Prosocial behaviors in rodents.

Prosocial behaviors (i.e., actions that benefit others) are central for social interactions in humans and other animals, by fostering social bonding and cohesion. To study prosociality in rodents, scientists have developed behavioral paradigms where animals can display actions that benefit conspecifics in distress or need. These paradigms have provided insights into the role of social interactions and transfer of emotional states in the expression of prosociality, and increased knowledge of its neural bases. However, prosociality levels are variable: not all tested animals are prosocial. Such variation has been linked to differences in animals' ability to process another's state as well as to contextual factors. Moreover, evidence suggests that prosocial behaviors involve the orchestrated activity of multiple brain regions and neuromodulators. This review aims to synthesize findings across paradigms both at the level of behavior and neural mechanisms. Growing evidence confirms that these processes can be studied in rodents, and intense research in the past years is rapidly advancing our knowledge. We discuss a strong bias in the field towards the study of these processes in negative valence contexts (e.g., pain, fear, stress), which should be taken as an opportunity to open new venues for future research.

The expression of empathy in human's closest relatives, bonobos and chimpanzees: current and future directions.

Empathy is a complex, multi-dimensional capacity that facilitates the sharing and understanding of others' emotions. As our closest living relatives, bonobos (Pan paniscus) and chimpanzees (P. troglodytes) provide an opportunity to explore the origins of hominin social cognition, including empathy. Despite certain assumptions that bonobos and chimpanzees may differ empathically, these species appear to overlap considerably in certain socio-emotional responses related to empathy. However, few studies have systematically tested for species variation in Pan empathic or socio-emotional tendencies. To address this, we synthesise the growing literature on Pan empathy to inform our understanding of the selection pressures that may underlie the evolution of hominin empathy, and its expression in our last common ancestor. As bonobos and chimpanzees show overlaps in their expression of complex socio-emotional phenomena such as empathy, we propose that group comparisons may be as or more meaningful than species comparisons when it comes to understanding the evolutionary pressures for such behaviour. Furthermore, key differences, such as how humans and Pan communicate, appear to distinguish how we experience empathy compared to our closest living relatives.

A study on the influence of community spiritual comfort service on the mental health of older people.

Background: China is experiencing rapid population aging, with the proportion of older adult people aged 60 and above reaching 19. 8% of the total population in 2022. With the growth of age, the physical function of older adults declines and their mental health is getting worse, with the increasing trend of empty nesting and childlessness, older adults lack information and social interaction with others and fall into social isolation, loneliness and some mental health problems, the proportion of older adults with mental health problems gradually rises and the mortality rate increases incrementally, requiring effective ways to intervene in the mental health of older adults and promote healthy aging. Aim of the study: The present study investigated the influence of spiritual comfort older adult services on the mental health of 12,624 older adults aged 60 years or older in 23 provinces in China from 2017 to 2018, with the aim of providing a case for the development of more targeted mental health strategies for older people. Methods: Using the data from the 2018 CLHLS Survey, the influencing factors of the mental health of older people were analyzed using chi-square test and the logit regression model. The mechanism of the health care facilities and the spiritual comfort services on mental health was analyzed using the chain mediation effect. Results: The spiritual comfort services decreased the risk of negative emotion and mental health of older adults, with female (OR = 1.168), rural residents (OR = 1.385), no drinking (OR = 1.255), not exercising (OR = 1.543), not having pension insurance (OR = 1.233), and low annual household income (OR = 1.416) being the characteristics as risk factors. The mediating effect results showed that the health care facilities had a partial mediating effect between the spiritual comfort services and the mental health status of older people, and the mediating effect accounted for 40.16% of the total effect. Conclusions: The use of spiritual comfort services can effectively reduce and alleviate the adverse mental health symptoms of older people, promote guidance and health education for healthy older people and chronically ill patients, and improve the good health perception of older people in order to improve their quality of life and mental health status.

Dorsomedial prefrontal hypoexcitability underlies lost empathy in frontotemporal dementia.

Empathic function is essential for the well-being of social species. Empathy loss is associated with various brain disorders and represents arguably the most distressing feature of frontotemporal dementia (FTD), a leading form of presenile dementia. The neural mechanisms are unknown. We established an FTD mouse model deficient in empathy and observed that aged somatic transgenic mice expressing GGGGCC repeat expansions in C9orf72, a common genetic cause of FTD, exhibited blunted affect sharing and failed to console distressed conspecifics by affiliative contact. Distress-induced consoling behavior activated the dorsomedial prefrontal cortex (dmPFC), which developed profound pyramidal neuron hypoexcitability in aged mutant mice. Optogenetic dmPFC inhibition attenuated affect sharing and other-directed consolation in wild-type mice, whereas chemogenetically enhancing dmPFC excitability rescued empathy deficits in mutant mice, even at advanced ages when substantial cortical atrophy had occurred. These results establish cortical hypoexcitability as a pathophysiological basis of empathy loss in FTD and suggest a therapeutic strategy.

Consolation Without Religious Topoi: Thematic Analysis of References to "Afterlife" in Secular Eulogies in Communist Czechoslovakia (1948-1989).

The article focuses on practice of secular eulogistic rhetoric in communist Czechoslovakia (1948-1989), specifically it investigates references to "afterlife" used in eulogies, which is generally conceived as allowing the bereaved to positively reappraise the death of the loved one. However, in secular eulogies, the use of "afterlife" in religious interpretation ("resurrection", "heaven") is explicitly excluded. Based on the thematic analysis of 61 secular eulogies, 2 secular interpretations of "afterlife" are identified: symbolic continuity (continuity of memory of the deceased, through his/her legacy/work done, as a part of Nature, through children/future generations, or continuity of the deceased's love) and factual (but deprived) continuity (death is presented as a sleep, journey, or pain relief).

Flexible signalling strategies by victims mediate post-conflict interactions in bonobos.

Compared to other animals, humans supposedly excel at voluntarily controlling and strategically displaying emotional signals. Yet, new data shows that nonhuman great apes' emotion expressions may also be subject to voluntary control. A key context to further explore this is during post-conflict (PC) periods, where signalling by distressed victims may influence bystander responses, including the offering of consolation. To address this, our study investigates the signalling behaviour of sanctuary-living bonobo victims following aggression and its relation to audience composition and PC interactions. Results show that the production of paedomorphic signals by victims (regardless of age) increased their chances of receiving consolation. In adults, the production of such signals additionally reduced the risk of renewed aggression from opponents. Signal production also increased with audience size, yet strategies differed by age: while immatures reduced signalling in proximity of close-social partners, adults did so especially after receiving consolation. These results suggest that bonobos can flexibly adjust their emotion signalling to influence the outcome of PC events, and that this tendency has a developmental trajectory. Overall, these findings highlight the potential role that flexible emotion communication played in the sociality of our last common ancestor with Pan. This article is part of the theme issue 'Cognition, communication and social bonds in primates'.

Mice Cohabiting With Familiar Conspecific in Chronic Stress Condition Exhibit Methamphetamine-Induced Locomotor Sensitization and Augmented Consolation Behavior.

Recognizing and sharing emotions are essential for species survival, but in some cases, living with a conspecific in distress condition may induce negative emotional states through empathy-like processes. Studies have reported that stressors promote psychiatric disorders in both, those who suffer directly and who witness these aversive episodes, principally whether social proximity is involved. However, the mechanisms underlying the harmful outcomes of emotional contagion need more studies, mainly in the drug addiction-related behaviors. Here, we investigated the relevance of familiarity and the effects of cohabitation with a partner submitted to chronic stress in the anxiety-like, locomotor sensitization, and consolation behaviors. Male Swiss mice were housed in pairs during different periods to test the establishment of familiarity and the stress-induced anxiety behavior in the elevated plus maze. Another cohort was housed with a conspecific subjected to repeated restraint stress (1 h/day) for 14 days. During chronic restraint the allogrooming was measured and after the stress period mice were tested in the open field for evaluation of anxiety and locomotor cross-sensitization induced by methamphetamine. We found that familiarity was established after 14 days of cohabitation and the anxiogenic behavior appeared after 14 days of stress. Repeated restraint stress also increased anxiety in the open field test and induced locomotor cross-sensitization in the stressed mice and their cagemates. Cagemates also exhibited an increase in the consolation behavior after stress sessions when compared to control mice. These results indicate that changes in drug abuse-related, consolation, and affective behaviors may be precipitated through emotional contagion in familiar conspecifics.

Consoling the Bereaved: Exploring How Sympathy Cards Influence What People Say.

In a sequential mixed methods study, interview data showed that bereaved parents consistently reported "stupid" or "insensitive" things people said that were not helpful when their child died. Subsequently, a discourse analysis of 170 sympathy cards was conducted to assess societal messaging that may influence people's insensitive words. The findings reflected two distinct time periods in the bereavement process: a time of sadness followed by a time of peace. Possible links to insensitive things people said included: suggestions that the sad time should only last a few days, suggestions of "healing," religious statements such as the deceased being with God or advice to pray to God, and notions of celebration in some contexts. Very few excerpts were similar to the advice-giving quotes that interviewees considered to be insensitive. Many cards reflected the love and memories inherent in creating a continuing bond with the deceased and achieving peace.

Music as Consolation-The Importance of Music at Farewells and Mourning.

In this study we examined a case where twelve participants conducted farewell ceremonies for their deceased relatives. Taking a qualitative approach, we used interviews and questionnaires to focus on life stories that involve grieving. Specifically, we asked about experiences of the grief process as related to choice of music. Our results indicate that the role of music in farewell rituals is important for the grieving process in several ways. Firstly, music was associated with positive memories of the loved one and gave rise to experiences of recognition. Secondly, music facilitated active participation in the grieving process through choosing farewell music together with a relative. Thirdly, selecting music for the funeral in advance, together with their loved ones, was also experienced as hopeful, comforting and consoling before, during and after the bereavement.

Monkey see, monkey feel? Marmoset reactions towards conspecifics' arousal.

Consolation has been observed in several species, including marmoset monkeys, but it is often unclear to what extent they are empathy-based. Marmosets perform well in at least two of three components of empathy-based consolation, namely understanding others and prosociality, but it is unknown to what extent they show matching with others. We, therefore, tested whether non-aroused individuals would become aroused themselves when encountering an aroused group member (indicated by piloerection of the tail). We found a robust contagion effect: group members were more likely to show piloerection themselves after having encountered an aroused versus relaxed conspecific. Moreover, group members offered consolation behaviours (affiliative approaches) towards the aroused fellow group members rather than the latter requesting it. Importantly, this pattern was shown by both aroused and non-aroused individuals, which suggests that they did not do this to reduce their own arousal but rather to console the individual in distress. We conclude that marmosets have all three components of empathy-based consolation. These results are in line with observations in another cooperative breeder, the prairie vole.

Examining Empathy Through Consolation Behavior in Prairie Voles.

Empathy is an affective and cognitive event in which an organism experiences an approximation of the physical or psychological state of another organism. The phenomenon has been well-studied in humans but is not as widely researched in other animals. Burkett and colleagues in a 2016 article published in Science measured empathy in prairie voles (Microtus ochrogaster) and meadow voles (Microtus pennsylvanicus) by observing consolation behavior between non-stressed and stressed individuals. Their data from behavioral analyses and histochemistry support their hypothesis that consolation behavior in prairie voles shares similar behavioral characteristics and conserved biological mechanisms with human empathy. Prairie voles match anxiety and fear states as well as groom stressed familiar conspecifics to lessen their stress. An oxytocin receptor antagonist abolished this empathetic response. This research impacted the field of neuroscience by demonstrating human-like empathy in rodents, and thereby supporting the value of animal models to investigations of higher order human experiences. The paper is also a valuable and accessible resource to undergraduate neuroscience students-from introductory courses to advanced seminars. In the classroom, this research provides a foundational look at the expanding field of social neuroscience. Empathy in prairie voles raises thought-provoking discussion concerning emotions, social behavior, and human nature.

CRF-CRFR1 system within the dorsal medial prefrontal cortex are involved in consolation deficits under acute restraint stress in mandarin voles.

Consolation is a complex empathic behavior that has recently been observed in some socially living rodents. Despite the growing body of literature suggesting that stress affects some simple form of empathy, the relationship between stress and consolation remains largely understudied. Using monogamous mandarin voles, we found that an acute restraint stress exposure significantly reduced consolation-like behaviors and induced anxiety-like behaviors. Along with these behavioral changes, corticotropin-releasing factor (CRF) and CRF receptor 1 (CRFR1) neurons were activated within the anterior cingulate cortex (ACC) and prelimbic cortex (PrL) but not within the infralimbic cortex (IL). Chemogenetic activation of CRF neurons in the ACC and PrL, recaptured acute stress-induced behavioral dysfunctions. We further observed that intracellular PKA and PKC signaling pathways mediate CRF-induced behavioral dysfunctions, but they work in a regional-specific, sex-biased manner. Together, these results suggest that the local CRF-CRFR1 system within the ACC and PrL is involved in the consolation deficits and anxiety induced by acute stress.

Wild jackdaws respond to their partner's distress, but not with consolation.

Individuals are expected to manage their social relationships to maximize fitness returns. For example, reports of some mammals and birds offering unsolicited affiliation to distressed social partners (commonly termed 'consolation') are argued to illustrate convergent evolution of prosocial traits across divergent taxa. However, most studies cannot discriminate between consolation and alternative explanations such as self-soothing. Crucially, no study that controls for key confounds has examined consolation in the wild, where individuals face more complex and dangerous environments than in captivity. Controlling for common confounds, we find that male jackdaws (Corvus monedula) respond to their mate's stress-states, but not with consolation. Instead, they tended to decrease affiliation and partner visit rate in both experimental and natural contexts. This is striking because jackdaws have long-term monogamous relationships with highly interdependent fitness outcomes, which is precisely where theory predicts consolation should occur. Our findings challenge common conceptions about where consolation should evolve, and chime with concerns that current theory may be influenced by anthropomorphic expectations of how social relationships should be managed. To further our understanding of the evolution of such traits, we highlight the need for our current predictive frameworks to incorporate the behavioural trade-offs inherent to life in the wild.

Dorsal raphe nucleus to anterior cingulate cortex 5-HTergic neural circuit modulates consolation and sociability.

Consolation is a common response to the distress of others in humans and some social animals, but the neural mechanisms underlying this behavior are not well characterized. By using socially monogamous mandarin voles, we found that optogenetic or chemogenetic inhibition of 5-HTergic neurons in the dorsal raphe nucleus (DR) or optogenetic inhibition of serotonin (5-HT) terminals in the anterior cingulate cortex (ACC) significantly decreased allogrooming time in the consolation test and reduced sociability in the three-chamber test. The release of 5-HT within the ACC and the activity of DR neurons were significantly increased during allogrooming, sniffing, and social approaching. Finally, we found that the activation of 5-HT1A receptors in the ACC was sufficient to reverse consolation and sociability deficits induced by the chemogenetic inhibition of 5-HTergic neurons in the DR. Our study provided the first direct evidence that DR-ACC 5-HTergic neural circuit is implicated in consolation-like behaviors and sociability.

Act of Consoling Helps Those Who Console: A Multicenter Observational Survey Study in Hospice Services.

Background: The contemporary scientific literature documents a lack of attention toward the act of consoling put into practice by health care professionals (HCPs) in hospice services. Objective: To describe the act of consoling and its meaning for hospice-employed HCPs. Design: A multicenter observational survey study was performed through a paper-based questionnaire. Subjects: The study was sent to the directors of 10 hospices in Northern Italy and distributed to 232 HCPs. Results: A total of 218 HCPs responded (94%). The results showed that most HCPs consider the practice of consoling to be essential to their profession, but they also underscored the extreme complexity of the process. Conclusion: The act of consoling is not simply a professional duty defined in contractual clauses. Rather, it is a set of specific communicative practices and skills required of HCPs.

Paraventricular thalamic nucleus plays a critical role in consolation and anxious behaviors of familiar observers exposed to surgery mice.

Background: Consolation behaviors toward the sick are common in humans. Anxiety in the relatives of the sick is also common. Anxiety can cause detrimental effects on multiple systems. However, our understanding on the neural mechanisms of these behaviors is limited because of the lack of small animal models. Methods: Five of 6- to 8-week-old CD-1 male mice were housed in a cage. Among them, 2 mice had right common artery exposure (surgery) and the rest were without surgery. Allo-grooming and performance in light and dark box and elevated plus maze tests of the mice were determined. Results: Mice without surgery had increased allo-grooming toward mice with surgery but decreased allo-grooming toward non-surgery intruders. This increased allo-grooming toward surgery mice was higher in familiar observers of surgery mice than that of mice that were not cage-mates of surgery mice before the surgery. Familiar observers developed anxious behavior after being with surgery mice. Surgery mice with familiar observers had less anxious behavior than surgery mice without interacting with familiar observers. Multiple brain regions including paraventricular thalamic nucleus (PVT) were activated in familiar observers. The activated cells in PVT contained orexin receptors. Injuring the neurons with ibotenic acid, antagonizing orexin signaling with an anti-orexin antibody or inhibiting neurons by chemogenetic approach in PVT abolished the consolation and anxious behaviors of familiar observers. Conclusions: Mice show consolation behavior toward the sick. This behavior attenuates the anxious behavior of surgery mice. The orexin signaling in the PVT neurons play a critical role in the consolation of familiar observers toward surgery mice and their anxious behavior. Considering that about 50 million patients have surgery annually in the United States, our study represents the initial attempt to understand neural mechanisms for consolation and anxiety of a large number of people.

Third-party prosocial behavior in adult female rats is impaired after perinatal fluoxetine exposure.

SSRIs are commonly used to treat pregnant women with depression. However, SSRIs can cross the placenta and affect the development of the fetus. The effects of perinatal SSRI exposure, and especially the effects on social behavior, are still incompletely documented. This study first aims to investigate whether rats show prosocial behavior in the form of consolation behavior. Secondly, it aims to investigate whether perinatal SSRI exposure affects this prosocial behavior. At last, we investigate whether the behavior changed after the rats had been exposed to an additional white-noise stressor. Rat dams received 10 mg/kg/d fluoxetine (FLX) or vehicle (CTR) via oral gavage from gestational day 1 until postnatal day 21. At adulthood, the rat offspring were housed in four cohorts of 4 females and 4 males in a seminatural environment. As prosocial behaviors are more prominent after stressful situations, we investigated the behavioral response of rats immediately after natural aggressive encounters (fights). Additionally, we studied whether a stressful white-noise exposure would alter this response to the aggressive encounters. Our study indicates that CTR-female rats are able to show third party prosocial behavior in response to witnessing aggressive encounters between conspecifics in a seminatural environment. In addition, we showed that perinatal FLX exposure impairs the display of prosocial behavior in female rats. Moreover, we found no signs of prosocial behavior in CTR- and FLX-males after natural aggressive encounters. After white-noise exposure the effects in third party prosocial behavior of CTR-females ceased to exist. We conclude that female rats are able to show prosocial behavior, possibly in the form of consolation behavior. In addition, the negative effects of perinatal fluoxetine exposure on prosocial behavior could provide additional evidence that SSRI treatment during pregnancy could contribute to the risk for social impairments in the offspring.

The similar past pain experience evokes both observational contagious pain and consolation in stranger rat observers.

Laboratory rodents have been shown to have an ability to recognize the injury site and negative emotional state of their conspecifics in pain, resulting in empathic consoling behaviors and observational contagious pain (OCP). However, these empathic responses have been shown to be familiarity-dependent. In this report, we further explored whether the past pain experience could evoke empathic response in stranger observers. In our rodent model, two types of empathic response have been identified from naive cagemate observer (COnaive) during and after a priming dyadic social interaction (PDSI) with a cagemate demonstrator in pain (CDpain): the consolation and OCP. Consolation is represented by allolicking and allogrooming behaviors toward the CDpain, while the OCP is represented by a long-term mechanical pain hypersensitivity. The current results showed that: (1) neither the consolation nor OCP could be identified in the naive noncagemate observer (NCOnaive) during and after a PDSI with a noncagemate demonstrator in pain (NCDpain); (2) nor were the two types of empathic response seen in the NCO, who had just experienced acute pain (NCOpainexp), during and after a PDSI with a naive unfamiliar conspecific (NCDnaive). However, both the consolation and OCP were dramatically identified in the NCOpainexp during and after a PDSI with a NCD in pain (NCDpain). The current results demonstrated that the past pain experience can evoke both consolation and OCP in stranger rat observers when witnessing a conspecific in pain, implicating that the processing of empathy for pain can be modulated by past negative mood experience.

Challenges in the comparative study of empathy and related phenomena in animals.

The aim of this review is to discuss recent arguments and findings in the comparative study of empathy. Based on a multidisciplinary approach including psychology and ethology, we review the non-human animal literature concerning theoretical frameworks, methodology, and research outcomes. One specific objective is to highlight discrepancies between theory and empirical findings, and to discuss ambiguities present in current data and their interpretation. In particular, we focus on emotional contagion and its experimental investigation, and on consolation and targeted helping as measures for sympathy. Additionally, we address the feasibility of comparing across species with behavioural data alone. One main conclusion of our review is that animal research on empathy still faces the challenge of closing the gap between theoretical concepts and empirical evidence. To advance our knowledge, we propose to focus more on the emotional basis of empathy, rather than on possibly ambiguous behavioural indicators, and we provide suggestions to overcome the limitations of previous research .

Do chimpanzees (Pan troglodytes) console a bereaved mother?

Comparative thanatology encompasses the study of death-related responses in non-human animals and aspires to elucidate the evolutionary origins of human behavior in the context of death. Many reports have revealed that humans are not the only species affected by the death of group members. Non-human primates in particular show behaviors such as congregating around the deceased, carrying the corpse for prolonged periods of time (predominantly mothers carrying dead infants), and inspecting the corpse for signs of life. Here, we extend the focus on death-related responses in non-human animals by exploring whether chimpanzees are inclined to console the bereaved: the individual(s) most closely associated with the deceased. We report a case in which a chimpanzee (Pan troglodytes) mother experienced the loss of her fully developed infant (presumed stillborn). Using observational data to compare the group members' behavior before and after the death, we found that a substantial number of group members selectively increased their affiliative expressions toward the bereaved mother. Moreover, on the day of the death, we observed heightened expressions of species-typical reassurance behaviors toward the bereaved mother. After ruling out several alternative explanations, we propose that many of the chimpanzees consoled the bereaved mother by means of affiliative and selective empathetic expressions.