The Octopus Mind

Sources

Bibliography

104 sources synthesized across this resource, spanning primary research, scholarly reviews, books, policy reports, preprints, and scientific declarations.

89 entries resolve directly through DOI. Unresolved or non-DOI items link to a bibliographic lookup rather than pretending to be primary papers.

  1. journal articleAditi Pophale, Kazumichi Shimizu, Tomoyuki Mano, Leenoy Meshulam, Sam Reiter, et al. (2023). Wake-like skin patterning and neural activity during octopus sleep. NatureDOI 10.1038/s41586-023-06203-4 ↗
  2. journal articleAlbertin CB, Medina-Ruiz S, Mitros T, Schmidbaur H, et al. (Rokhsar, Rosenthal, Simakov) (2022). Genome and transcriptome mechanisms driving cephalopod evolution. Nature CommunicationsDOI 10.1038/s41467-022-29748-w ↗
  3. journal articleAlexandra K. Schnell, Jonathan Birch, et al. (2026). Sentience in cephalopod molluscs: an updated assessment. Biological ReviewsDOI 10.1002/brv.70125 ↗
  4. journal articleAlexandra K. Schnell, Piero Amodio, Markus Boeckle & Nicola S. Clayton (2021). How intelligent is a cephalopod? Lessons from comparative cognition. Biological Reviews 96(1):162–178DOI 10.1111/brv.12651 ↗
  5. journal articleAmodio P, Boeckle M, Schnell AK, Ostojíc L, Fiorito G, Clayton NS (2019). Grow Smart and Die Young: Why Did Cephalopods Evolve Intelligence?. Trends in Ecology & EvolutionDOI 10.1016/j.tree.2018.10.010 ↗
  6. declarationAndrews K, Birch J, Sebo J, et al. (2024). The New York Declaration on Animal Consciousness. NYU (500+ signatories)Find source ↗
  7. journal articleBidel F., Meirovitch Y., Schalek R., ... Lichtman J.W., Hochner B. (2023). Connectomics of the Octopus vulgaris vertical lobe provides insight into conserved and novel principles of a memory acquisition network. eLifeDOI 10.7554/elife.84257 ↗
  8. journal articleBoal JG, Dunham AW, Williams KT, Hanlon RT (2000). Experimental evidence for spatial learning in octopuses (Octopus bimaculoides). Journal of Comparative PsychologyDOI 10.1037/0735-7036.114.3.246 ↗
  9. articleBrian Key (2018). Designing Brains for Pain: Human to Mollusc (and related work). Frontiers in PhysiologyFind source ↗
  10. journal articleBublitz A, Dehnhardt G, Hanke FD (2021). Reversal of a Spatial Discrimination Task in the Common Octopus (Octopus vulgaris). Frontiers in Behavioral NeuroscienceDOI 10.3389/fnbeh.2021.614523 ↗
  11. journal articleBublitz A, Weinhold SR, Strobel S, Dehnhardt G, Hanke FD (2017). Reconsideration of Serial Visual Reversal Learning in Octopus from a Methodological Perspective. Frontiers in PhysiologyDOI 10.3389/fphys.2017.00054 ↗
  12. journal articleBurghardt, G.M. (2005). The Genesis of Animal Play: Testing the Limits. MIT PressDOI 10.1002/ajhb.20433 ↗
  13. journal articleCaroline B. Albertin, Oleg Simakov, ..., Daniel S. Rokhsar, Clifton W. Ragsdale (2015). The octopus genome and the evolution of cephalopod neural and morphological novelties. NatureDOI 10.1038/nature14668 ↗
  14. journal articleChung, W.-S. & Marshall, N. J. (2016 / 2023). Comparative visual ecology of cephalopods / The neural basis of visual processing and behavior in cephalopods. Current Biology / Current BiologyDOI 10.1016/j.cub.2023.08.093 ↗
  15. articleCrawford K, Diaz Quiroz JF, Koenig KM, Ahuja N, Albertin CB, Rosenthal JJC (2020). Highly efficient knockout of a squid pigmentation gene (first CRISPR knockout in a cephalopod). Current BiologyFind source ↗
  16. articleDavid M. Meisel, Ruth A. Byrne, et al. (2011). Contribution of the visual system of the octopus to determination of sleep-like behavior. Journal of Experimental Marine BiologyFind source ↗
  17. journal articleE. R. Brown, S. Piscopo, R. De Stefano, A. Giuditta (2006). Brain and behavioural evidence for rest-activity cycles in Octopus vulgaris. Behavioural Brain ResearchDOI 10.1016/j.bbr.2006.05.009 ↗
  18. journal articleEduardo Sampaio, Catarina S. Ramos, Bruna L. M. Bernardino, et al. (Rui Rosa) (2021). Neurally underdeveloped cuttlefish newborns exhibit social learning. Animal Cognition 24(1):23–32DOI 10.1007/s10071-020-01411-1 ↗
  19. articleFiorito G, Affuso A, Basil J, et al. (CephRes/FELASA/Boyd Group) (2015). Guidelines for the Care and Welfare of Cephalopods in Research. Laboratory AnimalsFind source ↗
  20. journal articleFiorito G, Scotto P (1992). Observational Learning in Octopus vulgaris. ScienceDOI 10.1126/science.256.5056.545 ↗
  21. journal articleGerald B. Biederman & Vaughan A. Davey (1993). Social Learning in Invertebrates. Science 259(5101):1627–1628DOI 10.1126/science.259.5101.1627 ↗
  22. bookGodfrey-Smith P (2016). Other Minds: The Octopus, the Sea, and the Deep Origins of Consciousness. Farrar, Straus and GirouxFind source ↗
  23. journal articleGraziano Fiorito, Gerald B. Biederman, Vaughan A. Davey & Francesca Gherardi (1998). The role of stimulus preexposure in problem solving by Octopus vulgaris. Animal Cognition 1(2):107–112DOI 10.1007/s100710050015 ↗
  24. journal articleGutnick T, Byrne RA, Hochner B, Kuba M (2011). Octopus vulgaris Uses Visual Information to Determine the Location of Its Arm. Current Biology 21(6):460-462DOI 10.1016/j.cub.2011.01.052 ↗
  25. journal articleGutnick T, Neef A, Cherninskyi A, Ziadi-Künzli F, Di Cosmo A, Lipp HP, Kuba MJ (2023). Recording electrical activity from the brain of behaving octopus. Current BiologyDOI 10.1016/j.cub.2023.02.006 ↗
  26. journal articleGutnick T, Zullo L, Hochner B, Kuba MJ (2020). Use of Peripheral Sensory Information for Central Nervous Control of Arm Movement by Octopus vulgaris. Current Biology 30(21):4322-4327DOI 10.1016/j.cub.2020.08.037 ↗
  27. journal articleHanke, F. D. & Kelber, A. (2020). The Eye of the Common Octopus (Octopus vulgaris). Frontiers in PhysiologyDOI 10.3389/fphys.2019.01637 ↗
  28. bookHanlon RT, Messenger JB (2018). Cephalopod Behaviour (2nd ed.) and Cephalopod Camouflage (Nature Scitable). Cambridge Univ. Press / Nature EducationFind source ↗
  29. journal articleHanlon, R.T. (2007). Cephalopod dynamic camouflage. Current Biology 17(11):R400-R404DOI 10.1016/j.cub.2007.03.034 ↗
  30. bookHanlon, R.T. & Messenger, J.B. (1996/2018). Cephalopod Behaviour (1st & 2nd eds.). Cambridge University PressDOI 10.1017/9780511843600 ↗
  31. journal articleHochner B (2012). An Embodied View of Octopus Neurobiology. Current BiologyDOI 10.1016/j.cub.2012.09.001 ↗
  32. journal articleHochner B., Brown E.R., Langella M., Shomrat T., Fiorito G. (2003). A Learning and Memory Area in the Octopus Brain Manifests a Vertebrate-Like Long-Term Potentiation. Journal of NeurophysiologyDOI 10.1152/jn.00645.2003 ↗
  33. journal articleHuang, Y.-H., Lin, H.-J., Lin, L.-Y. & Chiao, C.-C. (2019). Do cuttlefish have fraction number sense?. Animal CognitionDOI 10.1007/s10071-018-01232-3 ↗
  34. journal articleJ.Z. Young (1963). The number and sizes of nerve cells in Octopus. Proceedings of the Zoological Society of LondonDOI 10.1111/j.1469-7998.1963.tb01862.x ↗
  35. journal articleJarmoluk, K. & Pelled, G. (2025). Evidence of play behavior in captive California two-spot octopuses (Octopus bimaculoides). PLOS ONEDOI 10.1371/journal.pone.0326379 ↗
  36. journal articleJean Alupay, Stavros Hadjisolomou, Robyn J. Crook (2013). Arm injury produces long-term behavioral and neural hypersensitivity in octopus. Neuroscience LettersDOI 10.1016/j.neulet.2013.11.002 ↗
  37. journal articleJennifer A. Mather (1994). 'Home' choice and modification by juvenile Octopus vulgaris (Mollusca: Cephalopoda): specialized intelligence and tool use?. Journal of Zoology (London) 233:359–368DOI 10.1111/j.1469-7998.1994.tb05270.x ↗
  38. journal articleJonas N. Richter, Binyamin Hochner, Michael J. Kuba (2016). Pull or Push? Octopuses Solve a Puzzle Problem. PLOS ONE 11(3):e0152048DOI 10.1371/journal.pone.0152048 ↗
  39. reportJonathan Birch, Charlotte Burn, Alexandra Schnell, Heather Browning, Andrew Crump (2021). Review of the Evidence of Sentience in Cephalopod Molluscs and Decapod Crustaceans (LSE report for DEFRA). London School of Economics / DEFRAFind source ↗
  40. journal articleJoshua J. C. Rosenthal, Eli Eisenberg (2023). Extensive Recoding of the Neural Proteome in Cephalopods by RNA Editing. Annual Review of Animal BiosciencesDOI 10.1146/annurev-animal-060322-114534 ↗
  41. journal articleJozet-Alves C, Bertin M, Clayton NS (2013). Evidence of episodic-like memory in cuttlefish. Current BiologyDOI 10.1016/j.cub.2013.10.021 ↗
  42. journal articleJulian K. Finn, Tom Tregenza, Mark D. Norman (2009). Defensive tool use in a coconut-carrying octopus. Current Biology 19(23):R1069–R1070DOI 10.1016/j.cub.2009.10.052 ↗
  43. articleKang G, Allard CAH, et al. (Bellono & Hibbs labs) (2023). Structural basis of sensory receptor evolution in octopus / Structural basis for the evolution of cephalopod chemotactile receptors. NatureFind source ↗
  44. journal articleKang R. et al. (2024). Neuronal segmentation in cephalopod arms. Nature CommunicationsDOI 10.1038/s41467-024-55475-5 ↗
  45. journal articleKawashima, S. & Ikeda, Y. (2025). Cross-Modal Object Recognition and Reliability Between Visual and Tactile Senses in Octopus (Callistoctopus aspilosomatis). Zoological Science, 42(3):260–269DOI 10.2108/zs240069 ↗
  46. journal articleKuan-Ling Huang & Chuan-Chin Chiao (2013). Can cuttlefish learn by observing others?. Animal Cognition 16(3):481–490DOI 10.1007/s10071-012-0573-z ↗
  47. journal articleKuba, M.J., Byrne, R.A., Meisel, D.V. & Mather, J.A. (2006). When do octopuses play? Effects of repeated testing, object type, age, and food deprivation on object play in Octopus vulgaris. Journal of Comparative Psychology 120(3):184–190DOI 10.1037/0735-7036.120.3.184 ↗
  48. journal articleKuba, M.J., Byrne, R.A., Meisel, D.V. & Mather, J.A. (2006). Exploration and habituation in intact free-moving Octopus vulgaris. International Journal of Comparative Psychology 19(4)DOI 10.46867/ijcp.2006.19.04.02 ↗
  49. journal articleKuuspalu A, Cody S, Hale ME (2022). Multiple nerve cords connect the arms of octopuses, providing alternative paths for inter-arm signaling. Current Biology 32(24):5415-5421DOI 10.1016/j.cub.2022.11.007 ↗
  50. journal articleLaan, A., Gutnick, T., Kuba, M.J. & Laurent, G. (2014). Behavioral analysis of cuttlefish traveling waves and its implications for neural control. Current Biology 24(15):1737-1742DOI 10.1016/j.cub.2014.06.027 ↗
  51. journal articleLangridge, K.V., Broom, M. & Osorio, D. (2007). Selective signalling by cuttlefish to predators (deimatic displays). Current Biology 17(24):R1044-R1045DOI 10.1016/j.cub.2007.10.028 ↗
  52. journal articleLevy G, Flash T, Hochner B (2015). Arm Coordination in Octopus Crawling Involves Unique Motor Control Strategies. Current Biology 25(9):1195-1200DOI 10.1016/j.cub.2015.02.064 ↗
  53. declarationLow P, Panksepp J, Edelman D, Koch C, et al. (2012). The Cambridge Declaration on Consciousness. Francis Crick Memorial ConferenceFind source ↗
  54. journal articleMarcos G. Frank, R. J. Waldrop, M. Dumoulin, et al. (2012). A preliminary analysis of sleep-like states in the cuttlefish Sepia officinalis. PLOS ONEDOI 10.1371/journal.pone.0038125 ↗
  55. journal articleMaselli, V., Al-Soudy, A.-S., Buglione, M., et al. (2022). A preliminary attempt to investigate mirror self-recognition in Octopus vulgaris. Frontiers in Physiology, 13:951808DOI 10.3389/fphys.2022.951808 ↗
  56. journal articleMather, J.A. (2004). Apparent movement in a visual display: the 'passing cloud' of Octopus cyanea. Journal of Zoology 263(1):89-94DOI 10.1017/s0952836904004911 ↗
  57. journal articleMather, J.A. & Anderson, R.C. (1993). Personalities of octopuses (Octopus rubescens). Journal of Comparative Psychology 107(3):336–340DOI 10.1037/0735-7036.107.3.336 ↗
  58. journal articleMather, J.A. & Anderson, R.C. (1999). Exploration, play, and habituation in octopuses (Octopus dofleini). Journal of Comparative Psychology 113(3):333–338DOI 10.1037/0735-7036.113.3.333 ↗
  59. journal articleMatthew A. Birk, Noa Liscovitch-Brauer, ..., Eli Eisenberg, Joshua J. C. Rosenthal (2023). Temperature-dependent RNA editing in octopus extensively recodes the neural proteome. CellDOI 10.1016/j.cell.2023.05.004 ↗
  60. posted contentMBL Cephalopod Program (Rosenthal lab and colleagues) (2023). Creation of an albino squid line by CRISPR-Cas9 and its application for in vivo functional imaging of neural activity. Current BiologyDOI 10.2139/ssrn.4369821 ↗
  61. journal articleMessenger, J.B. (2001). Cephalopod chromatophores: neurobiology and natural history. Biological Reviews 76(4):473-528DOI 10.1017/s1464793101005772 ↗
  62. journal articleNoa Liscovitch-Brauer, Shahar Alon, Hagit T. Porath, ..., Eli Eisenberg, Joshua J. C. Rosenthal (2017). Trade-off between Transcriptome Plasticity and Genome Evolution in Cephalopods. CellDOI 10.1016/j.cell.2017.03.025 ↗
  63. journal articleOgura, A., Ikeo, K. & Gojobori, T. (2004). Comparative Analysis of Gene Expression for Convergent Evolution of Camera Eye Between Octopus and Human. Genome ResearchDOI 10.1101/gr.2268104 ↗
  64. journal articlePackard, A. & Sanders, G.D. (1971). Body patterns of Octopus vulgaris and maturation of the response to disturbance. Animal Behaviour 19(4):780-790DOI 10.1016/s0003-3472(71)80181-1 ↗
  65. journal articlePeter Godfrey-Smith, David Scheel, Stephanie Chancellor, Stefan Linquist, Matthew Lawrence (2022). In the line of fire: Debris throwing by wild octopuses. PLOS ONE 17(11):e0276482DOI 10.1371/journal.pone.0276482 ↗
  66. journal articlePetrosino G, Ponte G, Volpe M, et al. (Sanges R, Fiorito G) (2022). Identification of LINE retrotransposons and long non-coding RNAs expressed in the octopus brain. BMC BiologyDOI 10.1186/s12915-022-01303-5 ↗
  67. articlePiero Amodio et al. (2019). Octopus intelligence: the importance of being agnostic (commentary on Mather). Animal SentienceFind source ↗
  68. journal articleRamirez MD, Oakley TH (2015). Eye-independent, light-activated chromatophore expansion (LACE) and expression of phototransduction genes in the skin of Octopus bimaculoides. Journal of Experimental Biology 218:1513–1520DOI 10.1242/jeb.110908 ↗
  69. journal articleRobyn J. Crook (2021). Behavioral and neurophysiological evidence suggests affective pain experience in octopus. iScience (Cell Press)DOI 10.1016/j.isci.2021.102229 ↗
  70. journal articleRobyn J. Crook, Kellie Dickson, Roger T. Hanlon, Edgar T. Walters (2014). Nociceptive sensitization reduces predation risk. Current BiologyDOI 10.1016/j.cub.2014.03.043 ↗
  71. journal articleRobyn J. Crook, Roger T. Hanlon, Edgar T. Walters (2013). Squid have nociceptors that display widespread long-term sensitization and spontaneous activity after bodily injury. Journal of NeuroscienceDOI 10.1523/jneurosci.0646-13.2013 ↗
  72. reportRoland C. Anderson, Jennifer A. Mather (popularized) (2010s). Jar/childproof-bottle opening in giant Pacific octopus (Seattle Aquarium). Seattle Aquarium / popular science reportingFind source ↗
  73. journal articleSampaio, E., et al. (2024). Multidimensional social influence drives leadership and composition-dependent success in octopus-fish hunting groups. Nature Ecology & EvolutionDOI 10.1038/s41559-024-02525-2 ↗
  74. journal articleSampaio, E., Seco, M.C., Rosa, R., Gingins, S. (2021). Octopuses punch fishes during collaborative interspecific hunting events. EcologyDOI 10.1002/ecy.3266 ↗
  75. journal articleSanders GD (1970). Long-term memory of a tactile discrimination in Octopus vulgaris and the effect of vertical lobe removal. Brain ResearchDOI 10.1016/0006-8993(70)90154-x ↗
  76. journal articleSandra Garrett, Joshua J. C. Rosenthal (2012). RNA Editing Underlies Temperature Adaptation in K+ Channels from Polar Octopuses. ScienceDOI 10.1126/science.1212795 ↗
  77. journal articleScheel, D., Chancellor, S., Hing, M., Lawrence, M., Linquist, S., Godfrey-Smith, P. (2017). A second site occupied by Octopus tetricus at high densities, with notes on their ecology and behavior (Octlantis). Marine and Freshwater Behaviour and PhysiologyDOI 10.1080/10236244.2017.1369851 ↗
  78. journal articleScheel, D., Godfrey-Smith, P., Lawrence, M. (2016). Signal Use by Octopuses in Agonistic Interactions. Current BiologyDOI 10.1016/j.cub.2015.12.033 ↗
  79. journal articleScheel, D., Godfrey-Smith, P., Linquist, S., Chancellor, S., Hing, M., Lawrence, M. (2018). Octopus engineering, intentional and inadvertent. Communicative & Integrative BiologyDOI 10.1080/19420889.2017.1395994 ↗
  80. journal articleSchnell AK, Boeckle M, Rivera M, Clayton NS, Hanlon RT (2021). Cuttlefish exert self-control in a delay of gratification task. Proceedings of the Royal Society BDOI 10.1098/rspb.2020.3161 ↗
  81. journal articleShahar Alon, Sara C. Garrett, ..., Eli Eisenberg, Joshua J. C. Rosenthal (2015). The majority of transcripts in the squid nervous system are extensively recoded by A-to-I RNA editing. eLifeDOI 10.7554/elife.05198 ↗
  82. journal articleShashar, N., Rutledge, P. S. & Cronin, T. W. (1996). Polarization Vision in Cuttlefish – A Concealed Communication Channel?. Journal of Experimental BiologyDOI 10.1242/jeb.199.9.2077 ↗
  83. journal articleShomrat T., Zarrella I., Fiorito G., Hochner B. (2008). The Octopus Vertical Lobe Modulates Short-Term Learning Rate and Uses LTP to Acquire Long-Term Memory. Current BiologyDOI 10.1016/j.cub.2008.01.056 ↗
  84. journal articleSinn, D.L. & Moltschaniwskyj, N.A. (2005). Personality traits in dumpling squid (Euprymna tasmanica): context-specific traits and their correlation with biological characteristics. Journal of Comparative Psychology 119(1):99–110DOI 10.1037/0735-7036.119.1.99 ↗
  85. journal articleSinn, D.L., Perrin, N.A., Mather, J.A. & Anderson, R.C. (2001). Early temperamental traits in an octopus (Octopus bimaculoides). Journal of Comparative Psychology 115(4):351–364DOI 10.1037//0735-7036.115.4.351-364 ↗
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  89. journal articleSumbre G, Fiorito G, Flash T, Hochner B (2006). Octopuses Use a Human-like Strategy to Control Precise Point-to-Point Arm Movements. Current BiologyDOI 10.1016/j.cub.2006.02.069 ↗
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  95. posted contentTurchetti-Maia A.L., Stern-Mentch N., Bidel F., Nesher N., Shomrat T., Hochner B. (2018/2024). A novel NO-dependent 'molecular-memory-switch' mediates presynaptic expression and postsynaptic maintenance of LTP in the octopus brain. bioRxivDOI 10.1101/491340 ↗
  96. preprintTurchetti-Maia AL, Stern-Mentch N, Hochner B, et al. (2024 (bioRxiv)). A novel nitric oxide (NO)-dependent molecular switch mediating LTP in the Octopus vulgaris brain. bioRxiv preprintFind source ↗
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