近年來人們對於探索無脊椎動物的類情緒狀態有顯著成長的趨勢。在頭足類動物中，烏賊擁有複雜的大腦以及靈活優秀的認知能力，但是對於烏賊是否可以經歷正向類情緒狀態的研究仍然缺乏，此篇研究基於2016年的熊蜂實驗概念(Solvi et al., 2016)改良了適用於烏賊的判斷偏誤測試。我們進行了三種版本的關聯性學習行為訓練，其中先對虎斑烏賊使用圖案與位置作為正向與負向刺激學習，在達到適合的訓練標準後，會將烏賊分成控制組與非預期獎勵組，以介於正向與負向的模糊刺激進行測試，從兩組間是否存在顯著差異探查烏賊的類情緒狀態存在與否。結果表明，使用五通道裝置可以有效促進亞成體烏賊學習，然而非預期獎勵是否可以誘導正向類情緒狀態目前仍沒有明確結論，需要更多數據去證明。而判斷偏誤測試中使用動物追蹤分析烏賊路徑與頭部方向則只有顯示出個體差異。此外圖案對調測試顯示在五通道裝置中，烏賊可能更加依賴以空間概念作學習，可能使用視覺提示作為地點標記。而幼年烏賊則由於訓練失敗，未能進入判斷偏誤測試，這表明關聯性學習訓練需要進一步修正。雖然數據有限，本篇研究為不同種類的動物情緒研究提供啟發，並在實驗方法與結果上提供參考資料，希望對未來烏賊認知與類情緒能力研究能有些幫助與理解。

In recent years, interest in exploring emotion-like states in invertebrates has significantly increased. Cephalopods, particularly cuttlefish, are renowned for their intricate brains and advanced cognitive behaviors. However, research on whether cuttlefish experience positive emotion-like states remains limited. In this study, we developed a judgment bias test based on the paradigm used in bumblebee research in 2016 (Solvi et al., 2016). We conducted three versions of associative training using patterns as positive and negative stimuli to examine potential differences in responses to ambiguous stimuli between the control and pre-reward groups of cuttlefish. The ambiguous stimuli consisted of patterns transitioning between positive and negative cues, and the unexpected shrimp given before each test trial represented a pre-reward intended to induce a positive bias in cuttlefish. Our results indicated that a five-tunnel apparatus effectively facilitated learning in subadult cuttlefish. However, the potential for a pre-reward to induce positive emotion-like behavior remains unclear. Thus, it requires additional data to determine if there is a significant difference between the control and pre-reward groups. Animal tracking was utilized to analyze the cuttlefish’s routes and head orientations during testing trials, and it revealed only individual differences. Additionally, the pattern-switching test suggested that cuttlefish may prioritize spatial stimuli, potentially using visual cues as place markers. Juvenile cuttlefish were unable to complete the judgment bias test due to the failed training, indicating the need for further modifications in the associative training protocol for juveniles. Despite the limited data, this study provides insights into emotional research across phyla and offers references for experimental methods and results to better understand the cognitive and emotion-like capacities of cuttlefish in the future.

Agin, V., Chichery, R., Dickel, L., & Chichery, M.-P. (2006). The “prawn-in-the-tube” procedure in the cuttlefish: habituation or passive avoidance learning? Learning & Memory, 13(1), 97-101.
Agin, V., Dickel, L., Chichery, R., & Chichery, M.-P. (1998). Evidence for a specific short-term memory in the cuttlefish, Sepia. Behavioural processes, 43(3), 329-334.
Alem, S., Perry, C. J., Zhu, X., Loukola, O. J., Ingraham, T., Søvik, E., & Chittka, L. (2016). Associative mechanisms allow for social learning and cultural transmission of string pulling in an insect. PLoS biology, 14(10), e1002564.
Allen-Hermanson, S. (2008). Insects and the problem of simple minds: Are bees natural zombies? The Journal of Philosophy, 105(8), 389-415.
Anderson, D. J., & Adolphs, R. (2014). A framework for studying emotions across species. Cell, 157(1), 187-200.
Baciadonna, L., & McElligott, A. G. (2015). The use of judgement bias to assess welfare in farm livestock. Animal Welfare, 24(1), 81-91.
Bacqué-Cazenave, J., Cattaert, D., Delbecque, J.-P., & Fossat, P. (2017). Social harassment induces anxiety-like behaviour in crayfish. Scientific reports, 7(1), 39935.
Bateson, M., & Nettle, D. (2015). Development of a cognitive bias methodology for measuring low mood in chimpanzees. PeerJ, 3, e998.
Billard, P., Clayton, N. S., & Jozet-Alves, C. (2020). Cuttlefish retrieve whether they smelt or saw a previously encountered item. Scientific reports, 10(1), 5413.
Boissy, A., Manteuffel, G., Jensen, M. B., Moe, R. O., Spruijt, B., Keeling, L. J., Winckler, C., Forkman, B., Dimitrov, I., & Langbein, J. (2007). Assessment of positive emotions in animals to improve their welfare. Physiology & behavior, 92(3), 375-397.
Boleij, H., van’t Klooster, J., Lavrijsen, M., Kirchhoff, S., Arndt, S. S., & Ohl, F. (2012). A test to identify judgement bias in mice. Behavioural brain research, 233(1), 45-54.
Bowers, J., Nimi, T., Wilson, J., Wagner, S., Amarie, D., & Sittaramane, V. (2020). Evidence of learning and memory in the juvenile dwarf cuttlefish Sepia bandensis. Learning & Behavior, 48, 420-431.
Burani, C., Pelosi, A., & Valsecchi, P. (2022). A promising novel judgement bias test to evaluate affective states in dogs (Canis familiaris). Anim Cogn, 25(4), 837-852.
Cahill, L., & McGaugh, J. L. (1996). Modulation of memory storage. Curr Opin Neurobiol, 6(2), 237-242.
Cole, P. D., & Adamo, S. A. (2005). Cuttlefish (Sepia officinalis: Cephalopoda) hunting behavior and associative learning. Animal Cognition, 8, 27-30.
Cooke, G. M., & Tonkins, B. M. (2015). Behavioural indicators of welfare exhibited by the common European cuttlefish (Sepia officinalis). Journal of Zoo and Aquarium Research, 3(4), 157-162.
Cooper, S. M., Holekamp, K. E., & Smale, L. (1999). A seasonal feast: long‐term analysis of feeding behaviour in the spotted hyaena (Crocuta crocuta). African Journal of Ecology, 37(2), 149-160.
Crook, R. J. (2021). Behavioral and neurophysiological evidence suggests affective pain experience in octopus. iScience, 24(3), 102229.
Darwin, C. (1872). The expression of the emotions in man and animals. J. Murray.
Dawkins, M. (2015). Animal welfare and the paradox of animal consciousness. In Advances in the Study of Behavior (Vol. 47, pp. 5-38). Elsevier.
de Vere, A. J., & Kuczaj, S. A., 2nd. (2016). Where are we in the study of animal emotions? Wiley Interdiscip Rev Cogn Sci, 7(5), 354-362.
de Waal, F. B., & Andrews, K. (2022). The question of animal emotions. Science, 375(6587), 1351-1352.
Désiré, L., Boissy, A., & Veissier, I. (2002). Emotions in farm animals:: a new approach to animal welfare in applied ethology. Behavioural processes, 60(2), 165-180.
Dickel, L., Boal, J. G., & Budelmann, B. U. (2000). The effect of early experience on learning and memory in cuttlefish. Developmental Psychobiology: The Journal of the International Society for Developmental Psychobiology, 36(2), 101-110.
Dickel, L., Chichery, M.-P., & Chichery, R. (1998). Time differences in the emergence of short-and long-term memory during post-embryonic development in the cuttlefish, Sepia. Behavioural processes, 44(1), 81-86.
Dickel, L., Chichery, M. P., & Chichery, R. (2001). Increase of learning abilities and maturation of the vertical lobe complex during postembryonic development in the cuttlefish, Sepia. Developmental Psychobiology: The Journal of the International Society for Developmental Psychobiology, 39(2), 92-98.
Dolensek, N., Gehrlach, D. A., Klein, A. S., & Gogolla, N. (2020). Facial expressions of emotion states and their neuronal correlates in mice. Science, 368(6486), 89-94.
Douglas, C., Bateson, M., Walsh, C., Bédué, A., & Edwards, S. A. (2012). Environmental enrichment induces optimistic cognitive biases in pigs. Applied Animal Behaviour Science, 139(1-2), 65-73.
Ekman, P. (1992). Are There Basic Emotions. Psychological Review, 99(3), 550-553. 10.1037/0033-295x.99.3.550
Ellsworth, P. C. (1991). Some implications of cognitive appraisal theories of emotion.
Espigares, F., Abad-Tortosa, D., Varela, S. A. M., Ferreira, M. G., & Oliveira, R. F. (2021). Short telomeres drive pessimistic judgement bias in zebrafish. Biol Lett, 17(3), 20200745.
Eysenck, M. W., Mogg, K., May, J., Richards, A., & Mathews, A. (1991). Bias in interpretation of ambiguous sentences related to threat in anxiety. J Abnorm Psychol, 100(2), 144-150.
Fossat, P., Bacqué-Cazenave, J., De Deurwaerdère, P., Delbecque, J.-P., & Cattaert, D. (2014). Anxiety-like behavior in crayfish is controlled by serotonin. Science, 344(6189), 1293-1297.
Fredrickson, B. L. (1998). What good are positive emotions? Review of general psychology, 2(3), 300-319.
Frijda, N. H. (1987). Emotion, cognitive structure, and action tendency. Cognition and emotion, 1(2), 115-143.
Galpayage Dona, H. S., Solvi, C., Kowalewska, A., Mäkelä, K., MaBouDi, H., & Chittka, L. (2022). Do bumble bees play? Animal Behaviour, 194, 239-251.
Gendron, M. (2010). Defining Emotion: A Brief History. Emotion Review, 2(4), 371-372.
Giancola-Detmering, S., & Crook, R. J. (2024). A judgment bias task reveals stress produces a negative affective state in cuttlefish. bioRxiv, 2024.2004. 2015.589613.
Giurfa, M., Zhang, S., Jenett, A., Menzel, R., & Srinivasan, M. V. (2001). The concepts of ‘sameness’ and ‘difference’in an insect. Nature, 410(6831), 930-933.
Hamann, S. B., Ely, T. D., Grafton, S. T., & Kilts, C. D. (1999). Amygdala activity related to enhanced memory for pleasant and aversive stimuli. Nat Neurosci, 2(3), 289-293.
Hanlon, R. T., & Messenger, J. B. (2018). Cephalopod behaviour. Cambridge University Press.
Harding, E. J., Paul, E. S., & Mendl, M. (2004). Cognitive bias and affective state. Nature, 427(6972), 312-312.
Horváth, Z., Dóka, A., & Miklósi, Á. (2008). Affiliative and disciplinary behavior of human handlers during play with their dog affects cortisol concentrations in opposite directions. Hormones and behavior, 54(1), 107-114.
Izard, C. E. (2010). The many meanings/aspects of emotion: Definitions, functions, activation, and regulation. Emotion Review, 2(4), 363-370.
Jaksić, F. M. (1989). Opportunism vs selectivity among carnivorous predators that eat mammalian prey: a statistical test of hypotheses. Oikos, 427-430.
Jozet-Alves, C., Bertin, M., & Clayton, N. S. (2013). Evidence of episodic-like memory in cuttlefish. Curr Biol, 23(23), R1033-1035. https://doi.org/10.1016/j.cub.2013.10.021
Kuo, T.-H., Sneddon, L. U., Spencer, J. W., & Chiao, C.-C. (2022). Impact of lidocaine on pain-related grooming in cuttlefish. Biology, 11(11), 1560.
Kuo, T. H., & Chiao, C. C. (2020). Learned valuation during forage decision-making in cuttlefish. R Soc Open Sci, 7(12), 201602.
Lagisz, M., Zidar, J., Nakagawa, S., Neville, V., Sorato, E., Paul, E. S., Bateson, M., Mendl, M., & Lovlie, H. (2020). Optimism, pessimism and judgement bias in animals: A systematic review and meta-analysis. Neurosci Biobehav Rev, 118, 3-17.
Lazarus, R. S. (1991). Progress on a cognitive-motivational-relational theory of emotion. American psychologist, 46(8), 819.
LeDoux, J. (2012). Rethinking the emotional brain. Neuron, 73(4), 653-676.
Lee, C., Verbeek, E., Doyle, R., & Bateson, M. (2016). Attention bias to threat indicates anxiety differences in sheep. Biol Lett, 12(6).
Lin, I.-R., & Chiao, C.-C. (2017). Visual equivalence and amodal completion in cuttlefish. Frontiers in Physiology, 8, 40.
Loukola, O. J., Solvi, C., Coscos, L., & Chittka, L. (2017). Bumblebees show cognitive flexibility by improving on an observed complex behavior. Science, 355(6327), 833-836.
Mathews, A., & MacLeod, C. (1994). Cognitive approaches to emotion and emotional disorders. Annual review of psychology, 45, 25.
Mendl, M., Brooks, J., Basse, C., Burman, O., Paul, E., Blackwell, E., & Casey, R. (2010). Dogs showing separation-related behaviour exhibit a 'pessimistic' cognitive bias. Curr Biol, 20(19), R839-840.
Mendl, M., Burman, O. H., & Paul, E. S. (2010). An integrative and functional framework for the study of animal emotion and mood. Proc Biol Sci, 277(1696), 2895-2904.
Mogg, K., & Bradley, B. P. (1998). A cognitive-motivational analysis of anxiety. Behaviour research and therapy, 36(9), 809-848.
Nettle, D., & Bateson, M. (2012). The evolutionary origins of mood and its disorders. Curr Biol, 22(17), R712-721.
Oatley, K., & Johnson-Laird, P. N. (1987). Towards a cognitive theory of emotions. Cognition and emotion, 1(1), 29-50.
Olmstead, M. C., & Kuhlmeier, V. A. (2015). Comparative cognition. Cambridge University Press.
Panksepp, J. (1982). Toward a general psychobiological theory of emotions. Behavioral and Brain sciences, 5(3), 407-422.
Panksepp, J. (2011). The basic emotional circuits of mammalian brains: do animals have affective lives? Neuroscience & Biobehavioral Reviews, 35(9), 1791-1804.
Panksepp, J., & Burgdorf, J. (2003). “Laughing” rats and the evolutionary antecedents of human joy? Physiology & behavior, 79(3), 533-547.
Paul, E. S., Harding, E. J., & Mendl, M. (2005). Measuring emotional processes in animals: the utility of a cognitive approach. Neurosci Biobehav Rev, 29(3), 469-491.
Paul, E. S., & Mendl, M. T. (2018). Animal emotion: Descriptive and prescriptive definitions and their implications for a comparative perspective. Appl Anim Behav Sci, 205, 202-209.
Pearce, J. M. (2013). Animal learning and cognition: an introduction. Psychology press.
Perry, C. J., & Baciadonna, L. (2017). Studying emotion in invertebrates: what has been done, what can be measured and what they can provide. J Exp Biol, 220(Pt 21), 3856-3868.
Pinczon du Sel, G., Blanc, A., & Daguzan, J. (2000). The diet of the cuttlefish Sepia officinalis L.(Mollusca: Cephalopoda) during its life cycle in the Northern Bay of Biscay (France). Aquatic Sciences, 62, 167-178.
Rey, S., Huntingford, F. A., Boltana, S., Vargas, R., Knowles, T. G., & Mackenzie, S. (2015). Fish can show emotional fever: stress-induced hyperthermia in zebrafish. Proc Biol Sci, 282(1819).
Roelofs, S., Boleij, H., Nordquist, R. E., & van der Staay, F. J. (2016). Making Decisions under Ambiguity: Judgment Bias Tasks for Assessing Emotional State in Animals. Front Behav Neurosci, 10, 119.
Russell, J. A. (1980). A circumplex model of affect. Journal of personality and social psychology, 39(6), 1161.
Scherer, K. R. (2001). Appraisal Considered as a Process of Multilevel Sequential Checking. Appraisal Processes in Emotion, 92-120.
Schlosberg, H. (1954). Three dimensions of emotion. Psychological Review, 61(2), 81.
Schnell, A. K., Amodio, P., Boeckle, M., & Clayton, N. S. (2021). How intelligent is a cephalopod? Lessons from comparative cognition. Biol Rev Camb Philos Soc, 96(1), 162-178.
Schnell, A. K., Boeckle, M., Rivera, M., Clayton, N. S., & Hanlon, R. T. (2021). Cuttlefish exert self-control in a delay of gratification task. Proc Biol Sci, 288(1946), 20203161.
Schnell, A. K., Clayton, N. S., Hanlon, R. T., & Jozet-Alves, C. (2021). Episodic-like memory is preserved with age in cuttlefish. Proc Biol Sci, 288(1957), 20211052.
Solvi, C., Baciadonna, L., & Chittka, L. (2016). Unexpected rewards induce dopamine-dependent positive emotion-like state changes in bumblebees. Science, 353(6307), 1529-1531.
Tinbergen, N. (1951). The study of instinct. Clarendon Press.
Walters, E. T., Carew, T. J., & Kandel, E. R. (1981). Associative Learning in <i>Aplysia</i>: Evidence for Conditioned Fear in an Invertebrate. Science, 211(4481), 504-506.
Wu, J. J.-S., & Chiao, C.-C. (2023). Switching by cuttlefish of preying tactics targeted at moving prey. iScience, 26(11).
Yang, T. I., & Chiao, C. C. (2016). Number sense and state-dependent valuation in cuttlefish. Proc Biol Sci, 283(1837).
Yasumuro, H., & Ikeda, Y. (2018). Environmental enrichment affects the ontogeny of learning, memory, and depth perception of the pharaoh cuttlefish Sepia pharaonis. Zoology, 128, 27-37.