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Form Approved OMB No. 0704-0188REPORT DOCUMENTATION PAGEPublic reporting burden for this collection of information is estimated to average 1 hour per response, including the time for reviewing instructions, searching existing data sources, gathering andmaintaining the data needed, and completing and reviewing the collection of information. Send comments regarding this burden estimate or any other aspect of this collection of information,including suggestions for reducing the burden, to Department of Defense, Washington Headquarters Services, Directorate for Information Operations and Reports (0704-0188), 1215 JeffersonDavis Highway, Suite 1204, Arlington, VA 22202-4302. Respondents should be aware that notwithstanding any other provision of law, no person shall be subject to any penalty for failing to complywith a collection of information if it does not display a currently valid OMB control number.PLEASE DO NOT RETURN YOUR FORM TO THE ABOVE ADDRESS.1. REPORT DATE (DD-MM-YYYY)2. REPORT TYPE19 September 20133. DATES COVERED (From – To)Conference Proceedings29 August 2012 – 12 September 20134. TITLE AND SUBTITLE5a. CONTRACT NUMBERNeural Coding: Information Beyond ShannonFA8655-12-1-20065b. GRANT NUMBERCSP 12-20065c. PROGRAM ELEMENT NUMBER61102F6. AUTHOR(S)5d. PROJECT NUMBER5d. TASK NUMBER5e. WORK UNIT NUMBER7. PERFORMING ORGANIZATION NAME(S) AND ADDRESS(ES)8. PERFORMING ORGANIZATIONREPORT NUMBERFYZIOLOGICKY USTAV AV CR, V.V.I.VIDENSKA 1083PRAHA 4, 14220 Czech RepublicN/A9. SPONSORING/MONITORING AGENCY NAME(S) AND ADDRESS(ES)10. SPONSOR/MONITOR’S ACRONYM(S)EOARDUnit 4515APO AE 09421-4515AFRL/AFOSR/IOE (EOARD)11. SPONSOR/MONITOR’S REPORT NUMBER(S)AFRL-AFOSR-UK-PC-2013-001712. DISTRIBUTION/AVAILABILITY STATEMENTDistribution A: Approved for public release; distribution is unlimited.13. SUPPLEMENTARY NOTES14. ABSTRACTOne of the primary, and still largely unresolved, goals of computational neuroscience is 12 to understand how neurons processand convey information. Since the early studies, the challenges of understanding the principles of neuronal coding haveattracted increasing number of scientists from different fields. Among the main themes of the workshop is: What can information theory tell us about information processing in neurons? Influence of environmental statistics on information coding in sensory neurons. Noise and its role in neuronal coding.The main impact of the workshop lies in the distribution of ideas and approaches that, although applied to different systems,share the common language of information theory. In particular, we believe that the computational neuroscience communityhas not yet fully benefited from the richness and generality of information theory as originally proposed by Shannon himself.Consequentially, workshops of this kind have the potential to change the perception of certain topics and perhaps, hopefully,provide solutions to some of the long-lasting questions in neuronal information processing.15. SUBJECT TERMSEOARD, Cyber Operations, Network Security16. SECURITY CLASSIFICATION OF:a. REPORTUNCLASb. ABSTRACTUNCLASc. THIS PAGEUNCLAS17. LIMITATION OFABSTRACTSAR18, NUMBEROF PAGES3119a. NAME OF RESPONSIBLE PERSONJames H. Lawton19b. TELEPHONE NUMBER (Include area code) 44 (0)1895 616187Standard Form 298 (Rev. 8/98)Prescribed by ANSI Std. Z39-18

Neural Coding:Information beyond ShannonProgramme and AbstractsPrague, Czech Republic,July 3–4, 2013'LVWULEXWLRQ SSURYHG IRU SXEOLF UHOHDVH GLVWULEXWLRQ LV XQOLPLWHG

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We wish to thank the following for their contribution to the success of this conference:European Office of Aerospace Research and Development,Air Force Office of Scientific Research,United States Air Force Research LaboratoryGrant No.: FA8655-12-1-2006Local Organising Agency:CONFORG, Ltd.http://www.conforg.czCelebrating the 60th anniversary of its foundation'LVWULEXWLRQ SSURYHG IRU SXEOLF UHOHDVH GLVWULEXWLRQ LV XQOLPLWHG

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ContentsForeword (Petr Lansky)A Random Walk through Computational Neuroscienceand Sixty Years of the Institute of Physiology, Prague1Workshop Information3Programme4Abstracts (ordered by the name of the first author)7Computing SurpriseAlberto Apostolico7Mathematical and Neuroscientific Arguments for Generalized Inverse Gaussian ISIDurations of Cortical NeuronsToby Berger8On the Functional and Structural Characterization of Hubs in Protein-protein Interaction NetworksConcettina Guerra9A Simple Model That Can Accurately Predict Spike Timings Generated by VariousKinds of Neurons10Ryota KobayashiAdaptation Improves Information Transfer in Hodgkin-Huxley NeuronsLubomir Kostal and Ryota Kobayashi11The Effect of Interspike Interval Statistics on the Information Gain under the RateCoding Hypothesis12Shinsuke KoyamaStochastic Neuronal Models and Information Transfer in NeuronsPetr Lansky13The t-Transform and Its Inverse in Modeling Neural Encoding and DecodingAurel A. Lazar14Information Filtering by Spiking Neurons as a Simple Form of Information Processing15Benjamin Lindner'LVWULEXWLRQ SSURYHG IRU SXEOLF UHOHDVH GLVWULEXWLRQ LV XQOLPLWHG

Maximizing Information Divergence from Exponential FamiliesFrantisek Matus16On the Calculation of Fisher Information and Entropy by Penalized ML DensityEstimation17Ondrej PokoraInformation from Molecules: a Pluridisciplinary Approach to the Sense of SmellJean-Pierre Rospars18Non-parametric Estimation of Mutual Information of d-dimensional VectorsLaura Sacerdote, Roberta Sirovich and Maria T. Giraudo19(Topic to be specified later)Daniel Suta20Beyond Shannon Information in Quantum CommunicationVladyslav Usenko21List of Participants'LVWULEXWLRQ SSURYHG IRU SXEOLF UHOHDVH GLVWULEXWLRQ LV XQOLPLWHG 23

ForewordA Random Walk through Computational Neuroscienceand Sixty Years of the Institute of Physiology, Prague“A neuron is an electrically excitable cell that processes and transmits information . . . ” is theopening of the Wikipedia article if inquiring about neuron. This fact automatically impliesan everlasting close relationship between neuroscience and information theory. Because ofthe celebration of the sixtieth anniversary of the Institute of Physiology we wish to remind theparticipants of this workshop that our Institute has always been active in this close interdisciplinary interaction, only the labels have changed over the decades. Of course, withoutclaiming completeness and precedences, we wish to mention some names close to us and theirroles in the history of the Institute and in the history of the field in general.The concept of randomness is crucial to information theory, and its application camelater, after the famous formalizations made in neuroscience by McCulloch-Pitts and HodgkinHuxley. George Gerstein published a paper called “Mathematical models for all-or-none activity of some neurons” in the journal IRA (now IEEE) Transactions on Information Theory. Simultaneously many other papers aiming to find how information is coded by neurons opened thegolden era in this direction of research. Personally, the sixties were great in so many respectsso why should not theoretical neuroscience also be included. In this decade, papers on thetopic of stochastic neuronal activity started to appear in large numbers being published in themost prestigious journals.Two immensely important meetings were organized during that period. Eduardo Caianiello attracted prominent researchers to Ravello to attend a School on Neural Networks in June,1967. The beauty of the Amalfi coast surely stimulated the participants and the contributionsof Luigi Ricciardi, Peter Johanesma or Michael Arbib’s. Some ideas on information processingin the cerebellum, should be mentioned. In the following year, January, 1968, and at the almostas beautiful location as Ravello, in San Diego, the workshop on Neural Coding was organizedby Donald Perkel and Theodore Bullock. Such a long time after the event, it is difficult tospeculate why no Europeans, except Donald McKay from UK, who also attended the Ravellomeeting, were invited to San Diego. Among the participants were – Jack Cowan, GeorgeGerstein, Vernon Mountcastle, Wilfried Rall, Jose (Pepe) Segundo, Richard Stein and others,several of whom also attended the meeting in Ravello. No one from Prague participated inthese meetings, nevertheless, what was going on here was at least part of the general trend.George Gerstein spent his sabbatical here at the Institute and it was a few years after publication of his seminal article, coauthored by Benoit Mandelbrot, on Random walk models forspike activity of a single neuron. At the same time two young researchers Jan Bures and TomasRadil arranged that a LINC-8 was purchased and used in recording and analyzing the spikingactivity of neurons. (The LINC-8 was built as a laboratory computer. It was small enoughto fit in a laboratory environment and included hardware capabilities necessary to monitorand control experiments.) Even younger fellows Ivan Krekule, Jan Skvaril and Josef Syka didboth experimental and theoretical research in that direction. The end of the sixties was soturbulent in Prague that this so promising effort did not last very long. At least, the paper1'LVWULEXWLRQ SSURYHG IRU SXEOLF UHOHDVH GLVWULEXWLRQ LV XQOLPLWHG

in Kybernetika (that time computational neuroscience was usually labeled as Cybernetics) onSpontanous discharge patterns of mesencephalic neurons was published in 1971.After this period, history started to move much faster, as I personally recall. However, thehope in cracking the neural code decreased. In the seventies, we had several visitors. GeorgeMoore came to visit Ivan Krekule and gave a seminar at the Institute. Renato Capocelli,a student of Luigi Ricciardi brought us a new book by his teacher on stochastic diffusionprocesses. Later on, Henry Tuckwell shortly visited Jan Bures, the former one being thespreading depression modeler, the latter an expert on its experimental studies. The timechanged into the eighties and at this point the third meeting has to be mentioned. LuigiRicciardi organized it in May of 1987. The opening was held in Palazzo Cassano in Napoliand the meeting at hotel Caesar Augustus on the island of Capri. The beauty of the locationprobably surmounted the previous meetings and the impulse for future research was at leastequally important. Among many participants were Sun-ichi Amari, Arun Holden and JohnRinzel together with others who had already attended Ravello twenty years earlier. At thesame time, the label “computational” became fashionable. Actually, the honour of first usingthis term is claimed by Eric Schwartz who, at the invitation of Charles Smith and Carl York,organized a meeting and later edited a book called Computational Neuroscience. However,simultaneously Neuroinformatics as a field which stands at the intersection of neuroscienceand information science was established. So, we have several labels suitable for this workshopbut the labels are not as important as the content.The end of the eighties brought enormous changes in our personal lives. Suddenly webecame a real part of events. This period is so recent that it is hard to recall it in a properperspective. In the autumn of 1995 we organized the Neural Coding workshop and from thattime every second year the event has moved all around the world coming back to Prague ayear ago. Pepe Segundo is always with us and it ensures the continuity with the workshoporganized in San Diego fifty-five years ago. Pepe is a permanent attender but the otherlegendary figures, at least occasionally, join these workshops. George Gerstein was withus in Tainan, John Rinzel in Plymouth, Luigi Ricciardi in Prague and Versailles, a sessionremembering the work of Donald Perkel was organized in Montevideo, Sun-ichi Amari andHenry Tuckwell come more frequently. Many, many other old and new friends attend but tomention all of them would make the list too long for this short Foreword. However, thosewho extended their contacts into a long lasting cooperation deserve to be acknowledged –Jean-Pierre Rospars, Susanne Ditlevsen, Laura Sacerdote, Roger Rodrigues, Charlie Smith,Henry Tuckwell, Cindy Greenwood and recently two young colleagues – Ryota Kobayashiand Massimilliano Tamborrino.Now we meet on a new occasion and we hope that the location and the interaction amongthe participants will again remain in your hearts and minds for a long time.Petr Lansky2'LVWULEXWLRQ SSURYHG IRU SXEOLF UHOHDVH GLVWULEXWLRQ LV XQOLPLWHG

Workshop InformationMain TopicsOne of the primary, and still largely unresolved, goals of computational neuroscience is tounderstand how neurons process and convey information. Since the early studies, the challenges ofunderstanding the principles of neuronal coding have attracted increasing number of scientistsfrom different fields. Among the main themes of the workshop is: What can information theory tell us about information processing in neurons? Influence of environmental statistics on information coding in sensory neurons. Noise and its role in neuronal coding.The main impact of the workshop lies in the distribution of ideas and approaches that,although applied to different systems, share the common language of information theory.In particular, we believe that the computational neuroscience community has not yet fullybenefited from the richness and generality of information theory as originally proposed byShannon himself. Consequentially, workshops of this kind have the potential to change theperception of certain topics and perhaps, hopefully, provide solutions to some of the longlasting questions in neuronal information processing.Conference Venue and OrganizersVenueVilla VoytaK Novemu Dvoru 124/54Praha 4, Lhotka142 00Czech RepublicOrganizersLubomir Kostal, Petr LanskyInstitu