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VANDERBILT HON 182 - On Building a Bridge Between Brain and Behavior

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18 Nov 2003 14:3 AR AR207-PS55-02.tex AR207-PS55-02.sgm LaTeX2e(2002/01/18) P1: GCE10.1146/annurev.psych.55.090902.141907Annu. Rev. Psychol. 2004. 55:23–50doi: 10.1146/annurev.psych.55.090902.141907Copyrightc° 2004 by Annual Reviews. All rights reservedFirst published online as a Review in Advance on October 20, 2003ON BUILDING A BRIDGE BETWEENBRAIN AND BEHAVIORJeffrey D. SchallCenter for Integrative & Cognitive Neuroscience, Vanderbilt Vision Research Center,Department of Psychology, Vanderbilt University, Nashville, Tennessee 37203;email: [email protected] Words linking proposition, linking hypothesis, stop signal, countermanding,response preparation, intention, eye field, reaction time, response time, saccade■ Abstract Cognitive neuroscience is motivated by the precept that a discoverablecorrespondence exists between mental states and brain states. This precept seemsto be supported by remarkable observations and conclusions derived from event-related potentials and functional imaging with humans and neurophysiology withbehaving monkeys. This review evaluates specific conceptual and technical limitsof claims of correspondence between neural events, overt behavior, and hypothe-sized covert processes examined using data on the neural control of saccadic eyemovements.CONTENTSINTRODUCTION ..................................................... 24Inferring Mechanism from Behavior ..................................... 24Inferring Function from Neuronal Properties .............................. 25LINKING PROPOSITIONS ............................................. 26Testing Linking Propositions ........................................... 27LINKING PROPOSITIONS ABOUT SACCADE PRODUCTION .............. 29Evidence for Covert Response Preparation ................................ 31Response Preparation and Intention ..................................... 31Bridge Locus for Response Preparation .................................. 32Control of Saccade Initiation ........................................... 33Relation of Neural Activity to Response Time ............................. 34Relation of Neural Activity to Movement Cancellation ...................... 37Alternative Propositions Mapping GO and STOP onto Neural Processes ........ 38EVALUATING LINKING PROPOSITIONS FOR THE PRODUCTIONOF SACCADES ..................................................... 40Do Identical Neural States Map onto Identical Saccades? .................... 40Do Identical Saccades Map onto Identical Neural States? .................... 42SUMMARY AND CONCLUSIONS ...................................... 430066-4308/04/0204-0023$14.002318 Nov 2003 14:3 AR AR207-PS55-02.tex AR207-PS55-02.sgm LaTeX2e(2002/01/18) P1: GCE24 SCHALLINTRODUCTIONMany authors write with conviction that the correspondence of the mental withthe neural is so secure that an ultimate theory of mental phenomena will reduce toneural terms (e.g., Churchland 1986, Crick 1994). Others argue that mental statesdepend on but are not reducible to the physical states of the brain (e.g., Davidson1970, Fodor 1981, Pylyshyn 1984). Determining whether the mental reduces to oremerges from the neural cannot be accomplished without correctly describing themapping between the two.Inferring Mechanism from BehaviorBefore the development of methods to monitor brain states during behavior, physi-ological mechanisms could be inferred only from behavioral testing. Nevertheless,in the nineteenth century investigators began to articulate the correspondence be-tween mental and physical processes. For example, Mach wrote, “To every psychi-cal there corresponds a physical, and conversely. Like psychical processes corre-spondtolike physical, unliketounlike....Particularsof thephysical correspond toall the particulars of the psychic” (Boring 1942). Even philosophers who advocatea nonreductionist position acknowledge a mapping between mental and physi-cal processes—“Although the position I describe denies there are psychophysicallaws, it is consistent with the view that mental characteristics are in some sensedependent, or supervenient, on physical characteristics. Such supervenience mightbe taken to mean that there cannot be two events alike in all physical respects butdiffering in some mental respects, or that an object cannot alter in some mental re-spect without altering in some physical respect” (Davidson 1970). Such a positioncan be translated into an effective research strategy according to the propositionthat “...whenever two stimuli cause physically indistinguishable signals to be sentfrom the sense organs to the brain, the sensations produced by these stimuli, as re-ported by the subject in words, symbols or actions, must also be indistinguishable”(Brindley 1970). Application of this principle in sensory detection or discrimina-tion experiments permits testing hypotheses about physiological processes.Another approach to understanding the mechanisms responsible for behaviorhas been through mathematically precise models of cognitive processes testedagainst detailed measurements of performance (e.g., Townsend & Ashby 1983;Luce 1986, 1995). Unfortunately, cognitive psychology abounds with alternativemodels with mutually exclusive architectures oralgorithms, manyof which are dif-ficult or impossible to distinguish through behavioral testing. For example, choicebehavior can be accounted for by sequential sampling models in which a singleaccumulator represents the relative evidence for two alternatives (e.g., Ratcliff &Rouder 1998). An alternative to a random walk of a single accumulator betweenalternatives is a race among multiple accumulators representing each alternative(e.g., Bundesen 1990, Logan 2002). In fact, models with single or multiple ac-cumulators can account for common sets of data (Van Zandt & Ratcliff 1995,18 Nov 2003 14:3 AR AR207-PS55-02.tex AR207-PS55-02.sgm LaTeX2e(2002/01/18) P1: GCEBRIDGING BRAIN AND BEHAVIOR 25Van Zandt et al. 2000), highlighting the limitations of arriving at secure inferencesabout mechanism based only on behavior (e.g., Uttal 1997). The theoretical issuehas been articulated most definitely in the theory of finite automata (Moore 1956).It has been proven that given any computer with a finite number of inputs, outputs,and internal states and any experiment that determines the mapping of outputs toinputs, there exist other computers that are experimentally distinguishable fromthe original computer for which the


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