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One influential conjecture is that consciousness is to do with some form of causality, measurable through information. The integrated information theory of consciousness (IIT) proposes that conscious experience, filled with rich and specific content, corresponds directly to a hierarchically organised, irreducible pattern of causal interactions; i.e. an integrated informational structure among elements of a system. Here, we tested this conjecture in a simple biological system (fruit flies), estimating the information structure of the system during wakefulness and general anesthesia. Consistent with this conjecture, we found that integrated interactions among populations of neurons during wakefulness collapsed to isolated clusters of interactions during anesthesia. We used classification analysis to quantify the accuracy of discrimination between wakeful and anesthetised states, and found that informational structures inferred conscious states with greater accuracy than a scalar summary of the structure, a measure which is generally championed as the main measure of IIT. In stark contrast to a view which assumes feedforward architecture for insect brains, especially fly visual systems, we found rich information structures, which cannot arise from purely feedforward systems, occurred across the fly brain. Further, these information structures collapsed uniformly across the brain during anesthesia. Our results speak to the potential utility of the novel concept of an \u201cinformational structure\u201d as a measure for level of consciousness, above and beyond simple scalar values.<\/jats:p>","DOI":"10.1371\/journal.pcbi.1008722","type":"journal-article","created":{"date-parts":[[2021,2,26]],"date-time":"2021-02-26T13:42:21Z","timestamp":1614346941000},"page":"e1008722","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":36,"title":["Integrated information structure collapses with anesthetic loss of conscious arousal in Drosophila melanogaster"],"prefix":"10.1371","volume":"17","author":[{"ORCID":"https:\/\/orcid.org\/0000-0003-0926-5534","authenticated-orcid":true,"given":"Angus","family":"Leung","sequence":"first","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5077-2207","authenticated-orcid":true,"given":"Dror","family":"Cohen","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6552-7418","authenticated-orcid":true,"given":"Bruno","family":"van Swinderen","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4216-8701","authenticated-orcid":true,"given":"Naotsugu","family":"Tsuchiya","sequence":"additional","affiliation":[],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"340","published-online":{"date-parts":[[2021,2,26]]},"reference":[{"key":"pcbi.1008722.ref001","doi-asserted-by":"crossref","first-page":"435","DOI":"10.2307\/2183914","article-title":"What is it like to be a bat?","volume":"83","author":"T. Nagel","year":"1974","journal-title":"Philos Rev"},{"key":"pcbi.1008722.ref002","author":"DJ Chalmers","year":"1996"},{"key":"pcbi.1008722.ref003","article-title":"Consciousness and the brain: Deciphering how the brain codes our thoughts","author":"S Dehaene","year":"2014","journal-title":"Penguin"},{"key":"pcbi.1008722.ref004","article-title":"The crack of dawn: perceptual functions and neural mechanisms that mark the transition from unconscious processing to conscious vision","author":"V. Lamme","year":"2014","journal-title":"Open MIND. 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Herculano-Houzel","year":"2009","journal-title":"Front Hum Neurosci"},{"key":"pcbi.1008722.ref075","doi-asserted-by":"crossref","first-page":"730","DOI":"10.1016\/j.cell.2018.06.019","article-title":"A complete electron microscopy volume of the brain of adult Drosophila melanogaster","volume":"174","author":"Z Zheng","year":"2018","journal-title":"Cell"},{"key":"pcbi.1008722.ref076","article-title":"A connectome of the adult Drosophila central brain","author":"CS Xu","year":"2020","journal-title":"bioRxiv"},{"key":"pcbi.1008722.ref077","doi-asserted-by":"crossref","first-page":"663","DOI":"10.1097\/00000542-200403000-00029","article-title":"Isoflurane inhibits transmitter release and the presynaptic action potential","volume":"100","author":"X-S Wu","year":"2004","journal-title":"Anesthesiology"},{"key":"pcbi.1008722.ref078","doi-asserted-by":"crossref","first-page":"489","DOI":"10.1113\/jphysiol.2004.065748","article-title":"Isoflurane depresses glutamate release by reducing neuronal excitability at the Drosophila neuromuscular junction","volume":"558","author":"DJ Sandstrom","year":"2004","journal-title":"J Physiol"},{"key":"pcbi.1008722.ref079","doi-asserted-by":"crossref","first-page":"1069","DOI":"10.1016\/j.concog.2009.04.004","article-title":"The directionality and functional organization of frontoparietal connectivity during consciousness and anesthesia in humans","volume":"18","author":"U Lee","year":"2009","journal-title":"Conscious Cogn"},{"key":"pcbi.1008722.ref080","doi-asserted-by":"crossref","first-page":"35","DOI":"10.1016\/S0896-6273(00)81135-6","article-title":"Ion channels and synaptic organization: analysis of the Drosophila genome","volume":"26","author":"JT Littleton","year":"2000","journal-title":"Neuron"},{"key":"pcbi.1008722.ref081","doi-asserted-by":"crossref","first-page":"1934","DOI":"10.1016\/S0960-9822(02)01300-3","article-title":"Electrophysiological correlates of rest and activity in Drosophila melanogaster","volume":"12","author":"DA Nitz","year":"2002","journal-title":"Curr Biol"},{"key":"pcbi.1008722.ref082","doi-asserted-by":"crossref","first-page":"1703","DOI":"10.1152\/jn.00414.2013","article-title":"Multichannel brain recordings in behaving Drosophila reveal oscillatory activity and local coherence in response to sensory stimulation and circuit activation","volume":"110","author":"AC Paulk","year":"2013","journal-title":"J Neurophysiol"},{"key":"pcbi.1008722.ref083","doi-asserted-by":"crossref","first-page":"96","DOI":"10.3389\/fnint.2012.00096","article-title":"Competing visual flicker reveals attention-like rivalry in the fly brain","volume":"6","author":"B. Van Swinderen","year":"2012","journal-title":"Front Integr Neurosci"},{"key":"pcbi.1008722.ref084","author":"PP Mitra","year":"2007"},{"key":"pcbi.1008722.ref085","author":"C. Koch","year":"2004"},{"key":"pcbi.1008722.ref086","first-page":"1871","article-title":"LIBLINEAR: a library for large linear classification.","volume":"9","author":"R-E Fan","year":"2008","journal-title":"J Mach Learn Res"},{"key":"pcbi.1008722.ref087","doi-asserted-by":"crossref","first-page":"1540023","DOI":"10.1142\/S0218213015400230","article-title":"Performance-estimation properties of cross-validation-based protocols with simultaneous hyper-parameter optimization","volume":"24","author":"I Tsamardinos","year":"2015","journal-title":"Int J Artif Intell Tools"},{"key":"pcbi.1008722.ref088","doi-asserted-by":"crossref","first-page":"166","DOI":"10.1016\/j.neuroimage.2016.10.038","article-title":"Assessing and tuning brain decoders: cross-validation, caveats, and guidelines","volume":"145","author":"G Varoquaux","year":"2017","journal-title":"NeuroImage"},{"key":"pcbi.1008722.ref089","doi-asserted-by":"crossref","first-page":"1","DOI":"10.18637\/jss.v067.i01","article-title":"Fitting linear mixed-effects models using lme4","volume":"67","author":"D Bates","year":"2015","journal-title":"J Stat Softw"},{"key":"pcbi.1008722.ref090","doi-asserted-by":"crossref","first-page":"e4794","DOI":"10.7717\/peerj.4794","article-title":"A brief introduction to mixed effects modelling and multi-model inference in ecology","volume":"6","author":"XA Harrison","year":"2018","journal-title":"PeerJ"}],"updated-by":[{"DOI":"10.1371\/journal.pcbi.1008722","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2021,3,10]],"date-time":"2021-03-10T00:00:00Z","timestamp":1615334400000}}],"container-title":["PLOS Computational Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1008722","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,3,10]],"date-time":"2021-03-10T13:53:11Z","timestamp":1615384391000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1008722"}},"subtitle":[],"editor":[{"given":"Daniele","family":"Marinazzo","sequence":"first","affiliation":[],"role":[{"role":"editor","vocabulary":"crossref"}]}],"short-title":[],"issued":{"date-parts":[[2021,2,26]]},"references-count":90,"journal-issue":{"issue":"2","published-online":{"date-parts":[[2021,2,26]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pcbi.1008722","relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/2020.05.17.090001","asserted-by":"object"}]},"ISSN":["1553-7358"],"issn-type":[{"value":"1553-7358","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,2,26]]}}}