Research Notes in Neural Computing Managing Editor Bart KosKo Editorial Board S. Amari M. A. Arbib C. von der Malsburg Advisory Board Y. Abu-Mostafa A. G. Barto E. Bienenstock J. D. Cowan M. Cynader W. Freeman G. Gross U. an der Heiden M. Hirsch T. Kohonen J. W. Moore L. Optic an A. I. Selverston R. Shapley B. Soffer P. Treleaven W. von Seelen B. Widrow S. Zucker Pablo Rudomin Michael A. Arbib Francisco Cervantes-Perez Ranulfo Romo Editors Neuroscience: From Neural Networks to Artificial Intelligence Proceedings of a U.S.-Mexico Seminar held in the city of Xalapa in the state of Veracruz on December 9-11, 1991 Springer-Verlag Berlin Heidelberg New York London Paris Tokyo Hong Kong Barcelona Budapest Michael A. Arbib Francisco Cervantes-Perez Center for Neural Engineering Centro de Instrumentos University of Southern California Universidad Nacional Aut6noma de Mexico Los Angeles, CA 90089-2520 Apartado Postal 70-183 USA Mexico D.F., CP 04510 Mexico Ranulfo Romo Instituto de Fisiologfa Celular Pablo Rudomin UNAM Centro de Investigaci6n Mexico D.F. y de Estudios Avanzados del I.P.N. Mexico Apartado Postal 14-740 Mexico 07000 D.F. Mexico Managing Editor Bart Kosko Engineering Image Processing Institute University of Southern California University Park Los Angeles, CA 90089-0782 USA ISBN-13:978-3-540-56501-7 e-ISBN-13:978-3-642-78102-5 DOl: 10.1007/978-3-642-78102-5 Library of Congress Cataloging-in-Publication Data Neuroscience: from neural networks to artificial intelligence: proceedings of a U.S. Mexico seminar held in the city of Xalapa in the state of Veracruz on December 9-11, 1991/ edited by Pablo Rudomin ... ret al.l. p. cm. - (Research notes in neural computing; v. 4) Includes bibliographical references and index. ISBN-13:978-3-540-56501-7 I. Neurosciences-Congresses. 2. Neural networks (Computer science )-Congresses. 3. Artificial intelligence-Congresses. I. Rudomin, Pablo, II. Series. QP35l.N4317 1993 006.3-dc20 93-18575 CIP This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustra- tions, recitation, broadcasting, reproduction on microfilms or in any other way, and storage in data banks. Duplication of this publication or parts thereof is permitted only under the provisions of the German Copyright Law of September 9, 1965, in its current version, and permission for use must always be obtained from Springer-Verlag. Violations are liable for prosecution under the German Copyright Law. © Springer-Verlag Berlin Heidelberg 1993 The use of registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. Typesetting: camera-ready by authors 33/3140 -543 2 1 0 -Printed on acid-free paper Introduction The Central Nervous System can be considered as an aggregate of neurons specialized in both the transmission and transformation of information. Information can be used for many purposes, but probably the most important one is to generate a representation of the "external" world that allows the organism to react properly to changes in its external environment. These functions range from such basic ones as detection of changes that may lead to tissue damage and eventual destruction of the organism and the implementation of avoidance reactions, to more elaborate representations of the external world implying recognition of shapes, sounds and textures as the basis of planned action or even reflection. Some of these functions confer a clear survival advantage to the organism (prey or mate recognition, escape reactions, etc.). Others can be considered as an essential part of cognitive processes that contribute, to varying degrees, to the development of individuality and self-consciousness. How can we hope to understand the complexity inherent in this range of functionalities? One of the distinguishing features of the last two decades has been the availability of computational power that has impacted many areas of science. In neurophysiology, computation is used for experiment control, data analysis and for the construction of models that simulate particular systems. Analysis of the behavior of neuronal networks has transcended the limits of neuroscience and is now a discipline in itself, with potential applications both in the neural sciences and in computing sciences. Pattern recognition, robotics and computer vision constitute interesting lines of research which often utilize concepts arising from the neurosciences. However, a limitation of the explosive growth of these closely related disciplines is the decreased communication between the different groups, a decrease potentiated by increasing specialization both in language and methods. Within this general framework it seems necessary to make some effort to cross boundaries and to have active scientists from different backgrounds and disciplines explain the basic principles which guide their investigations, with the hope that this will lay the basis for a future collaborative effort. Reflecting on this, a group of Mexican scientists determined that the Mexican scientific community was ripe for an integrative activity of this kind. It was further VI agreed that this meeting would have the greatest impact if it were organized in the form of a U.S.-Mexico Seminar since this would provide a broader range of disciplinary expertise and ensure the international impact of the proceedings. It should be noted that current work in artificial intelligence has a wide span, from computer vision and robotics to natural language processing and symbolic problem solving. Similarly, neuroscience spans from the finest details of neurochemistry to the findings of the neurological clinic. It was felt that, in each case, there were areas that were not yet ripe for dialog. In the end, we picked sensory perception, motor control and learning in the field of artificial intelligence, and the study of synaptic interactions and the neurophysiology of defined neural circuitry from neuroscience, as the focus for the meeting, and designed the program to ensure a rich dialog between experts in these two areas. Our planning committee discussed the one other area of possible interaction between artiticial intelligence and neuroscience, namely that between modeling of cognitive processes and cognitive neuroscience, but we concluded that the Mexican scientific community did not yet have enough experts in this area to justify its inclusion in a U.S.-Mexico Seminar. On the contrary, there was no problem in identifying a critical mass of distinguished researchers from both the U.S. and Mexico for all the topics we have chosen for the proposed program. The meeting took place in the city of Xalapa in the state of Veracruz on December 9 - 11, 1991, and brought together active neurophysiologists with experts in neuronal network analysis and artificial intelligence to further the integration of their disciplines by reviewing briefly the state of the art in their fields, defining which are the basic principles and concepts in which their disciplines are founded, and providing some perspective on the main problems whose solution will signify an important advance in knowledge. The meeting was held in the archaeological museum of Xalapa, and the record of 5000 years of Mexican cultures provided a stimulating backdrop to a meeting which saw an unusually successful level of interdisciplinary discussion. We have selected about a quarter of that discussion to close the volume, to share with our readers some sense of the lively intellectual exchange which united scientists from very different disciplines in their quest for mutual understanding. It is our hope that the present volume will further this exchange. Based on our experience at the meeting, we have rearranged the order of the papers to provide a more coherent flow of ideas, forming six groups: I. Scales of m. Analysis; II. Processing of sensory information; Visual Processing; IV. Learning and Knowledge Representation; V. Neuronal systems for motor integration; and VI. VII Robotics and Control. To this we have added a new Section VII, a Concluding Perspective which combines an essay on "Methodological Considerations in Cognitive Science" with a summary, "Viewpoints and Controversies", which preserves something of the liveliness (but not the length!) of the discussions at the Xalapa meeting. Each section is preceded by a short introduction, and the reader may wish to turn to these for a more detailed overview of what the book has to offer as it explores, at different scales of analysis, the transition from sensory to motor systems, via the central processes which are continually updated by the processes of learning. We express our thanks to the other members of the Program Committee - Pablo Noriega, Ofelia Cervantes, and Hugo Arechiga - for their help in the planning and organization of the meeting. We would also like to thank the institutions who sponsored the Xalapa meeting: Academia de la Investigacion Cientifica (AIC), Consejo Nacional de Ciencia y Tecnologia (CONACYT), and Instituto Nacional de Estadistica, Geografia e Informatica (INEGI) in Mexico; the National Science Foundation (NSF) in the United States; and UNESCO and Red Latinoamericana de Biologia (RLB) internationally. The Laboratorio Nacional de Informatica Avanzada (LANIA) and Gobierno del Estado de Veracruz provided the local support for the meeting in Xalapa and did much to create a most productive and convivial atmosphere for scientific exchange. July, 1992 Pablo Rudomin Mexico, D.F., and Los Angeles Michael A. Arbib Francisco Cervantes-Perez Ranulfo Romo Table of Contents I. Scales of Analysis......................................................................................... 1 Neuronal networks of the mammalian brain have functionally different classes of neurons: Suggestions for a taxonomy of membrane ionic conductances J. Bargas; E. Galarraga; D.J. Surmeier ............................................................. 3 Electrical coupling in networks containing oscillators E. Marder; L.F. Abbott; A.A. Sharp; N. Kopell ............................................. 33 Dynamical approach to collective brain M. Zak ........................................................................................................... 43 Schema-theoretic models of arm, hand, and eye movements M.A. Arbib ...................................... ;. ............................................................ 61 Cooperative distributed problem solving between (and within) intelligent agents E.H. Durfee .................................................................................................... 84 II. Processing of Sensory Information ............................................................ 99 Spinal processing of impulse trains from sensory receptors L.M. Mendell; H.R. Koerber ...................................................................... 103 Central control of sensory information P. Rudomin ................................................................................................. 116 Parallel and serial processing in the somatosensory system J.H. Kaas .................................................................................................... 136 Cortical representation of touch R. Romo; S. Rufz; P. Crespo ...................................................................... 154 An introduction to human haptic exploration and recognition of objects for neuroscience and AI S. J. Lederman; R. L. Klatzky .................................................................... 171 Common principles in auditory and visual processing S.A. Shamma.............................................................................................. 189 III. Visual Processing..................................................................................... 206 Neuronal substrate of ligth-induced attraction and withdrawal in crayfish: A case of behavioral selection F. Fernandez de Miguel; H. Arechiga ......................................................... 209 Neural and psychophysical models of chromatic and achromatic visual processes E. Martfnez-Uriegas.................................................................................... 231 Computational vision: A probabilistic view of the multi-module paradigm J.L. Marroquin ............................................................................................ 252 State of the art in image processing M. Lee; e.H. Anderson; R. J. Weidner.. ..................................................... 267 Shape recognition in mind, brain, and machine I. Biederman; J.E. Hummel; E.E. Cooper; P.e. Gerhardstein ...................... 282 x IV. Learning And Knowledge Representation ............................................. 294 Contrasting properties of NMDA-dependent and NMDA-independent forms of LTP in hippocampal pyramidal cells R.A. Nicoll; R.A. Zalutsky ........... .... ................................................ .......... 298 Kindling A. Fernandez-Guardiola; R. Gutierrez; A. Martinez; R. Fernandez-Mas ..... 312 Learning automata: An alternative to artificial neural networks A. Sanchez Aguilar ..................................................................................... 326 Learning, from a logical point of view P. Noriega. .................................................................................................. 340 Knowledge representation for speech processing O. Cervantes ............................................................................................... 359 Data management and inference strategies in a human gait pathology expert system G.A. Bekey ................................................................................................. 371 V. Neuronal Systems For Motor Integration ................................................ 385 Entrainment of the spinal neuronal network generating locomotion G. Viana di Prisco; P. Wallen; S. Grillner.. ................................................. 388 Cortical representation of intended movements A.P. Georgopoulos ..................................................................................... 398 Saccadic and fixation sytems of oculomotor control in monkey superior colliculus R. H. Wurtz; D.P. Munoz ........................................................................... 413 Modulatory effects on prey-recognition in amphibia: A theoretical experimental study F. Cervantes-Perez; A. Herrera-Becerra; M. Garcia-Ruiz ........................... 426 VI. Robotics And ControL ............................................................................ 450 Outline for a theory of motor behavior: Involving cooperative actions of the cerebellum, basal ganglia, and cerebral cortex J.C. Houk; S.P. Wise .................................................................................. 452 Neural networks and adaptive control A. G. Barto; V. Gullapalli... ........................................................................ 471 Robustness issues in robot manipulators C. Verde ...................................................................................................... 494 Symbolic planning versus neural control in robots C. Torras Genis .......................................................................................... 509 Divine inheritance vs. experience in the world: Where does the knowledge base come from? E.M. Riseman; A. R. Hanson ...................................................................... 524 VII. A Concluding Perspective .... .... .... .......................................................... 532 Methodological considerations in Cognitive Science N. Lara; F. Cervantes-Perez ........................................................................ 533 Viewpoints and controversies ..................................................................... 546 I. Scales of Analysis The study of how the architectonic characteristics of Central Nervous Systems of different animals might underlie their different physiological properties, which in turn may be postulated to subserve behavioral responses, spans from detailed studies of the ionic conductances of the membrane of a single neuron to neuroethological experiments in freely moving animals, passing through the analysis of the performance shown by small and large neural networks. In the first part of this section we have three papers in this stream. Jose Bargas, Elvira Galarraga, and James Surmeier provide the reader with a feel for the richness and subtlety of processing that may occur within a single neuron. They present a functional taxonomy of membrane ionic conductances and then explain how these conductances modulate the activation dynamics produced by specific input signals on a single neuron, as well as how these conductances underlie various neuronal properties. Persistent or slowly inactivating inward currents produce sustained steady firing, bistable properties, reverse electrotonic decay of synaptic potentials, dendritic or high threshold spikes, slow depolarizing potentials and after-potentials, and neurotransmitter release. This becomes very important if we take into account that in physiology it is assumed that part of the information is coded in the neuronal response frequency. Eve Marder, L.F. Abbott, Andrew Sharp, and Nancy Kopell then show how small networks of such neurons can exhibit intricate dynamics which are subject to a range of chemical modulations. Following a hybrid experimental-theoretical approach, analyze the activation dynamics of a small neural network, and show how an oscillatory network, considered as part of a bigger network and electrically coupled to other circuits with different membrane properties, enhances the variety of dynamic behaviors that may be displayed by the overall network. Through theoretical analysis, they found that the frequency of a network containing an oscillator can be either faster or slower than the frequency of the oscillator, the detailed properties of the oscillator being critically important to control the frequency of the network. This theoretical result guided an experiment in the stomatogastric ganglion where the authors modulated the oscillator response frequency when coupled to a non-oscillatory neuron, showing how the duty cycle of an oscillator can be dynamically regulated by electrically coupled neurons. 2 In the third paper, Michail Zak abstracts away from the detailed properties of neurons to study the collective properties of large networks. He links two different concepts: a) the collective brain, as a set of simple units of processors (neurons) that interact by exchanging signals without explicit global control, and where the objective of each unit may be partially compatible or contradictory, Le., the units may cooperate or compete; and b) a dynamical system, where a dynamical model is represented by a system of ordinary differential equations with terminal attractors and repellers, and it can be implemented only by analog elements. Zak takes advantage of the mathematical tools that have been developed to analyze dynamical systems, to propose a mathematical framework to study a dynamical system which mimics collective purposeful activities of a set of units which process a "knowledge base", in the form of a joint density function, without global control. In this system global coordination is replaced by the probabilistic correlations between the units. These correlations are learned during a long-term period of performing collective tasks, and they are stored in the joint density function. The understanding of the brain requires not only the explanation of phenomena occurring at the level of neural networks dynamics, but also at another different levels of complexity, which include overt animal behavior. In the fourth paper, Michael A. Arbib presents a methodology he defines as "Schema Theory" to attempt to bridge the gap between the different levels of analysis. This methodology offers the possibility of developing studies under three different approaches: "Top-Down", from behavior to neural mechanisms, "Bottom-Up", from neural mechanisms towards explaining behavior, and "Middle-Out", bridging intermediate levels of analysis. Although Arbib focuses on schema-theoretic models of sensorimotor coordination phenomena in living beings, it is clear that the same tools can be used to analyze similar phenomena in the design of complex automata (e.g., sensory based robots). Thus, an important characteristic of Arbib's Schema Theory is that its explanation of intelligent behavior in terms of the interactions among different functional processes provides a bridge between Neuroscience and AI. Finally, to complete this section's progression from membrane properties to the interactions of more and more complex systems, Edmund Durfee offers a point of view from Distributed Artificial Intelligence, showing how intelligent agents may coordinate their activities. Such agents may be like schemas competing and cooperating within the "head" of a single animal or robot, or may indeed be part of a truly social organization bringing together many people (and their machines) to collectively solve problems that are beyond their individual capabilities.