alarm pheromone in both the modified open-field test and acoustic startle reflex test. In addition, the results of habituation/dishabituation test and soybean agglutinin binding to the accessory olfactory bulb suggested that the vomeronasal organ-excising surgery completely ablated the vomeronasal organ while preserving the functioning of the main olfactory system. From the above results, we showed that the vomeronasal organ plays an important role in alarm pheromone effects in the modified open-field test and acoustic startle reflex test.

Parallel Olfactory Systems Synergistically Activate the Posteroventral Part of the Medial Amygdala Upon Alarm Pheromone Detection in Rats.

In rats, a mixture of hexanal and 4-methylpentanal is a main component of the alarm pheromone. When detected by the main olfactory system (MOS) and the vomeronasal system, respectively, they activate the anterior part of the bed nucleus of the stria terminalis (BNSTa). Therefore, the information from the two olfactory systems is expected to be integrated before being transmitted to the BNSTa. To specify the integration site, we examined Fos expression in 16 brain regions in response to water (n = 10), hexanal (n = 9), 4-methylpentanal (n = 9), the mixture (n = 9), or the alarm pheromone (n = 9) in male rats. The posteroventral part of the medial amygdala showed increased Fos expression to hexanal and 4-methylpentanal. The expression was further increased by the mixture. Therefore, this region is suggested as the integration site. In addition, the BNSTa, paraventricular nucleus of the hypothalamus, and anteroventral, anterodorsal, and posterodorsal parts of the medial amygdala were suggested to be located downstream of the integrated site because only the mixture increased Fos expression. We suggest that the posterolateral part of the cortical amygdala is upstream of the integration site in the MOS because all stimuli increased Fos expression. The posterior part of the bed nucleus of the stria terminalis and posteromedial part of the cortical amygdala were suggested as being located upstream in the vomeronasal system because 4-methylpentanal and the mixture increased Fos expression. These results provide information about the neural pathway underlying the alarm pheromone effects.

Glutamatergic Projections from the Posterior Complex of the Anterior Olfactory Nucleus to the Amygdala Complexes.

Social buffering is a phenomenon where stress responses are ameliorated by an affiliative conspecific. Our previous findings suggest that the posterior complex of the anterior olfactory nucleus (AOP) is well positioned to participate in the neural mechanisms underlying social buffering. However, the lack of anatomical information prevents us from further estimating the role of the AOP. Here, we obtained anatomical information regarding the AOP in male rats. In Experiment 1 (n = 5), among 4',6-diamidino-2-phenylindole-positive cells in the AOP, the proportion of glutamic acid decarboxylase 67 (GAD67)-positive cells was 13.8% ± 1.2%. In Experiment 2 (n = 5), among the cells that were labeled by a retrograde tracer injected into the basolateral complex of the amygdala (BLA), the proportion of GAD67-positive cells was 18.6% ± 0.8%. In Experiment 3 (n = 5), we demonstrated the existence of cells that were labeled by the retrograde tracer injected into the posterior part of the medial amygdala (MeP), mostly into the ventral part of the MeP. In addition, the proportion of GAD67-positive cells among the tracer-labeled cells was 21.7% ± 1.7%. In Experiment 4 (n = 3), the retrograde tracers were injected into the BLA and MeP, mostly into the ventral part of the MeP. The proportion of double-labeled cells among the tracer-labeled cells was 2.1% ± 1.2%. Taken together, these results suggest that the AOP is predominantly composed of glutamatergic neurons. In addition, the AOP sends mutually independent glutamatergic-predominant projections to the BLA and MeP.

An appeasing pheromone ameliorates fear responses in the brown rat (Rattus norvegicus).

The brown rat (Rattus norvegicus) is one of the major animals both in the laboratory and in urban centers. Brown rats communicate various types of information using pheromones, the chemicals that mediate intra-species communication in minute amounts. Therefore, analyses of pheromones would further our understanding of the mode of life of rats. We show that a minute amount of 2-methylbutyric acid (2-MB) released from the neck region can ameliorate fear responses both in laboratory rats and in wild brown rats. Based on these findings, we conclude that 2-MB is an appeasing pheromone in the brown rat. A better understanding of rats themselves would allow us to perform more effective ecologically based research on social skills and pest management campaigns with low animal welfare impacts, which might contribute to furthering the advancement of science and improving public health.

The neural pathway underlying social buffering of conditioned fear responses in male rats.

In social animals, the presence of an affiliative conspecific alleviates acute stress responses, and this is termed social buffering. However, the neural mechanisms underlying social buffering have not been elucidated. We have reported that the main olfactory system mediates social buffering of conditioned fear responses in male rats, and this is accompanied by suppression of the lateral and central amygdala. Therefore, olfactory signals are probably transmitted from the main olfactory system to the amygdala. Because the lateral and central amygdala do not receive projections from the main olfactory bulb, the site that links the main olfactory bulb and amygdala was presumed to be located within the main olfactory system. To find the linkage site, we generated lesions within the main olfactory system, and found that a bilateral lesion in the posteromedial region of the olfactory peduncle (pmOP) blocked social buffering. Next, we determined that the pmOP receives direct projections from the main olfactory bulb. Finally, we demonstrated that the connection between the pmOP and ipsilateral amygdala is important for social buffering, and that the pmOP projects directly to the ipsilateral amygdala, including the lateral and central amygdala. On the basis of these results, we suggest that the pmOP links the main olfactory blub to the amygdala and enables social buffering of conditioned fear responses. These results provide the first comprehensive picture of the neural pathway underlying the social buffering phenomenon.

Social distancing measures differentially affected rats in North America and Tokyo.

Because rats are commensal organisms that depend on human activities for food, shifts in human behavior will have pronounced effects on local rat populations. In the spring of 2020, social distancing measures were implemented globally to curtail the spread of SARS-CoV-2. This presented a unique opportunity to obtain information regarding the immediate effects of shifts in human behavior on rat populations in a variety of countries. In response to increased sightings of rats in the USA that were reported in American media, we analyzed the changes in the number of public service calls in Tokyo, Japan. We found that the number of calls increased after the implementation of social distancing measures, suggesting that rat sightings had also increased in Tokyo. We then surveyed the changes in the business activities of pest management professionals in the USA, Canada, and Tokyo. We found that the activities were increased in 50 to 60% of the respondents from the USA and Canada. In contrast, 60 to 70% of the respondents from Tokyo answered that their activities were not changed. These results implied that, following the implementation of social distancing measures, rat infestations increased in North America, but not in Tokyo. The survey also suggested that roof rats were considered to be the predominant rodent species in Tokyo. This may account for the limited infestations in Tokyo because roof rats are more sedentary than brown rats. Taken together, our findings suggest that social distancing measures differentially affected rat populations in North America and Tokyo. Supplementary Information: The online version contains supplementary material available at 10.1007/s10340-021-01405-z.

Mapping of c-Fos expression in the medial amygdala following social buffering in male rats.

Social buffering is the phenomenon in which an affiliative conspecific (associate) ameliorates stress responses of a subject. We previously found that social buffering in Wistar subject rats is induced if the strain of the associate is Wistar or a strain derived from Wistar rats. In the present study, we assessed the possible role of medial amygdala (Me) in this strain-dependent induction of social buffering. The subjects were exposed to the conditioned stimulus (CS) that had been paired or unpaired with a foot shock either alone, with an unfamiliar Wistar associate, or with an unfamiliar Fischer 344 (F344) associate. We found that the Wistar associates, but not F344 associates, ameliorated increased freezing and Fos expression in the paraventricular nucleus of the hypothalamus and lateral amygdala caused by the CS. In addition, Fos expression in the posterior complex of the anterior olfactory nucleus and lateral intercalated cell mass of the amygdala was increased simultaneously. These results suggest that Wistar associates, but not F344 associates, induced social buffering. In the Me, we did not find any differences associated with stress responses or amelioration of stress responses. In contrast, a comparison among the unpaired subjects found that the Wistar associates, but not F344 associates, increased exploratory behavior and Fos expression in the posteroventral subdivision of the Me (MePV). Based on these results, we propose that the MePV is involved in the recognition of social similarity with the associates. Taken together, the present study provides information about the possible role of Me in social buffering.

The strain of unfamiliar conspecifics affects stress identification in rats.

Humans show distinct social behaviours when we evaluate an individual as being a member of the same group and recognize social similarity to the individual. One example is more accurate identification of emotion in that individual. Our previous studies proposed that rats recognize social similarity to certain strains of unfamiliar rats. It is therefore possible that the strain of unfamiliar conspecifics affects stress identification in rats. Wistar subject rats were allowed to explore a pair of unfamiliar Wistar, Sprague-Dawley (SD), Long-Evans (LE), or Fischer344 (F344) stimulus rats. To induce differences in stress, one of the stimulus rats had received foot shocks immediately before the test. It was found that the subjects showed biased interaction towards the shocked Wistar and SD stimulus rats, but not toward the shocked LE or F344 stimulus rats. Subsequent experiments confirmed that the biased interaction towards the shocked Wistar and SD stimulus rats was driven by stress in these stimulus rats. In addition, the lack of biased interaction towards the shocked LE and F344 stimulus rats did not appear to be due to procedural reasons. The experiment using LE subject rats further confirmed that the shocked LE stimulus rats emitted distress signals. These results suggested that Wistar rats could identify stress in unfamiliar Wistar and SD rats, but not in unfamiliar LE or F344 rats. Therefore, rats appear to recognize social similarity to certain unfamiliar strains of rats.

Pretreatment with CP-154526 blocks the modifying effects of alarm pheromone on components of sexual behavior in male, but not in female, rats.

We previously demonstrated that an alarm pheromone released from male donor Wistar rats evoked several physiological and behavioral responses in recipient rats. However, the pheromone effects on social behavior were not analyzed. In the present study, we examined whether the alarm pheromone affects sexual behavior in male or female rats. When a pair of male and female subjects was exposed to the alarm pheromone during sexual behavior, the ejaculation latency was elongated, the number of mounts was increased, and the hit rate (number of intromissions/number of mounts and intromissions) was decreased in the male subject. In contrast, female sexual behavior was not affected by the alarm pheromone. When we exposed only the male or female subject of the pair to the pheromone just before sexual behavior, the results were similar: the pheromone effects were evident in male, but not in female, subjects. In addition, when we pretreated with corticotropin-releasing factor (CRF) antagonist (CP-154526) before exposing the male subject to the alarm pheromone, the pheromone effects were attenuated in a dose-dependent manner. These results indicate that the alarm pheromone modifies male, but not female, components of sexual behavior and that CRF participates in the effects.

Rats do not consider all unfamiliar strains to be equivalent.

Humans show distinct social behaviours when we recognise social similarity in opponents that are members of the same social group. However, little attention has been paid to the role of social similarity in non-human animals. In the Wistar subject rats, the presence of an unfamiliar Wistar rat mitigated stress responses, suggesting the importance of social similarity in this stress-buffering phenomenon. We subsequently found that the presence of unfamiliar Sprague-Dawley (SD) or Long-Evans (LE) rats, but not an unfamiliar Fischer 344 (F344) rat, similarly mitigated stress responses in the subject rats. It is therefore possible that the subject rats recognised social similarity to unfamiliar SD and LE rats. In this study, we demonstrated that the Wistar subject rats were capable of categorizing unfamiliar rats based on their strain, and that the Wistar subjects showed a preference for unfamiliar Wistar, SD, and LE rats over F344 rats. However, the subject rats did not show a preference among Wistar, SD, and LE rats. In addition, the results were not due to an aversion to F344 rats, and preference was not affected when anaesthetised rats were presented to the subject rats. The findings suggested that rats recognise social similarity to certain unfamiliar strains of rats.

Existence of wild brown rats (Rattus norvegicus) that are indifferent to novel objects.

Exposure to novel objects typically evokes avoidance behavior in wild animals, which is called neophobia. We previously found that wild brown rats (Rattus norvegicus) that were trapped in a park in downtown Tokyo, Japan, exhibited neophobia. We also found that this behavior was accompanied by the activation of the basolateral complex of the amygdala (BLA). Previous studies have suggested that genetic factors are the primary determinants of neophobia. Since rats in cities form populations with distinct genetic characteristics, it is reasonable to assume that wild rats caught at different locations in urban centers will exhibit different levels of neophobia. Here we assessed the intensity of neophobia in wild rats trapped at a wholesale market in Tokyo. Although we performed exactly the same experiment in which neophobia was observed in wild rats trapped at the park, the presence of novel objects did not affect the behaviors of wild rats trapped at the market. Conversely, laboratory rats showed approach and exploratory behaviors as seen in the previous study, suggesting that the experiment was performed appropriately. Compared to the laboratory rats, the lack of behavioral changes in the wild rats was accompanied by fewer Fos immunoreactive cells in the BLA. In addition, the numbers of Fos immunoreactive cells in the bed nucleus of the stria terminalis and ventromedial hypothalamus were similar between the two types of rats. The results demonstrated the existence of wild rats that were indifferent to novel objects.

Prevalence of 17 feline behavioral problems and relevant factors of each behavior in Japan.

Undesirable behaviors exhibited by cats are regarded as feline behavioral problems. Like canine behavioral problems, feline behavioral problems are commonly reported by owners. Thus, in order to advise cat owners appropriately, veterinarians, particularly general veterinary practitioners, are encouraged to have sufficient knowledge of these problems. However, in ordinary Japanese homes, only a small number of feline behavioral problems have been investigated in terms of prevalence and relevant factors. Comprehensive survey data may improve understanding of feline behavioral problems. In the present study, we distributed an online survey including general information and questions about 17 feline behaviors which can be problematic for owners, to investigate: (1) the prevalence of feline behavioral problems, (2) the most common behavioral problems, and (3) relevant factors for each behavior. We collected 1,376 valid responses. Among all cats, 75.7% exhibited at least one behavioral problem. "Pica" and "Showing fear while on the examination table" were the most frequently reported behavioral problems. To identify factors related to the expression of each behavior, we conducted multiple logistic regression analyses, with the presence of behaviors as objective variables and cats' general information as explanatory variables. Age, sex, breed, number of cats in the household, and type of residence were each associated with more than one behavior. The present study revealed the prevalence of 17 feline behavioral problems in Japanese homes and relevant factors of each behavior. These findings will help veterinarians understand and address feline behavioral problems.