Abstract
This chapter studies the second component of the functional architecture of primary visual areas , namely what are called cortico-cortical ‘horizontal’ connections. They join together neurons that detect orientations at different points which are parallel, and even approximately coaxial. This coaxial ‘parallel transport’ has been confirmed by psychophysical experiments on what David Field, Anthony Hayes, , and Robert Hess called the association field. To a first approximation, coaxial parallel transport implements what is known in differential geometry as the contact structure of the fibre bundle \(\mathbb {V}_\mathrm{J}\) of 1 -jets of curves in the plane. It is invariant under the action of the group SE(2) of isometries of the Euclidean plane. The contact structure is associated with a non-commutative group structure on \(\mathbb {V}_\mathrm{J}\), the action of SE(2) then being identified with the left-translations of \(\mathbb {V}_\mathrm{J}\) on itself. This group is isomorphic to the ‘polarized’ Heisenberg group. It is a nilpotent group belonging to the class of what are called Carnot groups. The definition of an SE(2)-invariant metric on the contact planes thus defines what is called a sub-Riemannian geometry. Using this, one obtains a natural explanation for the phenomenon whereby the visual system constructs long-range illusory contours. For illusory contours with curvature, this leads to variational models. At the end of the 1980s, David Mumford already proposed a first model, defined in the plane \(\mathbb {R}^{2}\) of the visual field, which minimized a certain functional of curve length and curvature . We propose a model in the contact bundle \(\mathbb {V}_\mathrm{J}\) that rests on the hypothesis that illusory contours are sub-Riemannian geodesics. The Cartan ‘prolongation’ of this structure, called the Engel structure, corresponds to the 2 -jets of curves in the plane \(\mathbb {R}^{2}\). It must be taken into account if we want to report on the experimental data showing that, in the primary visual system, there exist not only orientation detectors (tangents and 1-jets), but also curvature detectors (osculating circles and 2-jets). We then point out a few properties of the functional architecture of areas V2, V4 (for colour), V5, and MT (for motion). We also discuss Swindale’s model for directions. Finally, we briefly describe the genetic control of the neural morphogenesis and axon guidance of the functional architectures.
This is a preview of subscription content, log in via an institution.
Buying options
Tax calculation will be finalised at checkout
Purchases are for personal use only
Learn about institutional subscriptionsNotes
- 1.
Recall that \(\log \left( 0\right) =-\infty \), so if \(p\in \left[ 0,1\right] \) is small, then \(-\log \left( p\right) \) is a large positive number.
- 2.
\(90\times 90\) is the number of ways to position the little square, 1/2 is the probability of black or white for each pixel, \(\left( 1/2\right) \!{}^{100}\) is the probability that the 100 pixels of the small square are black, and \((1/2){}^{900}\) is the probability that the 900 pixels remaining in the large square are white.
- 3.
Named after George Boole.
- 4.
In the rest of this section, A and B will denote the ends of curves.
- 5.
The reader should not confuse \(\pi \) the projection \(\pi :R\times P\rightarrow R\) and \(\pi \) the component of a tangent vector t.
- 6.
If \(\omega _{1}\) and \(\omega _{2}\) are two 1-forms, \(\omega _{2} \wedge \omega _{1}=-\omega _{1} \wedge \omega _{2}\) (and hence \(\omega \wedge \omega =-\omega \wedge \omega =0\)), and if \(\omega =\mathrm{d}f\) is the differential of a function f(x, y, z), then \(\mathrm{d}\omega =\mathrm{d}^{2}f=0\).
- 7.
The term ‘pacman’ comes from a famous Japanese video game in which a disc with a variable angular sector representing a stylized mouth (the ‘pac-man’) has to find its way through a maze while ‘eating pac-dots’ and at the same time avoiding enemies.
- 8.
Friedrich Engel was one of the main disciples and collaborators of Sophus Lie.
- 9.
There are some technical problems in the definition of the energy, but these are well known in signal theory. To obtain good results, the even/odd filters must be refined, for example, taking the Hilbert transform of \(G_{2}\) instead of \(G_{3} \).
- 10.
For an introduction to structuralism in biology from Waddington to Thom, see [78].
References
Eysel, U.: Turning a corner in vision research. Nature 399, 641–644 (1999)
Alexander, D.M., Bourke, P.D., Sheridan, P., Konstandatos, O., Wright, J.J.: Intrinsic connections in tree shrew \(V1\) imply a global to local mapping. Vision Res. 44, 857–876 (2004)
Ts’o, D.Y., Gilbert, C.D., Wiesel, T.N.: Relationships between horizontal interactions and functional architecture in cat striate cortex as revealed by cross-correlation analysis. J. Neurosci. 6(4), 1160–1170 (1986)
Das, A., Gilbert, C.D.: Long range horizontal connections and their role in cortical reorganization revealed by optical recording of cat primary visual cortex. Nature 375, 780–784 (1995)
Malach, R., Amir, Y., Harel, M., Grinvald, A.: Relationship between intrinsic connections and functional architecture revealed by optical imaging and in vivo targeted biocytin injections in primate striate cortex. Proc. Nat. Acad. Sci. 90, 10469–10473 (1993)
Bosking, W.H., Zhang, Y., Schofield, B., Fitzpatrick, D.: Orientation selectivity and the arrangement of horizontal connections in tree shrew striate cortex. J. Neurosci. 17(6), 2112–2127 (1997)
Mitchison, G., Crick, F.: Long axons within the striate cortex: their distribution, orientation, and patterns of connections. Proc. Nat. Acad. Sci. 79, 3661–3665 (1982)
Rockland, K.S., Lund, J.S.: Widespread periodic intrinsic connections in the tree shrew visual cortex. Science 215, 532–534 (1982)
Rockland, K.S., Lund, J.S.: Intrinsic laminar lattice connections in the primate visual cortex. J. Comp. Neurol. 216(3), 303–318 (1993)
Grinvald, A., Lieke, E.E., Frostig, R.D., Hildesheim, R.: Cortical point-spread function and long-range lateral interactions revealed by real-time optical imaging of macaque monkey primary visual cortex. J. Neurosci. 14(5), 2545–2568 (1994)
Field, D.J., Hayes, A., Hess, R.F.: Contour integration by the human visual system: evidence for a local ‘association field’. Vision Res. 33(2), 173–193 (1993)
Kapadia, M.K., Ito, M., Gilbert, C.D., Westheimer, G.: Improvement in visual sensitivity by changes in local context: parallel studies in human observers and in \(V1\) of alert monkeys. Neuron 15, 843–856 (1995)
Kapadia, M.K., Westheimer, G., Gilbert, C.D.: Dynamics of spatial summation in primary visual cortex of alert monkey. Proc. Nat. Acad. Sci. 96(21), 12073–12078 (1999)
Kovács, I., Julesz, B.: A closed curve is much more than an incomplete one: effect of closure in figure-ground segmentation. Proc. Nat. Acad. Sci. 90, 7495–7497 (1993)
Sajda, P., Han, F.: Perceptual salience as novelty detection in cortical pinwheel space. In: Proceedings of the First International IEEE EMBS Conference on Neural Engineering (2003)
Desolneux, A., Moisan, L., Morel, J.-M.: From Gestalt Theory to Image Analysis: A Probabilistic Approach, Interdisciplinary Applied Mathematics, vol. 34. Springer, Heidelberg (2008)
Kourtzi, Z., Tolias, A.S., Altmann, C.F., Augath, M., Logothetis, N.K.: Integration of local features into global shapes: monkey and human fMRI studies. Neuron 37, 333–346 (2003)
Georges, S., Seriès, P., Frégnac, Y., Lorenceau, J.: Orientation dependent modulation of apparent speed: psychophysical evidence. Vision Res. 42, 2757–2772 (2002)
Georges, S., Seriès, P., Lorenceau, J.: Contrast dependence of high-speed apparent motion. http://www.perceptionweb.com/ecvp00/0300.html (2000)
Frégnac, Y., Baudot, P., Chavane, F., Lorenceau, J., Marre, O., Monier, C., Pananceau, M., Carelli, P., Sadoc, G.: Multiscale functional imaging in \(V1\) and cortical correlates of apparent motion. In: Masson, G.S. Ilg, U.J. (eds.) Dynamics of Visual Motion Processing. Neuronal, Behavioral, and Computational Approaches, pp. 73–94. Springer, Berlin (2010)
Seriès, P., Georges, S., Lorenceau, J., Frégnac, Y.: Orientation dependent modulation of apparent speed: a model based on the dynamics of feed-forward and horizontal connectivity in \(V1\) cortex. Vision Res. 42, 2781–2797 (2002)
Alais, D., Lorenceau, J., Arrighi, R., Cass, J.: Contour interactions between pairs of Gabors engaged in binocular rivalry reveal a map of the association field. Vision Res. 46, 1473–1487 (2006)
Paradis, A.-L., Morel, S., Seriès, P., Lorenceau, J.: Speeding up the brain: when spatial facilitation translates into latency shortening. Frontiers Human Neurosci. 6, 330 (2012)
Petitot, J., Tondut, Y.: Géométrie de contact et champ d’association dans le cortex visuel. CREA reports # 9725, École Polytechnique, Paris (1997)
Lee, T.S., Nguyen, M.: Dynamics of subjective contour formation in the early visual cortex. Proc. Nat. Acad. Sci. 98(4), 1907–1911 (2001)
Polat, U., Sagi, D.: Lateral interactions between spatial channels: suppression and facilitation revealed by lateral masking experiment. Vision Res. 33(7), 993–999 (1993)
Gilbert, C.D., Das, A., Ito, M., Kapadia, M., Westheimer, G.: Spatial integration and cortical dynamics. Proc. Nat. Acad. Sci. 93, 615–622 (1996)
Zucker, S.W., David, C., Dobbins, A., Iverson, L.: The organization of curve detection: coarse tangent fields and fine spline covering. In: Proceedings of the COST-13 Workshop (European Cooperation in Science and Technology), Bonas, France, 1988. Republished in Simon, J.C. (ed.) From Pixels to Features, pp. 75–90. North Holland, New York (1989)
Parent, P., Zucker, S.W.: Trace inference, curvature consistency, and curve detection. IEEE Trans. Pattern Anal. Mach. Intell. II(8), 823–839 (1989)
Grossberg, S., Mingolla, E.: Neural dynamics of form perception: boundary completion, illusory figures and neon color spreading. Psychol. Rev. 92, 173–211 (1985)
Fitzpatrick, D.: Seeing beyond the receptive field in primary visual cortex. Curr. Opin. Neurobiol. 10, 438–443 (2000)
Wright, J.J., Alexander, D.M., Bourke, P.D.: Contribution of lateral interactions in \(V1\) to organization of response properties. Vision Res. 46, 2703–2720 (2006)
Jancke, D., Chavane, F., Naaman, S., Grinvald, A.: Imaging cortical correlates of illusion in early visual cortex. Nature 428, 423–426 (2004)
Rangan, A.V., Cai, D., McLaughlin, D.W.: Modeling the spatiotemporal cortical activity associated with the line-motion illusion in primary visual cortex. Proc. Nat. Acad. Sci. 102(52), 18793–18800 (2005)
Gilbert, C.D., Wiesel, T.N.: Columnar specificity of intrinsic horizontal and corticocortical connections in cat visual cortex. J. Neurosci. 9(7), 2432–2442 (1989)
Gilbert, C.D.: Horizontal integration and cortical dynamics. Neuron 9, 1–13 (1992)
Berthoz, A.: La simplexité. Odile Jacob, Paris (2009)
Petitot, J.: La simplexité de la notion géométrique de jet. In: Berthoz, A., Petit, J.-L. (eds). Simplexité-Complexité. Coll‘ege de France, OpenEdition Books, Paris (2014)
Ullman, S.: Filling in the gaps: the shape of subjective contours and a model for their generation. Biol. Cybern. 25, 1–6 (1976)
Petitot, J., (with a contribution of Tondut, Y.): Vers une Neurogéométrie. Fibrations corticales, structures de contact et contours subjectifs modaux. Mathématiques, Informatique et Sciences Humaines 145, 5–101 (1999)
Bryant, R., Griffiths, P.: Reduction for constrained variational problems and \(\int {\kappa ^{2}}{\rm d}s/2\). Am. J. Math. 108, 525–570 (1986)
Ben-Shahar, O., Zucker, S.: Geometrical computations explain projection patterns of long-range horizontal connections in visual cortex. Neural Comput. 16(3), 445–476 (2004)
Ben-Shahar, O., Huggins, P.S., Izo, T., Zucker, S.: Cortical connections and early visual function: intra- and inter-columnar processing. In: Petitot. J., Lorenceau, J. (eds.) Neurogeometry and Visual Perception. J. Physiol. Paris 97, 191–208 (2003)
Sigman, M., Cecchi, G.A., Gilbert, C.D., Magnasco, M.O.: On a common circle: natural scenes and Gestalt rules. Proc. Nat. Acad. Sci. 98(4), 1935–1940 (2001)
Mallat, S., Peyré, G.: Traitements géométriques des images par bandelettes. In: Journée annuelle de la SMF, 24 June 2006, Mathématiques et Vision, pp. 39–67 (2006)
Le Pennec, E., Mallat, S.: Sparse geometrical image representation with bandelets. In: IEEE Transactions on Image Processing (2003)
Ts’o, D.Y., Zarella, M., Burkitt, G.: Whither the hypercolumn? J. Physiol. 587(12), 2791–2805 (2009)
Roe, A.W.: Modular complexity of area \(V2\) in the macaque monkey. In: Kaas, J.H., Collins, C.E. (eds). The Primate Visual System, Chap. 5, pp. 109–138. CRC Press LLC, Boca Raton, FL (2003)
Sit, Y.F., Miikkulainen, R.: Computational prediction on the receptive fields and organization of \(V2\) for shape processing. Neural Comput. 21(3), 762–785 (2009)
Lu, H.D., Chen, G., Tanigawa, H., Roe, A.W.: A motion direction map in macaque \(V2\). Neuron 68, 1–12 (2010)
Peterhans, E., von der Heydt, R.: Subjective contours: bridging the gap between psychophysics and psychology. Trends Neurosci. 14(3), 112–119 (1991)
von der Heydt, R., Peterhans, E.: Mechanisms of contour perception in monkey visual cortex. I. Lines of pattern discontinuity. J. Neurosci. 9(5), 1731–1748 (1989)
Sheth, B.R., Sharma, J., Rao, S.C., Sur, M.: Orientation maps of subjective contours in visual cortex. Nature 274, 2110–2115 (1996)
Zeki, S.: A Vision of the Brain. Wiley-Blackwell, Oxford (1993)
Thom, R.: Esquisse d’une Sémiophysique. InterEditions, Paris (1988)
Petitot, J.: ‘Le hiatus entre le logique et le morphologique’. Prédication et perception. In: Wildgen, W., Brandt, P.A. (eds). Semiosis and Catastrophes. René Thom’s Semiotic Heritage, pp. 141–166. Peter Lang, Bern (2010)
Shapley, R., Hawken, M.J.: Color in the cortex: single- and double-opponents cells. Vision Res. 51, 701–717 (2011)
Tanigawa, H., Lu, H.D., Roe, A.W.: Functional organization for color and orientation in macaque \(V4\). Nature Neurosci. 13, 1542–1548 (2010)
Thompson, E., Palacios, A., Varela, F.: Ways of coloring: comparative color vision as a case study in cognitive science. Behav. Brain Sci. 15, 1–74 (1992)
Petitot, J.: Morphodynamical enaction: the case of color. Biol. Res. (In: Bacigalupo, J., Palacios, A.G. (eds). A Tribute to Francisco Varela) 36(1), 107–112 (2003)
Gur, M., Snodderly, D.M.: Direction selectivity in \(V1\) of alert monkeys: evidence for parallel pathways for motion processing. J. Physiol. 585(2), 383–400 (2007)
Born, R.T., Bradley, D.C.: Structure and function of visual area \(MT\). Annu. Rev. Neurosci. 28, 157–189 (2005)
DeAngelis, G.C., Newsome, W.T.: Organization of disparity-selective neurons in macaque area MT. J. Neurosci. 19(4), 1398–1415 (1999)
Tanaka, S., Shinbata, H.: Mathematical model for self-organization of direction columns in the primate middle temporal area. Bio. Cybern. 70, 227–234 (1994)
Swindale, N.V., Grinvald, A., Shmuel, A.: The spatial pattern of response magnitude and selectivity for orientation and direction in cat visual cortex. Cereb. Cortex 13(3), 225–235 (2003)
Petitot, J.: Éléments de théorie des singularités. http://jean.petitot.pagesperso-orange.fr/ArticlesPDF_new/Petitot_Sing.pdf (1982)
Prochiantz, A.: Machine-esprit. Odile Jacob, Paris (2001)
Chedotal, A., Richards, L.J.: Wiring the Brain: The Biology of Neuronal Guidance. In: Cold Spring Harbor Perspectives in Biology. http://cshperspectives.cshlp.org (2010)
Kolodkin, A.L., Tessier-Lavigne, M.: Mechanisms and molecules of neuronal wiring: a primer. In: Cold Spring Harbor Perspectives in Biology. http://cshperspectives.cshlp.org (2011)
GeneCards: Human Gene Compendium. Weizmann Institute. www.genecards.org
Reese, B.E.: Development of the retina and optic pathway. Vision Res. 51, 613–632 (2011)
Baye, L.M., Link, B.A.: Nuclear migration during retinal development. Brain Res. 2008, 29–36 (1992)
Ohsawa, R., Kageyama, R.: Regulation of retinal cell fate specification by multiple transcription factors. Brain Res. 1192, 90–98 (2008)
Martins, R.A.P., Pearson, R.A.: Control of cell proliferation by neurotransmitters in the developing vertebrate retina. Brain Res. 1192, 37–60 (2008)
Li, S., Mo, Z., Yang, X., Price, S.M., Shen, M.M., Xiang, M.: Foxn4 controls the genesis of amacrine and horizontal cells by retinal progenitors. Neuron 43, 795–807 (2004)
Schulte, D., Bumsted-O’Brien, K.M.: Molecular mechanisms of vertebrate retina development: Implications for ganglion cell and photorecptor patterning. Brain Res. 1192, 151–164 (2008)
Feldheim, D.A., O’Leary, D.D.: Visual map development: bidirectional signaling, bifunctional guidance, molecules and competition. Cold Spring Harb. Perspect. Biol. http://cshperspectives.cshlp.org (2010)
Petitot, J.: Morphogenèse du Sens, Presses Universitaires de France, Paris, 1985. English Translation: Manjali, F. Morphogenesis of Meaning. Bern, Peter Lang (2003)
McLaughlin, T., Torborg, C.L., Feller, M.B., O’Leary, D.D.M.: Retinotopic map refinement requires spontaneous retinal wave during a brief period of development. Neuron 40, 1147–1160 (2003)
Scicolone, G., Ortalli, A.L., Carri, N.G.: Key roles of Ephs and ephrins in retinotectal topographic map formation. Brain Res. Bull. 79, 227–247 (2009)
Luo, L., Flanagan, J.G.: Development of continuous and discrete neural maps. Neuron 56, 284–300 (2007)
Turing, A.M.: The chemical basis of morphogenesis. Philos. Trans. R. Soc. B, Bio. Sci. 237(641), 37–72 (1952)
Turing, A.M., Wardlaw, C.W.: A diffusion reaction theory of morphogenesis in plants. New Phytol. 52, 40–47. Also in Collected Works of A.M. Turing, P.T. Saunders, Amsterdam 1953, 37–47 (1952)
Cai, A.Q., Landman, K.A., Hughes, B.D.: Modelling directional guidance and motility regulation in cell migration. Bull. Math. Biol. 68, 25–52 (2006)
Author information
Authors and Affiliations
Corresponding author
Rights and permissions
Copyright information
© 2017 Springer International Publishing AG
About this chapter
Cite this chapter
Petitot, J. (2017). Functional Architectures II: Horizontal Connections and Contact Structure. In: Elements of Neurogeometry. Lecture Notes in Morphogenesis. Springer, Cham. https://doi.org/10.1007/978-3-319-65591-8_5
Download citation
DOI: https://doi.org/10.1007/978-3-319-65591-8_5
Published:
Publisher Name: Springer, Cham
Print ISBN: 978-3-319-65589-5
Online ISBN: 978-3-319-65591-8
eBook Packages: Mathematics and StatisticsMathematics and Statistics (R0)