An integrative and functional framework for the study of animal emotion and mood
Open accessAbstract
A better understanding of animal emotion is an important goal in disciplines ranging from neuroscience to animal welfare science. The conscious experience of emotion cannot be assessed directly, but neural, behavioural and physiological indicators of emotion can be measured. Researchers have used these measures to characterize how animals respond to situations assumed to induce discrete emotional states (e.g. fear). While advancing our understanding of specific emotions, this discrete emotion approach lacks an overarching framework that can incorporate and integrate the wide range of possible emotional states. Dimensional approaches that conceptualize emotions in terms of universal core affective characteristics (e.g. valence (positivity versus negativity) and arousal) can provide such a framework. Here, we bring together discrete and dimensional approaches to: (i) offer a structure for integrating different discrete emotions that provides a functional perspective on the adaptive value of emotional states, (ii) suggest how long-term mood states arise from short-term discrete emotions, how they also influence these discrete emotions through a bi-directional relationship and how they may function to guide decision-making, and (iii) generate novel hypothesis-driven measures of animal emotion and mood.
The claims and statistics of this paper haven’t been checked yet.
Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.
Integrity signals
Registry
Cites
Statistics
Claims the paper makes
The claims and statistics of this paper haven’t been checked yet.
Run the check — free, no account needed. It reads the open-access text and checks the sources it cites.
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
Fecal corticosterone may reflect an increased state of arousal, but the arousal may not be negative.
“Although fecal corticosterone may reflect an increased state of arousal, the arousal may not be negative.29”
From Ultrasonic Vocalization Analysis as a Novel Metric to Assess Cage Enrichment in Rats (2022) · cites this paper as [29]
Section 4, “Implications for the Measurement of Animal Emotion” (page unavailable), says physiological measures, including hypothalamic-pituitary-adrenal activity, may reflect arousal without distinguishing positive from negative states. The paper does not specifically discuss fecal corticosterone, so it supports the general caution but not its application to that measure. (LLM, conf=0.85)
Experimental — accuracy not yet measured
Self-check
Percentages, sums and reported differences re-computed from the paper’s own numbers.
The self-check needs the paper’s full text.
Cites
Core affect and the psychological construction of emotion.
Russell, J. A. (2003) · Psychological Review · doi:10.1037/0033-295x.110.1.145 · Cited by 6,113 · see citing works
Schwarz, N., & Clore, G. L. (1983) · Journal of Personality and Social Psychology · doi:10.1037/0022-3514.45.3.513 · Cited by 4,952 · see citing works
POSNER, J., RUSSELL, J. A., & PETERSON, B. S. (2005) · Development and Psychopathology · doi:10.1017/s0954579405050340 · Cited by 2,602 · see citing works
Emotion, attention, and the startle reflex
Lang, P. J., Bradley, M. M., & Cuthbert, B. N. (1990) · Psychological Review · doi:10.1037/0033-295x.97.3.377 · Cited by 2,205 · see citing works
Barrett, L. F. (2006) · Perspectives on Psychological Science · doi:10.1111/j.1745-6916.2006.00003.x · Cited by 1,859 · see citing works
Comorbidity of anxiety and unipolar mood disorders
Mineka, S., Watson, D., & Clark, L. A. (1998) · Annual Review of Psychology · doi:10.1146/annurev.psych.49.1.377 · Cited by 1,573 · see citing works
Affective style and affective disorders: Perspectives from affective neuroscience
Davidson, R. J. (1998) · Cognition & Emotion · doi:10.1080/026999398379628 · Cited by 1,496 · see citing works
Assessment of positive emotions in animals to improve their welfare
Boissy, A., Manteuffel, G., Jensen, M. B., Moe, R. O., Spruijt, B., Keeling, L. J., Winckler, C., Forkman, B., Dimitrov, I., Langbein, J., et al. (2007) · Physiology & Behavior · doi:10.1016/j.physbeh.2007.02.003 · Cited by 1,486 · see citing works
Barrett, L. F., Mesquita, B., Ochsner, K. N., & Gross, J. J. (2007) · Annual Review of Psychology · doi:10.1146/annurev.psych.58.110405.085709 · Cited by 1,425 · see citing works
Cognitive bias and affective state
Harding, E. J., Paul, E. S., & Mendl, M. (2004) · Nature · doi:10.1038/427312a · Cited by 835 · see citing works
Self-citationA common neurobiology for pain and pleasure
Leknes, S., & Tracey, I. (2008) · Nature Reviews Neuroscience · doi:10.1038/nrn2333 · Cited by 784 · cited 1× · see citing works
“…opioidergic) appear to be prime candidates for the latter reward/punishment systems (Gray 1994; Burgdorf & Panksepp 2006; Berridge 2007; Leknes & Tracey 2008), and it is possible that they, and higher cortical areas (e.g. anterior cingulate cortex, orbitofrontal cortex), somehow…”
BackgroundNatural selection and the regulation of defenses
Nesse, R. M. (2005) · Evolution and Human Behavior · doi:10.1016/j.evolhumbehav.2004.08.002 · Cited by 365 · see citing works
Rolls, E. T. (2005) · doi:10.1093/acprof:oso/9780198570035.001.0001 · Cited by 339 · see citing works
Sex, trauma, stress hormones and depression
Young, E., & Korszun, A. (2010) · Molecular Psychiatry · doi:10.1038/mp.2009.94 · Cited by 235 · see citing works
Flagged first, then by citations
Research works printed but not linked: 46
- Bechara, A., Damasio, H. & Damasio, A. R. 2000 Emotion, decision-making and the orbitofrontal cortex. Cereb. Cortex 10 , 295-307. (doi:10.1093/cercor/10.3.295) · DOI · Paper page
- Beck, A. T. 2008 The evolution of the cognitive model of depression and its neurobiological correlates. Am. J. Psychiatry 165 , 969-977. (doi:10.1176/appi.ajp. 2008.08050721) · DOI
- Berridge, K. C. 1996 Food reward: brain substrates of wanting and liking. Neurosci. Biobehav. Rev. 20 , 1-25. (doi:10.1016/0149-7634(95)00033-B) · DOI · Paper page
- Berridge, K. C. 2003 Comparing the emotional brains of humans and other animals. In Handbook of affective sciences (eds R. J. Davidson, K. R. Scherer & H. H. Goldsmith), pp. 25-51. Oxford, UK: Oxford University Press.
- Berridge, K. C. 2007 The debate over dopamine's role in reward: the case for incentive salience. Psychopharmacology 191 , 391-431. (doi:10.1007/s00213-006-0578-x) · DOI · Paper page
- Boissy, A. 1995 Fear and fearfulness in animals. Q. Rev. Biol. 70 , 165-191. (doi:10.1086/418981) · DOI · Paper page
- Burgdorf, J. & Panksepp, J. 2006 The neurobiology of positive emotions. Neurosci. Biobehav. Rev. 30 , 173-187. (doi:10.1016/j.neubiorev.2005.06.001) · DOI · Paper page
- Burman, O. H. P., Parker, R., Paul, E. S. & Mendl, M. 2008 A spatial judgement task to determine background emotional state in laboratory rats, Rattus norvegicus . Anim. Behav. 76 , 801-809. (doi:10.1016/j.anbehav. 2008.02.014) · DOI
- Cabanac, M. 1992 Pleasure-the common currency. J. Theor. Biol. 155 , 173-200. (doi:10.1016/S00225193(05)80594-6) · DOI
- Cacioppo, J. T., Gardner, W . L. & Berntson, G. G. 1999 The affect system has parallel and integrative processing components: form follows function. J. Pers. Soc. Psychol. 76 , 839-855. (doi:10.1037/0022-3514.76.5.839) · DOI · Paper page
- Corr, P. J. 2008 The reinforcement sensitivity theory of personality. Cambridge, UK: Cambridge University Press.
- Dalgleish, T. 2004 The emotional brain. Nat. Rev. Neurosci. 5 , 582-589. (doi:10.1038/nrn1432) · DOI · Paper page
- Davidson, R. J. 1994 On emotion, mood, and related affective constructs. In The nature of emotion (eds P. Ekman & R. J. Davidson), pp. 51-55. New York, NY: Oxford University Press.
- Davidson, R. J. & Irwin, W. 1999 The functional neuroanatomy of emotion and affective style. Trends Cogn. Sci. 3 , 11-21. (doi:10.1016/S1364-6613(98)01265-0) · DOI · Paper page
- Dawkins, R. & Krebs, J. R. 1979 Arms races between and within species. Proc. R. Soc. Lond. B 205 , 489-511. (doi:10.1098/rspb.1979.0081) · DOI
- Desire ´, L., Boissy, A. & Veissier, I. 2002 Emotions in farm animals: a new approach to animal welfare in applied ethology. Behav. Proc. 60 , 165-180. (doi:10.1016/ S0376-6357(02)00081-5)
- Doyle, R. E., Fisher, A. D., Hinch, G. N., Boissy, A. & Lee, C. 2010 Release from restraint generates a positive judgement bias in sheep. Appl. Anim. Behav. Sci. 122 , 28-34. (doi:10.1016/j.applanim.2009.11.003) · DOI
- Ekman, P. 1992 Are there basic emotions. Psychol. Rev. 99 , 550-553. (doi:10.1037/0033-295X.99.3.550) · DOI
- Ellsworth, P. C. & Scherer, K. E. 2003 Appraisal processes in emotion. In Handbook of affective sciences (eds R. J. Davidson, K. R. Scherer & H. H. Goldsmith), pp. 572595. Oxford, UK: Oxford University Press.
- Forkman,B.,Boissy,A.,Meunier-Salauen,M.C.,Canali,E.& Jones, R. B. 2007 A critical review of fear tests used on cattle, pigs, sheep, poultry and horses. Physiol. Behav. 92 , 340-374. (doi:10.1016/j.physbeh.2007.03.016) · DOI
- Frijda, N. 1994 Emotions are functional, most of the time. In The nature of emotion (eds P. Ekman & R. J. Davidson), pp. 112-122. New York, NY: Oxford University Press.
- Grandjean, D. & Scherer, K. R. 2008 Unpacking the cognitive architecture of emotion processes. Emotion 8 , 341-351. (doi:10.1037/1528-3542.8.3.341) · DOI
- Gray, J. A. 1994 Three fundamental emotion systems. In The nature of emotion (eds P. Ekman & R. J. Davidson), pp. 243-247. New York, NY: Oxford University Press.
- Izard, C. E. 2007 Basic emotions, natural kinds, emotion schemas and a new paradigm. Perspect. Psychol. Sci. 2 , 260-280. (doi:10.1111/j.1745-6916.2007.00044.x) · DOI · Paper page
- Loewenstein, G. F., Weber, E. U., Hsee, C. K. & Welch, N. 2001 Risk as feelings. Psychol. Bull. 127 , 267-286. (doi:10.1037/0033-2909.127.2.267) · DOI · Paper page
- Macphail, E. M. 1998 The evolution of consciousness. Oxford, UK: Oxford University Press.
- McNamara, J. M. & Houston, A. I. 1986 The common currency for behavioral decisions. Am. Nat. 127 , 358-378. (doi:10.1086/284489) · DOI
- McNaughton, N. & Corr, P. J. 2008 The neuropsychology of fear and anxiety: a foundation for reinforcement sensitivity theory. In The reinforcement sensitivity theory of personality (ed. P. J. Corr), pp. 44-94. Cambridge, UK: Cambridge University Press.
- Miller, K. A., Garner, J. P . & Mench, J. A. 2006 Is fearfulness a trait that can be measured with behavioural tests? A validation of four fear tests for Japanese quail. Anim. Behav. 71 , 1323-1334. (doi:10.1016/j.anbehav.2005.08.018) · DOI
- Mogg, K. & Bradley, B. P. 1998 A cognitive-motivational analysis of anxiety. Behav. Res. Ther. 36 , 809-848. (doi:10.1016/S0005-7967(98)00063-1) · DOI · Paper page
- Nesse, R. M. 2000 Is depression an adaptation? Arch. Gen. Psychiat. 57 , 14-20. (doi:10.1001/archpsyc.57.1.14) · DOI · Paper page
- Nesse, R. M. 2004 Natural selection and the elusiveness of happiness. Phil. Trans. R. Soc. Lond. B 359 , 1333-1347. (doi:10.1098/rstb.2004.1511) · DOI
- Nesse, R. M. & Ellsworth, P. C. 2009 Evolution, emotions, and emotional disorders. Am. Psychol. 64 , 129-139. (doi:10.1037/a0013503) · DOI
- Nygren, T. E., Isen, A. M., Taylor, P . J. & Dulin, J. 1996 The influence of positive affect on the decision rule in risky situations. Org. Behav. Hum. Decis. Process 66 , 79-91. (doi:10.1006/obhd.1996.0038) · DOI
- Panksepp, J. 1998 In Affective neuroscience. The foundations of human and animal emotion. New York, NY: Oxford University Press.
- Panksepp, J. 2007 Neurologizing the psychology of affects. How appraisal-based constructivism and basic emotion theory can coexist. Perspect. Psychol. Sci 2 , 281-296. (doi:10.1111/j.1745-6916.2007.00045.x) · DOI
- Paul, E. S., Harding, E. J. & Mendl, M. 2005 Measuring emotional processes in animals: the utility of a cognitive approach. Neurosci. Biobehav. Rev. 29 , 469-491. (doi:10.1016/j.neubiorev.2005.01.002) · DOI
- Porges, S. W. 2001 Is there a major stress system at the periphery other than the adrenals? In Coping with challenge. Welfare in animals including humans (ed. D. M. Broom), pp. 135-149. Berlin, Germany: Dahlem University Press.
- Prinz, J. J. 2004 Gut reactions. A perceptual theory of emotion. Oxford, UK: Oxford University Press.
- Ramos, A. & Mormede, P. 1998 Stress and emotionality: a multidimensional and genetic approach. Neurosci. Biobehav. Rev. 22 , 33-57. (doi:10.1016/S01497634(97)00001-8) · DOI
- Russell, J. A. & Barrett, L. F. 1999 Core affect, prototypical emotional episodes, and other things called emotion: dissecting the elephant. J. Pers. Soc. Psychol. 76 , 805-819. (doi:10.1037/0022-3514.76.5.805) · DOI
- Scherer, K. R. 2001 Appraisal considered as a process of multi-level sequential checking. In Appraisal process in emotion: theory, methods, research (eds K. R. Scherer, A. Schorr & T. Johnstone), pp. 92-120. Oxford, UK: Oxford University Press.
- Spruijt, B. M., van den Bos, R. & Pijlman, F. T. A. 2001 A concept of welfare based on reward evaluating mechanisms in the brain: anticipatory behaviour as an indicator for the state of reward systems. Appl. Anim. Behav. Sci. 72 , 145-171. (doi:10.1016/S0168-1591(00)00204-5) · DOI · Paper page
- Stanley, D. J. & Meyer, J. P . 2009 Two-dimensional affective space: a new approach to orienting the axes. Emotion 9 , 214-237. (doi:10.1037/a0014612) · DOI
- Tellegen, A., Watson, D. & Clark, L. A. 1999 On the dimensional and hierarchical structure of affect. Psychol. Sci. 10 , 297-303. (doi:10.1111/1467-9280. 00157) · DOI
- Wemelsfelder, F. 1997 The scientific validity of subjective concepts in models of animal welfare. Appl. Anim. Behav. Sci. 53 , 75-88. (doi:10.1016/S01681591(96)01152-5) · DOI
Cited by
Sorted by citations
The role of sleep in emotional brain function
Goldstein, A. N., & Walker, M. P. (2014) · Annual Review of Clinical Psychology · doi:10.1146/annurev-clinpsy-032813-153716 · Cited by 1,052 · see citing works
“…less well characterized, interaction between sleep and mood states; the latter which we and others considered distinct from emotions (emotions being shortlived events, often in response to external stimuli, while mood states are more sustained events, often internally generated (Mendl et al 2010)).”
BackgroundA framework for studying emotions across species
Anderson, D. J., & Adolphs, R. (2014) · Cell · doi:10.1016/j.cell.2014.03.003 · Cited by 691 · see citing works
Vocal expression of emotions in mammals: mechanisms of production and evidence
Briefer, E. F. (2012) · Journal of Zoology · doi:10.1111/j.1469-7998.2012.00920.x · Cited by 541 · see citing works
Adolphs, R. (2013) · Current Biology · doi:10.1016/j.cub.2012.11.055 · Cited by 524 · see citing works
Panksepp, J. (2011) · PLoS ONE · doi:10.1371/journal.pone.0021236 · Cited by 342 · see citing works
The evolutionary origins of mood and its disorders
Nettle, D., & Bateson, M. (2012) · Current Biology · doi:10.1016/j.cub.2012.06.020 · Cited by 230 · see citing works
Assessment of welfare in zoo animals: towards optimum quality of life
Wolfensohn, S., Shotton, J., Bowley, H., Davies, S., Thompson, S., & Justice, W. S. M. (2018) · Animals · doi:10.3390/ani8070110 · Cited by 197 · see citing works
Social dimension of emotions and its implication for animal welfare
Špinka, M. (2012) · Applied Animal Behaviour Science · doi:10.1016/j.applanim.2012.02.005 · Cited by 147 · see citing works
Godlewska, B. R., & Harmer, C. J. (2021) · Psychopharmacology · doi:10.1007/s00213-019-05448-0 · Cited by 123 · see citing works
Staley, M., Conners, M. G., Hall, K., & Miller, L. J. (2018) · Hormones and Behavior · doi:10.1016/j.yhbeh.2018.04.011 · Cited by 100 · see citing works
Assessing measures of animal welfare
Browning, H. (2022) · Biology & Philosophy · doi:10.1007/s10539-022-09862-1 · Cited by 85 · see citing works
Diffusion Modelling Reveals the Decision Making Processes Underlying Negative Judgement Bias in Rats
Hales, C. A., Robinson, E. S. J., & Houghton, C. J. (2016) · PLOS ONE · doi:10.1371/journal.pone.0152592 · Cited by 49 · see citing works
Affective Biases in Humans and Animals
Robinson, E. S. J., & Roiser, J. P. (2015) · Current Topics in Behavioral Neurosciences · doi:10.1007/7854_2015_5011 · Cited by 37 · see citing works
Veasey, J. S. (2019) · Animals · doi:10.3390/ani10010039 · Cited by 36 · see citing works
Raoult, C. M. C., Moser, J., & Gygax, L. (2017) · Frontiers in Psychology · doi:10.3389/fpsyg.2017.02197 · Cited by 29 · see citing works
Ultrasonic Vocalization Analysis as a Novel Metric to Assess Cage Enrichment in Rats
Bigelow, L. J., Cohen, A. J., Pimm, R., Knight, J. B., & Bernard, P. B. (2022) · Journal of the American Association for Laboratory Animal Science · doi:10.30802/aalas-jaalas-21-000024 · Cited by 9 · see citing works
Rethinking core affect: the role of dominance in animal behaviour and welfare research
Carranza-Pinedo, V. (2024) · Synthese · doi:10.1007/s11229-024-04591-2 · Cited by 5 · see citing works
- Authors
- Mendl, M., Burman, O. H. P., & Paul, E. S.
- Affiliations
- University of Bristol (GB)
- Venue
- Proceedings of the Royal Society B: Biological Sciences
- Publisher
- The Royal Society
- Published
- 2010-08-04
- Volume
- 277
- Issue
- 1696
- Pages
- 2895-2904
- ISSN
- 0080-4649 (ISSN-L)
- 0962-8452
- 1471-2954
- PMC
- PMC2982018
- OpenAlex
- W2127099931
- Cited by
- 944
- References
- 14
- Citations by year
- 2026: 63 · 2025: 120 · 2024: 76 · 2023: 67 · 2022: 70 · 2021: 85 · 2020: 74 · 2019: 67 · 2018: 67 · 2017: 54 · 2016: 43 · 2015: 38 · 2014: 38 · 2013: 28 · 2012: 26
- FWCI
- 4.94
- Type
- journal article
- Language
- en
- Open access
- green
- www.ncbi.nlm.nih.gov
- Locations
- Proceedings of the Royal Society B Biological Sciences (publishedVersion)
- Europe PMC (PubMed Central) (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- Bristol Research (University of Bristol) (publishedVersion)
- Lincoln Repository (University of Lincoln) (acceptedVersion)
- Figshare (submittedVersion, cc-by)
- Funders
- Biotechnology and Biological Sciences Research Council (BB/C518949/1)
- Topics
- Health Sciences › Veterinary › Small Animals › Animal Behavior and Welfare Studies
- Social Sciences › Psychology › Social Psychology › Primate Behavior and Ecology
- Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
- Keywords
- animal emotions, discrete emotions, dimensional emotion models, valence, arousal, core affect, decision-making, animal welfare, emotion measurement
- MeSH terms
- Affect, Animals, Emotions, Motivation, Research Design, Reward
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
Are you an author, or do you see a problem with this page? Request a review