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PLOS ONE  2007 

On the Origin of the Functional Architecture of the Cortex

DOI: 10.1371/journal.pone.0000251

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Abstract:

The basic structure of receptive fields and functional maps in primary visual cortex is established without exposure to normal sensory experience and before the onset of the critical period. How the brain wires these circuits in the early stages of development remains unknown. Possible explanations include activity-dependent mechanisms driven by spontaneous activity in the retina and thalamus, and molecular guidance orchestrating thalamo-cortical connections on a fine spatial scale. Here I propose an alternative hypothesis: the blueprint for receptive fields, feature maps, and their inter-relationships may reside in the layout of the retinal ganglion cell mosaics along with a simple statistical connectivity scheme dictating the wiring between thalamus and cortex. The model is shown to account for a number of experimental findings, including the relationship between retinotopy, orientation maps, spatial frequency maps and cytochrome oxidase patches. The theory's simplicity, explanatory and predictive power makes it a serious candidate for the origin of the functional architecture of primary visual cortex.

References

[1]  Hubel DH, Wiesel TN (1959) Receptive Fields of Single Neurones in the Cats Striate Cortex. Journal of Physiology-London 148: 574–591.
[2]  Hubel DH, Wiesel TN (1962) Receptive Fields, Binocular Interaction and Functional Architecture in Cats Visual Cortex. Journal of Physiology-London 160: 106–&.
[3]  Mountcastle VB (1957) Modality and Topographic Properties of Single Neurons of Cats Somatic Sensory Cortex. Journal of Neurophysiology 20: 408–434.
[4]  Mountcastle VB (1997) The columnar organization of the neocortex. Brain 120: 701–722.
[5]  Grinvald A, Lieke E, Frostig RD, Gilbert CD, Wiesel TN (1986) Functional Architecture of Cortex Revealed by Optical Imaging of Intrinsic Signals. Nature 324: 361–364.
[6]  Blasdel GG, Salama G (1986) Voltage-Sensitive Dyes Reveal a Modular Organization in Monkey Striate Cortex. Nature 321: 579–585.
[7]  Bonhoeffer T, Grinvald A (1991) Iso-Orientation Domains in Cat Visual-Cortex Are Arranged in Pinwheel-Like Patterns. Nature 353: 429–431.
[8]  Weliky M, Bosking WH, Fitzpatrick D (1996) A systematic map of direction preference in primary visual cortex. Nature 379: 725–728.
[9]  Basole A, White LE, Fitzpatrick D (2003) Mapping multiple features in the population response of visual cortex. Nature 423: 986–990.
[10]  Issa NP, Trepel C, Stryker MP (2000) Spatial frequency maps in cat visual cortex. Journal of Neuroscience 20: 8504–8514.
[11]  Crair MC, Ruthazer ES, Gillespie DC, Stryker MP (1997) Relationship between the ocular dominance and orientation maps in visual cortex of monocularly deprived cats. Neuron 19: 307–318.
[12]  Bartfeld E, Grinvald A (1992) Relationships between Orientation-Preference Pinwheels, Cytochrome-Oxidase Blobs, and Ocular-Dominance Columns in Primate Striate Cortex. Proceedings of the National Academy of Sciences of the United States of America 89: 11905–11909.
[13]  Tso DY, Frostig RD, Lieke EE, Grinvald A (1990) Functional-Organization of Primate Visual-Cortex Revealed by High-Resolution Optical Imaging. Science 249: 417–420.
[14]  Bonhoeffer T, Grinvald A (1993) The Layout of Iso-Orientation Domains in Area-18 of Cat Visual-Cortex - Optical Imaging Reveals a Pinwheel-Like Organization. Journal of Neuroscience 13: 4157–4180.
[15]  Maldonado PE, Godecke I, Gray CM, Bonhoeffer T (1997) Orientation selectivity in pinwheel centers in cat striate cortex. Science 276: 1551–1555.
[16]  Rao SC, Toth LJ, Sur M (1997) Optically imaged maps of orientation preference in primary visual cortex of cats and ferrets. Journal of Comparative Neurology 387: 358–370.
[17]  Shmuel A, Grinvald A (2000) Coexistence of linear zones and pinwheels within orientation maps in cat visual cortex. Proceedings of the National Academy of Sciences of the United States of America 97: 5568–5573.
[18]  Everson R, Kaplan E, Knight B, Obrien EV, Orbach D, et al. (1995) Functional-Organization of Orientation, Direction, and Binocular Patches in Cat and Macaque Visual-Cortex Revealed by Optical Imaging. Investigative Ophthalmology & Visual Science 36: S873–S873.
[19]  Landisman CE, Ts'o DY (2002) Color processing in macaque striate cortex: Relationships to ocular dominance, cytochrome oxidase, and orientation. Journal of Neurophysiology 87: 3126–3137.
[20]  Xiao YP, Wang Y, Felleman DJ (2003) A spatially organized representation of colour in macaque cortical area V2. Nature 421: 535–539.
[21]  Blasdel G, Campbell D (2001) Functional retinotopy of monkey visual cortex. Journal of Neuroscience 21: 8286–8301.
[22]  Obermayer K, Blasdel GG (1997) Singularities in primate orientation maps. Neural Computation 9: 555–575.
[23]  Blasdel GG (1992) Orientation Selectivity, Preference, and Continuity in Monkey Striate Cortex. Journal of Neuroscience 12: 3139–3161.
[24]  Ohki K, Chung SY, Kara P, Hubener M, Bonhoeffer T, et al. (2006) Highly ordered arrangement of single neurons in orientation pinwheels. Nature 442: 925–928.
[25]  Ohki K, Chung S, Ch'ng YH, Kara P, Reid RC (2005) Functional imaging with cellular resolution reveals precise micro-architecture in visual cortex. Nature 433: 597–603.
[26]  Horton JC, Adams DL (2005) The cortical column: a structure without a function. Philosophical Transactions of the Royal Society B-Biological Sciences 360: 837–862.
[27]  Purves D, Riddle DR, Lamantia AS (1992) Iterated Patterns of Brain Circuitry (or How the Cortex Gets Its Spots). Trends in Neurosciences 15: 362–368.
[28]  Swindale NV (1990) Is the Cerebral-Cortex Modular. Trends in Neurosciences 13: 487–492.
[29]  Soodak RE (1987) The Retinal Ganglion-Cell Mosaic Defines Orientation Columns in Striate Cortex. Proceedings of the National Academy of Sciences of the United States of America 84: 3936–3940.
[30]  Ringach DL (2004) Haphazard wiring of simple receptive fields and orientation columns in visual cortex. Journal of Neurophysiology 92: 468–476.
[31]  Swindale NV, Grinvald A, Shmuel A (2003) The spatial pattern of response magnitude and selectivity for orientation and direction in cat visual cortex. Cerebral Cortex 13: 225–238.
[32]  Muller T, Stetter M, Hubener M, Sengpiel E, Bonhoeffer T, et al. (2000) An analysis of orientation and ocular dominance patterns in the visual cortex of cats and ferrets. Neural Computation 12: 2573–2595.
[33]  Hubener M, Shoham D, Grinvald A, Bonhoeffer T (1997) Spatial relationships among three columnar systems in cat area 17. Journal of Neuroscience 17: 9270–9284.
[34]  Shoham D, Hubener M, Schulze S, Grinvald A, Bonhoeffer T (1997) Spatio-temporal frequency domains and their relation to cytochrome oxidase staining in cat visual cortex. Nature 385: 529–533.
[35]  Shmuel A, Grinvald A (1996) Functional organization for direction of motion and its relationship to orientation maps in cat area 18. Journal of Neuroscience 16: 6945–6964.
[36]  Crair MC, Gillespie DC, Stryker MP (1998) The role of visual experience in the development of columns in cat visual cortex. Science 279: 566–570.
[37]  Crair MC, Ruthazer ES, Gillespie DC, Stryker MP (1997) Ocular dominance peaks at pinwheel center singularities of the orientation map in cat visual cortex. Journal of Neurophysiology 77: 3381–3385.
[38]  Ruthazer ES, Stryker MP (1996) The role of activity in the development of long-range horizontal connections in area 17 of the ferret. Journal of Neuroscience 16: 7253–7269.
[39]  Schmidt KF, Lowel S (2006) The layout of functional maps in area 18 of strabismic cats. Neuroscience 141: 1525–1531.
[40]  Schmidt KF, Lowel S (2006) Optical imaging in cat area 18: Strabismus does not enhance the segregation of ocular dominance domains. Neuroimage 29: 439–445.
[41]  Kaschube M, Wolf F, Geisel T, Lowel S (2002) Genetic influence on quantitative features of neocortical architecture. Journal of Neuroscience 22: 7206–7217.
[42]  Wolf F, Pawelzik K, Scherf O, Geisel T, Lowel S (2000) How can squint change the spacing of ocular dominance columns? Journal of Physiology-Paris 94: 525–537.
[43]  Movshon JA, Vansluyters RC (1981) Visual Neural Development. Annual Review of Psychology 32: 477–522.
[44]  Fregnac Y, Imbert M (1984) Development of Neuronal Selectivity in Primary Visual-Cortex of Cat. Physiological Reviews 64: 325–434.
[45]  Alonso JM, Usrey WM, Reid RC (2001) Rules of connectivity between geniculate cells and simple cells in cat primary visual cortex. Journal of Neuroscience 21: 4002–4015.
[46]  Reid RC, Alonso JM (1995) Specificity of Monosynaptic Connections from Thalamus to Visual-Cortex. Nature 378: 281–284.
[47]  Wassle H, Peichl L, Boycott BB (1981) Morphology and Topography of on-Alpha and Off-Alpha Cells in the Cat Retina. Proceedings of the Royal Society of London Series B-Biological Sciences 212: 157–&.
[48]  Zhan XJ, Troy JB (2000) Modeling cat retinal beta-cell arrays. Visual Neuroscience 17: 23–39.
[49]  Wassle H, Boycott BB, Illing RB (1981) Morphology and Mosaic of on-Beta and Off-Beta Cells in the Cat Retina and Some Functional Considerations. Proceedings of the Royal Society of London Series B-Biological Sciences 212: 177–&.
[50]  Eglen SJ (2006) Development of regular cellular spacing in the retina: theoretical models. Mathematical Medicine and Biology-a Journal of the Ima 23: 79–99.
[51]  Eglen SJ, Diggle PJ, Troy JB (2005) Homotypic constraints dominate positioning of on- and off-center beta retinal ganglion cells. Visual Neuroscience 22: 859–871.
[52]  Hubel DH, Wiesel TN (1963) Receptive Fields of Cells in Striate Cortex of Very Young, Visually Inexperienced Kittens. Journal of Neurophysiology 26: 994–&.
[53]  Albus K, Wolf W (1984) Early Post-Natal Development of Neuronal Function in the Kittens Visual-Cortex - a Laminar Analysis. Journal of Physiology-London 348: 153–185.
[54]  Sherk H, Stryker MP (1976) Quantitative Study of Cortical Orientation Selectivity in Visually Inexperienced Kitten. Journal of Neurophysiology 39: 63–70.
[55]  Crair MC, Horton JC, Antonini A, Stryker MP (2001) Emergence of ocular dominance columns in cat visual cortex by 2 weeks of age. Journal of Comparative Neurology 430: 235–249.
[56]  Miller KD (1994) A Model for the Development of Simple Cell Receptive-Fields and the Ordered Arrangement of Orientation Columns through Activity-Dependent Competition between on- and Off-Center Inputs. Journal of Neuroscience 14: 409–441.
[57]  Kayser AS, Miller KD (2002) Opponent inhibition: A developmental model of layer 4 of the neocortical circuit. Neuron 33: 131–142.
[58]  Swindale NV (1996) The development of topography in the visual cortex: A review of models. Network-Computation in Neural Systems 7: 161–247.
[59]  Miller KD, Erwin E, Kayser A (1999) Is the development of orientation selectivity instructed by activity? Journal of Neurobiology 41: 44–57.
[60]  Dayan P (2003) Pattern formation and cortical maps. Journal of Physiology-Paris 97: 475–489.
[61]  Kohonen T (1985) Self-Organized Feature Maps. Journal of the Optical Society of America a-Optics Image Science and Vision 2: P16–P16.
[62]  Malsburg CV (1973) Self-Organization of Orientation Sensitive Cells in Striate Cortex. Kybernetik 14: 85–100.
[63]  Malsburg CV (1973) Model for Self-Organization of Orientation Sensitivity and Columns in Visual-Cortex. Pflugers Archiv-European Journal of Physiology 339: 95–95.
[64]  Goodhill GJ, Cimponeriu A (2000) Analysis of the elastic net model applied to the formation of ocular dominance and orientation columns. Network-Computation in Neural Systems 11: 153–168.
[65]  Goodhill GJ, Willshaw DJ (1994) Elastic Net Model of Ocular Dominance - Overall Stripe Pattern and Monocular Deprivation. Neural Computation 6: 615–621.
[66]  Miller KD (1994) Models of Activity-Dependent Neural Development. Self-Organizing Brain: From Growth Cones to Functional Networks 102: 303–318.
[67]  Miller KD (1992) Development of Orientation Columns Via Competition between on-Center and Off-Center Inputs. Neuroreport 3: 73–76.
[68]  Crowley JC, Katz LC (2002) Ocular dominance development revisited. Current Opinion in Neurobiology 12: 104–109.
[69]  Yuste R, Nelson DA, Rubin WW, Katz LC (1995) Neuronal Domains in Developing Neocortex - Mechanisms of Coactivation. Neuron 14: 7–17.
[70]  Wong ROL, Meister M, Shatz CJ (1993) Transient Period of Correlated Bursting Activity during Development of the Mammalian Retina. Neuron 11: 923–938.
[71]  Weliky M (1999) Recording and manipulating the in vivo correlational structure of neuronal activity during visual cortical development. Journal of Neurobiology 41: 25–32.
[72]  Shatz CJ (1996) Emergence of order in visual system development. Proceedings of the National Academy of Sciences of the United States of America 93: 602–608.
[73]  Shatz CJ (1994) Role for Spontaneous Neural Activity in the Patterning of Connections between Retina and Lgn during Visual-System Development. International Journal of Developmental Neuroscience 12: 531–546.
[74]  Price DJ, Kennedy H, Dehay C, Zhou LB, Mercier M, et al. (2006) The development of cortical connections. European Journal of Neuroscience 23: 910–920.
[75]  Skaliora I, Adams R, Blakemore C (2000) Morphology and growth patterns of developing thalamocortical axons. Journal of Neuroscience 20: 3650–3662.
[76]  Sur M, Leamey CA (2001) Development and plasticity of cortical areas and networks. Nature Reviews Neuroscience 2: 251–262.
[77]  Crowley JC, Katz LC (2000) Early development of ocular dominance columns. Science 290: 1321–1324.
[78]  Crowley JC, Katz LC (1999) Development of ocular dominance columns in the absence of retinal input. Nature Neuroscience 2: 1125–1130.
[79]  Stein JJ, Johnson SA, Berson DM (1996) Distribution and coverage of beta cells in the cat retina. J Comp Neurol 372: 597–617.
[80]  Cleland BG, Dubin MW, Levick WR (1971) Simultaneous Recording of Input and Output of Lateral Geniculate Neurones. Nature-New Biology 231: 191–&.
[81]  Cleland BG, Lee BB (1985) A Comparison of Visual Responses of Cat Lateral Geniculate-Nucleus Neurons with Those of Ganglion-Cells Afferent to Them. Journal of Physiology-London 369: 249–268.
[82]  Usrey WM, Reppas JB, Reid RC (1999) Specificity and strength of retinogeniculate connections. Journal of Neurophysiology 82: 3527–3540.
[83]  Ringach DL (2002) Spatial structure and symmetry of simple-cell receptive fields in macaque primary visual cortex. Journal of Neurophysiology 88: 455–463.
[84]  Jones JP, Palmer LA (1987) An evaluation of the two-dimensional Gabor filter model of simple receptive fields in cat striate cortex. J Neurophysiol 58: 1233–1258.
[85]  Movshon JA, Thompson ID, Tolhurst DJ (1978) Spatial Summation in Receptive-Fields of Simple Cells in Cats Striate Cortex. Journal of Physiology-London 283: 53–77.
[86]  Chapman B, Godecke I (2000) Cortical cell orientation selectivity fails to develop in the absence of ON-center retinal ganglion cell activity. Journal of Neuroscience 20: 1922–1930.
[87]  Wassle H, Grunert U, Rohrenbeck J, Boycott BB (1989) Cortical Magnification Factor and the Ganglion-Cell Density of the Primate Retina. Nature 341: 643–646.
[88]  Das A, Gilbert CD (1997) Distortions of visuotopic map match orientation singularities in primary visual cortex. Nature 387: 594–598.
[89]  Buzas P, Volgushev M, Eysel UT, Kisvarday ZF (2003) Independence of visuotopic representation and orientation map in the visual cortex of the cat. European Journal of Neuroscience 18: 957–968.
[90]  Bosking WH, Crowley JC, Fitzpatrick D (2002) Spatial coding of position and orientation in primary visual cortex. Nature Neuroscience 5: 874–882.
[91]  Yu H, Farley BJ, Jin DZ, Sur M (2005) The coordinated mapping of visual space and response features in visual cortex. Neuron 47: 267–280.
[92]  Ernst U, Pawelzik K, Tsodyks M, Sejnowski TJ (1999) Relation between retinotopical and orientation maps in visual cortex. Neural Computation 11: 375–379.
[93]  Marino J, Schummers J, Lyon DC, Schwabe L, Beck O, et al. (2005) Invariant computations in local cortical networks with balanced excitation and inhibition. Nature Neuroscience 8: 194–201.
[94]  Schummers J, Marino J, Sur M (2004) Local networks in visual cortex and their influence on neuronal responses and dynamics. Journal of Physiology-Paris 98: 429–441.
[95]  Schummers J, Marino J, Sur M (2002) Synaptic integration by V1 neurons depends on location within the orientation map. Neuron 36: 969–978.
[96]  McLaughlin D, Shapley R, Shelley M (2003) Large-scale modeling of the primary visual cortex: influence of cortical architecture upon neuronal response. Journal of Physiology-Paris 97: 237–252.
[97]  Xing DJ, Ringach DL, Shapley R, Hawken MJ (2004) Correlation of local and global orientation and spatial frequency tuning in macaque V1. Journal of Physiology-London 557: 923–933.
[98]  Horton JC (1984) Cytochrome-Oxidase Patches - a New Cytoarchitectonic Feature of Monkey Visual-Cortex. Philosophical Transactions of the Royal Society of London Series B-Biological Sciences 304: 199–253.
[99]  Murphy KM, Vansluyters RC, Jones DG (1991) Cytochrome-Oxidase Blobs in Cat Visual-Cortex. Investigative Ophthalmology & Visual Science 32: 1116–1116.
[100]  Murphy KM, Jones DG, Vansluyters RC (1995) Cytochrome-Oxidase Blobs in Cat Primary Visual-Cortex. Journal of Neuroscience 15: 4196–4208.
[101]  Murphy KM, Duffy KR, Jones DG, Mitchell DE (2001) Development of cytochrome oxidase blobs in visual cortex of normal and visually deprived cats. Cerebral Cortex 11: 122–135.
[102]  Adams DL, Horton JC (2003) The representation of retinal blood vessels in primate striate cortex. Journal of Neuroscience 23: 5984–5997.
[103]  Adams DL, Horton JC (2003) A precise retinotopic map of primate striate cortex generated from the representation of angioscotomas. Journal of Neuroscience 23: 3771–3789.
[104]  Sakitt B (1982) Why the Cortical Magnification Factor in Rhesus Can Not Be Isotropic. Vision Research 22: 417–421.
[105]  Matsuda Y, Ohki K, Saito T, Ajima A, Kim DS (2000) Coincidence of ipsilateral ocular dominance peaks with orientation pinwheel centers in cat visual cortex. Neuroreport 11: 3337–3343.
[106]  Ohzawa I, DeAngelis GC, Freeman RD (1996) Encoding of binocular disparity by simple cells in the cat's visual cortex. Journal of Neurophysiology 75: 1779–1805.
[107]  Erwin E, Miller KD (1998) Correlation-based development of ocularly matched orientation and ocular dominance maps: Determination of required input activities. Journal of Neuroscience 18: 9870–9895.
[108]  Weliky M, Katz LC (1999) Correlational structure of spontaneous neuronal activity in the developing lateral geniculate nucleus in vivo. Science 285: 599–604.
[109]  Van Hooser SD, Heimel JAF, Chung S, Nelson SB, Toth LJ (2005) Orientation selectivity without orientation maps in visual cortex of a highly visual mammal. Journal of Neuroscience 25: 19–28.
[110]  Niebur E, Worgotter F (1994) Design Principles of Columnar Organization in Visual-Cortex. Neural Computation 6: 602–614.
[111]  Miller KD (1998) Equivalence of a sprouting-and-retraction model and correlation-based plasticity models of neural development. Neural Computation 10: 529–547.
[112]  Miller KD, Erwin E (2001) Effects of monocular deprivation and reverse suture on orientation maps can be explained by activity-instructed development of geniculocortical connections. Visual Neuroscience 18: 821–834.
[113]  Miller KD (2003) Understanding layer 4 of the cortical circuit: A model based on cat V1. Cerebral Cortex 13: 73–82.
[114]  Soodak RE (1991) Reverse-Hebb Plasticity Leads to Optimization and Association in a Simulated Visual-Cortex. Visual Neuroscience 6: 507–518.
[115]  Miller KD, Mackay DJC (1994) The Role of Constraints in Hebbian Learning. Neural Computation 6: 100–126.
[116]  Obermayer K, Sejnowski T, Blasdel GG (1995) Neural Pattern-Formation Via a Competitive Hebbian Mechanism. Behavioural Brain Research 66: 161–167.
[117]  Wimbauer S, Wenisch OG, Miller KD, van Hemmen JL (1997) Development of spatiotemporal receptive fields of simple cells .1. Model formulation. Biological Cybernetics 77: 453–461.
[118]  Andrade MA, Moran F (1997) Receptive field map development by anti-Hebbian learning. Neural Networks 10: 1037–1052.
[119]  Bartsch AP, van Hemmen JL (2001) Combined Hebbian development of geniculocortical and lateral connectivity in a model of primary visual cortex. Biological Cybernetics 84: 41–55.
[120]  Bednar JA, Miikkulainen R (2006) Joint maps for orientation, eye, and direction preference in a self-organizing model of V1. Neurocomputing 69: 1272–1276.
[121]  Ohshiro T, Weliky M (2006) Simple fall-off pattern of correlated neural activity in the developing lateral geniculate nucleus. Nat Neurosci 9: 1541–1548.
[122]  Durbin R, Mitchison G (1990) A Dimension Reduction Framework for Understanding Cortical Maps. Nature 343: 644–647.
[123]  Obermayer K, Blasdel GG, Schulten K (1992) Statistical-Mechanical Analysis of Self-Organization and Pattern-Formation during the Development of Visual Maps. Physical Review A 45: 7568–7589.
[124]  Swindale NV (2000) How many maps are there in visual cortex? Cerebral Cortex 10: 633–643.
[125]  Swindale NV (2004) How different feature spaces may be represented in cortical maps. Network-Computation in Neural Systems 15: 217–242.
[126]  Kohonen T (1982) Self-Organized Formation of Topologically Correct Feature Maps. Biological Cybernetics 43: 59–69.
[127]  Hubener M, Grinvald A, Shoham D, Bonhoeffer T, Swindale NV (2000) Coverage optimization as a principle for the arrangement of functional maps in the visual cortex. European Journal of Neuroscience 12: 194–194.
[128]  Swindale NV (1991) Coverage and the Design of Striate Cortex. Biological Cybernetics 65: 415–424.
[129]  Swindale NV, Shoham D, Grinvald A, Bonhoeffer T, Hubener M (2000) Visual cortex maps are optimized for uniform coverage. Nature Neuroscience 3: 822–826.
[130]  Cowey A (1979) Cortical Maps and Visual-Perception - Grindley Memorial Lecture. Quarterly Journal of Experimental Psychology 31: 1–17.
[131]  Chklovskii DB, Koulakov AA (2004) Maps in the brain: What can we learn from them? Annual Review of Neuroscience 27: 369–392.
[132]  Chklovskii DB, Koulakov AA (2000) A wire length minimization approach to ocular dominance patterns in mammalian visual cortex. Physica a-Statistical Mechanics and Its Applications 284: 318–334.
[133]  Thomas PJ, Cowan JD (2006) Simultaneous constraints on pre- and post-synaptic cells couple cortical feature maps in a 2D geometric model of orientation preference. Mathematical Medicine and Biology-a Journal of the Ima 23: 119–138.
[134]  Naegele JR, Jhaveri S, Schneider GE (1988) Sharpening of topographical projections and maturation of geniculocortical axon arbors in the hamster. J Comp Neurol 277: 593–607.
[135]  Kanold PO, Kara P, Reid RC, Shatz CJ (2003) Role of subplate neurons in functional maturation of visual cortical columns. Science 301: 521–525.
[136]  Ferster D, Levay S (1978) Axonal Arborizations of Lateral Geniculate Neurons in Striate Cortex of Cat. Journal of Comparative Neurology 182: 923–944.
[137]  Humphrey AL, Sur M, Uhlrich DJ, Sherman SM (1985) Termination Patterns of Individual X-Cell and Y-Cell Axons in the Visual-Cortex of the Cat - Projections to Area-18, to the 17/18 Border Region, and to Both Area-17 and Area-18. Journal of Comparative Neurology 233: 190–212.
[138]  Weng C, Yeh CI, Stoelzel CR, Alonso JM (2005) Receptive field size and response latency are correlated within the cat visual thalamus. Journal of Neurophysiology 93: 3537–3547.
[139]  Yeh CI, Stoelzel CR, Alonso JM (2003) Two different types of Y cells in the cat lateral geniculate nucleus. Journal of Neurophysiology 90: 1852–1864.
[140]  Daniels JD, Pettigrew JD, Norman JL (1978) Development of Single-Neuron Responses in Kittens Lateral Geniculate-Nucleus. Journal of Neurophysiology 41: 1373–1393.
[141]  Ikeda H, Tremain KE (1978) Development of Spatial Resolving Power of Lgn Cells and Its Susceptibility to Blur and Strabismus. Archives Italiennes De Biologie 116: 375–384.
[142]  Ikeda H, Tremain KE (1978) Development of Spatial Resolving Power of Lateral Geniculate Neurons in Kittens. Experimental Brain Research 31: 193–206.
[143]  Tavazoie SF, Reid RC (2000) Diverse receptive fields in the lateral geniculate nucleus during thalamocortical development. Nature Neuroscience 3: 608–616.
[144]  Ahmed B, Hammond P (1991) Orientation Bias of Cat Dorsal Lateral Geniculate Cells - Directional Analysis of the Major Axis of the Receptive-Field Center. Experimental Brain Research 84: 676–679.
[145]  Schall JD, Leventhal AG (1987) Relationships between Ganglion-Cell Dendritic Structure and Retinal Topography in the Cat. Journal of Comparative Neurology 257: 149–159.
[146]  Schall JD, Perry VH, Leventhal AG (1986) Retinal Ganglion-Cell Dendritic Fields in Old-World Monkeys Are Oriented Radially. Brain Research 368: 18–23.
[147]  Leventhal AG, Schall JD (1983) Structural Basis of Orientation Sensitivity of Cat Retinal Ganglion-Cells. Journal of Comparative Neurology 220: 465–475.
[148]  Sasaki Y, Rajimehr R, Kim BW, Ekstrom LB, Vanduffel W, et al. (2006) The Radial Bias: A Different Slant on Visual Orientation Sensitivity in Human and Nonhuman Primates. Neuron 51: 661–670.
[149]  Goodhill GJ, Bates KR, Montague PR (1997) Influences on the global structure of cortical maps. Proceedings of the Royal Society of London Series B-Biological Sciences 264: 649–655.
[150]  Jones DG, Van Sluyters RC, Murphy KM (1991) A computational model for the overall pattern of ocular dominance. J Neurosci 11: 3794–3808.
[151]  Wolf F, Bauer HU, Pawelzik K, Geisel T (1996) Organization of the visual cortex. Nature 382: 306–307.
[152]  Li Y, Fitzpatrick D, White LE (2006) The development of direction selectivity in ferret visual cortex requires early visual experience. Nature Neuroscience 9: 676–681.
[153]  Blumenfeld B, Bibitchkov D, Tsodyks M (2006) Neural network model of the primary visual cortex: From functional architecture to lateral connectivity and back. Journal of Computational Neuroscience 20: 219–241.
[154]  Ernst UA, Pawelzik KR, Sahar-Pikielny C, Tsodyks MV (2001) Intracortical origin of visual maps. Nature Neuroscience 4: 431–436.
[155]  Mooser F, Bosking WH, Fitzpatrick D (2004) A morphological basis for orientation tuning in primary visual cortex. Nature Neuroscience 7: 872–879.
[156]  Carandini M, Ferster D (2000) Membrane potential and firing rate in cat primary visual cortex. Journal of Neuroscience 20: 470–484.
[157]  Boycott BB, Wassle H (1974) Morphological Types of Ganglion-Cells of Domestic Cats Retina. Journal of Physiology-London 240: 397–&.
[158]  Peters A, Payne BR (1993) Numerical Relationships between Geniculocortical Afferents and Pyramidal Cell Modules in Cat Primary Visual-Cortex. Cerebral Cortex 3: 69–78.
[159]  Olshausen BA, Field DJ (1996) Natural image statistics and efficient coding. Network-Computation in Neural Systems 7: 333–339.

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