creators_name: Hillenbrand, Ulrich creators_name: van Hemmen, J. Leo type: journalp datestamp: 2002-08-09 lastmod: 2011-03-11 08:54:59 metadata_visibility: show title: Does Corticothalamic Feedback Control Cortical Velocity Tuning? ispublished: pub subjects: neuro-mod subjects: comp-neuro-sci subjects: bio-theory full_text_status: public keywords: corticothalamic feedback, velocity tuning, visual motion, lateral geniculate nucleus, visual cortex abstract: The thalamus is the major gate to the cortex and its contribution to cortical receptive field properties is well established. Cortical feedback to the thalamus is, in turn, the anatomically dominant input to relay cells, yet its influence on thalamic processing has been difficult to interpret. For an understanding of complex sensory processing, detailed concepts of the corticothalamic interplay need yet to be established. To study corticogeniculate processing in a model, we draw on various physiological and anatomical data concerning the intrinsic dynamics of geniculate relay neurons, the cortical influence on relay modes, lagged and nonlagged neurons, and the structure of visual cortical receptive fields. In extensive computer simulations we elaborate the novel hypothesis that the visual cortex controls via feedback the temporal response properties of geniculate relay cells in a way that alters the tuning of cortical cells for speed. date: 2001 date_type: published publication: Neural Computation volume: 13 pagerange: 327-355 refereed: TRUE referencetext: \bibitem[Ahmed et~al., 1994]{Ahmed_etal94} Ahmed, B., Anderson, J.~A., Douglas, R.~J., Martin, K. A.~C., \& Nelson, J.~C. (1994). \newblock Polyneuronal innervation of spiny stellate neurons in cat visual cortex. \newblock {\em J.\ Comp.\ Neurol.}, 341, 39--49. \bibitem[Albrecht \& Geisler, 1991]{Albrecht&Geisler91} Albrecht, D.~G. \& Geisler, W.~S. (1991). \newblock Motion selectivity and the contrast-response function of simple cells in the visual cortex. \newblock {\em Vis.\ Neurosci.}, 7, 531--546. \bibitem[Alonso et~al., 1996]{Alonso_etal96} Alonso, J.~M., Usrey, W.~M., \& Reid, R.~C. (1996). \newblock Precisely correlated firing in cells of the lateral geniculate nucleus. \newblock {\em Nature}, 383, 815--819. \bibitem[Bal et~al., 1995]{Bal_etal95a} Bal, T., von Krosigk, M., \& McCormick, D.~A. (1995). \newblock Synaptic and membrane mechanisms underlying synchronized oscillations in the ferret {LGNd} in vitro. \newblock {\em J.\ Physiol.\ (Lond.)}, 483, 641--663. \bibitem[Bloomfield \& Sherman, 1988]{Bloomfield&Sherman88} Bloomfield, S.~A. \& Sherman, S.~M. (1988). \newblock Postsynaptic potentials recorded in neurons of the cat's lateral geniculate nucleus following electrical stimulation of the optic chiasm. \newblock {\em J.\ Neurophysiol.}, 60, 1924--1945. \bibitem[Cleland \& Harding, 1983]{Cleland&Harding83} Cleland, B.~G. \& Harding, T.~H. (1983). \newblock Response to the velocity of moving visual stimuli of the brisk classes of ganglion cells in the cat retina. \newblock {\em J.\ Physiol.\ (Lond.)}, 345, 47--63. \bibitem[Cleland et~al., 1979]{Cleland_etal79} Cleland, B.~G., Harding, T.~H., \& Tulunay-Keesey, U. (1979). \newblock Visual resolution and receptive field size: {E}xamination of two kinds of cat retinal ganglion cell. \newblock {\em Science}, 205, 1015--1017. \bibitem[Contreras et~al., 1993]{Contreras_etal93} Contreras, D., Dossi, R.~C., \& Steriade, M. (1993). \newblock Electrophysiological properties of cat reticular thalamic neurones in vivo. \newblock {\em J.\ Physiol.\ (Lond.)}, 470, 273--294. \bibitem[Cox \& Sherman, 1999]{Cox&Sherman99} Cox, C.~L. \& Sherman, S.~M. (1999). \newblock Glutamate inhibits thalamic reticular neurons. \newblock {\em J.\ Neurosci.}, 19, 6694--6699. \bibitem[Cox et~al., 1998]{Cox_etal98} Cox, C.~L., Zhou, Q., \& Sherman, S.~M. (1998). \newblock Glutamate locally activates dendritic outputs of thalamic interneurons. \newblock {\em Nature}, 394, 478--482. \bibitem[Crick \& Koch, 1998]{Crick&Koch98} Crick, F. \& Koch, C. (1998). \newblock Constraints on cortical and thalamic projections: The no-strong-loops hypothesis. \newblock {\em Nature}, 391, 245--250. \bibitem[Crook et~al., 1998]{Crook_etal98} Crook, J.~M., Kisvarday, Z.~F., \& Eysel, U.~T. (1998). \newblock Evidence for a contribution of lateral inhibition to orientation tuning and direction selectivity in cat visual cortex: reversible inactivation of functionally characterized sites combined with neuroanatomical tracing techniques. \newblock {\em Eur.\ J.\ Neurosci.}, 10, 2056--2075. \bibitem[Crunelli \& Leresche, 1991]{Crunelli&Leresche91} Crunelli, V. \& Leresche, N. (1991). \newblock A role for gabab receptors in excitation and inhibition of thalamocortical cells. \newblock {\em Trends Neurosci.}, 14, 16--21. \bibitem[DeAngelis et~al., 1993]{DeAngelis_etal93} DeAngelis, G.~C., Ohzawa, I., \& Freeman, R.~D. (1993). \newblock Spatiotemporal organization of simple-cell receptive fields in the cat's striate cortex. {II}. {L}inearity of temporal and spatial summation. \newblock {\em J.\ Neurophysiol.}, 69, 1118--1135. \bibitem[DeAngelis et~al., 1995]{DeAngelis_etal95} DeAngelis, G.~C., Ohzawa, I., \& Freeman, R.~D. (1995). \newblock Receptive-field dynamics in the central visual pathways. \newblock {\em Trends Neurosci.}, 18, 451--458. \bibitem[Douglas et~al., 1995]{Douglas_etal95} Douglas, R.~J., Koch, C., Mahowald, M., Martin, K. A.~C., \& Suarez, H.~H. (1995). \newblock Recurrent excitation in neocortical circuits. \newblock {\em Science}, 269, 981--985. \bibitem[Emerson, 1997]{Emerson97} Emerson, R.~C. (1997). \newblock Quadrature subunits in directionally selective simple cells: spatiotemporal interactions. \newblock {\em Vis.\ Neurosci.}, 14, 357--371. \bibitem[Ferster et~al., 1996]{Ferster_etal96} Ferster, D., Chung, S., \& Wheat, H. (1996). \newblock Orientation selectivity of thalamic input to simple cells of cat visual cortex. \newblock {\em Nature}, 380, 249--252. \bibitem[Field, 1994]{Field94} Field, D.~J. (1994). \newblock What is the goal of sensory coding? \newblock {\em Neural Comput.}, 6, 559--601. \bibitem[Funke \& Eysel, 1992]{Funke&Eysel92} Funke, K. \& Eysel, U.~T. (1992). \newblock {EEG}-dependent modulation of response dynamics of cat {dLGN} relay cells and the contribution of corticogeniculate feedback. \newblock {\em Brain Res.}, 573, 217--227. \bibitem[Gerstner \& van Hemmen, 1992]{Gerstner&vanHemmen92} Gerstner, W. \& van Hemmen, J.~L. (1992). \newblock Associative memory in a network of `spiking' neurons. \newblock {\em Network}, 3, 139--164. \bibitem[Godwin et~al., 1996]{Godwin_etal96} Godwin, D.~W., Vaughan, J.~W., \& Sherman, S.~M. (1996). \newblock Metabotropic glutamate receptors switch visual response mode of lateral geniculate nucleus cells from burst to tonic. \newblock {\em J.\ Neurophysiol.}, 76, 1800--1816. \bibitem[Guido et~al., 1992]{Guido_etal92} Guido, W., Lu, S.-M., \& Sherman, S.~M. (1992). \newblock Relative contributions of burst and tonic responses to the receptive field properties of lateral geniculate neurons in the cat. \newblock {\em J.\ Neurophysiol.}, 68, 2199--2211. \bibitem[Guido et~al., 1995]{Guido_etal95} Guido, W., Lu, S.-M., Vaughan, J.~W., Godwin, D.~W., \& Sherman, S.~M. (1995). \newblock Receiver operating characteristic ({ROC}) analysis of neurons in the cat's lateral geniculate nucleus during tonic and burst response mode. \newblock {\em Vis.\ Neurosci.}, 12, 723--741. \bibitem[Guido \& Weyand, 1995]{Guido&Weyand95} Guido, W. \& Weyand, T. (1995). \newblock Burst responses in thalamic relay cells of the awake, behaving cat. \newblock {\em J.\ Neurophysiol.}, 74, 1782--1786. \bibitem[Guillery, 1995]{Guillery95} Guillery, R.~W. (1995). \newblock Anatomical evidence concerning the role of the thalamus in corticocortical communication: {A} brief review. \newblock {\em J.\ Anat.}, 187, 583--592. \bibitem[Gulyas et~al., 1990]{Gulyas_etal90} Gulyas, B., Lagae, L., Eysel, U., \& Orban, G.~A. (1990). \newblock Corticofugal feedback influences the responses of geniculate neurons to moving stimuli. \newblock {\em Exp.\ Brain.\ Res.}, 79, 441--446. \bibitem[Hammond \& Pomfrett, 1990]{Hammond&Pomfrett90} Hammond, P. \& Pomfrett, C.~J. (1990). \newblock Directionality of cat striate cortical neurones: Contribution of suppression. \newblock {\em Exp.\ Brain.\ Res.}, 81, 417--425. \bibitem[Hartveit \& Heggelund, 1990]{Hartveit&Heggelund90} Hartveit, E. \& Heggelund, P. (1990). \newblock Neurotransmitter receptors mediating excitatory input to cells in the cat lateral geniculate nucleus. {II}. {N}onlagged cells. \newblock {\em J.\ Neurophysiol.}, 63, 1361--1372. \bibitem[Hassenstein \& Reichardt, 1956]{Hassenstein&Reichardt56} Hassenstein, B. \& Reichardt, W. (1956). \newblock Systemtheoretische {A}nalyse {d}er {Z}eit-, {R}eihenfolgen- {u}nd {V}orzeichenauswertung {b}ei {d}er {B}ewegungsperzeption {d}es {R}{\"u}sselk{\"a}fers {\em {c}hlorophanus}. \newblock {\em Zeitschrift f\"ur Naturforschung}, 11 b, 513--524. \bibitem[Heggelund \& Hartveit, 1990]{Heggelund&Hartveit90} Heggelund, P. \& Hartveit, E. (1990). \newblock Neurotransmitter receptors mediating excitatory input to cells in the cat lateral geniculate nucleus. {I}. {L}agged cells. \newblock {\em J.\ Neurophysiol.}, 63, 1347--1360. \bibitem[Hille, 1992]{Hille92} Hille, B. (1992). \newblock {\em Ionic channels of excitable membranes}. \newblock Sunderland, Massachusetts: Sinauer Associates Inc. \bibitem[Hillenbrand \& {van Hemmen}, 2000]{Hillenbrand&vanHemmen00} Hillenbrand, U. \& {van Hemmen}, J.~L. (2000). \newblock Spatiotemporal adaptation through corticothalamic loops: {A} hypothesis. \newblock {\em Vis.\ Neurosci.}, 17, 107--118. \bibitem[Hirsch et~al., 1998]{Hirsch_etal98} Hirsch, J.~A., Alonso, J.-M., Reid, R.~C., \& Martinez, L.~M. (1998). \newblock Synaptic integration in striate cortical simple cells. \newblock {\em J.\ Neurosci.}, 18, 9517--9528. \bibitem[Huguenard \& McCormick, 1992]{Huguenard&McCormick92} Huguenard, J.~R. \& McCormick, D.~A. (1992). \newblock Simulation of the currents involved in rythmic oscillations in thalamic relay neurons. \newblock {\em J.\ Neurophysiol.}, 68, 1373--1383. \bibitem[Humphrey \& Weller, 1988a]{Humphrey&Weller88a} Humphrey, A.~L. \& Weller, R.~E. (1988a). \newblock Functionally distinct groups of {X}-cells in the lateral geniculate nucleus of the cat. \newblock {\em J.\ Comp.\ Neurol.}, 268, 429--447. \bibitem[Humphrey \& Weller, 1988b]{Humphrey&Weller88b} Humphrey, A.~L. \& Weller, R.~E. (1988b). \newblock Structural correlates of functionally distinct {X}-cells in the lateral geniculate nucleus of the cat. \newblock {\em J.\ Comp.\ Neurol.}, 268, 448--468. \bibitem[Jagadeesh et~al., 1993]{Jagadeesh_etal93} Jagadeesh, B., Wheat, H.~S., \& Ferster, D. (1993). \newblock Linearity of summation of synaptic potentials underlying direction selectivity in simple cells of the cat visual cortex. \newblock {\em Science}, 262, 1901--1904. \bibitem[Jagadeesh et~al., 1997]{Jagadeesh_etal97} Jagadeesh, B., Wheat, H.~S., Kontsevich, L.~L., Tyler, C.~W., \& Ferster, D. (1997). \newblock Direction selectivity of synaptic potentials in simple cells of the cat visual cortex. \newblock {\em J.\ Neurophysiol.}, 78, 2772--2789. \bibitem[Jahnsen \& Llin{\'a}s, 1984a]{Jahnsen&Llinas84a} Jahnsen, H. \& Llin{\'a}s, R. (1984a). \newblock Electrophysiological properties of guinea-pig thalamic neurons: {A}n in vitro study. \newblock {\em J.\ Physiol.\ (Lond.)}, 349, 205--226. \bibitem[Jahnsen \& Llin{\'a}s, 1984b]{Jahnsen&Llinas84b} Jahnsen, H. \& Llin{\'a}s, R. (1984b). \newblock Ionic basis for the electroresponsiveness and oscillatory properties of guinea-pig thalamic neurons in vitro. \newblock {\em J.\ Physiol.\ (Lond.)}, 349, 227--247. \bibitem[Katz, 1987]{Katz87} Katz, L.~C. (1987). \newblock Local circuitry of identified projection neurons in cat visual cortex brain slices. \newblock {\em J.\ Neurosci.}, 7, 1223--1249. \bibitem[Kistler et~al., 1997]{Kistler_etal97} Kistler, W.~M., Gerstner, W., \& van Hemmen, J.~L. (1997). \newblock Reduction of the {H}odgkin-{H}uxley equations to a single-variable threshold model. \newblock {\em Neural Comput.}, 9, 1015--1045. \bibitem[Koch, 1987]{Koch87} Koch, C. (1987). \newblock The action of the corticofugal pathway on sensory thalamic nuclei: A hypothesis. \newblock {\em Neurosci.}, 23, 399--406. \bibitem[Kontsevich, 1995]{Kontsevich95} Kontsevich, L.~L. (1995). \newblock The nature of the inputs to cortical motion detectors. \newblock {\em Vision Res.}, 35, 2785--2793. \bibitem[Kwon et~al., 1991]{Kwon_etal91} Kwon, Y.~H., Esguerra, M., \& Sur, M. (1991). \newblock {NMDA} and non-{NMDA} receptors mediate visual responses of neurons in the cat's lateral geniculate nucleus. \newblock {\em J.\ Neurophysiol.}, 66, 414--428. \bibitem[Lo \& Sherman, 1994]{Lo&Sherman94} Lo, F.~S. \& Sherman, S.~M. (1994). \newblock Feedback inhibition in the cat's lateral geniculate nucleus. \newblock {\em Exp.\ Brain.\ Res.}, 100, 365--368. \bibitem[Lu et~al., 1992]{Lu_etal92} Lu, S.-M., Guido, W., \& Sherman, S.~M. (1992). \newblock Effects of membrane voltage on receptive field properties of lateral geniculate neurons in the cat: {C}ontributions of the low-treshold {C}a$^{2+}$ conductance. \newblock {\em J.\ Neurophysiol.}, 68, 2185--2198. \bibitem[Maex \& Orban, 1996]{Maex&Orban96} Maex, R. \& Orban, G.~A. (1996). \newblock Model circuit of spiking neurons generating directional selectivity in simple cells. \newblock {\em J.\ Neurophysiol.}, 75, 1515--1545. \bibitem[Mastronarde, 1987a]{Mastronarde87a} Mastronarde, D.~N. (1987a). \newblock Two classes of single-input {X}-cells in cat lateral geniculate nucleus. {I}. {R}eceptive-field properties and classification of cells. \newblock {\em J.\ Neurophysiol.}, 57, 357--380. \bibitem[Mastronarde, 1987b]{Mastronarde87b} Mastronarde, D.~N. (1987b). \newblock Two classes of single-input {X}-cells in cat lateral geniculate nucleus. {II}. {R}etinal inputs and the generation of receptive-field properties. \newblock {\em J.\ Neurophysiol.}, 57, 381--413. \bibitem[McClurkin et~al., 1994]{McClurkin_etal94} McClurkin, J.~W., Optican, L.~M., \& Richmond, B.~J. (1994). \newblock Cortical feedback increases visual information transmitted by monkey parvocellular lateral geniculate nucleus neurons. \newblock {\em Vis.\ Neurosci.}, 11, 601--617. \bibitem[McCormick, 1992]{McCormick92} McCormick, D.~A. (1992). \newblock Neurostransmitter actions in the thalamus and cerebral cortex and their role in neuromodulation of thalamocortical activity. \newblock {\em Prog.\ Neurobiol.}, 39, 337--388. \bibitem[McCormick \& Huguenard, 1992]{McCormick&Huguenard92} McCormick, D.~A. \& Huguenard, J.~R. (1992). \newblock A model of the electrophysiological properties of thalamocortical relay neurons. \newblock {\em J.\ Neurophysiol.}, 68, 1384--1400. \bibitem[McCormick \& {von Krosigk}, 1992]{McCormick&vonKrosigk92} McCormick, D.~A. \& {von Krosigk}, M. (1992). \newblock Corticothalamic activation modulates thalamic firing through glutamate ``metabotropic'' receptors. \newblock {\em Proc.\ Natl.\ Acad.\ Sci.\ USA}, 89, 2774--2778. \bibitem[McLean \& Palmer, 1989]{McLean&Palmer89} McLean, J. \& Palmer, L.~A. (1989). \newblock Contribution of linear spatiotemporal receptive field structure to velocity selectivity of simple cells in area 17 of the cat. \newblock {\em Vision Res.}, 29, 675--679. \bibitem[McLean et~al., 1994]{McLean_etal94} McLean, J., Raab, S., \& Palmer, L.~A. (1994). \newblock Contribution of linear meachanisms to the specification of local motion by simple cells in areas 17 and 18 of the cat. \newblock {\em Vis.\ Neurosci.}, 11, 295--306. \bibitem[Movshon, 1975]{Movshon75} Movshon, J.~A. (1975). \newblock The velocity tuning of single units in cat striate cortex. \newblock {\em J.\ Physiol.\ (Lond.)}, 249, 445--468. \bibitem[Mukherjee \& Kaplan, 1995]{Mukherjee&Kaplan95} Mukherjee, P. \& Kaplan, E. (1995). \newblock Dynamics of neurons in the cat lateral geniculate nucleus: {I}n vivo electrophysiology and computational modeling. \newblock {\em J.\ Neurophysiol.}, 74, 1222--1243. \bibitem[Murthy \& Humphrey, 1999]{Murthy&Humphrey99} Murthy, A. \& Humphrey, A.~L. (1999). \newblock Inhibitory contributions to spatiotemporal receptive-field structure and direction selectivity in simple cells of cat area 17. \newblock {\em J.\ Neurophysiol.}, 81, 1212--1224. \bibitem[Murthy et~al., 1998]{Murthy_etal98} Murthy, A., Humphrey, A.~L., Saul, A.~B., \& Feidler, J.~C. (1998). \newblock Laminar differences in the spatiotemporal structure of simple cell receptive fields in cat area 17. \newblock {\em Vis.\ Neurosci.}, 15, 239--256. \bibitem[Orban et~al., 1981a]{Orban_etal81b} Orban, G.~A., Kennedy, H., \& Maes, H. (1981a). \newblock Response to movement of neurons in areas 17 and 18 of the cat: Direction selectivity. \newblock {\em J.\ Neurophysiol.}, 45, 1059--1073. \bibitem[Orban et~al., 1981b]{Orban_etal81a} Orban, G.~A., Kennedy, H., \& Maes, H. (1981b). \newblock Response to movement of neurons in areas 17 and 18 of the cat: Velocity sensitivity. \newblock {\em J.\ Neurophysiol.}, 45, 1043--1058. \bibitem[Pape et~al., 1994]{Pape_etal94} Pape, H.~C., Budde, T., Mager, R., \& Kisvarday, Z.~F. (1994). \newblock Prevention of {Ca(2+)}-mediated action potentials in {GABAergic} local circuit neurones of rat thalamus by a transient {K+} current. \newblock {\em J.\ Physiol.\ (Lond.)}, 478, 403--422. \bibitem[Press et~al., 1992]{Press_etal92} Press, W.~H., Teukolsky, S.~A., Vetterling, W.~T., \& Flannery, B.~P. (1992). \newblock {\em Numerical recipes in {C}: {The} art of scientific computing}. \newblock Cambridge University Press. \bibitem[Reid \& Alonso, 1995]{Reid&Alonso95} Reid, R.~C. \& Alonso, J.~M. (1995). \newblock Specificity of monosynaptic connections from thalamus to visual cortex. \newblock {\em Nature}, 378, 281--284. \bibitem[Reid et~al., 1991]{Reid_etal91} Reid, R.~C., Soodak, R.~E., \& Shapley, R.~M. (1991). \newblock Directional selectivity and spatiotemporal structure of receptive fields of simple cells in cat striate cortex. \newblock {\em J.\ Neurophysiol.}, 66, 505--529. \bibitem[Reinagel et~al., 1999]{Reinagel_etal99} Reinagel, P., Godwin, D., Sherman, S.~M., \& Koch, C. (1999). \newblock Encoding of visual information by {LGN} bursts. \newblock {\em J.\ Neurophysiol.}, 81, 2558--2569. \bibitem[Saul \& Humphrey, 1990]{Saul&Humphrey90} Saul, A.~B. \& Humphrey, A.~L. (1990). \newblock Spatial and temporal response properties of lagged and nonlagged cells in the cat lateral genicualte nucleus. \newblock {\em J.\ Neurophysiol.}, 64, 206--224. \bibitem[Saul \& Humphrey, 1992a]{Saul&Humphrey92a} Saul, A.~B. \& Humphrey, A.~L. (1992a). \newblock Evidence of input from lagged cells in the lateral geniculate nucleus to simple cells in cortical area 17 of the cat. \newblock {\em J.\ Neurophysiol.}, 68, 1190--1207. \bibitem[Saul \& Humphrey, 1992b]{Saul&Humphrey92b} Saul, A.~B. \& Humphrey, A.~L. (1992b). \newblock Temporal-frequency tuning of direction selectivity in cat visual cortex. \newblock {\em Vis.\ Neurosci.}, 8, 365--372. \bibitem[Sherman, 1996]{Sherman96} Sherman, S.~M. (1996). \newblock Dual response modes in lateral geniculate neurons: {M}echanisms and functions. \newblock {\em Vis.\ Neurosci.}, 13, 205--213. \bibitem[Sherman \& Guillery, 1996]{Sherman&Guillery96} Sherman, S.~M. \& Guillery, R.~W. (1996). \newblock Functional organization of thalamocortical relays. \newblock {\em J.\ Neurophysiol.}, 76, 1367--1395. \bibitem[Sherman \& Koch, 1986]{Sherman&Koch86} Sherman, S.~M. \& Koch, C. (1986). \newblock The control of retinogeniculate transmission in the mammalian lateral geniculate nucleus. \newblock {\em Exp.\ Brain.\ Res.}, 63, 1--20. \bibitem[Sherman \& Koch, 1990]{Sherman&Koch90} Sherman, S.~M. \& Koch, C. (1990). \newblock Thalamus. \newblock In G.~M. Shepherd (Ed.), {\em The synaptic organization of the brain} (pp.\ 246--278). Oxford University Press. \bibitem[Sillito et~al., 1994]{Sillito_etal94} Sillito, A.~M., Jones, H.~E., Gerstein, G.~L., \& West, D.~C. (1994). \newblock Feature-linked synchronization of thalamic relay cell firing induced by feedback from the visual cortex. \newblock {\em Nature}, 369, 479--482. \bibitem[Singer, 1994]{Singer94} Singer, W. (1994). \newblock A new job for the thalamus. \newblock {\em Nature}, 369, 444--445. \bibitem[Suarez et~al., 1995]{Suarez_etal95} Suarez, H., Koch, C., \& Douglas, R. (1995). \newblock Modeling direction selectivity of simple cells in striate visual cortex within the framework of the canonical microcircuit. \newblock {\em J.\ Neurosci.}, 15, 6700--6719. \bibitem[Toth et~al., 1997]{Toth_etal97} Toth, L.~J., Kim, D.~S., Rao, S.~C., \& Sur, M. (1997). \newblock Integration of local inputs in visual cortex. \newblock {\em Cereb.\ Cortex}, 7, 703--710. \bibitem[Tuckwell, 1988a]{Tuckwell88a} Tuckwell, H.~C. (1988a). \newblock {\em Introduction to theoretical neurobiology, Volume 1: Linear cable theory and dendritic structure}. \newblock Cambridge: Cambridge University Press. \bibitem[Tuckwell, 1988b]{Tuckwell88b} Tuckwell, H.~C. (1988b). \newblock {\em Introduction to theoretical neurobiology, Volume 2: Nonlinear and stochastic theories}. \newblock Cambridge: Cambridge University Press. \bibitem[Valois \& Cottaris, 1998]{DeValois&Cottaris98} Valois, R. L.~D. \& Cottaris, N.~P. (1998). \newblock Inputs to directionally selective simple cells in macaque striate cortex. \newblock {\em Proc.\ Natl.\ Acad.\ Sci.\ USA}, 95, 14488--14493. \bibitem[{von Krosigk} et~al., 1999]{vonKrosigk_etal99} {von Krosigk}, M., Monckton, J.~E., Reiner, P.~B., \& McCormick, D.~A. (1999). \newblock Dynamic properties of corticothalamic excitatory postsynaptic potentials and thalamic reticular inhibitory postsynaptic potentials in thalamocortical neurons of the guinea-pig dorsal lateral geniculate nucleus. \newblock {\em Neurosci.}, 91, 7--20. \bibitem[Watson \& Ahumada, 1985]{Watson&Ahumada85} Watson, A.~B. \& Ahumada, A.~J. (1985). \newblock Model of human visual-motion sensing. \newblock {\em J.\ Opt.\ Soc.\ Am.}, A2, 322--342. \bibitem[Wimbauer et~al., 1997a]{Wimbauer_etal97a} Wimbauer, S., Wenisch, O.~G., Miller, K.~D., \& {van Hemmen}, J.~L. (1997a). \newblock Development of spatiotemporal receptive fields of simple cells: {I}.~{M}odel formulation. \newblock {\em Biol.\ Cybern.}, 77, 453--461. \bibitem[Wimbauer et~al., 1997b]{Wimbauer_etal97b} Wimbauer, S., Wenisch, O.~G., {van Hemmen}, J.~L., \& Miller, K.~D. (1997b). \newblock Development of spatiotemporal receptive fields of simple cells: {II}.~{S}imulation and analysis. \newblock {\em Biol.\ Cybern.}, 77, 463--477. \bibitem[W\"org\"otter et~al., 1998]{Worgotter_etal98} W\"org\"otter, F., Suder, K., Zhao, Y., Kerscher, N., Eysel, U.~T., \& Funke, K. (1998). \newblock State-dependent receptive-field restructuring in the visual cortex. \newblock {\em Nature}, 396, 165--168. citation: Hillenbrand, Dr. Ulrich and van Hemmen, Prof. Dr. J. Leo (2001) Does Corticothalamic Feedback Control Cortical Velocity Tuning? [Journal (Paginated)] document_url: http://cogprints.org/2394/1/Hillenbrand_vanHemmen_01.pdf