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UCSD COGS 107B - Entorhinal Cortex

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perience correlate with subsequent choices offersstrong evidence for the existence of intrinsic pref-erences. Although it is not clear how malleablethese preferences are, their existence may havehealth implications for the way in which in-dividuals deal with events that are known to beunpleasant—for example, going to the doctor forpainful procedures. The neurobiological mecha-nisms governing dreading behavior may holdclues for both better pain management andimprovements in public health.References and Notes1. P. A. Samuelson, Rev. Econ. Stud. 4, 155 (1937).2. The opposite of utility is generally referred to as disutility,but for the sake of clarity, we refer to ‘‘utility’’ aspossessing both positive and negative domains.3. G. Loewenstein, Econ. J. 97 , 666 (1987).4. A. Caplin, J. Leahy, Q. J. Econ. 116, 55 (2001).5. R.-R. Ji, T. Kohno, K. A. Moore, C. J. Woolf, TrendsNeurosci. 26, 696 (2003).6. A. Ploghaus et al., Science 284, 1979 (1999).7. T. Koyama, J. G. McHaffie, P. J. Laurienti, R. C. Coghill,Proc. Natl. Acad. Sci. U.S.A. 102, 12950 (2005).8. T. T. Raij, J. Numminen, S. Narvanen, J. Hiltunen, R. Hari,Proc. Natl. Acad. Sci. U.S.A. 102, 2147 (2005).9. I. Tracey, Curr. Opin. Neurobiol. 15, 478 (2005).10. A. D. Craig, Annu. Rev. Neurosci. 26, 1 (2003).11. R. Peyron, B. Laurent, L. Garcia-Larrea, Neurophysiol.Clin. 30, 263 (2000).12. Materials and methods are available as supportingmaterial on Science Online.13. H. R. Varian, Intermediate Microeconomics: A ModernApproach (Norton & Co., New York, ed. 6, 2002).14. K. C. Berridge, in Well-Being. The Foundations of HedonicPsychology, D. Kahneman, E. Diener, N. Schwarz, Eds.(Russell Sage Foundation, New York, 1999), pp. 525–557.15. P. Petrovic, K. M. Petersson, P. Hansson, M. Ingvar,Neuroimage 16, 1142 (2002).16. B. A. Vogt, Nat. Rev. Neurosci. 6, 533 (2005).17. D. E. Bentley et al., Clin. Neurophysiol. 115, 1846(2004).18. D. E. Bentley, S. W. G. Derbyshire, P. D. Youell,A. K. P. Jones, Pain 102, 265 (2003).19. E. A. Phelps, Annu. Rev. Psychol. 57, 27 (2006).20. P. Rainville, B. Carrier, R. K. Hofbauer, M. C. Bushnell,G. H. Duncan, Pain 82, 159 (1999).21. S. M. McClure, D. I. Laibson, G. Loewenstein, J. D. Cohen,Science 306, 503 (2004).22. D. Kahneman, P. P. Wakker, R. Sarin, Q. J. Econ. 112, 375(1997).23. C. Camerer, G. Loewenstein, D. Prelec, J. Econ. Lit. 43,9(2005).24. R. A. Rescorla, A. R. Wagner, in Classical Conditioning 2:Current Research and Theory,A.H.Black,W.F.Prokasy,Eds.(Appleton Century-Crofts, New York, 1972), pp. 64–69.25. R. S. Sutton, Mach. Learn. 3, 9 (1988).26. C. K. Morewedge, D. T. Gilbert, T. D. Wilson, Psychol. Sci.16, 626 (2005).27. A. Ploghaus et al., Proc. Natl. Acad. Sci. U.S.A. 97, 9281(2000).28. A. Ploghaus, L. Becerra, C. Borras, D. Borsook, TrendsCogn. Sci. 7, 197 (2003).29. T. V. Salomons, T. Johnstone, M.-M. Backonja, R. J. Davidson,J. Neurosci. 24, 7199 (2004).30. T. D. Wager et al., Science 303, 1162 (2004).31. C. A. Porro et al., J. Neurosci. 22, 3206 (2002).32. A. Ferretti et al., Neuroimage 23, 1217 (2004).33. U. Bingel et al., Neuroimage 23, 224 (2004).34. We thank C. M. Capra, C. Noussair, A. Rangel, andA. Rustichini for comments on this paper. Supported bygrants from the National Institute on Drug Abuse(DA00367 and DA016434).Supporting Online Materialwww.sciencemag.org/cgi/content/full/312/5774/754/DC1Materials and MethodsFigs. S1 to S6Tables S1 to S3References12 December 2005; accepted 17 March 200610.1126/science.1123721Conjunctive Representation ofPosition, Direction, and Velocity inEntorhinal CortexFrancesca Sargolini,1Marianne Fyhn,1Torkel Hafting,1Bruce L. McNaughton,1,2Menno P. Witter,1,3May-Britt Moser,1Edvard I. Moser1*Grid cells in the medial entorhinal cortex (MEC) are part of an environment-independent spatialcoordinate system. To determine how information about location, direction, and distance isintegrated in the grid-cell network, we recorded from each principal cell layer of MEC in rats thatexplored two-dimensional environments. Whereas layer II was predominated by grid cells, grid cellscolocalized with head-direction cells and conjunctive grid  head-direction cells in the deeperlayers. All cell types were modulated by running speed. The conjunction of positional, directional,and translational information in a single MEC cell type may enable grid coordinates to be updatedduring self-motion–based navigation.The MEC is the hub of a widespread brainnetwork for spatial navigation (1–11).Layer II of the MEC contains a two-dimensional (2D), ensemble-encoded metricmap of relative spatial location (6–8)thatisindependent of the specific environment andthe external sensory cues (7, 11). The elementsof the map are Bgrid[ cells, which fire when-ever the animal_s position coincides with thevertices of a periodic triangular grid span-ning the complete surface of the environ-ment, with different cells having differentfiring coordinates on the unit grid (7, 12).The regular structure of the grid field, and theenvironmentally invariant relationshipsamong simultaneously recorded grid fields(13), implicates the grid cell as part of auniversal, path-integration–based spatial met-ric, but its interaction with other cell types inMEC is not understood. To investigate theintegration of metric spatial information in themultilayered entorhinal network (10, 14, 15),we compared the activity of cell populations inits four principal cell layers while rats wererunning in a 2D environment (16). Recordingswere made from the most dorsal 23% of MECin 17 rats (Fig. 1).Grid cells with tessellating firing fields (7)were observed in all principal cell layers (Fig.1, A and B). To compare their prevalence, weestimated the periodicityoftheratemapofeach cell by computing a 2D autocorrelationmatrix for the rate distribution (Fig. 1C, left),rotating the autocorrelation map in steps of 6-,and calculating the correlation between eachrotated map and the original. Grid structure wasapparent as a sinusoidal modulation of thiscorrelation, with peaks recurring at multiples of60- (Fig. 1C, right, and fig. S1) (12). The de-gree of Bgridness[ was expressed as the differ-ence between the correlations at the expectedpeaks (60- and 120-) and the expected troughs(30-,90-, and 150-) of the function. The pro-portion of cells with a sinusoidal modulationwas layer-dependent (Fig. 1D and table S1).Whereas most well-separated layer II cells hadstrongly periodic firing fields, only a smallerproportion of the deeper


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