Neural pathways from the central nucleus of the amygdala to the paraventricular nucleus of the hypothalamus are involved in induction of yawning behavior due to emotional stress in rats.

As yawning is often observed in stressful or emotional situations such as tension and anxiety, this suggests that yawning can be considered to be an emotional behavior. However, the neural mechanisms underlying emotion-induced yawning remain unclear. It is well known that the hypothalamic paraventricular nucleus (PVN) is the most important brain structure for induction of yawning behavior. We previously showed that induction of yawning involves the central nucleus of the amygdala (CeA), as well as the PVN. Therefore, emotion-induced yawning could potentially be induced through activation of the direct/indirect neural pathways from the CeA to the PVN. Our present study used a combination of retrograde tracing (injection of Fluoro-Gold (FG) into the PVN) and c-Fos immunohistochemistry to examine the neural pathways that evoke emotion-induced yawning. We additionally performed lesion experiments on the CeA using ibotenic acid, a neurotoxin, to determine whether the CeA is involved in the induction of emotion-induced yawning. Emotional stress by fear conditioning induced yawning behavior, and induced expression of double-labeled cells for c-Fos and FG in the bed nucleus of the stria terminalis (BNST), but not in the CeA. Furthermore, the CeA lesions caused by ibotenic acid abolished the induction of emotion-induced yawning. These results suggest that a neural pathway from the CeA to the PVN via the BNST may be primarily involved in the induction of emotion-induced yawning behavior.

Familiarity modulates both intra- and interspecific yawn contagion in red-capped mangabeys.

Yawn contagion (YC) is, compared to spontaneous yawning, an evolutionary recent phenomenon probably linked to behavioral synchronization in highly social species that is more likely when it involves familiar subjects. Here, we investigate for the first time in monkeys which factors modulate intra- and interspecific YC. Through an experimental approach, we exposed 17 red-capped mangabeys to video stimuli (Yawn vs Control) depicting familiar/unfamiliar red-capped mangabeys and humans, and unfamiliar hamadryas. We found that mangabeys yawned more often in response to Yawn than Control videos independently from the species depicted, demonstrating both intra- and interspecific YC in the tested species. Moreover, both mangabey and human familiar yawning stimuli evoked a stronger yawning response in the subjects compared to the unfamiliar counterparts. Neither the amount of time spent looking frontally at the screen (probability of stimulus perception) nor the levels of self-directed behaviors (a proxy of anxiety) accounted for the results. In conclusion, we provide the first evidence that in non-human primate familiarity modulates both intra- and inter-specific YC. Stimuli emitted by familiar faces somehow ease the mechanisms underlying YC, and this modulation can also apply to heterospecific subjects when previous shared experiences provide the prerequisites for the development of social bonds.

Parakinesia Brachialis Oscitans - a Rare Post-Stroke Phenomenon.

Background: Abnormal involuntary movement of paralyzed upper limb during yawning is a rare phenomenon termed as parakinesia brachialis oscitans. Case Report: We describe a 59-year-old gentleman with abnormal involuntary movement of paralyzed right upper limb during yawning 2 weeks following ischemic stroke of left middle cerebral artery territory. Discussion: This is a rare post-stroke phenomenon and its pathophysiological mechanism is poorly understood but this entity highlights possible preserved extrapyramidal pathway which might help in rehabilitating stroke survivors.

Yawning and airway physiology: a scoping review and novel hypothesis.

BACKGROUND AND PURPOSE: Yawning is a stereotypical complex muscular movement and is commonly executed by most vertebrates. In seconds, the entire airway is fully dilated and surrounding muscles are powerfully stretched, most prominently around the pharynx. To date, yawning has been rarely studied, and as of yet there is no consensus on its main function. MATERIAL AND METHODS: To investigate a mechanical airway function for yawning, a literature search was conducted to relate the frequency of yawning and obstructive airway conditions. RESULTS: The results show that changes in obstructive airway conditions and alteration of the frequency of yawning are temporally related. INTERPRETATION: These relationships, however, cannot be interpreted as causal, nor can they be extrapolated to explain the function of yawning. Yet airway management and yawning share many physiological characteristics. We therefore propose a novel hypotheses: yawning plays a significant role in airway physiology by muscle repositioning and widening the airway lumen, thereby securing long-term oxygenation.

Mating-induced analgesia is dependent of copulatory male pattern in high- and low- yawning male rats.

Mating behavior in rodents can modulate pain sensations in both sexes. In males, the execution of mounts, intromissions, and ejaculations induced a progressive increase in their vocalization thresholds induced by tail shocks and other types of noxious stimuli. We selectively inbred two sublines from Sprague-Dawley (SD) rats that differed in their spontaneous yawning frequency. The high-yawning (HY) subline had a mean of 20 yawns/h and a different pattern of sexual behavior characterized by longer interintromission intervals and more sexual bouts that delayed ejaculation. The low-yawning (LY) subline and SD rats yawned as a mean 2 and 1 yawns/h, respectively. So, we determine mating-induced analgesia in HY, LY, and SD male rats by measuring vocalization thresholds in response to noxious electric tail shocks. Our results showed that the magnitude of mating-induced analgesia was lower in HY and LY rats with respect to SD rats. When the rats performed different components of male sexual pattern, both sublines exhibited a significantly lower increase in their vocalization thresholds with respect to SD rats-being sublines less responsive regarding mating-induced analgesia. Pain modulation mechanisms depend on responses to stress, so the low levels of analgesia obtained in the yawning sublines may be due either to differences in their response to stress in other paradigms, or to atypical performance of male sexual behavior during mating, an event which as a stressful event in rats. Therefore, the yawning sublines are a suitable model for analyzing how a different temporal pattern in the display of male sexual behavior affects analgesia mechanisms. Our results concur with Wistar rats with different endophenotypes that could apply to humans as well.

Yawn contagion in bonobos: Another group, another story.

In primates, yawn contagion (the yawning response elicited by others' yawn) is variably influenced by individual (e.g., sex, age) and social factors (e.g., familiarity) and possibly linked to interindividual synchronization, coordination, and emotional contagion. Two out of three studies on yawn contagion in bonobos (Pan paniscus), found the presence of the phenomenon with mixed results concerning the effect of familiarity and no replication on its modulating factors. To address this puzzling issue, we recorded all occurrences data on yawn contagion in a captive bonobo group (March-June 2021; 18 individuals; La Vallée des Singes, France). Contrary to chimpanzees and humans, the number of triggering yawns increased contagion, possibly owing to a higher stimulus threshold. This aspect may explain the interindividual variability observed in yawn contagion rates. In subjects under weaning, we did not detect yawn contagion and, as it occurs in certain human cohorts, yawn contagion declined with age, possibly due to reduced sensitivity to others. Females responded more than males and elicited more responses from females when showing sexual swelling. As reproductive females are central in bonobo society, our results support the hypothesis that-as in other Hominini-the most influential sex can influence yawn contagion. The relationship quality (measured via grooming/play) did not affect yawn contagion, possibly due to bonobos' xenophilic nature. Overall, this study confirms the presence of yawn contagion in bonobos and introduces new elements on its modulating factors, pointing toward the necessity of cross-group studies.

First evidence of yawn contagion in a wild monkey species.

Yawn contagion occurs when individuals yawn in response to the yawn of others (triggers). This is the first account of yawn contagion in wild geladas (Theropithecus gelada), a monkey species that shows yawn contagion in captivity and is organized in core units (one-male/bachelor groups) forming multilevel associations. In a population of geladas from the Kundi plateau (Ethiopia) we found that the yawning response was highest when geladas could perceive a triggering yawn, which confirms that yawn contagion is present in the wild. Yawn duration, mouth-opening degree and presence/absence of vocalisation (possibly modulating yawn detectability) did not affect the likelihood of contagion. Males and females, known to be both implicated in movement initiation within groups, were similarly powerful as yawn triggers. Instead, group membership and responder sex had a significant role in shaping the phenomenon. Yawn contagion was highest between individuals belonging to different core units and males were most likely to respond to others' yawns. Because males have a non-negligible role in inter-group coordination, our results suggest that yawn contagion may have a communicative function that goes beyond the basic unit level.

Earlier than previously thought: Yawn contagion in preschool children.

Yawning is a primitive and stereotyped motor action involving orofacial, laryngeal, pharyngeal, thoracic and abdominal muscles. Contagious yawning, an involuntarily action induced by viewing or listening to others' yawns, has been demonstrated in human and several non-human species. Previous studies with humans showed that infants and preschool children, socially separated during video experiments, were not infected by others' yawns. Here, we tested the occurrence of yawn contagion in 129 preschool children (ranging from 2.5 to 5.5 years) belonging to five different classes by video recording them in their classrooms during the ordinary school activities. As it occurs in adult humans, children of all ages were infected by others' yawns within the 2 min after the perception of the stimulus. The yawn contagion occurred earlier than previously thought. For children, it appears that the natural social setting is more conducive to yawn contagion than the inherently artificial experimental approach. Moreover, children's gender did not affect the level of contagious yawning. The neural, emotional and behavioural traits of preschool children are probably not sufficiently mature to express variability between boys and girls; nevertheless, children appeared to be already well equipped with the 'neural toolkit' necessary for expressing yawn contagion.

Yawn contagion in domestic pigs (Sus scrofa).

Contrary to spontaneous yawning-an ancient phenomenon common to vertebrates-contagious yawning (elicited by others' yawns) has been found only in highly social species and may reflect an emotional inter-individual connection. We investigated yawn contagion in the domestic pig, Sus scrofa. Owing to the complex socio-emotional and cognitive abilities of Sus scrofa, we posited that yawn contagion could be present in this species (Prediction 1) and influenced by individual/social factors (Prediction 2). In June-November 2018, on 104 semi-free ranging adolescent/adult pigs, 224 videos were recorded for video analysis on yawning. Kinship information was refined via genetic analyses. Statistical elaboration was conducted via GLMMs and non-parametric/randomization/cross-tabulation tests. We found yawn contagion in Sus scrofa, as it was more likely that pigs yawned when perceiving rather than not perceiving (yawning/control condition) others' yawns (response peak in the first out of three minutes). Yawn contagion was more likely: (1) in response to males' yawns; (2) as the age increased; (3) within short distance (1 m); (4) between full siblings, with no significant association between kinship and distance. The influence of kinship suggests that-as also hypothesized for Homo sapiens-yawn contagion might be linked with emotional communication and possibly contagion.

On the Emergence of Tremor in Prodromal Parkinson's Disease.

Clinical, neuropathological and neuroimaging research suggests that pathological changes in Parkinson's disease (PD) start many years before the emergence of motor signs. Since disease-modifying treatments are likely to be most effective when initiated early in the disease process, there has been significant interest in characterizing prodromal PD. Some people with PD describe autonomic symptoms at the time of diagnosis suggesting that autonomic dysfunction is a common feature of prodromal PD. Furthermore, subtle motor signs may be present and emerge prior to the time of diagnosis. We present a series of patients who, in the prodromal phase of PD, experienced the emergence of tremor initially only while yawning or straining at stool and discuss how early involvement of autonomic brainstem nuclei could lead to these previously unreported phenomena. The hypothalamic paraventricular nucleus (PVN) plays a central role in autonomic control including bowel/bladder function, cardiovascular homeostasis and yawning and innervates multiple brainstem nuclei involved in autonomic functions (including brainstem reticular formation, locus ceruleus, dorsal raphe nucleus and motor nucleus of the vagus). The PVN is affected in PD and evidence from related phenomena suggest that the PVN could increase tremor either by increasing downstream cholinergic activity on brainstem nuclei such as the reticular formation or by stimulating the locus ceruleus to activate the cerebellothalamocortical network via the ventrolateral nucleus of the thalamus. Aberrant cholinergic/noradrenergic transmission between these brainstem nuclei early in PD couldlead to tremor before the emergence of other parkinsonian signs, representing an early clinical clue to prodromal PD.

An Adaptive Face Tracker with Application in Yawning Detection.

In this work, we propose an adaptive face tracking scheme that compensates for possible face tracking errors during its operation. The proposed scheme is equipped with a tracking divergence estimate, which allows to detect early and minimize the face tracking errors, so the tracked face is not missed indefinitely. When the estimated face tracking error increases, a resyncing mechanism based on Constrained Local Models (CLM) is activated to reduce the tracking errors by re-estimating the tracked facial features' locations (e.g., facial landmarks). To improve the Constrained Local Model (CLM) feature search mechanism, a Weighted-CLM (W-CLM) is proposed and used in resyncing. The performance of the proposed face tracking method is evaluated in the challenging context of driver monitoring using yawning detection and talking video datasets. Furthermore, an improvement in a yawning detection scheme is proposed. Experiments suggest that our proposed face tracking scheme can obtain a better performance than comparable state-of-the-art face tracking methods and can be successfully applied in yawning detection.

Contagious yawning is not a signal of empathy: no evidence of familiarity, gender or prosociality biases in dogs.

Contagious yawning has been suggested to be a potential signal of empathy in non-human animals. However, few studies have been able to robustly test this claim. Here, we ran a Bayesian multilevel reanalysis of six studies of contagious yawning in dogs. This provided robust support for claims that contagious yawning is present in dogs, but found no evidence that dogs display either a familiarity or gender bias in contagious yawning, two predictions made by the contagious yawning-empathy hypothesis. Furthermore, in an experiment testing the prosociality bias, a novel prediction of the contagious yawning-empathy hypothesis, dogs did not yawn more in response to a prosocial demonstrator than to an antisocial demonstrator. As such, these strands of evidence suggest that contagious yawning, although present in dogs, is not mediated by empathetic mechanisms. This calls into question claims that contagious yawning is a signal of empathy in mammals.

The neuroethology of spontaneous mimicry and emotional contagion in human and non-human animals.

Spontaneous mimicry appears fundamental to emotional perception and contagion, especially when it involves facial emotional expressions. Here we cover recent evidence on spontaneous mimicry from ethology, psychology and neuroscience, in non-human and human animals. We first consider how mimicry unfolds in non-human animals (particularly primates) and how it relates to emotional contagion. We focus on two forms of mimicry-related phenomena: facial mimicry and yawn contagion, which are largely conserved across mammals and useful to draw evolutionary scenarios. Next, we expand on the psychological evidence from humans that bears on current theoretical debates and also informs non-human animal research. Finally, we cover the neural bases of facial mimicry and yawn contagion. We move beyond the perception/expression/experience trichotomy and from the correlational to the causal evidence that links facial mimicry to emotional contagion by presenting evidence from neuroimaging, direct manipulation, neuro-stimulation and lesion studies. In conclusion, this review proposes a bottom-up, multidisciplinary approach to the study of spontaneous mimicry that accounts for the evolutionary continuity linking non-human and human animals.

Emotional contagion in children with autism spectrum disorder varies with stimulus familiarity and task instructions.

Although deficits in cognitive empathy are well established in individuals with autism spectrum disorder (ASD), the literature on emotional empathy, or emotional contagion, in individuals with ASD is sparse and contradictory. The authors tested susceptibility to contagious yawning and laughter in children with ASD (n = 60) and typically developing (TD) children (n = 60), ages 5-17 years, under various conditions, to elucidate factors that may affect emotional contagion in these populations. Although TD children showed equal amounts of emotional contagion across conditions, children with ASD were highly influenced by the familiarity of the target stimulus, as well as task instructions that encourage eye gaze to target. More specifically, children with ASD exhibited less contagious yawning and laughter than their TD peers except when their attention was explicitly directed to the eyes or (and even more so) when their parents served as the stimulus targets. The authors explore the implications of these findings for theories about the mechanisms underlying empathic deficits in ASD as well as the clinical implications of having parents involved in treatment.

Spontaneous Yawning and its Potential Functions in South American Sea Lions (Otaria flavescens).

Spontaneous yawning is a widespread behaviour in vertebrates. However, data on marine mammals are scarce. In this study, we tested some hypotheses on the functions of yawning in a captive group of South American sea lions (Otaria flavescens). According to the Dimorphism Hypothesis, species showing low levels of sexual dimorphism in canine size do not show sex differences in yawning distribution; this was supported by our findings, since yawning did not differ between the sexes. Yawning was more frequently performed during resting/sleeping contexts, thus supporting the Drowsiness Hypothesis. Yawning and self-scratching are considered reliable indicators of short-term anxiety in sea lions, since they immediately increased after conflicts both in aggressors and victims (Social Distress Hypothesis supported). In the long-term, yawning was not correlated with individuals' dominance status, thus showing that anxiety is similarly experienced by dominants and subordinates. The last two findings can be explained by the social competition of this species, that involves individuals independently from their sex, age or ranking status. Therefore, the exposure to frequent stressful events can induce similar levels of anxiety in all the subjects (Resource Inequity Hypothesis supported). In conclusion, spontaneous yawning in sea lions seems to share similar functions with other social mammals, suggesting that this behaviour is a possible plesiomorphic trait.

Pathological Yawning in Patients with Acute Middle Cerebral Artery Infarction: Prognostic Significance and Association with the Infarct Location

Background: Pathological yawning is a compulsive, frequent, repetitive yawning triggered by a specific reason besides fatigue or boredom. It may be related to iatrogenic, neurologic, psychiatric, gastrointestinal, or metabolic disorders. Moreover, it could also be seen in the course of an ischemic stroke. Aims: To determine whether pathological yawning is a prognostic marker of middle cerebral artery stroke and evaluate its relationship with the infarct location. Study Design: Cross-sectional study. Methods: We examined 161 patients with acute middle cerebral artery stroke, consecutively admitted to emergency department. Demographic information, stroke risk factors, stroke type according to Trial of Org 10172 in Acute Stroke Treatment classification, blood oxygen saturation, body temperature, blood pressure, heart rate, glucose levels, daytime of stroke onset, National Institutes of Health Stroke Scale score (National Institutes of Health Stroke Scale score, at admission and 24 h), modified Rankin scale (at 3 months), and infarct locations were documented. Pathological yawning was defined as ≥3 yawns/15 min. All patients were observed for 6 hours to detect pathological yawning. National Institutes of Health Stroke Scale score >10 was determined as severe stroke. The correlation between the presence of pathological yawning and stroke severity, infarct location, and the short- and long-term outcomes of the patients were evaluated. Results: Sixty-nine (42.9%) patients had pathological yawning and 112 (69.6%) had cortical infarcts. Insular and opercular infarcts were detected in 65 (40.4%) and 54 (33.5%) patients, respectively. Pathological yawning was more frequently observed in patients with cortical, insular, and opercular infarcts (p<0.05). Pathological yawning was related to higher National Institutes of Health Stroke Scale scores. Patients with severe stroke (National Institutes of Health Stroke Scale score ≥10) presented with more pathological yawning than those with mild to moderate strokes (p<0.05). The clinical outcomes and mortality rates showed no significant relationship with the occurrence of pathological yawning. Conclusion: Pathological yawning in middle cerebral artery stroke was associated with stroke severity, presence of cortical involvement, and insular and opercular infarcts. However, no association was found with long-term outcome and mortality.

Observational data reveal evidence and parameters of contagious yawning in the behavioral repertoire of captive-reared chimpanzees (Pan troglodytes).

There is robust experimental evidence for contagious yawning, yet observational studies of naturalistic behavior have been fewer. Without data from real-world behavior, researchers have questioned the existence of contagious yawning and made assumptions about some parameters (e.g., the duration of the effect). We observed contagious yawning in chimpanzees to confirm/disconfirm its existence in the behavioral repertoire of this species, and if present, provide some of the missing descriptives. We recorded yawns on an all-occurrence basis from 18 captive-reared chimpanzees at the Los Angeles Zoo. We recorded identity, time, and individuals who could have been affected. We calculated a threshold for contagion by taking the mean and adding 1.96 standard deviations, constructing a response curve. Across multiple measures we see a consistent pattern in which there is a strong effect of contagion for 1.5 minutes, a less strong but still significant effect lasting up to 3.5 minutes in some measures, and no evidence of contagion beyond 3.5 minutes. From the time stamp on each yawn we were able to rule out temporal synchrony as an alternative hypothesis. Thus, contagious yawning appears to be a natural phenomenon in chimpanzees lending support to the myriad experimental and observational studies to date.