FREE SHIPPING over $40
OVER 9000 FIVE-STAR REVIEWS
5% OFF ALL SUBSCRIPTIONS
100% HAPPINESS GUARANTEE
Analyze Diet
0
Home / Equine Research Bank / Sci Rep / Effect of horse sleep behavior on performance in a field-sid...
Scientific reports2026; 16(1); 4265; doi: 10.1038/s41598-025-34463-9

Effect of horse sleep behavior on performance in a field-side spatial reversal learning test.

Abstract: Reduced rapid-eye-movement (REM) sleep is associated with impaired learning in many species. We developed a reversal learning test (RLT) suitable for field conditions to explore this association in 16 healthy horses. Nocturnal REM-like sleep behavior was recorded five times for 48 h over a six-week period. The horses performed a target training task followed by an RLT using two objects. When the horses reached a predefined frequency of touching the object, the spatial location was reversed. Mean test parameters were statistically analysed using GENLIN models, longitudinal continuous variables were analysed using linear repeated measures models, and dichotomous repeated measures were analysed using GEE models and Kaplan-Meier method. Altogether 15/16 horses completed RLT by reversing three or more times. Most errors occurred before the second and third reversals. Overall, REM sleep duration (mean ± SE) was 46.1 ± 8.7 min. However, there were ten horses that exhibited REM-like sleep for less than 30 min (10.6 ± 2.2 min, range 0.0–28.0 min), while six horses exhibited REM-like sleep for at least 30 min (42.3 ± 4.8 min, range 36.6–65.8 min). Longer REM sleep was associated with a greater number of reversals ( = 0.04), while no relationship was found with error rate. Survival analysis further indicated a significant difference in progression probability between groups, with horses with shorter REM sleep having a 50% chance of progressing after five reversals, compared to six reversals for longer REM sleepers ( < 0.05). We present here a method to test horses’ reversal learning ability on site over a single day. We found that short REM-like sleep duration without clinical signs of sleep disturbances in horses was associated with poorer performance and lower perseverance during the RLT. The online version contains supplementary material available at 10.1038/s41598-025-34463-9.
Get Full Text
Publication Date: 2026-01-06 PubMed ID: 41495171PubMed Central: PMC12858803DOI: 10.1038/s41598-025-34463-9Google 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.
LinkedInCopy
  • Journal Article

Cite This Article

APA
Hämäläinen MJ, Brotherus IL, Wigren HM, Kaimio TE, Suomala H, Olbricht AM, Hänninen LT, Mykkänen AK. (2026). Effect of horse sleep behavior on performance in a field-side spatial reversal learning test. Sci Rep, 16(1), 4265. https://doi.org/10.1038/s41598-025-34463-9

Publication

Scientific reports
ISSN: 2045-2322
NlmUniqueID: 101563288
Country: England
Language: English
Volume: 16
Issue: 1
Pages: 4265
PII: 4265

Researcher Affiliations

Hämäläinen, Mira Joanna
  • Department of Equine and Small Animal Medicine, Faculty of Veterinary Medicine, University of Helsinki, Helsinki, 00014, Finland. mira.hamalainen@helsinki.fi.
Brotherus, Iina Liisa
  • Department of Equine and Small Animal Medicine, Faculty of Veterinary Medicine, University of Helsinki, Helsinki, 00014, Finland.
Wigren, Henna-Kaisa Margareta
  • Molecular and Integrative Biosciences Research Programme, Faculty of Biological and Environmental Sciences, University of Helsinki, Helsinki, 00014, Finland.
  • SLEEPWELL Research Program, Faculty of Medicine, University of Helsinki, 00014, Helsinki, Finland.
Kaimio, Tuire Eriikka
  • Vahviste, Vantaa, 01690, Finland.
Suomala, Heli
  • Department of Equine and Small Animal Medicine, Faculty of Veterinary Medicine, University of Helsinki, Helsinki, 00014, Finland.
  • Equine Information Centre, Kiuruvesi, 74700, Finland.
Olbricht, Anna-Mari
  • Equine Information Centre, Kiuruvesi, 74700, Finland.
Hänninen, Laura Talvikki
  • Department of Production Animal Medicine, Faculty of Veterinary Medicine, University of Helsinki, University of Helsinki, Helsinki, 00014, Finland.
Mykkänen, Anna Kristina
  • Department of Equine and Small Animal Medicine, Faculty of Veterinary Medicine, University of Helsinki, Helsinki, 00014, Finland.

Conflict of Interest Statement

Declarations. Competing interests: The authors declare no competing interests.

References

This article includes 74 references
  1. Brodt, S., Inostroza, M., Niethard, N. & Born, J. Sleep—A brain-state serving systems memory consolidation. , 1050–1075 (2023).n
    doi: 10.1016/j.neuron.2023.03.005pubmed: 37023710google scholar: lookup
  2. Diekelmann, S. & Born, J. The memory function of sleep. , 114–126 (2010).n
    doi: 10.1038/nrn2762pubmed: 20046194google scholar: lookup
  3. Hudson, A. N., Van Dongen, H. P. A. & Honn, K. A. Sleep deprivation, vigilant attention, and brain function: a review. , 21–30 (2020).n
    doi: 10.1038/s41386-019-0432-6pmc: PMC6879580pubmed: 31176308google scholar: lookup
  4. Tkachenko, O. & Dinges, D. F. Interindividual variability in neurobehavioral response to sleep loss: A comprehensive review. , 29–48 (2018).n
    doi: 10.1016/j.neubiorev.2018.03.017pubmed: 29563066google scholar: lookup
  5. Yuksel, C. et al. Both slow wave and rapid eye movement sleep contribute to emotional memory consolidation. , 485 (2025).
    pmc: PMC11930935pubmed: 40123003
  6. Boyce, R., Williams, S. & Adamantidis, A. REM sleep and memory. , 167–177 (2017).n
    doi: 10.1016/j.conb.2017.05.001pubmed: 28544929google scholar: lookup
  7. Ruckebusch, Y., Barbey, P. & Quillemot, P. Stages of sleep in the horse (Equus caballus). Les etats de sommeil Chez Le cheval (Equus caballus). , 658–665 (1970).
    pubmed: 4322113
  8. Belling, T. H. Sleep patterns in the horse. , 22–27 (1990).
  9. Williams, D. C. et al. Qualitative and quantitative characteristics of the electroencephalogram in normal horses during spontaneous drowsiness and sleep. , 630–638 (2008).n
    doi: 10.1111/j.1939-1676.2008.0096.xpubmed: 18466241google scholar: lookup
  10. Schuurman, S. O., Kersten, W. & Weijs, W. A. The equine Hind limb is actively stabilized during standing. , 355–362 (2003).n
    doi: 10.1046/j.1469-7580.2003.00166.xpmc: PMC1571089pubmed: 12739613google scholar: lookup
  11. Peever, J. & Fuller, P. M. The biology of REM sleep. , R1237–R1248 (2017).n
    doi: 10.1016/j.cub.2017.10.026pubmed: 29161567google scholar: lookup
  12. Dallaire, A., Rest & Behavior , 591–607 (1986).
    pubmed: 3492247
  13. Chung, E. L. T., Khairuddin, N. H., Azizan, T. R. P. T. & Adamu, L. Sleeping patterns of horses in selected local horse stables in Malaysia. , 1–4 (2018).
    doi: 10.1016/j.jveb.2018.03.014google scholar: lookup
  14. Kalus, M. (Ludwig-Maximilian University Munich, 2014).
  15. Lima, S. L., Rattenborg, N. C., Lesku, J. A. & Amlaner, C. J. Sleeping under the risk of predation. , 723–736 (2005).
    doi: 10.1016/j.anbehav.2005.01.008google scholar: lookup
  16. Lyamin, O. I., Siegel, J. M. & Sleep Giving it up to get it on. , R213–R216 (2024).n
    doi: 10.1016/j.cub.2024.01.061pubmed: 38471454google scholar: lookup
  17. Lyamin, O. I., Mukhametov, L. M. & Siegel, J. M. Sleep in the Northern fur seal. , 144–151 (2017).n
    doi: 10.1016/j.conb.2017.04.009pmc: PMC5609733pubmed: 28505502google scholar: lookup
  18. Raabymagle, P. & Ladewig, J. Lying behavior in horses in relation to box size. , 11–17 (2006).
    doi: 10.1016/j.jevs.2005.11.015google scholar: lookup
  19. Greening, L., Downing, J., Amiouny, D., Lekang, L. & McBride, S. The effect of altering routine husbandry factors on sleep duration and memory consolidation in the horse. , 105229 (2021).
    doi: 10.1016/j.applanim.2021.105229google scholar: lookup
  20. Kjellberg, L., Sassner, H. & Yngvesson, J. Horses’ resting behaviour in shelters of varying size compared with single boxes. , 105715 (2022).
    doi: 10.1016/j.applanim.2022.105715google scholar: lookup
  21. Suomala, H., Brotherus, I., Hänninen, L., Ternman, E. & Mykkänen, A. K. Risk factors associated with owner-reported sleep disturbances in nordic horses. 10.1111/evj.14560 (2025).
    pubmed: 40704824
  22. Barbosa, Â. P. et al. Sleep pattern interference in the cognitive performance of lusitano horses. , 334 (2024).n
    doi: 10.3390/ani14020334pmc: PMC10812765pubmed: 38275793google scholar: lookup
  23. Uddin, L. Q. Cognitive and behavioural flexibility: neural mechanisms and clinical considerations. , 167–179 (2021).n
    doi: 10.1038/s41583-021-00428-wpmc: PMC7856857pubmed: 33536614google scholar: lookup
  24. Bertone, J. J. Excessive drowsiness secondary to recumbent sleep deprivation in two horses. , 157–162 (2006).n
    doi: 10.1016/j.cveq.2005.12.020pubmed: 16627113google scholar: lookup
  25. Bertone, J. J. Sleep and sleep disorders in horses. in Equine Neurology (eds Furr, M. & Reed, S.) 123–129 (Wiley Blackwell, (2015).
  26. Haavisto, M. L. et al. Sleep restriction for the duration of a work week impairs multitasking performance: sleep restriction and multitasking. , 444–454 (2010).n
    doi: 10.1111/j.1365-2869.2010.00823.xpubmed: 20408942google scholar: lookup
  27. Honn, K. A., Hinson, J. M., Whitney, P. & Van Dongen, H. P. A. Cognitive flexibility: A distinct element of performance impairment due to sleep deprivation. , 191–197 (2019).n
    doi: 10.1016/j.aap.2018.02.013pubmed: 29549968google scholar: lookup
  28. Sun, X. et al. The effects of sleep deprivation on cognitive flexibility: a scoping review of outcomes and biological mechanisms. , 1626309. 10.3389/fnins.2025.1626309 (2025).n
    doi: 10.3389/fnins.2025.1626309pmc: PMC12321868pubmed: 40766906google scholar: lookup
  29. Izquierdo, A., Brigman, J. L., Radke, A. K., Rudebeck, P. H. & Holmes, A. The neural basis of reversal learning: an updated perspective. , 12–26 (2017).n
    doi: 10.1016/j.neuroscience.2016.03.021pmc: PMC5018909pubmed: 26979052google scholar: lookup
  30. Clark, L., Cools, R. & Robbins, T. W. The neuropsychology of ventral prefrontal cortex: Decision-making and reversal learning. , 41–53 (2004).n
    doi: 10.1016/S0278-2626(03)00284-7pubmed: 15134842google scholar: lookup
  31. Pavlov, I. P. Conditioned reflexes: an investigation of the physiological activity of the cerebral cortex. . , 1090 (1930).
    doi: 10.1001/archneurpsyc.1930.02220110256018pmc: PMC4116985pubmed: 25205891google scholar: lookup
  32. Skinner, B. F. The behavior of organisms at 50. In (B. F. Skinner Foundation, 2021).
  33. Harlow, H. F. The formation of learning sets. , 51–65 (1949).
    doi: 10.1037/h0062474pubmed: 18124807google scholar: lookup
  34. Bitterman, M. E. The evolution of intelligence. , 92–101 (1965).n
    doi: 10.1038/scientificamerican0165-92pubmed: 14252463google scholar: lookup
  35. Berg, E. L. & Silverman, J. L. Animal models of autism. in 157–196Academic Press, (2022).
  36. Warren, J. M. & Warren, H. B. Reversal learning by horse and raccoon. , 215–220 (1962).n
    doi: 10.1080/00221325.1962.10533590pubmed: 14005044google scholar: lookup
  37. Voith, V. L. (The Ohio State University, 1975).
  38. Fiske, J. & Potter, G. Discrimination reversal learning in yearling horses. , 583–588 (1979).
    doi: 10.2527/jas1979.492583xgoogle scholar: lookup
  39. Martin, T. I., Zentall, T. R. & Lawrence, L. Simple discrimination reversals in the domestic horse (Equus caballus): effect of discriminative stimulus modality on learning to learn. , 328–338 (2006).
    doi: 10.1016/j.applanim.2006.02.011google scholar: lookup
  40. Mancini, N. et al. Reversal learning in drosophila larvae. , 424–435 (2019).n
    doi: 10.1101/lm.049510.119pmc: PMC6796787pubmed: 31615854google scholar: lookup
  41. Meagher, R. K. et al. Effects of degree and timing of social housing on reversal learning and response to novel objects in dairy calves. , e0132828 (2015).
    pmc: PMC4537137pubmed: 26274583
  42. Strobel, B. K., Schmidt, M. A., Harvey, D. O. & Davis, C. J. Image discrimination reversal learning is impaired by sleep deprivation in rats: cognitive rigidity or fatigue? , 1052441 (2022).
    pmc: PMC9713940pubmed: 36467979
  43. Ineichen, C. et al. Establishing a probabilistic reversal learning test in mice: evidence for the processes mediating reward-stay and punishment-shift behaviour and for their modulation by serotonin. , 1012–1021 (2012).n
    doi: 10.1016/j.neuropharm.2012.07.025pubmed: 22824190google scholar: lookup
  44. Butter, C. M. Perseveration in extinction and in discrimination reversal tasks following selective frontal ablations in Macaca mulatta. , 163–171 (1969).
    doi: 10.1016/0031-9384(69)90075-4google scholar: lookup
  45. Rolls, E. T., Hornak, J., Wade, D. & McGrath, J. Emotion-related learning in patients with social and emotional changes associated with frontal lobe damage. . , 1518–1524 (1994).n
    doi: 10.1136/jnnp.57.12.1518pmc: PMC1073235pubmed: 7798983google scholar: lookup
  46. Kratzer, D. D., Netherland, W. M., Pulse, R. E. & Baker, J. P. Maze learning in quarter horses. , 896–902 (1977).
    doi: 10.2527/jas1977.454896xgoogle scholar: lookup
  47. Mongillo, P. et al. Spatial reversal learning is impaired by age in pet dogs. . , 2273–2282 (2013).
    doi: 10.1007/s11357-013-9524-0pmc: PMC3824977pubmed: 23529504google scholar: lookup
  48. Flannery, B. Relational discrimination learning in horses. , 267–280 (1997).
    doi: 10.1016/S0168-1591(97)00006-3google scholar: lookup
  49. Vigouroux, L., Jolivald, A., Ijichi, C. & Yarnell, K. 89 learning ability and physiological stress response of horses (Equus caballus) undergoing discrimination and reversal learning tasks using a touch screen. , 103552 (2022).
    doi: 10.1016/j.jevs.2021.103552google scholar: lookup
  50. McLean, A. N. & Christensen, J. W. The application of learning theory in horse training. , 18–27 (2017).
    doi: 10.1016/j.applanim.2017.02.020google scholar: lookup
  51. Sappington, B. K. F., McCall, C. A., Coleman, D. A., Kuhlers, D. L. & Lishak, R. S. A preliminary study of the relationship between discrimination reversal learning and performance tasks in yearling and 2-year-old horses. , 157–166 (1997).
    doi: 10.1016/S0168-1591(96)01157-4google scholar: lookup
  52. Samhita, L. & Gross, H. J. The ‘Clever Hans phenomenon’ revisited. , e27122 (2013).
    pmc: PMC3921203pubmed: 24563716
  53. Kaimio, T. & Tallberg, M. Hevosen käyttäytyminen (Behaviour of the horse). in Hevosen Kanssa 108–109 (WSOY, (2012).
  54. Hall, C. A., Cassaday, H. J. & Derrington, A. M. The effect of stimulus height on visual discrimination in horses. , 1715–1720 (2003).n
    doi: 10.2527/2003.8171715xpubmed: 12854807google scholar: lookup
  55. Blackmore, T. L., Foster, T. M., Sumpter, C. E. & Temple, W. An investigation of colour discrimination with horses (Equus caballus). , 387–396 (2008).
    doi: 10.1016/j.beproc.2008.02.003pubmed: 18359171google scholar: lookup
  56. Tomonaga, M. et al. A horse’s eye view: size and shape discrimination compared with other mammals. , 20150701 (2015).
    pmc: PMC4685539pubmed: 26601679
  57. Waring, G. H. 175–223 (Noyes, 1983).
  58. McDonnell, S. (A Division of the Blood-Horse, Inc., 2003).
  59. Wilson, R. C., Shenhav, A., Straccia, M. & Cohen, J. D. The 85% Rule for optimal learning. , 4646 (2019).
    pmc: PMC6831579pubmed: 31690723
  60. Buechel, S. D., Boussard, A., Kotrschal, A. & van Der Bijl, W. & Kolm, N. Brain size affects performance in a reversal-learning test. 285, (2018).
    pmc: PMC5805926pubmed: 29367391
  61. Pannewitz, L. & Loftus, L. Frustration in horses: investigating expert opinion on behavioural indicators and causes using a Delphi consultation. , 105818 (2023).
  62. Hartman, N. & Greening, L. M. A preliminary study investigating the influence of auditory stimulation on the occurrence of nocturnal equine Sleep-Related behavior in stabled horses. , 102782 (2019).
    pubmed: 31732116
  63. Durmer, J. S. & Dinges, D. F. Neurocognitive consequences of sleep deprivation. , 117–129 (2005).n
    doi: 10.1055/s-2005-867080pubmed: 15798944google scholar: lookup
  64. Johnsson, R. D. et al. Sleep loss impairs cognitive performance and alters song output in Australian magpies. , 6645 (2022).n
    doi: 10.1038/s41598-022-10162-7pmc: PMC9033856pubmed: 35459249google scholar: lookup
  65. Pinheiro-da-Silva, J., Tran, S. & Luchiari, A. C. Sleep deprivation impairs cognitive performance in zebrafish: A matter of fact? , 656–663 (2018).
    doi: 10.1016/j.beproc.2018.04.004pubmed: 29656092google scholar: lookup
  66. Samson, D. R., Vining, A. & Nunn, C. L. Sleep influences cognitive performance in lemurs. , 697–706 (2019).n
    doi: 10.1007/s10071-019-01266-1pubmed: 31055705google scholar: lookup
  67. Skinner, B. F. (B. F. Skinner Foundation, 1939).
  68. Sherry, C. J., Walters, T. J. & Rodney, G. G. Henry P.J. Behavioral Chaining in the goat (Capra hircus). , 241–251 (1994).
    doi: 10.1016/0168-1591(94)90065-5google scholar: lookup
  69. Skinner, B. F. Are theories of learning necessary? , 193–216 (1950).n
    doi: 10.1037/h0054367pubmed: 15440996google scholar: lookup
  70. Junttila, S. et al. Breed differences in social cognition, inhibitory control, and Spatial problem-solving ability in the domestic dog (). , 22529. 10.1038/s41598-022-26991-5 (2022).n
    doi: 10.1038/s41598-022-26991-5pmc: PMC9800387pubmed: 36581704google scholar: lookup
  71. Martin, P. & Bateson, P. (Cambridge University, 2007).
  72. Stamps, J. Motor learning and the value of familiar space. , 41–58 (1995).
    doi: 10.1086/285786google scholar: lookup
  73. Baragli, P. et al. Consistency and flexibility in solving Spatial tasks: different horses show different cognitive styles. , 16557 (2017).n
    doi: 10.1038/s41598-017-16729-zpmc: PMC5707407pubmed: 29185468google scholar: lookup
  74. Valenchon, M., Lévy, F., Górecka-Bruzda, A., Calandreau, L. & Lansade, L. Characterization of long-term memory, resistance to extinction, and influence of temperament during two instrumental tasks in horses. , 1001–1006 (2013).n
    doi: 10.1007/s10071-013-0648-5pubmed: 23743707google scholar: lookup

Citations

This article has been cited 0 times.
Subscribe to our newsletter
Join 99,001 horse owners like you who receive our email updates on horse health and nutrition.