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Frontiers in veterinary science2019; 6; 24; doi: 10.3389/fvets.2019.00024

Farm Animal Cognition-Linking Behavior, Welfare and Ethics.

Abstract: Farm animal welfare is a major concern for society and food production. To more accurately evaluate animal farming in general and to avoid exposing farm animals to poor welfare situations, it is necessary to understand not only their behavioral but also their cognitive needs and capacities. Thus, general knowledge of how farm animals perceive and interact with their environment is of major importance for a range of stakeholders, from citizens to politicians to cognitive ethologists to philosophers. This review aims to outline the current state of farm animal cognition research and focuses on ungulate livestock species, such as cattle, horses, pigs and small ruminants, and reflects upon a defined set of cognitive capacities (physical cognition: categorization, numerical ability, object permanence, reasoning, tool use; social cognition: individual discrimination and recognition, communication with humans, social learning, attribution of attention, prosociality, fairness). We identify a lack of information on certain aspects of physico-cognitive capacities in most farm animal species, such as numerosity discrimination and object permanence. This leads to further questions on how livestock comprehend their physical environment and understand causal relationships. Increasing our knowledge in this area will facilitate efforts to adjust husbandry systems and enrichment items to meet the needs and preferences of farm animals. Research in the socio-cognitive domain indicates that ungulate livestock possess sophisticated mental capacities, such as the discrimination between, and recognition of, conspecifics as well as human handlers using multiple modalities. Livestock also react to very subtle behavioral cues of conspecifics and humans. These socio-cognitive capacities can impact human-animal interactions during management practices and introduce ethical considerations on how to treat livestock in general. We emphasize the importance of gaining a better understanding of how livestock species interact with their physical and social environments, as this information can improve housing and management conditions and can be used to evaluate the use and treatment of animals during production.
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Publication Date: 2019-02-12 PubMed ID: 30838218PubMed Central: PMC6383588DOI: 10.3389/fvets.2019.00024Google Scholar: Lookup
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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 highlights the significance of better understanding the cognitive capacities of farm animals, such as cattle, horses, pigs, and small ruminants, to improve their welfare and contribute to ethical considerations in managing these animals. It also reflects on the current state of farm animal cognition research, identifying areas of cognition that require further study.

Farm Animal Cognition and its Importance

  • The article begins by expressing the necessity to thoroughly evaluate animal farming, which extends beyond behavioral observations and includes understanding the cognitive needs and capabilities of farm animals. The main motive behind this holistic approach is to prevent exposure of farm animals to poor welfare conditions.
  • Comprehending how animals perceive and interact with their surroundings is pivotal for various stakeholders, such as citizens, politicians, cognitive ethologists, and philosophers. Studying this can significantly improve animal farming strategies by catering to the animals’ cognitive needs.
  • The authors emphasize how animals’ physical cognition (like categorization, numerical ability, reasoning, tool use) and social cognition (like individual recognition, communication with humans, social learning) are important factors that need detailed exploration.

Farm Animal Cognition Research: Current State

  • The research focuses mainly on ungulate livestock species. The authors identify certain cognitive abilities like numerosity discrimination (understanding quantities) and object permanence (understanding that objects continue to exist even when they cannot be observed) where information is currently scarce.
  • The lack of such information invites further investigation into how livestock understand their physical environment and decipher causal relationships, which can be used to better tailor husbandry systems and enrichment items to the needs and preferences of these animals.

Socio-Cognitive Domain of Livestock

  • The article emphasizes that livestock possess advanced mental capacities like recognizing individuals among species and human handlers, and react to subtle behavioural cues. These abilities play a crucial role in human-animal interactions, affecting the management practices of these animals.
  • Understanding these socio-cognitive abilities also introduces important ethical considerations about the treatment of livestock.

Concluding Remarks

  • The article concludes by stressing the need to gain a deeper understanding of how livestock species interact with both their physical and social environments, which can further improve housing and management conditions.
  • Acquiring this knowledge will also be crucial in evaluating and revising the treatment and usage of animals during production, which further underlines the importance of a holistic approach towards animal farming.

Cite This Article

APA
Nawroth C, Langbein J, Coulon M, Gabor V, Oesterwind S, Benz-Schwarzburg J, von Borell E. (2019). Farm Animal Cognition-Linking Behavior, Welfare and Ethics. Front Vet Sci, 6, 24. https://doi.org/10.3389/fvets.2019.00024

Publication

Frontiers in veterinary science
ISSN: 2297-1769
NlmUniqueID: 101666658
Country: Switzerland
Language: English
Volume: 6
Pages: 24

Researcher Affiliations

Nawroth, Christian
  • Institute of Behavioural Physiology, Leibniz Institute for Farm Animal Biology (FBN), Dummerstorf, Germany.
  • Centre for Proper Housing of Ruminants and Pigs, Federal Food Safety and Veterinary Office, Agroscope Tänikon, Bern, Switzerland.
Langbein, Jan
  • Institute of Behavioural Physiology, Leibniz Institute for Farm Animal Biology (FBN), Dummerstorf, Germany.
Coulon, Marjorie
  • Cabinet EASIER, Clermont-Ferrand, France.
Gabor, Vivian
  • Department of Animal Sciences, University of Goettingen, Goettingen, Germany.
Oesterwind, Susann
  • Institute of Behavioural Physiology, Leibniz Institute for Farm Animal Biology (FBN), Dummerstorf, Germany.
  • Faculty of Agricultural and Environmental Sciences, University of Rostock, Rostock, Germany.
Benz-Schwarzburg, Judith
  • Unit for Ethics and Human-Animal Studies, Messerli Research Institute, Vetmeduni Vienna, University of Vienna, Medical University of Vienna, Vienna, Austria.
von Borell, Eberhard
  • Institute of Agricultural and Nutritional Sciences, Martin-Luther-University Halle-Wittenberg, Halle, Germany.

References

This article includes 223 references
  1. Farm Animal Welfare Council. The Five Freedoms. Surbiton: Farm Animal Welfare Council; (2013).
  2. Wathes C. Guarding the welfare of farm animals. Vet Rec (2010) 167:583–4.
    doi: 10.1136/vr.c4785pubmed: 21257423google scholar: lookup
  3. Špinka M. How important is natural behaviour in animal farming systems?. Appl Anim Behav Sci (2006) 100:117–28.
    doi: 10.1016/j.applanim.2006.04.006google scholar: lookup
  4. Désiré L, Boissy A, Veissier I. Emotions in farm animals: a new approach to animal welfare in applied ethology. Behav Process (2002) 60:165–80.
    doi: 10.1016/S0376-6357(02)00081-5pubmed: 12426068google scholar: lookup
  5. Mendl M, Burman OHP, Paul ES. An integrative and functional framework for the study of animal emotion and mood. Proc R Soc B (2010a) 277:2895–904.
    doi: 10.1098/rspb.2010.0303pmc: PMC2982018pubmed: 20685706google scholar: lookup
  6. Forkman B, Furuhaug IL, Jensen P. Personality, coping patterns, and aggression in piglets. Appl Anim Behav Sci (1995) 45:31–42.
    doi: 10.1016/0168-1591(95)00601-Ngoogle scholar: lookup
  7. Shettleworth SJ. Cognition, Evolution, and Behavior. Oxford: Oxford University Press; (2010).
  8. Tomasello M, Call J. Primate Cognition. 1st ed. New York, NY: Oxford University Press; (1997).
  9. Emery NJ. Cognitive ornithology: the evolution of avian intelligence. Phil Trans R Soc B (2006) 361:23–43.
    doi: 10.1098/rstb.2005.1736pmc: PMC1626540pubmed: 16553307google scholar: lookup
  10. Kaminski J, Nitzschner M. Do dogs get the point? A review of dog–human communication ability. Learn Motiv (2013) 44:294–302.
    doi: 10.1016/j.lmot.2013.05.001google scholar: lookup
  11. Shettleworth SJ. The evolution of comparative cognition: Is the snark still a boojum?. Behav Process (2009) 80:210–7.
    doi: 10.1016/j.beproc.2008.09.001pubmed: 18824222google scholar: lookup
  12. Coulon M, Baudoin C, Heyman Y, Deputte B. Cattle discriminate between familiar and unfamiliar conspecifics by using only head visual cues. Anim Cogn (2011) 14:279–90.
    doi: 10.1007/s10071-010-0361-6pubmed: 21132446google scholar: lookup
  13. Hanggi EB. Categorization learning in horses (Equus caballus). J Comp Psychol (1999) 113:243–52.
    doi: 10.1037/0735-7036.113.3.243google scholar: lookup
  14. Hanggi EB, Ingersoll JF. Long-term memory for categories and concepts in horses (Equus caballus). Anim Cogn (2009) 12:451–62.
    doi: 10.1007/s10071-008-0205-9pubmed: 19148689google scholar: lookup
  15. Ginane C, Dumont B. Do grazing sheep use species-based categorization to select their diet?. Behav Process (2010) 84:622–4.
    doi: 10.1016/j.beproc.2010.01.022pubmed: 20138975google scholar: lookup
  16. Meyer S, Nürnberg G, Puppe B, Langbein J. The cognitive capabilities of farm animals: categorisation learning in dwarf goats (Capra hircus). Anim Cogn (2012) 15:567–76.
    doi: 10.1007/s10071-012-0485-ypubmed: 22457061google scholar: lookup
  17. Gabor V, Gerken M. Shetland ponies (Equus caballus) show quantity discrimination in a matching-to-sample design. Anim Cogn (2014) 17:1233–43.
    doi: 10.1007/s10071-014-0753-0pubmed: 24831887google scholar: lookup
  18. Uller C, Lewis J. Horses (Equus caballus) select the greater of two quantities in small numerical contrasts. Anim Cogn (2009) 12:733–8.
    doi: 10.1007/s10071-009-0225-0pubmed: 19387706google scholar: lookup
  19. Proops L, Walton M, McComb K. The use of human-given cues by domestic horses, Equus caballus, during an object choice task. Anim Behav (2010) 79:1205–9.
    doi: 10.1016/j.anbehav.2010.02.015pubmed: 19588176google scholar: lookup
  20. Krueger K, Flauger B, Farmer K, Maros K. Horses (Equus caballus) use human local enhancement cues and adjust to human attention. Anim Cogn (2011) 14:187–201.
    doi: 10.1007/s10071-010-0352-7pubmed: 20845052google scholar: lookup
  21. Nawroth C, Ebersbach M, von Borell E. A note on pigs' knowledge of hidden objects. Arch Anim Breed (2013a) 56:861–72.
    doi: 10.7482/0003-9438-56-086google scholar: lookup
  22. Nawroth C, Ebersbach M, von Borell E. Juvenile domestic pigs (Sus scrofa domestica) use human-given cues in an object choice task. Anim Cogn (2014a) 17:701–13.
    doi: 10.1007/s10071-013-0702-3pubmed: 24197275google scholar: lookup
  23. Nawroth C, von Borell E, Langbein J. Object permanence in the dwarf goat (Capra aegagrus hircus): perseveration errors and the tracking of complex movements of hidden objects. Appl Anim Behav Sci (2015b) 167:20–6.
    doi: 10.1016/j.applanim.2015.03.010google scholar: lookup
  24. Nawroth C, von Borell E. Domestic pigs' (Sus scrofa domestica) use of direct and indirect visual and auditory cues in an object choice task. Anim Cogn (2015) 18:757–66.
    doi: 10.1007/s10071-015-0842-8pubmed: 25650328google scholar: lookup
  25. Nawroth C, von Borell E, Langbein J. Exclusion performance in dwarf goats (Capra aegagrus hircus) and sheep (Ovis orientalis aries). PLoS ONE (2014b) 9:e93534.
    doi: 10.1371/journal.pone.0093534pmc: PMC3973590pubmed: 24695781google scholar: lookup
  26. Coulon M, Deputte B, Heyman Y, Delatouche L. Visual discrimination by heifers (Bos taurus) of their own species. J Comp Psychol (2007) 121:198–204.
    doi: 10.1037/0735-7036.121.2.198pubmed: 17516798google scholar: lookup
  27. Krueger K, Flauger B. Olfactory recognition of individual competitors by means of faeces in horse (Equus caballus). Anim Cogn (2011) 14:245–57.
    doi: 10.1007/s10071-010-0358-1pubmed: 21132447google scholar: lookup
  28. Proops L, McComb K, Reby D. Cross-modal individual recognition in domestic horses (Equus caballus). Proc Natl Acad Sci USA (2009) 106:947–51.
    doi: 10.1073/pnas.0809127105pmc: PMC2630083pubmed: 19075246google scholar: lookup
  29. McLeman MA, Mendl M, Jones RB, White R, Wathes CM. Discrimination of conspecifics by juvenile domestic pigs, Sus scrofa. Anim Behav (2005) 70:451–61.
    doi: 10.1016/j.anbehav.2004.11.013google scholar: lookup
  30. McLeman MA, Mendl MT, Jones RB, Wathes CM. Social discrimination of familiar conspecifics by juvenile pigs, Sus scrofa: development of a non-invasive method to study the transmission of unimodal and bimodal cues between live stimuli. Appl Anim Behav Sci (2008) 115:123–37.
    doi: 10.1016/j.applanim.2008.06.010google scholar: lookup
  31. Kendrick KM, da Costa P, Leigh E, Hinton MR, Peirce JW. Sheep don't forget a face. Nature (2001) 414:165–6.
    doi: 10.1038/35102669pubmed: 11700543google scholar: lookup
  32. Keil NM, Imfeld-Mueller S, Aschwanden J, Wechsler B. Are head cues necessary for goats (Capra hircus) in recognising group members ?. Anim Cogn (2012) 15:913–21.
    doi: 10.1007/s10071-012-0518-6pubmed: 22644114google scholar: lookup
  33. Rybarczyk P, Koba Y, Rushen J, Tanida H, de Passillá AM. Can cows discriminate people by their faces?. Appl Anim Behav Sci (2001) 74:175–89.
    doi: 10.1016/S0168-1591(01)00162-9google scholar: lookup
  34. Stone SM. Human facial discrimination in horses: can they tell us apart?. Anim Cogn (2010) 13:51–61.
    doi: 10.1007/s10071-009-0244-xpubmed: 19533185google scholar: lookup
  35. Proops L, McComb K. Cross-modal individual recognition in domestic horses (Equus caballus) extends to familiar humans. Proc R Soc B (2012) 279:3131–8.
    doi: 10.1098/rspb.2012.0626pmc: PMC3385734pubmed: 22593108google scholar: lookup
  36. Koba Y, Tanida H. How do miniature pigs discriminate between people? The effect of exchanging cues between a non-handler and their familiar handler on discrimination. Appl Anim Behav Sci (1999) 61:239–52.
    doi: 10.1016/S0168-1591(98)00192-0google scholar: lookup
  37. Koba Y, Tanida H. How do miniature pigs discriminate between people? Discrimination between people wearing coveralls of the same colour. Appl Anim Behav Sci (2001) 73:45–58.
    doi: 10.1016/S0168-1591(01)00106-Xpubmed: 11356290google scholar: lookup
  38. Knolle F, Goncalves RP, Morton AJ. Sheep recognize familiar and unfamiliar human faces from two-dimensional images. R Soc Open Sci (2017) 4:171228.
    doi: 10.1098/rsos.171228pmc: PMC5717684pubmed: 29291109google scholar: lookup
  39. McKinley J, Sambrook TD. Use of human-given cues by domestic dogs (Canis familiaris) and horses (Equus caballus). Anim Cogn (2000) 3:13–22.
    doi: 10.1007/s100710050046google scholar: lookup
  40. Albiach-Serrano A, Bräuer J, Cacchione T, Zickert N, Amici F. The effect of domestication and ontogeny in swine cognition (Sus scrofa scrofa and S. s domestica). Appl Anim Behav Sci (2012) 141:25–35.
    doi: 10.1016/j.applanim.2012.07.005google scholar: lookup
  41. Kaminski J, Riedel J, Call J, Tomasello M. Domestic goats, Capra hircus, follow gaze direction and use social cues in an object choice task. Anim Behav (2005) 69:11–8.
    doi: 10.1016/j.anbehav.2004.05.008google scholar: lookup
  42. Nawroth C, von Borell E, Langbein J. “Goats that stare at men”: dwarf goats alter their behaviour in response to human head orientation, but do not spontaneously use head direction as a cue in a food-related context. Anim Cogn (2015a) 18:65–73.
    doi: 10.1007/s10071-014-0777-5pubmed: 24997158google scholar: lookup
  43. Malavasi R, Huber L. Evidence of heterospecific referential communication from domestic horses (Equus caballus) to humans. Anim Cogn (2016) 19:899–909.
    doi: 10.1007/s10071-016-0987-0pubmed: 27098164google scholar: lookup
  44. Nawroth C, Brett JM, McElligott AG. Goats display audience-dependent human-directed gazing behaviour in a problem-solving task. Biol Lett (2016b) 12:20160283.
    doi: 10.1098/rsbl.2016.0283pmc: PMC4971169pubmed: 27381884google scholar: lookup
  45. Langbein J, Krause A, Nawroth C. Human-directed behaviour in goats is not affected by short-term positive handling. Anim Cogn (2018) 21:795–803.
    doi: 10.1007/s10071-018-1211-1pubmed: 30173331google scholar: lookup
  46. Howery LD, Provenza FD, Banner RE, Scott CB. Social and environmental factors influence cattle distribution on rangeland. Appl Anim Behav Sci (1998) 55:231–44.
    doi: 10.1016/S0168-1591(97)00054-3google scholar: lookup
  47. Christensen JW. Early-life object exposure with a habituated mother reduces fear reactions in foals. Anim Cogn (2016) 19:171–9.
    doi: 10.1007/s10071-015-0924-7pubmed: 26395986google scholar: lookup
  48. Oostindjer M, Bolhuis JE, Mendl M, Held S, van den Brand H, Kemp B. Learning how to eat like a pig: effectiveness of mechanisms for vertical social learning in piglets. Anim Behav (2011) 82:503–11.
    doi: 10.1016/j.anbehav.2011.05.031google scholar: lookup
  49. Veit A, Wondrak M, Huber L. Object movement re-enactment in free-ranging Kune Kune piglets. Anim Behav (2017) 132:49–59.
    doi: 10.1016/j.anbehav.2017.08.004google scholar: lookup
  50. Glasser TA, Ungar ED, Landau SY, Perevolotsky A, Muklada H, Walker JW. Breed and maternal effects on the intake of tannin-rich browse by juvenile domestic goats (Capra hircus). Appl Anim Behav Sci (2009) 119:71–7.
    doi: 10.1016/j.applanim.2009.02.028google scholar: lookup
  51. Bailey DW, Howery LD, Boss DL. Effects of social facilitation for locating feeding sites by cattle in an eight-arm radial maze. Appl Anim Behav Sci (2000) 68:93–105.
    doi: 10.1016/S0168-1591(00)00091-5pubmed: 10771318google scholar: lookup
  52. Veissier I. Observational learning in cattle. Appl Anim Behav Sci (1993) 35:235–43.
    doi: 10.1016/0168-1591(93)90139-Ggoogle scholar: lookup
  53. Krueger K, Farmer K, Heinze J. The effects of age, rank and neophobia on social learning in horses. Anim Cogn (2014) 17:645–55.
    doi: 10.1007/s10071-013-0696-xpubmed: 24170136google scholar: lookup
  54. Rørvang MV, Ahrendt LP, Christensen JW. Horses fail to use social learning when solving spatial detour tasks. Anim Cogn (2015) 18:847–54.
    doi: 10.1007/s10071-015-0852-6pubmed: 25716720google scholar: lookup
  55. Figueroa J, Solà-Oriol D, Manteca X, Pérez JF. Social learning of feeding behaviour in pigs: effects of neophobia and familiarity with the demonstrator conspecific. Appl Anim Behav Sci (2013) 148:120–7.
    doi: 10.1016/j.applanim.2013.06.002google scholar: lookup
  56. Baciadonna L, McElligott AG, Briefer EF. Goats favour personal over social information in an experimental foraging task. PeerJ (2013) 1:e172.
    doi: 10.7717/peerj.172pmc: PMC3792185pubmed: 24109556google scholar: lookup
  57. Briefer EF, Haque S, Baciadonna L, McElligott AG. Goats excel at learning and remembering a highly novel cognitive task. Front Zool (2014) 11:20.
    doi: 10.1186/1742-9994-11-20pmc: PMC3987177pubmed: 24666734google scholar: lookup
  58. Burla JB, Siegwart J, Nawroth C. Human demonstration does not facilitate the performance of horses (Equus caballus) in a spatial problem-solving task. Animals (2018) 8:96.
    doi: 10.3390/ani8060096pmc: PMC6025305pubmed: 29899296google scholar: lookup
  59. Schuetz A, Farmer K, Krueger K. Social learning across species: horses (Equus caballus) learn from humans by observation. Anim Cogn (2017) 20:567–73.
    doi: 10.1007/s10071-016-1060-8pubmed: 27866286google scholar: lookup
  60. Nawroth C, Baciadonna L, McElligott AG. Goats learn socially from humans in a spatial problem-solving task. Anim Behav (2016a) 121:123–9.
    doi: 10.1016/j.anbehav.2016.09.004google scholar: lookup
  61. Proops L, McComb K. Attributing attention: the use of human-given cues by domestic horses (Equus caballus). Anim Cogn (2010) 13:197–205.
    doi: 10.1007/s10071-009-0257-5pubmed: 19588176google scholar: lookup
  62. Held S, Mendl M, Devereux C, Byrne RW. Behaviour of domestic pigs in a visual perspective taking task. Behaviour (2001) 138:1337–54.
    doi: 10.1163/156853901317367627google scholar: lookup
  63. Held S, Mendl M, Devereux C, Byrne RW. Foraging pigs alter their behaviour in response to exploitation. Anim Behav (2002a) 64:157–66.
    doi: 10.1006/anbe.2002.3044google scholar: lookup
  64. Held S, Mendl M, Laughlin K, Byrne RW. Cognition studies with pigs: livestock cognition and its implication for production. J Anim Sci (2002b) 80:E10–17.
    doi: 10.2527/animalsci2002.0021881200800ES10003xgoogle scholar: lookup
  65. Nawroth C, Ebersbach M, von Borell E. Are juvenile domestic pigs (Sus scrofa domestica) sensitive to the attentive states of humans? - The impact of impulsivity on choice behaviour. Behav Process (2013b) 96:53–8.
    doi: 10.1016/j.beproc.2013.03.002pubmed: 23500190google scholar: lookup
  66. Nawroth C, McElligott AG. Human head orientation and eye visibility as indicators of attention for goats (Capra hircus). PeerJ (2017) 5:e3073.
    doi: 10.7717/peerj.3073pmc: PMC5346283pubmed: 28289568google scholar: lookup
  67. Zentall TR, Galizio M, Critchfied TS. Categorization, concept learning, and behavior analysis: an introduction. J Exp Anal Behav (2002) 78:237–48.
    doi: 10.1901/jeab.2002.78-237pmc: PMC1284898pubmed: 12507002google scholar: lookup
  68. Pepperberg I. Grey parrot numerical competence: a review. Anim Cogn (2006) 9:377–91.
    doi: 10.1007/s10071-006-0034-7pubmed: 16909236google scholar: lookup
  69. Feigenson L, Dehaene S, Spelke E. Core systems of number. Trends Cogn Sci (2004) 8:307–14.
    doi: 10.1016/j.tics.2004.05.002pubmed: 15242690google scholar: lookup
  70. Jaakkola K. Do animals understand invisible displacement? A Critical Review. J Comp Psychol (2014) 128:1–15.
    doi: 10.1037/a0035675pubmed: 24611640google scholar: lookup
  71. Premack D. Cause/induced motion: intention/spontaneous motion. In: Changeux JP, Chavaillon J, editors. Origins of the Human Brain. Oxford: Oxford University Press; (1995). p. 286–309.
  72. Heyes CM. Anecdotes, training, trapping and triangulating: do animals attribute mental states?. Anim Behav (1993) 46:177–88.
    doi: 10.1006/anbe.1993.1173google scholar: lookup
  73. Penn DC, Povinelli DJ. Causal cognition in human and nonhuman animals: a comparative, critical review. Annu Rev Psychol (2007) 58:97–118.
    doi: 10.1146/annurev.psych.58.110405.085555pubmed: 17029564google scholar: lookup
  74. Bentley-Condit V, Smith EO. Animal tool use: current definitions and an updated comprehensive catalog. Behaviour (2010) 147:185–32A.
    doi: 10.1163/000579509X12512865686555google scholar: lookup
  75. Vaesen K. The cognitive bases of human tool use. Behav Brain Sci (2012) 35:203–18.
    doi: 10.1017/S0140525X11001452pubmed: 22697258google scholar: lookup
  76. Barton RA, Dunbar RIM. Evolution of the social brain. In: Whiten A, Byrne RW, editors. Machiavellian Intelligence II. Cambridge: Cambridge University Press; (1997). p. 240–63.
    doi: 10.1017/CBO9780511525636.010google scholar: lookup
  77. Tibbetts EA, Dale J. Individual recognition: it is good to be different. Trends Ecol Evol (2007) 22:529–37.
    doi: 10.1016/j.tree.2007.09.001pubmed: 17904686google scholar: lookup
  78. Adachi I. Cross-modal representations in primates and dogs: a new framework of recognition of social objects. Interact Stud (2009) 10:225–51.
    doi: 10.1075/is.10.2.07adagoogle scholar: lookup
  79. Bradbury JW, Vehrencamp SL. Principles of Animal Communication. Sunderland, MA: Sinauer Associates, Inc; (1998).
  80. Marshall-Pescini S, Colombo E, Passalacqua C, Merola I, Prato-Previde E. Gaze alternation in dogs and toddlers in an unsolvable task: evidence of an audience effect. Anim Cogn (2013) 16:933–43.
    doi: 10.1007/s10071-013-0627-xpubmed: 23543361google scholar: lookup
  81. Miklósi Á, Kubinyi E, Topál J, Gácsi M, Virányi Z, Csányi V. A simple reason for a big difference: wolves do not look back at humans, but dogs do. Curr Biol (2003) 13:763–6.
    doi: 10.1016/S0960-9822(03)00263-Xpubmed: 12725735google scholar: lookup
  82. Savalli C, Ades C, Gaunet F. Are dogs able to communicate with their owners about a desirable food in a referential and intentional way?. PLoS ONE (2014) 9:e108003.
    doi: 10.1371/journal.pone.0108003pmc: PMC4169500pubmed: 25232956google scholar: lookup
  83. Rørvang MV, Christensen JW, Ladewig J, McLean A. Social learning in horses—fact or fiction?. Front Vet Sci (2018) 5:212.
    doi: 10.3389/fvets.2018.00212pmc: PMC6135911pubmed: 30238009google scholar: lookup
  84. Huber L, Range F, Voelkl B, Szucsich A, Virányi Z, Miklosi A. The evolution of imitation: what do the capacities of non-human animals tell us about the mechanisms of imitation?. Phil Trans R Soc B (2009) 364:2299–309.
    doi: 10.1098/rstb.2009.0060pmc: PMC2865073pubmed: 19620102google scholar: lookup
  85. Galef BG, Laland KN. Social learning in animals: empirical studies and theoretical models. Bioscience (2005) 55:489.
    doi: 10.1641/0006-3568(google scholar: lookup
  86. Heyes CM. Social learning in animals: categories and mechanisms. Biol Rev (1994) 69:207–31.
    doi: 10.1111/j.1469-185X.1994.tb01506.xpubmed: 8054445google scholar: lookup
  87. Truskanov N, Shy R, Lotem A. Context-specific learning and its implications for social learning. Behav Ecol (2018) 29:1046–55.
    doi: 10.1093/beheco/ary078google scholar: lookup
  88. Call J, Tomasello M. Does the chimpanzee have a theory of mind? 30 years later. Trends Cogn Sci (2008) 12:187–92.
    doi: 10.1016/j.tics.2008.02.010pubmed: 18424224google scholar: lookup
  89. Horner V, Carter JD, Suchak M, de Waal FBM. Spontaneous prosocial choice by chimpanzees. Proc Natl Acad Sci USA (2011) 108:13847–51.
    doi: 10.1073/pnas.1111088108pmc: PMC3158226pubmed: 21825175google scholar: lookup
  90. Brosnan SF, De Waal FBM. Monkeys reject unequal pay. Nature (2003) 435:297–9.
    doi: 10.1038/nature01963pubmed: 13679918google scholar: lookup
  91. Rault JL. Be kind to others: prosocial behaviours and their implications for animal welfare. Appl Anim Behav Sci (2019) 210:113–23.
    doi: 10.1016/J.APPLANIM.2018.10.015google scholar: lookup
  92. Bailey DW, Sims PL. Association of food quality and locations by cattle. J Range Manag (1998) 51:2–6.
    doi: 10.2307/4003555google scholar: lookup
  93. Howery LD, Bailey DW, Ruyle GB, Renken WJ. Cattle use visual cues to track food locations. Appl Anim Behav Sci (2000) 67:1–14.
    doi: 10.1016/S0168-1591(99)00118-5pubmed: 10719185google scholar: lookup
  94. Bouissou MF, Andrieu S. Etablissement des relations préférentielles chez les bovins domestiques. Behaviour (1978) 64:148–57.
    doi: 10.1163/156853978X00486google scholar: lookup
  95. Bouissou MF, Andrieu S. Etablissement des relations de dominance-soumission chez les bovins domestiques. Biol Behav (1977) 2:97–107.
  96. Le Neindre P. Influence of cattle rearing conditions and breed on social relationships of mother and young. Appl Anim Behav Sci (1989) 23:117–27.
    doi: 10.1016/0168-1591(89)90012-9google scholar: lookup
  97. Hagen K, Broom DM. Cattle discriminate between individual familiar herd members in a learning experiment. Appl Anim Behav Sci (2003) 82:13–28.
    doi: 10.1016/S0168-1591(03)00053-4google scholar: lookup
  98. Baldwin BA. Ability of goats and calves to distinguish between conspecific urine samples using olfaction. Appl Anim Ethol (1977) 3:145–50.
    doi: 10.1016/0304-3762(77)90023-2pubmed: 1197391google scholar: lookup
  99. Cummins KA. Olfactory and visual cues. Individual recognition and social aggression in lactating cows. J Dairy Sci (1991) 74:301.
  100. Lensink J, Leruste H, Tillie M. L'Observation du troupeau bovin : Voir, interpréter, agir. Fr Agric. (2006). Available online at: https://www.unitheque.com/Livre/france_agricole/Agri_production/L_observation_du_troupeau_bovin-58428.html.
  101. Coulon M, Baudoin C, Abdi H, Heyman Y, Deputte BL. Social behavior and kin discrimination in a mixed group of cloned and non cloned heifers (Bos taurus). Theriogenology (2010) 74:1596–603.
    doi: 10.1016/j.theriogenology.2010.06.031pubmed: 20708240google scholar: lookup
  102. Taylor A, Davis H. Individual humans as discriminative stimuli for cattle (Bos taurus). Appl Anim Behav Sci (1998) 58:13–21.
    doi: 10.1016/S0168-1591(97)00061-0google scholar: lookup
  103. Munksgaard L, de Passillé A, Rushen J, Ladewig J. Dairy cows' use of colour cues to discriminate between people. Appl Anim Behav Sci (1999) 65:1–11.
  104. Ralphs MH, Olsen JD. Adverse influence of social facilitation and learning context in training cattle to avoid eating larkspur. J Anim Sci (1990) 68:1944–52.
    doi: 10.2527/1990.6871944xpubmed: 2166731google scholar: lookup
  105. Boissy A, Le Neindre P. Social influences on the reactivity of heifers: implications for learning abilities in operant conditioning. Appl Anim Behav Sci (1990) 25:149–65.
    doi: 10.1016/0168-1591(90)90077-Qgoogle scholar: lookup
  106. Henselek Y, Fischer J, Schloegl C. Does the stimulus type influence horses' performance in a quantity discrimination task?. Front Psychol (2012) 3:504.
    doi: 10.3389/fpsyg.2012.00504pmc: PMC3499915pubmed: 23181043google scholar: lookup
  107. Petrazzini ME. Trained quantity abilities in horses (Equus caballus): a preliminary investigation. Behav Sci (2014) 4:213–25.
    doi: 10.3390/bs4030213pmc: PMC4219264pubmed: 25379278google scholar: lookup
  108. Péron F, Ward R, Burman O. Horses (Equus caballus) discriminate body odour cues from conspecifics. Anim Cogn (2013) 17:1007–11.
    doi: 10.1007/s10071-013-0717-9pubmed: 24305997google scholar: lookup
  109. Lemasson A, Boutin A, Boivin S, Blois-Heulin C, Hausberger M. Horse (Equus caballus) whinnies: a source of social information. Anim Cogn (2009) 12:693–704.
    doi: 10.1007/s10071-009-0229-9pubmed: 19449192google scholar: lookup
  110. Sankey C, Henry S, André N, Richard-Yris MA, Hausberger M. Do Horses have a concept of person?. PLoS ONE (2011) 6:e18331.
    doi: 10.1371/journal.pone.0018331pmc: PMC3068175pubmed: 21479184google scholar: lookup
  111. Lampe J, Andre J. Cross-modal recognition of human individuals in domestic horses (Equus caballus). Anim Cogn (2012) 15:623–30.
    doi: 10.1007/s10071-012-0490-1pubmed: 22526687google scholar: lookup
  112. Pfungst O. Clever Hans (the Horse of Mr. von Osten): a contribution to experimental, animal, and human psychology. J Philos Psychol Sci Methods (1911) 8:663.
    doi: 10.2307/2012691google scholar: lookup
  113. Birke L, Hockenhull J, Creighton E, Pinno L, Mee J, Mills D. Horses' responses to variation in human approach. Appl Anim Behav Sci (2011) 134:56–63.
    doi: 10.1016/j.applanim.2011.06.002google scholar: lookup
  114. Lovrovich P, Sighieri C, Baragli P. Following human-given cues or not? Horses (Equus caballus) get smarter and change strategy in a delayed three choice task. Appl Anim Behav Sci (2015) 166:80–8.
    doi: 10.1016/j.applanim.2015.02.017google scholar: lookup
  115. Maros K, Gácsi M, Miklósi Á. Comprehension of human pointing gestures in horses (Equus caballus). Anim Cogn (2008) 11:457–66.
    doi: 10.1007/s10071-008-0136-5pubmed: 18247069google scholar: lookup
  116. Proops L, Rayner J, Taylor AM, McComb K. The responses of young domestic horses to human-given cues. PLoS ONE (2013) 8:e67000.
    doi: 10.1371/journal.pone.0067000pmc: PMC3686775pubmed: 23840572google scholar: lookup
  117. Wathan J, McComb K. The eyes and ears are visual indicators of attention in domestic horses. Curr Biol (2014) 24:R677–9.
    doi: 10.1016/j.cub.2014.06.023pmc: PMC4123162pubmed: 25093554google scholar: lookup
  118. Lesimple C, Sankey C, Richard Ma, Hausberger M. Do horses expect humans to solve their problems?. Front Psychol (2012) 3:306.
    doi: 10.3389/fpsyg.2012.00306pmc: PMC3426792pubmed: 22936923google scholar: lookup
  119. McGreevy PD, Cripps PJ, French NP, Green LE, Nicol CJ. Management factors associated with stereotypic and redirected behaviour in the Thoroughbred horse. Equine Vet J (1995) 27:86–91.
    doi: 10.1111/j.2042-3306.1995.tb03041.xpubmed: 7607155google scholar: lookup
  120. Lindberg A, Kelland A, Nicol C. Effects of observational learning on acquisition of an operant response in horses. Appl Anim Behav Sci (1999) 61:187–99.
    doi: 10.1016/S0168-1591(98)00184-1google scholar: lookup
  121. Baer KL, Potter GD, Friend TH, Beaver BV. Observation effects on learning in horses. Appl Anim Ethol (1983) 11:123–9.
    doi: 10.1016/0304-3762(83)90121-9google scholar: lookup
  122. Baker AEM, Crawford BH. Observational learning in horses. Appl Anim Behav Sci (1986) 15:7–13.
    doi: 10.1016/0168-1591(86)90017-1google scholar: lookup
  123. Clarke JV, Nicol CJ, Jones R, McGreevy PD. Effects of observational learning on food selection in horses. Appl Anim Behav Sci (1996) 50:177–84.
    doi: 10.1016/0168-1591(96)01071-4google scholar: lookup
  124. Krueger K, Heinze J. Horse sense: social status of horses (Equus caballus) affects their likelihood of copying other horses' behavior. Anim Cogn (2008) 11:431–9.
    doi: 10.1007/s10071-007-0133-0pubmed: 18183432google scholar: lookup
  125. Hanggi EB. The thinking horse: cognition and perception reviewed. In: AAEP Proceedings, Lexington, KY: (2005). p. 246–55.
  126. Ahrendt LP, Christensen JW, Ladewig J. The ability of horses to learn an instrumental task through social observation. Appl Anim Behav Sci (2012) 139:105–13.
    doi: 10.1016/j.applanim.2012.03.004google scholar: lookup
  127. Henry S, Hemery D, Richard MA, Hausberger M. Human–mare relationships and behaviour of foals toward humans. Appl Anim Behav Sci (2005) 93:341–62.
    doi: 10.1016/j.applanim.2005.01.008google scholar: lookup
  128. Stolba A, Wood-Gush DGM. The behaviour of pigs in a semi-natural environment. Anim Sci (1989) 48:419–25.
  129. Held S, Baumgartner J, Kilbride A, Byrne RW, Mendl M. Foraging behaviour in domestic pigs (Sus scrofa): remembering and prioritizing food sites of different value. Anim Cogn (2005) 8:114–21.
    doi: 10.1007/s10071-004-0242-ypubmed: 15871038google scholar: lookup
  130. Melotti L, Thomsen LR, Toscano MJ, Mendl M, Held S. Delay discounting task in pigs reveals response strategies related to dopamine metabolite. Physiol Behav (2013) 120:182–92.
    doi: 10.1016/j.physbeh.2013.08.009pubmed: 23954408google scholar: lookup
  131. Mendl M, Randle K, Pope S. Young female pigs can discriminate individual differences in odours from conspecific urine. Anim Behav (2002) 64:97–101.
    doi: 10.1006/anbe.2002.3040google scholar: lookup
  132. Gieling ET, Musschenga MA, Nordquist RE, van der Staay FJ. Juvenile pigs use simple geometric 2D shapes but not portrait photographs of conspecifics as visual discriminative stimuli. Appl Anim Behav Sci (2012) 142:142–53.
    doi: 10.1016/j.applanim.2012.10.018google scholar: lookup
  133. Tanida H, Nagano Y. The ability of miniature pigs to discriminate between a stranger and their familiar handler. Appl Anim Behav Sci (1998) 56:149–59.
    doi: 10.1016/S0168-1591(97)00095-6google scholar: lookup
  134. Miklósi Á, Soproni K. A comparative analysis of animals' understanding of the human pointing gesture. Anim Cogn (2006) 9:81–93.
    doi: 10.1007/s10071-005-0008-1pubmed: 16235075google scholar: lookup
  135. Held S, Mendl M, Devereux C, Byrne RW. Social tactics of pigs in a competitive foraging task: the “informed forager” paradigm. Anim Behav (2000) 59:569–76.
    doi: 10.1006/anbe.1999.1322pubmed: 10715179google scholar: lookup
  136. Coussi-Korbel S. Learning to outwit a competitor in mangabeys (Cercocebus torquatus torquatus). J Comp Psychol (1994) 108:164–71.
    doi: 10.1037/0735-7036.108.2.164pubmed: 8026168google scholar: lookup
  137. Held SDE, Byrne RW, Jones S, Murphy E, Friel M, Mendl MT. Domestic pigs, Sus scrofa, adjust their foraging behaviour to whom they are foraging with. Anim Behav (2010) 79:857–62.
    doi: 10.1016/j.anbehav.2009.12.035google scholar: lookup
  138. Ginane C, Dumont B. Do sheep (Ovis aries) categorize plant species according to botanical family?. Anim Cogn (2011) 14:369–76.
    doi: 10.1007/s10071-010-0371-4pubmed: 21203791google scholar: lookup
  139. Langbein J, Siebert K, Nürnberg G, Manteuffel G. Learning to learn during visual discrimination in group housed dwarf goats (Capra hircus). J Comp Psychol (2007) 121:447–56.
    doi: 10.1037/0735-7036.121.4.447pubmed: 18085929google scholar: lookup
  140. Roitberg E, Franz H. Oddity learning by African dwarf goats (Capra hircus). Anim Cogn (2004) 7:61–7.
    doi: 10.1007/s10071-003-0190-ypubmed: 13680403google scholar: lookup
  141. Hosoi E, Swift DM, Rittenhouse LR, Richards RW. Comparative foraging strategies of sheep and goats in a T-maze apparatus. Appl Anim Behav Sci (1995) 44:37–45.
    doi: 10.1016/0168-1591(95)00572-Agoogle scholar: lookup
  142. de la Lama GCM, Mattiello S. The importance of social behaviour for goat welfare in livestock farming. Small Rumin Res (2010) 90:1–10.
    doi: 10.1016/j.smallrumres.2010.01.006google scholar: lookup
  143. Stanley CR, Dunbar RIM. Consistent social structure and optimal clique size revealed by social network analysis of feral goats, Capra hircus. Anim Behav (2013) 85:771–9.
    doi: 10.1016/j.anbehav.2013.01.020google scholar: lookup
  144. Kendrick KM, Atkins K, Hinton MR, Broad KD, Fabre-Nys C, Keverne B. Facial and vocal discrimination in sheep. Anim Behav (1995) 49:1665–76.
    doi: 10.1016/0003-3472(95)90088-8google scholar: lookup
  145. Tate AJ, Fischer H, Leigh AE, Kendrick KM. Behavioural and neurophysiological evidence for face identity and face emotion processing in animals. Phil Trans R Soc B (2006) 361:2155–72.
    doi: 10.1098/rstb.2006.1937pmc: PMC1764842pubmed: 17118930google scholar: lookup
  146. Briefer EF, Padilla de la Torre M, McElligott AG. Mother goats do not forget their kids' calls. Proc R Soc B (2012) 279:3749–55.
    doi: 10.1098/rspb.2012.0986pmc: PMC3415910pubmed: 22719031google scholar: lookup
  147. Pitcher BJ, Briefer EF, Baciadonna L, McElligott AG. Cross-modal recognition of familiar conspecifics in goats. R Soc Open Sci (2017) 4:160346.
    doi: 10.1098/rsos.160346pmc: PMC5367292pubmed: 28386412google scholar: lookup
  148. Beausoleil NJ, Stafford KJ, Mellor DJ. Does direct human eye contact function as a warning cue for domestic sheep (Ovis aries)?. J Comp Psychol (2006) 120:269–79.
    doi: 10.1037/0735-7036.120.3.269pubmed: 16893264google scholar: lookup
  149. Nawroth C, von Borell E, Langbein J. “Goats that stare at men”—revisited: do dwarf goats alter their behaviour in response to eye visibility and head direction of a human?. Anim Cogn (2016c) 19:667–72.
    doi: 10.1007/s10071-016-0957-6pubmed: 26820558google scholar: lookup
  150. Gómez JC. Species comparative studies and cognitive development. Trends Cogn Sci (2005) 9:118–25.
    doi: 10.1016/j.tics.2005.01.004pubmed: 15737820google scholar: lookup
  151. Thorhallsdottir AG, Provenza FD, Balph DF. Ability of lambs to learn about novel foods while observing or participating with social models. Appl Anim Behav Sci (1990) 25:25–33.
    doi: 10.1016/0168-1591(90)90066-Mgoogle scholar: lookup
  152. Veissier I, Stefanova I. Learning to suckle from an artificial teat within groups of lambs: influence of a knowledgeable partner. Behav Process (1993) 30:75–82.
    doi: 10.1016/0376-6357(93)90013-Hpubmed: 24896473google scholar: lookup
  153. Broom DM. Cognitive ability and awareness in domestic animals and decisions about obligations to animals. Appl Anim Behav Sci (2010) 126:1–11.
    doi: 10.1016/j.applanim.2010.05.001google scholar: lookup
  154. Mendl M, Held S, Byrne RW. Pig cognition. Curr Biol (2010b) 20:R796–8.
    doi: 10.1016/j.cub.2010.07.018pubmed: 20869602google scholar: lookup
  155. Nawroth C. Invited review: socio-cognitive capacities of goats and their impact on human-animal interactions. Small Rumin Res (2017) 150:70–5.
    doi: 10.1016/j.smallrumres.2017.03.005google scholar: lookup
  156. Averós X, Brossard L, Dourmad J.-Y., de Greef KH, Edge HL, Edwards SA. A meta-analysis of the combined effect of housing and environmental enrichment characteristics on the behaviour and performance of pigs. Appl Anim Behav Sci (2010) 127:73–85.
    doi: 10.1016/j.applanim.2010.09.010google scholar: lookup
  157. Newberry RC. Environmental enrichment: increasing the biological relevance of captive environments. Appl Anim Behav Sci (1995) 44:229–43.
    doi: 10.1016/0168-1591(95)00616-Zgoogle scholar: lookup
  158. Mason G, Clubb R, Latham N, Vickery S. Why and how should we use environmental enrichment to tackle stereotypic behaviour?. Appl Anim Behav Sci (2007) 102:163–88.
    doi: 10.1016/j.applanim.2006.05.041google scholar: lookup
  159. Wemelsfelder F. The concept of animal boredom and its relationship to stereotyped behaviour. In: Lawrence AB, Rushen J, editors. Stereotypie Behavior: Fundamentals and Applications to Animal Welfare. Wallingford: CAB International; (1993). p. 65–95.
  160. Carlstead K, Shepherdson DJ. Alleviating stress in zoo animals with environmental enrichment. In: Moberg GP, Mench JA, editors. The Biology of Animal Stress. The Biology of Animal Stress:Basic Principles and Implications for Animal Welfare. Oxon, NY: CAB1 Publishing; (2000). p. 337–54.
    doi: 10.1079/9780851993591.0337google scholar: lookup
  161. van de Weerd HA, Day JEL. A review of environmental enrichment for pigs housed in intensive housing systems. Appl Anim Behav Sci (2009) 116:1–20.
    doi: 10.1016/j.applanim.2008.08.001google scholar: lookup
  162. Dantzer R. Can farm animal welfare be understood without taking into account the issues of emotion and cognition?. J Anim Sci (2002) 80:E1–9.
    doi: 10.2527/animalsci2002.0021881200800ES10002xgoogle scholar: lookup
  163. Greiveldinger L, Veissier I, Boissy A. Emotional experience in sheep: predictability of a sudden event lowers subsequent emotional responses. Physiol Behav (2007) 92:675–83.
    doi: 10.1016/j.physbeh.2007.05.012pubmed: 17588624google scholar: lookup
  164. Manteuffel G, Langbein J, Puppe B. From operant learning to cognitive enrichment in farm animal housing: bases and applicability. Anim Welf (2009) 18:87–95.
  165. Kalbe C, Puppe B. Long-term cognitive enrichment affects opioid receptor expression in the amygdala of domestic pigs. Genes Brain Behav (2010) 9:75–83.
    doi: 10.1111/j.1601-183X.2009.00536.xpubmed: 19804558google scholar: lookup
  166. Meyer S, Langbein J, Puppe B. Kognitive Umweltanreicherung bei Zoo- und Nutztieren - Implikationen für Verhalten und Wohlbefinden der Tiere. Berl Munch Tierarztl Wochenschr (2010) 12:446–56.
    doi: 10.2376/0005-9366-123-446pubmed: 21141273google scholar: lookup
  167. Puppe B, Ernst K, Schön PC, Manteuffel G. Cognitive enrichment affects behavioural reactivity in domestic pigs. Appl Anim Behav Sci (2007) 105:75–86.
    doi: 10.1016/j.applanim.2006.05.016google scholar: lookup
  168. Zebunke M, Langbein J, Manteuffel G, Puppe B. Autonomic reactions indicating positive affect during acoustic reward learning in domestic pigs. Anim Behav (2011) 81:481–9.
    doi: 10.1016/j.anbehav.2010.11.023google scholar: lookup
  169. Ernst K, Puppe B, Schön PC, Manteuffel G. A complex automatic feeding system for pigs aimed to induce successful behavioural coping by cognitive adaptation. Appl Anim Behav Sci (2005) 91:205–18.
    doi: 10.1016/j.applanim.2004.10.010google scholar: lookup
  170. Zebunke M, Puppe B, Langbein J. Effects of cognitive enrichment on behavioural and physiological reactions of pigs. Physiol Behav (2013) 118:70–9.
    doi: 10.1016/j.physbeh.2013.05.005pubmed: 23680428google scholar: lookup
  171. Langbein J, Nürnberg G, Manteuffel G. Visual discrimination learning in dwarf goats and associated changes in heart rate and heart rate variability. Physiol Behav (2004) 82:601–9.
    doi: 10.1016/j.physbeh.2004.05.007pubmed: 15327907google scholar: lookup
  172. Selye H. Stress in Health and Disease. Boston, MA: Butterworth-Heinemann; (1976).
  173. Langbein J, Siebert K, Nürnberg G. On the use of an automated learning device by group-housed dwarf goats: do goats seek cognitive challenges?. Appl Anim Behav Sci (2009) 120:150–8.
    doi: 10.1016/j.applanim.2009.07.006google scholar: lookup
  174. Osborne SR. The free food (contrafreeloading) phenomenon: a review and analysis. Anim Learn Behav (1977) 5:221–35.
    doi: 10.3758/BF03209232google scholar: lookup
  175. Hagen K, Broom DM. Emotional reactions to learning in cattle. Appl Anim Behav Sci (2004) 85:203–13.
    doi: 10.1016/j.applanim.2003.11.007google scholar: lookup
  176. McGowan RTS, Rehn T, Norling Y, Keeling LJ. Positive affect and learning: exploring the “Eureka Effect” in dogs. Anim Cogn (2014) 17:577–87.
    doi: 10.1007/s10071-013-0688-xpubmed: 24096703google scholar: lookup
  177. Meehan CL, Mench JA. The challenge of challenge: can problem solving opportunities enhance animal welfare?. Appl Anim Behav Sci (2007) 102:246–61.
    doi: 10.1016/j.applanim.2006.05.031google scholar: lookup
  178. Baymann U, Langbein J, Siebert K, Nürnberg G, Manteuffel G, Mohr E. Cognitive enrichment in farm animals-the impact of social rank and social environment on learning behaviour of dwarf goats. Berliner und Muenchener Tierarztl Wochenschrift (2007) 120:89–97.
    doi: 10.2376/0005-9366-120-89pubmed: 17416130google scholar: lookup
  179. Chapple RS, Wodzicka-Tomaszewska M, Lynch JJ. The learning behaviour of sheep when introduced to wheat. II Social transmission of wheat feeding and the role of the senses. Appl Anim Behav Sci (1987) 18:163–72.
    doi: 10.1016/0168-1591(87)90190-0google scholar: lookup
  180. Chua B, Coenen E, van Delen J, Weary DM. Effects of pair versus individual housing on the behavior and performance of dairy calves. J Dairy Sci (2002) 85:360–4.
    doi: 10.3168/jds.S0022-0302(02)74082-4pubmed: 11913695google scholar: lookup
  181. Costa JHC, Costa WG, Weary DM, Filho LCPM, von Keyserlingk MAG. Dairy heifers benefit from the presence of an experienced companion when learning how to graze. J Dairy Sci (2015) 99:562–8.
    doi: 10.3168/jds.2015-9387pubmed: 26547655google scholar: lookup
  182. Laland KN. Social learning strategies. Anim Learn Behav (2004) 32:4–14.
    doi: 10.3758/BF03196002pubmed: 15161136google scholar: lookup
  183. Hemsworth P. Human–animal interactions in livestock production. Appl Anim Behav Sci (2003) 81:185–98.
    doi: 10.1016/S0168-1591(02)00280-0google scholar: lookup
  184. Brajon S, Laforest JP, Bergeron R, Tallet C, Devillers N. The perception of humans by piglets: recognition of familiar handlers and generalisation to unfamiliar humans. Anim Cogn (2015) 18:1299–316.
    doi: 10.1007/s10071-015-0900-2pubmed: 26187347google scholar: lookup
  185. Fureix C, Jego P, Sankey C, Hausberger M. How horses (Equus caballus) see the world: humans as significant “objects.”. Anim Cogn (2009) 12:643–54.
    doi: 10.1007/s10071-009-0223-2pubmed: 19381698google scholar: lookup
  186. Destrez A, Coulon M, Deiss V, Delval E, Boissy A, Boivin X. The valence of the long-lasting emotional experiences with various handlers modulates discrimination and generalization of individual humans in sheep. J Anim Sci (2013) 91:5418–26.
    doi: 10.2527/jas.2012-5654pubmed: 24045468google scholar: lookup
  187. Sankey C, Richard-Yris MA, Leroy H, Henry S, Hausberger M. Positive interactions lead to lasting positive memories in horses, Equus caballus. Anim Behav (2010) 79:869–75.
    doi: 10.1016/j.anbehav.2009.12.037google scholar: lookup
  188. Jago J, Krohn C, Matthews L. The influence of feeding and handling on the development of the human–animal interactions in young cattle. Appl Anim Behav Sci (1999) 62:137–51.
    doi: 10.1016/S0168-1591(98)00219-6google scholar: lookup
  189. Lürzel S, Windschnurer I, Futschik A, Waiblinger S. Gentle interactions decrease the fear of humans in dairy heifers independently of early experience of stroking. Appl Anim Behav Sci (2016) 178:16–22.
    doi: 10.1016/j.applanim.2016.02.012google scholar: lookup
  190. Probst JK, Spengler Neff A, Leiber F, Kreuzer M, Hillmann E. Gentle touching in early life reduces avoidance distance and slaughter stress in beef cattle. Appl Anim Behav Sci (2012) 139:42–9.
    doi: 10.1016/j.applanim.2012.03.002google scholar: lookup
  191. Waiblinger S, Boivin X, Pedersen V, Tosi MV, Janczak AM, Visser EK. Assessing the human-animal relationship in farmed species: a critical review. Appl Anim Behav Sci (2006) 101:185–242.
    doi: 10.1016/j.applanim.2006.02.001google scholar: lookup
  192. Boissy A, Bouissou MF. Effects of early handling on heifers' subsequent reactivity to humans and to unfamiliar situations. Appl Anim Behav Sci (1988) 20:259–73.
  193. Schmied C, Boivin X, Waiblinger S. Stroking different body regions of dairy cows: effects on avoidance and approach behavior toward humans. J Dairy Sci (2008) 91:596–605.
    doi: 10.3168/jds.2007-0360pubmed: 18218746google scholar: lookup
  194. Stewart M, Shepherd HM, Webster JR, Waas JR, McLeay LM, Schütz KE. Effect of previous handling experiences on responses of dairy calves to routine husbandry procedures. Animal (2013) 7:828–33.
    doi: 10.1017/S175173111200225Xpubmed: 23218065google scholar: lookup
  195. Breuer K, Hemsworth PH, Barnett JL, Matthews LR, Coleman GJ. Behavioural response to humans and the productivity of commercial dairy cows. Appl Anim Behav Sci (2000) 66:273–88.
    doi: 10.1016/S0168-1591(99)00097-0pubmed: 10700627google scholar: lookup
  196. Caroprese M, Napolitano F, Albenzio M, Annicchiarico G, Musto M, Sevi A. Influence of gentling on lamb immune response and human-lamb interactions. Appl Anim Behav Sci (2006) 99:118–31.
    doi: 10.1016/j.applanim.2005.08.023google scholar: lookup
  197. Lürzel S, Münsch C, Windschnurer I, Futschik A, Palme R, Waiblinger S. The influence of gentle interactions on avoidance distance towards humans, weight gain and physiological parameters in group-housed dairy calves. Appl Anim Behav Sci (2015) 172:9–16.
    doi: 10.1016/j.applanim.2015.09.004google scholar: lookup
  198. Garner R. Animal Ethics. Cambridge: Polity Press; (2005).
  199. Palmer C. Animal Ethics in Context. New York, NY: Columbia University Press; (2010).
  200. Benz-Schwarzburg J, Knight A. Cognitive relatives yet moral strangers?. J Anim Ethics (2011) 1:9–36.
    doi: 10.5406/janimalethics.1.1.0009google scholar: lookup
  201. Fraser D. Animal ethics and animal welfare science: bridging the two cultures. Appl Anim Behav Sci (1999) 65:171–89.
    doi: 10.1016/S0168-1591(99)00090-8google scholar: lookup
  202. Jones RC. Science, sentience, and animal welfare. Biol Philos (2013) 28:1–30.
    doi: 10.1007/s10539-012-9351-1google scholar: lookup
  203. Rollin BE. The inseparability of science and ethics in animal welfare. J Agric Environ Ethics (2015) 28:759–65.
    doi: 10.1007/s10806-015-9558-7google scholar: lookup
  204. Cavalieri P, Singer P. The Great Ape Project: Equality Beyond Humanity. New York, NY: St. Martin's Griffin; (1993).
  205. Nussbaum MC. Beyond “compassion and humanity”: justice for nonhuman animals. In: Sunstein CR, Nussbaum MC, editors. Animal Rights: Current Debates and New Directions. New York, NY: Oxford University Press; (2004). p. 299–320.
  206. DeGrazia D. Taking Animals Seriously: Mental Life and Moral Status. Cambridge: Cambridge University Press; (1996).
    doi: 10.1017/CBO9781139172967google scholar: lookup
  207. Duncan IJH, Petherick JC. The implications of cognitive processes for animal welfare. J Anim Sci (1991) 69:5017–22.
    doi: 10.2527/1991.69125017xpubmed: 1808195google scholar: lookup
  208. Mendl M, Paul ES. Do animals live in the present? Current evidence and implications for welfare. Appl Anim Behav Sci (2008) 113:357–82.
    doi: 10.1016/j.applanim.2008.01.013google scholar: lookup
  209. Andrews K, Monsó S. Animal moral psychologies. In: Doris J, Vargas M, editors. The Oxford Handbook of Moral Psychology. New York, NY: Oxford University Press; (in press).
  210. Bekoff M, Pierce J. Wild Justice: The Moral Lives of Animals. Chicago, IL: University of Chicago Press; (2009).
    doi: 10.7208/chicago/9780226041667.001.0001google scholar: lookup
  211. Monsó S. Morality without mindreading. Mind Lang (2017) 32:338–57.
    doi: 10.1111/mila.12146google scholar: lookup
  212. Rowlands M. Can Animals Be Moral?. New York, NY: Oxford University Press; (2012).
    doi: 10.1093/acprof:oso/9780199842001.001.0001google scholar: lookup
  213. Cataldi SL. Animals and the concept of dignity: critical reflections on a circus performance. Ethics Environ (2002) 7:104–26.
    doi: 10.2979/ETE.2002.7.2.104google scholar: lookup
  214. Rutgers B, Heeger R. Inherent worth and respect for animal integrity. In: Dol M, Fentener Van Vlissingen M, Kasanmoentalib S, Visser T, Zwart H, editors. Recognizing the Intrinsic Value of Animals: Beyond Animal Welfare. Assen: Van Gorcum Ltd. (1999). p. 41–51.
  215. Regan T. The Case for Animal Rights. Berkeley, CA: University of California Press; (2004).
  216. Gutmann AK, Špinka M, Winckler C. Long-term familiarity creates preferred social partners in dairy cows. Appl Anim Behav Sci (2015) 169:1–8.
    doi: 10.1016/j.applanim.2015.05.007google scholar: lookup
  217. Cronin KA. Prosocial behaviour in animals: the influence of social relationships, communication and rewards. Anim Behav (2012) 84:1085–93.
    doi: 10.1016/j.anbehav.2012.08.009google scholar: lookup
  218. Monsó S, Benz-Schwarzburg J, Huber L. Morality in animals: what it means and why it matters. J Ethics (2018) 22:283–10.
    doi: 10.1007/s10892-018-9275-3pmc: PMC6404642pubmed: 30930677google scholar: lookup
  219. Baciadonna L, McElligott A. The use of judgement bias to assess welfare in farm livestock. Anim Welf (2015) 24:81–91.
    doi: 10.7120/09627286.24.1.081google scholar: lookup
  220. Spence C, Osman M, McElligott AG. Theory of animal mind: human nature or experimental artefact?. Trends Cogn Sci (2017) 21:333–43.
    doi: 10.1016/j.tics.2017.02.003pubmed: 28347613google scholar: lookup
  221. Ruby MB, Heine SJ. Too close to home. Factors predicting meat avoidance. Appetite (2012) 59:47–52.
    doi: 10.1016/j.appet.2012.03.020pubmed: 22465239google scholar: lookup
  222. Bastian B, Loughnan S, Haslam N, Radke HRM. Don't mind meat? The denial of mind to animals used for human consumption. Pers Soc Psychol Bull (2012) 38:247–56.
    doi: 10.1177/0146167211424291pubmed: 21980158google scholar: lookup
  223. Rosenthal R. The file drawer problem and tolerance for null results. Psychol Bull (1979) 86:638–41.
    doi: 10.1037/0033-2909.86.3.638google scholar: lookup

Citations

This article has been cited 45 times.
  1. Proudfoot KL, Ede T, Ryan CL, Neave HW. Cognition of dairy cattle: Implications for animal welfare and dairy science. JDS Commun 2025 Dec;6(Suppl 1):S37-S41.
    doi: 10.3168/jdsc.2025-0860pubmed: 41497384google scholar: lookup
  2. Liu S, Fujita N, Sasaki T, Chiba T, Konno S, Roh S, Fukasawa M. Associations Between REM Sleep-like Posture Expression and Cognitive Flexibility in 2-Month-Old Japanese Black Calves. Animals (Basel) 2025 Nov 28;15(23).
    doi: 10.3390/ani15233438pubmed: 41375496google scholar: lookup
  3. Perrone M, Mazzocchi C, Palladini N, Bava L, Ruggeri G. Farm Animal Welfare: Consumers' Perception Toward Different Breeds of Animals in Italy. Animals (Basel) 2025 Nov 25;15(23).
    doi: 10.3390/ani15233406pubmed: 41375463google scholar: lookup
  4. Iglesias Pastrana C, Navas González FJ, Ciani E, McLean AK, Delgado Bermejo JV. Cognitive performance and variability in dromedary camels: insights from a comparative psychometric approach. BMC Vet Res 2025 Dec 4;22(1):11.
    doi: 10.1186/s12917-025-05169-xpubmed: 41339887google scholar: lookup
  5. Gunkel J, Miller FG. Xenotransplantation: Injustice, Harm, and Alternatives for Addressing the Organ Crisis. Hastings Cent Rep 2025 Sep-Oct;55(5):7-17.
    doi: 10.1002/hast.5021pubmed: 41091031google scholar: lookup
  6. Dwulit AK, Morey RA, van der Staay FJ. Incorporating farm animal models for the study of neuropsychiatric diseases: Expansion of the possibilities. Cogn Affect Behav Neurosci 2026 Feb;26(1):16-32.
    doi: 10.3758/s13415-025-01332-xpubmed: 40775184google scholar: lookup
  7. Gao J, Yamanashi Y, Tanaka M. Touchscreen Tasks for Cognitive Testing in Domestic Goats (Capra hircus): A Pilot Study Using Odd-Item Search Training. Animals (Basel) 2025 Jul 17;15(14).
    doi: 10.3390/ani15142115pubmed: 40723578google scholar: lookup
  8. Oldham L, Agha S, Arnott G, Brims M, Camerlink I, Futro A, Lee VE, Doeschl-Wilson A, Turner SP. Cognitive and non-cognitive factors predict pigs' positions in an aggression social network. Sci Rep 2025 May 20;15(1):17439.
    doi: 10.1038/s41598-025-02023-wpubmed: 40394067google scholar: lookup
  9. Brosche K, Veit A, Fraga PP, Wondrak M, Andics A, Virányi Z. Human ostension enhances attentiveness but not performance in domestic pigs. Sci Rep 2025 May 9;15(1):16161.
    doi: 10.1038/s41598-025-00511-7pubmed: 40346262google scholar: lookup
  10. Hillerer KM, Gimsa U. Adult neurogenesis and the microbiota-gut-brain axis in farm animals: underestimated and understudied parameters for improving welfare in livestock farming. Front Neurosci 2024;18:1493605.
    doi: 10.3389/fnins.2024.1493605pubmed: 39664450google scholar: lookup
  11. Brelaz ÉCDO, Moura GAB, Fonsêca VFC, Almeida JAT, Simão BR, Maia ASC. Training Protocol for Nellore Cattle in Respirometry Flow Trials Using Non-Ventilated Facial Mask. Animals (Basel) 2024 Oct 8;14(19).
    doi: 10.3390/ani14192888pubmed: 39409836google scholar: lookup
  12. Schaffer A, Widdig A, Holland R, Amici F. Evidence of object permanence, short-term spatial memory, causality, understanding of object properties and gravity across five different ungulate species. Sci Rep 2024 Jun 14;14(1):13718.
    doi: 10.1038/s41598-024-64396-8pubmed: 38877059google scholar: lookup
  13. Williams E, Sadler J, Rutter SM, Mancini C, Nawroth C, Neary JM, Ward SJ, Charlton G, Beaver A. Human-animal interactions and machine-animal interactions in animals under human care: A summary of stakeholder and researcher perceptions and future directions. Anim Welf 2024;33:e27.
    doi: 10.1017/awf.2024.23pubmed: 38751800google scholar: lookup
  14. Torres Borda L, Roth P, Lumetzberger J, Auer U, Jenner F. Proximity tracking using ultra-wideband technology for equine social behaviour research. Sci Rep 2024 Apr 30;14(1):9971.
    doi: 10.1038/s41598-024-60805-0pubmed: 38693325google scholar: lookup
  15. Torres Borda L, Kelemen Z, Auer U, Jenner F. Video Ethogram of Equine Social Behaviour. Animals (Basel) 2024 Apr 14;14(8).
    doi: 10.3390/ani14081179pubmed: 38672327google scholar: lookup
  16. Moscovice LR, Sobczak B, Niittynen T, Koski SE, Gimsa U. Changes in salivary oxytocin in response to biologically-relevant events in farm animals: method optimization and usefulness as a biomarker. Front Physiol 2024;15:1370557.
    doi: 10.3389/fphys.2024.1370557pubmed: 38567114google scholar: lookup
  17. Ede T, Parsons TD. Cognitive tasks as measures of pig welfare: a systematic review. Front Vet Sci 2023;10:1251070.
    doi: 10.3389/fvets.2023.1251070pubmed: 38033647google scholar: lookup
  18. Masebo NT, Zappaterra M, Felici M, Benedetti B, Padalino B. Dromedary camel's welfare: literature from 1980 to 2023 with a text mining and topic analysis approach. Front Vet Sci 2023;10:1277512.
    doi: 10.3389/fvets.2023.1277512pubmed: 38026661google scholar: lookup
  19. Cozzolino F, Canè L, Sacchettino L, Gatto MC, Iacobucci I, Gatta C, De Biase D, Di Napoli E, Paciello O, Avallone L, Monti M, d'Angelo D, Napolitano F. Preliminary evaluation of the proteomic profiling in the hippocampus of aged grazing cattle. Front Aging Neurosci 2023;15:1274073.
    doi: 10.3389/fnagi.2023.1274073pubmed: 37965495google scholar: lookup
  20. Pollard-Wright H. The feelings of knowing - fundamental interoceptive patterns (FoK-FIP) system: connecting consciousness to physics. Commun Integr Biol 2023;16(1):2260682.
    doi: 10.1080/19420889.2023.2260682pubmed: 37779822google scholar: lookup
  21. Calapai A, Pfefferle D, Cassidy LC, Nazari A, Yurt P, Brockhausen RR, Treue S. A Touchscreen-Based, Multiple-Choice Approach to Cognitive Enrichment of Captive Rhesus Macaques (Macaca mulatta). Animals (Basel) 2023 Aug 24;13(17).
    doi: 10.3390/ani13172702pubmed: 37684966google scholar: lookup
  22. Paganelli A, Felici M, Turini L, Baragli P, Carlucci L, Recchia FA, Sgorbini M. Detour test performance of cloned minipigs from three different clone populations. Vet Res Commun 2023 Dec;47(4):2137-2144.
    doi: 10.1007/s11259-023-10168-0pubmed: 37452229google scholar: lookup
  23. Fuentes A, Han S, Nasir MF, Park J, Yoon S, Park DS. Multiview Monitoring of Individual Cattle Behavior Based on Action Recognition in Closed Barns Using Deep Learning. Animals (Basel) 2023 Jun 17;13(12).
    doi: 10.3390/ani13122020pubmed: 37370530google scholar: lookup
  24. Levey DJ, Poulsen JR, Schaeffer AP, Deochand ME, Oswald JA, Robinson SK, Londoño GA. Wild mockingbirds distinguish among familiar humans. Sci Rep 2023 Jun 24;13(1):10259.
    doi: 10.1038/s41598-023-36225-xpubmed: 37355713google scholar: lookup
  25. Torres Borda L, Auer U, Jenner F. Equine Social Behaviour: Love, War and Tolerance. Animals (Basel) 2023 Apr 26;13(9).
    doi: 10.3390/ani13091473pubmed: 37174510google scholar: lookup
  26. Tajonar K, Gonzalez-Ronquillo M, Relling A, Nordquist RE, Nawroth C, Vargas-Bello-Pérez E. Toward assessing the role of dietary fatty acids in lamb's neurological and cognitive development. Front Vet Sci 2023;10:1081141.
    doi: 10.3389/fvets.2023.1081141pubmed: 36865439google scholar: lookup
  27. Lee VE, Arnott G, Turner SP. Social behavior in farm animals: Applying fundamental theory to improve animal welfare. Front Vet Sci 2022;9:932217.
    doi: 10.3389/fvets.2022.932217pubmed: 36032304google scholar: lookup
  28. Nagy PP, Skidmore JA, Juhasz J. Intensification of camel farming and milk production with special emphasis on animal health, welfare, and the biotechnology of reproduction. Anim Front 2022 Aug;12(4):35-45.
    doi: 10.1093/af/vfac043pubmed: 35974793google scholar: lookup
  29. Garnham LC, Clarke C, Løvlie H. How Inhibitory Control Relates to Positive and Negative Affective States in Red Junglefowl. Front Vet Sci 2022;9:872487.
    doi: 10.3389/fvets.2022.872487pubmed: 35464350google scholar: lookup
  30. Celozzi S, Battini M, Prato-Previde E, Mattiello S. Humans and Goats: Improving Knowledge for a Better Relationship. Animals (Basel) 2022 Mar 18;12(6).
    doi: 10.3390/ani12060774pubmed: 35327171google scholar: lookup
  31. Parois SP, Eicher SD, Lindemann SR, Marchant JN. Potential improvements of the cognition of piglets through a synbiotic supplementation from 1 to 28 days via the gut microbiota. Sci Rep 2021 Dec 16;11(1):24113.
    doi: 10.1038/s41598-021-03565-5pubmed: 34916559google scholar: lookup
  32. Campbell DLM, Lee C. A Perspective on Strategic Enrichment for Brain Development: Is This the Key to Animal Happiness?. Front Vet Sci 2021;8:720422.
    doi: 10.3389/fvets.2021.720422pubmed: 34621808google scholar: lookup
  33. Nawroth C, Baciadonna L, Emery NJ. Editorial: Humans in an Animal's World-How Non-human Animals Perceive and Interact With Humans. Front Psychol 2021;12:733430.
    doi: 10.3389/fpsyg.2021.733430pubmed: 34539531google scholar: lookup
  34. Krause A, Kreiser M, Puppe B, Tuchscherer A, Düpjan S. The effect of age on discrimination learning and self-control in a marshmallow test for pigs. Sci Rep 2021 Sep 14;11(1):18287.
    doi: 10.1038/s41598-021-97770-xpubmed: 34521956google scholar: lookup
  35. Jardat P, Lansade L. Cognition and the human-animal relationship: a review of the sociocognitive skills of domestic mammals toward humans. Anim Cogn 2022 Apr;25(2):369-384.
    doi: 10.1007/s10071-021-01557-6pubmed: 34476652google scholar: lookup
  36. . Environmental enrichment and stress relief in pigs. J Anim Sci 2021 May 1;99(5).
    doi: 10.1093/jas/skab113pubmed: 33939814google scholar: lookup
  37. Fife-Cook I, Franks B. Koi (Cyprinus rubrofuscus) Seek Out Tactile Interaction with Humans: General Patterns and Individual Differences. Animals (Basel) 2021 Mar 5;11(3).
    doi: 10.3390/ani11030706pubmed: 33807873google scholar: lookup
  38. Croney CC, Boysen ST. Acquisition of a Joystick-Operated Video Task by Pigs (Sus scrofa). Front Psychol 2021;12:631755.
    doi: 10.3389/fpsyg.2021.631755pubmed: 33679560google scholar: lookup
  39. Neethirajan S, Reimert I, Kemp B. Measuring Farm Animal Emotions-Sensor-Based Approaches. Sensors (Basel) 2021 Jan 14;21(2).
    doi: 10.3390/s21020553pubmed: 33466737google scholar: lookup
  40. Christensen JW, Ahrendt LP, Malmkvist J, Nicol C. Exploratory behaviour towards novel objects is associated with enhanced learning in young horses. Sci Rep 2021 Jan 14;11(1):1428.
    doi: 10.1038/s41598-020-80833-wpubmed: 33446827google scholar: lookup
  41. Benz-Schwarzburg J, Monsó S, Huber L. How Dogs Perceive Humans and How Humans Should Treat Their Pet Dogs: Linking Cognition With Ethics. Front Psychol 2020;11:584037.
    doi: 10.3389/fpsyg.2020.584037pubmed: 33391102google scholar: lookup
  42. Trösch M, Flamand A, Chasles M, Nowak R, Calandreau L, Lansade L. Horses Solve Visible but Not Invisible Displacement Tasks in an Object Permanence Paradigm. Front Psychol 2020;11:562989.
    doi: 10.3389/fpsyg.2020.562989pubmed: 33117229google scholar: lookup
  43. 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
  44. Farrar BG, Boeckle M, Clayton NS. Replications in Comparative Cognition: What Should We Expect and How Can We Improve?. Anim Behav Cogn 2020 Feb;7(1):1-22.
    doi: 10.26451/abc.07.01.02.2020pubmed: 32626823google scholar: lookup
  45. Wilson V, Guenther A, Øverli Ø, Seltmann MW, Altschul D. Future Directions for Personality Research: Contributing New Insights to the Understanding of Animal Behavior. Animals (Basel) 2019 May 15;9(5).
    doi: 10.3390/ani9050240pubmed: 31096599google scholar: lookup
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