Analyze Diet
PloS one2015; 10(8); e0131738; doi: 10.1371/journal.pone.0131738

EquiFACS: The Equine Facial Action Coding System.

Abstract: Although previous studies of horses have investigated their facial expressions in specific contexts, e.g. pain, until now there has been no methodology available that documents all the possible facial movements of the horse and provides a way to record all potential facial configurations. This is essential for an objective description of horse facial expressions across a range of contexts that reflect different emotional states. Facial Action Coding Systems (FACS) provide a systematic methodology of identifying and coding facial expressions on the basis of underlying facial musculature and muscle movement. FACS are anatomically based and document all possible facial movements rather than a configuration of movements associated with a particular situation. Consequently, FACS can be applied as a tool for a wide range of research questions. We developed FACS for the domestic horse (Equus caballus) through anatomical investigation of the underlying musculature and subsequent analysis of naturally occurring behaviour captured on high quality video. Discrete facial movements were identified and described in terms of the underlying muscle contractions, in correspondence with previous FACS systems. The reliability of others to be able to learn this system (EquiFACS) and consistently code behavioural sequences was high--and this included people with no previous experience of horses. A wide range of facial movements were identified, including many that are also seen in primates and other domestic animals (dogs and cats). EquiFACS provides a method that can now be used to document the facial movements associated with different social contexts and thus to address questions relevant to understanding social cognition and comparative psychology, as well as informing current veterinary and animal welfare practices.
Publication Date: 2015-08-05 PubMed ID: 26244573PubMed Central: PMC4526551DOI: 10.1371/journal.pone.0131738Google 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.

The research article presents the development of EquiFACS, a Facial Action Coding System for horses, which was created by studying their facial musculature and movements. The system can be used in a wide range of studies and can be learned easily by people with no previous experience with horses.

Development of the Equine Facial Action Coding System (EquiFACS)

  • The research team developed EquiFACS by studying the anatomy of horse facial musculature and analyzing the facial movements in high-quality videos of naturally occurring horse behavior.
  • They successfully described discrete facial movements in terms of the underlying muscle contractions, in line with previous Facial Action Coding Systems.
  • This process facilitated the categorization and documentation of all possible horse facial movements, providing a comprehensive and systematic approach to analyse horse facial expressions.

Reliability of EquiFACS

  • The developed system proved to be reliable as it could be learned and applied consistently by individuals with diverse backgrounds, including those with no previous experience with horses.
  • This highlights the system’s accessibility and potential for widespread use in various research areas.

Identification of a Wide Range of Facial Movements

  • The researchers identified a wide variety of facial movements in horses, which included movements similar to those observed in primates, dogs, and cats.
  • This commonality suggests the potential for comparative studies across different species.

Potential Applications of EquiFACS

  • EquiFACS is not confined to a particular context or situation, instead it can be applied to document the facial movements associated with different social contexts.
  • It can thus contribute to studies aimed at understanding social cognition and comparative psychology and can offer valuable insights into horse behavior in different situations and emotional states.
  • In addition, EquiFACS can be used to inform veterinary procedures and animal welfare practices, potentially improving the care and understanding of horses and their emotional states.

Cite This Article

APA
Wathan J, Burrows AM, Waller BM, McComb K. (2015). EquiFACS: The Equine Facial Action Coding System. PLoS One, 10(8), e0131738. https://doi.org/10.1371/journal.pone.0131738

Publication

ISSN: 1932-6203
NlmUniqueID: 101285081
Country: United States
Language: English
Volume: 10
Issue: 8
Pages: e0131738
PII: e0131738

Researcher Affiliations

Wathan, Jen
  • Mammal Communication and Cognition Research, School of Psychology, University of Sussex, Brighton, United Kingdom.
Burrows, Anne M
  • Centre for Comparative and Evolutionary Psychology, Department of Psychology, University of Portsmouth, Portsmouth, United Kingdom.
Waller, Bridget M
  • Department of Physical Therapy, Duquesne University, Pittsburgh, Pennsylvania, United States of America.
McComb, Karen
  • Mammal Communication and Cognition Research, School of Psychology, University of Sussex, Brighton, United Kingdom.

MeSH Terms

  • Animals
  • Emotions / physiology
  • Face
  • Facial Expression
  • Facial Muscles / physiology
  • Horses
  • Social Behavior
  • Species Specificity

Grant Funding

  • Biotechnology and Biological Sciences Research Council

Conflict of Interest Statement

The authors have declared that no competing interests exist.

References

This article includes 48 references
  1. Ekman P, Friesen WV, Hager JC. Facial Action Coding System. Salt Lake City: Research Nexus; 2002.
  2. Gleerup KB, Forkman B, Lindegaard C, Andersen PH. An equine pain face.. Vet Anaesth Analg 2015 Jan;42(1):103-14.
    doi: 10.1111/vaa.12212pmc: PMC4312484pubmed: 25082060google scholar: lookup
  3. Dalla Costa E, Minero M, Lebelt D, Stucke D, Canali E, Leach MC. Development of the Horse Grimace Scale (HGS) as a pain assessment tool in horses undergoing routine castration.. PLoS One 2014;9(3):e92281.
  4. Diogo R, Wood BA, Aziz MA, Burrows A. On the origin, homologies and evolution of primate facial muscles, with a particular focus on hominoids and a suggested unifying nomenclature for the facial muscles of the Mammalia.. J Anat 2009 Sep;215(3):300-19.
  5. Bruce V, Young A. Face Perception. Taylor & Francis; 2013.
  6. Hole G, Bourne V. Face Processing: Psychological, Neuropsychological, and Applied Perspectives. OUP Oxford; 2010.
  7. Waller BM, Bard KA, Vick SJ, Smith Pasqualini MC. Perceived differences between chimpanzee (Pan troglodytes) and human (Homo sapiens) facial expressions are related to emotional interpretation.. J Comp Psychol 2007 Nov;121(4):398-404.
    doi: 10.1037/0735-7036.121.4.398pubmed: 18085923google scholar: lookup
  8. Vick SJ, Waller BM, Parr LA, Smith Pasqualini MC, Bard KA. A Cross-species Comparison of Facial Morphology and Movement in Humans and Chimpanzees Using the Facial Action Coding System (FACS).. J Nonverbal Behav 2007 Mar;31(1):1-20.
    doi: 10.1007/s10919-006-0017-zpmc: PMC3008553pubmed: 21188285google scholar: lookup
  9. Caeiro CC, Waller BM, Zimmermann E, Burrows AM, Davila-Ross M. OrangFACS: A Muscle-Based Facial Movement Coding System for Orangutans (Pongo spp.). International Journal of Primatology 2012;34(1):115–29.
    doi: 10.1007/s10764-012-9652-xgoogle scholar: lookup
  10. Parr LA, Waller BM, Burrows AM, Gothard KM, Vick SJ. Brief communication: MaqFACS: A muscle-based facial movement coding system for the rhesus macaque.. Am J Phys Anthropol 2010 Dec;143(4):625-30.
    doi: 10.1002/ajpa.21401pmc: PMC2988871pubmed: 20872742google scholar: lookup
  11. Waller BM, Lembeck M, Kuchenbuch P, Burrows AM, Liebal K. GibbonFACS: A Muscle-Based Facial Movement Coding System for Hylobatids. International Journal of Primatology 2012;33(4):809–21.
    doi: 10.1007/s10764-012-9611-6google scholar: lookup
  12. Waller BM, Peirce K, Caeiro CC, Scheider L, Burrows AM, McCune S, Kaminski J. Paedomorphic facial expressions give dogs a selective advantage.. PLoS One 2013;8(12):e82686.
  13. Caeiro CC, Burrows AM, Waller BM. CatFACS. Univeristy of Portsmouth; 2013.
  14. Waller BM, Misch A, Whitehouse J, Herrmann E. Children, but not chimpanzees, have facial correlates of determination.. Biol Lett 2014 Mar;10(3):20130974.
    doi: 10.1098/rsbl.2013.0974pmc: PMC3982434pubmed: 24598107google scholar: lookup
  15. Outram AK, Stear NA, Bendrey R, Olsen S, Kasparov A, Zaibert V, Thorpe N, Evershed RP. The earliest horse harnessing and milking.. Science 2009 Mar 6;323(5919):1332-5.
    doi: 10.1126/science.1168594pubmed: 19265018google scholar: lookup
  16. Clutton-Brock J. A Natural History of Domesticated Mammals. Cambridge University Press; 1999.
  17. Hare B, Brown M, Williamson C, Tomasello M. The domestication of social cognition in dogs.. Science 2002 Nov 22;298(5598):1634-6.
    doi: 10.1126/science.1072702pubmed: 12446914google scholar: lookup
  18. Trut L. Early canid domestication: The farm-fox experiment. American Scientist 1999;87(2):160–9.
  19. Hare B, Plyusnina I, Ignacio N, Schepina O, Stepika A, Wrangham R, Trut L. Social cognitive evolution in captive foxes is a correlated by-product of experimental domestication.. Curr Biol 2005 Feb 8;15(3):226-30.
    doi: 10.1016/j.cub.2005.01.040pubmed: 15694305google scholar: lookup
  20. Waring GH. Horse Behavior. Noyes Publications/William Andrew Pub.; 2003.
  21. Aureli F, Schaffner CM, Boesch C, Bearder SK, Call J, Chapman CA. Fission‐Fusion Dynamics: New Research Frameworks. Current Anthropology 2008;49(4):627–54.
    doi: 10.1086/586708google scholar: lookup
  22. Hinde RA. A conceptual framework. In: Hinde RA, editor. Primate social relationships;an integrated approach. Oxford, UK: Blackwells; 1983. p. 1–7.
  23. Harman AM, Moore S, Hoskins R, Keller P. Horse vision and an explanation for the visual behaviour originally explained by the 'ramp retina'.. Equine Vet J 1999 Sep;31(5):384-90.
  24. Timney B, Keil K. Visual acuity in the horse.. Vision Res 1992 Dec;32(12):2289-93.
    pubmed: 1288005doi: 10.1016/0042-6989(92)90092-wgoogle scholar: lookup
  25. Miller PE, Murphy CJ. Vision in dogs.. J Am Vet Med Assoc 1995 Dec 15;207(12):1623-34.
    pubmed: 7493905
  26. McDonnell S. The Equid Ethogram: A Practical Field Guide to Horse Behavior. ECLIPSE Press; 2003.
  27. Schilder MBH, van Hooff JARAM, van Geer-Plesman CJ, Wensing JB. A Quantitative Analysis of Facial Expression in the Plains Zebra. Zeitschrift für Tierpsychologie 1984;66(1):11–32.
  28. Klingel H. Communication in Perissodactyla. In: Sebeok TA, editor. How Animals Communicate. Bloomington & London: Indiana University Press; 1977.
  29. Parr LA, Waller BM, Vick SJ, Bard KA. Classifying chimpanzee facial expressions using muscle action.. Emotion 2007 Feb;7(1):172-81.
    doi: 10.1037/1528-3542.7.1.172pmc: PMC2826116pubmed: 17352572google scholar: lookup
  30. Waller BM, Micheletta J. Facial Expression in Nonhuman Animals. Emotion Review 2013;5(1):54–9.
    doi: 10.1177/1754073912451503google scholar: lookup
  31. Ashdown RR, Done SH. Color Atlas of Veterinary Anatomy, Volume 2, The Horse. Elsevier Health Sciences UK; 2011.
  32. Burrows AM, Smith TD. Muscles of facial expression in Otolemur, with a comparison to lemuroidea.. Anat Rec A Discov Mol Cell Evol Biol 2003 Sep;274(1):827-36.
    doi: 10.1002/ar.a.10093pubmed: 12923893google scholar: lookup
  33. Burrows AM, Waller BM, Parr LA, Bonar CJ. Muscles of facial expression in the chimpanzee (Pan troglodytes): descriptive, comparative and phylogenetic contexts.. J Anat 2006 Feb;208(2):153-67.
  34. Burrows AM, Waller BM, Parr LA. Facial musculature in the rhesus macaque (Macaca mulatta): evolutionary and functional contexts with comparisons to chimpanzees and humans.. J Anat 2009 Sep;215(3):320-34.
  35. Burrows AM, Diogo R, Waller BM, Bonar CJ, Liebal K. Evolution of the muscles of facial expression in a monogamous ape: evaluating the relative influences of ecological and phylogenetic factors in hylobatids.. Anat Rec (Hoboken) 2011 Apr;294(4):645-63.
    doi: 10.1002/ar.21355pubmed: 21370494google scholar: lookup
  36. Budras KD, Sack WO, Rock S, Horowitz A, Berg R. Anatomy of the Horse: with Aaron Horowitz and Rolf Berg. 6th ed. Hanover, Germany: Schlutersche Verlagsgesellschaft mbH & Co. KG; 2012.
  37. Sisson S, Grossman JD. The Anatomy of the Domestic Animals. 4th ed. Philadelphia and London: W. B. Saunders company; 1962.
  38. Ekman P, Friesen WV, Hager JC. Facial Action Coding System Investigator's Guide. Salt Lake City, USA: Research Nexus; 2002.
  39. Gregory WK. Our Face from Fish to Man: A Portrait Gallery of Our Ancient Ancestors and Kinsfolk Together with a Concise History of Our Best Features. G.P. Putnam's Sons; 1929.
  40. Huber E. Evolution of facial musculature and expression. Baltimore: John Hopkins Press; 1931.
  41. Land MF, Nilsson DE. Animal Eyes. Oxford University Press; 2002.
  42. Sandem AI, Braastad BO. Effects of cow–calf separation on visible eye white and behaviour in dairy cows—A brief report. Applied Animal Behaviour Science 2005;95(3–4):233–9.
  43. Whalen PJ, Kagan J, Cook RG, Davis FC, Kim H, Polis S, McLaren DG, Somerville LH, McLean AA, Maxwell JS, Johnstone T. Human amygdala responsivity to masked fearful eye whites.. Science 2004 Dec 17;306(5704):2061.
    doi: 10.1126/science.1103617pubmed: 15604401google scholar: lookup
  44. Tinbergen N. On aims and methods of ethology. Zeitschrift für Tierpsychologie 1963;20(4):410–33.
  45. Bussières G, Jacques C, Lainay O, Beauchamp G, Leblond A, Cadoré JL, Desmaizières LM, Cuvelliez SG, Troncy E. Development of a composite orthopaedic pain scale in horses.. Res Vet Sci 2008 Oct;85(2):294-306.
    doi: 10.1016/j.rvsc.2007.10.011pubmed: 18061637google scholar: lookup
  46. Jochle W, Moore JN, Brown J, Baker GJ, Lowe JE, Fubini S, Reeves MJ, Watkins JP, White NA. Comparison of detomidine, butorphanol, flunixin meglumine and xylazine in clinical cases of equine colic.. Equine Vet J Suppl 1989 Jun;(7):111-6.
  47. 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.
    doi: 10.1016/j.cub.2014.06.023pmc: PMC4123162pubmed: 25093554google scholar: lookup
  48. Boissy A, Manteuffel G, Jensen MB, Moe RO, Spruijt B, Keeling LJ, Winckler C, Forkman B, Dimitrov I, Langbein J, Bakken M, Veissier I, Aubert A. Assessment of positive emotions in animals to improve their welfare.. Physiol Behav 2007 Oct 22;92(3):375-97.
    doi: 10.1016/j.physbeh.2007.02.003pubmed: 17428510google scholar: lookup

Citations

This article has been cited 79 times.
  1. Marchand WR. Potential Mechanisms of Action and Outcomes of Equine-Assisted Services for Veterans with a History of Trauma: A Narrative Review of the Literature. Int J Environ Res Public Health 2023 Jul 16;20(14).
    doi: 10.3390/ijerph20146377pubmed: 37510609google scholar: lookup
  2. Carmo LG, Werner LC, Michelotto PV Jr, Daros RR. Horse behavior and facial movements in relation to food rewards. PLoS One 2023;18(6):e0286045.
    doi: 10.1371/journal.pone.0286045pubmed: 37307268google scholar: lookup
  3. Tomberg C, Petagna M, de Selliers de Moranville LA. Horses (Equus caballus) facial micro-expressions: insight into discreet social information. Sci Rep 2023 May 27;13(1):8625.
    doi: 10.1038/s41598-023-35807-zpubmed: 37244937google scholar: lookup
  4. Kim SM, Cho GJ. Analysis of Various Facial Expressions of Horses as a Welfare Indicator Using Deep Learning. Vet Sci 2023 Apr 10;10(4).
    doi: 10.3390/vetsci10040283pubmed: 37104439google scholar: lookup
  5. Kappel S, Ramirez Montes De Oca MA, Collins S, Herborn K, Mendl M, Fureix C. Do you see what I see? Testing horses' ability to recognise real-life objects from 2D computer projections. Anim Cogn 2023 Jul;26(4):1147-1159.
    doi: 10.1007/s10071-023-01761-6pubmed: 36864246google scholar: lookup
  6. Carvalho JRG, Trindade PHE, Conde G, Antonioli ML, Funnicelli MIG, Dias PP, Canola PA, Chinelatto MA, Ferraz GC. Facial Expressions of Horses Using Weighted Multivariate Statistics for Assessment of Subtle Local Pain Induced by Polylactide-Based Polymers Implanted Subcutaneously. Animals (Basel) 2022 Sep 13;12(18).
    doi: 10.3390/ani12182400pubmed: 36139260google scholar: lookup
  7. Merkies K, Sudarenko Y, Hodder AJ. Can Ponies (Equus Caballus) Distinguish Human Facial Expressions?. Animals (Basel) 2022 Sep 7;12(18).
    doi: 10.3390/ani12182331pubmed: 36139191google scholar: lookup
  8. Leconstant C, Spitz E. Integrative Model of Human-Animal Interactions: A One Health-One Welfare Systemic Approach to Studying HAI. Front Vet Sci 2022;9:656833.
    doi: 10.3389/fvets.2022.656833pubmed: 35968006google scholar: lookup
  9. Lou ME, Porter ST, Massey JS, Ventura B, Deen J, Li Y. The Application of 3D Landmark-Based Geometric Morphometrics towards Refinement of the Piglet Grimace Scale. Animals (Basel) 2022 Jul 30;12(15).
    doi: 10.3390/ani12151944pubmed: 35953933google scholar: lookup
  10. Kavanagh E, Kimock C, Whitehouse J, Micheletta J, Waller BM. Revisiting Darwin's comparisons between human and non-human primate facial signals. Evol Hum Sci 2022;4.
    doi: 10.1017/ehs.2022.26pubmed: 35821665google scholar: lookup
  11. Correia-Caeiro C, Burrows A, Wilson DA, Abdelrahman A, Miyabe-Nishiwaki T. CalliFACS: The common marmoset Facial Action Coding System. PLoS One 2022;17(5):e0266442.
    doi: 10.1371/journal.pone.0266442pubmed: 35580128google scholar: lookup
  12. Broomé S, Ask K, Rashid-Engström M, Haubro Andersen P, Kjellström H. Sharing pain: Using pain domain transfer for video recognition of low grade orthopedic pain in horses. PLoS One 2022;17(3):e0263854.
    doi: 10.1371/journal.pone.0263854pubmed: 35245288google scholar: lookup
  13. Mota-Rojas D, Marcet-Rius M, Ogi A, Hernández-Ávalos I, Mariti C, Martínez-Burnes J, Mora-Medina P, Casas A, Domínguez A, Reyes B, Gazzano A. Current Advances in Assessment of Dog's Emotions, Facial Expressions, and Their Use for Clinical Recognition of Pain. Animals (Basel) 2021 Nov 22;11(11).
    doi: 10.3390/ani11113334pubmed: 34828066google scholar: lookup
  14. Mielke A, Waller BM, Pérez C, Rincon AV, Duboscq J, Micheletta J. NetFACS: Using network science to understand facial communication systems. Behav Res Methods 2022 Aug;54(4):1912-1927.
    doi: 10.3758/s13428-021-01692-5pubmed: 34755285google scholar: lookup
  15. Lencioni GC, de Sousa RV, de Souza Sardinha EJ, Corrêa RR, Zanella AJ. Pain assessment in horses using automatic facial expression recognition through deep learning-based modeling. PLoS One 2021;16(10):e0258672.
    doi: 10.1371/journal.pone.0258672pubmed: 34665834google scholar: lookup
  16. Hintze S, Schanz L. Using the Judgment Bias Task to Identify Behavioral Indicators of Affective State: Do Eye Wrinkles in Horses Reflect Mood?. Front Vet Sci 2021;8:676888.
    doi: 10.3389/fvets.2021.676888pubmed: 34307525google scholar: lookup
  17. Andersen PH, Broomé S, Rashid M, Lundblad J, Ask K, Li Z, Hernlund E, Rhodin M, Kjellström H. Towards Machine Recognition of Facial Expressions of Pain in Horses. Animals (Basel) 2021 Jun 1;11(6).
    doi: 10.3390/ani11061643pubmed: 34206077google scholar: lookup
  18. Lundblad J, Rashid M, Rhodin M, Haubro Andersen P. Effect of transportation and social isolation on facial expressions of healthy horses. PLoS One 2021;16(6):e0241532.
    doi: 10.1371/journal.pone.0241532pubmed: 34086704google scholar: lookup
  19. de Oliveira AR, Gozalo-Marcilla M, Ringer SK, Schauvliege S, Fonseca MW, Esteves Trindade PH, Prospero Puoli Filho JN, Luna SPL. Development and validation of the facial scale (FaceSed) to evaluate sedation in horses. PLoS One 2021;16(6):e0251909.
    doi: 10.1371/journal.pone.0251909pubmed: 34061878google scholar: lookup
  20. Torcivia C, McDonnell S. Equine Discomfort Ethogram. Animals (Basel) 2021 Feb 23;11(2).
    doi: 10.3390/ani11020580pubmed: 33672338google scholar: lookup
  21. Correia-Caeiro C, Holmes K, Miyabe-Nishiwaki T. Extending the MaqFACS to measure facial movement in Japanese macaques (Macaca fuscata) reveals a wide repertoire potential. PLoS One 2021;16(1):e0245117.
    doi: 10.1371/journal.pone.0245117pubmed: 33411716google scholar: lookup
  22. Ask K, Rhodin M, Tamminen LM, Hernlund E, Haubro Andersen P. Identification of Body Behaviors and Facial Expressions Associated with Induced Orthopedic Pain in Four Equine Pain Scales. Animals (Basel) 2020 Nov 19;10(11).
    doi: 10.3390/ani10112155pubmed: 33228117google scholar: lookup
  23. Rashid M, Silventoinen A, Gleerup KB, Andersen PH. Equine Facial Action Coding System for determination of pain-related facial responses in videos of horses. PLoS One 2020;15(11):e0231608.
    doi: 10.1371/journal.pone.0231608pubmed: 33141852google scholar: lookup
  24. Csoltova E, Mehinagic E. Where Do We Stand in the Domestic Dog ( Canis familiaris ) Positive-Emotion Assessment: A State-of-the-Art Review and Future Directions. Front Psychol 2020;11:2131.
    doi: 10.3389/fpsyg.2020.02131pubmed: 33013543google scholar: lookup
  25. Neethirajan S. Transforming the Adaptation Physiology of Farm Animals through Sensors. Animals (Basel) 2020 Aug 26;10(9).
    doi: 10.3390/ani10091512pubmed: 32859060google scholar: lookup
  26. Mellor DJ. Mouth Pain in Horses: Physiological Foundations, Behavioural Indices, Welfare Implications, and a Suggested Solution. Animals (Basel) 2020 Mar 29;10(4).
    doi: 10.3390/ani10040572pubmed: 32235343google scholar: lookup
  27. van Dierendonck MC, Burden FA, Rickards K, van Loon JPAM. Monitoring Acute Pain in Donkeys with the Equine Utrecht University Scale for Donkeys Composite Pain Assessment (EQUUS-DONKEY-COMPASS) and the Equine Utrecht University Scale for Donkey Facial Assessment of Pain (EQUUS-DONKEY-FAP). Animals (Basel) 2020 Feb 22;10(2).
    doi: 10.3390/ani10020354pubmed: 32098391google scholar: lookup
  28. Trindade PHE, Hartmann E, Keeling LJ, Andersen PH, Ferraz GC, Paranhos da Costa MJR. Effect of work on body language of ranch horses in Brazil. PLoS One 2020;15(1):e0228130.
    doi: 10.1371/journal.pone.0228130pubmed: 31990951google scholar: lookup
  29. Semin GR, Scandurra A, Baragli P, Lanatà A, D'Aniello B. Inter- and Intra-Species Communication of Emotion: Chemosignals as the Neglected Medium. Animals (Basel) 2019 Oct 31;9(11).
    doi: 10.3390/ani9110887pubmed: 31683710google scholar: lookup
  30. Merkies K, Ready C, Farkas L, Hodder A. Eye Blink Rates and Eyelid Twitches as a Non-Invasive Measure of Stress in the Domestic Horse. Animals (Basel) 2019 Aug 15;9(8).
    doi: 10.3390/ani9080562pubmed: 31443315google scholar: lookup
  31. Finka LR, Luna SP, Brondani JT, Tzimiropoulos Y, McDonagh J, Farnworth MJ, Ruta M, Mills DS. Geometric morphometrics for the study of facial expressions in non-human animals, using the domestic cat as an exemplar. Sci Rep 2019 Jul 8;9(1):9883.
    doi: 10.1038/s41598-019-46330-5pubmed: 31285531google scholar: lookup
  32. Briefer Freymond S, Bardou D, Beuret S, Bachmann I, Zuberbühler K, Briefer EF. Elevated Sensitivity to Tactile Stimuli in Stereotypic Horses. Front Vet Sci 2019;6:162.
    doi: 10.3389/fvets.2019.00162pubmed: 31275947google scholar: lookup
  33. Benn AL, McLelland DJ, Whittaker AL. A Review of Welfare Assessment Methods in Reptiles, and Preliminary Application of the Welfare Quality(®) Protocol to the Pygmy Blue-Tongue Skink, Tiliqua adelaidensis, Using Animal-Based Measures. Animals (Basel) 2019 Jan 17;9(1).
    doi: 10.3390/ani9010027pubmed: 30658490google scholar: lookup
  34. Camerlink I, Coulange E, Farish M, Baxter EM, Turner SP. Facial expression as a potential measure of both intent and emotion. Sci Rep 2018 Dec 4;8(1):17602.
    doi: 10.1038/s41598-018-35905-3pubmed: 30514964google scholar: lookup
  35. 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.
    doi: 10.1038/s41598-018-30777-zpubmed: 30158532google scholar: lookup
  36. Nakamura K, Takimoto-Inose A, Hasegawa T. Cross-modal perception of human emotion in domestic horses (Equus caballus). Sci Rep 2018 Jun 21;8(1):8660.
    doi: 10.1038/s41598-018-26892-6pubmed: 29930289google scholar: lookup
  37. Sommerville R, Brown AF, Upjohn M. A standardised equine-based welfare assessment tool used for six years in low and middle income countries. PLoS One 2018;13(2):e0192354.
    doi: 10.1371/journal.pone.0192354pubmed: 29466391google scholar: lookup
  38. Caeiro C, Guo K, Mills D. Dogs and humans respond to emotionally competent stimuli by producing different facial actions. Sci Rep 2017 Nov 14;7(1):15525.
    doi: 10.1038/s41598-017-15091-4pubmed: 29138393google scholar: lookup
  39. Baragli P, Demuru E, Scopa C, Palagi E. Are horses capable of mirror self-recognition? A pilot study. PLoS One 2017;12(5):e0176717.
    doi: 10.1371/journal.pone.0176717pubmed: 28510577google scholar: lookup
  40. Briefer EF, Mandel R, Maigrot AL, Briefer Freymond S, Bachmann I, Hillmann E. Perception of emotional valence in horse whinnies. Front Zool 2017;14:8.
    doi: 10.1186/s12983-017-0193-1pubmed: 28203263google scholar: lookup
  41. Wathan J, Proops L, Grounds K, McComb K. Horses discriminate between facial expressions of conspecifics. Sci Rep 2016 Dec 20;6:38322.
    doi: 10.1038/srep38322pubmed: 27995958google scholar: lookup
  42. Finlayson K, Lampe JF, Hintze S, Würbel H, Melotti L. Facial Indicators of Positive Emotions in Rats. PLoS One 2016;11(11):e0166446.
    doi: 10.1371/journal.pone.0166446pubmed: 27902721google scholar: lookup
  43. Hintze S, Smith S, Patt A, Bachmann I, Würbel H. Are Eyes a Mirror of the Soul? What Eye Wrinkles Reveal about a Horse's Emotional State. PLoS One 2016;11(10):e0164017.
    doi: 10.1371/journal.pone.0164017pubmed: 27732647google scholar: lookup
  44. 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.
    doi: 10.1098/rsbl.2015.0907pubmed: 26864784google scholar: lookup
  45. Burrows AM, Li L, Waller BM, Micheletta J. Social variables exert selective pressures in the evolution and form of primate mimetic musculature. J Anat 2016 Apr;228(4):595-607.
    doi: 10.1111/joa.12440pubmed: 26750637google scholar: lookup
  46. Julle-Danière É, Micheletta J, Whitehouse J, Joly M, Gass C, Burrows AM, Waller BM. MaqFACS (Macaque Facial Action Coding System) can be used to document facial movements in Barbary macaques (Macaca sylvanus). PeerJ 2015;3:e1248.
    doi: 10.7717/peerj.1248pubmed: 26401458google scholar: lookup
  47. Wathan J, Burrows AM, Waller BM, McComb K. Correction: EquiFACS: The Equine Facial Action Coding System. PLoS One 2015;10(9):e0137818.
    doi: 10.1371/journal.pone.0137818pubmed: 26336096google scholar: lookup
  48. Górski K, Kondracki S, Kępka-Borkowska K. The Complexity of Communication in Mammals: From Social and Emotional Mechanisms to Human Influence and Multimodal Applications. Animals (Basel) 2026 Jan 15;16(2).
    doi: 10.3390/ani16020265pubmed: 41594455google scholar: lookup
  49. Panzera M, Statelli A. Heart Rate Variability Spectral Analysis for Monitoring Autonomic Activation in a Donkey Involved in Animal-Assisted Therapy: A Single Subject Design During Animal-Assisted Therapy Sessions. Vet Sci 2025 Nov 28;12(12).
    doi: 10.3390/vetsci12121131pubmed: 41472111google scholar: lookup
  50. Correia-Caeiro C, Zamansky A, Karl S, Bremhorst A. Research Methods for the Analysis of Visual Emotion Cues in Animals: A Workshop Report. Animals (Basel) 2025 Oct 29;15(21).
    doi: 10.3390/ani15213142pubmed: 41227471google scholar: lookup
  51. Hennes N, Tutin L, Foury A, Vancassel S, Bourguignon H, Duluard A, Ruet A, Lansade L. Exploring the association between stress-related hormonal changes, behaviours and facial movements after an interval training exercise in French Standardbred. PLoS One 2025;20(11):e0328430.
    doi: 10.1371/journal.pone.0328430pubmed: 41191577google scholar: lookup
  52. Martvel G, Riemer S. Automated analysis of emotional expressions in dogs based on geometric morphometrics. Sci Rep 2025 Sep 2;15(1):32331.
    doi: 10.1038/s41598-025-15741-ypubmed: 40897770google scholar: lookup
  53. Uldahl M, Mellor DJ. Regulatory Integrity and Welfare in Horse Sport: A Constructively Critical Perspective. Animals (Basel) 2025 Jun 30;15(13).
    doi: 10.3390/ani15131934pubmed: 40646833google scholar: lookup
  54. Gobbo E, Topal O, Novalija I, Mladenić D, Zupan Šemrov M. An iterative approach to identify key predictive features of fear reactivity and fearfulness in horses (Equus caballus). Sci Rep 2025 Jul 9;15(1):24590.
    doi: 10.1038/s41598-025-10725-4pubmed: 40628935google scholar: lookup
  55. Correia-Caeiro C, Kuchenbuch PH, Oña LS, Wegdell F, Leroux M, Schuele A, Taglialatela J, Townsend S, Surbeck M, Waller BM, Liebal K. Adapting the facial action coding system for chimpanzees (Pan troglodytes) to bonobos (Pan paniscus): the ChimpFACS extension for bonobos. PeerJ 2025;13:e19484.
    doi: 10.7717/peerj.19484pubmed: 40525106google scholar: lookup
  56. Portier K, Schiesari C, Gauthier L, Yeng LT, Tabacchi Fantoni D, Formenton MR. Comparison of the Prevalence and Location of Trigger Points in Dressage and Show-Jumping Horses. Animals (Basel) 2025 May 27;15(11).
    doi: 10.3390/ani15111558pubmed: 40509024google scholar: lookup
  57. 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.
    doi: 10.1038/s41598-025-98034-8pubmed: 40425817google scholar: lookup
  58. Jardat P, Yamamoto S, Ringhofer M, Tanguy-Guillo N, Parias C, Reigner F, Calandreau L, Lansade L. Emotional contagion of fear and joy from humans to horses using a combination of facial and vocal cues. Sci Rep 2025 May 21;15(1):17689.
    doi: 10.1038/s41598-025-98794-3pubmed: 40399542google scholar: lookup
  59. da Fé VCS, Dos Santos VMO, de Lima ACB, Hernandes MSP, Caldara FR, Gomes MNB. Auditory enrichment on facial and physiological responses of Pantaneiro geldings and mares under short-term stress. PLoS One 2025;20(5):e0323649.
    doi: 10.1371/journal.pone.0323649pubmed: 40392914google scholar: lookup
  60. Phelipon R, Bertrand L, Jardat P, Reigner F, Lewis K, Micheletta J, Lansade L. Characterisation of facial expressions and behaviours of horses in response to positive and negative emotional anticipation using network analysis. PLoS One 2025;20(5):e0319315.
    doi: 10.1371/journal.pone.0319315pubmed: 40367029google scholar: lookup
  61. Cordoni G, Brescini M, Pirarba L, Giaretto F, Norscia I. Functional and Morphological Differences in the Play Face and Full Play Face in Lowland Gorillas, a Hominid Species: Implications for the Evolutionary Roots of Smile and Laugh Face. Am J Biol Anthropol 2025 May;187(1):e70061.
    doi: 10.1002/ajpa.70061pubmed: 40341893google scholar: lookup
  62. Florkiewicz BN, Lazebnik T. Combinatorics and complexity of chimpanzee (Pan troglodytes) facial signals. Anim Cogn 2025 Apr 30;28(1):34.
    doi: 10.1007/s10071-025-01955-0pubmed: 40304773google scholar: lookup
  63. MacKechnie-Guire R, Clayton H, Williams J, Marlin D, Fisher M, Fisher D, Walker V, Murray R. Measuring Noseband Tightness on the Lateral Aspect of the Horse's Face. Animals (Basel) 2025 Feb 13;15(4).
    doi: 10.3390/ani15040537pubmed: 40003019google scholar: lookup
  64. Lundblad J, Rhodin M, Hernlund E, Bjarnestig H, Hidén Rudander S, Haubro Andersen P. Facial expressions during compound interventions of nociception, conspecific isolation, and sedation in horses. Sci Rep 2025 Feb 13;15(1):5373.
    doi: 10.1038/s41598-025-89329-xpubmed: 39948238google scholar: lookup
  65. Correia-Caeiro C, Costa R, Hayashi M, Burrows A, Pater J, Miyabe-Nishiwaki T, Richardson JL, Robbins MM, Waller B, Liebal K. GorillaFACS: The Facial Action Coding System for the Gorilla spp. PLoS One 2025;20(1):e0308790.
    doi: 10.1371/journal.pone.0308790pubmed: 39874277google scholar: lookup
  66. Mahmoud A, Scott L, Florkiewicz BN. Examining Mammalian facial behavior using Facial Action Coding Systems (FACS) and combinatorics. PLoS One 2025;20(1):e0314896.
    doi: 10.1371/journal.pone.0314896pubmed: 39869619google scholar: lookup
  67. Domínguez-Oliva A, Chávez C, Martínez-Burnes J, Olmos-Hernández A, Hernández-Avalos I, Mota-Rojas D. Neurobiology and Anatomy of Facial Expressions in Great Apes: Application of the AnimalFACS and Its Possible Association with the Animal's Affective State. Animals (Basel) 2024 Nov 26;14(23).
    doi: 10.3390/ani14233414pubmed: 39682379google scholar: lookup
  68. Chiavaccini L, Gupta A, Anclade N, Chiavaccini G, De Gennaro C, Johnson AN, Portela DA, Romano M, Vettorato E, Luethy D. Automated acute pain prediction in domestic goats using deep learning-based models on video-recordings. Sci Rep 2024 Nov 7;14(1):27104.
    doi: 10.1038/s41598-024-78494-0pubmed: 39511381google scholar: lookup
  69. 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 I: Fulfilment of species-specific needs. Anim Welf 2024;33:e40.
    doi: 10.1017/awf.2024.38pubmed: 39464387google scholar: lookup
  70. Bogossian PM, Pereira JS, da Silva NF, Hilgert AR, Seidel SRT, Fülber J, Belli CB, Fernandes WR. Social facilitation of trotting: Can horses perceive and adapt to the movement of another horse?. PLoS One 2024;19(8):e0309474.
    doi: 10.1371/journal.pone.0309474pubmed: 39186726google scholar: lookup
  71. Chiavaccini L, Gupta A, Chiavaccini G. From facial expressions to algorithms: a narrative review of animal pain recognition technologies. Front Vet Sci 2024;11:1436795.
    doi: 10.3389/fvets.2024.1436795pubmed: 39086767google scholar: lookup
  72. Feighelstein M, Riccie-Bonot C, Hasan H, Weinberg H, Rettig T, Segal M, Distelfeld T, Shimshoni I, Mills DS, Zamansky A. Automated recognition of emotional states of horses from facial expressions. PLoS One 2024;19(7):e0302893.
    doi: 10.1371/journal.pone.0302893pubmed: 39008504google scholar: lookup
  73. Hobkirk ER, Twiss SD. Domestication constrains the ability of dogs to convey emotions via facial expressions in comparison to their wolf ancestors. Sci Rep 2024 May 7;14(1):10491.
    doi: 10.1038/s41598-024-61110-6pubmed: 38714729google scholar: lookup
  74. Ask K, Rhodin M, Rashid-Engström M, Hernlund E, Andersen PH. Changes in the equine facial repertoire during different orthopedic pain intensities. Sci Rep 2024 Jan 2;14(1):129.
    doi: 10.1038/s41598-023-50383-ypubmed: 38167926google scholar: lookup
  75. Kil N, Ertelt K, Auer U. Development and Validation of an Automated Video Tracking Model for Stabled Horses. Animals (Basel) 2020 Nov 30;10(12).
    doi: 10.3390/ani10122258pubmed: 33266297google scholar: lookup
  76. Scopa C, Contalbrigo L, Greco A, Lanatà A, Scilingo EP, Baragli P. Emotional Transfer in Human-Horse Interaction: New Perspectives on Equine Assisted Interventions. Animals (Basel) 2019 Nov 26;9(12).
    doi: 10.3390/ani9121030pubmed: 31779120google scholar: lookup
  77. Baciadonna L, Briefer EF, Favaro L, McElligott AG. Goats distinguish between positive and negative emotion-linked vocalisations. Front Zool 2019;16:25.
    doi: 10.1186/s12983-019-0323-zpubmed: 31320917google scholar: lookup
  78. Schanz L, Krueger K, Hintze S. Sex and Age Don't Matter, but Breed Type Does-Factors Influencing Eye Wrinkle Expression in Horses. Front Vet Sci 2019;6:154.
    doi: 10.3389/fvets.2019.00154pubmed: 31192235google scholar: lookup
  79. Lansade L, Nowak R, Lainé AL, Leterrier C, Bonneau C, Parias C, Bertin A. Facial expression and oxytocin as possible markers of positive emotions in horses. Sci Rep 2018 Oct 2;8(1):14680.
    doi: 10.1038/s41598-018-32993-zpubmed: 30279565google scholar: lookup