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The ant colony as a test for scientific theories of consciousness

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Abstract

The appearance of consciousness in the universe remains one of the major mysteries unsolved by science or philosophy. Absent an agreed-upon definition of consciousness or even a convenient system to test theories of consciousness, a confusing heterogeneity of theories proliferate. In pursuit of clarifying this complicated discourse, we here interpret various frameworks for the scientific and philosophical study of consciousness through the lens of social insect evolutionary biology. To do so, we first discuss the notion of a forward test versus a reverse test, analogous to the normal and revolutionary phases of the scientific process. Contemporary theories of consciousness are forward tests for consciousness, in that they strive to become a means to classify the level of consciousness of arbitrary states and systems. Yet no such theory of consciousness has earned sufficient confidence such that it might be actually used as a forward test in ambiguous settings. What is needed now is thus a legitimate reverse test for theories of consciousness, to provide internal and external calibration of different frameworks. A reverse test for consciousness would ideally look like a method for referencing theories of consciousness to a tractable (and non-human) model system. We introduce the Ant Colony Test (ACT) as a rigorous reverse test for consciousness. We show that social insect colonies, though disaggregated collectives, fulfill many of the prerequisites for conscious awareness met by humans and honey bee workers. A long lineage of philosophically-neutral neurobehavioral, evolutionary, and ecological studies on social insect colonies can thus be redeployed for the study of consciousness in general. We suggest that the ACT can provide insight into the nature of consciousness, and highlight the ant colony as a model system for ethically performing clarifying experiments about consciousness.

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References

  • Aaronson, S. (2014). Giulio Tononi and me: A phi-nal exchange. Shtetl-optimized blog. https://www.scottaaronson.com/blog/?p=1823. Accessed May 30, 2018.

  • Adler, F. R., & Gordon, D. M. (2003). Optimization, conflict, and nonoverlapping foraging ranges in ants. The American Naturalist, 162(5), 529–543.

    Google Scholar 

  • Adler, F. R., Quinonez, S., Plowes, N., & Adams, E. S. (2018). Mechanistic models of conflict between ant colonies and their consequences for territory scaling. American Naturalist, 192(2), 204–216.

    Google Scholar 

  • Alem, S., Perry, C. J., Zhu, X., Loukola, O. J., Ingraham, T., Søvik, E., et al. (2016). Associative mechanisms allow for social learning and cultural transmission of string pulling in an insect. PLoS Biology, 14(10), e1002564.

    Google Scholar 

  • Alexander, S. (2017). Why are transgender people immune to optical illusions? Slate star codex. http://slatestarcodex.com/2017/06/28/why-are-transgender-people-immune-to-optical-illusions/. Accessed October 5, 2018.

  • Allen, C., & Trestman, M. (2017). Animal consciousness. In E. N. Zalta (Ed.) The stanford encyclopedia of philosophy. Metaphysics Research Lab, Stanford University. https://plato.stanford.edu/archives/win2017/entries/consciousness-animal/. Accessed January 2, 2019.

  • Animal welfare - Food Safety - European Commission. (2016). Food safety. https://ec.europa.eu/food/animals/welfare_en. Accessed October 23, 2018.

  • Baluška, F., & Levin, M. (2016). On having no head: Cognition throughout biological systems. Frontiers in Psychology, 7, 902.

    Google Scholar 

  • Barron, A. B., & Klein, C. (2016). What insects can tell us about the origins of consciousness. Proceedings of the National Academy of Sciences of the United States of America, 113(18), 4900–4908.

    Google Scholar 

  • Barron, A. B., Søvik, E., & Cornish, J. L. (2010). The roles of dopamine and related compounds in reward-seeking behavior across animal phyla. Frontiers in Behavioral Neuroscience, 4, 163.

    Google Scholar 

  • Bateson, M., Desire, S., Gartside, S. E., & Wright, G. A. (2011). Agitated honeybees exhibit pessimistic cognitive biases. Current Biology: CB, 21(12), 1070–1073.

    Google Scholar 

  • Bayne, T. (2018). On the axiomatic foundations of the integrated information theory of consciousness. Neuroscience of Consciousness, 2018(1), niy007.

    Google Scholar 

  • Bayne, T., & Carter, O. (2018). Dimensions of consciousness and the psychedelic state. Neuroscience of Consciousness, 2018(1), niy008.

    Google Scholar 

  • Bayne, T., Hohwy, J., & Owen, A. M. (2016). Are there levels of consciousness? Trends in Cognitive Sciences, 20(6), 405–413.

    Google Scholar 

  • Bethell, E. J. (2015). A “how-to” guide for designing judgment bias studies to assess captive animal welfare. Journal of Applied Animal Welfare science: JAAWS, 18(sup1), S18–S42.

    Google Scholar 

  • Bird, A. (2013). Thomas Kuhn. In E. N. Zalta (Ed.) The stanford encyclopedia of philosophy. Metaphysics Research Lab, Stanford University. https://plato.stanford.edu/archives/fall2013/entries/thomas-kuhn/. Accessed January 2, 2019.

  • Block, N. (1995a). On a confusion about a function of consciousness. The Behavioral and Brain Sciences, 18(2), 227–247.

    Google Scholar 

  • Block, N. (1995b). How many concepts of consciousness? The Behavioral and Brain Sciences, 18(2), 272–287.

    Google Scholar 

  • Block, N. (2007). Consciousness, accessibility, and the mesh between psychology and neuroscience. The Behavioral and Brain Sciences, 30(5–6), 481–499.

    Google Scholar 

  • Bochynek, T., Tanner, J. L., Meyer, B., & Burd, M. (2017). Parallel foraging cycles for different resources in leaf-cutting ants: A clue to the mechanisms of rhythmic activity. Ecological Entomology, 42(6), 849–852.

    Google Scholar 

  • Boomsma, J. J., & Gawne, R. (2017). Superorganismality and caste differentiation as points of no return: How the major evolutionary transitions were lost in translation. Biological Reviews of the Cambridge Philosophical Society. https://doi.org/10.1111/brv.12330.

    Article  Google Scholar 

  • Brancucci, A., Franciotti, R., D’Anselmo, A., Della Penna, S., & Tommasi, L. (2011). The sound of consciousness: Neural underpinnings of auditory perception. The Journal of Neuroscience: The Official Journal of the Society for Neuroscience, 31(46), 16611–16618.

    Google Scholar 

  • Bruineberg, J., Kiverstein, J., & Rietveld, E. (2016). The anticipating brain is not a scientist: The free-energy principle from an ecological-enactive perspective. Synthese, 195(6), 2417–2444.

    Google Scholar 

  • Calabrò, R. S., Cacciola, A., Bramanti, P., & Milardi, D. (2015). Neural correlates of consciousness: What we know and what we have to learn! Neurological Sciences: Official Journal of the Italian Neurological Society and of the Italian Society of Clinical Neurophysiology, 36(4), 505–513.

    Google Scholar 

  • Chalmers, D. (1996). On the search for the neural correlate of consciousness. Cogprints. http://cogprints.org/227/. Accessed October 4, 2018.

  • Chalmers, D. (2017). The hard problem of consciousness. In S. Schneider & M. Velmans (Eds.), The blackwell companion to consciousness (Vol. 4, pp. 32–42). Chichester, UK: Wiley.

    Google Scholar 

  • Cole, B. J. (1991). Short-term activity cycles in ants: Generation of periodicity by worker interaction. The American Naturalist, 137(2), 244–259.

    Google Scholar 

  • Cole, D. (2014). The Chinese room argument. Stanford encyclopedia of philosophy. https://stanford.library.sydney.edu.au/entries/chinese-room/. Accessed October 8, 2018.

  • Constant, A., Ramstead, M. J. D., Veissière, S. P. L., Campbell, J. O., & Friston, K. J. (2018). A variational approach to niche construction. Journal of the Royal Society, Interface/the Royal Society, 15(141), 20170685.

    Google Scholar 

  • Cooper, P. D., Schaffer, W. M., & Buchmann, S. L. (1985). Temperature regulation of honey bees (Apis mellifera) foraging in the Sonoran desert. The Journal of Experimental Biology, 114(1), 1–15.

    Google Scholar 

  • Crick, F., & Koch, C. (1990). Towards a neurobiological theory of consciousness. Seminars in the Neurosciences, 2, 263–275.

    Google Scholar 

  • Crick, F. C., & Koch, C. (2005). What is the function of the claustrum? Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 360(1458), 1271–1279.

    Google Scholar 

  • Crompton, J. (1954). Ways of the ant. Boston: Houghton Mifflin.

    Google Scholar 

  • Damasio, A. (2000). The feeling of what happens: Body and emotion in the making of consciousness (1st ed.). ‎Wilmington: Mariner Books.

    Google Scholar 

  • Davidson, J. D., Arauco-Aliaga, R. P., Crow, S., Gordon, D. M., & Goldman, M. S. (2016). Effect of interactions between harvester ants on forager decisions. Frontiers in Ecology and Evolution, 4, 115.

    Google Scholar 

  • Dawkins, M. S. (2012). Why animals matter: Animal consciousness, animal welfare, and human well-being (1st ed.). Oxford: Oxford University Press.

    Google Scholar 

  • De Sousa, A. (2013). Towards an integrative theory of consciousness: Part 1 (neurobiological and cognitive models). Mens Sana Monographs, 11(1), 100–150.

    Google Scholar 

  • Decker, K. S. (2013). Enders game and philosophy the logic gate is down. Hoboken: Wiley.

    Google Scholar 

  • Dehaene, S., Lau, H., & Kouider, S. (2017). What is consciousness, and could machines have it? Science, 358(6362), 486–492.

    Google Scholar 

  • Dennett, D. (2001). Are we explaining consciousness yet? Cognition, 79(1–2), 221–237.

    Google Scholar 

  • Dennett, D. C. (1995). Animal Consciousness: What matters and why. Social Research, 62(3), 691–710.

    Google Scholar 

  • Dennett, D. C. (1996). Darwin’s dangerous idea: Evolution and the meanings of life. New York City: Simon and Schuster.

    Google Scholar 

  • Dennett, D. C. (2017). From bacteria to bach and back: The evolution of minds. New York City: W.W. Norton.

    Google Scholar 

  • Doesburg, S. M., Green, J. J., McDonald, J. J., & Ward, L. M. (2009). Rhythms of consciousness: Binocular rivalry reveals large-scale oscillatory network dynamics mediating visual perception. PLoS ONE, 4(7), e6142.

    Google Scholar 

  • Entler, B. V., Cannon, J. T., & Seid, M. A. (2016). Morphine addiction in ants: A new model for self-administration and neurochemical analysis. The Journal of Experimental Biology, 219(Pt 18), 2865–2869.

    Google Scholar 

  • Fabbro, F., Aglioti, S. M., Bergamasco, M., Clarici, A., & Panksepp, J. (2015). Evolutionary aspects of self- and world consciousness in vertebrates. Frontiers in Human Neuroscience, 9, 157.

    Google Scholar 

  • Fazekas, P., & Overgaard, M. (2016). Multidimensional models of degrees and levels of consciousness. Trends in Cognitive Sciences, 20(10), 715–716.

    Google Scholar 

  • Feinerman, O., & Korman, A. (2017). Individual versus collective cognition in social insects. The Journal of Experimental Biology, 220(Pt 1), 73–82.

    Google Scholar 

  • Fekete, T., van Leeuwen, C., & Edelman, S. (2016). System, subsystem, hive: Boundary problems in computational theories of consciousness. Frontiers in Psychology, 7, 1041.

    Google Scholar 

  • Feuillet, L., Dufour, H., & Pelletier, J. (2007). Brain of a white-collar worker. The Lancet, 370(9583), 262.

    Google Scholar 

  • Flanagan, T. P., Pinter-Wollman, N. M., Moses, M. E., & Gordon, D. M. (2013). Fast and flexible: Argentine ants recruit from nearby trails. PLoS ONE, 8(8), e70888.

    Google Scholar 

  • Fleischmann, P. N., Rössler, W., & Wehner, R. (2018). Early foraging life: Spatial and temporal aspects of landmark learning in the ant Cataglyphis noda. Journal of Comparative Physiology. A, Neuroethology, Sensory, Neural, and Behavioral Physiology. https://doi.org/10.1007/s00359-018-1260-6.

    Article  Google Scholar 

  • Fossat, P., Bacqué-Cazenave, J., De Deurwaerdère, P., Delbecque, J.-P., & Cattaert, D. (2014). Comparative behavior. Anxiety-like behavior in crayfish is controlled by serotonin. Science, 344(6189), 1293–1297.

    Google Scholar 

  • Friedman, D. A., & Gordon, D. M. (2016). Ant genetics: Reproductive physiology, worker morphology, and behavior. Annual Review of Neuroscience, 39(1), 41–56.

    Google Scholar 

  • Friedman, D. A., Pilko, A., Skowronska-Krawczyk, D., Krasinska, K., Parker, J. W., Hirsh, J., et al. (2018). The role of dopamine in the collective regulation of foraging in harvester ants. iScience. https://doi.org/10.1016/j.isci.2018.09.001.

    Article  Google Scholar 

  • Friston, K. (2012). The history of the future of the Bayesian brain. NeuroImage, 62(2), 1230–1233.

    Google Scholar 

  • Friston, K. (2017a). Sentient dynamics: The kiss of chaos * through the markov blanket. Chaos and Complexity Letters; Hauppauge, 11(1), 117–140.

    Google Scholar 

  • Friston, K. (2017b). Consciousness is not a thing, but a process of inference—Karl Friston | Aeon Essays. Aeon. Aeon. https://aeon.co/essays/consciousness-is-not-a-thing-but-a-process-of-inference. Accessed October 19, 2017.

  • Friston, K. (2018). Am i self-conscious? (or does self-organization entail self-consciousness?). Frontiers in Psychology, 9, 579.

    Google Scholar 

  • Friston, K., Martin, F., & Friedman, D. A. (2018). Of woodlice and men. ALIUS Bulletin, 2, 17–43.

    Google Scholar 

  • Gilbert, S. F., Sapp, J., & Tauber, A. I. (2012). A symbiotic view of life: We have never been individuals. The Quarterly Review of Biology, 87(4), 325–341.

    Google Scholar 

  • Giurfa, M. (2013). Cognition with few neurons: Higher-order learning in insects. Trends in Neurosciences, 36(5), 285–294.

    Google Scholar 

  • Goff, P. (2017). Consciousness and fundamental reality. New York: Oxford University Press.

    Google Scholar 

  • Gordon, D. M. (1983). The relation of recruitment rate to activity rhythms in the harvester ant, Pogonomyrmex barbatus (F. Smith) (Hymenoptera: Formicidae). Journal of the Kansas Entomological Society, 56(3), 277–285.

    Google Scholar 

  • Gordon, D. M. (1986). The dynamics of the daily round of the harvester ant colony (Pogonomyrmex barbatus). Animal Behaviour, 34(5), 1402–1419.

    Google Scholar 

  • Gordon, D. M. (1989). Dynamics of task switching in harvester ants. Animal Behaviour, 38(2), 194–204.

    Google Scholar 

  • Gordon, D. M. (1992). Wittgenstein and ant-watching. Biology & Philosophy, 7(1), 13–25.

    Google Scholar 

  • Gordon, D. M. (1995). The development of an ant colony’s foraging range. Animal Behaviour, 49(3), 649–659.

    Google Scholar 

  • Gordon, D. M. (2010). Ant encounters: Interaction networks and colony behavior. Princeton: Princeton University Press.

    Google Scholar 

  • Gordon, D. M. (2013). The rewards of restraint in the collective regulation of foraging by harvester ant colonies. Nature, 498(7452), 91–93.

    Google Scholar 

  • Gordon, D. M. (2014). The ecology of collective behavior. PLoS Biology, 12(3), e1001805.

    Google Scholar 

  • Gordon, D. M., Goodwin, B. C., & Trainor, L. E. H. (1992). A parallel distributed model of the behaviour of ant colonies. Journal of Theoretical Biology, 156(3), 293–307.

    Google Scholar 

  • Gordon, D. M., & Mehdiabadi, N. J. (1999). Encounter rate and task allocation in harvester ants. Behavioral Ecology and Sociobiology, 45(5), 370–377.

    Google Scholar 

  • Grossberg, S. (2017). Towards solving the hard problem of consciousness: The varieties of brain resonances and the conscious experiences that they support. Neural Networks: The Official Journal of the International Neural Network Society, 87, 38–95.

    Google Scholar 

  • Guterstam, A., Björnsdotter, M., Gentile, G., & Ehrsson, H. H. (2015). Posterior cingulate cortex integrates the senses of self-location and body ownership. Current Biology: CB, 25(11), 1416–1425.

    Google Scholar 

  • Hameroff, S. (2012). How quantum brain biology can rescue conscious free will. Frontiers in Integrative Neuroscience, 6(October), 93.

    Google Scholar 

  • Hameroff, S., & Penrose, R. (2014). Consciousness in the universe: A review of the “Orch OR” theory. Physics of Life Reviews, 11(1), 39–78.

    Google Scholar 

  • Hasegawa, E., Ishii, Y., Tada, K., Kobayashi, K., & Yoshimura, J. (2016). Lazy workers are necessary for long-term sustainability in insect societies. Scientific Reports, 6, 20846.

    Google Scholar 

  • Hayashi, Y., Yuki, M., Sugawara, K., Kikuchi, T., & Tsuji, K. (2012). Rhythmic behavior of social insects from single to multibody. Robotics and Autonomous Systems, 60(5), 714–721.

    Google Scholar 

  • Hendricks, M. (2015). Neuroecology: Tuning foraging strategies to environmental variability. Current Biology: CB, 25(12), R498–R500.

    Google Scholar 

  • Hill, C. S. (2016). Insects: Still looking like zombies. Animal Sentience: An Interdisciplinary Journal on Animal Feeling, 1(9), 20.

    Google Scholar 

  • Hofstadter, D. R. (1981). Prelude… ant fugue. The Mind’s I: Fantasies and Reflections on Self and Soul, 149.

  • Hofstadter, D. R. (2007). I am a strange loop. New York: Basic Books.

    Google Scholar 

  • Hölldobler, B., & Wilson, E. O. (1990). The ants. Cambridge: Harvard University Press.

    Google Scholar 

  • Johnson, B. R., & Linksvayer, T. A. (2010). Deconstructing the superorganism: Social physiology, groundplans, and sociogenomics. The Quarterly Review of Biology, 85(1), 57–79.

    Google Scholar 

  • Jonas, E., & Kording, K. P. (2017). Could a neuroscientist understand a microprocessor? PLoS Computational Biology, 13(1), e1005268.

    Google Scholar 

  • Kellert, S. H., Longino, H. E., & Kenneth Waters, C. (2006). Scientific pluralism. Minneapolis: University of Minnesota Press.

    Google Scholar 

  • Key, B. (2016). Why fish do not feel pain. Animal Sentience, 1(3), 1.

    Google Scholar 

  • Key, B., & Brown, D. (2018). Designing brains for pain: Human to mollusc. Frontiers in Physiology, 9, 1027.

    Google Scholar 

  • Kim, I. S., & Dickinson, M. H. (2017). Idiothetic path integration in the fruit fly Drosophila melanogaster. Current Biology: CB, 27(15), 2227.e3–2238.e3.

    Google Scholar 

  • Kirchhoff, M., Parr, T., Palacios, E., Friston, K., & Kiverstein, J. (2018). The Markov blankets of life: Autonomy, active inference and the free energy principle. Journal of the Royal Society, Interface/the Royal Society, 15(138), 20170792.

    Google Scholar 

  • Klein, C., & Barron, A. B. (2016). Insect consciousness: Commitments, conflicts and consequences. Animal Sentience: An Interdisciplinary Journal on Animal Feeling, 1(9), 21.

    Google Scholar 

  • Koch, C. (2014). Neuronal “Superhub” might generate consciousness. Scientific American. https://doi.org/10.1038/scientificamericanmind1114-24.

    Article  Google Scholar 

  • Koch, C., Massimini, M., Boly, M., & Tononi, G. (2016). Neural correlates of consciousness: Progress and problems. Nature Reviews Neuroscience, 17(5), 307–321.

    Google Scholar 

  • Kurthen, M., Grunwald, T., & Elger, C. E. (1998). Will there be a neuroscientific theory of consciousness? Trends in Cognitive Sciences, 2(6), 229–234.

    Google Scholar 

  • Lacalli, T. (2018). Amphioxus neurocircuits, enhanced arousal, and the origin of vertebrate consciousness. Consciousness and Cognition. https://doi.org/10.1016/j.concog.2018.03.006.

    Article  Google Scholar 

  • LaRock, E. (2006). Why neural synchrony fails to explain the unity of visual consciousness. Behavior and Philosophy, 34, 39–58.

    Google Scholar 

  • Lasseter, J., & Stanton, A. (1998). A bug’s life. USA. https://www.imdb.com/title/tt0120623/. Accessed January 2, 2019.

  • Laubichler, M. D., Stadler, P. F., Prohaska, S. J., & Nowick, K. (2015). The relativity of biological function. Theory in Biosciences = Theorie in den Biowissenschaften, 134(3–4), 143–147.

    Google Scholar 

  • Linksvayer, T. A. (2015). Chapter eight—The molecular and evolutionary genetic implications of being truly social for the social insects. In A. Zayed & C. F. Kent (Eds.), Advances in insect physiology (Vol. 48, pp. 271–292). Cambridge: Academic Press.

    Google Scholar 

  • Longino, H. E. (2000). Toward an epistemology for biological pluralism. In Biology and Epistemology, pp. 261–286.

  • Longino, H. E. (2013). Studying human behavior (pp. 3430623–3430810). University of Chicago Press. http://www.press.uchicago.edu/ucp/books/book/chicago/S/bo13025491.html. Accessed December 11, 2017.

  • Low, P., Panksepp, J., Reiss, D., Edelman, D., Van Swinderen, B., & Koch, C. (2012). The Cambridge declaration on consciousness. In Francis crick memorial conference, Cambridge, England.

  • Mallatt, J., & Feinberg, T. E. (2016). Insect consciousness: Fine-tuning the hypothesis. Animal Sentience: An Interdisciplinary Journal on Animal Feeling, 1(9), 10.

    Google Scholar 

  • Mangan, B. (1993). Dennett, consciousness, and the sorrows of functionalism. Consciousness and Cognition, 2(1), 1–17.

    Google Scholar 

  • Marshall, J. A. R., Bogacz, R., Dornhaus, A., Planqué, R., Kovacs, T., & Franks, N. R. (2009). On optimal decision-making in brains and social insect colonies. Journal of the Royal Society, Interface/the Royal Society, 6(40), 1065–1074.

    Google Scholar 

  • Marvel. (2015). Ant–man (film). Wikipedia, the free encyclopedia. https://en.wikipedia.org/w/index.php?title=Ant-Man_(film)&oldid=862864493. Accessed October 9, 2018.

  • Mashour, G. A. (2018). The controversial correlates of consciousness. Science, 360(6388), 493–494.

    Google Scholar 

  • Mashour, G. A., & Alkire, M. T. (2013). Evolution of consciousness: Phylogeny, ontogeny, and emergence from general anesthesia. Proceedings of the National Academy of Sciences of the United States of America, 110(Suppl 2), 10357–10364.

    Google Scholar 

  • Mather, J. A., & Carere, C. (2016). Cephalopods are best candidates for invertebrate consciousness. Animal sentience: An. http://animalstudiesrepository.org/cgi/viewcontent.cgi?article=1127&context=animsent. Accessed January 2, 2019.

  • Matthes, J., Davis, C. S., & Potter, R. F. (Eds.). (2017). Normal science and paradigm shift. In The international encyclopedia of communication research methods (Vol. 55, pp. 1–17). Hoboken, NJ, USA: Wiley.

  • Mendl, M., Burman, O. H. P., Parker, R. M. A., & Paul, E. S. (2009). Cognitive bias as an indicator of animal emotion and welfare: Emerging evidence and underlying mechanisms. Applied Animal Behaviour Science, 118(3), 161–181.

    Google Scholar 

  • Mitchell, S. D. (2002). Integrative pluralism. Biology and Philosophy, 17(1), 55–70.

    Google Scholar 

  • Moor, J. H. (2012). The turing test: The elusive standard of artificial intelligence. Berlin: Springer.

    Google Scholar 

  • Morin, A. (2006). Levels of consciousness and self-awareness: A comparison and integration of various neurocognitive views. Consciousness and Cognition, 15(2), 358–371.

    Google Scholar 

  • Nagel, T. (1974). What is it like to be a bat? The Philosophical Review, 83(4), 435–450.

    Google Scholar 

  • Newen, A., & Vogeley, K. (2003). Self-representation: Searching for a neural signature of self-consciousness. Consciousness and Cognition, 12(4), 529–543.

    Google Scholar 

  • Noble, D. (2013). A biological relativity view of the relationships between genomes and phenotypes. Progress in Biophysics and Molecular Biology, 111(2–3), 59–65.

    Google Scholar 

  • Núñez, J. A., & Giurfa, M. (1996). Motivation and regulation of honey bee foraging. Bee World, 77(4), 182–196.

    Google Scholar 

  • O’Sullivan, J. (2010). Collective consciousness in science fiction. Foundations. http://search.proquest.com/openview/036b39d2d8caf299d7e01c8ed5ada97b/1?pq-origsite=gscholar&cbl=636386. Accessed January 2, 2019.

  • Overgaard, M. (2017). The status and future of consciousness research. Frontiers in Psychology, 8, 1719.

    Google Scholar 

  • Panksepp, J. (2004). Affective neuroscience: The foundations of human and animal emotions. Oxford: Oxford University Press.

    Google Scholar 

  • Panksepp, J. (2011). Toward a cross-species neuroscientific understanding of the affective mind: Do animals have emotional feelings? American Journal of Primatology, 73(6), 545–561.

    Google Scholar 

  • Pereboom, D. (2016). Illusionism and anti-functionalism about phenomenal consciousness. Journal of Consciousness Studies, 23(11–12), 172–185.

    Google Scholar 

  • Perry, C. J., Baciadonna, L., & Chittka, L. (2016). Unexpected rewards induce dopamine-dependent positive emotion–like state changes in bumblebees. Science, 353(6307), 1529–1531.

    Google Scholar 

  • Phillips, Z. I., Zhang, M. M., & Mueller, U. G. (2017). Dispersal of Attaphila fungicola, a symbiotic cockroach of leaf-cutter ants. Insectes Sociaux, 64(2), 1–8.

    Google Scholar 

  • Portha, S., Deneubourg, J.-L., & Detrain, C. (2002). Self-organized asymmetries in ant foraging: A functional response to food type and colony needs. Behavioral Ecology: Official Journal of the International Society for Behavioral Ecology, 13(6), 776–781.

    Google Scholar 

  • Proverbs. (n.d.). Bible. Bible. https://www.biblegateway.com/passage/?search=Proverbs+6%3A6-8&version=NIV. Accessed October 8, 2018.

  • Ramstead, M. J. D., Badcock, P. B., & Friston, K. J. (2017). Answering Schrödinger’s question: A free-energy formulation. Physics of Life Reviews. https://doi.org/10.1016/j.plrev.2017.09.001.

    Article  Google Scholar 

  • Richardson, T. O., Liechti, J. I., Stroeymeyt, N., Bonhoeffer, S., & Keller, L. (2017). Short-term activity cycles impede information transmission in ant colonies. PLoS Computational Biology, 13(5), e1005527.

    Google Scholar 

  • Robinson, E. J. H. (2014). Polydomy: The organisation and adaptive function of complex nest systems in ants. Current Opinion in Insect Science, 5, 37–43.

    Google Scholar 

  • Ross, D., & Spurrett, D. (2004). What to say to a skeptical metaphysician: A defense manual for cognitive and behavioral scientists. The Behavioral and Brain Sciences, 27(5), 603–627.

    Google Scholar 

  • Sakiyama, T., & Gunji, Y.-P. (2013). The Müller–Lyer illusion in ant foraging. PLoS ONE, 8(12), e81714.

    Google Scholar 

  • Sakiyama, T., & Gunji, Y.-P. (2016). The Kanizsa triangle illusion in foraging ants. BioSystems, 142–143, 9–14.

    Google Scholar 

  • Sapolsky, R. M. (2004). The frontal cortex and the criminal justice system. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 359(1451), 1787–1796.

    Google Scholar 

  • Sapolsky, R. M. (2017). Behave: The biology of humans at our best and worst. City of Westminster: Penguin Press.

    Google Scholar 

  • Schmidt, H., Avitabile, D., Montbrió, E., & Roxin, A. (2018). Network mechanisms underlying the role of oscillations in cognitive tasks. PLoS Computational Biology, 14(9), e1006430.

    Google Scholar 

  • Schwitzgebel, E. (2014). Tononi’s exclusion postulate would make consciousness (nearly) irrelevant. The splintered mind. http://schwitzsplinters.blogspot.com/2014/07/tononis-exclusion-postulate-would-make.html. Accessed October 11, 2018.

  • Schwitzgebel, E. (2015). If materialism is true, the United States is probably conscious. Philosophical Studies, 172(7), 1697–1721.

    Google Scholar 

  • Schwitzgebel, E. (2018a). An argument against every general theory of consciousness. The splintered mind. https://schwitzsplinters.blogspot.com/2018/05/an-argument-against-every-single.html? Accessed May 28, 2018.

  • Schwitzgebel, E. (2018b). Are garden snails conscious? Yes, no, or *gong*. The splintered mind. http://schwitzsplinters.blogspot.com/2018/09/are-garden-snails-conscious-yes-no-or.html. Accessed October 8, 2018.

  • Searle, J. R. (2002). Consciousness and language. Cambridge: Cambridge University Press.

    Google Scholar 

  • Seeley, T. D., & Levien, R. A. (1987). A colony of mind. The Sciences, 27(4), 39–42.

    Google Scholar 

  • Seth, A. K., Izhikevich, E., Reeke, G. N., & Edelman, G. M. (2006). Theories and measures of consciousness: An extended framework. Proceedings of the National Academy of Sciences of the United States of America, 103(28), 10799–10804.

    Google Scholar 

  • Shanahan, M. (2016). Consciousness as integrated perception, motivation, cognition, and action. Animal Sentience, 1(9), 12.

    Google Scholar 

  • Shoemaker, S. (1993). Functionalism and consciousness. Ciba Foundation Symposium, 174, 14–21.

    Google Scholar 

  • Shtonda, B. B., & Avery, L. (2006). Dietary choice behavior in Caenorhabditis elegans. The Journal of Experimental Biology, 209(Pt 1), 89–102.

    Google Scholar 

  • Sihn, W. (1997). Paradigm shift in the corporation: The fractal company. IFAC Proceedings Volumes, 30(19), 131–136.

    Google Scholar 

  • Silvertown, J., & Gordon, D. M. (1989). A framework for plant behavior. Annual Review of Ecology and Systematics, 20(1), 349–366.

    Google Scholar 

  • Singer, P. (1975). Animal liberation. New York City: Harper Collins.

    Google Scholar 

  • Singer, W. (1999). Neuronal synchrony: A versatile code for the definition of relations? Neuron, 24(1), 49–65.

    Google Scholar 

  • Smith, D. W. (2018). Phenomenology. In E. N. Zalta (Ed.) The stanford encyclopedia of philosophy. Metaphysics Research Lab, Stanford University. https://plato.stanford.edu/archives/sum2018/entries/phenomenology/. Accessed January 2, 2019.

  • Sokolowski, M. B. (2010). Social interactions in “simple” model systems. Neuron, 65(6), 780–794.

    Google Scholar 

  • Søvik, E., & Perry, C. (2016). The evolutionary history of consciousness. Animal Sentience: An Interdisciplinary Journal on Animal Feeling, 1(9), 19.

    Google Scholar 

  • Stern, S., Kirst, C., & Bargmann, C. I. (2017). Neuromodulatory control of long-term behavioral patterns and individuality across development. Cell, 171(7), 1649.e10–1662.e10.

    Google Scholar 

  • Tabery, J., Preda, A., & Longino, H. (2014). Pluralism, social action and the causal space of human behavior. Metascience, 23(3), 443–459.

    Google Scholar 

  • Theraulaz, G. (2014). Embracing the creativity of stigmergy in social insects. Architectural Design, 84(5), 54–59.

    Google Scholar 

  • Theraulaz, G., & Bonabeau, E. (1999). A brief history of stigmergy. Artificial Life, 5(2), 97–116.

    Google Scholar 

  • Tinbergen, N. (1963). On aims and methods of ethology. Ethology: Formerly Zeitschrift fur Tierpsychologie, 20(4), 410–433.

    Google Scholar 

  • Toadvine, T. (2018). Maurice Merleau-Ponty. In E. N. Zalta (Ed.) The stanford encyclopedia of philosophy. Metaphysics Research Lab, Stanford University. https://plato.stanford.edu/archives/spr2018/entries/merleau-ponty/. Accessed January 2, 2019.

  • Toda, K., & Platt, M. L. (2015). Animal cognition: Monkeys pass the mirror test. Current Biology: CB, 25(2), R64–R66.

    Google Scholar 

  • Tononi, G., Boly, M., Massimini, M., & Koch, C. (2016). Integrated information theory: From consciousness to its physical substrate. Nature Reviews. Neuroscience, 17(7), 450–461.

    Google Scholar 

  • Tononi, G., & Koch, C. (2015). Consciousness: Here, there and everywhere? Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences, 370(1668), 20140167.

    Google Scholar 

  • Tsuchiya, N., Wilke, M., Frässle, S., & Lamme, V. A. F. (2015). No-report paradigms: Extracting the true neural correlates of consciousness. Trends in Cognitive Sciences, 19(12), 757–770.

    Google Scholar 

  • Turing, A. M. (1950). I.—Computing machinery and intelligence. Mind; a Quarterly Review of Psychology and Philosophy, LIX(236), 433–460.

    Google Scholar 

  • Uhlhaas, P. J., Pipa, G., Lima, B., Melloni, L., Neuenschwander, S., Nikolić, D., et al. (2009). Neural synchrony in cortical networks: History, concept and current status. Frontiers in Integrative Neuroscience, 3, 17.

    Google Scholar 

  • Van Gulick, R. (2018). Consciousness. In E. N. Zalta (Ed.) The stanford encyclopedia of philosophy. Metaphysics Research Lab, Stanford University. https://plato.stanford.edu/archives/spr2018/entries/consciousness/. Accessed January 2, 2019.

  • Walter, H. (2009). Neurophilosophy of free will: From Libertarian illusions to a concept of natural autonomy. Cambridge: MIT Press.

    Google Scholar 

  • Ward, P. S. (2014). The phylogeny and evolution of ants. Annual Review of Ecology, Evolution, and Systematics, 45(1), 140822120436000.

    Google Scholar 

  • Wehner, R. (2003). Desert ant navigation: How miniature brains solve complex tasks. Journal of Comparative Physiology. A, Neuroethology, Sensory, Neural, and Behavioral Physiology, 189(8), 579–588.

    Google Scholar 

  • Wheeler, W. M. (1911). The ant-colony as an organism. Journal of Morphology, 22(2), 307–325.

    Google Scholar 

  • Wilson, R. A. (2017). Collective intentionality in non-human animals. Routledge handbook on collective. http://robwilson-philosophy.s3.amazonaws.com/mind/CollectIntentCorrectedProofsJuly2017.pdf. Accessed January 2, 2019.

  • Winston, M. L., Higo, H. A., & Slessor, K. N. (1990). Effect of various dosages of queen mandibular gland pheromone on the inhibition of queen rearing in the honey bee (Hymenoptera: Apidae). Annals of the Entomological Society of America, 83(2), 234–238.

    Google Scholar 

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Acknowledgements

We thank Tucker Chambers, Zach Phillips and Dr. Clint J. Perry for their comments on an earlier draft.

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Friedman, D.A., Søvik, E. The ant colony as a test for scientific theories of consciousness. Synthese 198, 1457–1480 (2021). https://doi.org/10.1007/s11229-019-02130-y

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