{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,6,23]],"date-time":"2026-06-23T17:22:30Z","timestamp":1782235350639,"version":"3.54.5"},"reference-count":77,"publisher":"MDPI AG","issue":"21","license":[{"start":{"date-parts":[[2025,11,5]],"date-time":"2025-11-05T00:00:00Z","timestamp":1762300800000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100015564","name":"Universit\u00e0 Italo-Francese","doi-asserted-by":"crossref","id":[{"id":"10.13039\/100015564","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Animals"],"abstract":"<jats:p>Current research on acoustic encoding of emotional content suggests that there are universal cues, allowing for decoding within and across taxa. This is particularly important for human\u2013animal relationships, wherein domestic animals are supposed to be particularly efficient in decoding human emotions. Here we investigated whether the decoding of the emotional content in human voices shared universal acoustic properties, or whether it could be influenced by experience. Emotional human voices were presented to two populations of horses, in which behavioral, cardiac, and brain responses were measured. The two populations differed in their living and working conditions: one population lived in naturalistic conditions (stable social groups in pastures) and were ridden occasionally for outdoor trail riding with one to a few different riders, while the other was kept in more restricted conditions (individual stalls) and participated in riding lessons involving many different riders. Assessment of the horses\u2019 welfare state (animal-based measures) and their relationships with humans, performed independently of the playback experiments, revealed that the populations differed in both aspects. Whereas both populations appeared to react to the angry human voice, the population with the best welfare state and relationship with humans showed little differentiation between the different emotional voices and exhibited low behavioral reactions. On the contrary, the other population showed high behavioral and cardiac reactions to all negative voices. Brain responses also differed, with the first population showing higher responses (increased gamma, i.e., excitation) for the happy voice and the second for fear and anger (increased theta, i.e., alarm). Thus, animals\u2019 affective state and past experiences appear very influential for their perception of (cross-taxa) acoustic emotional cues.<\/jats:p>","DOI":"10.3390\/ani15213217","type":"journal-article","created":{"date-parts":[[2025,11,5]],"date-time":"2025-11-05T16:09:47Z","timestamp":1762358987000},"page":"3217","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Chronic State and Relationship to Humans Influence How Horses Decode Emotions in Human Voices: A Brain and Behavior Study"],"prefix":"10.3390","volume":"15","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-6164-2598","authenticated-orcid":false,"given":"Serenella","family":"d\u2019Ingeo","sequence":"first","affiliation":[{"name":"Animal Physiology and Behavior Research Unit, Department of Veterinary Medicine, University of Bari Aldo Moro, 70121 Bari, Italy"},{"name":"Laboratoire Ethologie Animale et Humaine-EthoS, UMR 6552-CNRS, University of Rennes, 35042 Rennes Cedex, France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1592-6549","authenticated-orcid":false,"given":"Marcello","family":"Siniscalchi","sequence":"additional","affiliation":[{"name":"Animal Physiology and Behavior Research Unit, Department of Veterinary Medicine, University of Bari Aldo Moro, 70121 Bari, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7835-2485","authenticated-orcid":false,"given":"Angelo","family":"Quaranta","sequence":"additional","affiliation":[{"name":"Animal Physiology and Behavior Research Unit, Department of Veterinary Medicine, University of Bari Aldo Moro, 70121 Bari, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hugo","family":"Cousillas","sequence":"additional","affiliation":[{"name":"Laboratoire Ethologie Animale et Humaine-EthoS, UMR 6552-CNRS, University of Rennes, 35042 Rennes Cedex, France"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4867-5408","authenticated-orcid":false,"given":"Martine","family":"Hausberger","sequence":"additional","affiliation":[{"name":"Integrative Neuroscience and Cognition Center, UMR 8002-CNRS, University of Paris-Cit\u00e9, 75006 Paris, France"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2025,11,5]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"105282","DOI":"10.1016\/j.applanim.2021.105282","article-title":"Lateralized emotional functioning in domestic animals","volume":"237","author":"Siniscalchi","year":"2021","journal-title":"Appl. Anim. Behav. Sci."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Maigrot, A.-L., Hillmann, E., and Briefer, E.F. (2022). Cross-species discrimination of vocal expression of emotional valence by Equidae and Suidae. BMC Biol., 20.","DOI":"10.1186\/s12915-022-01311-5"},{"key":"ref_3","doi-asserted-by":"crossref","unstructured":"Filippi, P., Congdon, J.V., Hoang, J., Bowling, D.L., Reber, S.A., Pa\u0161ukonis, A., Hoeschele, M., Ocklenburg, S., de Boer, B., and Sturdy, C.B. (2017). Humans recognize emotional arousal in vocalizations across all classes of terrestrial vertebrates: Evidence for acoustic universals. Proc. R. Soc. B Biol. Sci., 284.","DOI":"10.1098\/rspb.2017.0990"},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"2895","DOI":"10.1098\/rspb.2010.0303","article-title":"An integrative and functional framework for the study of animal emotion and mood","volume":"277","author":"Mendl","year":"2010","journal-title":"Proc. R. Soc. B Biol. Sci."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1037\/0033-295X.110.1.145","article-title":"Core affect and the psychological construction of emotion","volume":"110","author":"Russell","year":"2003","journal-title":"Psychol. Rev."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"855","DOI":"10.1086\/283219","article-title":"On the Occurrence and Significance of Motivation-Structural Rules in Some Bird and Mammal Sounds","volume":"111","author":"Morton","year":"1977","journal-title":"Am. Nat."},{"key":"ref_7","doi-asserted-by":"crossref","unstructured":"Siniscalchi, M., d\u2019Ingeo, S., Fornelli, S., and Quaranta, A. (2018). Lateralized behavior and cardiac activity of dogs in response to human emotional vocalizations. Sci. Rep., 8.","DOI":"10.1038\/s41598-017-18417-4"},{"key":"ref_8","doi-asserted-by":"crossref","unstructured":"Quaranta, A., d\u2019Ingeo, S., Amoruso, R., and Siniscalchi, M. (2020). Emotion recognition in cats. Animals, 10.","DOI":"10.3390\/ani10071107"},{"key":"ref_9","doi-asserted-by":"crossref","unstructured":"Smith, A.V., Proops, L., Grounds, K., Wathan, J., and McComb, K. (2018). Domestic horses (Equus caballus) discriminate between negative and positive human nonverbal vocalisations. Sci. Rep., 8.","DOI":"10.1038\/s41598-018-30777-z"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"180491","DOI":"10.1098\/rsos.180491","article-title":"Goats prefer positive human emotional facial expressions","volume":"5","author":"Nawroth","year":"2018","journal-title":"R. Soc. Open Sci."},{"key":"ref_11","doi-asserted-by":"crossref","unstructured":"Tr\u00f6sch, M., Cuzol, F., Parias, C., Calandreau, L., Nowak, R., and Lansade, L. (2019). Horses categorize human emotions cross-modally based on facial expression and non-verbal vocalizations. Animals, 9.","DOI":"10.3390\/ani9110862"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"369","DOI":"10.1007\/s10071-021-01557-6","article-title":"Cognition and the human-animal relationship: A review of the sociocognitive skills of domestic mammals toward humans","volume":"25","author":"Jardat","year":"2021","journal-title":"Anim. Cogn."},{"key":"ref_13","doi-asserted-by":"crossref","unstructured":"Nakamura, K., Takimoto-Inose, A., and Hasegawa, T. (2018). Cross-modal perception of human emotion in domestic horses (Equus caballus). Sci. Rep., 8.","DOI":"10.1038\/s41598-018-26892-6"},{"key":"ref_14","doi-asserted-by":"crossref","unstructured":"Barber, A.L.A., Randi, D., M\u00fcller, C.A., and Huber, L. (2016). The processing of human emotional faces by pet and lab dogs: Evidence for lateralization and experience effects. PLoS ONE, 11.","DOI":"10.1371\/journal.pone.0152393"},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"210656","DOI":"10.1098\/rsos.210656","article-title":"Performance of goats in a detour and a problem-solving test following long-term cognitive test exposure","volume":"8","author":"Rosenberger","year":"2021","journal-title":"R. Soc. Open Sci."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"146","DOI":"10.1016\/j.applanim.2011.12.007","article-title":"Housing induced mood modulates reactions to emotional stimuli in sheep","volume":"136","author":"Reefmann","year":"2012","journal-title":"Appl. Anim. Behav. Sci."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"217","DOI":"10.1007\/s10071-021-01543-y","article-title":"Who is there? Captive western gorillas distinguish human voices based on familiarity and nature of previous interactions","volume":"25","author":"Salmi","year":"2021","journal-title":"Anim. Cogn."},{"key":"ref_18","doi-asserted-by":"crossref","unstructured":"D\u2019Ingeo, S., Quaranta, A., Siniscalchi, M., Stomp, M., Coste, C., Bagnard, C., Hausberger, M., and Cousillas, H. (2019). Horses associate individual human voices with the valence of past interactions: A behavioural and electrophysiological study. Sci. Rep., 9.","DOI":"10.1038\/s41598-019-47960-5"},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"339","DOI":"10.1016\/S0168-1591(02)00016-3","article-title":"A brief note on some possible factors involved in the reactions of horses to humans","volume":"76","author":"Hausberger","year":"2002","journal-title":"Appl. Anim. Behav. Sci."},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Sankey, C., Henry, S., Andr\u00e9, N., Richard-Yris, M.-A., and Hausberger, M. (2011). Do horses have a concept of person?. PLoS ONE, 6.","DOI":"10.1371\/journal.pone.0018331"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"643","DOI":"10.1007\/s10071-009-0223-2","article-title":"How horses (Equus caballus) see the world: Humans as significant \u201cobjects\u201d","volume":"12","author":"Fureix","year":"2009","journal-title":"Anim. Cogn."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"469","DOI":"10.1016\/j.neubiorev.2005.01.002","article-title":"Measuring emotional processes in animals: The utility of a cognitive approach","volume":"29","author":"Paul","year":"2005","journal-title":"Neurosci. Biobehav. Rev."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"312","DOI":"10.1038\/427312a","article-title":"Cognitive bias and affective state","volume":"427","author":"Harding","year":"2004","journal-title":"Nature"},{"key":"ref_24","doi-asserted-by":"crossref","unstructured":"Brajon, S., Laforest, J.-P., Schmitt, O., and Devillers, N. (2015). The way humans behave modulates the emotional state of piglets. PLoS ONE, 10.","DOI":"10.1371\/journal.pone.0133408"},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1016\/j.applanim.2015.08.036","article-title":"What can inactivity (in its various forms) reveal about affective states in non-human animals? A review","volume":"171","author":"Fureix","year":"2015","journal-title":"Appl. Anim. Behav. Sci."},{"key":"ref_26","doi-asserted-by":"crossref","unstructured":"Scheumann, M., Hasting, A.S., Kotz, S.A., and Zimmermann, E. (2014). The Voice of Emotion across Species: How Do Human Listeners Recognize Animals\u2019 Affective States?. PLoS ONE, 9.","DOI":"10.1371\/journal.pone.0091192"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"199","DOI":"10.1016\/j.ijpsycho.2004.07.004","article-title":"Changes in EEG and autonomic nervous activity during meditation and their association with personality traits","volume":"55","author":"Takahashi","year":"2005","journal-title":"Int. J. Psychophysiol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"73","DOI":"10.1016\/j.biopsycho.2009.10.008","article-title":"EEG theta\/beta ratio in relation to fear-modulated response-inhibition, attentional control, and affective traits","volume":"83","author":"Putman","year":"2010","journal-title":"Biol. Psychol."},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"293","DOI":"10.1111\/j.1469-8986.2007.00497.x","article-title":"Music and emotion: Electrophysiological correlates of the processing of pleasant and unpleasant music","volume":"44","author":"Sammler","year":"2007","journal-title":"Psychophysiology"},{"key":"ref_30","unstructured":"Davidson, R.J. (1995). Cerebral Asymmetry, Emotion, and Affective Style in Brain Asymmetry, MIT Press."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"487","DOI":"10.1080\/02699930126048","article-title":"Frontal brain electrical activity (EEG) distinguishes valence and intensity of musical emotions","volume":"15","author":"Schmidt","year":"2001","journal-title":"Cogn. Emot."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"297","DOI":"10.1080\/17470919.2015.1081400","article-title":"Vocal and visual stimulation, congruence and lateralization affect brain oscillations in interspecies emotional positive and negative interactions","volume":"11","author":"Balconi","year":"2016","journal-title":"J. Soc. Neuro."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"377","DOI":"10.1016\/j.neubiorev.2006.10.004","article-title":"Motivation, emotion, and their inhibitory control mirrored in brain oscillations","volume":"31","author":"Knyazev","year":"2007","journal-title":"Neurosci. Biobehav. Rev."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1802","DOI":"10.1016\/j.clinph.2004.03.009","article-title":"Cognitive tasks augment gamma EEG power","volume":"115","author":"Fitzgibbon","year":"2004","journal-title":"Clin. Neurophysiol."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"57","DOI":"10.1016\/S0304-3940(01)02094-8","article-title":"Human anterior and frontal midline theta and lower alpha reflect emotionally positive state and internalized attention: High-resolution EEG investigation of meditation","volume":"310","author":"Aftanas","year":"2001","journal-title":"Neurosci. Lett."},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"214","DOI":"10.1111\/j.1467-9450.1988.tb00794.x","article-title":"Immobility-related hippocampal theta activity in rats during reaction to novelty","volume":"29","author":"Myhrer","year":"1988","journal-title":"Scand. J. Psychol."},{"key":"ref_37","doi-asserted-by":"crossref","unstructured":"Hausberger, M., Cousillas, H., Meter, A., Karino, G., George, I., Lemasson, A., and Blois-Heulin, C. (2019). A crucial role of attention in lateralisation of sound processing?. Symmetry, 11.","DOI":"10.3390\/sym11010048"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"8","DOI":"10.1007\/s00114-016-1429-1","article-title":"Do horses with poor welfare show \u2018pessimistic\u2019 cognitive biases?","volume":"104","author":"Henry","year":"2017","journal-title":"Sci. Nat."},{"key":"ref_39","doi-asserted-by":"crossref","unstructured":"Lesimple, C. (2020). Indicators of Horse Welfare: State-of-the-Art. Animals, 10.","DOI":"10.3390\/ani10020294"},{"key":"ref_40","unstructured":"(2018). Saisie n\u00b0 2016-SA-0288 (Standard No. ANSES 2018)."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"247","DOI":"10.1016\/j.biopsycho.2007.11.006","article-title":"Affective Picture Processing: An Integrative Review of ERP Findings","volume":"77","author":"Olofsson","year":"2008","journal-title":"Biol. Psychol."},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"200851","DOI":"10.1098\/rsos.200851","article-title":"Event-Related Potentials Reveal Limited Readiness to Access Phonetic Details during Word Processing in Dogs","volume":"7","author":"Magyari","year":"2020","journal-title":"R. Soc. Open Sci."},{"key":"ref_43","doi-asserted-by":"crossref","unstructured":"Cousillas, H., Oger, M., Rochais, C., Pettoello, C., M\u00e9noret, M., Henry, S., and Hausberger, M. (2017). An ambulatory electroencephalography system for freely moving horses: An innovating approach. Front. Vet. Sci., 4.","DOI":"10.3389\/fvets.2017.00057"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"105271","DOI":"10.1016\/j.applanim.2021.105271","article-title":"Brain activity reflects (chronic) welfare state: Evidence from individual electroencephalography profiles in an animal model","volume":"236","author":"Stomp","year":"2021","journal-title":"Appl. Anim. Behav. Sci."},{"key":"ref_45","doi-asserted-by":"crossref","unstructured":"Rochais, C., S\u00e9billeau, M., Menoret, M., Oger, M., Henry, S., Hausberger, M., and Cousillas, H. (2018). Attentional state and brain processes: State-dependent lateralization of EEG profiles in horses. Sci. Rep., 8.","DOI":"10.1038\/s41598-018-28334-9"},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"611","DOI":"10.1007\/s10071-009-0220-5","article-title":"Socially dependent auditory laterality in domestic horses (Equus caballus)","volume":"12","author":"Basile","year":"2009","journal-title":"Anim. Cogn."},{"key":"ref_47","unstructured":"Waring, G.H. (2002). Horse Behavior, William Andrew."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1163\/156853974X00534","article-title":"Observational study of behavior: Sampling methods","volume":"49","author":"Altman","year":"1974","journal-title":"Behaviour"},{"key":"ref_49","doi-asserted-by":"crossref","unstructured":"Stomp, M., D\u2019iNgeo, S., Henry, S., Lesimple, C., Cousillas, H., and Hausberger, M. (2020). EEG individual power profiles correlate with tension along spine in horses. PLoS ONE, 15.","DOI":"10.1371\/journal.pone.0243970"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"291","DOI":"10.1007\/s00114-012-0902-8","article-title":"On the Significance of Adult Play: What Does Social Play Tell Us about Adult Horse Welfare?","volume":"99","author":"Hausberger","year":"2012","journal-title":"Naturwissenschaften"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1016\/j.tree.2008.10.008","article-title":"Generalized linear mixed models: A practical guide for ecology and evolution","volume":"24","author":"Bolker","year":"2009","journal-title":"Trends Ecol. Evol."},{"key":"ref_52","doi-asserted-by":"crossref","unstructured":"Albuquerque, N., Guo, K., Wilkinson, A., Savalli, C., Otta, E., and Mills, D. (2016). Dogs recognize dog and human emotions. Biol. Lett., 12.","DOI":"10.1098\/rsbl.2015.0883"},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"44","DOI":"10.1037\/0735-7036.117.1.44","article-title":"Classification of domestic cat (Felis catus) vocalizations by naive and experienced human listeners","volume":"117","author":"Nicastro","year":"2003","journal-title":"J. Comp. Psychol."},{"key":"ref_54","first-page":"87","article-title":"Owner\u2019s beliefs on the ability of their pet dogs to understand human verbal communication: A case of social understanding","volume":"20","year":"2001","journal-title":"Cogn. Psychol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"81","DOI":"10.1037\/a0017354","article-title":"Human perception of vocalizations of domestic piglets and modulation by experience with domestic pigs (Sus scrofa)","volume":"124","author":"Tallet","year":"2010","journal-title":"J. Comp. Psychol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"653","DOI":"10.1080\/14640748808402292","article-title":"Anxiety and the allocation of attention to threat","volume":"40","author":"MacLeod","year":"1988","journal-title":"Q. J. Exp. Psychol."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"359","DOI":"10.1007\/BF01183287","article-title":"Visual attention and depression: Cognitive biases in the eye fixations of the dysphoric and the nondepressed","volume":"16","author":"Matthews","year":"1992","journal-title":"Cogn. Ther. Res."},{"key":"ref_58","doi-asserted-by":"crossref","unstructured":"Fureix, C., Jego, P., Henry, S., Lansade, L., and Hausberger, M. (2012). Towards an ethological animal model of Depression? A study on Horses. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0039280"},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1016\/j.physbeh.2007.08.001","article-title":"Removing individual rats affects indicators of welfare in the remaining group members","volume":"93","author":"Burman","year":"2008","journal-title":"Physiol. Behav."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.applanim.2007.04.015","article-title":"A review of the human\u2013horse relationship","volume":"109","author":"Hausberger","year":"2007","journal-title":"Appl. Anim. Behav. Sci."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"869","DOI":"10.1016\/j.anbehav.2009.12.037","article-title":"Positive Interactions Lead to Lasting Positive Memories in Horses, Equus caballus","volume":"79","author":"Sankey","year":"2010","journal-title":"Anim. Behav."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"118480","DOI":"10.1016\/j.neuroimage.2021.118480","article-title":"Social relationship-dependent neural response to speech in dogs","volume":"243","author":"Andics","year":"2021","journal-title":"Neuroimage"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1016\/j.anbehav.2023.12.008","article-title":"Goats discriminate emotional valence in the human voice","volume":"209","author":"Mason","year":"2024","journal-title":"Anim. Behav."},{"key":"ref_64","doi-asserted-by":"crossref","unstructured":"Lesimple, C., Fureix, C., Menguy, H., and Hausberger, M. (2010). Human Direct Actions May Alter Animal Welfare, a Study on Horses (Equus caballus). PLoS ONE, 5.","DOI":"10.1371\/annotation\/530e1439-fee8-434c-97b1-7d2cd87e2e46"},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"435","DOI":"10.1007\/s10071-012-0584-9","article-title":"Dogs\u2019 attention towards humans depends on their relationship, not only on social familiarity","volume":"16","author":"Horn","year":"2012","journal-title":"Anim. Cogn."},{"key":"ref_66","doi-asserted-by":"crossref","unstructured":"D\u2019Ingeo, S., Siniscalchi, M., Straziota, V., Ventriglia, G., Sasso, R., and Quaranta, A. (2023). Relationship between asymmetric nostril use and human emotional odours in cats. Sci. Rep., 13.","DOI":"10.1038\/s41598-023-38167-w"},{"key":"ref_67","doi-asserted-by":"crossref","unstructured":"Siniscalchi, M., d\u2019Ingeo, S., Minunno, M., and Quaranta, A. (2022). Facial asymmetry in dogs with fear and aggressive behaviors towards humans. Sci. Rep., 12.","DOI":"10.1038\/s41598-022-24136-2"},{"key":"ref_68","doi-asserted-by":"crossref","unstructured":"Rogers, L.J., Vallortigara, G., and Andrew, R.J. (2013). Divided Brains: The Biology and Behavior of Brain Asymmetries, Cambridge University Press.","DOI":"10.1017\/CBO9780511793899"},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"34","DOI":"10.1016\/j.bbr.2016.02.011","article-title":"The dog nose \u201cKNOWS\u201d fear: Asymmetric nostril use during sniffing at canine and human emotional stimuli","volume":"304","author":"Siniscalchi","year":"2016","journal-title":"Behav. Brain Res."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"25","DOI":"10.1186\/1880-6805-33-21","article-title":"Time window for cognitive activity involved in emotional processing","volume":"33","author":"Motoi","year":"2014","journal-title":"J. Physiol. Anthropol."},{"key":"ref_71","doi-asserted-by":"crossref","unstructured":"Jardat, P., Destrez, A., Damon, F., Menard--Peroy, Z., Parias, C., Barri\u00e8re, P., Keller, M., Calandreau, L., and Lansade, L. (2023). Horses discriminate human body odors between fear and joy contexts in a habituation-discrimination protocol. Sci. Rep., 13.","DOI":"10.1038\/s41598-023-30119-8"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"1959","DOI":"10.1016\/j.neubiorev.2011.05.001","article-title":"Human brain EEG indices of emotions: Delineating responses to affective vocalizations by measuring frontal theta event-related synchronization","volume":"35","author":"Bekkedal","year":"2011","journal-title":"Neurosci. Biobehav. Rev."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"72","DOI":"10.1016\/j.brainres.2009.08.056","article-title":"Appetitive vs. defensive responses to emotional cues: Autonomic measures and brain oscillation modulation","volume":"1296","author":"Balconi","year":"2009","journal-title":"Brain Res."},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"174","DOI":"10.1016\/j.brainres.2008.06.108","article-title":"Anxiety and oscillatory responses to emotional facial expressions","volume":"1227","author":"Knyazev","year":"2008","journal-title":"Brain Res."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"e57313","DOI":"10.7554\/eLife.57313","article-title":"Different theta frameworks coexist in the rat hippocampus and are coordinated during memory-guided and novelty tasks","volume":"9","author":"Moratal","year":"2020","journal-title":"eLife"},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"209","DOI":"10.1016\/S0376-6357(97)00784-5","article-title":"Experimental tests to assess emotionality in horses","volume":"40","author":"Wolff","year":"1997","journal-title":"Behav. Processes"},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"70","DOI":"10.1016\/j.tvjl.2009.03.013","article-title":"Investigating horse\u2013human interactions: The effect of a nervous human","volume":"181","author":"Keeling","year":"2009","journal-title":"Vet. J."}],"container-title":["Animals"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2076-2615\/15\/21\/3217\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,11,7]],"date-time":"2025-11-07T05:11:34Z","timestamp":1762492294000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2076-2615\/15\/21\/3217"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2025,11,5]]},"references-count":77,"journal-issue":{"issue":"21","published-online":{"date-parts":[[2025,11]]}},"alternative-id":["ani15213217"],"URL":"https:\/\/doi.org\/10.3390\/ani15213217","relation":{},"ISSN":["2076-2615"],"issn-type":[{"value":"2076-2615","type":"electronic"}],"subject":[],"published":{"date-parts":[[2025,11,5]]}}}