Horses show individual level lateralisation when inspecting an unfamiliar and unexpected stimulus.
Abstract: Animals must attend to a diverse array of stimuli in their environments. The emotional valence and salience of a stimulus can affect how this information is processed in the brain. Many species preferentially attend to negatively valent stimuli using the sensory organs on the left side of their body and hence the right hemisphere of their brain. Here, we investigated the lateralisation of visual attention to the rapid appearance of a stimulus (an inflated balloon) designed to induce an avoidance reaction and a negatively valent emotional state in 77 Italian saddle horses. Horses' eyes are laterally positioned on the head, and each eye projects primarily to the contralateral hemisphere, allowing eye use to be a proxy for preferential processing in one hemisphere of the brain. We predicted that horses would inspect the novel and unexpected stimulus with their left eye and hence right hemisphere. We found that horses primarily inspected the balloon with one eye, and most horses had a preferred eye to do so, however, we did not find a population level tendency for this to be the left or the right eye. The strength of this preference tended to decrease over time, with the horses using their non-preferred eye to inspect the balloon increasingly as the trial progressed. Our results confirm a lateralised eye use tendency when viewing negatively emotionally valent stimuli in horses, in agreement with previous findings. However, there was not any alignment of lateralisation at the group level in our sample, suggesting that the expression of lateralisation in horses depends on the sample population and testing context.
Publication Date: 2021-08-05 PubMed ID: 34351986PubMed Central: PMC8341651DOI: 10.1371/journal.pone.0255688Google Scholar: Lookup
The Equine Research Bank provides access to a large database of publicly available scientific literature. Inclusion in the Research Bank does not imply endorsement of study methods or findings by Mad Barn.
- Journal Article
- Research Support
- Non-U.S. Gov't
Summary
This research summary has been generated with artificial intelligence and may contain errors and omissions. Refer to the original study to confirm details provided. Submit correction.
This research study focuses on the tendency of horses to use one eye more than the other when examining an unfamiliar and negative stimulus, in this case, an inflated balloon. There was no overall preference for left or right eye usage at the population level; the majority of the horses consistently used a specific eye to view the balloon, although this preference tended to decrease over time.
Research Context
- The research revolves around the concept of ‘lateralisation’, which refers to the preference of an animal to use one side of its body more than the other for certain tasks.
- The study highlights how this phenomenon occurs in horses when responding to an unexpected and unfamiliar stimulus, specifically an inflated balloon that would induce a negative emotional state and an avoidance reaction in the subjects.
Horses’ Visual Perception
- The unique positioning and physiology of a horse’s eyes, which are laterally positioned on the head and primarily project to the opposite hemisphere of the brain, was utilized as a measure for the study.
- This implies that the eye that a horse opts to use to inspect an object can indicate which brain hemisphere it prefers to process that particular information.
Research Findings
- The research observed that horses predominantly inspected the balloon using a single eye over the other, with most having a certain preference.
- The strength of this preference tended to decrease over time, as the horses used their non-preferred eye increasingly to inspect the balloon as the trial progressed.
- While individual horses displayed lateralized eye use, there was no overall preference for left or right eye use across the entire population of horses tested.
Implications and Conclusions
- The results align with previous findings, corroborating the existence of lateralized eye use when horses view negative stimuli.
- However, the lack of left or right eye preference at a population level indicates that lateralization in horses could significantly depend on the sample population and testing context.
- Overall, this study contributes to our understanding of how horses process environmental stimuli and may have potential implications in their training and welfare.
Cite This Article
APA
(2021).
Horses show individual level lateralisation when inspecting an unfamiliar and unexpected stimulus.
PLoS One, 16(8), e0255688.
https://doi.org/10.1371/journal.pone.0255688 Publication
Researcher Affiliations
MeSH Terms
- Animals
- Attention
- Emotions
- Eye Movements
- Functional Laterality
- Horses / physiology
- Horses / psychology
- Visual Perception
Conflict of Interest Statement
The authors have declared that no competing interests exist.
References
This article includes 84 references
- Rogers LJ, Vallortigara G. When and why did brains break symmetry?. Symmetry 2015; 7(4): 2181–2194.
- Cook ND. The brain code: Mechanisms of information transfer and the role of the corpus callosum. 1st ed. London: Routledge; 2018.
- Güntürkün O, Ocklenburg S. Ontogenesis of Lateralization.. Neuron 2017 Apr 19;94(2):249-263.
- Basile M, Boivin S, Boutin A, Blois-Heulin C, Hausberger M, Lemasson A. Socially dependent auditory laterality in domestic horses (Equus caballus).. Anim Cogn 2009 Jul;12(4):611-9.
- Parthasarathy K, Bhalla US. Laterality and symmetry in rat olfactory behavior and in physiology of olfactory input.. J Neurosci 2013 Mar 27;33(13):5750-60.
- van der Ham IJ, Postma A, Laeng B. Lateralized perception: the role of attention in spatial relation processing.. Neurosci Biobehav Rev 2014 Sep;45:142-8.
- Siniscalchi M, d’Ingeo S, Quaranta A. Lateralized emotional functioning in domestic animals. Applied Animal Behaviour Science 2021; 105282: 1–12.
- Andrew RJ, Watkins JAS. Evidence for cerebral lateralization from senses other than vision. In: Andrew RJ, Rogers LJ, editors. Comparative vertebrate lateralization. Cambridge: Cambridge University Press; 2002. pp 365–382.
- MacNeilage PF, Rogers LJ, Vallortigara G. Origins of the left & right brain.. Sci Am 2009 Jul;301(1):60-7.
- Rogers LJ. Relevance of brain and behavioural lateralization to animal welfare. Appl Anim Behav Sci 2010; 127(1–2): 1–11.
- Rogers LJ. Asymmetry of brain and behavior in animals: Its development, function, and human relevance.. Genesis 2014 Jun;52(6):555-71.
- Baijal S, Srinivasan N. Emotional and hemispheric asymmetries in shifts of attention: an ERP study.. Cogn Emot 2011 Feb;25(2):280-94.
- Bartolomeo P, Seidel Malkinson T. Hemispheric lateralization of attention processes in the human brain.. Curr Opin Psychol 2019 Oct;29:90-96.
- Chapelain AS, Blois-Heulin C. Lateralization for visual processes: eye preference in Campbell's monkeys (Cercopithecus c. campbelli).. Anim Cogn 2009 Jan;12(1):11-9.
- Braccini SN, Lambeth SP, Schapiro SJ, Fitch WT. Eye preferences in captive chimpanzees.. Anim Cogn 2012 Sep;15(5):971-8.
- Güntürkün O, Ströckens F, Ocklenburg S. Brain Lateralization: A Comparative Perspective.. Physiol Rev 2020 Jul 1;100(3):1019-1063.
- Karenina K, Giljov A, Baranov V, Osipova L, Krasnova V, Malashichev Y. Visual laterality of calf-mother interactions in wild whales.. PLoS One 2010 Nov 3;5(11):e13787.
- Siniscalchi M, Bertino D, d’Ingeo S, Quaranta A. Relationship between motor laterality and aggressive behavior in sheepdogs. Symmetry 2019; 11(2): 233.
- Schnell AK, Hanlon RT, Benkada A, Jozet-Alves C. Lateralization of Eye Use in Cuttlefish: Opposite Direction for Anti-Predatory and Predatory Behaviors.. Front Physiol 2016;7:620.
- Baciadonna L, Nawroth C, Briefer EF, McElligott AG. Perceptual lateralization of vocal stimuli in goats.. Curr Zool 2019 Feb;65(1):67-74.
- Galuret S, Lumineau S, Pouzol D, George I. Mothering influences domestic chick’s laterality. Anim Behav 2020; 159: 69–79.
- Miletto Petrazzini ME, Pecunioso A, Dadda M, Agrillo C. The Impact of Brain Lateralization and Anxiety-Like Behaviour in an Extensive Operant Conditioning Task in Zebrafish (Danio rerio). Symmetry 2019; 11(11): 1395.
- Halpern ME, Güntürkün O, Hopkins WD, Rogers LJ. Lateralization of the vertebrate brain: taking the side of model systems.. J Neurosci 2005 Nov 9;25(45):10351-7.
- Leliveld L. From Science to Practice: A Review of Laterality Research on Ungulate Livestock. Symmetry 2019; 11(9): 1157.
- Timney B, Macuda T. Vision and hearing in horses.. J Am Vet Med Assoc 2001 May 15;218(10):1567-74.
- Hanggi EB, Ingersoll JF. Lateral vision in horses: a behavioral investigation.. Behav Processes 2012 Sep;91(1):70-6.
- Schmidt MJ, Knemeyer C, Heinsen H. Neuroanatomy of the equine brain as revealed by high-field (3Tesla) magnetic-resonance-imaging.. PLoS One 2019;14(4):e0213814.
- Johnson PJ, Janvier V, Luh WM, FitzMaurice M, Southard T, Barry EF. Equine Stereotaxtic Population Average Brain Atlas With Neuroanatomic Correlation.. Front Neuroanat 2019;13:89.
- Rogers LJ, Andrew R. Comparative vertebrate lateralization. 1st ed. Cambridge: Cambridge University Press; 2002.
- Rogers LJ, Vallortigara G, Andrew RJ. Divided brains: the biology and behaviour of brain asymmetries. Cambridge: Cambridge University Press; 2013.
- Larose C, Richard-Yris MA, Hausberger M, Rogers LJ. Laterality of horses associated with emotionality in novel situations.. Laterality 2006 Jul;11(4):355-67.
- Austin NP, Rogers LJ. Asymmetry of flight and escape turning responses in horses.. Laterality 2007 Sep;12(5):464-74.
- Austin NP, Rogers LJ. Limb preferences and lateralization of aggression, reactivity and vigilance in feral horses, Equus caballus. Anim Behav 2012; 83: 239–247.
- Farmer K, Krueger K, Byrne RW. Visual laterality in the domestic horse (Equus caballus) interacting with humans.. Anim Cogn 2010 Mar;13(2):229-38.
- Sankey C, Henry S, Clouard C, Richard-Yris MA, Hausberger M. Asymmetry of behavioral responses to a human approach in young naive vs. trained horses.. Physiol Behav 2011 Sep 1;104(3):464-8.
- Smith AV, Proops L, Grounds K, Wathan J, McComb K. Functionally relevant responses to human facial expressions of emotion in the domestic horse (Equus caballus).. Biol Lett 2016 Feb;12(2):20150907.
- Phillips CJ, Oevermans H, Syrett KL, Jespersen AY, Pearce GP. Lateralization of behavior in dairy cows in response to conspecifics and novel persons.. J Dairy Sci 2015 Apr;98(4):2389-400.
- Camerlink I, Menneson S, Turner SP, Farish M, Arnott G. Lateralization influences contest behaviour in domestic pigs.. Sci Rep 2018 Aug 14;8(1):12116.
- Raoult CMC, Gygax L. Valence and Intensity of Video Stimuli of Dogs and Conspecifics in Sheep: Approach-Avoidance, Operant Response, and Attention.. Animals (Basel) 2018 Jul 17;8(7).
- Siniscalchi M, Padalino B, Aubé L, Quaranta A. Right-nostril use during sniffing at arousing stimuli produces higher cardiac activity in jumper horses.. Laterality 2015;20(4):483-500.
- Farmer K, Krüger K, Byrne RW, Marr I. Sensory laterality in affiliative interactions in domestic horses and ponies (Equus caballus).. Anim Cogn 2018 Sep;21(5):631-637.
- Smith AV, Proops L, Grounds K, Wathan J, Scott SK, McComb K. Domestic horses (Equus caballus) discriminate between negative and positive human nonverbal vocalisations.. Sci Rep 2018 Aug 29;8(1):13052.
- Komárková M, Bartošová J. Lateralized suckling in domestic horses (Equus caballus).. Anim Cogn 2013 May;16(3):343-9.
- Shivley C, Grandin T, Deesing M. Behavioral laterality and facial hair whorls in horses. J Equine Vet Sci 2016; 44: 62–66.
- De Boyer Des Roches A, Richard-Yris MA, Henry S, Ezzaouïa M, Hausberger M. Laterality and emotions: visual laterality in the domestic horse (Equus caballus) differs with objects' emotional value.. Physiol Behav 2008 Jun 9;94(3):487-90.
- de Boyer des Roches A, Durier V, Richard-Yris MA, Blois-Heulin C, Ezzaouïa M, Hausberger M, Henry S. Differential outcomes of unilateral interferences at birth.. Biol Lett 2011 Apr 23;7(2):177-80.
- Esch L, Wöhr C, Erhard M, Krüger K. Horses' (Equus Caballus) Laterality, Stress Hormones, and Task Related Behavior in Innovative Problem-Solving.. Animals (Basel) 2019 May 22;9(5).
- Hausberger M, Cousillas H, Meter A, Karino G, George I, Lemasson A. A crucial role of attention in lateralisation of sound processing?. Symmetry 2019; 11(1), 48.
- Wei XX, Stocker AA. Lawful relation between perceptual bias and discriminability.. Proc Natl Acad Sci U S A 2017 Sep 19;114(38):10244-10249.
- Moscatelli A, Scaleia B, Zago M, Lacquaniti F. Motion direction, luminance contrast, and speed perception: An unexpected meeting.. J Vis 2019 Jun 3;19(6):16.
- Schouten B, Davila A, Verfaillie K. Further explorations of the facing bias in biological motion perception: perspective cues, observer sex, and response times.. PLoS One 2013;8(2):e56978.
- Armario A. The hypothalamic-pituitary-adrenal axis: what can it tell us about stressors?. CNS Neurol Disord Drug Targets 2006 Oct;5(5):485-501.
- Koolhaas JM, Bartolomucci A, Buwalda B, de Boer SF, Flügge G, Korte SM, Meerlo P, Murison R, Olivier B, Palanza P, Richter-Levin G, Sgoifo A, Steimer T, Stiedl O, van Dijk G, Wöhr M, Fuchs E. Stress revisited: a critical evaluation of the stress concept.. Neurosci Biobehav Rev 2011 Apr;35(5):1291-301.
- Baragli P, Vitale V, Banti L, Sighieri C. Effect of aging on behavioural and physiological responses to a stressful stimulus in horses (Equus caballus). Behaviour 2014; 151(11): 1513–1533.
- Scopa C, Palagi E, Sighieri C, Baragli P. Physiological outcomes of calming behaviors support the resilience hypothesis in horses.. Sci Rep 2018 Nov 30;8(1):17501.
- Désiré L, Boissy A, Veissier I. Emotions in farm animals: a new approach to animal welfare in applied ethology.. Behav Processes 2002 Nov;60(2):165-180.
- Heffner RS, Heffner HE. Visual factors in sound localization in mammals.. J Comp Neurol 1992 Mar 15;317(3):219-32.
- Wathan J, McComb K. The eyes and ears are visual indicators of attention in domestic horses.. Curr Biol 2014 Aug 4;24(15):R677-9.
- Brown C, Bibost AL. Laterality is linked to personality in the black-lined rainbowfish, Melanotaenia nigrans. Behav Ecol Sociobiol 2014; 68(6): 999–1005.
- Karolis VR, Corbetta M, Thiebaut de Schotten M. The architecture of functional lateralisation and its relationship to callosal connectivity in the human brain.. Nat Commun 2019 Mar 29;10(1):1417.
- Vallortigara G, Rogers LJ. A function for the bicameral mind.. Cortex 2020 Mar;124:274-285.
- Bisazza A, Cantalupo C, Capocchiano M, Vallortigara G. Population lateralisation and social behaviour: a study with 16 species of fish.. Laterality 2000 Jul;5(3):269-84.
- Vallortigara G, Bisazza A. How ancient is brain lateralization?. In: Rogers LJ, Andrew RJ, editors. Comparative vertebrate lateralization. Cambridge: Cambridge University Press; 2002. pp. 9–69.
- Brown C, Gardner C, Braithwaite VA. Population variation in lateralized eye use in the poeciliid Brachyraphis episcopi.. Proc Biol Sci 2004 Dec 7;271 Suppl 6(Suppl 6):S455-7.
- Corballis MC. The evolution and genetics of cerebral asymmetry.. Philos Trans R Soc Lond B Biol Sci 2009 Apr 12;364(1519):867-79.
- Vallortigara G, Rogers LJ. Survival with an asymmetrical brain: advantages and disadvantages of cerebral lateralization.. Behav Brain Sci 2005 Aug;28(4):575-89; discussion 589-633.
- Bisazza A, De santi A, Vallortigara G. Laterality and cooperation: mosquitofish move closer to a predator when the companion is on their left side.. Anim Behav 1999 May;57(5):1145-1149.
- Ghirlanda S, Vallortigara G. The evolution of brain lateralization: a game-theoretical analysis of population structure.. Proc Biol Sci 2004 Apr 22;271(1541):853-7.
- Ghirlanda S, Frasnelli E, Vallortigara G. Intraspecific competition and coordination in the evolution of lateralization.. Philos Trans R Soc Lond B Biol Sci 2009 Apr 12;364(1519):861-6.
- Sankey C, Richard-Yris MA, Henry S, Fureix C, Nassur F, Hausberger M. Reinforcement as a mediator of the perception of humans by horses (Equus caballus).. Anim Cogn 2010 Sep;13(5):753-64.
- Mendl M, Burman OH, Paul ES. An integrative and functional framework for the study of animal emotion and mood.. Proc Biol Sci 2010 Oct 7;277(1696):2895-904.
- Forkman B, Boissy A, Meunier-Salaün MC, Canali E, Jones RB. A critical review of fear tests used on cattle, pigs, sheep, poultry and horses.. Physiol Behav 2007 Oct 22;92(3):340-74.
- Robins A, Phillips C. Lateralised visual processing in domestic cattle herds responding to novel and familiar stimuli.. Laterality 2010 Sep;15(5):514-34.
- Ocklenburg S, Korte SM, Peterburs J, Wolf OT, Güntürkün O. Stress and laterality - The comparative perspective.. Physiol Behav 2016 Oct 1;164(Pt A):321-9.
- Panksepp J. Cross-species affective neuroscience decoding of the primal affective experiences of humans and related animals.. PLoS One 2011;6(9):e21236.
- Cohen H, Zohar J, Matar MA, Zeev K, Loewenthal U, Richter-Levin G. Setting apart the affected: the use of behavioral criteria in animal models of post traumatic stress disorder.. Neuropsychopharmacology 2004 Nov;29(11):1962-70.
- Bisazza A, Facchin L, Pignatti R, Vallortigara G. Lateralization of detour behaviour in poeciliid fish: the effect of species, gender and sexual motivation.. Behav Brain Res 1998 Mar;91(1-2):157-64.
- Sovrano VA, Bisazza A, Vallortigara G. Lateralization of response to social stimuli in fishes: a comparison between different methods and species.. Physiol Behav 2001 Sep 1-15;74(1-2):237-44.
- Marr I, Preisler V, Farmer K, Stefanski V, Krueger K. Non-invasive stress evaluation in domestic horses (Equus caballus): impact of housing conditions on sensory laterality and immunoglobulin A.. R Soc Open Sci 2020 Feb;7(2):191994.
- McGreevy PD, Rogers LJ. Motor and sensory laterality in thoroughbred horses. Appl Anim Behav Sci 2005; 92(4): 337–352.
- Roche DG, Amcoff M, Morgan R, Sundin J, Andreassen AH, Finnøen MH. Behavioural lateralization in a detour test is not repeatable in fishes. Anim Behav 2020; 167: 55–64.
- McLean S, Morrell LJ. Consistency in the strength of laterality in male, but not female, guppies across different behavioural contexts.. Biol Lett 2020 May;16(5):20190870.
- Vinogradov IM, Jennions MD, Neeman T, Fox RJ. Repeatability of lateralisation in mosquitofish Gambusia holbrooki despite evidence for turn alternation in detour tests.. Anim Cogn 2021 Jul;24(4):765-775.
- Zucca P, Cerri F, Carluccio A, Baciadonna L. Space availability influence laterality in donkeys (Equus asinus).. Behav Processes 2011 Sep;88(1):63-6.
Citations
This article has been cited 7 times.- Felici M, Reddon AR, Maglieri V, Lanatà A, Baragli P. Heart and brain: Change in cardiac entropy is related to lateralised visual inspection in horses. PLoS One 2023;18(8):e0289753.
- Kieson E, Goma AA, Radi M. Tend and Befriend in Horses: Partner Preferences, Lateralization, and Contextualization of Allogrooming in Two Socially Stable Herds of Quarter Horse Mares. Animals (Basel) 2023 Jan 7;13(2).
- Roth LSV, McGreevy P. Horse vision through two lenses: Tinbergen's Four Questions and the Five Domains. Front Vet Sci 2025;12:1647911.
- Loftus L, Newman A, Leach M, Asher L. Exploring the induction and measurement of positive affective state in equines through a personality-centred lens. Sci Rep 2025 May 27;15(1):18550.
- Haussler KK, le Jeune SS, MacKechnie-Guire R, Latif SN, Clayton HM. The Challenge of Defining Laterality in Horses: Is It Laterality or Just Asymmetry?. Animals (Basel) 2025 Jan 21;15(3).
- Hall C, Kay R. Living the good life? A systematic review of behavioural signs of affective state in the domestic horse (Equus caballus) and factors relating to quality of life. Part 2: Horse-human interactions. Anim Welf 2024;33:e41.
- Gainotti G. Mainly Visual Aspects of Emotional Laterality in Cognitively Developed and Highly Social Mammals-A Systematic Review. Brain Sci 2024 Jan 5;14(1).
Use Nutrition Calculator
Check if your horse's diet meets their nutrition requirements with our easy-to-use tool Check your horse's diet with our easy-to-use tool
Talk to a Nutritionist
Discuss your horse's feeding plan with our experts over a free phone consultation Discuss your horse's diet over a phone consultation
Submit Diet Evaluation
Get a customized feeding plan for your horse formulated by our equine nutritionists Get a custom feeding plan formulated by our nutritionists