- Email: [email protected]
Methods in Neuronal Modelling. From Synapses to Networks
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Computational Neurosciences Laboratory, Departmentof Psychiatry, New York UniversityMedical Center,550 First Avenue, New York, NY 10016, USA,and Departmentof ComputerScience, Courantlnstitute, New YorkUniversity, WashinEtonSquare, New York,NY 10003, USA.
436
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.
and neuroscientific perspectives on PDP modelling. In principle, PDP models allows a bridge to be built between the cognitive sciences and the neurosciences. From reading the book it is evident that something has been accomplished in bringing together the psychologist who, until recently, was explaining complex cognitive functioning solely in terms of condition-action rules or as the formal manipulation of symbols, and the neuroscientist, who painstakingly recorded singleneurone activity with a microelectrode or analysed the biochemical activity at synaptic clefts. However, it also becomes
clear that much more is needed. There is still a large gap between the psychologist developing neurally inspired networks to simulate learning of the past tense of English verbs, and the neuroscientist exploring the behaviour of neurally constrained networks and speculating about the possible role of the hippocampus in cognitive functioning. If our goal is to understand the brain and to fill the gap between the algorithmic and the implementational levels of describing cognition, PDP modellers from psychology and the neurosciences still have a lot to learn from each other; this book will assist in this process.
it becomes necessary to divide the membrane into small patches or 'compartments', which are assumed to be isopotential. One edited by Christof Koch and Idan then applies Kirchhoff's circuit Segev, MIT Press, 1989. £40.50 (xii laws at each node, or compart+ 524 pages) ISBN 0 262 11133 0 ment, resulting in a large set of coupled ordinary differential equations. Excellent chapters by This book presents a collection of Rail, and by Segev, Fleshman and 12 papers that were used in lieu Burke establish the basics of this of course notes in a summer approach, and some of the course taught in 1988 at Cold mathematical and computational Spring Harbor, NY, USA by C. aspects are reviewed in chapters Koch and J, Bower. A careful by Mascagni and by Nelson, Furreading shows that it consists of manski and Bower. This approach to neural modeltwo distinct parts. Eight chapters cover the theory and applications ing is most effective when applied of a style of 'compartmental to physiologically (and anatomimodeling' that seeks to model cally) well-understood, and very neurons and neural systems in small-scale problems. An excelterms of membrane voltages and lent example in this volume is currents. The remaining four Getting's analysis of escape chapters cover a miscellany of behavior in Tritonia. However, topics: there is an excellent chap- Getting is careful to point out a ter by Shamma on auditory mod- problem with this method in that els, and three chapters discussing the specification of an appropri'neural network' models. Here, I ate set of parameters to deterwill concentrate on the book's mine the active membrane propmajor focus, which is the appli- erties of neurons is 'largely a matcation of compartmental model- ter of conjecture' (p. 176). Thus, even in the context of a ing to various neural systems. Compartmental modeling de- small, well-studied invertebrate rives from the early work of system, it is problematic to apply Rail, at the National Institutes of these methods fully. In several of Health (NIH), who applied one- the later chapters of this book, dimensional cable theory to applications to large-scale mamdescribe current flow in a passive malian systems are presented, but dendritic tree. In order to account without the benefit of Getting's for synaptic currents and voltage- warning. The most exciting, but dependent membrane properties, potentially dangerous, of these
are provided by several chapters that model large systems, such as macroscopic portions of the piriform and visual cortices. Wilson and Bower's ambitious model of piriform cortex produces impressive images of 'waves' of neural activity. However, the only comparison to data was for single neuron phenomena. How do these large-scale simulations compare to existing experimental data? No direct comparison is made to the relevant experimental work of Walter Freeman
biology may be less drastic, but they are important nevertheless. Neural network modelling allows the neural sciences to make the step from gathering ever more knowledge about the brain at a microscopic level, to integrating this knowledge at a level where it can be used to understand cognitive behavioural phenomena. From the neurobiology contributions it becomes clear that also in this field differences in opinion are the rule rather than the exception. What makes the book important and distinguishable from many other books on this topic is that it combines psychological
Eric L. Schwartz
,,
Methods in Neuronal Modelling. From Synapses to Networks
(Mass Action in the Nervous System Academic Press, 1975), who
pioneered the large-scale mathematical modeling of piriform cortex. It is thus not possible to evaluate Wilson and Bower's model. Even more ambitious is a model of the retino-geniculo-striate system, by Wehmeir eta/., which seeks to understand the generation of orientation specificity in cat V1. Their chapter cites a large number of biological parameters, such as cell densities and conductances, and it incorporates several well-known mechanisms to generate orientation tuning, including the original HubelWiesel model of summed excitatory input. It does indeed provide orientation tuning, which is reassuring, since several mechanisms for this purpose were built into the model in the first place. However, there is again no sigTINS, Vol. 13, No. 10, 1990
nificant comparison to experiments, or suggestion for further experiments that might be used to assess the reliability of the model. This chapter clearly illustrates the dangers and limitations of the use of compartmental-style modeling in the context of large systems. First, the attempt to extend knowledge about synapses and membranes, sketchy as it is in an area such as visual cortex, to a macroscopic model, is a form of radical reductionism. The sub-text of this book is that the computer resources (and by implication the neural parameters and knowledge) are finally available to enable such an attempt. But the results reviewed above do not
provide much support for this hope, at least in the sense of demonstrating a strong relationship between these simulations and available experimental data. In summary, this book is a valuable treatise on methods for applying fine-grained cable and compartmental neural modeling techniques, particularly as they are currently practised at the California Institute of Technology and the NIH, the institutions at which most of the authors are based. It is not, as is claimed on the dustjacket, a 'handbook of computational neuroscience', since it covers only a small and highly focused part of this subject. It is unfortunate that the book re-
Books Received
Alan A. Boulton, Glen B. Baker Michael W. Eysenck and Mark T. and Roger F. Butterworth (eds) Keane Cognitive Psychology: A Neuromethods 11. Carbohydrates Student's Handbook Lawrence and Energy Metabolism Humana Erlbaum Associates, 1990. £25.00 Press, 1989. £71.45 (xx + 383 (x + 557 pages) ISBN 0 86377 pages) ISBN 0 89603 143 8 153 X
David J. Aidley The Physiology of Excitable Cells (3d edn) Cambridge University Press, 1989. $34.50 (ix + 508 pages) ISBN 0 521 38863 5 Anon. Environmental Health Criteria, No. 101 World Health Organization Publications, 1990. Sw. fr. 16.00/$12.80 (Sw. fr. 11.20 in developing countries) (144 pages) ISBN 92 4 157101 2
Roger F. Butterworth and Gilles Pomier Layrargues (eds) Hepatic Encephalopathy: Pathophysiology and Treatment Humana Press, 1989. £83.85 (xix + 621 pages) ISBN 0 89603 164 0
peatedly represents its own, somewhat idiosyncratic style as identical to 'neural modeling' as a whole. From the point of view of students this is unsatisfactory since compartmental modeling techniques are not widely used: they represent a useful, but small part of the techniques available for modeling the nervous system. However, this book is an important source of practical information for those interested in constructing highly detailed, electrical circuit models of neurons and neural systems, and would provide an excellent source book for a course specifically entitled 'Methods of compartmental neural modeling'.
J. James Frost and Henry N. Wagner, Jr (eds) Quantitative Imaging. Neuroreceptors, Neurotransmitters and Enzymes Raven Press, 1990. $118.00 (x + 199 pages) ISBN 0 88167 611 X
Francesco Clementi and Jacopo Albert M. Galaburda (ed.) From Meldolesi (eds) Neurotransmitter Reading to Neurons MIT Press, N. G. Bazan (ed.) Lipid Mediators Release: The Neuromuscular 1989. £40.50 (xxii + 545 pages) in Ischemic Brain Damage and Ex- Junction Academic Press, 1990. ISBN 0 262 07115 0 perimental Epilepsy (New Trends in £19.50 (xv + 232 pages) ISBN Lipid Mediators Research Vol. 4) 0 1 2 1764605 Mark A. Gluck and David E. S. Karger, 1990. Sw. ft. 238.00/ Rumelhart (eds) Neuroscience DM 285.00/£99.20/$158.75 James Darnell, Harvey Lodish and and Connectionist Theory Law(vi + 262 pages) ISBN 3 8055 David Baltimore Molecular Ceil rence Erlbaum Associates, 1990. 5068 5 Biology (2nd edn) Scientific £49.95 (xiv + 405 pages) ISBN American Books, 1990. £25.95 0 8058 0504 4 Anders Bj6rklund, Albert J. (xl + 1105 pages) ISBN 0 7167 Aguayo and David Ottoson Brain 2078 9 Elisabeth A. H. Hall Biosensors Repair (Proceedings of an InterOpen University Press, 1990. national Symposium at the Stephen I. Deutsch, Abraham £19.99 (viii + 351 pages) ISBN Wenner-Gren Center, Stockholm, Weizman and Ronit Weizman 0335 15161 2 24-27 May, 1989) Macmillan (eds) Application of Basic NeuroPress, 1990. £55.00 (xii + 385 science to Child Psychiatry Plenum George A. Hashim and Mario pages) ISBN 0 935859 96 9 Press, 1990. $59.50 (xx + 400 Moscarello (eds) Dynamic Interpages) ISBN 0 306 43357 5 actions of Myelin Proteins (ProA. B. Bo~kovec and Edward P. gress in Clinical and Biological Masler (eds) Insect Neuro- Philippe Evrard and Alexandre Interactions Vol. 336)Wiley-Liss, chemistry and Neurophysiology Minkowski (eds) Developmental 1990. $59.50 (xi + 196 pages) 1989 Humana Press, 1990. Neurobiology (Nestl6 Nutrition ISBN 0 471 56729 9 $79.50 in USA ($89.50 else- Workshop Series Vol. 12) Raven where) (xii+480 pages) ISBN Press, 1989. $62.50 (xviii + 315 Robert D. Hawkins and Gordon 089603 1683 pages) ISBN 0 88167 544 X H. Bower (eds) Computational
TINS, Vol. 13, No. 10, 1990
437
i~
......
,~ .
.
.
.
.
.
.
.
.
.
Computational Neurosciences Laboratory, Departmentof Psychiatry, New York UniversityMedical Center,550 First Avenue, New York, NY 10016, USA,and Departmentof ComputerScience, Courantlnstitute, New YorkUniversity, WashinEtonSquare, New York,NY 10003, USA.
436
~
.
.
.
.
.
and neuroscientific perspectives on PDP modelling. In principle, PDP models allows a bridge to be built between the cognitive sciences and the neurosciences. From reading the book it is evident that something has been accomplished in bringing together the psychologist who, until recently, was explaining complex cognitive functioning solely in terms of condition-action rules or as the formal manipulation of symbols, and the neuroscientist, who painstakingly recorded singleneurone activity with a microelectrode or analysed the biochemical activity at synaptic clefts. However, it also becomes
clear that much more is needed. There is still a large gap between the psychologist developing neurally inspired networks to simulate learning of the past tense of English verbs, and the neuroscientist exploring the behaviour of neurally constrained networks and speculating about the possible role of the hippocampus in cognitive functioning. If our goal is to understand the brain and to fill the gap between the algorithmic and the implementational levels of describing cognition, PDP modellers from psychology and the neurosciences still have a lot to learn from each other; this book will assist in this process.
it becomes necessary to divide the membrane into small patches or 'compartments', which are assumed to be isopotential. One edited by Christof Koch and Idan then applies Kirchhoff's circuit Segev, MIT Press, 1989. £40.50 (xii laws at each node, or compart+ 524 pages) ISBN 0 262 11133 0 ment, resulting in a large set of coupled ordinary differential equations. Excellent chapters by This book presents a collection of Rail, and by Segev, Fleshman and 12 papers that were used in lieu Burke establish the basics of this of course notes in a summer approach, and some of the course taught in 1988 at Cold mathematical and computational Spring Harbor, NY, USA by C. aspects are reviewed in chapters Koch and J, Bower. A careful by Mascagni and by Nelson, Furreading shows that it consists of manski and Bower. This approach to neural modeltwo distinct parts. Eight chapters cover the theory and applications ing is most effective when applied of a style of 'compartmental to physiologically (and anatomimodeling' that seeks to model cally) well-understood, and very neurons and neural systems in small-scale problems. An excelterms of membrane voltages and lent example in this volume is currents. The remaining four Getting's analysis of escape chapters cover a miscellany of behavior in Tritonia. However, topics: there is an excellent chap- Getting is careful to point out a ter by Shamma on auditory mod- problem with this method in that els, and three chapters discussing the specification of an appropri'neural network' models. Here, I ate set of parameters to deterwill concentrate on the book's mine the active membrane propmajor focus, which is the appli- erties of neurons is 'largely a matcation of compartmental model- ter of conjecture' (p. 176). Thus, even in the context of a ing to various neural systems. Compartmental modeling de- small, well-studied invertebrate rives from the early work of system, it is problematic to apply Rail, at the National Institutes of these methods fully. In several of Health (NIH), who applied one- the later chapters of this book, dimensional cable theory to applications to large-scale mamdescribe current flow in a passive malian systems are presented, but dendritic tree. In order to account without the benefit of Getting's for synaptic currents and voltage- warning. The most exciting, but dependent membrane properties, potentially dangerous, of these
are provided by several chapters that model large systems, such as macroscopic portions of the piriform and visual cortices. Wilson and Bower's ambitious model of piriform cortex produces impressive images of 'waves' of neural activity. However, the only comparison to data was for single neuron phenomena. How do these large-scale simulations compare to existing experimental data? No direct comparison is made to the relevant experimental work of Walter Freeman
biology may be less drastic, but they are important nevertheless. Neural network modelling allows the neural sciences to make the step from gathering ever more knowledge about the brain at a microscopic level, to integrating this knowledge at a level where it can be used to understand cognitive behavioural phenomena. From the neurobiology contributions it becomes clear that also in this field differences in opinion are the rule rather than the exception. What makes the book important and distinguishable from many other books on this topic is that it combines psychological
Eric L. Schwartz
,,
Methods in Neuronal Modelling. From Synapses to Networks
(Mass Action in the Nervous System Academic Press, 1975), who
pioneered the large-scale mathematical modeling of piriform cortex. It is thus not possible to evaluate Wilson and Bower's model. Even more ambitious is a model of the retino-geniculo-striate system, by Wehmeir eta/., which seeks to understand the generation of orientation specificity in cat V1. Their chapter cites a large number of biological parameters, such as cell densities and conductances, and it incorporates several well-known mechanisms to generate orientation tuning, including the original HubelWiesel model of summed excitatory input. It does indeed provide orientation tuning, which is reassuring, since several mechanisms for this purpose were built into the model in the first place. However, there is again no sigTINS, Vol. 13, No. 10, 1990
nificant comparison to experiments, or suggestion for further experiments that might be used to assess the reliability of the model. This chapter clearly illustrates the dangers and limitations of the use of compartmental-style modeling in the context of large systems. First, the attempt to extend knowledge about synapses and membranes, sketchy as it is in an area such as visual cortex, to a macroscopic model, is a form of radical reductionism. The sub-text of this book is that the computer resources (and by implication the neural parameters and knowledge) are finally available to enable such an attempt. But the results reviewed above do not
provide much support for this hope, at least in the sense of demonstrating a strong relationship between these simulations and available experimental data. In summary, this book is a valuable treatise on methods for applying fine-grained cable and compartmental neural modeling techniques, particularly as they are currently practised at the California Institute of Technology and the NIH, the institutions at which most of the authors are based. It is not, as is claimed on the dustjacket, a 'handbook of computational neuroscience', since it covers only a small and highly focused part of this subject. It is unfortunate that the book re-
Books Received
Alan A. Boulton, Glen B. Baker Michael W. Eysenck and Mark T. and Roger F. Butterworth (eds) Keane Cognitive Psychology: A Neuromethods 11. Carbohydrates Student's Handbook Lawrence and Energy Metabolism Humana Erlbaum Associates, 1990. £25.00 Press, 1989. £71.45 (xx + 383 (x + 557 pages) ISBN 0 86377 pages) ISBN 0 89603 143 8 153 X
David J. Aidley The Physiology of Excitable Cells (3d edn) Cambridge University Press, 1989. $34.50 (ix + 508 pages) ISBN 0 521 38863 5 Anon. Environmental Health Criteria, No. 101 World Health Organization Publications, 1990. Sw. fr. 16.00/$12.80 (Sw. fr. 11.20 in developing countries) (144 pages) ISBN 92 4 157101 2
Roger F. Butterworth and Gilles Pomier Layrargues (eds) Hepatic Encephalopathy: Pathophysiology and Treatment Humana Press, 1989. £83.85 (xix + 621 pages) ISBN 0 89603 164 0
peatedly represents its own, somewhat idiosyncratic style as identical to 'neural modeling' as a whole. From the point of view of students this is unsatisfactory since compartmental modeling techniques are not widely used: they represent a useful, but small part of the techniques available for modeling the nervous system. However, this book is an important source of practical information for those interested in constructing highly detailed, electrical circuit models of neurons and neural systems, and would provide an excellent source book for a course specifically entitled 'Methods of compartmental neural modeling'.
J. James Frost and Henry N. Wagner, Jr (eds) Quantitative Imaging. Neuroreceptors, Neurotransmitters and Enzymes Raven Press, 1990. $118.00 (x + 199 pages) ISBN 0 88167 611 X
Francesco Clementi and Jacopo Albert M. Galaburda (ed.) From Meldolesi (eds) Neurotransmitter Reading to Neurons MIT Press, N. G. Bazan (ed.) Lipid Mediators Release: The Neuromuscular 1989. £40.50 (xxii + 545 pages) in Ischemic Brain Damage and Ex- Junction Academic Press, 1990. ISBN 0 262 07115 0 perimental Epilepsy (New Trends in £19.50 (xv + 232 pages) ISBN Lipid Mediators Research Vol. 4) 0 1 2 1764605 Mark A. Gluck and David E. S. Karger, 1990. Sw. ft. 238.00/ Rumelhart (eds) Neuroscience DM 285.00/£99.20/$158.75 James Darnell, Harvey Lodish and and Connectionist Theory Law(vi + 262 pages) ISBN 3 8055 David Baltimore Molecular Ceil rence Erlbaum Associates, 1990. 5068 5 Biology (2nd edn) Scientific £49.95 (xiv + 405 pages) ISBN American Books, 1990. £25.95 0 8058 0504 4 Anders Bj6rklund, Albert J. (xl + 1105 pages) ISBN 0 7167 Aguayo and David Ottoson Brain 2078 9 Elisabeth A. H. Hall Biosensors Repair (Proceedings of an InterOpen University Press, 1990. national Symposium at the Stephen I. Deutsch, Abraham £19.99 (viii + 351 pages) ISBN Wenner-Gren Center, Stockholm, Weizman and Ronit Weizman 0335 15161 2 24-27 May, 1989) Macmillan (eds) Application of Basic NeuroPress, 1990. £55.00 (xii + 385 science to Child Psychiatry Plenum George A. Hashim and Mario pages) ISBN 0 935859 96 9 Press, 1990. $59.50 (xx + 400 Moscarello (eds) Dynamic Interpages) ISBN 0 306 43357 5 actions of Myelin Proteins (ProA. B. Bo~kovec and Edward P. gress in Clinical and Biological Masler (eds) Insect Neuro- Philippe Evrard and Alexandre Interactions Vol. 336)Wiley-Liss, chemistry and Neurophysiology Minkowski (eds) Developmental 1990. $59.50 (xi + 196 pages) 1989 Humana Press, 1990. Neurobiology (Nestl6 Nutrition ISBN 0 471 56729 9 $79.50 in USA ($89.50 else- Workshop Series Vol. 12) Raven where) (xii+480 pages) ISBN Press, 1989. $62.50 (xviii + 315 Robert D. Hawkins and Gordon 089603 1683 pages) ISBN 0 88167 544 X H. Bower (eds) Computational
TINS, Vol. 13, No. 10, 1990
437