Animals : an open access journal from MDPI2023; 13(9); 1507; doi: 10.3390/ani13091507

Mental Experiences in Wild Animals: Scientifically Validating Measurable Welfare Indicators in Free-Roaming Horses.

Abstract: The mental experiences of animals are what characterises their welfare status. The Five Domains Model for assessing welfare aligns with the understanding that physical and mental states are linked. Following measurement of indicators within each of the four physical/functional Domains (1. Nutrition; 2. Physical environment; 3. Health; and 4. Behavioural interactions), the anticipated negative or positive affective consequences (mental experiences) are cautiously inferred and assigned to Domain 5. Those inferences derive credibility from validated knowledge of the underlying systems of physiology, neurophysiology, neuroethology and affective neuroscience. Any indicators used for assessing welfare need to be scientifically validated. This requires, firstly, evidence of the links between a measurable/observable indicator and the physical/functional impact (in Domains 1 to 4), and secondly, a demonstrable relationship between the physical/functional impact and the mental experience it is inferred the indicators reflect (in Domain five). This review refers to indicators of physical/functional states in Domains 1 to 4, which have been shown to be measurable in free-roaming wild horses, and then evaluates the scientific evidence linking them to inferred mental experiences in Domain 5. This is the first time that the scientific evidence validating a comprehensive range of welfare indicators has been synthesised in this way. Inserting these indicators into the Five Domains Model enables transparently justifiable assessment and grading of welfare status in free-roaming horses.
Publication Date: 2023-04-28 PubMed ID: 37174544PubMed Central: PMC10177449DOI: 10.3390/ani13091507Google 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
  • Review

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 discusses how the mental experiences of wild animals, particularly free-roaming horses, can be scientifically validated as indicators of their welfare status. It does so by utilizing the Five Domains Model which examines both physical and mental factors to ascertain an animal’s wellbeing.

Understanding the Mental Experiences in Animals

  • The research strongly emphasizes the importance of mental experiences in understanding the welfare state of animals. These experiences are identified as essential because they are a reflection of how the animal responds to its environment which, in turn, affects its overall wellness.
  • The Five Domains Model is used as a framework in the research to systematically observe and evaluate animal welfare. This model categorizes the indicators into five domains: nutrition, physical environment, health, behavioural interactions, and the resulting mental experiences from the first four domains.

Finding Credibility in Inference

  • The researchers assert that the credibility of their inferences is validated by existing scientific knowledge. Studies and data from the fields of physiology, neurophysiology, neuroethology, and affective neuroscience serve as a base that lends these inferences empirical veracity.
  • This provides a strong scientific foundation that allows for the inference of the mental experience (Domain 5) from the physical/functional indicators (Domains 1 to 4).

The Validation Process

  • The validation of welfare indicators process consists of two parts. Firstly, evidence is needed to establish a link between a measurable/observable indicator and its physical/functional impact. Secondly, there must be a demonstrable relationship between this physical/functional impact and the inferred mental experience.
  • The validation process ensures that the chosen indicators are firmly rooted in empirical evidence, rendering their application in assessing welfare both accurate and scientifically grounded.

Applying the Model to Free-Roaming Horses

  • The research focuses specifically on free-roaming horses, using these indicators measured in Domains 1 to 4. It is the first time scientific evidence is synthesized this way, providing an insightful perspective into the mental experiences of these animals.
  • The findings allow for a transparent and justified assessment of welfare status in free-roaming horses – essentially providing a method by which the physical and mental states of these animals can be accurately gauged.

Cite This Article

APA
Harvey AM, Beausoleil NJ, Ramp D, Mellor DJ. (2023). Mental Experiences in Wild Animals: Scientifically Validating Measurable Welfare Indicators in Free-Roaming Horses. Animals (Basel), 13(9), 1507. https://doi.org/10.3390/ani13091507

Publication

ISSN: 2076-2615
NlmUniqueID: 101635614
Country: Switzerland
Language: English
Volume: 13
Issue: 9
PII: 1507

Researcher Affiliations

Harvey, Andrea M
  • Centre for Compassionate Conservation, TD School, University of Technology Sydney, Ultimo, NSW 2007, Australia.
Beausoleil, Ngaio J
  • Animal Welfare Science and Bioethics Centre, School of Veterinary Science, Massey University, Palmerston North 4442, New Zealand.
Ramp, Daniel
  • Centre for Compassionate Conservation, TD School, University of Technology Sydney, Ultimo, NSW 2007, Australia.
Mellor, David J
  • Animal Welfare Science and Bioethics Centre, School of Veterinary Science, Massey University, Palmerston North 4442, New Zealand.

Conflict of Interest Statement

The authors declare no conflict of interest.

References

This article includes 152 references
  1. Beausoleil NJ, Mellor DJ, Baker L, Baker SE, Bellio M, Clarke AS, Dale A, Garlick S, Jones B, Harvey A, Pitcher BJ, Sherwen S, Stockin KA, Zito S. "Feelings and Fitness" Not "Feelings or Fitness"-The Raison d'u00eatre of Conservation Welfare, Which Aligns Conservation and Animal Welfare Objectives.. Front Vet Sci 2018;5:296.
    doi: 10.3389/fvets.2018.00296pmc: PMC6277474pubmed: 30538995google scholar: lookup
  2. Harvey AM, Beausoleil NJ, Ramp D, Mellor DJ. A Ten-Stage Protocol for Assessing the Welfare of Individual Non-Captive Wild Animals: Free-Roaming Horses (Equus Ferus Caballus) as an Example.. Animals (Basel) 2020 Jan 16;10(1).
    doi: 10.3390/ani10010148pmc: PMC7022444pubmed: 31963232google scholar: lookup
  3. Mellor DJ, Beausoleil NJ, Littlewood KE, McLean AN, McGreevy PD, Jones B, Wilkins C. The 2020 Five Domains Model: Including Human-Animal Interactions in Assessments of Animal Welfare.. Animals (Basel) 2020 Oct 14;10(10).
    doi: 10.3390/ani10101870pmc: PMC7602120pubmed: 33066335google scholar: lookup
  4. Mellor D.J., Beausoleil N.J. Extending the u2018Five Domainsu2019 model for animal welfare assessment to incorporate positive welfare states. Anim. Welf. 2015;24:241u2013253. doi: 10.7120/09627286.24.3.241.
    doi: 10.7120/09627286.24.3.241google scholar: lookup
  5. Mellor DJ. Updating Animal Welfare Thinking: Moving beyond the "Five Freedoms" towards "A Life Worth Living".. Animals (Basel) 2016 Mar 14;6(3).
    doi: 10.3390/ani6030021pmc: PMC4810049pubmed: 27102171google scholar: lookup
  6. Littlewood KE, Mellor DJ. Changes in the Welfare of an Injured Working Farm Dog Assessed Using the Five Domains Model.. Animals (Basel) 2016 Sep 21;6(9).
    doi: 10.3390/ani6090058pmc: PMC5035953pubmed: 27657140google scholar: lookup
  7. Mellor DJ. Operational Details of the Five Domains Model and Its Key Applications to the Assessment and Management of Animal Welfare.. Animals (Basel) 2017 Aug 9;7(8).
    doi: 10.3390/ani7080060pmc: PMC5575572pubmed: 28792485google scholar: lookup
  8. Harvey AM, Ramp D, Mellor DJ. Review of the Foundational Knowledge Required for Assessing Horse Welfare.. Animals (Basel) 2022 Dec 1;12(23).
    doi: 10.3390/ani12233385pmc: PMC9736110pubmed: 36496906google scholar: lookup
  9. Harvey AM, Morton JM, Mellor DJ, Russell V, Chapple RS, Ramp D. Use of Remote Camera Traps to Evaluate Animal-Based Welfare Indicators in Individual Free-Roaming Wild Horses.. Animals (Basel) 2021 Jul 15;11(7).
    doi: 10.3390/ani11072101pmc: PMC8300222pubmed: 34359229google scholar: lookup
  10. Harvey AM, Meggiolaro MN, Hall E, Watts ET, Ramp D, u0160lapeta J. Wild horse populations in south-east Australia have a high prevalence of Strongylus vulgaris and may act as a reservoir of infection for domestic horses.. Int J Parasitol Parasites Wildl 2019 Apr;8:156-163.
  11. Hockenhull J., Whay H.R. A review of approaches to assessing equine welfare. Equine Vet. Educ. 2014;26:159u2013166. doi: 10.1111/eve.12129.
    doi: 10.1111/eve.12129google scholar: lookup
  12. Beausoleil N.J., Mellor D.J. Validating indicators of sheep welfare. In: Greyling J., editor. Achieving Sustainable Production of Sheep. Burleigh Dodds Science Publishing; London, UK: 2017.
  13. Denton DA, McKinley MJ, Farrell M, Egan GF. The role of primordial emotions in the evolutionary origin of consciousness.. Conscious Cogn 2009 Jun;18(2):500-14.
    doi: 10.1016/j.concog.2008.06.009pubmed: 18701321google scholar: lookup
  14. Sufit E, Houpt KA, Sweeting M. Physiological stimuli of thirst and drinking patterns in ponies.. Equine Vet J 1985 Jan;17(1):12-6.
  15. Pritchard JC, Burn CC, Barr AR, Whay HR. Validity of indicators of dehydration in working horses: a longitudinal study of changes in skin tent duration, mucous membrane dryness and drinking behaviour.. Equine Vet J 2008 Sep;40(6):558-64.
    doi: 10.2746/042516408X297462pubmed: 18356129google scholar: lookup
  16. Mullan S., Szmaraged C., Hotchkiss I., Whay H.R. The welfare of long-line tethered and free-ranging horses kept on public grazing land in South Wales. Anim. Welf. 2014;23:25u201337. doi: 10.7120/09627286.23.1.025.
    doi: 10.7120/09627286.23.1.025google scholar: lookup
  17. Dalla Costa E., Murray L., Dai F., Canali E., Minero M. Equine on-farm welfare assessment: A review of animal-based indicators. Anim Welf. 2014;23:323u2013341.
  18. McKiernan F, Houchins JA, Mattes RD. Relationships between human thirst, hunger, drinking, and feeding.. Physiol Behav 2008 Aug 6;94(5):700-8.
  19. Carroll CL, Huntington PJ. Body condition scoring and weight estimation of horses.. Equine Vet J 1988 Jan;20(1):41-5.
  20. Burkholder WJ. Use of body condition scores in clinical assessment of the provision of optimal nutrition.. J Am Vet Med Assoc 2000 Sep 1;217(5):650-4.
    doi: 10.2460/javma.2000.217.650pubmed: 10976293google scholar: lookup
  21. Pritchard JC, Lindberg AC, Main DC, Whay HR. Assessment of the welfare of working horses, mules and donkeys, using health and behaviour parameters.. Prev Vet Med 2005 Jul 12;69(3-4):265-83.
  22. Burn C., Pritchard J., Whay H. Observer reliability for working equine welfare assessment: Problems with high prevalenceu2019s of certain results. Anim. Welf. 2009;18:177u2013187. doi: 10.1017/S0962728600000324.
    doi: 10.1017/S0962728600000324google scholar: lookup
  23. Burn C.C., Dennison T.L., Whay H.R. Relationships between behaviour and health in working horses, donkeys, and mules in developing countries. Appl. Anim. Behav. Sci. 2010;126:109u2013118. doi: 10.1016/j.applanim.2010.06.007.
  24. Henneke DR, Potter GD, Kreider JL, Yeates BF. Relationship between condition score, physical measurements and body fat percentage in mares.. Equine Vet J 1983 Oct;15(4):371-2.
  25. Dugdale AH, Grove-White D, Curtis GC, Harris PA, Argo CM. Body condition scoring as a predictor of body fat in horses and ponies.. Vet J 2012 Nov;194(2):173-8.
    doi: 10.1016/j.tvjl.2012.03.024pubmed: 22578691google scholar: lookup
  26. Kirkden R.D., Pajor E.A. Using preference, motivation and aversion tests to ask scientific questions about animalsu2019 feelings. Appl. Anim. Behav. Sci. 2006;100:29u201347. doi: 10.1016/j.applanim.2006.04.009.
  27. Toates F.M. Motivational Systems. Cambridge University Press; Cambridge, UK: 1986.
  28. Alexander G. Cold thermogenesis.. Int Rev Physiol 1979;20:43-155.
    pubmed: 387638
  29. Cymbaluk NF, Christison GI. Environmental effects on thermoregulation and nutrition of horses.. Vet Clin North Am Equine Pract 1990 Aug;6(2):355-72.
    doi: 10.1016/S0749-0739(17)30546-1pubmed: 2202497google scholar: lookup
  30. Cymbaluk N.F. Thermoregulation of horses in cold, winter weather: A review. Livest. Prod. Sci. 1994;40:15u201371. doi: 10.1016/0301-6226(94)90266-6.
  31. EVANS CL, SMITH DF. Sweating responses in the horse.. Proc R Soc Lond B Biol Sci 1956 Mar 27;144(918):61-83.
    pubmed: 13310585doi: 10.1098/rspb.1956.0018google scholar: lookup
  32. Ashley FH, Waterman-Pearson AE, Whay HR. Behavioural assessment of pain in horses and donkeys: application to clinical practice and future studies.. Equine Vet J 2005 Nov;37(6):565-75.
    doi: 10.2746/042516405775314826pubmed: 16295937google scholar: lookup
  33. Mejdell C.M., Bu00f8e K.E. Responses to climatic variables of horses housed outdoors under Nordic winter conditions. Can. J. Anim. Sci. 2005;85:307u2013308. doi: 10.4141/A04-066.
    doi: 10.4141/A04-066google scholar: lookup
  34. Heleski C.R., Murtazashvili I. Daytime shelter-seeking behavior in domestic horses. J. Vet. Behav. 2010;5:276u2013282. doi: 10.1016/j.jveb.2010.01.003.
  35. Hausberger M., Fureix C., Lesimple C. Detecting horsesu2019 sickness: In search of visible signs. Appl. Anim. Behav. Sci. 2016;175:41u201349. doi: 10.1016/j.applanim.2015.09.005.
  36. IASP International Association for the Study of Pain. Pain. 1983;16:109u2013110.
    pubmed: 0
  37. Guthrie D.M. Neuroethology: An Introduction. Blackwell Scientific Publications; Oxford, UK: 1980.
  38. Murrell JC, Johnson CB. Neurophysiological techniques to assess pain in animals.. J Vet Pharmacol Ther 2006 Oct;29(5):325-35.
  39. Gregory N.G. Physiology and Behaviour of Animal Suffering. Blackwell Science; Oxford, UK: 2004.
    doi: 10.1002/9780470752494google scholar: lookup
  40. Mellor D.J., Cook C.J., Stafford K.J. Quantifying some responses to pain as a stressor. In: Moberg G.P., Mench J.A., editors. The Biology of Animal Stress: Basic Principles and Implications for Animal Welfare. CABI; Wallingford, UK: 2000. pp. 171u2013198.
  41. Ru00e9dua MA, Valadu00e3o CA, Duque JC, Balestrero LT. The pre-emptive effect of epidural ketamine on wound sensitivity in horses tested by using von Frey filaments.. Vet Anaesth Analg 2002 Oct;29(4):200-206.
  42. Olbrich VH, Mosing M. A comparison of the analgesic effects of caudal epidural methadone and lidocaine in the horse.. Vet Anaesth Analg 2003 Jul;30(3):156-64.
  43. 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/ani10040572pmc: PMC7222381pubmed: 32235343google scholar: lookup
  44. Lane J.G. A review of dental disorders of the horse, their treatment, and possible fresh approaches to management. Equine Vet. Educ. 1994;6:13u201321. doi: 10.1111/j.2042-3292.1994.tb01074.x.
  45. Easley J.K. Dental and oral examination. In: Baker G.J., Easley J.K., editors. Equine Dentistry. W.B Saunders Co.; London, UK: 1999. pp. 114u2013117.
  46. Graham BP. Dental care in the older horse.. Vet Clin North Am Equine Pract 2002 Dec;18(3):509-22.
    doi: 10.1016/S0749-0739(02)00031-7pubmed: 12516931google scholar: lookup
  47. Coneglian M.M., Borges T.D., Weber S.H., Bertagnon H.G., Michelotto P.V. Use of the horse grimace scale to identify and quantify pain due to dental disorders in horses. Appl. Anim. Behav. Sci. 2020;225:104970. doi: 10.1016/j.applanim.2020.104970.
  48. Haggard P, de Boer L. Oral somatosensory awareness.. Neurosci Biobehav Rev 2014 Nov;47:469-84.
  49. Mantyh PW. The neurobiology of skeletal pain.. Eur J Neurosci 2014 Feb;39(3):508-19.
    doi: 10.1111/ejn.12462pmc: PMC4453827pubmed: 24494689google scholar: lookup
  50. 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.
  51. 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
  52. Yang AY, Chow J, Liu J. Corneal Innervation and Sensation: The Eye and Beyond.. Yale J Biol Med 2018 Mar;91(1):13-21.
    pmc: PMC5872636pubmed: 29599653
  53. Brooks D.E., Matthews A.G. Chapter 25: Equine Ophthalmology. In: Gelatt K.N., editor. Veterinary Ophthalmology. 4th ed. Blackwell Pub; Ames, IA, USA: 2007. pp. 1165u20131274.
  54. Hood DM, Wagner IP, Taylor DD, Brumbaugh GW, Chaffin MK. Voluntary limb-load distribution in horses with acute and chronic laminitis.. Am J Vet Res 2001 Sep;62(9):1393-8.
    doi: 10.2460/ajvr.2001.62.1393pubmed: 11560266google scholar: lookup
  55. DuPreez P. Proceedings of the 41st British Equine Veterinary Association Congress. Equine Veterinary Journal Ltd.; Newmarket, ON, Canada: 2002. Pelvic fractures: Diagnosis and management; pp. 71u201372.
  56. Dyson S. Proceedings of the 41st British Equine Veterinary Association Congress. Equine Veterinary Journal Ltd.; Newmarket, ON, Canada: 2002. Itu2019s just the way I walk! A review of gait characteristics; p. 69.
  57. Dyson S, Marks D. Foot pain and the elusive diagnosis.. Vet Clin North Am Equine Pract 2003 Aug;19(2):531-65, viii.
    doi: 10.1016/S0749-0739(03)00004-Xpubmed: 14575173google scholar: lookup
  58. Dyson S, Murray R. Pain associated with the sacroiliac joint region: a clinical study of 74 horses.. Equine Vet J 2003 May;35(3):240-5.
    doi: 10.2746/042516403776148255pubmed: 12755425google scholar: lookup
  59. Dabareiner RM, Carter GK. Diagnosis, treatment, and farriery for horses with chronic heel pain.. Vet Clin North Am Equine Pract 2003 Aug;19(2):417-41.
    doi: 10.1016/S0749-0739(03)00025-7pubmed: 14575167google scholar: lookup
  60. Hewetson M. Proceedings of the 42nd British Equine Veterinary Association Congress. Equine Veterinary Journal Ltd.; Newmarket, ON, Canada: 2003. Reproducibility of a visual analogue scale, numerical rating scale and verbal rating scale for the assessment of lameness in the horse; p. 295.
  61. Bussiu00e8res G, Jacques C, Lainay O, Beauchamp G, Leblond A, Cadoru00e9 JL, Desmaiziu00e8res 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
  62. Stashak T.S. Examination for lameness. In: Stashak T.S., editor. Adamsu2019 Lameness in Horses. 5th ed. Lippincott, Williams and Wilkins; Philadelphia, PA, USA: 2002. pp. 113u2013183.
  63. Owens JG, Kamerling SG, Stanton SR, Keowen ML. Effects of ketoprofen and phenylbutazone on chronic hoof pain and lameness in the horse.. Equine Vet J 1995 Jul;27(4):296-300.
  64. Graubner C, Gerber V, Doherr M, Spadavecchia C. Clinical application and reliability of a post abdominal surgery pain assessment scale (PASPAS) in horses.. Vet J 2011 May;188(2):178-83.
    doi: 10.1016/j.tvjl.2010.04.029pubmed: 20627635google scholar: lookup
  65. de Grauw JC, van Loon JP. Systematic pain assessment in horses.. Vet J 2016 Mar;209:14-22.
    doi: 10.1016/j.tvjl.2015.07.030pubmed: 26831169google scholar: lookup
  66. Gregory N.G. Physiological mechanisms causing sickness behaviour and suffering in diseased animals. Anim. Welf. 1998;7:293u2013305. doi: 10.1017/S0962728600020728.
    doi: 10.1017/S0962728600020728google scholar: lookup
  67. Fureix C, Menguy H, Hausberger M. Partners with bad temper: reject or cure? A study of chronic pain and aggression in horses.. PLoS One 2010 Aug 26;5(8):e12434.
  68. Broom D.M. Welfare, stress, and the evolution of feelings. Adv. Study Behav. 1998;27:371u2013403. doi: 10.1016/S0065-3454(08)60369-1.
  69. Ledger RA, Mellor DJ. Forensic Use of the Five Domains Model for Assessing Suffering in Cases of Animal Cruelty.. Animals (Basel) 2018 Jun 25;8(7).
    doi: 10.3390/ani8070101pmc: PMC6071132pubmed: 29941781google scholar: lookup
  70. Wagner AE. Effects of stress on pain in horses and incorporating pain scales for equine practice.. Vet Clin North Am Equine Pract 2010 Dec;26(3):481-92.
    doi: 10.1016/j.cveq.2010.07.001pubmed: 21056295google scholar: lookup
  71. Taylor PM, Pascoe PJ, Mama KR. Diagnosing and treating pain in the horse. Where are we today?. Vet Clin North Am Equine Pract 2002 Apr;18(1):1-19, v.
    doi: 10.1016/S0749-0739(02)00009-3pubmed: 12064173google scholar: lookup
  72. Ransom J.I., Cade B.S. U.S. Geological Survey Techniques and Methods Report 2-A9. United States Geological Survey; Reston, VA, USA: 2009. Quantifying equid behavior: A research ethogram for free-roaming feral horses.
  73. McDonnell S.M. The Equid Ethogram: A Practical Field Guide to Horse Behaviour. CAB International; Cambridge, MA, USA: 2003.
  74. Kiley-Worthington M. The tail of ungulates, canids, and felids with particular reference to their causation and function as displays. Behavior. 1976;56:69u2013114. doi: 10.1163/156853976X00307.
    doi: 10.1163/156853976X00307google scholar: lookup
  75. Wolff A, Hausberger M, Le Scolan N. Experimental tests to assess emotionality in horses.. Behav Processes 1997 Sep;40(3):209-21.
    doi: 10.1016/S0376-6357(97)00784-5pubmed: 24895882google scholar: lookup
  76. Waring G. Horse Behavior. 2nd ed. Noyes Publications; Norwich, NY, USA: William Andrew Publishing; Norwich, NY, USA: 2003.
  77. Torcivia C, McDonnell S. Equine Discomfort Ethogram.. Animals (Basel) 2021 Feb 23;11(2).
    doi: 10.3390/ani11020580pmc: PMC7931104pubmed: 33672338google scholar: lookup
  78. Price J, Catriona S, Welsh EM, Waran NK. Preliminary evaluation of a behaviour-based system for assessment of post-operative pain in horses following arthroscopic surgery.. Vet Anaesth Analg 2003 Jul;30(3):124-37.
  79. Pritchett L.C., Ulibarri C., Roberts M.C., Schneider R.K., Sellon D.C. Identification of potential physiological and behavioral indicators of postoperative pain in horses after exploratory celiotomy for colic. Appl. Anim. Behav. Sci. 2003;80:31u201343. doi: 10.1016/S0168-1591(02)00205-8.
  80. Viu00f1uela-Fernu00e1ndez I, Jones E, Chase-Topping ME, Price J. Comparison of subjective scoring systems used to evaluate equine laminitis.. Vet J 2011 May;188(2):171-7.
    doi: 10.1016/j.tvjl.2010.05.011pubmed: 20541956google scholar: lookup
  81. Mellor D.J., Patterson-Kane E., Stafford K.J. The Sciences of Animal Welfare. Wiley-Blackwell; Oxford, UK: 2009. Sleep, Sleep Deprivation and the Welfare of Mature Animals; pp. 181u2013185.
  82. Fleming P. Nontraditional approaches to pain management.. Vet Clin North Am Equine Pract 2002 Apr;18(1):83-105, vii.
    doi: 10.1016/S0749-0739(01)00006-2pubmed: 12064184google scholar: lookup
  83. Dyson S., Berger J.M., Ellis A.D., Mullard J. Can the presence of musculoskeletal pain be determined from the facial expressions of ridden horses (FEReq)? J. Vet. Behav. 2017;19:78u201389. doi: 10.1016/j.jveb.2017.03.005.
  84. Wemelsfelder F. The scientific validity of subjective concepts in models of animal welfare. Appl. Anim. Behav. Sci. 1997;53:75u201388. doi: 10.1016/S0168-1591(96)01152-5.
  85. Popescu S., Diugan E.A. The relationship between behavioral and other welfare indicators of working horses. J. Equine Vet. Sci. 2013;33:1u201312. doi: 10.1016/j.jevs.2012.04.001.
  86. Lesimple C. Indicators of Horse Welfare: State-of-the-Art.. Animals (Basel) 2020 Feb 13;10(2).
    doi: 10.3390/ani10020294pmc: PMC7070675pubmed: 32069888google scholar: lookup
  87. Fureix C, Jego P, Henry S, Lansade L, Hausberger M. Towards an ethological animal model of depression? A study on horses.. PLoS One 2012;7(6):e39280.
  88. Henneke D.R., Potter G.D., Kreiger J.L. Body condition during pregnancy and lactation, and reproductive efficiency of mares. Theriogenology. 1984;21:897u2013909. doi: 10.1016/0093-691X(84)90383-2.
  89. De Kloet ER, Oitzl MS, Schu00f6bitz B. Cytokines and the brain corticosteroid receptor balance: relevance to pathophysiology of neuroendocrine-immune communication.. Psychoneuroendocrinology 1994;19(2):121-34.
    doi: 10.1016/0306-4530(94)90002-7pubmed: 8190832google scholar: lookup
  90. Beausoleil NJ, Mellor DJ. Introducing breathlessness as a significant animal welfare issue.. N Z Vet J 2015 Jan;63(1):44-51.
    doi: 10.1080/00480169.2014.940410pubmed: 25004795google scholar: lookup
  91. Mellor DJ, Beausoleil NJ. Equine Welfare during Exercise: An Evaluation of Breathing, Breathlessness and Bridles.. Animals (Basel) 2017 May 26;7(6).
    doi: 10.3390/ani7060041pmc: PMC5483604pubmed: 28587125google scholar: lookup
  92. Lansing RW, Gracely RH, Banzett RB. The multiple dimensions of dyspnea: review and hypotheses.. Respir Physiol Neurobiol 2009 May 30;167(1):53-60.
    doi: 10.1016/j.resp.2008.07.012pmc: PMC2763422pubmed: 18706531google scholar: lookup
  93. Parshall MB, Schwartzstein RM, Adams L, Banzett RB, Manning HL, Bourbeau J, Calverley PM, Gift AG, Harver A, Lareau SC, Mahler DA, Meek PM, O'Donnell DE. An official American Thoracic Society statement: update on the mechanisms, assessment, and management of dyspnea.. Am J Respir Crit Care Med 2012 Feb 15;185(4):435-52.
    doi: 10.1164/rccm.201111-2042STpmc: PMC5448624pubmed: 22336677google scholar: lookup
  94. Mellema M.S. The neurophysiology of dyspnea. J. Vet. Emerg. Crit. Care. 2008;18:561u2013571. doi: 10.1111/j.1476-4431.2008.00372.x.
  95. Banzett RB, Pedersen SH, Schwartzstein RM, Lansing RW. The affective dimension of laboratory dyspnea: air hunger is more unpleasant than work/effort.. Am J Respir Crit Care Med 2008 Jun 15;177(12):1384-90.
    doi: 10.1164/rccm.200711-1675OCpmc: PMC2427058pubmed: 18369200google scholar: lookup
  96. Gilbert FF, Gofton N. Terminal dives in mink, muskrat and beaver.. Physiol Behav 1982 May;28(5):835-40.
    doi: 10.1016/0031-9384(82)90200-1pubmed: 7048357google scholar: lookup
  97. Nehashi S, Nishino T, Ide T. Inhaled furosemide inhibits behavioral response to airway occlusion in anesthetized cats.. Anesthesiology 2001 Nov;95(5):1234-7.
  98. Niel L., Weary D.M. Rats avoid exposure to carbon dioxide and argon. Appl. Anim. Behav. Sci. 2007;107:100u2013109. doi: 10.1016/j.applanim.2006.08.002.
  99. O'Donnell DE, Ora J, Webb KA, Laveneziana P, Jensen D. Mechanisms of activity-related dyspnea in pulmonary diseases.. Respir Physiol Neurobiol 2009 May 30;167(1):116-32.
    doi: 10.1016/j.resp.2009.01.010pubmed: 19450767google scholar: lookup
  100. Dalmau A., Rodriguez P., Llonch P., Velarde A. Stunning pigs with different gas mixtures: Aversion in pigs. Anim. Welf. 2010;19:325u2013333. doi: 10.1017/S096272860000172X.
    doi: 10.1017/S096272860000172Xgoogle scholar: lookup
  101. Packer R.M.A., Hendricks A., Burn C.C. Do dog owners perceive the clinical signs related to conformation inherited disorders as u2019normalu2019 for the breed? A potential constraint to improving canine welfare. Anim. Welf. 2012;21:81u201393. doi: 10.7120/096272812X13345905673809.
  102. Buchanan GF, Richerson GB. Role of chemoreceptors in mediating dyspnea.. Respir Physiol Neurobiol 2009 May 30;167(1):9-19.
    doi: 10.1016/j.resp.2008.12.002pmc: PMC4486073pubmed: 19118647google scholar: lookup
  103. Davenport PW, Vovk A. Cortical and subcortical central neural pathways in respiratory sensations.. Respir Physiol Neurobiol 2009 May 30;167(1):72-86.
    doi: 10.1016/j.resp.2008.10.001pubmed: 18977463google scholar: lookup
  104. Evans KC. Cortico-limbic circuitry and the airways: insights from functional neuroimaging of respiratory afferents and efferents.. Biol Psychol 2010 Apr;84(1):13-25.
  105. Banzett RB, Mulnier HE, Murphy K, Rosen SD, Wise RJ, Adams L. Breathlessness in humans activates insular cortex.. Neuroreport 2000 Jul 14;11(10):2117-20.
  106. Evans KC, Banzett RB, Adams L, McKay L, Frackowiak RS, Corfield DR. BOLD fMRI identifies limbic, paralimbic, and cerebellar activation during air hunger.. J Neurophysiol 2002 Sep;88(3):1500-11.
    doi: 10.1152/jn.2002.88.3.1500pubmed: 12205170google scholar: lookup
  107. Peiffer C. Dyspnea and emotion: what can we learn from functional brain imaging?. Am J Respir Crit Care Med 2008 May 1;177(9):937-9.
    doi: 10.1164/rccm.200802-298EDpubmed: 18434331google scholar: lookup
  108. von Leupoldt A, Sommer T, Kegat S, Baumann HJ, Klose H, Dahme B, Bu00fcchel C. The unpleasantness of perceived dyspnea is processed in the anterior insula and amygdala.. Am J Respir Crit Care Med 2008 May 1;177(9):1026-32.
    doi: 10.1164/rccm.200712-1821OCpubmed: 18263796google scholar: lookup
  109. Mellor D.J., Beausoleil N.J. Ch 5: Moving beyond a problem-based focus on poor welfare towards creating opportunities to have positive welfare experiences. In: McMillan F.D., editor. Mental Health and Well-Being in Animals. 2nd ed. CAB International; Wallingford, UK: 2020. pp. 50u201366.
  110. McMillan FD. The psychobiology of social pain: Evidence for a neurocognitive overlap with physical pain and welfare implications for social animals with special attention to the domestic dog (Canis familiaris).. Physiol Behav 2016 Dec 1;167:154-171.
    doi: 10.1016/j.physbeh.2016.09.013pubmed: 27640131google scholar: lookup
  111. Alexander R.D. The evolution of social behavior. Annu. Rev. Ecol. Syst. 1974;5:325u2013383. doi: 10.1146/annurev.es.05.110174.001545.
  112. Cacioppo S, Capitanio JP, Cacioppo JT. Toward a neurology of loneliness.. Psychol Bull 2014 Nov;140(6):1464-504.
    doi: 10.1037/a0037618pmc: PMC5130107pubmed: 25222636google scholar: lookup
  113. Sondergaard E., Jensen M.B., Nical C.J. Motivation for social contact in horses measured by operant conditioning. Appl. Anim. Behav. Sci. 2011;132:131u2013137. doi: 10.1016/j.applanim.2011.04.007.
  114. Schatzmann U. Winter pasturing of sport horses in Switzerland: An experimental study. Equine Vet. J. Suppl. 1998;27:53u201354.
  115. Panksepp J. Affective consciousness: Core emotional feelings in animals and humans.. Conscious Cogn 2005 Mar;14(1):30-80.
    doi: 10.1016/j.concog.2004.10.004pubmed: 15766890google scholar: lookup
  116. 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
  117. Mellor DJ. Welfare-aligned Sentience: Enhanced Capacities to Experience, Interact, Anticipate, Choose and Survive.. Animals (Basel) 2019 Jul 13;9(7).
    doi: 10.3390/ani9070440pmc: PMC6680886pubmed: 31337042google scholar: lookup
  118. Mellor DJ. Positive animal welfare states and encouraging environment-focused and animal-to-animal interactive behaviours.. N Z Vet J 2015 Jan;63(1):9-16.
    doi: 10.1080/00480169.2014.926800pubmed: 24875367google scholar: lookup
  119. Mellor DJ. Enhancing animal welfare by creating opportunities for positive affective engagement.. N Z Vet J 2015 Jan;63(1):3-8.
    doi: 10.1080/00480169.2014.926799pubmed: 24875268google scholar: lookup
  120. Fraser D., Duncan I.J.H. u2018Pleasuresu2019, u2018painsu2019 and animal welfare; towards a natural history of affect. Anim. Welf. 1998;7:383u2013396. doi: 10.1017/S0962728600020935.
    doi: 10.1017/S0962728600020935google scholar: lookup
  121. Ikemoto S, Panksepp J. The role of nucleus accumbens dopamine in motivated behavior: a unifying interpretation with special reference to reward-seeking.. Brain Res Brain Res Rev 1999 Dec;31(1):6-41.
    doi: 10.1016/S0165-0173(99)00023-5pubmed: 10611493google scholar: lookup
  122. Spinka M, Newberry RC, Bekoff M. Mammalian play: training for the unexpected.. Q Rev Biol 2001 Jun;76(2):141-68.
    doi: 10.1086/393866pubmed: 11409050google scholar: lookup
  123. Scherer K.R., Dan E.S., Flykt A. What determines a feelingu2019s position in affective space? A case for appraisal. Cogn. Emot. 2006;20:92u2013113. doi: 10.1080/02699930500305016.
    doi: 10.1080/02699930500305016google scholar: lookup
  124. Yeates JW, Main DC. Assessment of positive welfare: a review.. Vet J 2008 Mar;175(3):293-300.
    doi: 10.1016/j.tvjl.2007.05.009pubmed: 17613265google scholar: lookup
  125. Held S.D.E., Spinka M. Animal play and animal welfare. Anim. Behav. 2011;81:891u2013899. doi: 10.1016/j.anbehav.2011.01.007.
  126. Grill HJ, Norgren R. The taste reactivity test. I. Mimetic responses to gustatory stimuli in neurologically normal rats.. Brain Res 1978 Mar 24;143(2):263-79.
    doi: 10.1016/0006-8993(78)90568-1pubmed: 630409google scholar: lookup
  127. Berridge KC. Food reward: brain substrates of wanting and liking.. Neurosci Biobehav Rev 1996;20(1):1-25.
    doi: 10.1016/0149-7634(95)00033-Bpubmed: 8622814google scholar: lookup
  128. Deag J.M. Behavioural ecology and the welfare of extensively farmed animals. Appl. Anim. Behav. Sci. 1996;49:9u201322. doi: 10.1016/0168-1591(95)00663-X.
  129. Steiner JE, Glaser D, Hawilo ME, Berridge KC. Comparative expression of hedonic impact: affective reactions to taste by human infants and other primates.. Neurosci Biobehav Rev 2001 Jan;25(1):53-74.
    doi: 10.1016/S0149-7634(00)00051-8pubmed: 11166078google scholar: lookup
  130. Kelley AE, Baldo BA, Pratt WE, Will MJ. Corticostriatal-hypothalamic circuitry and food motivation: integration of energy, action and reward.. Physiol Behav 2005 Dec 15;86(5):773-95.
    doi: 10.1016/j.physbeh.2005.08.066pubmed: 16289609google scholar: lookup
  131. Balcombe J.P. Animal pleasure and its moral significance. Appl. Anim. Behav. Sci. 2009;118:208u2013216. doi: 10.1016/j.applanim.2009.02.012.
  132. Ninomiya S., Sato S., Kusunose R., Mitumasu T., Obara Y. A note on a behavioural indicator of satisfaction in stabled horses. Appl. Anim. Behav. Sci. 2007;106:184u2013189. doi: 10.1016/j.applanim.2006.06.011.
  133. Ninomiya S., Kusunose R., Obara Y., Sato S. Effect of an open window and conspecifics within view on the welfare of stabled horses, estimated on the basis of positive and negative behavioural indicators. Anim. Welf. 2008;17:351u2013354. doi: 10.1017/S0962728600027846.
    doi: 10.1017/S0962728600027846google scholar: lookup
  134. Cabanac M. The experience of pleasure in animals. In: McMillan F.D., editor. Mental Health and Well-Being in Animals. Blackwell; Clive, IA, USA: 2005. pp. 29u201346.
  135. Freire R, Buckley P, Cooper JJ. Effects of different forms of exercise on post inhibitory rebound and unwanted behaviour in stabled horses.. Equine Vet J 2009 May;41(5):487-92.
    doi: 10.2746/095777309X383883pubmed: 19642410google scholar: lookup
  136. Chaplin SJ, Gretgrix L. Effect of housing conditions on activity and lying behaviour of horses.. Animal 2010 May;4(5):792-5.
    doi: 10.1017/S1751731109991704pubmed: 22444135google scholar: lookup
  137. u0160pinka M. Animal agency, animal awareness and animal welfare. Anim. Welf. 2019;28:11u201320. doi: 10.7120/09627286.28.1.011.
    doi: 10.7120/09627286.28.1.011google scholar: lookup
  138. Spinka M., Wemelsfelder F. Environmental challenge and animal agency. In: Appleby M.C., Mench J.A., Olsson I.A.S., Hughes B.O., editors. Animal Welfare. 2nd ed. CAB International; Wallingford, UK: 2011. pp. 27u201343.
  139. Panksepp J., Zellner M.R. Towards a neurologically based unified theory of aggression. Int. Rev. Soc. Psychol. 2004;17:37u201361.
  140. Nelson EE, Panksepp J. Brain substrates of infant-mother attachment: contributions of opioids, oxytocin, and norepinephrine.. Neurosci Biobehav Rev 1998 May;22(3):437-52.
    doi: 10.1016/S0149-7634(97)00052-3pubmed: 9579331google scholar: lookup
  141. Carter C.S., Keverne E.B. The neurobiology of social affiliation and pair bonding. Horm. Brain Behav. 2002;1:299u2013337.
  142. Lim MM, Young LJ. Neuropeptidergic regulation of affiliative behavior and social bonding in animals.. Horm Behav 2006 Nov;50(4):506-17.
    doi: 10.1016/j.yhbeh.2006.06.028pubmed: 16890230google scholar: lookup
  143. Pfaff D.W. Drive: Neurobiological and Molecular Mechanisms of Sexual Behavior. MIT Press; Cambridge, MA, USA: 1999.
  144. Fisher HE, Aron A, Brown LL. Romantic love: a mammalian brain system for mate choice.. Philos Trans R Soc Lond B Biol Sci 2006 Dec 29;361(1476):2173-86.
    doi: 10.1098/rstb.2006.1938pmc: PMC1764845pubmed: 17118931google scholar: lookup
  145. Panksepp J. Emotional endophenotypes in evolutionary psychiatry.. Prog Neuropsychopharmacol Biol Psychiatry 2006 Jul;30(5):774-84.
    doi: 10.1016/j.pnpbp.2006.01.004pubmed: 16554114google scholar: lookup
  146. Vanderschuren LJ, Niesink RJ, Van Ree JM. The neurobiology of social play behavior in rats.. Neurosci Biobehav Rev 1997 May;21(3):309-26.
    doi: 10.1016/S0149-7634(96)00020-6pubmed: 9168267google scholar: lookup
  147. Burgdorf J, Panksepp J. The neurobiology of positive emotions.. Neurosci Biobehav Rev 2006;30(2):173-87.
  148. Mellor DJ. Positive animal welfare states and reference standards for welfare assessment.. N Z Vet J 2015 Jan;63(1):17-23.
    doi: 10.1080/00480169.2014.926802pubmed: 24875152google scholar: lookup
  149. McMillan F.D. Mental health and well-being benefits of social contact and social support in animals. In: McMillan F.D., editor. Mental Health and Well-Being in Animals. 2nd ed. CAB International; Wallingford, UK: 2019. pp. 96u2013110.
  150. Hausberger M, Lerch N, Guilbaud E, Stomp M, Grandgeorge M, Henry S, Lesimple C. On-Farm Welfare Assessment of Horses: The Risks of Putting the Cart before the Horse.. Animals (Basel) 2020 Feb 25;10(3).
    doi: 10.3390/ani10030371pmc: PMC7143857pubmed: 32106531google scholar: lookup
  151. Harley JJ, Stack JD, Braid H, McLennan KM, Stanley CR. Evaluation of the Feasibility, Reliability, and Repeatability of Welfare Indicators in Free-Roaming Horses: A Pilot Study.. Animals (Basel) 2021 Jul 2;11(7).
    doi: 10.3390/ani11071981pmc: PMC8300213pubmed: 34359108google scholar: lookup
  152. Harvey A.M. Ph.D. Thesis. University of Technology Sydney; Sydney, NSW, Australia: 2022. Wild Horse Welfare: Assessment and Associations with Population and Behavioural Ecology.

Citations

This article has been cited 0 times.