in

Calves fed more milk prioritise play over feeding in a hole-board test


Abstract

Dairy calves are typically feed-restricted. Depending on its severity, hunger can improve or impair animals’ cognitive abilities, but the effect of chronic feed restriction on calf cognition remains underexplored. We used a spatial foraging task (hole-board test) to assess whether chronic feed restriction impairs cognitive performance in dairy calves provided 6 L/day of milk (restricted) compared to calves provided up to 12 L/day (enhanced). Calves had to recall 4 milk reward locations among 15 possibilities; locations remained constant for 14 trials (initial learning), and were then changed and kept constant for 6 trials (re-learning). We measured calves’ ability to remember locations already visited within trials (working and general working memory) and recall reward locations between trials (reference memory). We also measured play behaviour to explore trade-offs between foraging and playing in the testing arena (larger than calves’ home-pen). Restricted calves had better working and reference memory, shorter latencies to reach the first bucket, and completed trials faster, indicating higher food motivation. Enhanced calves were less motivated to engage in the test but played more than restricted ones. This highlights how hunger can shift priorities to foraging. Future research should explore trade-offs between important behaviours like play and feeding to understand the value calves place on different activities.

Similar content being viewed by others

Effects of group housing and incremental hay supplementation in calf starters at different ages on growth performance, behavior, and health

Effects of reward type and previous social experience on cognitive testing outcomes of weaned dairy calves

Effects of different milk feeding levels and frequencies on performance of Holstein heifers during weaning and first lactation

Data availability

Data is provided within the manuscript or supplementary information files.

References

  1. Mahendran, S., Wathes, D., Booth, R. & Blackie, N. A survey of calf management practices and farmer perceptions of calf housing in UK dairy herds. J. Dairy Sci. 105, 409–423. https://doi.org/10.3168/jds.2021-20638 (2022).

    Google Scholar 

  2. Appleby, M. C., Weary, D. M. & Chua, B. Performance and feeding behaviour of calves on ad libitum milk from artificial teats. Appl. Anim. Behav. Sci. 74 (3), 191–201. https://doi.org/10.1016/S0168-1591(01)00171-X (2001).

    Google Scholar 

  3. Miller-Cushon, E. K., Bergeron, R., Leslie, K. E. & DeVries, T. J. Effect of milk feeding level on development of feeding behavior in dairy calves. J. Dairy Sci. 96 (1), 551–564. https://doi.org/10.3168/JDS.2012-5937 (2013).

    Google Scholar 

  4. Welk, A., Otten, N. D. & Jensen, M. B. Invited review: The effect of milk feeding practices on dairy calf behavior, health, and performance—A systematic review. J. Dairy Sci. 106 (9), 5853–5879. https://doi.org/10.3168/JDS.2022-22900 (2023).

    Google Scholar 

  5. Khan, M. A., Weary, D. M. & Von Keyserlingk, M. A. G. Invited review: Effects of milk ration on solid feed intake, weaning, and performance in dairy heifers. J. Dairy Sci. 94 (3), 1071–1081. https://doi.org/10.3168/JDS.2010-3733 (2011).

    Google Scholar 

  6. De Paula Vieira, A., Guesdon, V., de Passille, A. M., von Keyserlingk, M. A. G. & Weary, D. M. Behavioural indicators of hunger in dairy calves. Appl. Anim. Behav. Sci. 109, 180–189. https://doi.org/10.1016/j.applanim.2007.03.006 (2008).

    Google Scholar 

  7. Ackermans, M. A., Jonker, N. C., Bennik, E. C. & de Jong, P. J. Hunger increases negative and decreases positive emotions in women with a healthy weight. Appetite 168, 105746. https://doi.org/10.1016/J.APPET.2021.105746 (2022).

    Google Scholar 

  8. hangry Oxford English Dictionary. (2023). https://doi.org/10.1093/OED/1081659877

  9. D’Eath, R. B., Tolkamp, B. J., Kyriazakis, I. & Lawrence, A. B. Freedom from hunger’ and preventing obesity: the animal welfare implications of reducing food quantity or quality. Anim. Behav. 77 (2), 275–288. https://doi.org/10.1016/J.ANBEHAV.2008.10.028 (2009).

    Google Scholar 

  10. Duncan, I. J. H. A Concept of Welfare Based on Feelings. Well-Being Farm. Animals: Challenges Solutions. 85–101. https://doi.org/10.1002/9780470344859.CH5 (2004).

  11. Paul, E. S., Harding, E. J. & Mendl, M. Measuring emotional processes in animals: the utility of a cognitive approach. Neurosci. Biobehavioral Reviews. 29 (3), 469–491. https://doi.org/10.1016/J.NEUBIOREV.2005.01.002 (2005).

    Google Scholar 

  12. Pizzagalli, D. A., Iosifescu, D., Hallett, L. A., Ratner, K. G. & Fava, M. Reduced hedonic capacity in major depressive disorder: Evidence from a probabilistic reward task. J. Psychiatr. Res. 43 (1), 76–87. https://doi.org/10.1016/J.JPSYCHIRES.2008.03.001 (2008).

    Google Scholar 

  13. Slaney, C. L., Hales, C. A. & Robinson, E. S. J. Rat models of reward deficits in psychiatric disorders. Curr. Opin. Behav. Sci. 22, 136–142. https://doi.org/10.1016/J.COBEHA.2018.05.001 (2018).

    Google Scholar 

  14. Baciadonna, L. & McElligott, A. G. The use of judgement bias to assess welfare in farm livestock. Anim Welf. 24 (1), 81–91. https://doi.org/10.7120/09627286.24.1.081 (2015).

    Google Scholar 

  15. Lagisz, M. et al. Optimism, pessimism and judgement bias in animals: A systematic review and meta-analysis. Neurosci. Biobehavioral Reviews. 118, 3–17. https://doi.org/10.1016/J.NEUBIOREV.2020.07.012 (2020).

    Google Scholar 

  16. Neville, V. et al. Pharmacological manipulations of judgement bias: A systematic review and meta-analysis. Neurosci. Biobehavioral Reviews. 108, 269–286. https://doi.org/10.1016/J.NEUBIOREV.2019.11.008 (2020).

    Google Scholar 

  17. Hinchcliffe, J. K. & Robinson, E. S. J. The Affective Bias Test and Reward Learning Assay: Neuropsychological Models for Depression Research and Investigating Antidepressant Treatments in Rodents. Curr. Protocols. 4 (6), e1057. https://doi.org/10.1002/CPZ1.1057 (2024).

    Google Scholar 

  18. van der Staay, F. J., Gieling, E. T., Pinzón, N. E., Nordquist, R. E. & Ohl, F. The appetitively motivated cognitive holeboard: A family of complex spatial discrimination tasks for assessing learning and memory. Neurosci. Biobehavioral Reviews. 36 (1), 379–403. https://doi.org/10.1016/J.NEUBIOREV.2011.07.008 (2012).

    Google Scholar 

  19. Conrad, C. D. A critical review of chronic stress effects on spatial learning and memory. Prog. Neuropsychopharmacol. Biol. Psychiatry. 34 (5), 742–755. https://doi.org/10.1016/J.PNPBP.2009.11.003 (2010).

    Google Scholar 

  20. Mendl, M., Burman, O., Laughlin, K. & Paul, E. Animal Memory and Animal Welfare. Anim Welf. 10 (S1), S141–S159. https://doi.org/10.1017/S0962728600023587 (2001).

    Google Scholar 

  21. Day, J. E. L., Kyriazakis, I. & Lawrence, A. B. The effect of food deprivation on the expression of foraging and exploratory behaviour in the growing pig. Appl. Anim. Behav. Sci. 42 (3), 193–206. https://doi.org/10.1016/0168-1591(95)93889-9 (1995).

    Google Scholar 

  22. Smith, N. K. & Grueter, B. A. Hunger-driven adaptive prioritization of behavior. FEBS J. 289 (4), 922–936. https://doi.org/10.1111/FEBS.15791 (2022).

    Google Scholar 

  23. Krashes, M. J. et al. Rapid, reversible activation of AgRP neurons drives feeding behavior in mice. J. Clin. Invest. 121 (4), 1424–1428. https://doi.org/10.1172/JCI46229 (2011).

    Google Scholar 

  24. Lawrence, A. B., Appleby, M. C. & Macleod, H. A. Measuring hunger in the pig using operant conditioning: The effect of food restriction. Anim. Sci. 47 (1), 131–137. https://doi.org/10.1017/S0003356100037132 (1988).

    Google Scholar 

  25. Mendl, M. Performing under pressure: stress and cognitive function. Appl. Anim. Behav. Sci. 65 (3), 221–244. https://doi.org/10.1016/S0168-1591(99)00088-X (1999).

    Google Scholar 

  26. Buckley, L. A. et al. Too hungry to learn? Hungry broiler breeders fail to learn a Y-maze food quantity discrimination task. Anim Welf. 20 (4), 469–481. https://doi.org/10.1017/S0962728600003110 (2011).

    Google Scholar 

  27. Fu, Y. et al. Food restriction affects Y-maze spatial recognition memory in developing mice. Int. J. Dev. Neurosci. 60, 8–15. https://doi.org/10.1016/j.ijdevneu.2017.03.010 (2017).

    Google Scholar 

  28. Lecorps, B., Woodroffe, R. E., von Keyserlingk, M. A. G. & Weary, D. M. Hunger affects cognitive performance of dairy calves. Biology Letter. 19, 20220475. https://doi.org/10.1098/rsbl.2022.0475 (2023).

    Google Scholar 

  29. Jensen, M. B. & Kyhn, R. Play behaviour in group-housed dairy calves, the effect of space allowance. Appl. Anim. Behav. Sci. 67 (1–2), 35–46. https://doi.org/10.1016/S0168-1591(99)00113-6 (2000).

    Google Scholar 

  30. Rushen, J. & de Passille, A. M. Locomotor play of veal calves in an arena: Are effects of feed level and spatial restriction mediated by responses to novelty? Appl. Anim. Behav. Sci. 155, 34–41. https://doi-org.bris.idm.oclc.org/10.1016/j.applanim.2014.03.009

  31. Sutherland, M. A., Worth, G. M., Schütz, K. E. & Stewart, M. Rearing substrate and space allowance influences locomotor play behaviour of dairy calves in an arena test. Appl. Anim. Behav. Sci. 154, 8–14. https://doi.org/10.1016/J.APPLANIM.2014.02.008 (2014).

    Google Scholar 

  32. Jensen, M. B., Duve, L. R. & Weary, D. M. Pair housing and enhanced milk allowance increase play behavior and improve performance in dairy calves. J. Dairy Sci. 98 (4), 2568–2575. https://doi.org/10.3168/JDS.2014-8272 (2015).

    Google Scholar 

  33. Krachun, C., Rushen, J. & Marie De Passillé, A. Play behaviour in dairy calves is reduced by weaning and by a low energy intake. Appl. Anim. Behav. Sci. 122, 71–76. https://doi.org/10.1016/j.applanim.2009.12.002 (2010).

    Google Scholar 

  34. Lecorps, B., Woodroffe, R. E., von Keyserlingk, M. A. G. & Weary, D. M. Assessing cognitive performance in dairy calves using a modified hole-board test. Anim. Cogn. 25 (5), 1365–1370. https://doi.org/10.1007/s10071-022-01617-5 (2022).

    Google Scholar 

  35. Yoo, S., von Keyserlingk, M. A. G. & Weary, D. M. The effects of pain following disbudding on calf memory. J. Dairy Sci. 106, 9507–9513. https://doi.org/10.3168/jds.2023-23604 (2023).

    Google Scholar 

  36. Friard, O. & Gamba, M. BORIS: a free, versatile open-source event-logging software for video/audio coding and live observations. Methods Ecol. Evol. 7, 1325–1330 (2016).

    Google Scholar 

  37. Posit team. RStudio: Integrated Development Environment for R. Posit Software (PBC, 2024). http://www.posit.co/

  38. Mazza, S., Frot, M. & Rey, A. E. A comprehensive literature review of chronic pain and memory. Progress Neuropsychopharmacol. Biol. Psychiatry. 87, 183–192. https://doi.org/10.1016/j.pnpbp.2017.08.006 (2018). https://doi.org/https://doi-org.bris.idm.oclc.

    Google Scholar 

  39. Johnston, I. N., Maier, S. F., Rudy, J. W. & Watkins, L. R. Post-conditioning experience with acute or chronic inflammatory pain reduces contextual fear conditioning in the rat. Behav. Brain. Res. 226 (2), 361–368. https://doi.org/10.1016/J.BBR.2011.08.048 (2012).

    Google Scholar 

  40. Grimberg-Henrici, C. G. E., Vermaak, P., Bolhuis, J. E., Nordquist, R. E. & Van Der Staay, F. J. Effects of environmental enrichment on cognitive performance of pigs in a spatial holeboard discrimination task. Anim. Cogn. 19, 271–283. https://doi.org/10.1007/s10071-015-0932-7 (2016).

    Google Scholar 

  41. Laughlin, K., Huck, M. & Mendl, M. Disturbance effects of environmental stimuli on pig spatial memory. Appl. Anim. Behav. Sci. 64 (3), 169–180. https://doi.org/10.1016/S0168-1591(99)00036-2 (1999).

    Google Scholar 

  42. Arts, J. W. M., van der Staay, F. J. & Ekkel, E. D. Working and reference memory of pigs in the spatial holeboard discrimination task. Behav. Brain. Res. 205 (1), 303–306. https://doi.org/10.1016/J.BBR.2009.06.014 (2009).

    Google Scholar 

  43. Tahamtani, F. M., Nordgreen, J., Nordquist, R. E. & Janczak, A. M. Early life in a barren environment adversely affects spatial cognition in laying hens (Gallus gallus domesticus). Front. Veterinary Sci. 2 (3). https://doi.org/10.3389/FVETS.2015.00003/BIBTEX (2015).

  44. Dumontier, L., Janczak, A. M., Smulders, T. V. & Nordgreen, J. Effects of the rearing environment complexity on laying hens’ spatial cognition: A holeboard test approach. Appl. Anim. Behav. Sci. 260, 105878. https://doi.org/10.1016/J.APPLANIM.2023.105878 (2023).

    Google Scholar 

  45. Franchi, G. A., Herskin, M. S. & Jensen, M. B. Dairy cows fed a low energy diet before dry-off show signs of hunger despite ad libitum access. Sci. Rep. 9 (1), 1–9. https://doi.org/10.1038/s41598-019-51866-7 (2019).

    Google Scholar 

  46. Schütz, K., Davison, D. & Matthews, L. Do different levels of moderate feed deprivation in dairy cows affect feeding motivation? Appl. Anim. Behav. Sci. 101 (3–4), 253–263. https://doi.org/10.1016/J.APPLANIM.2006.02.008 (2006).

    Google Scholar 

  47. van Horik, J. O. & Madden, J. R. A problem with problem solving: motivational traits, but not cognition, predict success on novel operant foraging tasks. Anim. Behav. 114, 189–198. https://doi.org/10.1016/J.ANBEHAV.2016.02.006 (2016).

    Google Scholar 

  48. Dorrance, B. R. & Zentall, T. R. Imitative learning in Japanese quail (Coturnix japonica) depends on the motivational state of the observer quail at the time of observation. J. Comp. Psychol. 115 (1), 62–67. https://doi.org/10.1037/0735-7036.115.1.62 (2001).

    Google Scholar 

  49. Verbeek, E., Ferguson, D. & Lee, C. Are hungry sheep more pessimistic? The effects of food restriction on cognitive bias and the involvement of ghrelin in its regulation. Physiol. Behav. 123, 67–75. https://doi.org/10.1016/J.PHYSBEH.2013.09.017 (2014).

    Google Scholar 

  50. Burnett, C. J. et al. Hunger-Driven Motivational State Competition. Neuron 92(1), 187–201. (2016). https://doi.org/10.1016/J.NEURON.2016.08.032/ATTACHMENT/BC41658A-7E5D-4258-8C5D-C202D16A5E5B/MMC2.PDF

  51. Muller-Schwarze, D., Stagge, B. & Muller-Schwarze, C. Play behavior: Persistence, decrease, and energetic compensation during food shortage in deer fawns. Science 215 (4528), 85–87. https://doi.org/10.1126/SCIENCE.215.4528.85 (1982).

    Google Scholar 

  52. Špinka, M., Newberry, R. C. & Bekoff, M. Mammalian play: Training for the unexpected. Q. Rev. Biology. 76 (2), 141–168. https://doi.org/10.1086/393866;WGROUP:STRING:PUBLICATION (2001).

    Google Scholar 

  53. Wallis, A., Mendl, M., Lecorps, B. & Held, S. From welfare indicator to welfare contributor: the role of play in building flexibility and resilience in captive animals. Proceedings B 292(2059), 20251962. (2025). https://doi.org/10.1098/rspb.2025.1962

  54. Ahloy-Dallaire, J., Espinosa, J. & Mason, G. Play and optimal welfare: Does play indicate the presence of positive affective states? Behavioural Processes 156, 3–15. (2018). https://doi.org/10.1016/J.BEPROC.2017.11.011

  55. Held, S. D. E. & Špinka, M. Animal play and animal welfare. Anim. Behav. 81 (5), 891–899. https://doi.org/10.1016/J.ANBEHAV.2011.01.007 (2011).

    Google Scholar 

  56. Mintline, E. M. et al. Play behavior as an indicator of animal welfare: Disbudding in dairy calves. Appl. Anim. Behav. Sci. 144 (1–2), 22–30. https://doi.org/10.1016/J.APPLANIM.2012.12.008 (2013).

    Google Scholar 

  57. Winder, C. B. et al. Clinical trial of local anesthetic protocols for acute pain associated with caustic paste disbudding in dairy calves. J. Dairy Sci. 100 (8), 6429–6441. https://doi.org/10.3168/JDS.2017-12724 (2017).

    Google Scholar 

  58. Sutton Hickey, A. K. & Krashes, M. J. Integrating Hunger with Rival Motivations. Trends Endocrinol. Metabolism. 31 (7), 495–507. https://doi.org/10.1016/J.TEM.2020.04.006 (2020).

    Google Scholar 

Download references

Acknowledgements

ACKNOWLEDGEMENTS We thank Celine Kretschmann (University of Münster), Xiyi Hu and Hui Wang (Shandong Agricultural University), Marko Palumbo (University of Bristol), Maria Fiori (University of Franche-Comté Besancon), the farmers at Wyndhurst Farm, and Langford Vets Farm Animal Practice for their help with this study.

Author information

Authors and Affiliations

Authors

Contributions

JH: Study conceptualisation, data collection and analysis, manuscript drafting and revisions. TL: Data collection and analysis, manuscript revisions. MTM: Study conceptualisation, manuscript revisions. BL: Study conceptualisation, manuscript revisions. All authors reviewed and approved the final version of the manuscript.

Corresponding author

Correspondence to
Benjamin Lecorps.

Ethics declarations

Competing interests

The authors declare no competing interests.

Additional information

Publisher’s note

Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Supplementary Information

Below is the link to the electronic supplementary material.

Supplementary Material 1 (download ZIP )

Rights and permissions

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.

Reprints and permissions

About this article

Cite this article

Hendricks, J., Leroy, T., Mendl, M.T. et al. Calves fed more milk prioritise play over feeding in a hole-board test.
Sci Rep (2026). https://doi.org/10.1038/s41598-026-47403-y

Download citation

  • Received:

  • Accepted:

  • Published:

  • DOI: https://doi.org/10.1038/s41598-026-47403-y

Keywords

  • animal welfare
  • animal wellbeing
  • dairy cattle
  • food motivation
  • positive animal welfare


Source: Ecology - nature.com

A fundamental model for oxygen consumption of Atlantic salmon

Smallholder farmers’ adaptation strategies among coffee farming household heads in the Kafa biosphere reserve, Ethiopia

Back to Top