ABSTRACT "CROSS-FIELD" VISUAL MASKING BY TACHISTOSCOPIC PRESENTATION OF TARGET AND NOISE PATTERNS TO OPPOSITE CEREBRAL HEMISPHERES BY David P. Goff Methods for studying hemispheric asymmetry in the processing of sensory information by normal human subjects involve unilateral presenta- tion of stimuli which then project to the opposite hemisphere. In the employment of these techniques, detection of laterality differences is often difficult. A reason for this is that several milliseconds after initial projection to one hemisphere, information becomes available for processing by the other hemisphere due to transfer across the commissures. The goals of this study were to develop a new technique for increas- ing observed laterality differences in the visual modality and test the underlying explanatory constructs. The procedure employed tachistoscopic presentation of target information to one hemisphere and a pattern "mask" to the other. Although target and mask information then transfer to apposite hemispheres, it was predicted that the mask would produce less interference in one hemisphere than in the other due to attenuation through the corpus callosum. Consequently, a greater laterality difference would be observed under masking than with conventional procedures. Target stimuli were single alphabetic letters, and recognition accuracy was the dependent variable. When the pattern was projected initially to the left (language) hemisphere, masking should have been greater than when mask was projected to the right hemisphere. David P. Gaff Findings indicated that the predicted phenomenon was: statistically confirmed, not influenced by differences in energy levels between mask and target stimuli, significantly affected by stimulus onset asynchrony between target and mask, dependent on size of the mask, and not reproducible when diffuse light was substituted for the pattern. Various aspects of these results supported the theory that the cross-field masking effect resulted from interaction of central (hemispheric) rather than peripheral neural mechanisms. ‘Hethods of improving the paradigm and for testing related theoretical issues were discussed. "CROSS-FIELD" VISUAL MASKING BY TACHISTOSCOPIC PRESENTATION OF TARGET AND NOISE PATTERNS To OPPOSITE CEREBRAL HEMISPHERES By \ David PP"‘co££ A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Department of Psychology 1973 g. I: a- ’9} fly; r 2; e Copyright by DAVID PAUL.GOFF 1973 DEDICATION To Dr. Charles Henley ii ACKNOWLEDGMENTS I would like to thank Dr. Charles Henley, Committee Chairman, for his assistance with this project and for his support as mentor, counselor, and confidant throughout my past four years as a graduate student at Michigan State. My appreciation also goes to Committee Members Dr. Lester H. Hyman, Dr. Hiram E. Fitzgerald, and Dr. Robert L. Raisler for their support. Dr. Paul Bakan has greatly contributed to the develOpment of my interests which evolved into conducting this project, and Dr. James L. Zacks has provided valuable technical assistance, consultation time, and literature sources. Finally, I am indebted to my wife, Anna-Karin, for the laborious job of typing and proof-reading this manuscript, for her moral support, and the many personal sacrifices she has made in my behalf over the past four years. 111 TABLE OF CONTENTS CHAPTER 1: INTRODUCTION ............................................. 1 CHAPTER 2: EXPERIMENT I ... .......................................... 17 Method ............ ......................... . ................... 18 Results and discussion ................. . ........... . ........... 25 CHAPTER 3: EXPERIMENT II ............................................ 39 Method ............... .......... . ............ ... ......... ....... 46 Results and discussion ..... ......................... .......... . 47 CHAPTER 4: EXPERIMENT III ......... ........... . ............. .... ..... 54 Method ................... ......... ............................. 56 Results and discussion .. ..... ........... ....... ................ 59 CHAPTER 5: EXPERIMENT IV ..... 70 Method ......................................................... 73 Results and discussion .......................... ...... ......... 75 CHAPTER 6: GENERAL SUMMARY AND CONCLUSION ........................... 81 BIBLIOGRAPHY ........................................................ 88 APPENDIX A: Brief Review of Controversial Issues Concerning Left Versus Right Field Differences in Tachistoscopically Presented Vigual Stimli O... ...... OOOOOOOOOOOOOOO......OOOOOOOOOOOOOOOOOO 93 APPENDIX B: Errors (Guesses) Classified as "Stimulus Dependent" for Each Target Letter Used in Experiment I (Error Analyses 1 and 2) 94 APPENDIX C: Summary Table and Analysis of Variance for Error Analysis 1 Based on Percentage of Stimulus-Dependent Errors ‘. the Depend‘nt V‘ri.b1. O0.0.0.0.0..........OOOOOOOOOOOO...OO. 95 APPENDIX D: Summary Table and Analysis of Variance for Error Analysis 2 Based on Percentage of Stimulus-Dependent Errors 8' the Dependent variable 00.0.0000........OOIOOOOOOOCOOOOOO0.00 96 iv LIST OF TABLES Table 1. Percentage of correct responses for left versus right hemis- phere and mask versus no-mask conditions for Groups I-IV, per— centage decrement produced by the mask, average across hemis- phere conditions, difference in percentage decrement, and mean target exposure time at threshold ................................ Table 2. Analysis of variance summary based on percentage correct .core. for Experimentl O....OOOOOOOOOOOOOOOOOO00.00.000.000....... Table 3. Percentage of stimulus-dependent errors for left versus right hemisphere and mask versus no-mask conditions averaged for Groups I-IV, percentage decrement produced by the mask, average across hemisphere conditions, and difference in percentage decre- ment (Error analysis 1) ........................................... Table 4. Percentage of stimulus-dependent errors for left versus right hemisphere and mask versus no-mask conditions averaged for Groups I-IV, percentage decrement produced by the mask, average across hemisphere conditions, and difference in percentage decre- mnt<£rroranaly.1.2) O00............OOOOOIOOOOOOOOO......OOOOOOO Table 5. Percentage of correct responses for left versus right hemis- phere and mask versus no-mask conditions for Groups III and V, percentage decrement produced by the mask, average across hemis- phere conditions, difference in percentage decrement, and mean target exposure time at threshold ................................ Table 6. Analysis of variance summary based on percentage correct scores for Experiment II (Groups III and V) ....................... Table 7. Average difference in percentage correct scores, between no-mask and mask conditions, for left and right hemisphere versus two "stimulus-onset-asynchrony" conditions ........................ Table 8. One—tailed orthogonal comparison test of the interaction between hemisphere and stimulus-onset-asynchrony conditions based on the average difference in percent correct scores (no-mask minus m.kcond1t1°n8)eeeeee ..... ......ICOOOO............OOOOOOOOOOOO0.0 Table 9. Percentage of correct responses for left and right hemisphere conditions as a function of the no-mask, large mask, small corre- sponding, and small non-corresponding mask conditions of Experi- ment III (Group VI) .............................................. 25 27 33 35 47 48 49 50 S9 Table 10. Table 11. Analysis of variance summary based on percentage correct scores for Experiment III ......................................... Statistical tests of the paired comparisons of all possible combinations of masking conditions (averaged across hemispheres) Table 12. for Experiment III by the method of Scheffé ....................... Statistical tests of the two-way interaction effects between left-right hemisphere conditions and all possible pairs of mask conditions for Experiment III by the method of Scheffé ............ Table 13. Table Table Table Table Table Table 14. 15. l6. 17. 18. 19. Percent correct responses for left-right and mask - no—mask conditions for three individual subjects receiving only the small "corresponding" mask stimulus, percentage decrement produced by the mask, average across hemisphere conditions, difference in per- cent decrement, and mean target exposure time at threshold ........ Percentage of correct responses for left-right hemisphere and no-mask - mask conditions for pattern versus light flash mask- ing stimuli, percentage decrement produced by masking, average across hemisphere conditions, difference in percentage decrement, and mean target exposure time at threshold ........................ Analysis of variance summary based on percentage correct scores for Groups V and VII of Experiment IV ...................... Summary table for error analysis 1 .......................... Analysis of variance for error analysis 1 ................... Summary table for error analysis 2 .......................... Analysis of variance for error analysis 2 ................... vi 59 6O 61 75 76 95 95 96 96 LIST OF FIGURES Figure l. Afferent pathways of the visual system of the rhesus monkey Figure 2. Relative strength of target and noise information arriving in the respective hemispheres as predicted by the model ........... Figure 3. Predicted changes in recognition accuracy for verbal target stimuli resulting from projection of the mask pattern to the hemis- phere Opposite that of the target ................................. Figure 4. Pathways in the visual system of the retinal images of two objects falling on the vertical meridian, one in front of and one behind the fixation point, for a human with sagittal section of the Optic chiasm .................................................. Figure 5. Example of a single neuron in the right visual cortex which was driven by a stimulus crossing the vertical meridian for a cat with sagittal section of the Optic chiasm ......................... Figure 6. Fixation stimulus with brackets for parallax adjustment ..... Figure 7. Example of the target, mask, and no-mask stimuli ............ Figure 8. Composite view of the stimuli as seen through the tachisto- scope on a "masked" trial with target projecting to the right hemi.Pher. 0.0.0.............OOOOOOOOOOOOOOOOOOO......OCOOOOOOOOOOO Figure 9. Expected stimulus arrival asynchrony of target and mask stimuli at the two hemispheres for onset of a left-field target preceding onset of a right-field mask by 8 ms with an assumed callosal transmission time of 8 ms ................................ Figure 10. Expected stimulus arrival asynchrony of target and mask stimuli at the two hemispheres for onset of a right-field target preceding onset of a left-field mask by 8 ms with an assumed callosal transmission time of 8 ms ................................ Figure 11. Corresponding and non-corresponding heteronymous mask locations for the right-field (left hemisphere) mask conditions ... vii 4 6 7 l4 16 20 21 4O 41 56 Chapter 1: INTRODUCTION Over the past 50 years a large volume of evidence has accrued in support of the concept of lateral specialisation of functioning in the two cerebral hemispheres of humans. This evidence has derived from a variety of sources which include: direct cortical stimulation, study of patients with unilateral brain damage, postoperative case studies, gross evoked potential recordings, sodium amytal anesthetisation of one or the other of the hemispheres as a preoperative investigation technique, unilateral presentation of visual, auditory, and tactile stimuli to normal human subjects, and most dramatic, the recent work of Sperry and associates with "split-brain" epileptic patients. Among the most consistent generaliza- tions made are that the left (language) hemisphere in most right-handers specialises in symbolic and analytic functions while the right (mute) hemisphere demonstrates a superiority in Gestalt-type processing of spatial information (Hecaen, 1962; Gaszaniga et al., 1965; Geschwind, 1965; Levy, 1969; Levy-Agresti & Sperry, 1968; Sperry & Levy, 1970). Nbst of the above methods for investigating asymmetry of hemispheric functioning comprise the drawback that few brain-damaged or surgical patients are available to the general scientific community. Furthermore, the condi- tions leading to the use of such surgical procedures involve pre-existing structural damage to the normal central nervous system of a rather critical nature. The choice, therefore, of some supplementary paradigms that permit the study of "normal" subjects seems desirable. The most popular methods 2 which fall in this category are the dichotic listening paradigm for the auditory mode and tachistoscOpic presentation of visual stimuli to the right or left hemisphere for the visual mode. Ximura (1961) showed a left hemisphere superiority for verbal stimuli via the right ear, and a right hemisphere superiority for the perception of melodies (1964). Although the hemisphere contralateral to the stimulated ear is favored for receiving the stimulus, each ear nevertheless has a direct cochlea-cortical projection route to both hemispheres (Davis, 1951), a "disadvantage" not shared by the visual system. For higher mammals, the two right hemiretinae (left visual field) contain direct retina-cortical projections to the right hemisphere only while the two left hemiretinae project directly to the left visual cortex (see Thompson, 1967, p. 250). Thus visual stimuli can be projected, at least initially, to only one hemisphere by tachistoscopic exposure to the contralateral (Opposite) visual field. The vision literature on laterality is extensive and not without controversy (see review by White, 1969). The general findings, however, support a left hemisphere superiority for verbal and a right hemisphere superiority for spatial processing as indicated by more accurate recogni- tion or shorter reaction times to tachistoscOpic presentation of the appropriate material in the appropriate visual half-field (Ximura, 1966; White, 1969; Rissolatti et al., 1971)1. While some of these vision studies show significant differences in support of the lateralisation of function hypothesis, the differences are nevertheless quite small and many of the studies are either contradictory or totally inconclusive. The most outstanding difficulty in this approach with normal subjects is, of course, the fact that 6-10 ms (Efron, 1963a; lThis literature has been reviewed and the troublesome issues dis- cussed in Appendix A. 3 Jeeves & Dickson, l970)after one hemisphere has been stimulated, the in- formation has become available for processing by the Opposite hemisphere via the corpus callosum. The function of the callosum in this transfer of visual information has been demonstrated with commissurotomized animal and human patients (Sperry, 1961; Geschwind, 1965). ElectrOphysiological evidence has been provided by Berlucchi et a1. (1967). Hence, either or some combination of both hemispheres may still do the processing. Clearly, if one is to investigate more complex modes of unilateral "information processing" in "normal" human subjects, a method is needed for isolating, as much as possible, the processing of Specific information to a given hemiSphere. One means by which this may be possible would be to present target information to one hemisphere while presenting "conflicting” or noise pattern stimuli to the contralateral side and perhaps interfere with its processing. The specific aim of this project has been to develOp and test the feasibility of such a model for the visual system. The empir- ical data will provide some tests of the hypothetico-deductive assumptions on which the model is based. Following is a detailed description of the model along with the behavioral, anatomical, and neuTOphysiological evidence which seem to support its tenability. grief anatomy. Figure l is a block diagram of the afferent pathways of the visual system of the rhesus monkey which relate specifically to the processing of patterned stimuli. For simplicity, only the pathways from right hemiretinae corresponding to the left field of vision have been depicted. In primates, the retina-cortical projection (for the contralateral half-field of vision) goes exclusively to area 17 (Wilson a Cragg, 1967) of the visual cortex. Area 17 then projects to 18 and 19 on both sides (Cragg et al., 1969; Zeki, 1969; Zeki, 1970); 18 and 19 from both sides then project to the Eight hemiretinae temporal nasal I I ‘t. Optic , \ chiasm ’ \ I \ I \> I ’I lateral geniculate .2, ® nuc leus Z////////””—_- anterior _—~§‘\\\\\\\\\\5 cogeismtzf [inf::::::poral __IL-"TL:_¢7 ‘ \ f I \w /‘i /1 19/ 2:32;: \“’ ><\ 718/ 4 L _____ hl/ spleneum of 17 corpus callosum Figure l. Afferent pathways of the visual system of the rhesus monkey (most like men). Connections to structures (e.g. pulvinar and superior colliculus) not related to pattern processing are excluded. (Drawn from a description by Gross, et. al., 1972) S inferotemporal cortex (Cragg et al., 1969). All the cortico-cortical path- ways just described go through the spleneum of the corpus callosum. The two inferotemporal cortices connect through the anterior commissure (Fox et al., 1948). Present evidence suggests that these pathways from area to area of a given side represent successive stages of pattern processing (Hubel & Wiesel, 1965; Gross et al., 1972). The model Given that the two eyes are focused on a fixation point, a "target" stimulus tachistoscopically flashed - for a time shorter than the 200 ms central reaction time for saccadic eye movements (Robinson, 1968) - to, say, the left half-field of vision, will go to area 17, the "primary" receiving area, in the right visual cortex (refer to Figure 1). The information will then go to the "secondary" receiving areas 18 and 19 on the ipsilateral side and to the secondary areas via the longer transcallosal pathway on the contralateral side. The writer proposes that the "raw" stimulus informa- tion, coded in electrophysiological form, will be more attenuated when it arrives at the contralateral as compared to the ipsilateral secondary re- ceiving area as a result of the longer transcallosal pathway. (These assumptions will be discussed below.) Considering, now, the other half of the system, a visual "noise" pattern simultaneously flashed in the right visual field will arrive in the secondary areas with greater intensity in the left as compared to the right hemisphere. The result, for this partic- ular example, would be that the right hemisphere receives a strong target signal (compared to the left) and a weak noise pattern (compared to the left). Correspondingly, the left side receives a weaker target signal and a stronger noise pattern (see Figure 2). Coined in different terms a high "signal-to-noise" ratio is obtained on the right side, and a low 6 signal-to-noise ratio on the left. If the target and visual noise pattern are located in corresponding heteronymous (mirror symmetric) areas of the half visual fields, and not too far from the vertical meridian (less than 7° visual angle), they should project to a common location in visual area 18 (see for example Berlucchi & Rizzolatti, 1968; and Blakemore, 1970). It is proposed that the noise pattern will interfere with (mask) the target stimulus in such a way as to decrease recognition accuracy for the target. This "masking" effect should occur on both sides (since both stimuli go to both sides), but it should be greater on the side with the lowest signal- to-noise ratio, i.e. the left hemisphere for this particular example. A G) I \ l \ I \ I, \ (.1 L) weak strong target target 1 strong weak noise noise 1 left right hemisphere hemisphere Figure 2. Relative strength of target and noise information arriving in the respective hemispheres as predicted by the model. Employing, now, the previously described concepts of lateral specialisation of function of the two hemispheres, it should be a simple matter to derive the behavioral predictions generated by the model. Take, for example, the left (language) hemisphere as the required processor. It is a given at this point that a verbally coded response to verbal stimulus material requires that the information reach the left hemisphere. 7 A stimulus (letter or word) flashed to the right hemisphere must cross the transcallosal pathway, and the signal attenuation hypothesis predicts that recognition accuracy should be less than when the stimulus is flashed directly to the left hemisphere (see Figure 3. no mask condition). These are the standard observations as described in the above studies. With the recognition X accuracy I k tar'get tar'get to left h to right h Figure 3. Predicted changes in recognition accuracy for verbal target stimuli resulting from projection of the mask pattern to the hemisphere opposite that of the target. (The finding Xj>k would satisfy the model) target flashed to the right and noise pattern now flashed to the left hemis- phere, recognition accuracy should decrease by some amount X, as compared to the right hemispher "no mask" condition (Figure 3). The responding hemisphere now has the low signal-to-noise ratio. To assess the undesirable (for present purposes) effect of the mask projecting to the opposite hemis- phere via the cortico-cortical pathway, we reverse (laterally) the above stimuli and flash the target to the left hemisphere and the noise pattern to the right. Recognition accuracy should now decrease by some smaller amount k (from left hemisphere - no mask condition), since the left or 8 responding hemiSphere now has the larger signal/noise ratio. Thus, if the X:>k prediction is the empirical finding. then the data will support the model. Secondly, and as an algebraic consequence of the above statements, the difference in recognition accuracy between left and right hemisphere presentations should increase under the masking condition, and the magni- tude of that increase will determine the utility of the paradigm as a re- search tool. Qichoptic visual masking That interference between a target and noise pattern projected by separate pathways can be produced at the cortical level receives strong sup- port from the dichOptic visual masking studies. Schiller (1965) flashed a letter to one eye and either a flash of light or a visual noise pattern to the other at corresponding retinal areas (foveally) and found a significant increase in recognition threshold (in ms exposure time) above the no- masking condition for the noise pattern - but not the diffuse light. These results indicate an interaction between stimuli presented to the two eyes at some central location subserving a binocular interaction between the two eyes. Area 17, which receives a direct retina-cortical projection from both eyeg.serves as a good candidate.1 Hubel and Wiesel (1965, cat; 1968, monkey) find that many "simple" cells in area 17 can be binocularly driven from corresponding homonymous points of the two retinae and that they respond only to lines and edges (pattern fragments), not diffuse light. 1In their spatial vision review article, BishOp and Henry (1971, p. 145) state: "Relatively little interaction between the pathways from the two eyes takes place below the level of the striate cortex (Sanderson et al., 1969; Singer, 1970)." The authors they cite, of whom Bishop is one, found that some inhibitory binocular interaction does occur in the lateral geniculate body of the cat. The relevant pathways are unknown and no speculation has been made concerning their purpose. It should also be noted that the inter- action is (a) extremely small and (b) exists only for corresponding homonymous points. 9 Referring to Figure 1, we see that the difference between the dichOptic and "cross-field" masking paradigms is as follows: in the dichoptic case, the stimuli may interact in area 17 which receives direct retino-cortical projections from homonymous retinal areas; in the "cross- field" case interaction cannot occur until area 18 which receives projec- tions from ipsilateral 17 (right hemiretinae in the diagram) and contra- lateral 17 (left hemiretinae) via the transcallosal route. In the latter instance, individual cells in area 18 would have to be driven from corre— sponding heteronymous points in the two half-fields of vision. Direct evidence for this latter possibility has recently been obtained by Berlucchi & Rizzolatti (1968) who found neurons in area 18 that could be binocularly driven from corresponding heteronymous areas in the split- chiasm cat (Figure 5). Some functions of the corpus callosum This area is too extensive to receive a detailed review here (see Ettlinger, 1965; BishOp & Henry, 1971); however, a few points relevant to the theoretical development of the above model require discussion. They concern the nature of information transfer between the visual cortices of the two hemispheres and its relation to the issues of interhemispheric masking. The following introductory paragraph will facilitate the discus- sion of these points. Hubel and Wiesel (1965), using the retrograde degeneration technique, found that fibers going from area 17 to ipsilateral and contralateral l8 and 19 projected to very specific areas; the area in 18 immediately adjacent to the 17-18 border, and the area in 19 immediately adjacent to the 18-19 border. The retinotopic projection to these areas represents the areas adjacent to the vertical midline of the field of vision. For example, 10 they found (by microelectrode recording; Hubel & Wiesel, 1967) that the "receptive fields" of all cells on the 18 side Of the 17-18 border ex- tended to the vertical meridian. Secondly, Berlucchi and Rizzolatti (1968) found, as mentioned previously, that cortical cells in this same section of area 18 could be binocularly driven from correSponding heter- onymous retinal areas adjacent to the vertical midline. This evidence suggests that two of the "natural" functions of this callosal crossover network may be to (a) provide continuity between the two half-fields of vision (Hubel & Wiesel, 1967; Berlucchi & Rizzolatti, 1967) and (b) pro- vide a stereOptic mechanism for making depth judgments about objects falling on the vertical meridian and thus projecting to Opposite hemis- pheres (see discussion and review by Bishop & Henry, 1971). It is by this particular crossover network through the spleneum of the corpus callosum that the prOposed interference phenomenon between the hemispheres might occur. One feature of the cross-field masking model described above was the prOposal that interference might be produced in the 18-19 areas by a visual noise pattern. This presupposes that, since a noise pattern is to be the effective masking stimulus, it must be interfering with "basic sensory data" (e.g. pattern fragments) of the target stimulus projecting from contralateral 17 rather than some more highly processed code for the stimulus (e.g. in engram form). Is there any evidence to support the con- tention that the spleneum transmits basic sensory data as Opposed to more complex codes? Myers (1962) and Sperry (1964) argue that this is, in fact, the case but their support is based on some rather indirect inferences deriving from the interhemispheric transfer of pattern discrimination learn- ing in chiasm-sectioned animals. Far more direct evidence was provided by ll Berlucchi and Rizzolatti (1967). In an effort to specifically answer the above question they conducted a microelectrode recording analysis Of the receptive fields of the axon fibers in the spleneum (posterior third) of the corpus callosum of the cat. They found all the fields to have pro- perties identical to those of the simple, complex, and hypercomplex cells found by Hubel and Wiesel (1965) in areas 17, 18, and 19 of the cat. The fields extended from the vertical midline to as far as 200 into the peri- phery. While the above obviously supports the hypothesis that the Spleneum may be specialized for the interhemiSpheric transfer of sensory-type visual information, it does not exclude the possibility that yet another commis~ sural structure may transmit more highly coded information between "higher" visual centers of the two hemispheres - in particular, the anterior commis- sure which links the two temporal lobes. The evidence on this point is short but rather dramatic. Gorden, Bogen, and Sperry (1971) describe the testing of two human patients having received partial commissurotomy Opera- tions for the control of epilepsy. Both had only the anterior commissure sectioned with all other structures remaining intact. These patients could easily give a verbal description of visual stimuli flashed to the right hemisphere. However, a third patient (described in the same report) who had received only a sectioning of the spleneum - all other structures re- maining intact - was totally unable to describe any visual material flashed to the right hemisphere. In sum, it appears that interhemispheric communica- tion of visual information depends on the existence of an intact spleneum which has been shown, at least in a lower species, to carry only sensory- type codes. The last events in the processing chain that would lead to the Observation of a cross-field masking phenomenon have involved the implicit 12 assumption that interference with the processing of target information in areas 18 and 19 would necessarily disrupt "higher" processing in the visual centers of the temporal lobe and therefore the output in response form produced by the reSponding hemisphere. Support for this contention derives from a series of cross-lesion experiments on monkeys performed by Mishken (1966). He demonstrated rather conclusively that the contribution of the inferotemporal cortex to the pattern recognition process depends on the neural processing that occurs in areas 18 and 19 (see also Stone & Freeman, 1973). While the above argument makes a case for the hypothesis that mask- ing in areas 18 and 19 could disrupt processing and output by higher centers, it does not exclude the mutually igclusive possibility that yet another pathway, the projection of 18 and 19 to the inferotemporal cortex via the spleneum could provide the target information for continued processing by the "masked hemisphere" (refer to Figure 1). Although the visual noise pattern also projects to this site from the ipsilateral side, the above suggests that masking, in a sense. "engram with engram" could yet be a more effective procedure than masking "engram" with "noise."1 If this were tenable, then masking, for example. a target letter or word with a matrix of random letters would have relatively EBEE effect than masking with a visual noise pattern. A testable hypothesis. Finally, it was mentioned previously that a "natural" callosal func- tion which could potentially account for an interhemispheric interaction 1 For lack of a better word, the term "engram" is used, not in its strictly conventional meaning as a stored bit as a result of learning, but to mean the terminal stage to which a given pattern is processed by the visual system. The word "code" also seems inapprOpriate since the "basic sensory information" is obviously in coded form even in the peripheral energy transducers. 13 between target and mask, was the mechanism providing for a stereOptic inter- action for stimuli that projected binocularly to Opposite hemispheres (a situation that is identically simulated by the cross-field visual masking paradigm). As much of the above data are based on cats and monkeys, and since, as we ascend the phylogenetic scale, the visual system becomes more precise (e.g. receptive fields and nasotemporal overlap are smaller for monkey than for cat etc.), the question arises as to how far into the peri- phery, on opposite sides of the fixation point, can we safely assume a binocular interaction from heteronymous points in the human subject. The data are provided by the unfortunate case of a 14-year-old boy who fell Off his bike and Split his Optic chiasm (Blakemore, 1970). The subject was able to make depth judgments when a slit was briefly exposed in front of his fixa- tion point although its images projected to the temporal retinae of both eyes and therefore projected separately to the two hemispheres (see Figure 4). The amount of disparity that could be tolerated and still give the stereo- sCOpic effect was 6°, i.e. 3° to the temporal side of each fovea. This study provides not only behavioral evidence for an interhemispheric pathway for binocular interaction in man (which correlates with the electrophysio- logical findings of Berlucchi and Rizzolatti, 1968), but an assessment of the spatial parameters relevant to the present project.1 The signal attenuation hypothesis Attenuation of signal and mask due to the longer transcallosal route is perhaps the most critical assumption on which this model is based. Three 1The question arises as to the possible contribution of the "naso- temporal overlap" (the small band at the fovea from which fibers might pass to either Optic tract) which was found to be 0.9° wide in the cat (Stone, 1966). In man and monkey it is purported to be considerable smaller (van Buren, 1963). Functionally, it does not seem to be an important factor in man since Mitchell and Blakemore (1970) found that section of the corpus callosum in man (chiasm intact), "disrupts depth perception entirely in the middle of the visual field." lateral ‘ ' geniculate 13 43 nucleus ‘ I I Q I I I i : I I ..---.u. ' . I I ' . I ' I I I I I I I I I I I . . A : ' A O)--—1---:----O 4: ‘visual I.) 1I I 4) cortex A I I mm. L--- corpus 4 callosum Figure 4. Pathways in the visual system of the retinal images of two objects falling on the vertical meridian, one in front of and one behind the fixation point, for a human with sagittal section of the Optic chiasm. The Object at A, directly in front of fixation point F, projects to the two temporal hemi- retinae and drive the neurons indicated by the solid circles. The neuron at A, left hemisphere, receives an input from corresponding heteronymous areas Of the two eyes, one input deriving from the corpus callosum. Object B can- not be seen due to interruption of decussating fibers at the Optic chiasm. (figure redrawn from Blakemore, 1970) 15 kinds of evidence will be offered for its support. Bremer (1965) electrically stimulated the lgfg lateral geniculate body (see Figure 1) of the brain of a cat and measured the gross evoked potential responses from symmetrical areas of the left and right visual cortex. The oscilloscOpe traces Obtained from the lgfg hemisphere re- presented the direct geniculo-cortical pathway while those obtained from the E£EE£ hemisphere represented the geniculo-cortico-cortical pathway via the corpus callosum (Ibid. 1965, p. 286). The two wave forms were not only similar, as might be expected, but the amplitude of the signal from the sigh; hemisphere (longer pathway) had decreased by a factor of about five when compared to the left hemisphere (direct pathway). Behavioral evidence of the "inefficiency" of the callosal trans- fer mechanism comes from studies by Myers (1962). He found that split- chiasm animals could learn a pattern discrimination task through one eye (while the other is occluded) and then perform the learned task using the Opposite eye even after post-training sectioning of the corpus callosum. In other words, both hemispheres learned the discrimination, one of them through the callosum. However, the "untrained hemisphere" always performed more poorly than the one which had received the discriminanda via the direct retina-cortical route. At the single cell level, Berlucchi and Rizsolatti (1968) provide the most dramatic evidence. These researchers found that cells in area 18 of the chiasm-sectioned cat (see Figure 5) could be binocularly driven from corresponding heteronymous retinal areas (as previously described). Further- more, a cell when stimulated (by natural visual stimuli) via the callosal pathway gave responses that "were in general less brisk and more fatigable" than when stimulated by equal stimuli via the direct retinO-cortical path. 16 left lateral geniculate nucleus r""“l | I I I I I g I I I I I I I I I I I I T L-_-J corpus .callosum microelectrode Figure 5. Example Of a single neuron in the right visual cortex which was driven (Berlucchi & Riszolatti, 1968) by a stimulus crossing the vertical meridian for a cat with sagittal section of the Optic chiasm. The recorded cell could be driven either by the part of the stimulus falling in the left visual field (white area) and projecting to the right temporal hemiretina or by the stimulus in the right visual field projecting to the left temporal hemiretina (black area) and across the corpus callosum. The line at A in- dicates the division between the nasal and temporal hemiretina of each eye. (figure redrawn from Berlucchi and Rissolatti, 1968) Compare Figures 4 and 5 and note that similar events in the visual system are produced by two different stimulus configurations. Figure 4 applies to the case that would represent binocular stereOpsis and Figure 5 represents the situation encountered in the cross-field visual masking paradigm. Chapter 2: EXPERIMENT I For all studies reported in this paper, the "target" stimuli em- ployed were single capital letters for which the left (language) hemisphere is presumed to be the primary, or final, processor for naming the letters and initiating a verbal response. To briefly reiterate the basic paradigm, single letters are to be flashed unilaterally and in random sequence to both left and right visual hemifields under both no-mask and mask (noise pattern in hemifield Opposite the letter) conditions. The main prediction is that with target letter projected to the right and noise pattern to the left hemisphere, decrement in response accuracy should be greater - com- pared tO the no-mask condition - than when the situation is reversed, i.e. letter to left and mask to right hemisphere (Figure 3). The former is pur- ported to represent the low "signal-to-noise" and the latter the high "signal-to-noise" stimulus conditions for the lg££_hemisphere. The results of exploratory pilot work supported, though not con- clusive statistically, the above predictions and provided information for establishing the base-line parameters to be used for the present studies. The goals of the experiments reported here were (a) to establish, if possible, statistical confirmation of the predicted "effect," (b) to provide a system- atic investigation of some stimulus parameters which may be relevant to in- creasing or decreasing the "effect," and (c) to test some related theoretical assumptions about the neural mechanisms potentially involved. The purposes of Experiment I were (1) to accumulate sufficient data to make a statistical determination of the tenability of the "effect" and (2) to investigate the possible effects Of varying brightness relationships between target and noise pattern. Brightness was varied by holding target 17 l8 intensity constant for different noise-pattern intensities and by holding noise intensity constant while varying target intensity. The preliminary supposition was that, as with other masking paradigms, the greater the brightness difference between target and mask, the greater would be the masking effect. In the present case, however, the issue becomes consider- ably complicated by the fact that the (presumed) masking site is exclusively cortical. The present goal was to find a set of relative brightness para- meters which might suggest the existence of an Optimum set for obtaining the differential effect. METHOD Subjects The subjects were 16 paid volunteers from two undergraduate psychology classes at Michigan State University. All were males, right- handed, as determined by the Crovitz and Zener (1962) questionnaire for hand dominance, and naive to the purpose of the experiment. Apparatus A three-channel Scientific Prototype (Model GB) Auto-TachistoscOpe was used. Each field was back-lit through the diffusion screen (with color-matching filters) by the standard fluorescent bulbs (GE-FTA-S with painted silver stripe between the electrodes). Although not measured, the rise and decay times are claimed to be considerably less than 1 ms each. Each of the two "stimulus" fields were modified to obtain more equal left-right half-field brightness by repositioning the right member of the vertically mounted pair of bulbs and by inserting a mat white strip of cardboard behind each bulb pair. Maximum and minimum stimulus field bright- Iaesses, measured through the split-beam binocular eyepiece with a Salford lilectrical Instruments Company spot-photometer, were 2.50 footlamberts (ft.l.) l9 and 0.40 ft.l. respectively. For two Of the fields, stimuli were presented on 24x36 umlslide transparencies automatically fed from lOO-slide "rototrays." Stimulus materials The fixation stimulus viewed by the subject was a dark field, and a lighted circle in the center (0.44° visual angle in diameter) within which was an "x" containing a "dot" (0.088° visual angle) in its center (Figure 6). The subject also saw a small lighted bracket in each of the four corners of the rectangular field. The area within the brackets re- presents the unoccluded portion of any stimulus field as seen through the Figure 6. Fixation stimulus with brackets for parallax adjustment. binocular eyepiece and permitted the subject to make minor self-centering head adjustments before each exposure sequence. The magnified (through the scope Optical system) portion of this unoccluded viewing area had an "apparent" viewing distance of 30 inches and projected a visual angle of 2.9°x5.25° (laterally) at the retina. The lighted areas (center circle and brackets) Of the fixation field measured 0.40 ft.l. while the "black" area of the transparency (found under higher luminance conditions to trans- mit 0.81 Of the back light) was 0.0032 ft.l. The target stimuli (see example in Figure 7) consisted of a rototray Of 100 slide transparencies, each with a single black letter Of the alphabet 20 against a "white" background on the left or right half of the slide while the half-aide Opposite the letter was "black". The center point of each letter was 1.70 to the left or right of fixation as viewed through the scope. The letters subtended an angle of 0.340 in height and a maximum of 0.340 in width. Thickness of the letter features was 0.030 visual angle. L______- target stimulus mask stimulus no—mask stimulus Figure 7. Example of the target, mask, and no-mask stimuli. A second rototray, for the remaining stimulus channel, contained a set of 50 blank slides, blank the entire area, and a set of 50 masking stimuli (Figure 7) which were black on either the left or right side and contained a visual noise pattern, the area of which was 501 white and 50% black, on the Opposite side. This pattern is identical to that used by Schiller (1965) for dichoptic masking Of single letters. A third rototray identical to the second except for a one to one reversal in order of mask and blank stimuli was also used. 21 All stimuli were prepared in black-white reverse on mat white paper with black marking pen and photographed with Kodak High Contrast Copy Film 5069. The negatives then comprised the stimulus material. The black areas of the mask and blank slides were exposed and develOped to the same density so that they would pass an equal amount of light from the rear diffusion screen in their specific channel.1 In the center of the target and masking fields, a black vertical strip, projecting a visual angle of 1.00 at the eyepiece, was mounted immediately in front of the stimulus position. This strip was to block foveal transmission from either half-field resulting from whatever naso- temporal overlap exists in man (estimated to be considerably less than the 0.90 found in the cat; van Buren, 1963), or from microsaccadic eye movements, 7 to 10 minutes to either side of fixation (Ditchborn & Gins- burg, 1953; St. Cyr & Fender, 1969). The resultant composite viewed by the subject on a "mask" trial and target presentation on, say, the left side is illustrated in Figure 8. The remaining unoccluded portions of the Figure 8. Composite view of the stimuli as seen thrOugh the tachistoscOpe On a "masked" trial with target projecting to the right hemisphere. LThis was crucial for the purpose of maintaining a constant back- ground illumination in the half-field of the superimposed target letter for both mask and no-mask trials. 22 mask projected a visual angle of 2.130 in width and 2.900 in height. A no- mask trial appears identical except for replacement of the noise pattern by a "black" half-field. Procedure As the subject sat in a chair facing the tachistoscOpe, a head rest, chin rest, and the interocular separation of the eyepieces were all adjusted so that the subject, by occluding first one eye, then the other, could see all four brackets of the fixation field with each eye separately. The room lights were then turned off and the subject was given a brief period to dark-adapt to the brightness level of the fixation field. At the beginning of each trial, the subject binocularly viewed the fixation slide. At the signal "ready", the subject made minor head adjustments (if necessary) to insure a clear view of all four reference brackets. About 1% seconds after the ready signal, the subject was signalled to "focus" on the center dot, and within another second the stimulus fields were fired by the experimenter. On a "masking" trial, the noise pattern fired on one side for a total duration of 200 ms. Fifty ms after onset of the noise pattern the target flashed on the Opposite side for a duration of 3.5 to 9.0 ms depend- ing on brightness conditions and threshold level for a particular subject. On a "no mask" trial, all conditions were the same except for replacement of the noise pattern with an all "black" slide (i.e. the masking field still fired for the same time duration to maintain constant target back- ground for 2252 "mask" and "no mask" trials). The fixation stimulus remained continuously lit. At the termination of a 750 ms interval (from trial onset), the slide changers automatically advanced the target and Inask - no-mask stimuli for the next trial. The subject's task was to 23 call out a letter as quickly as possible and within the 750 ms interval, i.e. before he heard the slide changers advance. The subject's response was manually recorded during an intertrial interval of approximately 4 seconds. Three aspects of the experiment were emphasized to the sub- ject as having the following order of importance: (1) he must constantly maintain the fixation point for the period beginning with the "focus" command and ending with the sound of the slide changers, (2) for each trial, he was to make his best guess as to what the letter might have been, regardless of the stimulus conditions and regardless of whether or not he actually "saw" a letter, and (3) he was to call out his best guess before he heard the changers advance. The target set Of stimuli contained 25 letters of the alphabet (excluding Q), each letter appearing 4 times, twice on the right and twice on the left. The order of the letters and their right-left posi- tion was randomized with the restriction that the same letter did not appear more than twice in succession, and not more than three right or three left presentations occured in succession. The "mask - no mask" stimulus set was randomly arranged with the restrictions that (a) each letter was "masked" once on each side, and (b) no more than three mask or three nO-mask conditions occurred in succession. The second "mask - no mask" set (100) was identical to the first except for a one to one reversal of the 50 noise and 50 blank stimuli. In other words, a letter in a given serial position that was masked on the first set of 100 trials, was not masked the second time around. In sum, the right-left, mask - no mask, and serial position of the mask - no mask conditions were counter- balanced for each letter, with conditions and letters Otherwise occurring in random sequence. 24 At the beginning of the first session, 50 to 75 practice trials were given to acclimate the subject and to determine the target eXposure time which elicited a probability of correct responding at approximately 0.5 averaged across all conditions. After a brief rest period, 100 test trials were administered with the first set of "mask - no mask" stimuli. Occasionally, the exposure times were varied by 5% increments to maintain the threshold responding level. Following a 5-10 minute rest period, another 100 trials were given with the second set of "mask - no mask" stimuli and rototrays Operating in the reverse direction. This ended the first session of 200 test trials. A second session was given within 3-4 days (but never on the same day) with the direction of the rototray for a particular set of "mask - no mask" stimuli reversed from what it ‘was the first testing day. A 30-40 trial warm-up was required to obtain a threshold level of responding at the previously determined target ex- posure time. For gpph of the four 100 trial sets, the stimulus rototray ‘was started from a random position. Each of the two daily sessions lasted approximately 1% hours. The 16 subjects were randomly assigned to four groups with 4 sub- jects in each group. The groups (conditions) varied with respect to the relative brightness difference between the noise pattern and target stimulus fields as follows. Group I: masking field - 2.50 ft.l., target field - 0.40 ft.l.; Group II: mask - 1.05 ft.l., target - 0.40 ft.l.; Group III: mask - 2.50 ft.l., target - 0.20 ft.l.; Group IV: mask - 2.50 ft.l., target - 0.10 ft.l. For Groups I and II, target brightness was held constant while mask brightness was decreased by 501 of the difference between the two. For conditions I, III, and IV mask brightness was held constant while target brightness was decreased in 50% increments from 0.40 to 0.10 ft.l. by Kodak neutral density filters 0.30 (50% transmission) and 0.60 (25% transmission) respectively. 25 RESULTS AND DISCUSSION Percent correct, based on 100 Observations per condition per sub- ject, was averaged across the four subjects in each group. These data are tabulated for each of the four groups in Table 1 (within the rec~ tangles). A comparison of the mask versus nO-mask conditions averaged across hemispheres is made in column 3. For each group, the third row Table 1. Percentage of correct responses for left versus right hemisphere and mask versus no-mask conditions for Groups I-IV, percentage decrement produced by the mask, average across hemisphere conditions, difference in percentage decrement, and mean target exposure time at threshold. diff. in mean T exp. T to T to average I decr. time (ms) @ left H right H across H RH-LH threshold Group I no-mask 63.8 59.0 61.4 til-2.50 ft.l. mask, 43.3 34.3 38.8 9.7 4.9 'T-0.40 ft.l. Z decr. 32.2 41.9 37.0 Group II no-mask 56.8 55.8 56.3 bi-l.05 ft.l. mask 46.8 40.8 43.8 9.3 4.4 'T-0.4O ft.l. Z decr. 17.6 26.9 22.3 Group III nO-mask 67.5 57.0 62.3 III-2.50 ft.l. mask 47.0 34.8 40.9 8.6 8.3 ‘T-0.20 ft.1. 2 decr. 30.4 39.0 34.7 Group IV no-mask 64.5 53.0 58.8 ‘M-2.50 ft.l. mask 49.0 34.8 41.9 10.4 10.6 T-0.10 ft.l. 2 door. 24.0 34.4 29.2 Groups I—IV no-mask 63.1 56.2 59.7 average mask 46.5 36.1 41.3 9.4 1 decr. 26.4 35.8 31.1 Note. H-mask; T-target; H-hemisphere; Rsright; L-left; Zdecr.=percentage decrement produced by mask in the table shows the percentage decrement in responding level produced by masking - using the no-mask condition as baseline responding level. The fourth column shows the difference in percent decrement, right minus 26 left hemisphere, and the fifth column shows the average target exposure time required to maintain threshold level of responding. A "one between - and two within - subjects variables" analysis of variance (see Myers, 1967, pp. 198-209) is summarised in Table 2. Across hemispheres and groups, the average percent correct for the no-mask condition was 59.71 and for the mask condition 41.32, a decrement from the no-mask baseline of 31.11 (p410.0005). The non-significant AC interaction term indicates that the magnitude of the over-all masking effect did not differ for Groups I-IV, the different mask-to-target bright- ness ratios. This point will be discussed subsequently. Although not of primary interest to the purpose of this study, the usual left hemisphere (right field) superiority for recognition of alphabetic material was Observed (p410.005). The non-significant AB interaction term indicates that this effect did not differ for Groups I-IV. Normally, such simple alphabetic material as single alphabetic letters presented over periods of time which afford extended practice (as in this study) shows considerably smaller left-right effects than more complex stimuli such as words (left hemisphere) and spatial stimuli (right hemis- phere) presented less frequently. Some of the pilot subjects receiving 400-800 presentations and following experiments employing a smaller number of subjects did not show a significant over-all left hemisphere superiority. The crucial interaction, BC (Table 2), hypothesized from the model, was significant in the direction predicted (p410.05). The masking decre- ment, averaged across groups, for the left hemisphere (target to left h, mask to right h) was 26.41 below the no-mask baseline while a 35.82 de- crease was obtained for the right hemisphere (target to right h, mask to left h). The former represents the high and the latter the low Table 2. for Experiment I Source Total Between subjects (8) Groups41A) S/A Within subjects Hemispheres (B) A! SB/A Mask conditions (C) 59 sc/A g9 £59 SIC/A *p<0.05 (one-tailed); U 12 1 3 12 **p<0.005 (one-tailed); 27 Analysis of variance summary SS 9257.980 541.230 25.668 515.563 8716.750 1198.886 219.801 871.563 5383.886 255.301 570.063 47.270 6.418 163.563 1Since variances decrease as proportions deviate from 0.5, the la 8.556 42.964 1198.886 73.267 72.630 5383.886 85.100 47.505 47.270 2.139 13.630 I'II 0.119 (ns) 16.507 1.009 (ns) 113.333 1.791 (ns) 3.468 0.157 (ns) based on percentage correct scoresl, 2 ln 4.063** 10.646*** 1.862* *** p<0.0005 (one—tailed) analysis should more appropriately be made on the arc sin transforms of the Such analyses were conducted on the transformed data for this and succeeding experiments and led to no difference in conclusions drawn or significance levels reported observed proportions (tabulated here as percentages). herein. are based on the data in their originally observed form. 2The one-tailed t hypotheses tested in this dictions derived from the left hemisphere should be right, and masking should one of all possible within-subjects interactions had been predicted, i.e. that masking should be greater when targets are projected to the right hemis- The apprOpriate analysis of this interaction for this model (Myers, pp. 337-345) is a one—tailed test of the apprOpriate phere as compared to the left. orthogonal comparison of the four interaction means. produce a decrement. For ease of interpretation, then, all statistical analyses reported tests are reported for the following reason: all analysis are based on "one-tailed" (a priori) pre- model, e.g. targets projected initially to the more accurately reported than when projected to the Most important, one and only Since that test as well as the preceding two are based on one degree of freedom, they should be algebraically identical to taking the square root of P and doing a one- tailed t test. and the orthogonal comparisons and producing identical results. This was confirmed for these data by doing both the anova (See Hays, 1963, Ch. 14, for a more thorough discussion of orthogonal comparisons for interaction terms.) 28 "signal-to-noise conditions" for the left hemisphere. The average differ- ence in percent decrement, right minus left hemisphere, was 9.42. This difference, a single index of the direction and magnitude of the inter- action, is tabulated in column 4, Table l, for each of the groups. This column, and the non-significant interaction between groups (A) and the within-subjects (BC) interaction, i.e. A(BC), shows that the hemisphere)( mask interaction did not differ with varying target-to-mask brightness ratios. The significant hemispheres x masking interaction effect, and its prediction in advance of the empirical observation, supports the major con- tention of the model develOped herein, that visual masking may occur inter- hemispherically as well as intrahemispherically (the dichOptic paradigm). However, the small magnitude of the effect, as observed so far, would seem to preclude its utility as a general research tool for the investigation of unilateral information processing. Each group taken separately (n-4) and all but two individual subjects1 failed to produce a statistically significant interaction. It should also be noted here that two of the 16 subjects in this sample (12.51) produced a non-significant interaction in the direction apposite to that predicted by the model. This is in close agreement with the observation (Hilner, et al., 1964; Kimura, 1961) that approximately 101 of right-handed males have the language center in the right hemisphere as determined by unilateral sodium amytal injection into the left or right carotid artery. The failure to find that varying mask-to-target brightness ratios had an influence on the hemispheres x mask interaction (column 4, Table l) 1For individual subjects, the orthogonal comparison was made on the arc sin transform of the observed prOportions. In this situation, analysis based on single data points (angles) is made possible by the fact that the error variance is known to be 821/n where n-number of observa- tions making up each prOportion (100 in this case). 29 as well as on the over-all masking effect (column 3, Table l) was initially disappointing in light of the original goal to find a relevant stimulus parameter for controlling or increasing the magnitude of the effect. How- ever, some work by Schiller (1969 - unknown to this writer at the time of this research) and a l9—experiment report by Turvey (1973) established rather conclusively that relative energy level between mask and target is a totally irrelevant dimension for "central" (cortical) visual masking - via the dichOptic procedure. The parameters that do influence cortical masking will be discussed in later sections. 0n the other hand, they find that "peripheral" (retina and neural pathways leading to the visual cortex) visual masking is most dependent on the relative energy level between mask and target - via the monOptic paradigm (the brighter the mask in relation to the target, the stronger the effect). The failure to find group differ- ences in this experiment supports, in view of the above findings, the thesis that the observed masking was central (and therefore interhemis- pheric in this case) rather than peripheral. Briefly, two additional aspects of the between-groups comparison deserve mention. For Groups I, III, and IV, mask intensity was held con- stant while target intensity was decreased by 501 increments. One might predict that the effect of halving target intensity might simply be that of requiring a doubling of the target exposure time required to maintain threshold responding levels. This statement is a parallel to Block's law which states that the time and intensity of a light flash required to produce a constant visual effect are reciprocally related. What was not known, of course, was whether the same relationship would apply to the detection of information from briefly exposed targets. If the law does hold, then the total stimulus energy of the target would remain constant 30 as well as the mask-to-target ratio for Groups III and IV when compared to I. Examination of the mean target exposure time required to maintain threshold for Groups I, III, and IV (column 5, Table 1) reveals that for Groups I and III the law holds approximately; but not so well for the I and IV or the III and IV comparisons. A possible explanation for this could be that if the light intensity of the bulbs was still rising over the time intervals used, then a longer exposure would also be a more intense one.1 For Groups I and II, target brightness was held constant while mask brightness was decreased by 501 of the mask - target difference. The numbers in column 3 (Table 1) show that over-all masking was somewhat less for Group II (not significant) than for any other group which suggests, according to the findings of Schiller (1968) and Turvey (1973), that some small percentage of the variance attributable to masking may have been "peripheral." The most plausible peripheral effect in this eXperiment would be that due to stray illumination within the eyeball. (Lateral in- hibition is discarded because of the large separation retinally between mask and target.) Nevertheless, target exposure time to maintain threshold remained unchanged, and more important, the differential effect between hemispheres (column 4, Table 1) did not change. Obviously, it is difficult to account for the differential masking effect with a peripheral hypothesis. Error analysis 1 Central neurOphysiological or parallel "cognitive" mechanisms that might account for "how" the masking effect is accomplished has not lBrightness levels of the stimulus fields were measured with bulbs constantly lit. 31 been a major (though admittedly important) theoretical issue of this paper except for the following implicit assumptions which determined the choice of the masking pattern itself. It was assumed that "cross-field" visual masking could occur relatively early in the succession of central processing stations, that is in the areas immediately succeeding the primary (1?) visual projection area (18 and possibly 19). It follows, if we may project Hubel and Wiesel's (1968) findings on the monkey to man, that these areas are still involved in processing the pattern ele- ments of a visual stimulusl. Consequently, the selected mask contained pattern elements whose size and density, when superimposed on the target, effectively "broke up" the raw stimulus elements of the target. Alterna- tive masks would, when coded, be those which would produce confusability of sounds, for example, or of meanings, for instance a matrix composed of random alphabetic letters. From the above, then, it was supposed that "masking" would some- how degrade or decrease the amount of physical stimulus information avail- able from the target pattern. If that is the case, it might be expected that the "quality" of a subject's forced guess when an error is committed might depend on the amount of physical stimulus information available to him, and therefore on whether a particular trial was a mask or no-mask condition. Specifically, it could be hypothesised that of the total number of errors made, the percentage of errors that appear to be depend- ent on physical stimulus information (report of letters with similar shape 1Not until the infero-temporal cortex has been reached (in monkey), has it been found that receptive fields of individual neurons attain a specificity for stimuli more complex than single lines, bars, and edges. For example, the silhouette of a hand - more specifically, a monkey hand (Gross et al., 1972). 32 but actually different from the target letters) might be higher under the no-mask as compared to the mask conditions. 0n the other hand, if masking affected primarily a distraction of attention, say, then errors related to the similarity-in-shape dimension might distribute randomly across mask conditions. It should, however, be emphasized that such an analysis based on confusability of shapes per se does not permit dis- crimination between central versus peripheral mechanisms as explanatory constructs. It can be argued for example that a perceived decrease in contrast between target and "ground" produced by stray illumination with- in the eye could account for a decrease in information about target features. Rather, it should support or reject the notion that some process degraded the content of physical stimulus information contained in the target. For present purposes, then, all errors were divided into two categories: (a) those in which there appeared to be a basis for physical similarity between the letter guessed and the actual target, the stimulus- dependent category; and (b) all others, the stimulus-independent category. The letters classified as stimulus dependent (listed in Appendix B) were selected without prior knowledge of experimental conditions into which they fell1 and by the following criteria: similarities (judged by the experimenter) based on addition or subtraction of a small number of frag- ments, e.g. E and B, C and G and 0, F and P, etc.; similarities based on "almost alike shapes," e.g. A and H, K and x, U and V; mirror image re- versals, e.g. A and V, M and W, S and Z, J and L; and letter pairs which upon examination of the response data showed a high frequency of 1The raw data sheets contained only the target letters, the re- sponse letter for each target, and a "right" or "wrong" code for template scoring. 33 confusability but could also be justified on the basis of some physical similarity between these particular hand-drawn letters, e.g. H and M or W, S and B, R and B, O and U, etc. For each subject and each "within-subjects" condition the per- centage of total errors falling into the stimulus-dependent category were computed. These data averaged across subjects and groups are presented in Table 3. To reiterate, the larger the percentage of stimulus-dependent errors, the greater the stimulus (target) information is presumed to have been received by the subject. Employing this new index as dependent vari- able, an analysis of variance, identical to that reported earlier in this section, was performed. Table 3. Percentage of stimulus-dependent errors for left versus right hemisphere and mask versus no-mask conditions averaged for Groups I-IV, percentage decrement produced by the mask, average across hemisphere con- ditions, and difference in percentage decrement.(Error analysis 1) diff. in T to T to average 2 decr. left fl right R across R RB-LH Groups I-IV no-mask 50.0 42.6 46.3 average mask 39.4 33.4 36.4 0.4 2 decr. 21.2 21.6 21.8 Note. T=target; H=hemisphere; R-right; L=left; 1 decr.-percentage decre- ment produced by mask The results showed that masking produced a 21.81 decrease (p4:0.001)1 in stimulus-dependent errors, and a left hemisphere superiority was ob- tained (p4(0.0005). It was not expected that this index would prove sensitive to left-right field differences. The within-subjects hemis- pheres x mask conditions interaction term was not significant, but the triple interaction was (p4<0.005). All other terms were not significant. lAll significance levels reported for the error analyses are based on the directly obtained F ratios. 34 The significant triple interaction showed that the hemispheres X masking interaction differed for the varying mask-to-target brightness ratios. Examination of this variation and also the over-all masking effect for the different groups revealed no monotonic changes with increasing or decreasing mask-target brightness ratios.1 The above finding that masking produces a significant decrement in the percentage of stimulus-dependent errors supports the hypothesis that the neuronal representation of the visual stimulus elements of the target pattern has been degraded by the mask. Further support for the contention that the present index reflects changes in stimulus degrada- tion derives from the finding that the condition requiring the longer callosal transfer route of target information to the left hemisphere also produced a significantly lower percentage of stimulus-dependent errors . Error analysis 2 Even though the main effects variables were significant in error analysis 1, the between-groups comparisons made no particular sense, and consistency between subjects was quite poor. Although it is perhaps un- reasonable to expect an alternate dependent variable to behave as nicely as the original, it was thought, after reexamination of the original criteria for defining stimulus-dependent errors, that the index might be improved. It was noticed that many letter pairs which were similar on the shape dimension afforded a great deal of confusability on the auditory dimension as well, for example E and B, P and B, M and N, and C and G. A second set of error categories was constructed (Appendix B) in which auditorily confusable letter pairs were eliminated as well as 1Summary tables and the analysis of variance are available in Appendix C. 35 mirror image reversals and any others which seemed to have a more tenuous shape relationship. The final set of errors that could be classified as stimulus-dependent guesses was reduced from the original 72 to 28 in number. The results, averaged across subjects and groups appear in Table 4. An analysis of variance1 identical to the previous one was performed. The findings were, in essence, identical to those of the first error analysis. The masking effect was significant (p<:0.001), Table 4. Percentage of stimulus~dependent errors for left versus right hemisphere and mask versus no-mask conditions averaged for Groups I—IV, percentage decrement produced by the mask, average across hemisphere con- ditions, and difference in percentage decrement. (Error analysis 2) diff. in T to T to average 1 decr. left H right B across fl (RH-LN) Groups I-IV no-mask 29.8 26.2 28.0 average mask 23.7 18.3 21.0 9.7 2 decr. 20.5 30.2 25.4 Note. T=target; H-hemisphere; R-right; L=left; Z decr.- percentage decre— ment produced by mask the left hemisphere superiority was even more significant (p<:0.001 com- pared to the previous 0.005), and the triple interaction remained signifi- cant (p‘<0.05), though less so. The fourth column of Table 4 shows that the difference in percent decrement, right minus left hemisphere, was in the direction that would be predicted by the model and was as large in magnitude, 9.7%, as that obtained with the original dependent variable, probability correct. Nevertheless, the hemispheres X masking conditions interaction was non-significant, and the between-subjects variation was much larger for this index as compared with the original dependent variable. 1available in Appendix D Recent writers have offered some interesting speculations about the nature of the visual masking process. Turvey (1973) reviews two kinds of masking processes originally elucidated by Kahneman (1968), both of which have received wide support. The first view is that the mask mixes with the "sensory" information of the target stimulus. "The idea is that two stimuli which follow one another in rapid succession are effectively simultaneous within a single frame of 'psychological' time, analogous to a double exposure of a photographic plate" (Turvey, 1973). This is the integration hypothesis. The second, or interrup- tion hypothesis, states that the (usually aftercoming) mask simply terminates central processing of the target by occupying, so to speak, the same processors. In essence, the time needed to completely process the target has been cut short. While Turvey stipulates that these hypo- theses need not be mutually exclusive, he views the integration process as being primarily a function of "peripheral" mechanisms and inter- ruption as a function of "central" processes. He supports these views by the following argument: masking by interruption should depend prim- arily on stimulus onset asynchrony (80A) or the onset-onset time rela- tionship between target and mask, whereas integration should more likely be controlled by relative stimulus energies of target and mask. He then proceeded to demonstrate that SOA primarily affected central masking (via the dichOptic paradigm) and that relative mask-target energy levels affected only peripheral masking (via the monOptic procedure). Neither variable was effective with the alternate procedure (within the range of time and energy relationships employed). Though relevant to the question of masking mechanism introduced earlier, the basic model with which the above issues are concerned is 37 backward masking - a paradigm most useful for the study of information processing. The model of this paper (as presented so far) differs slightly in that it is concerned only with the limiting case of back- ward masking, that is, simultaneous or overlapping coexistence of target and mask representation in the central processors. The question of $2535- ruption in central mechanisms presently gives way to one of integration, and it is now obvious that the position taken for the present model (and supported by the error analysis) is an integration hypothesis. The con- cept of masking by integration requires that the iconic storage of the target be contaminated at the input stage by sensory information from the mask (Turvey, 1973) and, in the present case, it must be effected centrally by separate neural input pathways. Can such a position be rationalized on the basis of present neurOphysiological knowledge of the visual system? Recent work by Schiller (1968) provides a plausible explanation. In this study, he attempted to find electrOphysiological correlates to the visual masking process by measuring responses of single neurons in the lateral geniculate nucleus and of binocularly driven cells in area 17 of the cat's visual cortex. After plotting the receptive fields of individual neurons and studying their response to the target and then target plus mask, he found that by making the contours of the pattern mask more and more similar to those of the target, the cells failed to respond differentially to the two stimuli in combination. In other words, the target lost its separate "identity" in terms of the response charac- teristics of the cell. His conclusion: "Pattern masking is brought about primarily by a failure of units, whose receptive fields lie near the contours of the figure, to respond differentially to target plus 38 mask and mask alone. It appears that this effect may take place either cortically or at earlier levels in the visual pathway depending on the mode of stimulus presentation. Under dichOptic conditions, the effect occurs at the cortical level, while under monOptic or binocular condi- tions it may take place earlier." (Schiller, 1968, p. 162.) Behavioral evidence supporting a central (cortical) masking-by- integration hypothesis comes from this experiment, of course, and the studies by Schiller (1965) and Turvey (1973). As discussed earlier, Schiller found that dichoptic masking, while successful with a pattern stimulus, was ineffective when a bright flash of light was used as mask. Further, Turvey (1973) was unable to obtain a dichOptic effect with a random dot noise pattern. Only when the contours of the mask were re- designed to simulate the contours of his target letters (a random array of short straight lines with approximately the same thickness as the target letter features), was he able to obtain a dichoptic effect. These observations, then, that structural stimulus similarity between mask and target are critical for dichOptic masking, support the concept of "sensory" integration of stimulus elements at the cortical level. Chapter 3: EXPERIMENT II The results of Experiment I failed to satisfy one of the goals set for this series of experiments, that was to find a physical stimulus parameter that would control the magnitude of the differential cross- field masking effect. While the differences in mask-target energy levels failed to influence masking, this finding ironically (on the basis of unknown, Schiller, 1968, or unavailable, Turvey, 1973, research) sup- ported the hypothesis that the obtained effects were central rather than peripheral. The purpose of Experiment II was to investigate the stimu- lus onset asynchrony (SOA) variable, i. e. the onset-onset relationship between target and mask, as being possibly relevant to the control of the differential cross-field masking effect.1 The new model is basically simple in concept. Suppose, for pur- poses of illustration, that the target and mask intensities to be used are equal and that the callosal transfer time between hemispheres for stimuli of the particular intensity is known to be 8 ms. If the onset of a target in the left visual field precedes the onset of the mask in the right half-field by 8 ms, then their arrival in the 15;; hemisphere should be simultaneous (see Figure 9) due to the additional 8 ms required for the target to transfer from the right to the left hemisphere. For present purposes, the term stimulus arrival asynchrony (SAA) will refer LThe finding of Turvey (1973) that central effects depended only on SOA values (and not on relative energy levels) was not available at the time this experiment was conducted. 39 40 to the relative arrival times between two stimuli for a given hemisphere. In the preceding illustration SAA for the left hemisphere is zero. Pro- ceeding now to the right hemisphere, the mask arrives 16 ms after the target QD mask l O t (1" to+ 8 ms responding hemisphere SAA - 0 SAA ' 16 ms (target preceding mask) Figure 9. Expected stimulus arrival asynchrony of target and mask stimuli at the two hemispheres for onset of a left-field target preceding onset of a right-field mask by 8 ms with an assumed callosal transmission time of 8 ms. Note. t - arbitrary onset time of first (target) stimulus; tl- onset time of mask; SAA - stimulus arrival asynchrony target (Figure 9) because (a) mask gppp£_was 8 ms later than that of the target and (b) it takes the mask an additional 8 ms to cross from left to right hemisphere. Consequently, the condition that obtains for the 15;; hemisphere is one that simulates the limiting case (maximum effect) for backward masking with a noise pattern (Kahneman, 1968), i.e. simultaneity of target and mask. SAA for the right hemisphere now corresponds to the backward masking paradigm which is known to produce a smaller effect than when SAA is zero (Schiller, 1965; Kahneman, 1968; Turvey, 1973). Backward 41 masking with noise pattern is a monotonically decreasing function of SOA1 (Kahneman, 1968). To follow the same reasoning as with the first model, we also wish to directly assess the relative effect of the 16 ms SAA condition. This is accomplished by reversing the target and mask positions of the above example and maintaining the same target-mask onset-onset relation- ships (Figure 10). NOw, SAA-0 for the right hemisphere, but more import- ant, the mask follows the target by 16 ms in the left or responding mask qp target t1. t0+ 8 1118' I l l l l l l responding hemisphere SAA - 16 ms (target preceding mask) Figure 10. Expected stimulus arrival asynchrony of target and mask stimuli at the two hemispheres for onset of a right-field target preceding onset of a left-field mask by 8 ms with an assumed callosal transmission time of 8 ms. Note. to- arbitrary onset time of first (target) stimulus; tl' onset time of mask; SAA - stimulus arrival asynchrony 1In the dichoptic paradigm SOA is considered equal to SAA, as defined here, for stimuli of equal intensities because the pathways are of equivalent length. 42 hemisphere.1 The decrement in responding accuracy (no mask to mask) should now be less than when SAA-0 for the responding hemisphere. It follows that the conditions of Experiment 11 should be super- ior to those of Experiment I for producing the desired differential mask- ing effect. SOA for Experiment I was 50 ms, mask leading target. This value is so much greater than any estimates of callosal crossover time (6-10 ms, Efron, 1963; Jeeves & Dickson, 1970) that in effect mask should overlap the target timewise in both hemispheres. That is, the mask arrives before, is "on" during, and is present after the brief target representation in both hemispheres. Hence, the only differential ad- vantage would be that due to the predicted difference in signal-to-noise ratios. It is prOposed that the conditions of Experiment II superimpose a second advantage over that of Experiment I. With target in the left and mask in the right fields, the low signal-to-noise ratio for the left hemisphere is now combined with the simultaneity of arrival condition for that hemisphere. This pair of conditions (for the left hemisphere) is expected to produce little difference compared to Experiment I since simultaneity is little different from overlap. (Forward masking produces relatively small effects centrally, Kahneman, 1968; Turvey, 1973.) On the other hand, the pigg signal-to-noise condition for the right hemis- phere (or for left hemisphere with mask and target positions reversed) is combined with the asynchrony in target-mask arrival time,and the combina- tion should produce a smaller masking effect than for Experiment I. The predicted result is a larger differential masking effect between hemis- phere conditions. 1This hypothesis includes the implicit assumption that trans- mission time of target information from right to left hemisphere is the same as from left to right - and similarly for the mask. 43 The strategy for Experiment II was to select what appeared to be the best of the four groups in Experiment I on the basis of differen- tial masking effect, determine the apprOpriate SOA (to produce simultaneity in arrival at the left hemisphere) for the exact mask-target brightness. levels employed for that group, run a new group of four subjects with the new SOA value, and compare the two groups. All conditions for the two groups would be identical except for the different target-mask SOA values. Group III from Experiment I was selected for comparison pur- poses.1 All that remained was to determine the apprOpriate SOA value. The report by Efron (1963a) provided the theory and methodology. Determination of SOA value to produce left-hemisphere simultaneity Efron (1963a) prOposes that for two stimuli traversing different neural pathways to be judged as simultaneous in occurrence, they must meet at some common point in the central nervous system. He reviews the research of various investigators dating back to 1912 which shows that (a) estimates of callosal transmission time deriving from various reaction time experiments (stimulus to one hemisphere requiring response of either the ipsilateral or contralateral hand) appear to be on the order of 2-6 ms and (b) when bilaterally symmetrical points of the body are stimulated, the left must precede the right by 2-6 ms in order to be judged as simul- taneous for most peOple. He suggests therefore that the location for the judgment of simultaneity must be the hemisphere that is dominant for 1Time and subject scheduling required that this selection be made prior to the detailed statistical and error analyses. Although the four groups did not differ statistically, it turned out that Group II would have been a slightly better choice. Group II came closest to producing a significant hemispheres X mask conditions interaction term when the groups were analysed separately, and it produced a larger differential effect under error analysis scrutiny than any other group. However, the overall analysis suggests that for practical purposes, any of the four groups would have sufficed. 44 speech (the left) since the left-right SOA corresponds to the time re- quired for stimulus information to travel from the right to the left hemisphere. He supported these contentions by showing in his own series of experiments that for Eiggt-handers (presumed left hemisphere speech dominant) a visual stimulus flashed to the left field had to precede a stimulus flashed to the right field to be judged simultaneous. But for left-handers (30-401 presumed right hemisphere speech dominant) the right-field stimulus had to precede the left. In a companion paper (Efron, 1963b) he determined that visual stimuli of lower intensity re- quired a longer callosal transmission time than for higher intensity and that for left-right field stimulus pairs of unegual intensity, the SOA requiring a simultaneity judgment could be predicted by algebraic summa- tion (or subtraction) of callosal transmission times for the respective stimuli. To determine the appropriate SOA value required for Experiment II, Efron's (1963a,;nu 264-265) procedure was replicated on two graduate students - but employing the brightness values used for Group III, Experi- ment I. The average SOA value obtained would be used as a constant best estimate value for all subjects of Experiment II. It should be noted that for present purposes it is not necessary to obtain the callosal transmission time for the less intense versus the more intense stimulus and then calculate the 35355 SAA time for the right hemisphere. Rather, it is only required to find the SOA value for the two stimuli to produce simultaneity at the left hemisphere. Method and results. Two right-handed male graduate students with 20-20 corrected vision and who were experienced at making delicate psycho- physical judgments were run as subjects. Both were naive to the purpose 45 of the experiment. The apparatus was that used for Experiment I. The stimuli consisted of two small back-lit circles, one in each visual half- field. The diameter of each circle (prepared by cutting a hole in black light-impervious paper) was identical to the height of the target letters; and their location, equidistant from the fixation point, was identical to that of the target letters which appeared (Experiment I) in the left and right visual fields. The brightness of the left-field circle was always 0.20 ft.l. and of the right was 2.50 ft.l. (the brightness of target and mask respectively of Group III, Experiment I). Each circle would be flashed for a duration of 7 ms. The procedure employed was (after Efron, 1963a, pp. 264-265) the method of decending limits from alternative directions, i.e. left on first, then right on first. The subject while viewing the fixation point decided which circle came on first by responding "left-first," "right- first," or "same." Each run started with, say, the left preceding the right by 100 ms. For each succeeding trial, the interval was reduced by 5 ms steps until a simultaneity judgment was made. Following this, a new run was started with the apposite stimulus appearing first. For any interval setting, the subject could view the stimuli as many times as he wished before making a judgment. After four practice runs, two from each direction, each left-rightdescending;series was repeated ten times. The starting points were random so as to vary the length of each series. The average results for the two subjects were 11.5 ms and 9.25 ms respectively - the left coming on before the right to obtain a judgment of simultaneity. The between-subjects mean value of 10.38 ms was the SOA value, target preceding mask, to be used for the four subjects of Experi- ment II. The SAA value for the right hemisphere must be somewhat less 46 than 2x10.38 ms or 20.76 ms because the two stimuli are of unequal in- tensity. A reasonable estimate based on Efron's work would place it somewhere between 15 and 20.76 ms. METHOD Subjects The subjects were four paid volunteers from the same two under- graduate psychology classes from which the subjects in Experiment I were drawn. All were males, right-handed, as determined by the Crovits and Zener (1962) questionnaire, and naive to the purpose of the experiment. Apparatus Same as Experiment I. Stimulus materials Same as Experiment 1. Procedure The procedure was identical to that of Experiment I except for the following: all subjects obtained the mask-target brightness condition as for Group III of Experiment I, i.e. target-0.20 ft.l.; mask-2.50 ft.l.; onset of the target field preceded onset of the mask field by 10.38 ms for all four stimulus conditions. Consequently, for the "right hemisphere- mask" condition (target to right hemisphere - mask to left), SAA for the left (responding) hemisphere is estimated to be zero, and for the "left hemisphere—mask" condition (target to left hemisphere - mask to right), SAA for the left hemisphere is estimated to be between 15 and 20.76 ms.1 1As a limiting condition, it must be greater than 10.38 ms by the amount of time it takes for the high intensity stimulus information to cross the corpus callosum. 47 RESULTS AND DISCUSSION The results for this experiment (Group V) as well as those for Group III, Experiment I, are summarized in Table 5 and the "one between- two within subjects" analysis of variance in Table 6. The nested between- groups factor now represents the different SOA values.1 Table 5. Percentage of correct responses for left versus right hemis- phere and mask versus no-mask conditions for Groups III and V, percentage decrement produced by the mask, average across hemisphere conditions, difference in percentage decrement, and mean target exposure time at threshold. diff. in mean T exp. T to T to average 1 decr. time (ms) @ left H right H across H (RH-LN) threshold Group III no-mask 67.5 57.0 62.3 SOA-50 ms mask 47.0 34.8 40.9 8.6 8.3 H before T Z decr. 30.4 39.0 34.7 Group V no-mask 53.3 57.3 55.3 SOA-10.38 ms mask 48.8 41.8 45.3 18.6 5.0 T before N. 1 decr. 8.5 27.1 17.8 Note. H-mask; T-target; H-hemisphere; R-right; L-left; SOA-stimulus onset asynchrony; 1 decr.-percentage decrement produced by mask Considering first the results of Group V alone (anova not tabled), the difference in percent correct between hemispheres was not significant, the masking effect was significant (p<<0.025), and the hemispheres x mask conditions interaction was significant (p‘<0.01) in the direction predicted by the model. It is noteworthy that this is the only one of the five groups 1Strictly speaking, the design requirements of this analysis were not fully met, i.e. subjects were not randomly assigned to the nested groups. However, since the subjects were drawn from the same classes during the same term, it was not felt that there was any systematic factor that could have influenced the content of the two groups. A better pro- cedure, had it been practical, would have been to include four additional subjects under replicated Group III conditions. Table 6. 48 Analysis of variance summary for Experiment II (Groups III and V) Source Total Between subjects (8) Groups4(A) S/A Within subjects Hemispheres (B) A; SB/A Mask conditions (C) Ag SC/A BE égg SBC/A °p < 0.05 (two-tailed) gg 31 7 1 6 24 O‘HHO‘HHO‘HH SS 4014.719 288.469 13.781 274.688 3726.250 331.531 195.031 602.188 1968.781 258.781 206.188 81.281 42.781 39.688 13.781 45.781 331.531 195.031 100.365 1968.781 258.781 34.365 81.281 42.781 6.615 based on percentage correct scores 2 5 0.301 (ns) 3.303 (ns) 1.943 (ns) 57.291 7.569*** 7.5310 12.288 3.506** 6.468 2.543* *p<0.025 (one-tailed); 1“:p< 0.01 (one-tailed); mp<0.0005 (one-tailed) which produced a significant hemispheres X mask conditions interaction when analyzed separately (on the basis of four subjects). The most important comparison for the issues of Experiment II is made in column 4, Table 5. It is seen that the difference in percentage decrement right minus left hemisphere increases from 8.6 to 18.6 with the change in the SOA parameter. To reiterate, the above index is a summary number which expresses the magnitude and direction of the double interaction (hemispheres x mask conditions). The significance of the change in that index betweenpgroups is therefore expressed by the triple interaction term of Table 6 which is significant at the 0.025 level (one-tailed). However, the manner in which the change in the double interaction (i.e. the triple interaction) occurs is also important for present issues. 49 It was predicted that the primary change would be produced by a smaller masking effect resulting from the SAA of 15 - 20.76 ms or backward mask- ing condition at the left hemisphere for the target-to-left h. mask-to- right h. condition. The third row for each group (Table 5) shows that this was the case. Perhaps a more lucid picture of this change, and the nature of this one-tailed test of the triple interaction term can be given in the following manner: for each subject we can find the difference between his "no mask" and "mask" percent correct score for each hemisphere (the larger the difference, the larger the masking effect). The average of those dif- ferences, across subjects, appears in Table 7 (these values can also be obtained by direct subtraction of the means in Table 5). A decrease from Table 7. Average difference in percentage correct scores, between no-mask and mask conditions, for left and right hemisphere versus two "stimulus- onset-asynchrony" conditions T to T to 122281281841 Group III SOA-50 ms 20.5 22.3 N before T Group V SOA=10.38 ms 4.5 15.5 T before M Note. H-mask; T-target; H-hemisphere; SOA-stimulus onset asynchrony 22.31 to 15.51 is obtained for the "right hemisphere" condition, while a much larger decrease in masking effect, from 20.51 to 4.51, is observed for the "left hemisphere" condition. The triple interaction has now been reduced "in form" to a double interaction. The one-tailed test of the orthogonal comparison of these four means, based on the a priori 50 prediction that the largest change in masking occurs in the "left hemis- phere - mask" condition, is made in Table 8 (after Myers, 1967, p. 344). Note that the F and t values in Table 8 are identical to those for the triple interaction term in Table 6. Table 8. One-tailed orthogonal comparison test of the interaction between hemisphere and stimulus-onset-asynchrony conditions based on the average difference in percent correct scores (no-mask minus mask conditions) ____5°ur¢° ii £92. 115 I .t. p(A) x q(B) 1 85.563 85.563 6.468 2.543’” S x q(B)/A 6 79.375 13.229 *p40.025 Note. A-groups; B-hemispheres; s-subjects From‘Table 6 a significant groups x masking (AC) condition inter- action effect was also obtained. Columns 3 and 5 of Table 5 show that this interaction is due to a decrease in the overall masking effect from a 34.71 decrement to a 17.81 decrement with a change in the SOA parameter. Also, the average target exposure time required to maintain threshold decreased from 8.3 to 5.0 ms. Clearly, the targets were more detectable 'with target preceding mask by 10.38 ms. While most of this change is reflected by the predicted decrease in masking for the left hemisphere condition, a notable but smaller decrease from 22.31 to 15.51 (Table 7) was obtained for the right hemisphere condition as wall. This result could possibly be accounted for by the elimination of the forward mask- ing eggpgnent of the total masking effect when the mask-target overlap condition of Group III is changed to the SAA-O condition in Group V (for the right hemisphere condition). While a central forward masking effect 51 does exist (Turvey, 1973), it is probably too small to totally account for the above change. What would seem to be a more likely hypothesis is that a possible alteration in peripheral phenomena produced a small but £3221 decrease in masking effect for pggh hemisphere conditions. As mentioned previously, stray illumination within the eyeball is, for this paradigm, the most likely source of peripheral interference between mask and target. (See Bartley & Fry, 1934; Bartley, 1935; Boynton et al., 1954, for a discussion of stray illumination phenomena.) This possibility seems especially likely in the context of present experiments because of the semi-dark adapted state of the eyes. However, the effect, whatever its magnitude, is ex- pected to be equal with regard to the left or right hemisphere conditions described here.1 If we now compare the stimulus conditions of Group III with those of Group V, it can be seen that they differ greatly with respect to possible stray illumination effects. For Group III, the on— set of the 200 ms mask preceded that of the target by 50 ms so that for both eyes, stray light was physically present during target exposure. For Group V, however, target onset preceded mask onset by 10.38 ms and the averege tagger exposure time was 5.0 T 1,3 ms. Consequently, stray illu— mination from the mask could not have become physically present until approximately 5 ms after offset of the target. In effect, whatever con- sequence that could have resulted from stray light would have had to be due to "backward masking" by stray illumination with an interstimulus 1The only laterality difference known to exist in the visual system prior to the cerebral cortex is the difference in transmission efficiency of the nasal versus the temporal hemi-retina of each eye. This variable is, of course, held constant in the experiments reported here be- cause both eyes are stimulated at the same time during any one stimulus condition. A stimulus appearing in the left visual field, for example, stimulates the temppral hemi-retina of the right eye and the nasal hemi- retina of the left. 52 interval of 5 ms. In sum, it is prOposed that for Groups I to IV, a per- centage of the total masking variance may have been due to stray illumina- tion and that it made an equal contribution to both left and right hemis- phere conditions. Thus stimulus conditions for Group V were such as to considerably reduce the magnitude of a possible stray illumination effect while producing an increase in the central differential effect by control- ling the SAA intervals for the two hemispheres. The general goals of the first two experiments were to establish the tenability of the predicted left-right hemisphere differential mask- ing effect and to identify the stimulus parameters that might be relevant to increasing that effect. Both experiments provided statistical con- firmation of the effect although its magnitude must, admittedly, be con- sidered "small." Experiment I failed to show that mask-target brightness relationships influenced either the overall or the differential masking effect. The results of Experiment II established that the desired dif- ferential effect could be increased by manipulation of the SOA parameter and therefore the arrival-arrival intervals of target and mask informa- tion at the two hemispheres. If it is assumed that the effects observed in these experiments are "central" in origin, then the above findings are in total agreement with Turvey's (1973) extensive analysis of peripheral and central masking processes. Turvey found that masking via presumed peripheral mechanisms was very insensitive to the SOA parameter while for "central" masking, via the dichoptic paradigm, it was most crucial. Experiment II is in agreement with those findings. On the other hand, his finding that relative mask-target energy levels influenced only "peri- pheral" but not "central" masking is in agreement with the results of Experiment I . 53 While Experiments I and II were concerned with attempts to in- crease the magnitude of the phenomenon, the remaining experiments deal in a more direct manner with some theoretical issues about the neural mechanisms involved. Experiment III tests the assumption about inter- action between mirror-symmetric heteronymous areas of the visual half- fields, and Experiment IV assumptions about single cell prOperties of the neural substrate (and therefore the stimulus characteristics of the mask), potentially responsible for the central masking effect. Chapter 4: EXPERIMENT III In the general introduction to this study it was prOposed that the potential mechanism responsible for producing a masking effect bet- ‘ween the hemispheres was that which provided for continuity between the left and right half-fields of vision and which also may be responsible for stereoscOpic vision along the vertical meridian, i.e. the inter- hemispheric connections through the spleneum of the callosum from areas 17 to 18 and 19. One of the features of this system as described by BishOp and Henry (1971) and Blakemore (1970) is that the hemispheric (stereosc0pic) interaction depends on stimulation of correspondipg heteronymous retinal areas. The rationale for this requirement can easily be seen by the following example. For a human fixating on some object in his environment, objects which fall in front of or behind his point of fixation but to his left or right side project sighs; to his left or right hemisphere. Information from the disparate images of the two eyes have a common meeting point (i.e. can fire a common cell) in area 17 of the contralateral hemisphere. (See Bishop, et al., 1971, for a discussion of how binocular interaction is coded by these units.) On the other hand, objects which lie immediately in front of or behind the fixation point along the vertical meridian stimulate corresponding heteronymous areas of the two retinae and hence project to separate hemispheres (refer to Figure 4). A stereOpsis effect resulting from this particular kind of interocular disparity would, so the argument goes, have to result from a combining of coded information from both 54 55 hemispheres via the corpus callosum. The earliest possible site for such an interaction is, of course, the 18 side of the 17-18 border. As dis- cussed in the general introduction, the behavioral support for this con- tention was provided by Blakemore's (1970) study of a patient with a sagittal section of his Optic chiasm, and neurOphysiological evidence came from Berlucchi and Rizzolatti's (1968) demonstration that cells in the apprOpriate part of area 18 could only be binocularly driven from corresponding heteronymous areas (see Figure 5). If this mechanism is responsible, at least in part, for the cross- field visual masking phenomenon (which artificially simulates the above stimulus conditions), it ought to be possible to show a difference in masking effect produced by masks which are presented in correppondipg versus non-corresponding heteronymous areas in relation to the target stimulus. This was the goal of Experiment III. In order to define corresponding versus non-corresponding visual areas within the confines of the maximum stimulus field dimensions avail- able in the tachistosc0pe used (2.130 wide x 2.900 high per visual half- field after central occlusion for possible naso-temporal overlap), it was necessary to substantially reduce the area of the noise pattern used to about l/lOth the original size so that mask location could be varied. The scheme for varying the location is depicted in Figure 11 for the right field mask conditions. The geometric center of each mask is the same distance from fixation as is the target on the opposite side. The mask in position M-2 is considered the correppondipg heteronymous location while those in positions M-1 and M-3 are defined as falling in non-cor- responding heteronymous locations. The large mask (MeL), comprising the entire visual half-field on either side, as well as the no-mask condi- tions were retained for control purposes. 56 Figure 11. Corresponding and non-corresponding heteronymous mask loca- tions for the right-field (left hemisphere) mask conditions. Note. M-2 - corresponding; M-1 8 M-3 - non-corresponding The predictions are that the small M-Z mask will produce the same differential masking effect obtained in the earlier experiments but that the "overall" effect (across left-right conditions) will be smaller than that produced by the large mask due to a decrease in stray illumination or possibly due to a decrease in total retinotOpic projec- tion area represented in the visual cortex. Masks M-1 and M-3, while providing equivalent extrafoveal stimulation, should produce a much smaller or no effect compared to mask M-2 because they project to non- corresponding heteronymous areas. METHOD Subjects The subjects were four paid graduate students in non-perception areas of experimental psychology at Michigan State University. All were right-handed males with 20-20 or better corrected vision in each eye and naive to the purpose of the experiment. 57 Apparatus Same as for previous experiments. Stimulus materials The set of 100 target letters used in the previous experiments were retained unaltered as well as the no-mask and large mask stimuli. The small masks were prepared by photographing a cropped version of the original noise pattern so that the entire stimulus slide was "black" except for the mask appearing in its appropriate location. Six sets of the new mask stimuli were prepared, three with masks appearing in the M-1, M—2, or M—3 positions in the right visual field (see Figure 11), and three for the same positions in the left field. The dimensions of each small mask were 0.850 in width by 0.74° in height - a little more than twice the height and width dimensions of the target letters and approximately l/lOth the visible area of the large (M—L) masks. The geometric center of these masks fell on an imaginary circle with a radius of 1.70 measured from the fixation point (the same distance as the target centers from the fixation point). The angular elevation of the M-1 masks (left or right field) was 38.2° from the horizontal and depression of the M-3 masks from horizontal was of the same magnitude. The masks M-1 and M-3 together comprise the stimuli for the non-corresponding mask condition. Consequently, there were half as many Mel or M-3 slides (25 each for each field) as there were M-2 slides (50 for each field). As a result, the total number of stimuli for each condition (non-corresponding, corresponding, no mask, and large mask) was the same. 58 Procedure The general procedure was the same as that for the preceding experiments except as noted below. The mask-target brightness ratios and the controlled SAA conditions were the same as for Group V (that group having produced the best results) except that the apprOpriate SOA value was independently determined for each subject by the psycho- physical procedures described in Experiment II. The four within-subjects conditions defining the previous ex- periments have here been expanded to eight. They included the no-mask, large mask, non-corresponding small mask, and corresponding small mask conditions for both the left and right visual half-fields. The random- izing and counterbalancing procedures for presentation of target and mask stimuli were the same as described earlier. Any one of the (now 4) lOO-slide rototrays (for masking) contained either 12 or 13 stimuli for each of the eight conditions. Administration of Egg 100 trial sets gave the subject 25 trials for each of the eight conditions with each of the 25 target letters appearing once per condition. Excluding Day 1, which was used for the psychophysical determination of a subject's SOA value, a subject received 200 trials per day on each of four separate days (usually within one calendar week). Consequently, the four different sets of 100 masking stimuli were administered twice but in reverse order. In sum, each unfortunate subject received a total of 100 trials for each of the eight conditions. 59 RESULTS AND DISCUSSION The percentage of correct reaponses averaged across subjects for each of the eight conditions appear in the first two columns of Table 9. The averages across hemispheres for each of the masking conditions are tabulated in column 3. Results of the 2 x 4 within-subjects analysis of variance are summarized in Table 10. Table 9. Percentage of correct responses for left and right hemisphere conditions as a function of the no-mask, large mask, small corresponding, and small non-corresponding mask conditions of Experiment III (Group VI) T to T to average left H right H across H no mask 49.5 53.0 51.3 large mask 38.8 37.0 37.9 (M-L) small mask 48.3 50.3 49.3 (M-Z) small mask 47.5 54.8 51.1 (M-1-3) Note. T=target; H=hemisphere; M-2-corresponding; M-1 and M-3-non-corre- sponding Table 10. Analysis of variance summary based on percentage correct scores for Experiment III Source pg §§ MS E Total 31 1377.500 Hemispheres (A), 1 60.500 60.500 3.924 (ns) Mask conditions (B) 3 982.750 327.580 26.568 (p