p u II I? .91";th1 U!“ “I“’Q‘/~\T‘Qi\ 1N Tm. a “*‘F' CQMRALAI’EM‘ EYE ;;t"1b Thesis I'm “he Dogma of M 1%. 'fiICBIGAN STATE UNI JERSITY rharry Lee IIiwey 19268 THESIS b LIB .3 3‘1 3.’ MiChigf‘."> ‘1. «its University ABSTRACT FOVEAL CRITICAL FLICKER FREQUENCY AS A FUNCTION OF SYNCHKONOUS STIMULATION IN THE PERIPHERY OF THE CONTRALATERAL EYE By Terry Lee Hickey A number of studies have indicated that, under certain conditions, visual thresholds of one eye are affected by stimulation of the other eye. Of most relevance to the present investigation were those studies dealing with critical flicker frequency. The majority of these studies report a depressive effect when the stimulation in the two eyes is out-of-phase and a summative effect when the two sources of stimulation are in-phase. The purpose of the present study was to systematically investigate the effects of in-phase stimulation of various points along the periphery of one eye on the CFF thresholds measured at the fovea of the other eye. The method of serial exploration was used to present two synchronous, intermittent stimulus targets to three subjects. Each subject made CFF determinations at the fovea of the left eye for each of the following points of stimulation: fovea-left eye only; fovea-left eye, fovea- Terry Lee Hickey right eye; fovea-left eye, 10 degrees-right eye; fovea- left eye, 20 degrees-right eye; fovea-left eye, 30 degrees- right eye; fovea-left eye, 40 degrees-right eye; fovea- left eye, 50 degrees-right eye; fovea-left eye, 60 degrees- right eye; and fovea-left eye, 70 degrees-right eye. The results of the present study were in agreement with earlier investigations. The greatest summative effect was found when both eyes were stimulated foveally; the greatest depressive effect, at 10 degrees. Beyond 10 degrees, the slight summative effect found was attributed to differences in the latencies of the rods and cones. The results of the present study were interpreted as a further indication of the effects of timing of inputs on the visual system and, thus, afforded further evidence that alternation of response can occur in the visual system. Approved: gain—.43 “@1— Date: gays. .35? FOVEAL CRITICAL FLICKER FREQUENCY AS A FUNCTION OF SYNCHRONOUS STIMULATION IN THE PERIPHERY OF THE CONTRALATERAL EYE By Terry Lee Hickey A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of MASTER OF ARTS Department of Psychology 1968 $03 5*: r 9’" /¢?'3‘uo ACKNOWLEDGEMENTS I wish to thank Dr. 5. Howard Bartley whose continuous support and assistance in this endeavor have been invalu- able. . I wish to also thank Dr. Charles Hanley and Dr. Mark Rilling for reading and criticizing the manuscript. 1 wish to thank Mrs. Le Ann Slicer for her invaluable assistance in typing and preparing the manuscript. 1 am especially grateful to my wife for her encourage- ment and support and, most of all, for her willingness to give up so much that I might have the Opportunity to com- plete this endeavor. ii II III IV VI Introduction . Method . . . . Subjects Apparatus Procedure Results . . . Discussion . . Summary . . . Bibliography . TABLE OF CONTENTS iii 13 16 24 29 30 LIST OF FIGURES FIGURE Page 1. Layout of eXperimental apparatus employed . . . . 10 2. Diagrammatic drawing of stimulus presentation apparatus 0 O O C O O O O O O O O O O O O O O O O 12 3. Critical flicker frequency determinations for individual subjects and determinations averaged over subjects . . . . . . . . . . . . . . . . . . l7 4. Learning or practice effect over days averaged overBUbjGCtSeeeeeeeeeeeIeeeeeee20 5. CFF determinations for subject I. . . . . . . . . 21 6. OFF determinations for subject 2. . . . . . . . . 22 7. CFF determinations for subject 3. . . . . . . . . 23 iv LIST OF TABLES Table Page 1 Summary of Analysis of Variance . . . . . . 18 INTRODUCTION Since the early work of such people as Sherrington (1904), Abney and Watson (1916), Piper (1904), and McDougall (1901), investigators have been concerned with the possibility that the central convergence of the path- ways from the two retinae may exert some influence on binocular'perception. "Since homonymous halves of both retinae are connected with the same half of the cerebrum, it is usually assumed that cor- responding points upon the two retinae out- side of the fovea are represented in the visual projection areas of theocortex by single patterns. It is therefore generally accepted that the two retinae function inte- gratively, i.e., that the singleness of vision with binocular observation is made possible by the unity of their central con- nections.” (DeSilva and Bartley, 1930; 241). While comparing the brightness of an object seen both monocularly and binocularly, DeSilva and Bartley (1930) demonstrated that in order for the object seen monocularly to appear just as brilliant as the object seen binocularly, it must be 1.27 to 1.44 times as bright. Fry and Bartley (1933) presented further evidence that binocular vision results in an increase in brilliance over monocular vision. They accounted for the fact on the hypothesis "that path- ways from each pair of corresponding points in the two 2 retinae converge upon a common pathway in the brain and that summation takes place." (Fry and Bartley, 1933; 693). Several other investigators (Valerius; Piper; Aubert; and McDougall) have proposed and shown a facilitative, or summative, effect upon brightness resulting from the unity of the central projections. A summative, and inhibitive, effect has also been demonstrated with temporally manipulated stimuli. It has been shown that synchronous flashes delivered to correspond- ing points on the two retinae yield a higher critical flicker frequency (OFF) than is obtained with monocular regard or with binocular regard and alternate flashes. Sherrington (1904) studying binocular summation by the flicker method, concluded that: ' "As far as sensual (sic) effect goes, the light phases at the one eye practically do not, therefore, interfere or combine at all with the coincident dark phases at the other; and conversely. Nor do they, in the alter- nate left and right arrangement, add them- selves as a series of additional stimuli to the like series of stimuli applied at the other eye." (Sherrington, 1904; 37). The mechanical device employed by Sherrington lacked pre- cision and Ireland (1950) partially replicated it utilizing more modern electronic equipment. Ireland tested twenty- four subjects under four conditions of stimulation: 3 (a) monocular flicker, dominant eye; (b) monocular flicker, non-dominant eye; (c) binocular flicker, out-of-phase; and (d) binocular flicker, in-phase. The results clearly indicated the CFF for binocular in-phase stimulation was reliably higher than the monocular. In view of the dif- ferences between monocular and binocular stimulation, Ireland postulated some interaction between the two eyes which appeared possible only by way of some central mech- anism of the brain. Thomas (1954) was interested in determining the effect of interocular differences in intensity on CFF values re- ported with binocular regard. His measures were made with both in- and out-of-phase flashes in the two eyes with the stimuli located centrally and 10 degrees peripher- ally from the fovea. The results indicated a general subtractive effect of a less intense stimulus in one eye on the CFF of a brighter stimulus on the corresponding area of the other eye. He found the subtractive effect to be even greater when both light sources were placed 10 degrees peripherally from the fovea. Thomas (1955) reported comparing CFF measured under three conditions: uniocular regard; binocular regard, flashes in-phase; and binocular regard, flashes out-of-phase. 4 He postulated that if the impulses arriving from each eye were additive, then the synchronous stimulation should yield a visual effect identical to doubling the flash luminance of‘a stimulus projected on a given region of one retina. His results showed that over the range of luminance studied, binocular CFF with synchronous flashes was significantly higher than the binocular CFF with out- of-phase flashes or the uniocular CFF. The difference was small "being only about one half to one third as much as would result from doubling the flash luminance of a stimulus viewed with one eye." (Thomas, 1955; 52-53). The magnitude of summation was independent of flash lumin- ance and flash frequency. In the first of a series of studies dealing with binocular CFF, Baker (1952) replicated Sherrington's (1904) study. The ranges between the synchronous and alternate flicker rates found in the Baker and Sherrington studies differed in magnitude with Baker's evidence definitely indicating the presence of some alliance and antagonism between flicker processes initiated at corresponding retinal points. Baker's results led him to the following conclu- sion: "Binocular fusion involves, in part at least, some central process which combines and inte- 5 grates the neural processes arising from stimulation of corresponding retinal areas, so that the resultant sensation differs from that arising from either eye alone." (Baker, 1952; 10). In a sequel to the first paper, Baker (1952a) described six additional eXperiments concerned with monocular and binocular CFF. The concensus of the six studies briefly described by Baker was that binocular fusion involved some central process which combined and integrated the neural processes arising from stimulation of correspond- ing, or non-corresponding, retinal areas of similar size. The resultant binocular percept is different from that arising from stimulation of either eye alone. Baker (1952b) demonstrated a central connection by utilizing "progressive“ and "instantaneous" stimulus occlusion. In the first of the two eXperiments reported, Baker demonstrated that higher CFF values could be obtained using an "instantaneous," rather than a "progressive," source; "...the receptor and Optic nerve discharge frequencies for the 'instantaneous' and 'progressive' methods of stimulus occlusion are dis- similar at similar stimulus intermittence rates." (Baker, 1952; 126). In the second experiment, Baker used an "instantaneously" occluded source of stimulation for one eye and a "progressively" occluded source of stimulation 6 for the other. His hypothesis was that the OFF value. obtained under this condition should fall somewhere between that value obtained with ”instantaneous" or "pro- gressive" occlusion alone. The eXperimental results con- firmed the hypothesis that central factors are active in "binocularly fusing neural impulses initiated by stimula- tion of the respective foveas." (Baker, 1952; 128). Perrin (1954) found that for a 2 degree visual field and dark surround, as the field luminance was varied the increase in binocular in-phase CFF was preportional to the mean critical frequency for the two eyes. Perrin was also concerned with the effect of field size on binocular summation. As would be expected, he found that increasing the stimulated area - at constant luminance - increased the critical frequency. He also noted that as the area of the stimulus target increased from 0 to 12 degrees, binocular summation increased. "Specifically, the summa- tion for corresponding areas was about 8 percent, while for noncorresponding areas it was of the order of 0.4 percent." (Perrin, 1954; 69). A study by Wolf and Zigler (1958) determined monocular and binocular thresholds for various points along the peripheral retina. Their measures extended from 30 degrees METHOD Subjects Three students (1 male and 2 females), including the author, served as subjects. All subjects had considerable eXperience in making flicker discriminations. Apparatus The apparatus shown in Figure 1 consisted of a metal rod curved into a half circle with a radius of 16 inches; a septum which divided the left and right visual fields; and a chin rest placed just to the left of the center of the arc. This placement of the chin rest positioned the entrance pupil of the right eye exactly at the center of the arc. Placed on the arc were two clamps each supporting a rod which held one stimulus target. Each stimulus target was a circular aperture 1/2 inch in diameter which pro- jected a visual angle of 1° 47' at 16 inches. A piece of Opal glass served as the target surface. The two stimulus sources were equated for brightness by means of a Macbeth photometer. The luminance was 1.5311 candles/ftz. The same level of luminance was maintained throughout the eXperiment. A small red "seed" lamp, positioned in the 9 7 to the right of fixation on a horizontal meridian and on a parallel line 1.5 degrees below the meridian. They found that in the binocular curves the thresholds were high at the center, dropped to low levels between 5 and 20 degrees, and gradually rose farther in the periphery. Thresholds were maximal at approximately 17.5 degrees. The monocular curves were asymmetrical due to the blind spots in the left and right fields. They also found that the binocular thresholds were slightly lower than the monocular thresholds, indicating slight binocular summation.. The Wolf and Zigler study was not, however, concerned with either CFF or inter- mittent stimulation. With the exception of Thomas (1954), Perrin (1954), and Wolf and Zigler (1958), the studies previously cited were concerned only with foveal stimulation. After a more than cursory search of the literature, the present author has concluded that very little, if any, work has been done investigating binocular summation by systematically stimulating points along the peripheral retina. PURPOSE OF THE STUDY It is, therefore, the purpose of the present study to partially duplicate and extend the work of previous inves- tigators dealing with binocular summation. Using inter- 8 mittent light to stimulate the fovea of one eye and various points toward the periphery of the other eye it should be possible to determine the effect of peripheral in-phase stimulation on foveal CFF with binocular regard. 10 4. Fig. 1. Layout of experimental apparatus employed. ll center Of the left stimulus target, served as a fixation point. The stimulus target for the left eye remained station- ary throughout the investigation while the stimulus target Ifor the right eye was moved along the rod. The right hand portion of the rod was calibrated in units of 10 de- grees. 1 Figure 2 represents a diagrammatic drawing of the stimulus presentation apparatus employed. The intermit- tent stimulus lights were produced by two Sylvania Rll3lC glow-modulator tubes activated by square-wave inputs. Two separate Model 8-4 variable-frequency squaregwave stimu- lators made by Grass Instrument Company furnished the square-wave oscillations for the two glowamodulator tubes. The rate of intermittency of both glow-modulator tubes was always equal and in-phase. The Operation of the glow- modulator tubes was completely silent at all times. Adjust- ment of the frequency of both square-wave stimulators was always made by the eXperimenter. Flicker frequency was monitored by a Beckman digital counter. The input channels were controlled by a series of Hunter Model lOO-C decade relay timers. The timers acti- vated both glow-modulator tubes simultaneously for three Fig. 12 2. [1'3 0—0 H Diagrammatic drawing of stimulus presentation apparatus. buzzer six volt transformer timers glow modulator glow modulator driver driver Grass 3-4 Grass 8-4 HID'TJFZUOtrIP 13 seconds, after which both tubes extinguished for seven seconds. This presentation procedure was continued throughout the experiment. The timers also controlled a buzzer which was activated 1 second prior to the activation of the two glowbmodulators. This warning buzzer signaled the subject that he should fixate on the red ”seed" lamp and prepare for the succeeding stimulus presentation. Procedure Although all subjects had previous experience in making flicker discriminations, all subjects were given several trials prior to actual experhmentation in order to stabilize their criterion for CFF. During these trials only the left eye was stimulated. The data from these trials were not included in the final analysis. Prior to each eXperimental session, subjects were dark.adapted for One-half hour. Following dark adaptation each subject was instructed to place his head as close to the apparatus as possible with his chin in the chin rest and to fixate on the red light at the sound Of the tone. The red "seed" lamp, although not as bright for the right eye due to the polaroid paper used to block out the left stimulus light, could be seen by both eyes. Following the ‘warning signal both stimulus lights would come on for three 14 seconds. After the stimulus lights were extinguished, the subjects were required to report their judgments as to whether the stimulus target had been steady or flickering. This judgment was reported verbally. During the seven seconds the stimulus lights were off, the eXperimenter recorded the subject's response and changed the rate setting for the next trial. Between series of trials the subject was given a brief rest period while the eXperimenter changed the stimulus setting along the rod. This procedure was continued throughout the investigation. The method of serial exploration, with five ascending and five descending trials, was used for the presentation of the various rates Of intermittencies at each stimulation point. The order of presentation Of the various stimulation points was randomly determined during each eXperimental session. The following conditions were used for subjects: (1) fovea-left eye only; (2) fovea-left eye, fovea-right eye; (3) fovea-left eye, 10 degrees-right eye; (4) fovea- left eye, 20 degrees-right eye; (5) fovea-left eye, 30 degrees-right eye; (6) fovea-left eye, 40 degrees-right eye; (7) fovea-left eye, 50 degrees-right eye; (8) fovea- ‘ left eye, 60 degrees-right eye; and (9) fovea-left eye, 70 degrees-right eye. 15 Each experimental session lasted approximately two hours including the one-half dark adaptation period. A total of five sessions on consecutive days was conducted for each_subject. RESULTS The flicker contours plotted in cycles per second are shown in Fig. 3. Open squares represent thresholds for subject 1; solid squares, subject 2; Open circles, subject 3. Each point for the curves is the mean of 50 CFF deter- minations. The points denoted by solid circles represent thresholds averaged over the three subjects and represent 150 CFF determinations each. It is apparent that CFF thresholds measured with foveal stimulation Of both eyes are consistently a little higher than the thresholds deter- mined with either peripheral stimulation Of right eye or with stimulation of the left eye only: There is also evi- dent a general depressive effect for all subjects when the right eye is stimulated between 10 degrees and 20 degrees in the periphery. Beyond 20 degrees the CFF thresholds remain fairly constant at approximately the threshold level for the left,eye alone. In the curves of two subjects there is a drOp in the thresholds beyond 60 degrees; however, this tendency is reversed in the curve of the third subject. The results of a five-way analysis of variance com- puted are presented in Table I. All of the main effects ‘with the exception of "trials" were significant at the .01 level. One three-way interaction, days 5 trials 5 pre- 16 CFF 40 39 38 37 36 35 34 33 32 31 30 29 28 17 Subject 1: a Subject 2: I Subject 3: ° ‘ Means : e n j a .4 r I I I I I I I I BOTH LEFT 10 20 3O 4O 50 60 70 EYES EYES DEGREES Fig. 3. Critical flicker frequency determin- ations for individual subjects and determinations averaged over subjects. Table I. 14. 15. Source Subjects Point Day Trial Presentation Point X Day Point X Trial Point X Presentation Day X Trial Day X Presentation Trial X Presentation Point X Day X Trial Point X Day X Pre- sentation Day X Trial X Pre- sentation Point X Day X Trial X Presentation * Significant at .01 ** Significant at .05 18 SS 9824.4 309.9 1196.2 3.2 109.9 198.3 8.7 1.9 3.0 13.3 3.7 36.7 7.7 4.7 17.3 Level Level Summary Of Analysis of Variance '0 .p a~ co to m 32 32 16 128 32 16 128 M§_ 4912.2 38.7 299.0 .8 109.9 § 354.9* 4.6* 9.3* .8 422.6* l9 sentations, was significant at the .05 level. A test of individual comparisons was not made due to the small num- ber of subjects. Figure 4 shows the "learning" or "practice".effect over days. Each point is the mean of 270 CFF determinations. Note the curve becomes asymptotic at about day four. The significant "days" main effect was attributed to the rise in CFF thresholds over days. This shift in the CFF curve agrees with data presented by Ireland (1950). Figures 5, 6, and 7 show CFF curves for subjects 1, 2, and 3, respectively. Open circles represent an average of the CFF determinations during the first two days of the experiment; solid circles represent averages for the last two days. Note that although the curves for the last two days are typically displaced toward the higher frequencies, the overall shapes of the curves are generally similar. These curves tend to indicate that rather than differences in the general shape of the curve occurring over days, the subjects, as a whole, become more sensitive, as evi- denced by the higher CFF determinations. CFF 41 40 39 38 37 36 35 34 33 32 31 ”30 29 20 DAY Fig. 4. Learning or practice effect over days averaged over subjects. CFF 4O 39 38 37 36 35 34 33 32 31 30 29 . 28 21 Mean-days l and 2: O Mean-days 4 and S: O . ° '-v, .1 e 'IIIIIjII BOTH LEFT 10 20 30 4O 50 6O 70 EYES.EYES ‘ DEGREES. Fig. 5. CFF determinations for subject 1. 40 39 38 37 36 - 35 34 33 32 31 30 29 28 22 Mean-days 1 and 2:0 Mean-days 4 and 5:. I l I I I I 1 I I BOTH LEFT 10 20 3O 4O 50 60 7O EYES EYES DEGREES Fig. 6. CFF determinations for subject 2. 23 Mean-days 1 and 2:0 Mean-days 2 and 4:. 40 - 39 « 38 q 37 d 36. 35. 334 314 30- 29. 28* BOTH LEFT 1b 20 so '40 330 ‘60 '70 EYES EYES DEGREES Fig. 7. CFF determinations for subject 3. DISCUSSION The finding that synchronous stimulation Of the foveas of both eyes yields a significantly elevated CFF agrees with the results Obtained by several authors (Ireland, 1950; Thomas, 1955a, 1955b; and Baker, 1952a, 1952b, 1952c). These results, along with the results Of the present study, indicate a certain degree of binocular summation. Thomas (1955) reported average differences of 1.37 and 0.89 flashes per second between binocular and uniocular CFF thresholds. In the present study, the value Obtained for the same comparison - 1.78 flashes per second - closely resembles those reported by Thomas. The depressive effect found at 10 degrees in the pre- sent study at first appears to coincide with the depressive effect reported by Thomas (1954). However, in comparing the results from the two studies, certain procedural dif- ferences must be kept in mind. Thomas was concerned with CFF thresholds as a function of stimulus intensity and the depressive effect he reported was due to differences in intensity of the stimulus targets. Also in the Thomas study, both stimulus patches were displaced 10 degrees, whereas, in the present study only the right target was displaced and the CFF determinations were always made at 24 25 the fovea Of the left eye. The fact, however, that Thomas did find a greater depressive effect at 10 degrees than at the fovea appears to coincide with the results of the present study. Ross (1936) determined CFF thresholds at the fovea and at 5, 10, 20, 30, and 45 degrees in the retinal periphery using a single stimulus target. He found that fusion fre- quency values decreased as the stimulus target was moved toward the periphery. His curves showed that at 45 degrees the CFF values were about 50% as large as they had been at the fovea, with the greatest drop occurring between 0 and 10 degrees. Beyond 10 degrees the curve continued to drOp but more gradually. In contrast to the curves pre- sented for binocular thresholds in the present study, the curves presented by Ross indicate no increase in CFF thresholds beyond 10 degrees. The increase in threshold values beyond 10 degrees found in the present study would, thus, appear to indicate some degree of binocular summa- tion occurring during peripheral stimulation. Since the proportion Of rods to cones increases toward the periphery of the eye, the points Of stimulation beyond 10 degrees may be viewed as stimulation to two different populations of receptors. Although the stimulus inputs to 26 the two populations of receptors were synchronous in the present study, it is unlikely that their inputs to any central processes mediated by the receptors would be syn- chronous. Bartley (1942), working with single flashes of light, found that when the area of the stimulus target was increased, the subject reported seeing two flashes. Bartley deduced from his finding that the two flashes were obtained ”when both the rod and the cone pOpulations are activated, and that the one pOpulation reacts more quickly than the other population." (Bartley, 1958; 127). Gouras (1966) presented electroretinogram records showing rod and cone threshold response latencies in the dark adapted retina of the Rhesus monkey. The records indicated a con- siderable (100 msec.) time delay between the cone and rod responses with the cone response being the faster. There- fore, if there is a difference in the latencies of the rods and cones, as there well appears to be, then the points beyond 10 degrees on the curve in Fig. 3 may actually repre- sent CFF determinations under cortical out-of-phase stimu- lation rather than under synchronous stimulation as was originally assumed. Such an interpretation would seem plausible in light of the following data. Perrin (1953) found that with a 2 degree field, subjects' monocular CFF 27 threshold values were approximately midway between the binocular thresholds with alternate flashes and those with synchronous flashes. Thomas (1954) also reported that binocular thresholds with out-Of-phase flashes tended to be the same as the uniocular thresholds, and in some cases lower. The results of the present study indicate that the timing of stimulus inputs, and thus the receptor outputs, has a considerable influence on processes within the visual system. Baker (1952c) interpreted his results as offering additional evidence that alternation of response can occur in the visual system. Baker states that: "...the central factors active in effecting binocular vision can perform their task when non-corresponding retinal points are stimu- lated, for, within an fovea, the receptors (correSponding points alternate in respond- ing to intermittent stimulation." (Baker, 1952c; 129). An interpretation of the results of the present study along the lines of the interpretation given by Baker appears reasonable. I The present study is taken to be another example of the fact that certain gross temporal features of input to the cortex exercise a demonstrable effect on sensory out- come a 28 It is, therefore, another example of the temporal factor that has received so much study in this laboratory. 29 SUMMARY The method of serial exploration was used to deter- mine CFF thresholds for three subjects. With stimulation being supplied synchronously by two Sylvania glow-modulator tubes, the subjects made CFF determinations at the fovea of the left eye under the following nine conditions: fovea-left eye; fovea-left eye, fovea-right eye; fovea- left eye, 10 degrees-right eye; fovea-left eye, 20 de- grees-right eye; fovea-left eye, 30 degrees-right eye; fovea-left eye, 40 degrees-right eye; fovea-left eye, 50 degrees-right eye; fovea-left eye, 60 degrees-right eye; and fovea-left eye, 70 degrees-right eye. A five-way analysis of variance was computed and the results presented in tables and graphs. The basic find- ings were in accord with previous studies dealing with binocular summation and inhibition. BIBLIOGRAPHY Abney, W. and Watson, W. Phil, 15323. A, CCXVI, 91, 1916. In Parsons, J. H. Ag Introduction £9,5hg I Sggnggf Colour Vision. Cambridge: Univ. Press,‘ 1924. Baker, C. H. The dependence of binocular fusion on timing of peripheral stimuli and on central process, 1. Symmetrical flicker. ggggg. g, ggzgh., 1952a, 6: 1-10. Baker, C. H. The dependence of binocular fusion on timing of peripherai stimuli and on central process, 1. Symmetrical flicker (con't.). 93229. g. 23152,, 1952b, 6: 84-91. Baker, C. H. The dependence of binocular fusion on timing of peripheral stimuli and on central process, 2. Asymmetrical flicker. ggggg. g. nggh., 1952c, 6: 123-130. Bartley, S. H. The features Of the Optic nerve discharge underlying recurrent vision. g. 352, figygh,, 1942, 30: 125-134. Bartley, S. H. Principles gngerception. New York: Harper & Row, Inc., 1958. 30 31 DeSilva, H. R. and Bartley, S. H. Summation and subtrac- tion of brightness in binocular perception. Bgig. g. 23152,, 1930, 24: 538-541. Fry, G. A. and Bartley. The brilliance of an object seen binocul'arly. 5335. ,1. Ophthalmol., 1933, 16: 687-693. Gouras, P. Rod and cone independence in the electroretino- gram of the dark-adapted monkey's perifovea. g. Physiol., 1966, 187: 455-464. Ireland, F. H. A comparison of critical flicker frequencies under conditions Of monocular and binocular stimula- tion. g. 252, Egygh., 1950, 40: 282-286. McDougall, W. ngig, XXIV, 577, 1901. In Parsons, J. H. An Introduction £2,523 Sgggx_gf Colour Vision. Cam- bridge: Univ. Press, 1924. Parsons, J. H. §3_Introduction £2,5hg_§£ggy_gf Colour Vision. Cambridge: Univ. Press, 1924. Perrin, F. H. A study of binocular flicker. J. 925. S23. 5325., 1954, 44: 60-69. Piper, H. ggggh. f. Psycho . g, Physiol. g. Sinnesorg., XXXII, 161, 1904. In Parsons, J. H. 5g Introduction £2 Egg Sgggy 2; Colour Vision. Cambridge: Univ. . Press, 1924. Ross, R. T. The fusion frequency in different areas of the visual field: 11. The regional gradient of fusion 32 frequency. g. 539, 25122., 1936, 15: 161-170. Sherrington, C. S. On binocular flicker and the correla- tion of activity of ‘corresponding' retinal points. 2511, g. 23152., 1904, 1: 26-59. Thomas, G. J. The effect on critical flicker frequency of interocular differences in intensity and in phase relations of flashes of light. 5335. g. 2g1gh., 1954, 67: 632-646. Thomas, G. J. A comparison of uniocular and binocular critical flicker frequencies: simultaneous and alter- nate flashes. A932. g. 22152., 1955, 68: 37-53. Thomas, G. J. Effect Of contours on binocular CFF obtained with synchronous and alternate flashes. 5322. g. 22132., 1956, 69: 369-377. Wolf, E. and Zigler, M. J. Uniocular and binocular sco- topic responsiveness of the peripheral retina. g. 925. 223, Amgg., 1959, 49: 394-398. "'TITI'ITIQITILIMEIITIIE[Iiifljljl'l‘flflllfifliflfgfin‘m'Es