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Learning and the Evolution of Conscious Agents

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Abstract

The scientific study of consciousness or subjective experiencing is a rapidly expanding research program engaging philosophers of mind, psychologists, cognitive scientists, neurobiologists, evolutionary biologists and biosemioticians. Here we outline an evolutionary approach that we have developed over the last two decades, focusing on the evolutionary transition from non-conscious to minimally conscious, subjectively experiencing organisms. We propose that the evolution of subjective experiencing was driven by the evolution of learning and we identify an open-ended, representational, generative and recursive form of associative learning, which we call Unlimited Associative Learning (UAL), as an evolutionary transition marker of minimal consciousness. This evolutionary marker provides evidence that the evolutionary transition to consciousness has gone to completion and allows reverse-engineering from this learning capacity to the system that enables it – making possible the construction of a toy model of UAL. The model allows us to identify some of the key processes and structures that constitute minimal consciousness, points its taxonomic distribution and the ecological context in which it first emerged, highlights its function and suggests a framework for exploring developmental and evolutionary modifications of consciousness. We point to ways of experimentally testing the relationship between UAL and consciousness in human and in non-human animals and discuss the theoretical and ethical implications of our approach. The framework we offer allows the exploration of the evolutionary changes in agency, value systems, selective processes and goals that were involved in the transition to subjective experiencing from a perspective that resonates with the approaches of bio-semioticians.

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Notes

  1. We use the term teleology and teleological as general terms covering all goal-directed behavior (GDB). These include behaviors that do not depend on conscious will or preconceived design (teleonomic GDB) as well as GDB that is driven by mental intention, desires or reflectively and rationally guided goal-directed behavior. The term “mode of being” is used in this article within the Aristotelian teleological framework. For a discussion of an ecological notion of a mode of being (e.g. terrestrial aquatic, aerial) see Ginsburg & Jablonka 2020a.

  2. We are aware that our evolutionary approach is not universally shared – there are panpsychists who believe that all matter is conscious, dualists who separate mind and body, and biopsychists according to whom living entails sentience, so all living organisms are considered sentient. The discussion of these different approaches, their merits and problems, is beyond the scope of this article. As this article makes clear we are neither panpsychists nor biopsychists.

  3. The term “semiotic selection” suggested by Maran and Kleisner (2010), is a broader concept than the intentional selection concept of Noble, which explicitly assumes consciousness. The qualification and elaboration of the concept of selection within the extended evolutionary synthesis (EES) is discussed by Jablonka and Lamb (2014); Jablonka (2021), and Kull (2021) discuss the term in their commentaries on Noble 2021 target article.

  4. The two issues of The Philosophical Transactions of the Royal Society 2020 and 2021 https://royalsocietypublishing.org/toc/rstb/2021/376/1821 discuss basal cognition at depth and from multiple perspectives; we have discussed major transitions in cognition from the learning perspective in Ginsburg & Jablonka, 2021.

References

  • Adams, C. D., & Dickinson, A. (1981). Instrumental responding following reinforcer evaluation. The Quarterly Journal of Experimental Psychology, 33B(2), 109–121. https://doi.org/10.1080/14640748108400816

    Article  Google Scholar 

  • Ahissar, E., & Assa, E. (2016). A closed loop hypothesis of perception. eLife 5, e12830. https://doi.org/10.7554/eLife.12830xxx

  • Aristotle. (1984a). Generation of animals (J, Barnes, Ed., W. D. Ross, Trans.). In The complete works of Aristotle. The revised Oxford translation, vol. 2. Princeton University Press.

  • Aristotle. (1984b). Metaphysics, Book 1. (J, Barnes, Ed., W. D. Ross Trans.) In The complete works of Aristotle. The revised Oxford translation, vol. 2. Princeton University Press.

  • Aristotle. (1984c). On the soul (J. Barnes, Ed., J. A. Smith, Trans.). The Complete Works of Aristotle. The revised Oxford translation, vol. 1. Princeton University Press.

  • Baars, B. J. (2005a). Global workspace theory of consciousness: Toward a cognitive neuroscience of human experience. Progress in Brain Research, 150, 45–53. https://doi.org/10.1016/S0079-6123(05)50004-9

    Article  Google Scholar 

  • Baars, B. J. (2005b). Subjective experience is probably not limited to humans: The evidence from neurobiology and behavior. Consciousness and Cognition, 14, 7–21. https://doi.org/10.1016/j.concog.2004.11.002

    Article  Google Scholar 

  • Baddeley, A. D. (1986). Working memory. Clarendon Press.

    Google Scholar 

  • Balleine, B. W., & Dickinson, A. (1998a). Consciousness – the interface between affect and cognition. In J. Cornwell (Ed.), Consciousness and human identity (pp. 57–85). Oxford University Press.

    Google Scholar 

  • Balleine, B. W. & Dickinson, A. (1998b). Goal-directed instrumental action: contingency and incentive learning and their cortical substrates. Neuropharmacology 37, 407–419. https://doi.org/10.1016/S0028-3908(98)00033-1

  • Bangasser, D. A., Waxler, D. E., Santollo, J., & Shors, T. J. (2006). Trace conditioning and the hippocampus: The importance of contiguity. Journal of Neuroscience, 26, 8702–8706. https://doi.org/10.1523/JNEUROSCI.1742-06.2006

    Article  CAS  Google Scholar 

  • Barron, A. B., & Klein, C. (2016). What insects can tell us about the origins of consciousness. Proceedings of the Natural Academy of Science USA, 113, 4900–4908. https://doi.org/10.1073/pnas.1520084113

  • Bateson, P. (2006). The adaptability driver: Links between behavior and evolution. Biological Theory 1(4), 342–345. https://doi.org/10.1162/biot.2006.1.4.342

  • Bateson, P., & Laland, K. N. (2013). Tinbergen’s four questions: An appreciation and an update. Trends in Ecology and Evolution, 28(12), 712–718. https://doi.org/10.1016/j.tree.2013.09.013

  • Benard, J., Stach, S., & Giurfa, M. (2006). Categorization of visual stimuli in the honeybee Apis mellifera. Animal Cognition, 9(4), 257–270. https://doi.org/10.1007/s10071-006-0032-9

    Article  Google Scholar 

  • Ben-Haim, M. S., Dal Monte, O., Fagan, N. A., Dunham, Y. & Hassin, R. R. (2021). Disentangling perceptual awareness from nonconscious processing in rhesus monkeys (Macaca mulatta), Proceedings of the National Academy of Sciences, 118, 15. https://doi.org/10.1073/pnas.2017543118

  • Birch, J., Ginsburg, S., & Jablonka, E. (2020a). Unlimited associative learning and the origins of consciousness: A primer and some predictions. Biology and Philosophy, 35, 56. https://doi.org/10.1007/s10539-020-09772-0

  • Birch, J., Schnell, A.K. & Clayton, N.S. (2020b) Dimensions of animal consciousness. Trends in Cognitive Sciences, 24(10), 789–801. https://doi.org/10.1016/j.tics.2020.07.007

  • Birch, J., Ginsburg, S., & Jablonka, E. (2021a). The learning-consciousness connection. Biology and Philosophy, 36, 49. https://doi.org/10.1007/s10539-021-09802-5

    Article  Google Scholar 

  • Birch, J., Burn, C., Schnell, A., Heather Browning, H. & Andrew Crump A. (2021b). Review of the evidence of sentience in cephalopod molluscs and decapod crustaceans. LSE consulting. https://www.lse.ac.uk/news/news-assets/pdfs/2021b/sentience-in-cephalopod-molluscs-and-decapod-crustaceans-final-report-november-2021.pdf

  • Bray, D (2009). Wetware: A computer in every living cell. Yale University Press

  • Brembs, B. (2011). Towards a scientific concept of free will as a biological trait: spontaneous actions and decision-making in invertebrates. Proceedings of the Royal Society B, 278, 930–939. https://doi.org/10.1098/rspb.2010.2325

  • Bronfman, Z., Ginsburg, S., & Jablonka, E. (2016a). The evolutionary origins of consciousness: Suggesting a transition marker. Journal of Consciousness Studies, 23(9/10), 7–34.

    Google Scholar 

  • Bronfman, Z. Ginsburg, S., & Jablonka, E. (2016b). The transition to minimal consciousness through the evolution of associative learning. Frontiers in Psychology 7, 1954. https://doi.org/10.3389/fpsyg.2016.01954

  • Bronfman, Z. Z., Ginsburg, S., & Jablonka, E. (2021). When will robots be sentient? Journal of Artificial Intelligence and Consciousness, 8(2), 1–21. https://doi.org/10.1142/S2705078521500168

    Article  Google Scholar 

  • Buckner, R. L., & DiNicola, L. M. (2019). The brain’s default network: Updated anatomy, physiology and evolving insights. Nature Reviews Neuroscience, 20, 593–608. https://doi.org/10.1038/s41583-019-0212-7

    Article  CAS  Google Scholar 

  • Clark, R. E. & Squire, L. R. (1998). Classical conditioning and brain systems: the role of awareness. Science, 280, 77–81. https://doi.org/10.1126/science.280.5360.77

  • Clark, R. E., & Squire, L. R. (1999). Human eyeblink classical conditioning: Effects of manipulating awareness of the stimulus contingencies. Psychological Science, 10, 14–18. https://doi.org/10.1111/1467-9280.00099

    Article  Google Scholar 

  • Couvillon, P. A., & Bitterman, M. E. (1988). Compound-component and conditional discrimination of colors and odors by honeybees: Further tests of a continuity model. Animal Learning and Behavior, 16, 67–74. https://doi.org/10.3758/BF03209045

    Article  Google Scholar 

  • Crane, S. (1894). In the depth of a coal mine. McClure’s Magazine III(3). https://ehistory.osu.edu/exhibitions/gildedage/content/CraneDepths

  • Damasio, A. (2021). Feeling and knowing: Making minds conscious. Pantheon.

  • Darwin, C. (1871). The descent of man, and selection in relation to sex (1st ed.). John Murray.

    Book  Google Scholar 

  • Darwin, C. (1872). The expression of emotions in man and animals. John Murray.

    Book  Google Scholar 

  • Dehaene, S. (2014). Consciousness and the brain: Deciphering how the brain codes our thoughts. Penguin Books.

    Google Scholar 

  • Dehaene, S., & Changeux, J.-P. (2011). Experimental and theoretical approaches to conscious processing. Neuron, 70, 201–227. https://doi.org/10.1016/j.neuron.2011.03.018

    Article  CAS  Google Scholar 

  • Dennett, D. C. (1995). Darwin’s dangerous idea. Simon and Schuster.

    Google Scholar 

  • Dickinson, A. (2012a). Associative learning and animal cognition. Philosophical Transactions of the Royal Society of London B, 367, 2733–2742. https://doi.org/10.1098/rstb.2012.0220

  • Dickinson, A. (2012b). Why a rat is not a beast machine. Frontiers of Consciousness. Chichele Lectures. https://doi.org/10.1093/acprof:oso/9780199233151.003.0010

  • Dickinson, S., & Balleine, B. (1994). Motivational control of goal-directed action. Animal Learning and Behavior, 22(1), 1–18. https://doi.org/10.3758/BF03199951

    Article  Google Scholar 

  • Dickinson, A., & Balleine, B. W. (2000). Causal cognition and goal-directed action. In C. Heyes & L. Huber (Eds.), Vienna series in theoretical biology: The evolution of cognition (pp. 185–204). MIT Press.

    Google Scholar 

  • Dickinson, A., & Balleine, B. (2010). Hedonics – the cognitive-motivational interface. In M. L. Kringelbach & A. C. Berridge (Eds.), Pleasures of the brain (pp. 74–84). Oxford University Press.

    Google Scholar 

  • Dickinson, A., Watt, A., & Griffiths, W. J. H. (1992). Free-operant acquisition with delayed reinforcement. The Quarterly Journal of Experimental Psychology Section B, 45(3), 241–258. https://doi.org/10.1080/14640749208401019

    Article  Google Scholar 

  • Donald, M. (1991). Origins of the modern mind: Three stages in the evolution of culture and cognition. Harvard University Press.

  • Dor, D. (2015). The instruction of imagination: Language as a social communication technology. Oxford University Press.

  • Dor, D. (2022). Communication for collaborative computation: Two major transitions in human evolution. Philosophical Transaction of the Royal Society B, (in press).

  • Dor, D., & Jablonka, E. (2010). Canalization and plasticity in the evolution of linguistic communication. In R. K. Larson, V. Deprez, & H. Yamakido (Eds.), The evolution of human language (pp. 135–147). Cambridge University Press.

    Chapter  Google Scholar 

  • Feinberg,, T.E. & Mallatt, J. (2016). The ancient origins of consciousness: How the brain created experience. MIT Press.

  • Freeman, W. T. (2003). Neurodynamic models of brain in Psychiatry. Neuropsychopharmacology, 28, S54–S63. https://doi.org/10.1038/sj.npp.1300147

    Article  Google Scholar 

  • Fresco, N., Ginsburg, S., & Jablonka, E. (2020). Functional information: A graded taxonomy of difference makers. Review of Philosophy and Psychology, 11, 547–567. https://doi.org/10.1007/s13164-018-0410-7

    Article  Google Scholar 

  • Gánti, T. (1987). The principle of life. (L. Vekerd, Trans.). Omikk.

  • Gánti, T. (2003). The principles of life, with a commentary by James Griesemer and Eörs Szathmáry. Oxford University Press.

    Google Scholar 

  • Gewirtz, J. C., & Davis, M. (2000). Using Pavlovian higher-order conditioning paradigms to investigate the neural substrates of emotional learning and memory. Learning and Mem, 7, 257–266. https://doi.org/10.1101/lm.35200

    Article  CAS  Google Scholar 

  • Ginsburg, S. & Jablonka, E. (2010). The evolution of associative learning: A factor in the Cambrian explosion. Journal of Theoretical Biology 266, 11–20. https://doi.org/10.1016/j.jtbi.2010.06.017

  • Ginsburg, S., & Jablonka, E. (2015). The teleological transitions in evolution: A Gántian view. Journal of Theoretical Biology, 381, 55–60. https://doi.org/10.1016/j.jtbi.2015.04.007

  • Ginsburg, S., & Jablonka, E. (2019). The evolution of the sensitive soul: Learning and the Origins of Consciousness. MIT Press.

  • Ginsburg, S., & Jablonka, E. (2020a). Consciousness as a mode of being. Journal of Consciousness Studies, 27(9–10), 148–162.

    Google Scholar 

  • Ginsburg, S., & Jablonka, E. (2020b). Sentience in plants: A green red herring? Journal of Consciousness Studies, 28(1–2), 17–34.

    Google Scholar 

  • Ginsburg, S., & Jablonka, E. (2021). Evolutionary transitions in learning and cognition. Philosophical Transactions of the Royal Society B, 376, 20190766. https://doi.org/10.1098/rstb.2019.0766

    Article  Google Scholar 

  • Ginsburg, S., & Jablonka, E. (2022). Picturing the mind: Consciousness through the lens of evolution. MIT Press.

    Google Scholar 

  • Godfrey-Smith, P. (2020). Metazoa: Animal life and the birth of the mind. Collins

  • Hadar, R., & Menzel, R. (2010). Memory formation in reversal learning of the honeybee. Frontiers in Behavioral Neuroscience, 4, 186. https://doi.org/10.3389/fnbeh.2010.00186

    Article  Google Scholar 

  • Hardy, A. (1965). The living stream: A restatement of evolution theory and Its relation to the spirit of man. Collins.

  • Hendlin, Y. H. & Sharov, A. (2022). Biosemiotics (Journal home page). Retrieved August 14, 2022. https://www.springer.com/journal/12304

  • Holland, P. C., & Rescorla, R. A. (1975). Second-order conditioning with food unconditioned stimulus. Journal of Comparative and Physiological Psychology, 88, 459–467. https://doi.org/10.1037/h0076219

    Article  CAS  Google Scholar 

  • Humphrey, N. (2011), Soul dust: The magic of consciousness. Princeton University Press.

  • Hussaini, S. A., Komischke, B., Menzel, R., & Lachnit, H. (2007). Forward and backward second-order Pavlovian conditioning in honeybees. Learning and Memory, 14, 678–683. https://doi.org/10.1101/lm.471307

    Article  Google Scholar 

  • Jablonka, E. (2002). Information: its interpretation, its inheritance and its sharing. Philosophy of Science, 69, 578–605. https://doi.org/10.1086/344621

  • Jablonka, E, (2021). Signs of consciousness? Biosemiotics 14(1), 25–29. https://doi.org/10.1007/s12304-021-09419-x

  • Jablonka, E., & Ginsburg, S. (2022). From teleonomy to mentally-driven goal-directed behavior: Evolutionary considerations. MIT Press (in press). 

  • Jablonka, E., & Lamb, M. L. (2014). Evolution in four dimensions. Genetics, epigenetic, behavioral and symbolic variation in the history of life (2nd ed.). MIT Press.

  • James, W. (1890). The principles of psychology. Dover.

    Google Scholar 

  • Jonas, H. (1966). The phenomenon of life: Toward a philosophical biology. Harper and Row.

    Google Scholar 

  • Keijzer, F., Van Duijn, M., & Lyon, P. (2013). What nervous systems do: early evolution, input–output, and the skin brain thesis. Adaptive Behavior 21(2), 67–85. https://doi.org/10.1177/1059712312465330

  • Kröger, B., Vinther, J., & Fuchs, D. (2011). Cephalopod origin and evolution: A congruent picture emerging from fossils, development and molecules. BioEssays, 33, 602–613. https://doi.org/10.1002/bies.201100001

    Article  CAS  Google Scholar 

  • Kull, K. (2021). Natural selection and self-organization do not make meaning, while the agent’s choice does. Biosemiotics, 14, 49–53. https://doi.org/10.1007/s12304-021-09422-2

    Article  Google Scholar 

  • Lamme, V. (2020). Visual functions generating conscious seeing. Frontiers in Psychology, 11, 83. https://doi.org/10.3389/fpsyg.2020.00083

    Article  Google Scholar 

  • Longo, G., Montévil, M., Sonnenschein, C. & Soto, A. M. (2015). In search of principles for a theory of organisms. Journal of Biosciences, 40(5), 955–968. https://doi.org/10.1007/s12038-015-9574-9

  • Lucas, G. A., Deich, J. D., & Wasserman, E. A. (1981). Trace autoshaping: acquisition, maintenance, and path dependence at long trace intervals. Journal of the Experimental Analysis of Behavior 36, 61–74. https://doi.org/10.1901/jeab.1981.36-61

  • Man, K., & Damasio, A. (2019). Homeostasis and soft robotics in the design of feeling machines. Nature Machine Intelligence 1, 446–452. https://doi.org/10.1038/s42256-019-0103-7

  • Mansur, B. E., Rodrigues, J. R. V., & Mota, T. (2018). Bimodal patterning discrimination in harnessed honey bees. Frontiers in Psychology, 9, 1529. https://doi.org/10.3389/fpsyg.2018.01529

    Article  Google Scholar 

  • Maran, T., & Kleisner, K. (2010). Towards an evolutionary biosemiotics: Semiotic selection and semiotic co-option. Biosemiotics, 3, 189–200. https://doi.org/10.1007/s12304-010-9087-8

    Article  Google Scholar 

  • Mashour, G. A., Roelfsema, P, Changeux, J.-P., & Dehaene, S. (2020). Conscious processing and the global neuronal workspace hypothesis. Neuron, 105(5), 776–798. https://doi.org/10.1016/j.neuron.2020.01.026

  • Maynard Smith, J., & Szathmáry, E. (1995). The major transitions in evolution. Oxford University Press.

    Google Scholar 

  • McGilchrist, I. (2010). The master and his emissary: The divided brain and the making of the Western world. Yale University Press.

    Google Scholar 

  • McMillan, F. D. (2020). The mental health and well-being benefits of personal control in animals. In F. D. McMillan (Ed.), Mental health and well-being in animals, 2nd edition. CABI.

  • Merker, B. (2005). The liabilities of mobility: a selection pressure for the transition to consciousness in animal evolution. Consciousness and Cognition 14, 89–114. https://doi.org/10.1016/S1053-8100(03)00002-3

  • Merker, B. (2007). Consciousness without a cerebral cortex: A challenge for neuroscience and medicine. Behavioral and Brain Sciences 30, 63–134. https://doi.org/10.1017/S0140525X07000891

  • Metzinger, T. (2003). Being no one: The self-model theory of subjectivity. MIT Press.

    Book  Google Scholar 

  • Mizunami, M. (2021). What Is learned in Pavlovian conditioning in crickets? Revisiting the S-S and S-R learning theories. Frontiers in Behavioral Neuroscience, 15, 661225. https://doi.org/10.3389/fnbeh.2021.661225

    Article  CAS  Google Scholar 

  • Morsella, E. (2005). The function of phenomenal states: Supramodular interaction theory. Psychological Review, 112, 1000–1021. https://doi.org/10.1037/0033-295X.112.4.1000

    Article  Google Scholar 

  • Moyer, J. R., Deyo, R. A., & Disterhoft, J. F. (2015). Hippocampectomy disrupts trace eye blink conditioning in rabbits. Behavioral Neuroscience, 129, 523–532. https://doi.org/10.1037/bne0000079

    Article  Google Scholar 

  • Nesse, R. M. (2001). The smoke detector principle. Annals of the New York Academy of Science 935, 75‒85. https://doi.org/10.1111/j.1749-6632.2001.tb03472.x

  • Nicholson, D. (2013). Organisms ≠ machines. Studies in History and Philosophy of Science Part C: Studies in History and Philosophy of Biological and Biomedical Sciences 44, 669–678. https://doi.org/10.1016/j.shpsc.2013.05.014

  • Noble, D. (2021). The illusions of the modern synthesis. Biosemiotics, 14, 5–24. https://doi.org/10.1007/s12304-021-09405-3

    Article  Google Scholar 

  • Öhman, A. & Soares, J. J. (1993). On the automatic nature of phobic fear: conditioned electrodermal responses to masked fear-relevant stimuli. Journal of Abnormal Psychology 102, 121–132. https://doi.org/10.1037/0021-843X.102.1.121

  • Okasha, S. (2018). Agents and goals in evolution. Oxford University Press.

    Book  Google Scholar 

  • Pagni, E., & Simanke, R. T. (2021). Biosemiotics and evolution: The natural foundations of meaning and symbolism. Springer.

  • Panksepp, J. (2005). Affective consciousness: Core emotional feelings in animals and humans. Consciousness & Cognition, 14, 30–80.

    Article  Google Scholar 

  • Panksepp, J. (2011). Cross-species affective neuroscience decoding of the primal affective experiences of humans and related animals. PLoS ONE 6(9), e21236. https://doi.org/10.1371/journal.pone.0021236

  • Petersen, S. E., & Posner, M. I. (2012). The attention system of the human brain: 20 years after. Annual Review of Neuroscience, 35, 73–89. https://doi.org/10.1146/annurev-neuro-062111-150525

    Article  CAS  Google Scholar 

  • Popper, K. (1972) Objective knowledge: An evolutionary approach. Clarendon Press.

  • Prum, R. O. (2017). The evolution of beauty: How Darwin’s forgotten theory of mate choice shapes the animal world and us. Anchor Books.

  • Raichle, M. E., MacLeod, A. M., Snyder, A. Z., Powers, W. J., Gusnard, D. A., & Shulman, G. L. (2001). Inaugural article: A default mode of brain function. Proceedings of the National Academy of Sciences. 98(2), 676–682. https://doi.org/10.1073/pnas.98.2.676

  • Reber, A. S. (2019). The first minds: Caterpillars, 'karyotes and consciousness. Oxford University Press.

    Google Scholar 

  • Riskin, J. (2016) The restless clock: A history of the centuries-long argument over what makes living things tick. University of Chicago Press.

  • Rodríguez-Expósito, B., Gómez, A., Martín-Monzón, I., Reiriz, M., Rodríguez, F., & Salas, C. (2017). Goldfish hippocampal pallium is essential to associate temporally discontiguous events. Neurobiology of Learning and Memory, 139, 128–134. https://doi.org/10.1016/j.nlm.2017.01.002

    Article  Google Scholar 

  • Schon, K. R., Parker, A. P. J., & Woods, C. G (2018, updated 2020). Congenital insensitivity to pain overview. In M. P. Adam., G. M. Mirzaa, R. A. Pagon, et al., (Eds.). GeneReviews®. University of Washington, Seattle. https://www.ncbi.nlm.nih.gov/books/NBK481553/

  • Searle, J. (2004). Mind: A brief introduction. Oxford University Press.

    Google Scholar 

  • Seth, A. (2021). Being you: A new science of consciousness. Faber & Faber

  • Sharov, A., & Tønnessen, M. (2021). Semiotic agency: Science beyond mechanism. Springer.

  • Skora, L. I, Yeomans M. R., Crombag H. S., & Scott R. B. (2021) Evidence that instrumental conditioning requires conscious awareness in humans. Cognition, 208:104546. https://doi.org/10.1016/j.cognition.2020.104546.

  • Solms, M. (2021). The hidden spring: A journey to the source of consciousness. Profile Books

  • Solms, M., & Friston, K. (2018). How and why consciousness arises: Some considerations from physics and physiology. Journal of Consciousness Studies, 25(5–6), 202–238.

    Google Scholar 

  • Telles, F. J., Corcobado, G., Trillo, A., & Rodríguez-Gironés, M. A. (2017). Multimodal cues provide redundant information for bumblebees when the stimulus is visually salient, but facilitate red target detection in a naturalistic background. PLoS One1, 12(9), e0184760. https://doi.org/10.1371/journal.pone.0184760

    Article  CAS  Google Scholar 

  • Thompson, E. (2007). Mind in life: Biology, phenomenology, and the science of mind. Harvard University Press.

    Google Scholar 

  • Tinbergen, N. (1951). The study of instinct. Clarendon Press.

    Google Scholar 

  • Tinbergen, N. (1963). On aims and methods of ethology. Zeitschrift fürTierpsychologie, 20, 410–433.

    Article  Google Scholar 

  • Tomasello, M. (2014). A Natural history of human thinking. Harvard University Press.

    Book  Google Scholar 

  • Tononi, G., Boly, M., Massimini, M., & Koch, C. (2016). Integrated information theory: From consciousness to its physical substrate. Nature Reviews Neuroscience, 17, 450–461. https://doi.org/10.1038/nrn.2016.44

    Article  CAS  Google Scholar 

  • Vallar, G., & Ronchi, R. (2009). Somatoparaphrenia: A body delusion. A review of the neuropsychological literature. Experimental Brain Research, 192, 533–551. https://doi.org/10.1007/s00221-008-1562-y

    Article  Google Scholar 

  • von Bertalanffy, L. (1952). Problems of life—an evaluation of modern biological thought. Wiley.

    Google Scholar 

  • von Holst, E., & Mittelstaedt, H. (1950). The reaference principle. Interaction between the central nervous system and the periphery. In: Selected papers of Erich von Holst: the behavioural physiology of animals and man (R. Martin, Trans., pp. 39–73). Methuen.

  • West-Eberhard, M. J. (2003). Developmental plasticity and evolution. Oxford University Press.

    Book  Google Scholar 

  • Williford, K., Bennequin, D., Friston, K., & Rudrauf, D. (2018). The projective consciousness model and phenomenal selfhood. Frontiers in Psychology, 9, 2571. https://doi.org/10.3389/fpsyg.2018.02571

    Article  Google Scholar 

  • Wyles, J. S., Kunkel, J. G., & Wilson, A. C. (1983). Birds, behavior, and anatomical evolution. Proceedings of the National Academy of Sciences USA 80, 4394‒4397. https://doi.org/10.1073/pnas.80.14.4394

  • Zacks, O., Ginsburg, S., & Jablonka, E (2022). The futures of the past. Journal of Consciousness Studies, 29 (3–4), 29–61. https://doi.org/10.53765/20512201.29.3.029

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Acknowledgements

We are grateful to the referees and to the editors for their helpful suggestions. Adi Mark helped us with the figures and we are very grateful for her for her assistance. The paper is dedicated to the memory of Marion J. Lamb.

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Authors and Affiliations

  1. The Cohn Institute for the History and Philosophy of Science and Ideas, Tel Aviv University, Tel Aviv, Israel

    Eva Jablonka

  2. Centre for Philosophy of Natural and Social Science, London School of Economics and Political Science, London, UK

    Eva Jablonka

  3. Natural Science Department, The Open University of Israel, 1 University Road, POB 808, 4353701, Raanana, Israel

    Simona Ginsburg

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  1. Eva Jablonka
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  2. Simona Ginsburg
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Correspondence to Eva Jablonka.

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Jablonka, E., Ginsburg, S. Learning and the Evolution of Conscious Agents. Biosemiotics 15, 401–437 (2022). https://doi.org/10.1007/s12304-022-09501-y

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