Abstract
Developmental mechanisms for tuning of visual cortex are derived from adult learning mechanisms: an adaptational property of shunting on-center off-surround networks that prevents saturation of parallel processed patterns at high input intensities, a contrast enhancement and short-term memory mechanism, and plastic synaptic strengths that compute a time average of presynaptic signals and postsynaptic activities and multiplicatively gate signals. The mechanisms can generate fields of feature detectors; e.g., line or picture detectors. A developing hierarchy of such fields can be synthesized in which successive critical developmental periods are triggered as a dynamic equilibrium is established between shortterm memory and long-term memory at each stage. Shunting adaptation can account for some data on spatial frequency adaptation. Shunting network properties resemble properties of certain reaction-diffusion systems that have been used to model developmental data in various species; e.g., Hydra, Xenopus retina, slime molds. For example, positional information due to regulation in reaction-diffusion systems is analogous to constancies due to network adaptation, firing of a developmental gradient is analogous to contrast enhancement, and maintenance of a pattern of morphogens is analogous to short-term memory.
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Blakemore,C., Cooper,G.F.: Development of the brain depends on the visual environment. Nature (Lond.). 228, 477–478 (1970)
Creutzfeldt,O.D., Lux,H.D., Nacimiento,A.C.: Intracelluläre Reizung corticaler Nervenzellen. Pflügers Arch. ges. Physiol. 281, 129–151 (1964)
Dickson,R.C., Abelson,J., Barnes,W.M., Reznikoff,W.S.: Genetic regulation: the lac control region. Science 187, 27–35 (1975)
Eccles,J.C.: In: Kimble,G.A. (Ed.): Anatomy of memory, p. 57. Science and Behavior Books, Inc. 1965
Ellias,S.A., Grossberg,S.: Pattern formation, contrast control, and oscillations in the short-term memory of shunting on-center offsurround networks. Biol. Cybernetics 20, 69–98 (1975)
Gaze,R.M.: The formation of nerve connections. London-New York: Academic Press 1970
Gierer,A., Meinhardt,H.: A theory of biological pattern formation. Kybernetik 12, 30–39 (1972)
Grossberg,S.: Nonlinear difference-differential equations in prediction and learning theory. Proc. nat. Acad. Sci. (Wash.). 58, 1329–1334 (1967)
Grossberg,S.: Some networks that can learn, remember, and reproduce any number of complicated space-time patterns. II. Stud. appl. Math. 49, 135–166 (1970a)
Grossberg,S.: Neural pattern discrimination. J. theor. Biol. 27, 291–337 (1970b)
Grossberg,S.: On the dynamics of operant conditioning. J. theor. Biol. 33, 225–255 (1971a)
Grossberg,S.: Pavlovian pattern learning by nonlinear neural networks. Proc. nat. Acad. Sci. (Wash.) 68, 828–831 (1971b)
Grossberg,S.: A neural theory of punishment and avoidance. I. Qualitative theory. Math. Biosci. 15, 39–67 (1972a)
Grossberg,S.: A neural theory of punishment and avoidance. II. Quantitative theory. Math. Biosci. 15, 253–285 (1972b)
Grossberg,S.: Neural expectation: cerebellar and retinal analogs of cells fired by learnable or unlearned pattern classes. Kybernetik 10, 49–57 (1972c)
Grossberg,S.: Pattern learning by functional-differential neural networks with arbitrary path weights. In: Schmitt,K. (Ed.). Delay and functional-differential equations and their applications, pp. 121–160. New York: Academic Press 1972d
Grossberg,S.: Contour enhancement, short-term memory, and constancies in reverberating neural networks. Stud. appl. Math. 52, 213–257 (1973)
Grossberg,S.: Classical and instrumental learning by neural networks. In: Rosen,R., Snell,F. (Eds.). Progress in theoretical biology, pp. 51–141. New York: Academic Press 1974
Grossberg,S.: A neural model of attention, reinforcement, and discrimination learning. In: Pfeiffer,C.C. (Ed.). Int. Rev. Neurobiol. 18 (in press) (1975a)
Grossberg,S.: Adaptive pattern classification and universal recoding: parallel development of neural feature detectors (in preparation) (1975b)
Grossberg,S., Levine,D.S.: Some developmental and attentional biases in the contrast enhancement and short-term memory of recurrent neural networks. J. theor. Biol. (in press) (1975)
Hirsch,H.V.B., Spinelli,D.N.: Visual experience modifies distribution of horizontally and vertically oriented receptive fields in cats. Science 168, 869–871 (1970)
Hodgkin,A.L.: The conduction of the nervous impulse. Springfield, Ill.: C. C. Thomas 1964
Holliday,R., Pugh,J.E.: DNA modification mechanisms and gene activity during development. Science 187, 226–232 (1975)
Hubel,D.H., Wiesel,T.N.: Receptive fields, binocular interaction, and functional architecture in the cat's visual cortex. J. Physiol. (Lond.). 160, 106–154 (1962)
Hubel,D.H., Wiesel,T.N.: Binocular interaction in striate cortex of kittens reared with artificial squint. J. Neurophysiol. 28, 1041–1059 (1965)
Hubel,D.H., Wiesel,T.N.: The period of susceptibility to the physiological effects of unilateral eye closure in kittens. J. Physiol. (Lond.). 206, 419–436 (1970)
Hunt,R.K., Jacobson,M.: Development and stability of positional information in Xenopus retinal ganglion cells. Proc. nat. Acad. Sci. (Wash.). 69, 780–783 (1972)
Hunt,R.K., Jacobson,M.: Specification of positional information in retinal ganglion cells of Xenopus: assay of the unspecified state. Proc. nat. Acad. Sci. (Wash.). 70, 507–511 (1973a)
Hunt,R.K., Jacobson,M.: Neuronal locus specificity: altered pattern of spatial development in fused fragments of embryonic Xenopus eyes. Science 180, 509 (1973b)
Kernell,D.: The adaptation and the relation between discharge frequency and current strength of cat lumbosacral motoneurones stimulated by long-lasting injected currents. Acta physiol. scand. 65, 65–73 (1965a)
Kernell,D.: High-frequency repetitive firing of cat lumbosacral motoneurones stimulated by long-lasting injected currents. Acta Physiol. scand. 65, 74–85 (1965b)
Koenderink,J.J., van de Grind,W.A., Bouman,M.A.: Opponent color coding: a mechanistic model and a new metric for color space. Kybernetik 10, 78–98 (1972)
Kuffer,S.W.: Discharge patterns and functional organization of the mammalian retina. J. Neurophysiol. 16, 37–68 (1953)
Levine, D.S., Grossberg, S.: Visual illusions in neural networks: Line neutralization, tilt aftereffect, and angle expansion. Submitted for publication
Mansfield,R.J.W.: Neural basis of orientation perception in primate vision. Science 186, 1133–1135 (1974)
Mark,R.: Memory and nerve cell connections. New York: Clarendon Oxford University Press 1974
Meinhardt,H., Gierer,A.: Applications of a theory of biological pattern formation based on lateral inhibition. J. Cell Sci. 15, 321–346 (1974)
Pérez,R., Glass,L., Shlaer,R.: Development of specificity in the cat's visual cortex. J. math. Biol. (in press) (1974)
Phillips,G.C.: Actions of antidromic pyramidal volleys on single betz cells in the cat. Quart. J. exp. Physiol. 44, 1–25 (1959)
Rall,W.: Experimental monosynaptic input-output relations in the mammalian spinal cord. J. Cell Comp. Physiol. 46, 413–437 (1955)
Sperling,G.: Model of visual adaptation and contrast detection. Percept. Psychophys. 8, 143–157 (1970)
Sperling,G., Sondhi,M.M.: Model for visual luminance discrimination and flicker detection. J. Optic. Soc. Amer. 58, 1133–1145 (1968)
Stefanis,C.: Interneuronal mechanisms in the cortex. In: Brazier,M. (Ed.): The interneuron, pp. 497–526. Los Angeles: University of Calif. Press 1969
Von der Malsburg,C.: Self-organization of orientation sensitive cells in the striate cortex. Kybernetik 14, 85–100 (1973)
Watson,J.D.: Molecular biology of the gene (2nd ed.). New York: Benjamin 1970
Werblin,F.S.: Adaptation in a vertebrate retina: intracellular recording in Necturus. J. Neurophysiol. 34, 228–241 (1971)
Werblin,F.S., Dowling,J.E.: Organization of the retina of the mudpuppy. Necturus maculosus. II. Intracellular recording. J. Neurophysiol. 32, 339–355 (1969)
Wiesel,T.N., Hubel,D.H.: Single-cell responses in striate cortex of kittens deprived of vision in one eye. J. Neurophysiol. 26, 1003–1017 (1963)
Wiesel,T.N., Hubel,D.H.: Comparison of the effects of unilateral and bilateral eye closure on cortical unit responses in kittens. J. Neurophysiol. 28, 1029–1040 (1965)
Wilson,H.: A synaptic model for spatial frequency adaptation. J. theor. Biol. 49 (in press) (1975)
Wolpert,L.: Positional information and the spatial pattern of cellular differentiation. J. theor. Biol. 25, 1–47 (1969)
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Supported in part by the Advanced Research Projects Agency (DAHC15-73-C-0320) administered by Computer Corporation America.
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Grossberg, S. On the development of feature detectors in the visual cortex with applications to learning and reaction-diffusion systems. Biol. Cybernetics 21, 145–159 (1976). https://doi.org/10.1007/BF00337422
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DOI: https://doi.org/10.1007/BF00337422