Animal training relies heavily on an understanding of species-specific behaviour as it integrates with operant conditioning principles. Following on from recent studies showing that affective states and arousal levels may correlate with behavioural outcomes, we explore the contribution of both affective state and arousal in behavioural responses to operant conditioning. This paper provides a framework for assessing how affective state and arousal may influence the efficacy of operant training methods. It provides a series of three-dimensional conceptual graphs as exemplars to describing putative influences of both affective state and arousal on the likelihood of dogs and horses performing commonly desired behaviours. These graphs are referred to as response landscapes, and they highlight the flexibility available for improving training efficacy and the likely need for different approaches to suit animals in different affective states and at various levels of arousal. Knowledge gaps are discussed and suggestions made for bridging them.
References
[1]
McGreevy, P.; Boakes, R.A. Carrots and Sticks: Principles of Animal Training; Cambridge University Press: Cambridge, UK, 2007.
[2]
Panksepp, J. Affective Neuroscience the Foundations of Human and Animal Emotions; Oxford University Press: New York, NY, USA, 1998.
[3]
McGreevy, P.D.; Starling, M.; Branson, N.J.; Cobb, M.L.; Calnon, D. An overview of the dog-human dyad and ethograms within it. J. Vet. Behav. Clin. Appl. Res. 2012, 7, 103–117, doi:10.1016/j.jveb.2011.06.001.
[4]
Barrett, L.F.; Mesquita, B.; Ochsner, K.N.; Gross, J.J. The experience of emotion. Annu. Rev. Psychol. 2007, 58, 373–403, doi:10.1146/annurev.psych.58.110405.085709.
[5]
Russell, J.A.; Bullock, M. Multidimensional scaling of emotional facial expressions: Similarity from preschoolers to adults. J. Personal. Soc. Psychol. 1985, 48, 1290–1298, doi:10.1037/0022-3514.48.5.1290.
[6]
Moruzzi, G. Sleep and Instinctive Behaviour. Arch. Ital. Biol. 1969, 107, 175–216.
[7]
Bradley, M.M.; Miccoli, L.; Escrig, M.A.; Lang, P.J. The pupil as a measure of emotional arousal and autonomic activation. Psychophysiology 2008, 45, 602–607, doi:10.1111/j.1469-8986.2008.00654.x.
[8]
Jones, B.E. Arousal systems. Front. Biosci. 2003, 8, S438–S451, doi:10.2741/1074.
[9]
Williams, L.M.; Phillips, M.L.; Brammer, M.J.; Skerrett, D.; Lagopoulos, J.; Rennie, C.; Bahramali, H.; Olivieri, G.; David, A.S.; Peduto, A.; et al. Arousal dissociates amygdala and hippocampal fear responses: Evidence from simultaneous fMRI and skin conductance recording. NeuroImage 2001, 14, 1070–1079, doi:10.1006/nimg.2001.0904.
[10]
Yerkes, R.M.; Dodson, D.D. The relation of strength of stimulus to rapidity of habit-formation. Psychol. J. Comp. Neurol. 1908, 459–482, doi:10.1002/cne.920180503.
[11]
Eysenck, M. Attention and Aroual: Cognition and Performance; Springer: Berlin, Germany, 1982.
[12]
Hanoch, Y. When less is more: Information, emotional arousal and the ecological reframing of the yerkes-dodson law. Theory Psychol. 2004, 14, 427–452, doi:10.1177/0959354304044918.
[13]
Mair, R.G.; Onos, K.D.; Hembrook, J.R. Cognitive activation by central thalamic stimulation: The yerkes-dodson law revisited. Dose-Response 2011, 9, 313–331, doi:10.2203/dose-response.10-017.Mair.
[14]
Mendl, M.; Burman, O.; Parker, R.; Paul, E. Cognitive bias as an indicator of animal emotion and welfare: Emerging evidence and underlying mechanisms. Appl. Anim. Behav. Sci. 2009, 118, 161–181, doi:10.1016/j.applanim.2009.02.023.
[15]
Bethell, E.; Holmes, A.; Maclarnon, A. Cognitive bias in a non-human primate: Husbandry procedures influence cognitive indicators of psychological well-being in captive rhesus macaques. Anim. Welf. 2012, 21, 185–195, doi:10.7120/09627286.21.2.185.
[16]
Salmeto, A.L.; Hymel, K.A.; Carpenter, E.C.; Brilot, B.O.; Bateson, M.; Sufka, K.J. Cognitive bias in the chick anxiety-depression model. Brain Res. 2011, 1373, 124–130.
[17]
Burman, O.; Parker, R.; Paul, E.; Mendl, M. A spatial judgement task to determine background emotional state in laboratory rats, Rattus norvegicus. Anim. Behav. 2008, 76, 801–809, doi:10.1016/j.anbehav.2008.02.014.
[18]
Mendl, M.; Brooks, J.; Basse, C.; Burman, O.; Paul, E.; Blackwell, E.; Casey, R. Dogs showing separation-related behaviour exhibit a “pessimistic” cognitive bias. Curr. Biol. 2010, 20, R839–R840.
[19]
Matheson, S.; Asher, L.; Bateson, M. Larger, enriched cages are associated with “optimistic” response biases in captive European starlings (Sturnus vulgaris). Appl. Anim. Behav. Sci. 2008, 109, 374–383, doi:10.1016/j.applanim.2007.03.007.
[20]
Brilot, B.; Asher, L.; Bateson, M. Stereotyping starlings are more “pessimistic”. Anim. Cogn. 2010, 13, 721–731.
[21]
Bateson, M.; Desire, S.; Gartside, S.E.; Wright, G.A. Agitated honeybees exhibit pessimistic cognitive biases. Curr. Biol. 2011, 21, 1070–1073.
[22]
Bateson, M.; Matheson, S. Performance on a categorisation task suggests that removal of environmental enrichment induces “pessimism” in captive European starlings (Sturnus vulgaris). Anim. Welf. 2007, 16, 33–36.
[23]
Doyle, R.E.; Lee, C.; Deiss, V.; Fisher, A.D.; Hinch, G.N.; Boissy, A. Measuring judgement bias and emotional reactivity in sheep following long-term exposure to unpredictable and aversive events. Physiol. Behav. 2011, 102, 503–510.
[24]
Destrez, A.; Deiss, V.; Belzung, C.; Lee, C.; Boissy, A. Does reduction of fearfulness tend to reduce pessimistic-like judgment in lambs? Appl. Anim. Behav. Sci. 2012, 139, 233–241.
Bar-Haim, Y.; Lamy, D.; Pergamin, L.; Bakermans-Kranenburg, M.J.; van IJzendoorn, M.H. Threat-related attentional bias in anxious and nonanxious individuals: A meta-analytic study. Psychol. Bull. 2007, 133, 1–24.
[27]
Eysenck, M.; Mogg, K.; May, J.; Richards, A.; Mathews, A. Bias in interpretation of ambiguous sentences related to threat in anxiety. J. Abnorm. Psychol. 1991, 100, 144–150, doi:10.1037/0021-843X.100.2.144.
[28]
Mendl, M.; Burman, O.H.P.; Paul, E.S. An integrative and functional framework for the study of animal emotion and mood. Proc. R. Soc. B: Biol. Sci. 2010, 277, 2895–2904, doi:10.1098/rspb.2010.0303.
[29]
Jing, J.; Gillette, R.; Weiss, K.R. Evolving concepts of arousal: Insights from simple model systems. Rev. Neurosci. 2009, 20, 405–427.
[30]
Baron, A.; Galizio, M. Positive and negative reinforcement: Should the distinction be preserved? Behav. Anal. 2005, 28, 85–98.
[31]
Baker, T.B.; Piper, M.E.; McCarthy, D.E.; Majeskie, M.R.; Fiore, M.C. Addiction motivation reformulated: An affective processing model of negative reinforcement. Psychol. Rev. 2004, 111, 33–51, doi:10.1037/0033-295X.111.1.33.
[32]
Hickey, C.; Chelazzi, L.; Theeuwes, J. Reward guides vision when it’s your thing: Trait reward-seeking in reward-mediated visual priming. PLoS One 2010, 5, doi:10.1371/journal.pone.0014087.
[33]
Depue, R.; Collins, P. Neurobiology of the structure of personality: Dopamine, facilitation of incentive motivation, and extraversion. Behav. Brain Sci. 1999, 22, 491–517.
[34]
Bogdan, R.; Pizzagalli, D. The heritability of hedonic capacity and perceived stress: A twin study evaluation of candidate depressive phenotypes. Psychol. Med. 2008, 39, 211–218.
[35]
McGreevy, P.; McLean, A.; Keay, K.; Thomson, P. SMART: Sensitivity models for animals in response to training. Vet. J. 2009, 181, 72–73, doi:10.1016/j.tvjl.2009.03.014.
[36]
Nijhout, H.F. The importance of context in genetics. 2003, 91, 416–423.
[37]
Overall, K.L. Proceedings of the Dogs Trust Meeting on Advances in Veterinary Behavioural Medicine London; 4th–7th November 2004: Veterinary Behavioural Medicine: A Roadmap for the 21st Century. Vet. J. 2005, 169, 130–143, doi:10.1016/j.tvjl.2004.10.007.
[38]
Haverbeke, A.; Laporte, B.; Depiereux, E.; Giffroy, J.M.; Diederich, C. Training methods of military dog handlers and their effects on the team’s performances. Appl. Anim. Behav. Sci. 2008, 113, 110–122, doi:10.1016/j.applanim.2007.11.010.
[39]
Miller, H.C.; Pattison, K.F.; DeWall, C.N.; Rayburn-Reeves, R.; Zentall, T.R. Self-control without a “self?”: Common self-control processes in humans and dogs. Psychol. Sci. 2010, 21, 534–538, doi:10.1177/0956797610364968.
[40]
Miller, H.C.; DeWall, C.N.; Pattison, K.; Molet, M.; Zentall, T.R. Too dog tired to avoid danger: Self-control depletion in canines increases behavioral approach toward an aggressive threat. Psychon. Bull. Rev. 2012, 19, 535–540, doi:10.3758/s13423-012-0231-0.
[41]
Robbins, T.W. Arousal systems and attentional processes. Biol. Psychol. 1997, 45, 57–71, doi:10.1016/S0301-0511(96)05222-2.
[42]
Pfaff, D.; Ribeiro, A.; Matthews, J.; Kow, L.-M. Concepts and mechanisms of generalized central nervous system arousal. Ann. N.Y. Acad. Sci. 2008, 1129, 11–25.
Tortora, D.F. Safety training: The elimination of avoidance-motivated aggression in dogs. J. Exp. Psychol. Gen. 1983, 112, 176–214, doi:10.1037/0096-3445.112.2.176.
[45]
Rooney, N.J.; Cowan, S. Training methods and owner-dog interactions: Links with dog behaviour and learning ability. Appl. Anim. Behav. Sci. 2011, 132, 169–177, doi:10.1016/j.applanim.2011.03.007.
[46]
Herron, M.E.; Shofer, F.S.; Reisner, I.R. Survey of the use and outcome of confrontational and non-confrontational training methods in client-owned dogs showing undesired behaviors. Appl. Anim. Behav. Sci. 2009, 117, 47–54, doi:10.1016/j.applanim.2008.12.011.
[47]
Blackwell, E.J.; Twells, C.; Seawright, A.; Casey, R.A. The relationship between training methods and the occurrence of behavior problems, as reported by owners, in a population of domestic dogs. J. Vet. Behav. Clin. Appl. Res. 2008, 3, 207–217, doi:10.1016/j.jveb.2007.10.008.
[48]
Haverbeke, A.; De Smet, A.; Depiereux, E.; Giffroy, J.; Diederich, C. Assessing undesired aggression in military working dogs. Appl. Anim. Behav. Sci. 2009, 117, 55–62, doi:10.1016/j.applanim.2008.12.002.
[49]
Schalke, E.; Stichnoth, J.; Ott, S.; Jones-Baade, R. Clinical signs caused by the use of electric training collars on dogs in everyday life situations. Appl. Anim. Behav. Sci. 2007, 105, 369–380, doi:10.1016/j.applanim.2006.11.002.
[50]
Schilder, M.B.H.; van der Borg, J.A.M. Training dogs with help of the shock collar: Short and long term behavioural effects. Appl. Anim. Behav. Sci. 2004, 85, 319–334, doi:10.1016/j.applanim.2003.10.004.
[51]
McLean, A.N.; McGreevy, P.D. Horse-training techniques that may defy the principles of learning theory and compromise welfare. J. Vet. Behav.: Clin. Appl. Res. 2010, 5, 187–195, doi:10.1016/j.jveb.2010.04.002.
[52]
Jones, B.; McGreevy, P.D. Ethical equitation: Applying a cost-benefit approach. J. Vet. Behav.: Clin. Appl. Res. 2010, 5, 196–202, doi:10.1016/j.jveb.2010.04.001.
[53]
Range, F.; Heucke, S.L.; Gruber, C.; Konz, A.; Huber, L.; Virányi, Z. The effect of ostensive cues on dogs’ performance in a manipulative social learning task. Appl. Anim. Behav. Sci. 2009, 120, 170–178, doi:10.1016/j.applanim.2009.05.012.
[54]
Garland, E.L.; Fredrickson, B.; Kring, A.M.; Johnson, D.P.; Meyer, P.S.; Penn, D.L. Upward spirals of positive emotions counter downward spirals of negativity: Insights from the broaden-and-build theory and affective neuroscience on the treatment of emotion dysfunctions and deficits in psychopathology. Clin. Psychol. Rev. 2010, 30, 849–864.
[55]
Mathews, A.; MacLeod, C. Selective processing of threat cues in anxiety states. Behav. Res. Ther. 1985, 23, 563–569, doi:10.1016/0005-7967(85)90104-4.