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Consolation, Reconciliation, and a Possible Cognitive Difference between Macaques and Chimpanzees
Abstract
The authors introduce two main hypotheses to account for variation in postconflict behaviour, and discuss the social background against which this variation needs to be evaluated. This is followed by a review of available data on postconflict behaviour itself, and the pros and cons of each hypothesis. The reconciliation behaviour of macaques Macaca and chimpanzees Pan troglodytes differs only in degree, with a higher level of sophistication in the chimpanzee, but not necessarily a greater tendency to reconcile or a fundamentally different motivation. In both macaques and chimpanzees, reconciliation seems to serve the restoration of valuable relationships and the reduction of tension. The authors conclude that chimpanzees and macaques differ substantially in their response to individuals distressed by previous aggression, and that it is possible yet unclear at this point whether this difference is part of a suite of differences related to species-typical levels of cognition and empathic capacity.
... La riconciliazione non rappresenta l'unica strategia post-conflittuale che i membri di un gruppo di primati hanno a disposizione per risolvere un conflitto. Si possono, infatti, verificare contatti affiliativi post-conflittuali che hanno come target la vittima e/o l'aggressore (de Waal & van Roosmalen, 1979;de Waal, 1992;de Waal & Aureli, 1996;Cords, 1997;Das, 2000;Arnold & Whiten, 2001;Call et al., 2002;Wittig & Boesch, 2003;Palagi et al., 2005). I contatti triadici possono essere sollecitati o meno dalla vittima o dall'aggressore (Verbeek & de Waal, 1997). ...
... Con il termine consolazione e con quello di appeasement, si definiscono tutti quei contatti affiliativi offerti spontaneamente da un terzo individuo, non coinvolto in un precedente conflitto, rispettivamente alla vittima e/o all'aggressore (de Waal & van Roosmalen, 1979;de Waal, 1992;de Waal & Aureli, 1996;Das, 2000). L'empatia, ovvero la capacità cognitiva di percepire il disagio emotivo di un conspecifico, diviene un pre-requisito necessario per mettere in atto questo tipo di strategie post-conflittuali. ...
... La capacità empatica di percepire un disagio emotivo di un conspecifico è risultata essere un fattore determinante per la risposta prosociale ad un soggetto in difficoltà (Caplan & Hay, 1989;Aureli & de Waal, 1996;Eisenberg & Fabes, 1998;Fujisawa et al., 2006). Nel 1996 Aureli e de Waal formularono l'ipotesi della Social Cognition, secondo la quale l'empatia rappresenta un prerequisito indispensabile affinché il soggetto possa mettere in atto un contatto affiliativo post-conflittuale, specialmente di tipo consolatorio. ...
Come per le altre specie sociali, anche nell’uomo la vita di gruppo rappresenta un delicato equilibrio tra cooperazione e competizione. In particolar modo, la competizione per le risorse limitate (es. cibo, partner sessuali) può portare ad un forte “conflitto di interessi” che, in taluni casi, implica una reazione conflittuale se non addirittura aggressiva. Per ridurre le conseguenze negative di un conflitto, molte specie sociali hanno sviluppato delle strategie comportamentali post-conflittuali: la riconciliazione e i contatti triadici sollecitati e non sollecitati tra vittima o aggressore e terzi individui estranei al conflitto. Lo scopo di questo studio è stato quello investigare le dinamiche di gestione dei conflitti in bambini in età pre-scolare (3, 4, 5 anni) applicando un approccio prettamente etologico usato comunemente negli studi sui primati non-umani (metodo Post-Conflict/Matched-Control, PC-MC). Oltre alla quantificazione delle varie tipologie di strategie post-conflittuali (riconciliazione e consolazione), abbiamo anche tentato di verificare quali potessero essere le variabili implicate (età dei soggetti, sesso, legami affiliativi tra i bambini) nella distribuzione dei fenomeni. La ricerca è stata condotta su 80 bambini della scuola dell'infanzia "Florinda" dell’Istituto Comprensivo “Centro-Migliarina-Motto” (Viareggio, Lucca, Italia). La raccolta dati, effettuata utilizzando la tecnica della ripresa filmata, ha permesso una videoanalisi estremamente accurata di ogni evento agonistico e dell'eventuale meccanismo post-conflittuale (la finestra temporale utilizzate era di 10 minuti in accordo con la letteratura relativa ad altri studi su Homo sapiens e su grandi antropomorfe). I risultati, seppur preliminari, indicano la presenza di riconciliazione e consolazione spontanea nel nostro campione. La riconciliazione è influenzata dalla qualità della relazione che lega i soggetti anche se non sembra avere un ruolo nella riduzione dello stato d'ansia della vittima. Riconciliazione e consolazione sono due fenomeni ugualmente rappresentati. Come accade nei primati non umani, l'evento consolatorio sembra entrare in gioco quando la riconciliazione è assente, ciò indica che anche in Homo sapiens il contatto affiliativo spontaneo da parte di un terzo soggetto rivolto alla vittima agisce come sostituto dell'atto conciliatorio. La consolazione sembra inoltre essere dipendente dall'età e contestualmente dal sesso dei soggetti che entrano in gioco. Questo risultato merita sicuramente attenzione data la natura empatica del fenomeno consolatorio. La maturazione della sfera empatica è legata fortemente al processo di sviluppo del bambino che potrebbe quindi essere maggiormente coinvolto una volta maturata la competenza emotiva e sociale necessarie non solo per percepire l'ansia provata dal soggetto che ha appena subito un'aggressione, ma anche per mettere in atto quelle azioni atte a migliorarne lo stato emozionale (consolazione).
... Great apes generally outperform monkeys in cognitive tasks involving inferential reasoning, planning for the future and attributing perceptual or mental states to others (Call 2004;Mulcahy & Call 2006;Call & Tomasello 2008;Shettleworth 2010;Bourjade et al. 2012;Crockford et al. 2012;Anderson & Gallup 2015;Costes-Thiré et al. 2015;Marsh et al. 2015;Petit et al. 2015). As a corollary, great apes appear to be more sensitive to the attention and needs of others, and show more empathy in the form of consolation, targeted helping, and concern for injured partners (Cheney & Seyfarth 1990de Waal 1996de Waal , 2008de Waal & Aureli 1996;Silk 2007;Boesch 2012;Pérez-Manrique & Gomila 2018;Sato et al. 2019). ...
... Great apes generally outperform monkeys in cognitive tasks involving inferential reasoning, planning for the future and attributing perceptual or mental states to others (Call 2004;Mulcahy & Call 2006;Call & Tomasello 2008;Shettleworth 2010;Bourjade et al. 2012;Crockford et al. 2012;Anderson & Gallup 2015;Costes-Thiré et al. 2015;Marsh et al. 2015;Petit et al. 2015). As a corollary, great apes appear to be more sensitive to the attention and needs of others, and show more empathy in the form of consolation, targeted helping, and concern for injured partners (Cheney & Seyfarth 1990de Waal 1996de Waal , 2008de Waal & Aureli 1996;Silk 2007;Boesch 2012;Pérez-Manrique & Gomila 2018;Sato et al. 2019). ...
... To investigate what primates may understand about death, we will examine the behavioural events surrounding helpless individuals. We will not consider responses to sick, injured or disabled individuals who can still stand upright and/or communicate, nor the delayed effects of partner loss, as such occurrences cannot be equated with the behaviours induced by the fainting or death of a conspecific (for review see : Zeller 1991;de Waal 1996;King 2013;Anderson 2018;Gonçalves & Carvalho 2019;Sato et al. 2019). No conclusive evidence is available regarding the possible adaptive function of death-related behaviours (Piel & Stewart 2016;Watson & Matsuzawa 2018; Gonçalves & Carvalho 2019; Monsó 2020); they may not be adaptive, and could reflect the importance of social bonding in primate evolution (Bercovitch 2020). ...
It was long assumed that only humans can distinguish the living from the dead. Renewed interest in this question over the last decade has led several authors to assert that non-human primates are also aware of death. We investigate this issue by comparing the behaviours of monkeys and great apes toward helpless conspecifics, basing our analysis on published reports. We first examine the behaviours of mothers towards the body of their dead offspring. They may carry the corpse for days or more before abandoning it. They groom, inspect and protect it, sometimes allowing group members to explore it, and rare cases of cannibalism have been reported. No significant difference is observed in the way that monkeys and great apes treat the bodies of infants. We then examine responses to collapsed (still able to move and react) and inanimate (unresponsive or dead) conspecifics. Monkeys and great apes guard, care for and inspect their helpless partners, and also manipulate and mobilise them. Through these actions, individuals may inform themselves about the state of their partners, test their responsiveness and/or attempt to rouse them. It appears that only chimpanzees and gorillas show violent action such as display behaviours and the rough treatment of bodies. They can also make distress calls, and periods of “stunned silence” sometimes occur in chimpanzees, indicating that they are experiencing intense emotion. Finally, we argue that while both monkeys and great apes detect body dysfunction through the victims’ inability to wake up and move, only great apes can understand that something serious has happened. The signs of emotional disturbance reported in them indicate that they may believe that inanimate conspecifics have entered a state of “dormancy”, meaning that they are unlikely to regain wakefulness. However, there is no evidence that any non-human primates are aware of mortality.
... A central question in the literature on emotion processing in primates is whether there is a division between apes and monkeys with respect to their understanding of others' emotions (Clay et al., 2018). Based on observations of third party post-conflict affiliation (or 'consolation'), observed in apes but not in monkeys, de Waal and Aureli (1996) argued that monkeys are not capable of cognitive empathy. Anecdotes of prosocial behaviour in apes, such as a chimpanzee rescuing an unrelated infant chimpanzee from the water (Goodall, 1986) and a gorilla rescuing a 3-year-old boy who had fallen into her zoo enclosure (de Waal, 1997) served as additional support for this proposal. ...
... Possible clarifications in the debate on cognitive empathy in apes and monkeys can be provided if we include research in the vocal domain, which is often neglected in the present hypotheses (de Waal and Aureli, 1996;Gruber and Clay, 2016;de Waal and Preston, 2017;Koski and Sterck, 2010;Kret et al., 2020). Primates often convey their emotional state using different calls, which has the advantage of reaching others over larger distances and in forested habitats. ...
... We discussed evidence of apes, Old World monkeys and New World monkeys being able to relate the meaning of their conspecifics' expressions to external concepts such as food and future events. The literature shows that the prerequisites for cognitive empathy are present in different monkey species, in contrast to previous beliefs (de Waal and Aureli, 1996). Possibly, the ability to attribute meaning to others' emotional expressions provides such strong adaptive benefits that this capacity evolved independently in different species. ...
Recognising conspecifics’ emotional expressions is important for nonhuman primates to navigate their physical and social environment. We address two possible mechanisms underlying emotion recognition: emotional contagion, the automatic matching of the observer’s emotions to the emotional state of the observed individual, and cognitive empathy, the ability to understand the meaning and cause of emotional expressions while maintaining a distinction between own and others’ emotions. We review experimental research in nonhuman primates to gain insight into the evolution of emotion recognition. Importantly, we focus on how emotional contagion and cognitive empathy can be studied experimentally. Evidence for aspects of cognitive empathy in different nonhuman primate lineages suggests that a wider range of primates than commonly assumed can infer emotional meaning from emotional expressions. Possibly, analogous rather than homologous evolution underlies emotion recognition. However, conclusions regarding its exact evolutionary course requires more research in different modalities and species.
... Scientific REPORts | (2018) 8:15291 | DOI: 10.1038/s41598-018-33548-y Third party spontaneous affiliation towards the victim has been observed in a variety of mammalian and non-mammalian species (chimpanzees, Pan troglodytes 4,[25][26][27][28][29][30][31][32] ; gorillas, Gorilla spp. [33][34][35] , bonobos, Pan paniscus 24,[36][37][38] , humans, Homo sapiens 39 , wolves 18,40 , ravens 16 , rooks 41 and horses 42 , prairie voles 19 ). ...
... In primates, consolation has been demonstrated in humans 44 , chimpanzees 31,32 (although one study did not find evidence in this species 29 ), bonobos 24,37,38 and Tonkean macaques (Macaca tonkeana) 45,46 . It is interesting to note that consolation was not found in several macaque species whose social groups are based on more despotic relationships (Macaca fascicularis, M. fuscata, M. sylvanus, and M. nemestrina 25 ). In despotic macaques, the distribution of affiliative dyadic interactions (e.g., grooming) is determined by rigid hierarchical and nepotistic rules 47 . ...
... According to the empathic gradient hypothesis 71 , subjects sharing strong affiliative bonds are more prone to engage in consolatory contacts. Moreover, the victim's anxiety would decrease because closely bonded partners (kin and friends) are supposed to be more effective in relieving anxiety, due to the good relationship quality shared with Scientific REPORts | (2018) 8:15291 | DOI:10.1038/s41598-018-33548-y the victim 25,30,64,72 . If consolation is the driving mechanism of third party affiliation in geladas, we expect that it should be biased towards kin/friends and effective in reducing victim's anxiety. ...
Post-conflict affiliation is a mechanism favored by natural selection to manage conflicts in animal groups thus avoiding group disruption. Triadic affiliation towards the victim can reduce the likelihood of redirection (benefits to third-parties) and protect and provide comfort to the victim by reducing its post-conflict anxiety (benefits to victims). Here, we test specific hypotheses on the potential functions of triadic affiliation in Theropithecus gelada, a primate species living in complex multi-level societies. Our results show that higher-ranking geladas provided more spontaneous triadic affiliation than lower-ranking subjects and that these contacts significantly reduced the likelihood of further aggression on the victim. Spontaneous triadic affiliation significantly reduced the victim’s anxiety (measured by scratching), although it was not biased towards kin or friends. In conclusion, triadic affiliation in geladas seems to be a strategy available to high-ranking subjects to reduce the social tension generated by a conflict. Although this interpretation is the most parsimonious one, it cannot be totally excluded that third parties could also be affected by the negative emotional state of the victim thus increasing a third party’s motivation to provide comfort. Therefore, the debate on the linkage between third-party affiliation and emotional contagion in monkeys remains to be resolved.
... They stated that such interactions seemed to serve an important distress alleviation function in the short term, and a longer-term social homeostasis function. Because of the short duration of the post-conflict observation period and the lack of control data, de Waal & Aureli (1996) replicated this study. They proposed that, if post-conflict affiliative contact with bystanders alleviates the distressed party, three predictions follow: (i) more affiliative contacts are expected within a few minutes of the conflict than after longer time periods or during control periods; (ii) more contacts are expected following intense aggressive encounters (given that the distress of the victim is likely to be proportional to conflict intensity); and (iii) more contacts are expected between bystanders and victims of aggression than between bystanders and aggressors. ...
... de Waal & Aureli (1996) compared their chimpanzees' results with data on post-conflict behaviour of several species of macaques in the wild and in captivity, and found important differences in unsolicited bystander affiliation. Unlike chimpanzees, affiliative contacts between macaque bystanders and victims of aggression did not occur more often following a conflict that during control periods. ...
... In Tonkean macaques, unsolicited third-party affiliation reduced indicators of anxiety (scratching) in the conflict victims and was mainly directed towards friends (measured by the amount of grooming exchanged between individuals), especially towards females who experienced higher levels of anxiety compared to males. These results support the 'Social Constraints Hypothesis' (de Waal & Aureli, 1996), which proposes that unsolicited bystander affiliation depends on the social structure of the species, being more likely to be found in tolerant than in despotic species since the risks of further aggression for potential bystanders are lower (Fraser et al., 2009). ...
While empathy is a century-old psychological concept, its study in non-human animals has become the focus of much recent scientific interest, as it promises to provide the clues to understand the evolutionary origins of our social and moral nature. A review of the comparative study of empathy is thus timely to complement and constrain anthropocentric views, and to integrate current findings. However, this is not an easy task. The study of animal empathy has developed using different paradigms, different concepts of the phenomena involved, and the absence of a systematic program. Herein, we carry out a comprehensive review of the literature on complex forms of empathy in non-human animals: sympathetic concern and empathic perspective-taking. In particular, we focus on consolation and targeted helping, as the best examples of each category. In so doing, we try to shed light on the current debate concerning whether these phenomena are exclusively human traits. First, we try to clarify the terminology and taxonomy of forms of empathy, providing operative criteria for these phenomena that are applicable to both human and non-human animals. Second, we discuss whether the available evidence qualifies such behaviour as empathic. Third, we aim to provide an integrative view of the field, clarifying the challenges and conditions to satisfy. We also hope to highlight the importance of the study of these processes for elucidating the evolutionary history of this capacity across the animal kingdom.
... lupus), hyenas (Crocuta crocuta), dolphins (Tursiops truncatus), and rooks (Corvus frugilegus) (Cools, van Hout, & Nelissen, 2007;Cozzi, Sighieri, Gazzano, Nicol, & Baragli, 2010;Palagi & Cordoni, 2009;Schino, 1998;Seed, Clayton, & Emery, 2007;Wahaj, Guse, & Holekamp, 2001;Yamamoto et al., 2015). Also, the grooming of a victim of a conflict by a bystander was labeled as an act of consolation (de Waal & Aureli, 1996), and for long it has been considered indicative of cognitive empathy of which only apes were supposed to be capable. Recently " consolation " has also been demonstrated in stump-tailed macaques (Macaca arctoides) (Call, Aureli, & De Waal, 2002), Barbary macaques (Macaca sylvanus) (McFarland & Majolo, 2012), and Tonkean macaques (Macaca tonkeana) (Palagi, Dall'Olio, Demuru, & Stanyon, 2014;Puga-Gonzalez, Butovskaya, Thierry, & Hemelrijk, 2014), dogs (Cools et al., 2007), wolves (Palagi & Cordoni, 2009), horses (Cozzi et al., 2010), rodents (Microtus ochrogaster) (Burkett et al., 2016), and rooks (Seed et al., 2007). ...
... When such post-conflict affiliation is directed by a bystander to the victim of the original conflict, it is usually referred to as " consolation " and when it is directed to the former aggressor, it is referred to as " appeasement. " As to the cognitive mechanisms involved in post-conflict affiliation, those involved in " consolation " have been supposed to be particularly sophisticated, because this social pattern has been found initially only in apes and not in monkeys (de Waal & Aureli, 1996). Scientists supposed that a bystander recognizes the victim's distress and tries to alleviate it, thus displaying " cognitive empathy. ...
... Scientists supposed that a bystander recognizes the victim's distress and tries to alleviate it, thus displaying " cognitive empathy. " The lack of occurrence of " consolation " in monkeys has been attributed to their lack of " cognitive empathy " (Preston & de Waal, 2002), also referred to as the cognitive constraint hypothesis (de Waal & Aureli, 1996). The statistical pattern of " consolation, " however, has recently been confirmed also in several species whose cognitive abilities are supposed to be less developed than those of apes, such as rodents (Burkett et al., 2016), dogs (Cools et al., 2007), wolves (Palagi & Cordoni, 2009), horses (Cozzi et al., 2010), rooks (Seed et al., 2007), and three species of monkeys (Call et al., 2002;McFarland & Majolo, 2012;Palagi et al., 2014). ...
This review demonstrates that many patterns of complex social behavior in group-living animals, in particular primates, may emerge by self-organization from cognitively “simple” competitive and affiliative interactions among group members. We focus on the lessons learnt from the different versions of model DomWorld. In these computational models, virtual individuals are group-living, initiate aggression in a risk-sensitive way, and groom others if they think they will be defeated. We show that, unexpectedly, in these models almost all social patterns emerge automatically that have been assumed to be displayed for cognitively more sophisticated reasons. For example, the emerging triadic agonistic patterns comprise all types of coalitionary support (conservative, bridging, and revolutionary), its reciprocation, and reciprocation of contra-support or opposition. Further, grooming is reciprocated, exchanged for support, and shown in patterns of post-conflict affiliation, including those of “reconciliation” and “consolation,” with similar differences between a tolerant and intolerant dominance style as in empirical data. These patterns emerge mainly because agonistic interactions create a spatio-social structure within groups that influences the occurrence of other social interactions in unexpected ways. When these dominance interactions in the computer model are based on the winner-loser effect, inter-sexual dominance relations appear to depend on sex ratio and intensity of aggression. Females become more similar in rank to males the fiercer the aggression among group members is and, in groups with intense aggression, the more male-biased the Adult Sex Ratio is. Similar empirical patterns have been reported for fish, primates, and humans. In conclusion, the DomWorld models illustrate how individual-based models provide an excellent tool for finding cognitively simple explanations for complex patterns of social behavior. These models generate nonintuitive results by integrating social interactions and their consequences across different levels, i.e., those of the individual, relationships, and group.
... Solicited consolation has been observed in a number of primate species (see Arnold and Barton [2001], Call et al. [2002], and Palagi et al. [2004] for recent findings, and Watts et al. [2000] for a review of earlier findings). Unsolicited consolation, however, has been observed only in bonobos , stumptailed macaques [Call et al., 2002], and chimpanzees [Arnold & Whiten, 2001;de Waal & Aureli, 1996;de Waal & van Roosmalen, 1979]. ...
... Initially, the finding of unsolicited consolation in chimpanzees but not macaques led de Waal and Aureli [1996] to propose the social cognition hypothesis, which states that the higher ''cognitive ability'' of chimpanzees (compared to macaques) enables them to ''empathize'' with the victim of a conflict, and thus exhibit unsolicited consolation. However, a recent study by Call et al. [2002] found evidence for unsolicited consolation in stumptailed macaques. ...
... However, a recent study by Call et al. [2002] found evidence for unsolicited consolation in stumptailed macaques. These results can be explained using the social constraints hypothesis, which states that unsolicited consolation should be more common in egalitarian than despotic societies because third parties have a lower risk of being attacked by the former aggressor in such societies [de Waal & Aureli, 1996]. Bonobo, chimpanzee, and stumptailed macaque societies have all been described as egalitarian [Sterck et al., 1997], and have all been observed to exhibit unsolicited consolation. ...
... Redirected affection and solicited postconflict contact, wherein the aggressor actively affiliates with a third party, were first observed in rhesus macaques (de Waal and Yoshihara 1983). Consolation is another form of bystander affiliation (de Waal and Aureli 1996), and is initiated by an uninvolved group member and directed to a victim of conflict. Consolation, like reconciliation, may alleviate the recipient's distress (de Waal and Aureli 1996), especially if initiators are the opponents' kin (Aureli and van Schaik 1991), and may establish the potential for short-term coalitions and so reduce the risk of further aggression for both the consoler and the victim of aggression. ...
... Consolation is another form of bystander affiliation (de Waal and Aureli 1996), and is initiated by an uninvolved group member and directed to a victim of conflict. Consolation, like reconciliation, may alleviate the recipient's distress (de Waal and Aureli 1996), especially if initiators are the opponents' kin (Aureli and van Schaik 1991), and may establish the potential for short-term coalitions and so reduce the risk of further aggression for both the consoler and the victim of aggression. This behavior was first observed in chimpanzees (de Waal and Aureli 1996) and more recently in stump-tailed macaques (Call et al. 2002). ...
... Consolation, like reconciliation, may alleviate the recipient's distress (de Waal and Aureli 1996), especially if initiators are the opponents' kin (Aureli and van Schaik 1991), and may establish the potential for short-term coalitions and so reduce the risk of further aggression for both the consoler and the victim of aggression. This behavior was first observed in chimpanzees (de Waal and Aureli 1996) and more recently in stump-tailed macaques (Call et al. 2002). ...
Since reconciliation was first described more than 20 years ago, a large number of postconflict behaviors have been observed among females in many polygynous primate species. However, few studies have been conducted among males, perhaps due to the rarity with which they maintain friendly relationships with one another and their aggressive competition for resources. Although this is true for many primate males, Sichuan snub-nosed monkeys Rhinopithecus roxellana represent a potential exception as male cooperation has been known to occur. In this study, using postconflict/matched-control(PC–MC) and time-rule methods, we analyzed postconflict behavior among males and the possible occurrence of bystander affiliation or reconciliation. A total of 246 PC–MC pairs among leader males were obtained. On average, each leader male exhibited only 0.04 aggressive behaviors per observation hour, and conciliation among leader males occurred at a low rate (2.03%) relative to other primate species. The occurrence of consolation–affiliation interactions between focal males and group members other than former opponents differed significantly between PCs and MCs, which is the first time this has been confirmed among R. roxellana males. We discuss the results in light of recent theories concerning consolation in primates. The patterns of postconflict contact demonstrated that R. roxellana may be a unique species among colobines.
... Several species have developed post-conflict behavioral strategies to preserve group integrity (see Fig. 1), namely: (i) reconciliation, which is defined as the first affiliative contact exchanged between the two opponents right after the end of the aggression (de Waal and van Roosmaleen 1979); and (ii) triadic affiliation, which is defined as the first affiliative contact exchanged between one (or both) of the opponents and an uninvolved third party (Romero et al. 2009(Romero et al. , 2011. In particular, triadic affiliations can be divided into two types: (i) 'solicited' if initiated by the victim or the aggressor and directed towards a third party (de Waal and Aureli 1996;de Waal 2000;Palagi and Cordoni 2009); and (ii) 'unsolicited' if spontaneously initiated by a third party and directed towards the victim or the aggressor (de Waal and Preston 2017). These two types of triadic affiliation can underlie different functions and cognitive abilities (Fraser et al. 2009;Romero et al. 2009Romero et al. , 2011Cordoni and Palagi 2015;de Waal and Preston 2017). ...
... Triadic solicited contacts occur when one of the former opponents (victim or aggressor) approaches an uninvolved third party and starts an affiliative contact with her/him (de Waal and Aureli 1996;de Waal 2000;Palagi and Cordoni 2009). From a functional point of view, solicited contacts can indeed regulate the former opponent's experience because they can (i) decrease the probability to receive further aggression from other group members (Victim Protection Hypothesis, Palagi and Norscia 2013;Palagi et al. 2014) and/or (ii) reduce self-anxiety (especially if victims; Self Anxiety Reduction Hypothesis; Palagi and Cordoni 2009;McFarland and Majolo 2012;Puga-Gonzalez et al. 2014). ...
In social mammals, conflict resolution involves the reunion of former opponents (aggressor and victim) after an aggressive event (reconciliation) or post-conflict triadic contacts with a third party, started by either opponent (solicited-TSC) or spontaneously offered by the third party (unsolicited-TUC). These post-conflict strategies can serve different functions, including consolation (specifically when TUCs reduce the victim’s anxiety). We investigated the possible presence and modulating factors of such strategies on semi-free ranging pigs (Sus scrofa; N = 104), housed at the ethical farm Parva Domus (Cavagnolo, Italy). Kinship was known. Reconciliation was present and mainly occurred between weakly related pigs to possibly improve tolerant cohabitation. Triadic contacts (all present except aggressor TSCs) mostly occurred between close kin. TSCs enacted by victims reduced neither their post-conflict anxiety behaviors nor further attacks by the previous aggressor, possibly because TSCs remained largely unreciprocated. TUCs towards aggressors did not reduce aggressor post-conflict anxiety but limited aggression redirection towards third parties. TUCs towards the victim reduced the victim but not the third-party’s anxiety. However, TUCs may also provide inclusive fitness benefits to third parties by benefiting close kin. In sum, pigs engaged in non-random solicited/unsolicited triadic contacts, which suggests that pigs might possess socio-emotional regulation abilities to change their own or others’ experience and elements of social appraisal, necessary to detect the emotional arousal of relevant others and (in case of TUCs) take the agency to restore homeostasis.
... One type of sympathetic concern is consola-tion behavior, where a bystander makes affiliative contact with and offers reassurance to a distressed individual. For example, chimpanzees (Pan troglodytes) sometimes embrace an individual after it has been attacked by a third party (29,30). Such consolation has been shown to reduce stress in the victim of the attack (31,32). ...
... Such consolation has been shown to reduce stress in the victim of the attack (31,32). Despite an obvious benefit accrued by the consoler, and, indeed, in the face of possible retaliation by the aggressor against the consoler, this behavior has been documented in both wild and captive chimpanzees, as well as bonobos (Pan paniscus) and stump-tailed macaques (Macaca arctoides; 17,[29][30][31][32][33][34]. ...
We present a brief overview of the study of empathy in nonhumans. We begin with a historical perspective that focuses on early ideas about empathy developed by Peter Kropotkin and Adam Smith. From there we discuss the origin and evolution of the multiple layers of empathy—emotional contagion, sympathetic concern, and empathetic perspective-taking—casting that discussion within the “Russian doll model” of empathy developed by de Waal. For each layer we provide examples from the animal behavior literature.
... Given the prosocial nature of consolation, researchers faced challenges in explaining the evolution of this behavior. De Waal and Aureli (1996) generated two evolutionary hypotheses, the Social Cognition Hypothesis (SCGH) and the Social Constraints Hypothesis (SCSH). Under the SCGH, empathy and metalizing abilities facilitate the rise of consolation among nonhuman primates. ...
... This hypothesis also predicts that consolation will occur between parties if the potential gains are higher than the costs of performing this behavior. De Waal and Aureli (1996) also argued that the risks depended on the power imbalances between the parties involved. Societies that frequently enforce a strict dominance hierarchy and discourage affiliation with low-status individuals generally lack consolation. ...
... Initially, consolation was found to occur in apes but not in monkeys. This result has been interpreted in the light of the cognitive constraint hypothesis as indicating the limitations in cognitive capacity of monkeys ( de Waal, Aureli 1996). According to this hypothesis, 'consolation' happens if a bystander recognizes that the victim is in distress and tries to alleviate the victim's distress, and monkeys were seen to lack 'cognitive empathy' (Preston, de Waal 2002). ...
... These findings questioned the cognitive constraint hypothesis, and stimulated researchers to concentrate on the social constraint hypothesis. This hypothesis suggested that the occurrence of 'consolation' may be related to a difference in the risks of aggression in different societies when approaching a former opponent ( de Waal, Aureli 1996). The virtue is, that in species with a tolerant dominance style, the risks of further aggression after a conflict are lower than in species with an intolerant dominance style, making such affiliation more likely. ...
... Another important PC affiliation is an affiliation offered by a previously uninvolved third party or 'bystander' to one of the former opponents (Call et al., 2002;De Waal and van Roosmalen, 1979;Palagi et al., 2004;Fraser et al., 2009). Consolation is a PC affiliation directed from an uninvolved bystander to the former recipient of an aggression (De Waal and van Roosmalen, 1979), while a PC affiliation directed from an uninvolved bystander to the former aggressor is termed appeasement (De Waal and Aureli, 1996;Romero et al., 2011). In chimpanzees, suggested but mutually non-exclusive functions of consolation include social support (true consolation), third party relationship repair, and self-protection (Fraser et al., 2008(Fraser et al., , 2009Fraser and Bugnyar, 2010;Koski and Sterck, 2009;Romero et al., 2010;Wittig and Boesch, 2010). ...
... A PC affiliation between former opponents was considered as reconciliation (De Waal and van Roosmalen, 1979), while a PC affiliation between a former opponent and any other group member uninvolved in the aggression was regarded as bystander PC affiliation. Bystander PC affiliations were considered as such when the affiliation was initiated by the bystander and directed to one of the former opponents, aggressor or victim (De Waal and Aureli, 1996;Wittig and Boesch, 2010). Both aggressors and victims were part of reconciliations and bystander PC affiliations, depending on the role a focal individual had during a conflict. ...
Many animals living in social groups have evolved behaviors to resolve conflicts between group members, behaviors thought crucial for maintaining stable group life. Several hypotheses, based mainly on observational data, aim to explain how post-conflict (PC) affiliations, such as reconciliation and consolation, resolve conflicts by restoring relationships and/or alleviating anxiety. To examine a potential endocrinological mechanism of PC affiliations, we used an experimental-like procedure to investigate whether the oxytocinergic system is activated during naturally observed reconciliations, receiving bystander PC affiliations and aggressions not followed by PC affiliations in wild male chimpanzees. We compared urinary oxytocin (uOT) levels after reconciliations, receiving bystander PC affiliations or aggressions without affiliations with two control conditions: affiliations without previous aggression and after time periods without social interactions. We furthermore tested the 'valuable relationship' hypothesis of reconciliation, as well as the influence of relationship quality between individuals engaged in each of the three behavioral conditions involving aggression on uOT levels. We found that the probability to reconcile a conflict increased with increasing relationship quality between opponents, thus our results support the 'valuable relationship' hypothesis. However, relationship quality did not influence uOT levels, while behavioral condition had a significant effect on uOT levels. uOT levels after reconciliations, receiving bystander PC affiliations and affiliations not related to conflicts were higher than after aggressions alone and time periods without social interactions. Overall, our results indicate that the oxytocinergic system is activated during affiliative interactions, whether occurring as reconciliation, bystander PC affiliation or affiliation alone. We conclude that the oxytocinergic system, in addition to building and maintaining social relationships, also takes part in repairing them.
... While it is likely that some qualitative taxonomic differences exist in the empathic and perspective-taking abilities of great apes as compared to monkeys (de Waal & Aureli, 1996), more recent data from monkeys, which we discuss in a following section, suggest more similarities than previously assumed (Palagi et al., 2014a). ...
... As with consolation, another important spontaneous manifestation of more advanced empathic perspective-taking is so-called targeted helping, which is help fine-tuned to another's specific situation ( de Waal, 1996). This kind of helping requires that one individual understands the pre- dicament another one is in and provides the exact solution to the other's problem. ...
Given that the cognitive and affective processes underlying empathy do not fossilize, studies of the empathic capacities of nonhuman primates provide us with a critical window through which we can explore the evolutionary origins of human empathy. Specifically, the comparative method provides an opportunity to determine which features of empathy are uniquely human and which are shared within the primate lineage. Here, we use the ethological approach to explore the affective and cognitive layers underlying empathy in primates. We review recent research exploring evidence for different layers of empathy, from the more basic forms such as yawn contagion and rapid facial mimicry, to more cognitively complex forms such as sympathetic concern and targeted helping. Combining evidence from both observational and experimental approaches, we argue that many of the core components underlying human empathy are deeply rooted in our primate past.
... Nevertheless, we found that immature gorillas directed consolatory contacts also toward unrelated individuals at significant levels. Aureli and de Waal (1996) proposed the Social Cognition Hypothesis to explain in part the occurrence of consolation in chimpanzees: empathy-the cognitive ability to perceive the distress of a conspecific-seems to be a necessary prerequisite for the occurrence of consolation (Aureli and Schaffner, 2002;Aureli and Smucny, 2000;Cords, 1997;de Waal and Aureli, 1996;Preston and de Waal, 2002). Even if empathy may be the prime mover of consolatory contacts, other factors have to concur to the occurrence of consolation. ...
... Nevertheless, we found that immature gorillas directed consolatory contacts also toward unrelated individuals at significant levels. Aureli and de Waal (1996) proposed the Social Cognition Hypothesis to explain in part the occurrence of consolation in chimpanzees: empathy-the cognitive ability to perceive the distress of a conspecific-seems to be a necessary prerequisite for the occurrence of consolation (Aureli and Schaffner, 2002;Aureli and Smucny, 2000;Cords, 1997;de Waal and Aureli, 1996;Preston and de Waal, 2002). Even if empathy may be the prime mover of consolatory contacts, other factors have to concur to the occurrence of consolation. ...
... Given its tension-reducing properties, the act of consoling an individual in distress is thought to represent a behavioural marker of empathy [16]. Consolation has also been observed in non-human animals, including monkeys and non-human great apes [17][18][19]; birds [20]; and rodents [21]. In addition to consolation, distress signals may also play a role in eliciting other social interactions in post-conflict (PC) contexts, like reunions among former opponents, a behaviour known as reconciliation [18] and in reducing the risk of further aggression [22]. ...
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’.
... Placing ourselves in another's mind, then asking "How would I feel or wish to be treated?", is at the heart of empathy. Love and empathy, as well as kindness, generosity, forgiveness and compassion are not uniquely human traits: non-human primates exhibit these properties [37][38][39][40][41] as do other mammals [42,43]. As already argued, there is nothing in the brains of animals that does not follow the laws of physics, chemistry, and biology. ...
... Another type of interaction that is worth investigating for understanding secondary intersubjectivity could be found in the social dynamics following an aggressive encounter. Specifically, we refer here to unsolicited triadic contacts that occur when an uninvolved third party spontaneously offers an affiliative behaviour to either the aggressor or the victim (de Waal & Aureli 1996). Unsolicited triadic contacts require that the conflict between two individuals is processed by a third, uninvolved subject and that the third party takes agency in resolving the conflict. ...
• Intersubjectivity has often been lauded as one of the defining features that separates humans and other extant hominids.
• Intersubjectivity and empathy are different, yet related, phenomena.
• The study of emotions and empathy-related abilities can provide insights into the ontogeny and evolution of intersubjectivity.
Intersubjectivity, which refers to the capacity to create shared value or connection between individuals, is a complex and multi-faceted phenomenon involving both cognitive and affective components. Intersubjectivity has often been lauded as one of the defining features that separates the social minds and existence of humans and non-human animals. Despite the apparently profound importance of inter-subjectivity for the socio-cognitive functioning of our species, we know surprisingly little about its evolution, nor how its evolution relates to the evolution of other related phenomena, such as empathy. In this review, we embrace the “bottom-up” perspective to consider recent theoretical and empirical advances in the fields of non-human animal cognition and emotion and what they can tell us about how complex socio-emotional capacities evolve. In particular, we focus on great ape species. Given their close phylogenetic relationship to us, great apes (the non-human, extant hominids) offer a unique lens to identify which of our capacities may be evolutionarily derived or phylogenetically shared.
... But there are also powerful forms of social support that more implicitly communicate understanding and concern, such as when a loved one holds our hand [7,8]. This type of physical support, often referred to as consoling touch, is observed across species and across cultures [9], and has been shown to reliably reduce the experience of physical pain [10][11][12]. Notably, however, research has yet to experimentally assess whether touch reduces the subjective experience of emotional pain in the same way that it reduces the subjective experience of physical pain. ...
Consoling touch is a powerful form of social support that has been repeatedly demonstrated to reduce the experience of physical pain. However, it remains unknown whether touch reduces emotional pain in the same way that it reduces physical pain. The present research sought to understand how handholding with a romantic partner shapes experiences of emotional pain and comfort during emotional recollection, as well as how it shapes lasting emotional pain associated with emotional experiences. Participants recalled emotionally painful memories or neutral memories with their partners, while holding their partner’s hand or holding a squeeze-ball. They additionally completed a follow-up survey to report how much emotional pain they associated with the emotional experiences after recalling them in the lab with their partners. Although consoling touch did not reduce emotional pain during the task, consoling touch increased feelings of comfort. Moreover, participants later recalled emotional memories that were paired with touch as being less emotionally painful than those that were not paired with touch. These findings suggest that touch does not decrease the immediate experience of emotional pain and may instead support adaptive processing of emotional experiences over time.
... This risk may be rather small, though, as research in chimpanzees and mandrills shows that providing comforting behaviour to others actually decreases the risk of receiving redirected aggression as compared to other bystanders (see Koski and Sterck, 2009;Schino and Marini, 2012). Second, it has been noted that the consolers do not appear distressed (de Waal and Aureli, 1996), but thus far there is no quantitative data on the consolers' emotional state prior to consolation. It may well be that an affiliative contact would not be effective in comforting the consoler. ...
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.
... Macaca fuscata: Aureli et al., 1993; Papio hamadryas hamadryas: Romero et al., 2009). Such lack of consolation in R. roxellana could be due to one possible explanation that monkeys lack the appropriate degree of empathy to perceive and respond to distress in others (de Waal and Aureli, 1996;Preston and de Waal, 2002). ...
For group-living primates, the information on postconflict management is crucial for understanding primate competition and cooperation. However, such information is poorly known for snub-nosed monkeys, especially for wild populations. In this study, from September 2007 to June 2008, we investigated postconflict behavior among adult females Sichuan snub-nosed monkeys Rhinopithecus roxellana within one-male units in a wild, provisioned group in the Qinling Mountains of China by means of the time-rule method and the PC-MC method. We obtained a total of 81 PC-MC pairs and each individual was involved in only 0.004 aggressive behavior per observation hour. The first affiliative behavior was more likely to occur within the first minute after a conflict. The postconflict affiliative behaviors most often seen were contact-sit, embrace and grooming. The affiliative contacts between adult females occur due to selective attraction, i.e. reconciliation. The pattern of postconflict affiliation demonstrates that the R. roxellana belongs to a tolerant species.
... Juveniles were the first to approach and affiliate with Pipo, both from his own group immediately after injury and from the foster group. In bonobos and gorillas, immature individuals were most likely to console (Cordoni et al. 2006; Clay and de Waal 2013) and de Waal and Aureli (1996) suggested that young rhesus macaques have greater "consolatory disposition" than adults. ...
Illegal live trade is one of the greatest threats to endangered Barbary macaques. Since trade focuses on young and Barbary macaques show extensive alloparental care, rehabilitation and release into wild foster groups is a promising strategy to free space in sanctuaries to allow continued confiscations, improve individual welfare, minimize captive care costs, and reinforce wild populations. However, little documentation exists on Barbary macaque releases and the age at which released macaques would be accepted by foster groups is unknown. Here, I report on a wild, nearly 3-year-old Barbary macaque male that was separated from his group after serious injury from a car collision. A neighbouring group found the distressed juvenile 2 days later, approached, groomed, and affiliated with him, and accepted him into the group. He remained with the foster group for 4 months until returning to his natal group. This observation challenges concerns that only infants and young juveniles may be accepted into non-natal groups, suggesting that even older confiscated juveniles could be candidates for release. This also adds to the literature on behavioural responses to distressed and injured individuals, demonstrating the capacity of wild monkeys to exhibit affiliative behaviours even towards unfamiliar distressed conspecifics, providing a rare example of potential consolation of an unknown individual. As so little documentation currently exists on the acceptance of Barbary macaques into wild foster groups, this observation provides valuable information for developing rehabilitation and release strategies for confiscated individuals, which can be critical for continuing to fight illegal trade of this endangered species.
... Prairie vole ability to perceive the emotional state of conspecifics was recently highlighted in a study, which, for the first time, described consolation behavior in rodents (Burkett et al., 2016). Consolation, previously studied in humans, nonhuman primates, elephants and canines (de Waal & Aureli 1996;Clay and de Waal, 2013), is defined as an increase in affiliative behavior directed toward a distressed individual. ...
Emotion recognition represents the ability to encode an ensemble of sensory stimuli providing information about the emotional state of another individual. This ability is not unique to humans. An increasing number of studies suggests that many aspects of higher order social functions, including emotion recognition, might be present in species ranging from primates to rodents, indicating a conserved role of these abilities in social animals.
The aim of this review is to use a comparative approach to examine how emotions are communicated and perceived in humans and other animals, with the intent to highlight possible new behavioral approaches and research perspectives. We summarize the evidence from human emotion recognition, and latest advances in the development of non‐human animal behavioral tests, using or implying the use of this cognitive function. The differential implication of sensory modalities used by animals to communicate and decipher emotional states is also discussed. The opportunity to measure emotion recognition abilities in rodents may allow us to better identify the neural mechanisms mediating this complex function, thus promoting the development of new intervention strategies for several neuropsychiatric disorders characterized by social cognitive dysfunctions.
... Post-aggression consolation in chimpanzees is promoted by social closeness [9,32,33]. Although several earlier studies did not find evidence associating age and sex with the likelihood of providing consolation [6,34,35], a recent study that draws on data from an exceptionally large sample of spontaneous consolation in chimpanzees, offers compelling evidence that female individuals are more likely to provide post-aggression consolation than males [9]. ...
Post-aggression consolation is assumed to occur in humans as well as in chimpanzees. While consolation following peer aggression has been observed in children, systematic evidence of consolation in human adults is rare. We used surveillance camera footage of the immediate aftermath of nonfatal robberies to observe the behaviors and characteristics of victims and bystanders. Consistent with empathy explanations, we found that consolation was linked to social closeness rather than physical closeness. While females were more likely to console than males, males and females were equally likely to be consoled. Furthermore, we show that high levels of threat during the robbery increased the likelihood of receiving consolation afterwards. These patterns resemble post-aggression consolation in chimpanzees and suggest that emotions of empathic concern are involved in consolation across humans and chimpanzees.
... Distinct social relationships characterized by friendly social behavior, such as grooming of potential partners (Seyfarth, 1978), seem to facilitate the formation of small coalitions in many primate species . Once relationships are disturbed by aggressive interactions, regular coalition partners may use reconciliation to re-establish former tolerance levels, allowing for future coalitionary behavior (de Waal & Aureli, 1996;Wittig & Boesch, 2003b). Researchers recognize the occurrence of reconciliation when former opponents engage in a friendly interaction soon after a conflict. ...
Coalitions among males during within group conflicts have a strong influence on the competitive and social environment within social groups. To evaluate possible variation in the occurrence of such coalitions in our closest relatives, chimpanzees and bonobos, we compared male aggression and coalitionary behavior in two natural communities, one of each species, with a similar size and composition. Furthermore we compared affiliative behavior that might be related to coalition formation among males. We found higher frequencies of aggression and a greater likelihood to form coalitions during within-group conflicts among wild male chimpanzees at Taï compared to wild male bonobos at LuiKotale. The species differed in the predominant sex of the male coalition partners, with male bonobos forming coalitions more often with females, while male chimpanzees formed coalitions more often with other males. Compared to male bonobos, male chimpanzees showed higher rates of grooming and tended to reconcile more conflicts with other males. Overall our results showed lower frequencies of reconciliation among bonobos than those described in captivity and at artificial feeding sites. These findings add to the evidence that male cooperation and conflict resolution are potentially very different in bonobos and chimpanzees, despite the fact that these two species are closely related, live in multi-male, multi-female communities with a high degree of fission-fusion dynamics and have female-biased migration patterns. Given the correlation between aggressive, cooperative and some affiliative patterns within the species in our study, we hypothesize that the fitness benefits of male relationships are greater in chimpanzees compared to bonobos.
... Several anecdotal reports have shown that chimpanzees occasionally assist others, as is the case of the chimpanzee who rescued another nonrelated female from drowning in the moat of their enclosure (Fouts & Mills, 1997) or the reported cases of adoptions by chimpanzee males (Boesch, Bolé, Eckhardt, & Boesch, 2010). In addition, there is evidence that chimpanzees regularly engage in behaviors such as food sharing, coalitionary support, and consolation (e.g., Boesch & Boesch-Achermann, 2000;de Waal & Aureli, 1996;Muller & Mitani, 2005). However, the difficulty with many of these observations is that it is not always possible to rule out a selfish motivation behind actors' actions, since often these behaviors also lead to immediate benefits for the actors. ...
... Though most non-human primate studies investigating the relationship between personality and early social history have focused on macaques, chimpanzees are also an excellent model for addressing these questions due to their close evolutionary relationship to humans and the range of complex social needs that influence their well-being (de Waal & Aureli, 1996). ...
Although much research has been conducted to understand personality development in humans, there remain substantial gaps in our understanding of these processes, particularly in relation to social influences. As such, investigations into personality development in our closest living relative, the chimpanzee, may provide useful insight. We evaluated the impact of early social exposure (to both humans and conspecifics) on personality development by studying 88 chimpanzees, including former pets and entertainers, living in accredited zoos and sanctuaries. During infancy, subjects varied in the amount of time spent with conspecifics compared with humans. Caregivers familiar with the chimpanzees rated them using a modified version of the Hominoid Personality Questionnaire (HPQ) and the ratings were found to have strong inter-rater reliability. We used the published structure of the HPQ to evaluate our results in relation to differences in early life experience. Chimpanzees who as infants spent less time with conspecifics were rated as lower in Extraversion later in life in comparison with chimpanzees who as infants spent more time with conspecifics. These results suggest that a broad range of social influences should be considered when evaluating the impact of early social environment on later personality expression. © 2016 Wiley Periodicals, Inc. Dev Psychobiol 9999: 1-9, 2016.
... This provides stress reduction to the victim-as well as the consoler (Koski & Sterck, 2007). This behavior occurs in apes (Romero, Castellanos, & de Waal, 2010) but has not been seen in monkeys (de Waal & Aureli, 1996), leading to the hypothesis that some empathetic behaviors require the more advanced cognition seen in the apes. Additionally, contagious yawning, a behavior that has been linked to empathy in humans, is also seen in several other species. ...
Researchers have studied non-human primate cognition along different paths, including social cognition, planning and causal knowledge, spatial cognition and memory, and gestural communication, as well as comparative studies with humans. This volume describes how primate cognition is studied in labs, zoos, sanctuaries, and in the field, bringing together researchers examining similar issues in all of these settings and showing how each benefits from the others. Readers will discover how lab-based concepts play out in the real world of free primates. This book tackles pressing issues such as replicability, research ethics, and open science. With contributors from a broad range of comparative, cognitive, neuroscience, developmental, ecological, and ethological perspectives, the volume provides a state-of-the-art review pointing to new avenues for integrative research.
Many people define behavior in terms of the motivations and intentions of the actor, but this is not directly observed, and is instead based on subjective judgments. Niko Tinbergen's position distinguishes structure from the four “why” questions. Defining behavior based on its Consequents is to define something based on its presumed function. To ascribe the cause of behavior to its function is a Teleological error, as that which has not yet occurred cannot be the cause of the present. Here, we discuss the potential functions of aggression, defining and distinguishing between conflict, competition, and aggression, and reconciliation in non-human primates.
The evolutionary origins of human right-handedness remain poorly understood. Some have hypothesized that tool use served as an important preadaptation for the eventual evolution of population-level right-handedness. In contrast, others have suggested that complex gestural and vocal communication served as prerequisite for the evolution of human right-handedness. In this study, we tested these competing hypotheses by comparing the handedness of bonobos and chimpanzees, two closely related species of Pan, on three different measures of hand use including simple reaching, manual gestures and coordinated bimanual actions. Chimpanzees are well known for their tool using abilities whereas bonobos rarely use tools in the wild. In contrast, many have suggested that bonobos have a more flexible gestural and vocal communication system than chimpanzees. The overall results showed that chimpanzees were significantly more right-handed than bonobos for all three measures suggesting that adaptations for tool use rather than communication may have led to the emergence of human right-handedness. We further show that species differences in handedness may be linked to variation in the size and asymmetry of the motor-hand area of the precentral gyrus. The results are discussed within the context of evolutionary theories of handedness, as well as some limitations in the approach to handedness measurement in nonhuman primates.
En el terreno de la moral, el contraste entre lo que el filósofo Wilfrid Sellars denominó "la imagen manifiesta" y la "imagen científica" es enorme y tiene unas consecuencias prácticas ineludibles. En la imagen manifiesta se albergan una serie de ideas comunes y cotidianas que a menudo se tienen sobre qué es la moral, cómo suele actuarse en situaciones moralmente cargadas, como evoluciona o degenera moralmente el mundo, la existencia del mal, la naturaleza de la corrupción, la forma de las normas sociales, etc. La imagen científica de nuestra moral viene emergiendo poco a poco, a partir de los estudios de la conducta, la toma de decisiones, la ética de las organizaciones y el escrutinio científico y filosófico de las intuiciones morales. Este libro reduce la distancia entre esa imagen manifiesta y la imagen científica, dotando al lector de referencias fundamentales, enfoques diversos y un amplio abanico de temáticas a partir de investigaciones experimentales sobre nuestra conducta moral. La cooperación, las intuiciones sobre justicia, las diferencias culturales o las teorías intuitivas de la moral son solo algunos de los temas abordados en un libro que interesará por igual al estudioso de estas temáticas y al lector deseoso de iniciarse en la nueva filosofía experimental.
Humans are social animals. Human societies emerge from vast networks of
cooperative interactions between many different individuals. In this respect, humans
are similar to most other primates. However, human societies are unusual among
primates in the number of different types of cooperative relationships that are
involved. In humans, males and females form strong pair bonds within large multimale,
multi-female societies in which many other cooperative relationships are also
important. How and when did human social systems arise? Do males and females use
different types of cooperative strategies? Under what conditions does paternal care
evolve? Do males and females have different constraints, and how do these affect the
types of social strategies they employ? How do factors such as environment quality
and seasonality modify these strategies? This thesis seeks answers to these questions
using computer simulations based on the iterated Prisoner's Dilemma. The
hypotheses generated by these models are tested using data from living primates.
They are then used to investigate the kinds of societies that our hominid ancestors
may have lived in.
The theoretical and empirical evidence presented in this thesis suggests that
sex differences in the energetic cost of reproduction determine the cooperative
strategies, and ultimately the types of social groups, that evolve. It is proposed that
during hominid evolution female energetic costs increased greatly, in comparison to
male energetic costs, due to changes in body size dimorphism, diet and brain size. A
two-stage model of hominid social structure is developed. The first stage, at the
transition from the australopithecines to Homo erectus, would have involved an
increase in female cooperation, especially food sharing. The second stage, occurring
between 500,000 and 100,000 years ago, would have involved male care giving, the
formation of pair bonds and the sexual division of labour within the context of a
wider cooperative network.
Across different cultures, social touch is used to alleviate distress. Here we adopt a dual-brain approach with fMRI to examine whether social touch involves similar activations between the suffering ‘target’ and the empathizer in brain regions related to emotional sharing such as the observation-execution (mirror) network. To inspect the neural underpinnings of the effects of social touch on pain, we scanned romantic couples during a task that required one partner (the empathizer) to hold the target’s hand as the latter experienced painful thermal stimulation. Empathizers and target participants were scanned sequentially, in two counterbalanced phases. Results revealed that hand-holding reduced the pain of the target participant, compared to the severity of pain in a control condition (holding a rubber ball). Importantly, during social touch we found striking shared activations between the target and empathizer in the inferior parietal lobule (IPL), a region related to the observation-execution network. The brain-to-brain analysis further revealed a positive correlation of IPL activation levels between the target and the empathizer. Finally, psychophysiological interaction (PPI) analysis in the target showed that the IPL activity during social touch was positively coupled with activity in the dorsomedial prefrontal cortex, a region that has been implicated in emotion regulation, suggesting that the interaction between the observation-execution network and emotion regulation network may contribute to pain reduction during social touch.
This thesis represents the first extended attempt by an archaeologist to construct an evolutionary theory of emotion. The handful of attempts that have appeared since the 1990s have failed to gain any real traction with archaeologists caught in a theoretical deadlock over the way in which an ‘archaeology of emotion’ should be approached.
This thesis will attempt to break the deadlock by reframing the debate around a ‘deep history of emotion’. It will be argued that it is only through a comprehensive longue durée approach that emotion can be understood in a prehistoric context. This requires the construction of a theory that can explain both the early biological origins of emotion and the later cultural constructions that characterize modern human societies.
This will be achieved through an appraisal of the interdisciplinary literature on emotion in search of a definition of emotion amendable for the archaeological enterprise. It is argued that rather than seeking discrete emotions directly, archaeologists should focus on the process by which emotional experiences are psychologically constructed and the cognitive traits that combine to produce complex emotional experience. Child development will be proposed as a starting point to understand how emotions are constructed from more basic cognitive ingredients.
Ultimately, three hypothetical mindstates will be proposed as heuristics through which hominin emotional capacities may be approached. Archaeological evidence for life history patterns and the cognitive ingredients of emotion will be used to anchor these mindstates in the past, providing predictions for the emotional vocabulary of hominins and possible new ways to interpret behaviour and material culture.
This thesis demonstrates that archaeologists can consider the emotional abilities of ancestral hominin by using innovative theoretical methods. An approach of this sort can provide new ways of looking at old data with the objective of expanding our appreciation of the decision-making processes that inform action.
Empathy as a research topic is receiving increasing attention, although there seems some confusion on the definition of empathy across different fields. Frans de Waal (de Waal FBM. Putting the altruism back into altruism: the evolution of empathy. Annu Rev Psychol 2008, 59:279–300. doi:10.1146/annurev.psych.59.103006.093625) used empathy as an umbrella term and proposed a comprehensive model for the evolution of empathy with some of its basic elements in nonhuman animals. In de Waal's model, empathy consists of several layers distinguished by required cognitive levels; the perception-action mechanism plays the core role for connecting ourself and others. Then, human-like empathy such as perspective-taking develops in outer layers according to cognitive sophistication, leading to prosocial acts such as targeted helping. I agree that animals demonstrate many empathy-related phenomena; however, the species differences and the level of cognitive sophistication of the phenomena might be interpreted in another way than this simple linearly developing model. Our recent studies with chimpanzees showed that their perspective-taking ability does not necessarily lead to proactive helping behavior. Herein, as a springboard for further studies, I reorganize the empathy-related phenomena by proposing a combination model instead of the linear development model. This combination model is composed of three organizing factors: matching with others, understanding of others, and prosociality. With these three factors and their combinations, most empathy-related matters can be categorized and mapped to appropriate context; this may be a good first step to discuss the evolution of empathy in relation to the neural connections in human and nonhuman animal brains. I would like to propose further comparative studies, especially from the viewpoint of Homo-Pan (chimpanzee and bonobo) comparison. For further resources related to this article, please visit the WIREs website.
The notion that phenomenologically observable differences in the human eye are correlated with behavioral tendencies (other than gaze-following) has been addressed poorly in the psychological literature. Most notably, the proposed correlations are based on an arbitrary categorization in discrete categories of the continuous variability across various traits that could be contributing to individual eye morphologies. We review the relevant literature and assume a view of human eyes as sign stimuli, identifying the relative contrast between the iridal and scleral areas as the main contributor to the strength of the signal. Based on this view, we present a new method for the precise quantification of the relative luminosity of the iris (RLI) and briefly discuss its potential applications in psychological research.
Is moral behavior unique to humans? Although moral behavior is primarily discussed in relation to humans, if a function of moral behavior is to promote social cohesion and harmony within a social group, there is no a priori reason not to expect a similar set of behaviors in other social species (Bonnie and de Waal 2004; Flack and de Waal 2000; Haidt 2003). Although this certainly does not mean that what we see in other species need be identical to humans’ behavior, there may be a suite of related behaviors that have evolved for the same purposes in other species. Of course, this idea is not new. In The Descent of Man and Selection in Relation to Sex (Darwin 1871/1981: 71–72), Darwin argued that sociality is innate, rather than created by humankind, and provided a framework for the development of morality in any species. The question is, then, from which precursor behaviors did morality evolve, and how can we study this in other species?
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