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Anticipatory behaviour in animals: A critical review

Published online by Cambridge University Press:  01 January 2023

C Anderson
Affiliation:
Animal Welfare Program, Faculty of Land and Food Systems, 2357 Main Mall, University of British Columbia, Vancouver, BC, V6T 1Z6, Canada Department of Animal Environment and Health, Swedish University of Agricultural Sciences, Box 7068, SE-750 07 Uppsala, Sweden
MAG von Keyserlingk
Affiliation:
Animal Welfare Program, Faculty of Land and Food Systems, 2357 Main Mall, University of British Columbia, Vancouver, BC, V6T 1Z6, Canada
LM Lidfors
Affiliation:
Department of Animal Environment and Health, Swedish University of Agricultural Sciences, Box 7068, SE-750 07 Uppsala, Sweden
DM Weary*
Affiliation:
Animal Welfare Program, Faculty of Land and Food Systems, 2357 Main Mall, University of British Columbia, Vancouver, BC, V6T 1Z6, Canada
*
* Contact for correspondence: dan.weary@ubc.ca
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Abstract

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A number of studies have investigated anticipatory behaviour in animals as a measure of sensitivity to reward or as an expression of emotional state. A common feature of many studies is that they base inferences on seemingly arbitrary measures, for example, the frequency of behavioural transitions (ie number of times an animal switches between different behaviours). This paper critically reviews the literature and discusses various hypotheses for why specific behavioural responses occur in the anticipatory period between the signal and reward in conditioned animals. We argue that the specific behaviours shown may be the result of superstitious learning and thus highly variable, leaving behavioural transitions as the only response that can be scored consistently, and that sometimes these responses may relate more to frustration than to a positive emotional state. Finally, we propose new research approaches to avoid potential confounds and improve future studies on this topic.

Type
Research Article
Creative Commons
Creative Common License - CCCreative Common License - BY
This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (https://creativecommons.org/licenses/by/4.0/), which permits unrestricted re-use, distribution, and reproduction in any medium, provided the original work is properly cited.
Copyright
© The Author(s), 2023. Published by Cambridge University Press on behalf of The Universities Federation for Animal Welfare

References

Amsel, A 1992 Frustration Theory: An Analysis of Dispositional Learning and Memory. Cambridge University Press: Cambridge, UK. https://doi.org/10.1017/CBO9780511665561CrossRefGoogle Scholar
Anderson, C 2016 Investigating anticipatory behaviours in lambs. Doctoral dissertation, Swedish University of Agricultural Sciences, SwedenGoogle Scholar
Anderson, C, Yngvesson, J, Boissy, A, Uvnäs-Moberg, K and Lidfors, L 2015 Behavioural expression of positive anticipation for food or opportunity to play in lambs. Behavioural Processes 113: 152158. https://doi.org/10.1016/j.beproc.2015.02.003CrossRefGoogle ScholarPubMed
Antle, MC and Silver, R 2009 Neural basis of timing and antici-patory behaviors. European Journal of Neuroscience 30: 16431649. https://doi.org/10.1111/j.1460-9568.2009.06959.xCrossRefGoogle ScholarPubMed
Balsam, P, Sanchez-Castillo, H, Taylor, K, Van Volkinburg, H and Ward, RD 2009 Timing and anticipation: Conceptual and methodological approaches. European Journal of Neuroscience 30: 17491755. https://doi.org/10.1111/j.1460-9568.2009.06967.xCrossRefGoogle ScholarPubMed
Berridge, KC 1996 Food reward: brain substrates of wanting and liking. Neuroscience and Biobehavioral Reviews 20: 125. https://doi.org/10.1016/0149-7634(95)00033-BCrossRefGoogle ScholarPubMed
Berridge, KC 2007 The debate over dopamine’s role in reward: the case for incentive salience. Psychopharmacology 191: 391431. https://doi.org/10.1007/s00213-006-0578-xCrossRefGoogle ScholarPubMed
Bishop, JD, Malven, PV, Singleton, WL and Weesner, GD 1999 Hormonal and behavioral correlates of emotional states in sexually trained boars. Journal of Animal Science 77: 33393345. https://doi.org/10.2527/1999.77123339xCrossRefGoogle ScholarPubMed
Boissy, A, Aubert, A, Désiré, L, Greiveldinger, L, Delval, E and Veissier, I 2011 Cognitive sciences to relate ear postures to emotions in sheep. Animal Welfare 20: 475610.1017/S0962728600002426CrossRefGoogle Scholar
Burgdorf, J, Knutson, B, Panksepp, J and Ikemoto, S 2001 Nucleus accumbens amphetamine microinjections unconditionally elicit 50 kHz ultrasonic vocalizations in rats. Behavioral Neuroscience 115: 940944. https://doi.org/10.1037/0735-7044.115.4.940CrossRefGoogle ScholarPubMed
Carlstead, K 1986 Predictability of feeding: its effect on agonistic behaviour and growth in grower pigs. Applied Animal Behaviour Science 16: 2538. https://doi.org/10.1016/0168-1591(86)90037-7CrossRefGoogle Scholar
Chapagain, D, Uvnäs-Moberg, K and Lidfors, LM 2014 Investigating the motivation to play in lambs. Applied Animal Behaviour Science 160: 6474. https://doi.org/10.1016/j.applan-im.2014.08.004CrossRefGoogle Scholar
Clark, RE and Squire, LR 1998 Classical conditioning and brain systems: the role of awareness. Science 280: 7781. https://doi.org/10.1126/science.280.5360.77CrossRefGoogle ScholarPubMed
Clegg, ILK and Delfour, F 2018 Cognitive judgement bias is associated with frequency of anticipatory behavior in bottlenose dolphins. Zoo Biology 37: 6773. https://doi.org/10.1002/zoo.21400CrossRefGoogle ScholarPubMed
Clegg, ILK, Rödel, HG, Boivin, X and Delfour, F 2018 Looking forward to interacting with their caretakers: dolphins’ anticipato-ry behaviour indicates motivation to participate in specific events. Applied Animal Behaviour Science 202: 8593. https://doi.org/10.1016/j.applanim.2018.01.015CrossRefGoogle Scholar
Cornwell, JFM, Franks, B and Higgins, ET 2014 Truth, con-trol, and value motivations: the ‘what’, ‘how’, and ‘why’ of approach and avoidance. Frontiers in Systems Neuroscience 8: 194. https://doi.org/10.3389/fnsys.2014.00194CrossRefGoogle Scholar
Di Canio, P, Cardinal, RN, Cowell, RA, Little, SJ and Everitt, BJ 2001 Differential involvement of NMDA, AMPA/kainate, and dopamine receptors in the nucleus accumbens core in the acqui-sition and performance of Pavlovian approach behavior. The Journal of Neuroscience 21: 94719477. https://doi.org/10.1523/JNEUROSCI.21-23-09471.2001CrossRefGoogle Scholar
Dickinson, A and Balleine, B 1994 Motivational control of goal-directed action. Animal Learning and Behavior 22: 118. https://doi.org/10.3758/BF03199951CrossRefGoogle Scholar
Folkedal, O, Stien, LH, Torgersen, T, Oppedal, F, Olsen, RE, Fosseidengen, JE, Braithwaite, VA and Kristiansen, TS 2012 Food anticipatory behaviour as an indicator of stress response and recovery in Atlantic salmon post-smolt after exposure to acute temperature fluctuation. Physiology and Behavior 105: 250256. https://doi.org/10.1016/j.physbeh.2011.08.008CrossRefGoogle ScholarPubMed
Forkman, B, Boissy, A, Meunier-Salaün, M-C, Canali, E and Jones, RB 2007 A critical review of fear tests used on cattle, pigs, sheep, poultry and horses. Physiology and Behavior 92: 340374. https://doi.org/10.1016/j.physbeh.2007.03.016CrossRefGoogle ScholarPubMed
Gray, J 1987 The Psychology of Fear and Stress. Problems in Behavioural Sciences. Cambridge University Press: Cambridge, UKGoogle Scholar
Greiveldinger, L, Veissier, I and Boissy, A 2011 The ability of lambs to form expectations and the emotional consequences of a discrepancy from their expectations. Psychoneuroendocrinology 36: 806815. https://doi.org/10.1016/j.psyneuen.2010.11.002CrossRefGoogle ScholarPubMed
Gygax, L, Reefmann, N, Wolf, M and Langbein, JM 2013 Prefrontal cortex activity, sympatho-vagal reaction and behaviour distinguish between situations of feed reward and frustration in dwarf goats. Behavioural Brain Research 239: 104114. https://doi.org/10.1016/j.bbr.2012.10.052CrossRefGoogle ScholarPubMed
Hall, G 1994 Pavlovian conditioning. Laws of association. In: Mackintosh NJ (ed) Animal Learning and Cognition. Academic Press: San Diego, USA. https://doi.org/10.1016/B978-0-08-057169-0.50008-5Google Scholar
Hansen, SW and Jeppesen, LL 2006 Temperament, stereotypies and anticipatory behaviour as measures of welfare in mink. Applied Animal Behaviour Science 99: 172182. https://doi.org/10.1016/j.applanim.2005.10.005CrossRefGoogle Scholar
Haskell, MJ, Coerse, NCA, Taylor, PAE and McCorquodale, C 2004 The effect of previous experience over control of access to food and light on the level of frustration-induced aggression in the domestic hen. Ethology 110: 501513. https://doi.org/10.1111/j.1439-0310.2004.00995.xCrossRefGoogle Scholar
Heyse, NC, Brenes, JC and Schwarting, RKW 2015 Exercise reward induces appetitive 50-kHz calls in rats. Physiology and Behavior 147: 131140. https://doi.org/10.1016/j.physbeh.2015.04.021CrossRefGoogle ScholarPubMed
Higgins, ET 2006 Value from hedonic experience and engage-ment. Psychological Review 113: 439460. https://doi.org/10.1037/0033-295X.113.3.439CrossRefGoogle Scholar
Imfeld-Mueller, S and Hillman, E 2012 Anticipation of a food ball increases short-term activity levels in growing pigs. Applied Animal Behaviour Science 137: 2329. https://doi.org/10.1016/j.applanim.2012.01.012CrossRefGoogle Scholar
Imfeld-Mueller, S, Van Wezemael, L, Stauffacher, M, Gygax, L and Hillmann, E 2011 Do pigs distinguish between sit-uations of different emotional valences during anticipation? Applied Animal Behaviour Science 131: 8693. https://doi.org/10.1016/j.applanim.2011.02.009CrossRefGoogle Scholar
Jensen, ALM, Delfour, F and Carter, T 2013 Anticipatory behav-ior in captive bottlenose dolphins (Tursiops truncatus): A preliminary study. Zoo Biology 32: 436444. https://doi.org/10.1002/zoo.21077CrossRefGoogle Scholar
Kaye, H and Pearce, JM 1984 The strength of the orienting response during Pavlovian conditioning. Journal of Experimental Psychology: Animal Behavior Processes 10: 90109. https://doi.org/10.1037/0097-7403.10.1.90Google ScholarPubMed
Knutson, B, Burgdorf, J and Panksepp, J 1998 Anticipation of play elicits high-frequency ultrasonic vocalizations in young rats. Journal of Comparative Psychology 112: 6573. https://doi.org/10.1037/0735-7036.112.1.65CrossRefGoogle ScholarPubMed
Knutson, B, Burgdorf, J and Panksepp, J 2002 Ultrasonic vocal-izations as indices of affective states in rats. Psychological Bulletin 128: 961977. https://doi.org/10.1037/0033-2909.128.6.961CrossRefGoogle ScholarPubMed
Krebs, BL, Torres, E, Chesney, C, Kantoniemi Moon, V and Watters, JV 2017 Applying behavioral conditioning to identify anticipatory behaviors. Journal of Applied Animal Welfare Science 20: 155175. https://doi.org/10.1080/10888705.2017.1283225CrossRefGoogle ScholarPubMed
Kuhne, F, Sauerbrey, AFC and Adler, S 2013 The discrimination-learning task determines the kind of frustration-related behav-iours in laying hens Gallus gallus domesticus. Applied Animal Behaviour Science 148: 192200. https://doi.org/10.1016/j.applanim.2013.09.003CrossRefGoogle Scholar
Makowska, IJ and Weary, DM 2016 Differences in anticipatory behaviour between rats (Rattus norvegicus) housed in standard versus semi-naturalistic laboratory environments. PLoS One 11: e0147595. https://doi.org/10.1371/journal.pone.0147595CrossRefGoogle ScholarPubMed
Manning, A and Stamp Dawkins, M 1998 An Introduction to Animal Behaviour, Fifth Edition pp 241. Cambridge University Press: Cambridge, UKGoogle Scholar
McGrath, N, Burman, O, Dwyer, C and Philips, CJC 2016 Does the anticipatory behaviour of chickens communicate reward quality? Applied Animal Behaviour Science 184: 8090. https://doi.org/10.1016/j.applanim.2016.08.010CrossRefGoogle Scholar
Mendl, M, Burman, OHP and Paul, ES 2010 An integrative and functional framework for the study of animal emotion and mood. Proceedings of the Royal Society B 277: 28952904. https://doi.org/10.1098/rspb.2010.0303CrossRefGoogle Scholar
Moe, RO, Bakken, M, Kittilsen, S, Kingsley-Smith, H and Spruijt, BM 2006 A note on reward-related behaviour and emotional expressions in farmed silver foxes (Vulpes vulpes). Basis for a novel tool to study animal welfare. Applied Animal Behaviour Science 101: 362368. https://doi.org/10.1016/j.applanim.2006.02.004CrossRefGoogle Scholar
Moe, RO, Nordgreen, J, Janczak, AM, Bakken, M, Spruijt, BM and Jensen, P 2014 Anticipatory and foraging behaviors in response to palatable food reward in chickens: Effects of dopamine D2 recep-tor blockade and domestication. Physiology and Behavior 133: 170177. https://doi.org/10.1016/j.physbeh.2014.05.023CrossRefGoogle Scholar
Moe, RO, Nordgreen, J, Janczak, AM, Spruijt, BM and Bakken, M 2013 Effects of signalled reward type, food status and a µ-opioid receptor antagonist on cue-induced anticipatory behaviour in laying hens Gallus domesticus. Applied Animal Behaviour Science 148: 4653. https://doi.org/10.1016/j.applanim.2013.08.001CrossRefGoogle Scholar
Moe, RO, Nordgreen, J, Janczak, AM, Spruijt, BM, Kostal, L, Skjerve, E, Zanella, AJ and Bakken, M 2011 Effects of haloperi-dol, a dopamine D2-like receptor antagonist, on reward-related behaviors in laying hens. Physiology and Behavior 102: 400405. https://doi.org/10.1016/j.physbeh.2010.12.008CrossRefGoogle ScholarPubMed
Moe, RO, Nordgreen, J, Janczak, AM, Spruijt, BM, Zanella, AJ and Bakken, M 2009 Trace classical conditioning as an approach to the study of reward-related behaviour in laying hens: A methodological study. Applied Animal Behaviour Science 121: 171178. https://doi.org/10.1016/j.applanim.2009.10.002CrossRefGoogle Scholar
Murphy, E, Nordquist, RE and van der Staay, FJ 2014 A review of behavioural methods to study emotion and mood in pigs, Sus scrofa. Applied Animal Behaviour Science 159: 928. https://doi.org/10.1016/j.applanim.2014.08.002CrossRefGoogle Scholar
Niel, L and Weary, DM 2006 Behavioural responses of rats to gradual-fill carbon dioxide euthanasia and reduced oxygen con-centrations Applied Animal Behaviour Science 100: 295308. https://doi.org/10.1016/j.applanim.2005.12.001CrossRefGoogle Scholar
Panksepp, J and Burgdorf, J 2003 ‘Laughing’ rats and the evolutionary antecedents of human joy? Physiology and Behavior 79: 533547. https://doi.org/10.1016/S0031-9384(03)00159-8CrossRefGoogle ScholarPubMed
Pavlov, IP 1927 Conditioned Reflexes. Oxford University Press: London, UKGoogle Scholar
Pearce, JM 2008 Animal Learning and Cognition. An Introduction, Third Edition. Psychology Press: New York, USAGoogle Scholar
Pearce, JM and Kaye, H 1985 Strength of the orienting response during inhibitory conditioning. Journal of Experimental Psychology: Animal Behavior Processes 11: 405420. https://doi.org/10.1037/0097-7403.11.3.405Google Scholar
Peters, SM, Bleijenberg, EH, van Dierendonck, MC, van der Harst, JE and Spruijt, BM 2012 Characterization of anticipatory behav-iour in domesticated horses Equus caballus. Applied Animal Behaviour Science 138: 6069. https://doi.org/10.1016/j.applanim.2012.01.018CrossRefGoogle Scholar
Reimert, I, Bolhuis, JE, Kemp, B and Rodenburg, TB 2013 Indicators of positive and negative emotions and emotional con-tagion in pigs. Physiology and Behavior 109: 4250. https://doi.org/10.1016/j.physbeh.2012.11.002CrossRefGoogle ScholarPubMed
Rescorla, RA 1988 Pavlovian conditioning: It’s not what you think it is. American Psychologist 43: 151160. https://doi.org/10.1037/0003-066X.43.3.151CrossRefGoogle Scholar
Rolls, ET 2005 Emotions Explained. Oxford University Press: New York, USA. https://doi.org/10.1093/acprof:oso/9780198570035.001.0001CrossRefGoogle Scholar
Schultz, W 2007 Behavioral dopamine signals. TRENDS in Neurosciences 30: 203210. https://doi.org/10.1016/j.tins.2007.03.007CrossRefGoogle ScholarPubMed
Skinner, BF 1948 Superstition in the pigeon. Journal of Experimental Psychology 38: 168172. https://doi.org/10.1037/h0055873CrossRefGoogle ScholarPubMed
Spanagel, R, Herz, A, Bals-Kubik, R and Shippenberg, TS 1991 ß-endorphin-induced locomotor stimulation and reinforce-ment are associated with an increase in dopamine release in the nucleus accumbens. Psychopharmacology 104: 5156. https://doi.org/10.1007/BF02244553CrossRefGoogle Scholar
Spangenberg, EMF and Wichman, A 2018 Methods for investi-gating the motivation of mice to explore and access food rewards. Journal of the American Association for Laboratory Animal Science 57: 244252. https://doi.org/10.30802/AALAS-JAALAS-17-000080CrossRefGoogle ScholarPubMed
Spruijt, BM, van den Bos, R and Pijlman, FTA 2001 A con-cept of welfare based on reward evaluating mechanisms in the brain; anticipatory behaviour as an indicator for the state of reward systems. Applied Animal Behaviour Science 72: 145171. https://doi.org/10.1016/S0168-1591(00)00204-5CrossRefGoogle Scholar
Staddon, JER and Ayres, SL 1975 Sequential and temporal properties of behavior induced by a schedule of periodic food delivery. Behaviour 54: 2649. https://doi.org/10.1163/156853975X00317CrossRefGoogle Scholar
Staddon, JER and Simmelhag, VL 1971 The ‘superstition’ experiment: A re-examination of its implications for the principle of adaptive behaviour. Psychological Review 78: 343. https://doi.org/10.1037/h0030305Google Scholar
Timberlake, W and Washburne, DL 1989 Feeding ecology and laboratory predatory behavior toward live and artificial moving prey in seven rodent species. Animal Learning and Behavior 17: 211. https://doi.org/10.3758/BF03205206CrossRefGoogle Scholar
Tornatzky, W and Miczek, KA 1995 Alcohol, anxiolytics and social stress in rats. Psychopharmacology 121: 135144. https://doi.org/10.1007/BF02245600CrossRefGoogle ScholarPubMed
van den Berg, CL, Pijlman, FTA, Koning, HAM, Diergaarde, L, van Ree, JM and Spruijt, BM 1999 Isolation changes the incentive value of sucrose and social behaviour in juvenile and adult rats. Behavioural Brain Research 106: 133142. https://doi.org/10.1016/S0166-4328(99)00099-6CrossRefGoogle ScholarPubMed
van den Bos, R, Meijer, MK, van Renselaar, JP, van der Harst, JE and Spruijt, BM 2003 Anticipation is differently expressed in rats Rattus norvegicus and domestic cats Felis silvestris catus in the same Pavlovian conditioning paradigm. Behavioural Brain Research 141: 8389. https://doi.org/10.1016/S0166-4328(02)00318-2Google Scholar
van den Bos, R, van der Harst, J, Vijftigschild, N, Spruijt, B, van Luijtelaar, G and Maes, R 2004 On the relationship between anticipatory behaviour in a Pavlovian paradigm and Pavlovian-to-Instrumental Transfer in rats Rattus norvegicus. Behavioural Brain Research 153: 397408. https://doi.org/10.1016/j.bbr.2003.12.017CrossRefGoogle Scholar
van der Harst, JE, Baars, A-M and Spruijt, BM 2003b Standard housed rats are more sensitive to rewards than enriched housed rats as reflected by their anticipatory behaviour. Behavioural Brain Research 142: 151156. https://doi.org/10.1016/S0166-4328(02)00403-5CrossRefGoogle ScholarPubMed
van der Harst, JE, Baars, A-M and Spruijt, BM 2005 Announced rewards counteract the impairment of anticipatory behaviour in socially stressed rats. Behavioural Brain Research 161: 183189. https://doi.org/10.1016/j.bbr.2005.02.029CrossRefGoogle ScholarPubMed
van der Harst, JE, Fermont, PCJ, Bilstra, AE and Spruijt, BM 2003a Access to enriched housing is rewarding to rats as reflect-ed by their anticipatory behaviour. Animal Behaviour 66: 493504. https://doi.org/10.1006/anbe.2003.2201CrossRefGoogle Scholar
van der Harst, JE and Spruijt, BM 2007 Tools to measure and improve animal welfare: reward-related behaviour. Animal Welfare 16: 677310.1017/S0962728600031742CrossRefGoogle Scholar
Vinke, CM, Houx, BB, van den Bos, R and Spruijt, BM 2006 Anticipatory behaviour and stereotypical behaviour in farmed mink Mustela vision in the presence, absence and after removal of swimming water. Applied Animal Behaviour Science 96: 129142. https://doi.org/10.1016/j.applanim.2005.04.022CrossRefGoogle Scholar
Vinke, CM, van den Bos, R and Spruijt, BM 2004 Anticipatory activity and stereotypical behaviour in American mink Mustela vison in three housing systems differing in the amount of enrich-ment. Applied Animal Behaviour Science 89: 145161. https://doi.org/10.1016/j.applanim.2004.06.002CrossRefGoogle Scholar
Vivian, JA and Miczek, KA 1993 Diazepam and gepirone selectively attenuate either 20-32 or 32-64 kHz ultrasonic vocalizations during aggressive encounters. Psychopharmacology 112: 6673. https://doi.org/10.1007/BF02247364CrossRefGoogle ScholarPubMed
von Frijtag, JC, van den Bos, R and Spruijt, BM 2002 Imipramine restores the long-term impairment of appetitive behaviour in socially stressed rats. Psychopharmacology 162: 232238. https://doi.org/10.1007/s00213-002-1093-3CrossRefGoogle ScholarPubMed
Watters, JV 2014 Searching for behavioral indicators of welfare in zoos: Uncovering anticipatory behavior. Zoo Biology 33: 251256. https://doi.org/10.1002/zoo.21144CrossRefGoogle ScholarPubMed
Wichman, A, Keeling, LJ and Forkman, B 2012 Cognitive bias and anticipatory behaviour of laying hens housed in basic and enriched pens. Applied Animal Behaviour Science 140: 6269. https://doi.org/10.1016/j.applanim.2012.05.006CrossRefGoogle Scholar
Wise, RA 2008 Dopamine and reward: the anhedonia hypothesis 30 years on. Neurotoxicity Research 14: 169183. https://doi.org/10.1007/BF03033808Google Scholar
Yayou, K-I, Nakamura, M and Ito, S 2009 Effects of AVP V1a and CRH receptor antagonist on psychological stress responses to frustrating condition in sheep. Journal of Veterinary Medical Science 71: 431439. https://doi.org/10.1292/jvms.71.431CrossRefGoogle ScholarPubMed
Zimmerman, PH, Buijs, SAF, Bolhuis, JE and Keeling, LJ 2011 Behaviour of domestic fowl in anticipation of positive and negative stimuli. Animal Behaviour 81: 569577. https://doi.org/10.1016/j.anbehav.2010.11.028CrossRefGoogle Scholar
Zimmerman, PH and Koene, P 1998 The effect of frustrative non reward on vocalisations and behaviour in the laying hen, Gallus gallus domesticus. Behavioural Processes 44: 7379. https://doi.org/10.1016/S0376-6357(98)00035-7Google Scholar
Zimmerman, PH, Koene, P and van Hooff, JA 2000 Thwarting of behaviour in different contexts and the gakel-call in the laying hen. Applied Animal Behaviour Science 69: 255264. https://doi.org/10.1016/S0168-1591(00)00137-4CrossRefGoogle ScholarPubMed
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