Computational Explorations in Cognitive Neuroscience Randall C. O'Reilly, Yuko Munakata The goal of computational cognitive neuroscience is to understand how the brain embodies the mind by using biologically based computational models comprising networks of neuronlike units, This text, based on a course taught by Randall O'Reilly and Yuko Munakata over the past several years, provides an in-depth introduction to the main ideas in the field. The neural units in the simulations use equations based directly on the ion channels that govern the behavior of real neurons, and the neural networks incorporate anatomical and physiological properties of the neocortex. Thus the text provides the student with knowledge of the basic biology of the brain as well as the computational skills needed to simulate large-scale cognitive phenomena. The text consists of two parts. The first part covers basic neural computation mechanisms: individual neurons, neural networks, and learning mechanisms. The second part covers large-scale brain area organization and cognitive phenomena: perception and attention, memory, language, and higher-level cognition. The second part is relatively self-contained and can be used separately for mechanistically oriented cognitive neuroscience courses. Integrated throughout the text are more than forty different simulation models, many of them full-scale research-grade models, with friendly interfaces and accompanying exerci: The simulation software (PDP++, available for all major platforms) and simulations can be downloaded free of charge from the Web. Exercise solutions are available, and the text includes full information on the software. Foreword Preface Acknowledgments 1 Introduction and Overview I Basic Neural Computational Mechanisms 2 Individual Neurons 3 Networks of Neurons 4 Hebbian Model Learning 5 Error-Driver Task Learning 6 Combined Model and Task Learning, and Other Mechanisms II Large-Scale Brain Area Organization and Cognitive Phenomena 7 Large-Scale Brain Area Functional Organization8 Perception and Attention 9 Memory 10 Language 11 Higher-Level Cognition 12 Conclusions III Simulator Details A Introduction to the PDP++ Simulation Environment B Tutorial for Constructing Simulations in PDP++ C Leabra Implementation Reference References Author Index Subject Index mindwapreeForeword ‘The Role of Computational Models in Cognitive Neuroscience ‘The publication of O'Reilly and Munakata’s Computa tional Explorations in Cognitive Neuroscience comes at an opportune moment. The field is rapidly growing, ‘with contare and inctin os Researchers trom backgrounds ranging trom psychol- gy to molecular bology are poneing sata the held. Keke Kandel has suggested that “cognitive neuroscience— with its concern about perception. action, memory. lan- suaye and selective attention—will increasingly come (represent the central cus of all neuruseienves twwentyfirst century.” Today, quite a bit of the excitement in the ficld su ities hold great promise to allow us to visualize the brain while cagaition 1s occurring, and itis likely that there will be ongoing breakthroughs in spatial and ten poral resolution. Working upward from the molecular ievei, new genetic methods for creating animais with alterations in tie basic functional properties of specific mivups oF neuiony and synapses is alivwing dei exploration of how these cellular and synaptic pro- cesses impact higher processes such as spatial learning in animals. Within neurophysiology, the use of multi electrode array's to record from as many as 100 separate neurons ata time has led to new insights into the repre- sentation of information during behavior, and the later reactivation of these representations from memory dur- ing sleep With ali their marveious toois, the question arises. do we really need computational models in cognitive know about the neural basis of cognition through ex perimental investigation? Do we need a book like the present one to explore the principles of neural compu- tation and apply them fo the task of understanding how cognition arises from neuronal interactions? The answer is: Yes, we do need computational mod eis in cognitive neuroscience. To support this answer, I ‘will begin by describing what I take to be one of the cen- tai zoais of cognitive neuroscience, i witi dren Gescti what we mean by the phrase “a computational model and consider the rote such models ean play in address- ing the central goal. Along the way I hope to indicat some of the shortcomings of experimental research un dertaken without the aid of computational models and how models can be used to go beyond these limitations. ‘The goal is to make clear exactly what models are, and the role they are mtended to play. Htst, what 1s this central goal of cognitive neuro~ science? To me. and I think to O'Reilly. Munakata, and many researchers in the field, the goal is to understand how neural processes zive tise W cognition, Typically, coznition iy broadly consttued to inctude perception, at- tention, language, memory, problem solving, planning, reasoning, and the coordination and execution of ac- tion. And typically some task or tasks are used to make behavioral observations that tap these underlying pro cesses: aspects of conscious experience are included, to the extent that they can be subjected to. scientific scrutiny through ohservahles (including vert reports or other readout methorls). The processes considered may be ones that take place in a brief interval of time. such as the processes that occur when a human observer ready a visualiy presented word. Or they nay be ones that take place over longer periods of time, such as the processes that occur as a child progresses through var-