Pathogen disgust sensitivity protects against infection in a high pathogen environment.

Disgust is hypothesized to be an evolved emotion that functions to regulate the avoidance of pathogen-related stimuli and behaviors. Individuals with higher pathogen disgust sensitivity (PDS) are predicted to be exposed to and thus infected by fewer pathogens, though no studies have tested this directly. Furthermore, PDS is hypothesized to be locally calibrated to the types of pathogens normally encountered and the fitness-related costs and benefits of infection and avoidance. Market integration (the degree of production for and consumption from market-based economies) influences the relative costs/benefits of pathogen exposure and avoidance through sanitation, hygiene, and lifestyle changes, and is thus predicted to affect PDS. Here, we examine the function of PDS in disease avoidance, its environmental calibration, and its socioecological variation by examining associations among PDS, market-related lifestyle factors, and measures of bacterial, viral, and macroparasitic infection at the individual, household, and community levels. Data were collected among 75 participants (ages 5 to 59 y) from 28 households in three Ecuadorian Shuar communities characterized by subsistence-based lifestyles and high pathogen burden, but experiencing rapid market integration. As predicted, we found strong negative associations between PDS and biomarkers of immune response to viral/bacterial infection, and weaker associations between PDS and measures of macroparasite infection, apparently mediated by market integration-related differences. We provide support for the previously untested hypothesis that PDS is negatively associated with infection, and document variation in PDS indicative of calibration to local socioeconomic conditions. More broadly, findings highlight the importance of evolved psychological mechanisms in human health outcomes.

The evolution of masturbation is associated with postcopulatory selection and pathogen avoidance in primates.

Masturbation occurs throughout the animal kingdom. At first glance, however, the fitness benefits of this self-directed behaviour are unclear. Regardless, several drivers have been proposed. Non-functional hypotheses posit that masturbation is either a pathology, or a byproduct of high underlying sexual arousal, whereas functional hypotheses argue an adaptive benefit. The Postcopulatory Selection Hypothesis states that masturbation aids the chances of fertilization, while the Pathogen Avoidance Hypothesis states that masturbation helps reduce host infection by flushing pathogens from the genital tract. Here, we present comprehensive new data documenting masturbation across the primate order and use these, in conjunction with phylogenetic comparative methods, to reconstruct the evolutionary pathways and correlates of masturbation. We find that masturbation is an ancient trait within the primate order, becoming a more common aspect of the haplorrhine behavioural repertoire after the split from tarsiers. Our analyses provide support for both the Postcopulatory Selection and Pathogen Avoidance Hypotheses in male primates, suggesting that masturbation may be an adaptive trait, functioning at a macroevolutionary scale.

Behavioral and physiological sensitivity to natural sick faces.

The capacity to rapidly detect and avoid sick people may be adaptive. Given that faces are reliably available, as well as rapidly detected and processed, they may provide health information that influences social interaction. Prior studies used faces that were manipulated to appear sick (e.g., editing photos, inducing inflammatory response); however, responses to naturally sick faces remain largely unexplored. We tested whether adults detected subtle cues of genuine, acute, potentially contagious illness in face photos compared to the same individuals when healthy. We tracked illness symptoms and severity with the Sickness Questionnaire and Common Cold Questionnaire. We also checked that sick and healthy photos were matched on low-level features. We found that participants (N = 109) rated sick faces, compared to healthy faces, as sicker, more dangerous, and eliciting more unpleasant feelings. Participants (N = 90) rated sick faces as more likely to be avoided, more tired, and more negative in expression than healthy faces. In a passive-viewing eye-tracking task, participants (N = 50) looked longer at healthy than sick faces, especially the eye region, suggesting people may be more drawn to healthy conspecifics. When making approach-avoidance decisions, participants (N = 112) had greater pupil dilation to sick than healthy faces, and more pupil dilation was associated with greater avoidance, suggesting elevated arousal to threat. Across all experiments, participants' behaviors correlated with the degree of sickness, as reported by the face donors, suggesting a nuanced, fine-tuned sensitivity. Together, these findings suggest that humans may detect subtle threats of contagion from sick faces, which may facilitate illness avoidance. By better understanding how humans naturally avoid illness in conspecifics, we may identify what information is used and ultimately improve public health.

Disgust in animals and the application of disease avoidance to wildlife management and conservation.

Disgust is an adaptive system hypothesized to have evolved to reduce the risk of becoming sick. It is associated with behavioural, cognitive and physiological responses tuned to allow animals to avoid and/or get rid of parasites, pathogens and toxins. Little is known about the mechanisms and outcomes of disease avoidance in wild animals. Furthermore, given the escalation of negative human-wildlife interactions, the translation of such knowledge into the design of evolutionarily relevant conservation and wildlife management strategies is becoming urgent. Contemporary methods in animal ecology and related fields, using direct (sensory cues) or indirect (remote sensing technologies and machine learning) means, provide a flexible toolbox for testing and applying disgust at individual and collective levels. In this review/perspective paper, we provide an empirical framework for testing the adaptive function of disgust and its associated disease avoidance behaviours across species, from the least to the most social, in different habitats. We predict various trade-offs to be at play depending on the social system and ecology of the species. We propose five contexts in which disgust-related avoidance behaviours could be applied, including endangered species rehabilitation, invasive species, crop-raiding, urban pests and animal tourism. We highlight some of the perspectives and current challenges of testing disgust in the wild. In particular, we recommend future studies to consider together disease, predation and competition risks. We discuss the ethics associated with disgust experiments in the above contexts. Finally, we promote the creation of a database gathering disease avoidance evidence in animals and its applications.

Le dégoût est un système adaptatif supposé avoir évolué afin de réduire le risque de tomber malade. Il est associé à des réponses comportementales, cognitives et physiologiques adaptées pour permettre aux animaux d'éviter et/ou de se débarrasser des parasites, pathogènes et toxines. On sait peu de choses sur les mécanismes et les conséquences de l'évitement des maladies chez les animaux sauvages. Étant donné l'escalade des interactions négatives entre humains et faune, la traduction de ces connaissances dans la conception de stratégies de conservation et de gestion de la faune - prenant en considération l'évolution des espèces - devient urgente. Les méthodes contemporaines en écologie animale et dans les domaines connexes, utilisant des moyens directs (indices sensoriels) ou indirects (technologies de télédétection et apprentissage automatique), fournissent une boîte à outils flexible pour tester et appliquer le dégoût aux niveaux individuel et collectif. Dans cet article de revue/perspective, nous fournissons un cadre empirique pour tester la fonction adaptative du dégoût et les comportements associés d'évitement des maladies chez différentes espèces - des moins sociales aux plus sociales, et dans différents habitats. Nous prédisons divers compromis en fonction du système social et de l'écologie de l'espèce. Nous proposons cinq contextes dans lesquels les comportements d'évitement liés au dégoût pourraient être appliqués: la réhabilitation d'espèces menacées; les espèces envahissantes; les dommages aux cultures; les nuisibles urbains; et le tourisme animalier. Nous mettons en avant certaines perspectives et défis actuels de l'expérimentation sur le dégoût en milieu naturel. En particulier, nous recommandons la considération de plusieurs risques ensemble: maladie, prédation et compétition. Nous discutons également de l'éthique associée aux expériences sur le dégoût dans les contextes ci-dessus. Enfin, nous promouvons la création d'une base de données rassemblant les stratégies d'évitement des maladies chez les animaux et leurs applications.

Dissection of a sensorimotor circuit underlying pathogen aversion in C. elegans.

BACKGROUND: Altering animal behavior to reduce pathogen exposure is a key line of defense against pathogen attack. In Caenorhabditis elegans, alterations in intestinal physiology caused by pathogen colonization and sensation of microbial metabolites may lead to activation of pathogen aversive behaviors ranging from aversive reflexes to learned avoidance. However, the neural circuitry between chemosensory neurons that sense pathogenic bacterial cues and the motor neurons responsible for avoidance-associated locomotion remains unknown. RESULTS: Using C. elegans, we found that backward locomotion was a component of learned pathogen avoidance, as animals pre-exposed to Pseudomonas aeruginosa or Enterococcus faecalis showed reflexive aversion to drops of the bacteria driven by chemosensory neurons, including the olfactory AWB neurons. This response also involved intestinal distention and, for E. faecalis, required expression of TRPM channels in the intestine and excretory system. Additionally, we uncovered a circuit composed of olfactory neurons, interneurons, and motor neurons that controls the backward locomotion crucial for learned reflexive aversion to pathogenic bacteria, learned avoidance, and the repulsive odor 2-nonanone. CONCLUSIONS: Using whole-brain simulation and functional assays, we uncovered a novel sensorimotor circuit governing learned reflexive aversion. The discovery of a complete sensorimotor circuit for reflexive aversion demonstrates the utility of using the C. elegans connectome and computational modeling in uncovering new neuronal regulators of behavior.

Ranavirus infection-induced avoidance behaviour in wood frog juveniles: do amphibians socially distance?

Hosts may limit exposure to pathogens through changes in behaviour, such as avoiding infected individuals or contaminated areas. Here, we tested for a behavioural response to ranavirus infection in juvenile wood frogs (Rana sylvatica) because the majority of dispersal between populations occurs during this life stage. We hypothesized that if infections are transmissible and detectable at this life stage, then susceptibles would display avoidance behaviours when introduced to an infected conspecific. Despite no apparent signs of infection, we observed a greater distance between susceptible-infected pairs, compared to pairs of either two infected or two susceptible animals. Further, distances between susceptible-infected pairs were positively related to the infection intensity of the focal exposed frog, suggesting the cue to avoid infected conspecifics may become more detectable with more intense infections. Although we did not quantify whether the transmission was affected by their distancing, our findings suggest that juvenile frogs have the potential to reduce terrestrial transmission of ranaviruses through avoidance behaviours.

Danger appraisal and pathogen-avoidance mechanisms in stigma towards severe mental illness: the mediating role of affective responses.

BACKGROUND: Stereotypes of dangerousness are common predictors of stigmatising attitudes towards Severe Mental Illness (SMI). However less is known about pathogen avoidance mechanisms underlying stigma towards SMI, specially in samples of non-industrialised societies of Latin America and the Caribbean. The primary aim of this study was to examine pathogen-disgust sensitivity and danger appraisal mechanisms in responses of stigma towards SMI. METHODS: Cross-sectional design with convenience sampling. Using an online survey, volunteers at the Universidad del Norte in Colombia (N = 271) provided their sociodemographic data and completed the Three-Domain Disgust Scale (TDDS). Participants were randomised to different descriptions of someone with SMI that varied in terms of aggressiveness (with and without danger) and causes of the SMI. Then, following the attribution questionnaire (AQ-27), respondents reported affective and discriminatory responses to the person in the description. RESULTS: Increased disgust sensitivity to pathogen stimuli resulted in stronger reports of anger (ß = .14; p = .03), and fear (ß = 0.27; p < 0.001). The relationship between disgust sensitivity and discriminatory responses was indirectly mediated by fear towards SMI (Bootstrapped CI =-.04,-.009). Dangerousness attributions in the description of SMI predicted stronger feelings of anger (ß = .23; p = 0.001) and fear (ß = .40; p < .001), as well increased support for coercion-segregation of SMI (ß = .34; p = 0.04), but less intentions to help (ß = -.26; p = 0.003). The relationship between dangerousness and support for coercion was mediated by fear (Bootstrapped CI = .72, 1.37) and anger (Bootstrapped CI = .06, .44), whereas pity (Bootstrapped CI = .03, .38) and fear (Bootstrapped CI = -1.39, -.69) mediated responses of support for coercion-segregation of SMI. Attributions about causes and personal responsibility were not significantly linked to stigma towards SMI (p > 0.05). CONCLUSIONS: Findings suggested that pathogen avoidance and danger appraisal systems interplay in the generation of discriminatory behaviour towards SMI. Anti-stigma programs and policy makers would benefit from introducing strategies that challenge stereotypes of dangerousness and unpredictability by promoting positive contact with people with SMI.

Lack of avoidance of the fungal entomopathogen, Metarhizium brunneum, by male Agriotes obscurus beetles.

Fungal entomopathogens can greatly reduce the fitness of their hosts, and it is therefore expected that susceptible insects will be selected to avoid exposure to pathogens. Metarhizium brunneum is a fungal pathogen that can infect Agriotes obscurus, which in its larval form is a destructive agricultural pest and is repelled by the presence of M. brunneum conidia. Due to the subterranean nature of larval A. obscurus, recent research has focused on targeting adult A. obscurus with M. brunneum. No-choice and choice behavioural assays were conducted to determine if male adult A. obscurus avoid M. brunneum mycosed cadavers, or conidia applied to either food or soil. To further investigate the response of A. obscurus beetles to conspecific cadavers, the movement and behaviour of beetles placed at the centre of a semi-circular arrangement of mycosed or control cadavers was examined using motion tracking software. We found little evidence to suggest that A. obscurus male beetles avoid M. brunneum conidia or mycosed conspecific cadavers or alter their behaviour in their presence.

Control over pathogen exposure and basal immunological activity influence disgust and pathogen-avoidance motivation.

Disgust is reasoned to operate in conjunction with the immune system to help protect the body from illness. However, less is known about the factors that impact the degree to which individuals invest in pathogen avoidance (disgust) versus pathogen management (prophylactic immunological activity). Here, we examine the role that one's control over pathogen contact plays in resolving such investment trade-offs, predicting that (a) those from low control environments will invest less in pathogen-avoidance strategies and (b) investment in each of these two strategies will occur in a compensatory fashion (i.e. they will be traded off with one other). Across four studies, we found support for these predictions, using a variety of manipulations and measures. By providing novel insights into how one's control over pathogen exposure influences disgust sensitivity and immune system activity, the current research poses an important contribution to the literature on disgust, pathogen avoidance, and the immune system.

Keep your (social) distance: Pathogen concerns and social perception in the time of COVID-19.

Previous research suggests that individual differences in pathogen disgust sensitivity and social anxiety predict avoidance behavior, especially of pathogen cues, and reduced tolerance for social ambiguity. Conversely, generalized social trust is associated with approach behavior and a greater tolerance for social ambiguity. We conducted an online study (N = 1078) to test these predictions in the context of the COVID-19 global pandemic. Specifically, we assessed whether individual differences in pathogen disgust sensitivity, social anxiety and generalized social trust predicted judgments of trustworthiness, desired social distance and perceptions of sickness of target faces wearing surgical masks. Our results showed that (a) high sensitivity to pathogen disgust predicted lower judgments of trustworthiness and lower social desirability; (b) high social anxiety predicted higher perceptions of illness and lower judgments of trustworthiness; and (c) generalized social trust predicted higher judgments of trustworthiness and lower perceptions of illness of target faces. Further, we found that mask wearers were perceived as more likely to be ill, more trustworthy and more socially desirable than the same faces presented to a control group, without the surgical mask superimposed. Results are discussed in terms of perceived compliance with an emerging social norm overriding the intrinsic untrustworthiness of masked faces.

Evidence from hunter-gatherer and subsistence agricultural populations for the universality of contagion sensitivity.

The phenomenon of magical contagion - the unobserved passage of properties between entities that come into physical contact - was described by anthropologists over a century ago, yet questions remain about its origin, function, and universality. Contagion sensitivity, along with the emotion of disgust, has been proposed to be part of a biologically-evolved system designed to reduce exposure to pathogens by increasing the avoidance of "contaminated" objects. Yet this phenomenon has not been studied using systematic psychological comparison outside of industrialized populations. Here we document contagion sensitivity in two culturally, geographically, and economically distinct populations with little exposure to Western biomedicine and formal education: the Hadza hunter-gatherers of Tanzania and Tannese subsistence-agriculturalists of Vanuatu. In both populations, a majority of individuals rejected familiar and palatable foods when contaminating items touched the food but were subsequently removed. The Tannese children in our study showed a similar response, consistent with previous research with Western children. Our data support the proposal that contagion sensitivity is universal in human populations.

Healthy Out-Group Members Are Represented Psychologically as Infected In-Group Members.

A range of studies have demonstrated that people implicitly treat out-groups as the carriers of pathogens and that considerable prejudice against out-groups is driven by concerns about pathogens. Yet the psychological categories that are involved and the selection pressures that underlie these categories remain unclear. A common view is that human pathogen-avoidance psychology is specifically adapted to avoid out-groups because of their potentially different pathogens. However, the series of studies reported here shows that there is no dedicated category for reasoning about out-groups in terms of pathogens. Specifically, a memory-confusion experiment conducted with two large-scale samples of Americans (one nationally representative) yielded strong, replicable evidence that healthy out-group members are represented using the same psychological category that is used to represent manifestly infected in-group members. This suggests that the link between out-group prejudice and pathogen concerns is a by-product of general mechanisms for treating any unfamiliar appearance as an infection cue.

Contrasting invertebrate immune defense behaviors caused by a single gene, the Caenorhabditis elegans neuropeptide receptor gene npr-1.

BACKGROUND: The invertebrate immune system comprises physiological mechanisms, physical barriers and also behavioral responses. It is generally related to the vertebrate innate immune system and widely believed to provide nonspecific defense against pathogens, whereby the response to different pathogen types is usually mediated by distinct signalling cascades. Recent work suggests that invertebrate immune defense can be more specific at least at the phenotypic level. The underlying genetic mechanisms are as yet poorly understood. RESULTS: We demonstrate in the model invertebrate Caenorhabditis elegans that a single gene, a homolog of the mammalian neuropeptide Y receptor gene, npr-1, mediates contrasting defense phenotypes towards two distinct pathogens, the Gram-positive Bacillus thuringiensis and the Gram-negative Pseudomonas aeruginosa. Our findings are based on combining quantitative trait loci (QTLs) analysis with functional genetic analysis and RNAseq-based transcriptomics. The QTL analysis focused on behavioral immune defense against B. thuringiensis, using recombinant inbred lines (RILs) and introgression lines (ILs). It revealed several defense QTLs, including one on chromosome X comprising the npr-1 gene. The wildtype N2 allele for the latter QTL was associated with reduced defense against B. thuringiensis and thus produced an opposite phenotype to that previously reported for the N2 npr-1 allele against P. aeruginosa. Analysis of npr-1 mutants confirmed these contrasting immune phenotypes for both avoidance behavior and nematode survival. Subsequent transcriptional profiling of C. elegans wildtype and npr-1 mutant suggested that npr-1 mediates defense against both pathogens through p38 MAPK signaling, insulin-like signaling, and C-type lectins. Importantly, increased defense towards P. aeruginosa seems to be additionally influenced through the induction of oxidative stress genes and activation of GATA transcription factors, while the repression of oxidative stress genes combined with activation of Ebox transcription factors appears to enhance susceptibility to B. thuringiensis. CONCLUSIONS: Our findings highlight the role of a single gene, npr-1, in fine-tuning nematode immune defense, showing the ability of the invertebrate immune system to produce highly specialized and potentially opposing immune responses via single regulatory genes.

Fruit flies learn to avoid odours associated with virulent infection.

While learning to avoid toxic food is common in mammals and occurs in some insects, learning to avoid cues associated with infectious pathogens has received little attention. We demonstrate that Drosophila melanogaster show olfactory learning in response to infection with their virulent intestinal pathogen Pseudomonas entomophila. This pathogen was not aversive to taste when added to food. Nonetheless, flies exposed for 3 h to food laced with P. entomophila, and scented with an odorant, became subsequently less likely to choose this odorant than flies exposed to pathogen-laced food scented with another odorant. No such effect occurred after an otherwise identical treatment with an avirulent mutant of P. entomophila, indicating that the response is mediated by pathogen virulence. These results demonstrate that a virulent pathogen infection can act as an aversive unconditioned stimulus which flies can associate with food odours, and thus become less attracted to pathogen-contaminated food.

Loss of cilia in chemosensory neurons inhibits pathogen avoidance in Caenorhabditis elegans.

Pathogen avoidance is a crucial and evolutionarily conserved behavior that enhances survival by preventing infection in diverse species, including Caenorhabditis elegans (C. elegans). This behavior relies on multiple chemosensory neurons equipped with cilia that are exposed to the external environment. However, the specific role of neuronal cilia in pathogen avoidance has not been completely elucidated. Herein, we discovered that osm-3(p802) mutants, which lack chemosensory neuronal cilia, exhibit slower avoidance of the pathogen Pseudomonas aeruginosa PA14, but not Escherichia coli OP50. This observation was consistent when osm-3(p802) mutants were exposed to P. aeruginosa PAO1. Following an encounter with PA14, the pumping, thrashing, and defecation behaviors of osm-3 mutants were comparable to those of the wild-type. However, the osm-3 mutants demonstrated reduced intestinal colonization of PA14, suggesting that they have stronger intestinal clearance ability. We conducted RNA-seq to identify genes responding to external stimuli that were differentially expressed owing to the loss of osm-3 and PA14 infection. Using RNAi, we demonstrated that three of these genes were essential for normal pathogen avoidance. In conclusion, our findings demonstrate that the loss of chemosensory neuronal cilia reduces pathogen avoidance in C. elegans while delaying intestinal colonization.

Neuronal basis and diverse mechanisms of pathogen avoidance in Caenorhabditis elegans.

Pathogen avoidance behaviour has been observed across animal taxa as a vital host-microbe interaction mechanism. The nematode Caenorhabditis elegans has evolved multiple diverse mechanisms for pathogen avoidance under natural selection pressure. We summarise the current knowledge of the stimuli that trigger pathogen avoidance, including alterations in aerotaxis, intestinal bloating, and metabolites. We then survey the neural circuits involved in pathogen avoidance, transgenerational epigenetic inheritance of pathogen avoidance, signalling crosstalk between pathogen avoidance and innate immunity, and C. elegans avoidance of non-Pseudomonas bacteria. In this review, we highlight the latest advances in understanding host-microbe interactions and the gut-brain axis.

Genetic variation in parasite avoidance, yet no evidence for constitutive fitness costs.

Behavioral avoidance of parasites is a widespread strategy among animal hosts and in human public health. Avoidance has repercussions for both individual and population-level infection risk. Although most cases of parasite avoidance are viewed as adaptive, there is little evidence that the basic assumptions of evolution by natural selection are met. This study addresses this gap by testing whether there is a heritable variation in parasite avoidance behavior. We quantified behavioral avoidance of the bacterial parasite Serratia marcescens for 12 strains of the nematode host Caenorhabditis elegans. We found that these strains varied in their magnitude of avoidance, and we estimated the broad-sense heritability of this behavior to be in the range of 11%-26%. We then asked whether avoidance carries a constitutive fitness cost. We did not find evidence of one. Rather, strains with higher avoidance had higher fitness, measured as population growth rate. Together, these results direct future theoretical and empirical work to identify the forces maintaining genetic variation in parasite avoidance.

Antagonism between neuropeptides and monoamines in a distributed circuit for pathogen avoidance.

Pathogenic infection elicits behaviors that promote recovery and survival of the host. After exposure to the pathogenic bacterium Pseudomonas aeruginosa PA14, the nematode Caenorhabditis elegans modifies its sensory preferences to avoid the pathogen. Here, we identify antagonistic neuromodulators that shape this acquired avoidance behavior. Using an unbiased cell-directed neuropeptide screen, we show that AVK neurons upregulate and release RF/RYamide FLP-1 neuropeptides during infection to drive pathogen avoidance. Manipulations that increase or decrease AVK activity accelerate or delay pathogen avoidance, respectively, implicating AVK in the dynamics of avoidance behavior. FLP-1 neuropeptides drive pathogen avoidance through the G protein-coupled receptor DMSR-7, as well as other receptors. DMSR-7 in turn acts in multiple neurons, including tyraminergic/octopaminergic neurons that receive convergent avoidance signals from the cytokine DAF-7/transforming growth factor ß. Neuromodulators shape pathogen avoidance through multiple mechanisms and targets, in agreement with the distributed neuromodulatory connectome of C. elegans.

Do Experimental Manipulations of Pathogen Avoidance Motivations Influence Conformity?

By conforming to ingroup norms, individuals coordinate with other group members, preserve cohesion, and avoid costs of exclusion. Previous experiments have shown that increased concerns about infectious disease increase conformity. However, coordination with other group members has multiple benefits, most of which exist independent of pathogenic infection. Hence, a strong causal effect of pathogen avoidance motivations on conformity seems unlikely. Results from five experiments (N = 1,931) showed only limited support for the hypothesis that experimentally increasing pathogen avoidance motivations influences conformity. Overall, our findings are not consistent with the notion that the human mind contains a fast-acting psychological mechanism that regulates conformity as a function of short-term pathogen avoidance motivations.

First impressions of a new face are shaped by infection concerns.

Along with a classical immune system, we have evolved a behavioral one that directs us away from potentially contagious individuals. Here I show, using publicly available cross-cultural data, that this adaptation is so fundamental that our first impressions of a male stranger are largely driven by the perceived health of his face. Positive (likeable, capable, intelligent, trustworthy) and negative (unfriendly, ignorant, lazy) first impressions are affected by facial health in adaptively different ways, inconsistent with a mere halo effect; they are also modulated by one's current state of health and inclination to feel disgusted by pathogens. These findings, which replicated across two countries as different as the USA and India, suggest that instinctive perceptions of badness and goodness from faces are not two sides of the same coin but reflect the (nonsymmetrical) expected costs and benefits of interaction. Apparently, pathogens run the show-and first impressions come second. Lay Summary: Our first impressions of strangers (whether they seem trustworthy, intelligent, unfriendly, or aggressive) are shaped by how healthy their faces look and by our unconscious motivation to avoid infections. Bad and good impressions turn out to reflect the concrete, potentially vital, expected costs and benefits of interacting with our fellow humans. Apparently, pathogens run the show-and first impressions come second.