Chapter 7 Temporal Factors in Vision Spatial vision is not possible unless the retinal image changes with time Spatial vision is not possible unless the retinal image changes over time The Troxler effect: the fading of a large stimulus with blurred edges presented in the peripheral visual field X Five Parts to this Chapter Temporal Acuity (critical flicker frequency [CFF]) The Temporal Contrast Sensitivity Function Temporal Summation Masking Motion Detection (Real and Apparent) Part One Temporal Acuity: The critical flicker frequency (CFF) The critical flicker frequency (CFF) is a measure of the minimum temporal interval that can be resolved by the visual system. CFF is analogous to grating acuity as a measure of spatial resolution acuity Measure CFF using an episcotister (a rotating sectored disk used to produce squarewave flickering stimuli) the period is the length of time for one complete cycle of light and dark, and the flicker rate, or flicker frequency is the number of cycles per second (Hz) Duty cycle – the ratio of the time a temporal square-wave pattern is at Lmax to the time it is at Lmin How bright does a fused flickering light appear? The time-averaged luminance of a flickering light determines its brightness at flicker rates above the CFF (Talbot-Plateau Law) Talbot brightness = Lmin + ([Lmax – Lmin] x f) where f is the fraction of time that Lmax is present during the total period To convert duty cycle to f, divide the first number by the sum of the two numbers: 1:1 means f=0.5 Eq. 7.1 If a square-wave flickering light has a duty cycle of 4:1, what is f? 0% 0% 1. 0% .4 0% .2 0% .8 0.1 0.2 0.4 0.8 1.0 .1 1. 2. 3. 4. 5. To convert duty cycle to f, divide the first number by the sum of the two numbers: 4:1 would be 4/(4+1) so f=0.8 How bright does a flickering light appear? At flicker rates slightly below the CFF, brightness is enhanced beyond the mean luminance of the flicker (the Brücke-Bartley phenomenon) This is related to the Broca-Sulzer effect described later in the chapter The neural basis of the CFF is the modulation of firing rates of retinal neurons (ganglion cells) A Neural response Stimulus luminance B Neural response Stimulus luminance C Neural response Stimulus luminance Time Cone flicker response (pig). Contrast 0.49; mean light level 48,300 photon/ square micron Courtesy of Dr. Tim Kraft Rat ganglion cell responses showing CFF In order to see a light as flickering 1. The flicker rate must be above the CFF 2. The Troxler effect must occur 3. Retinal neurons must be able to respond with gaps in their firing pattern 4. All of the other answers are correct A e. . ra ns ot he ll o ft he ne w . m us t. ur on s ef fe c et in al R e Th Th e fli ck Tr ox er le r ra te m us t. . tm u. . 0% 0% 0% 0% How does this measure of temporal acuity (CFF) change under different conditions (changes in the stimulus dimensions listed in Chapter 1)? First: stimulus luminance (intensity) Important Stimulus Dimensions intensity wavelength size exposure duration frequency shape relative locations of elements of the stimulus cognitive meaning In addition,(NOT stimulus Dimensions!) location on the subject’s retina light adaptation of the subject’s visual system CFF is directly proportional to the log of stimulus luminance (Ferry -Porter Law) CFF = k log L + b where k is the slope of the function , b is a constant, and L is the luminance of the flickering stimulus Critical Flicker Frequency (Hz) 50 642 nm at the fovea 40 30 Note: if the luminance of the stimulus increases by one log unit, so does the retinal illuminance 20 10 0 -1 0 1 2 3 Log Retinal IIluminance (Td) 4 Critical Flicker Frequency (Hz) 50 642 nm at the fovea 40 30 3 1 20 2 10 demo 0 -1 0 1 2 3 4 Log Retinal IIluminance (Td) 1) Find CFF 2) Raise intensity (luminance) by 1 log unit. The more intense stimulus is below CFF (flicker is seen). 3) Have to increase the flicker rate to again find CFF. The Ferry-Porter law holds at all eccentricities. The slope is steeper in the periphery. At high luminance, CFF is higher in the periphery than at the fovea. Critical Flicker Frequency (Hz) 100 80 60 o 0 o 3 40 o 10 o 35 o 65 o 85 20 0 -4 -3 -2 -1 0 1 2 o 3 4 o Log Retinal Illuminance for 10 -85 (td) -1 0 1 2 3 4 5 o 6 o Log Retinal Illuminance for 0 and 3 (td) 7 The CFF is highest in the midperipheral retina at high luminance, but nearly constant across the retina at low luminance. Critical Flicker Frequency (Hz) Retinal Illuminance (td) 2500 250 100 25 2.5 0.25 80 This is why you can see flicker on some PC monitors if you look slightly to the side 60 40 20 0 0 20 40 60 Retinal Eccentricity (deg) 80 100 CFF increases 1. In direct proportion to the log of the stimulus luminance 2. In the periphery at all luminance levels 3. In response to the Brücke-Bartley phenomenon 4. None of the above th e N on e of to ns e po re s In ab ov B th e at he ry rip pe th e In r.. . l.. . al to ... or tio n pr op di re ct In e 0% 0% 0% 0% Second: area (size) CFF is directly proportional to the logarithm of the area of the flickering stimulus (the Granit-Harper Law) CFF = k logA + b Where k and b are constants and A is the area of the flickering stimulus Demo, since I haven’t found a good figure showing this relationship Demo – Granit-Harper 1) Find CFF 2) Increase stimulus area by 1 log unit. The more intense stimulus is below CFF (flicker is seen). 3) Have to increase the flicker rate to again find CFF. Chapter 7 – Temporal Factors in Vision Main points so far: 1) CFF is a measure of temporal acuity – analogous to VA (how small a temporal interval can you detect – in time)? 2) CFF increases linearly with log stimulus luminance (Ferry-Porter Law) 3) CFF increases linearly with log stimulus area (Granit-Harper) You will not be responsible for the material starting on page 188, “flicker sensitivity increases….” and including all of page 189 and 190 (Figs. 7.7 and 7.8). You will be responsible for material starting again on page 191, “Temporal Contrast Sensitivity” Five Parts to this Chapter Temporal Acuity (critical flicker frequency [CFF]) The Temporal Contrast Sensitivity Function Temporal Summation Masking Motion Detection (Real and Apparent) Contrast, modulation and amplitude The contrast of a temporal sine wave is defined the same way as the contrast of a spatial sine wave grating: Contrast = (Lmax - Lmin)/( Lmax + Lmin) In Figure 7-1, Lmax is 300, Lmin is 100, so contrast = (200)/(400) = 0.5 Another term, modulation (abbreviated asm),is sometimes used for sine-wave flicker, and may be used interchangeably with contrast. As illustrated in Figure 7-1, Lmaxis the maximum luminance of the flicker, and L minis the minimum luminance. Lmax and L min are symmetrically arranged around the mean or average luminanc defined as: Mean Luminance = Lm = ( Lmax + Lmin)/2 Hence, contrast or modulation can also be expressed as: Contrast = m = (Lmax - L m )/ L m In addition, Lmax - L m is also called the amplitude of the wave, and, therefore, Contrast = modulation = amplitude /Lm Referring again to the sine wave at the bottom of Figure 7-1, the mean luminance is 200 units, the amplitude is 100, and the contrast (modulation) therefore is 0.5. As was the case for spatial sine-wave gratings, contrast sensitivity is defined as the inverse of the threshold contrast. Temporal CSFs have several features in common with spatial CSFs: band pass shape, cutoff high frequency indicating the acuity limit, and a low frequency rolloff. Threshold Contrast Contrast Sensitivity 200 0.005 100 0.01 50 0.02 20 0.05 10 0.1 5 0.2 2 0.5 1 1 2 5 10 20 Frequency (Hz) This is like Figure 6.9 in the spatial domain 50 100 Retinal Illuminance (Td) 9300 850 77 7.1 0.65 0.06 Temporal CSF Demo http://psy.ucsd.edu/~sanstis/TMTF.html Change in the temporal CSF with luminance: As luminance decreases, Threshold Contrast Contrast Sensitivity 200 0.005 100 0.01 50 0.02 20 0.05 10 0.1 5 0.2 2 0.5 1 1 2 5 10 20 Frequency (Hz) the peak contrast sensitivity becomes lower the cutoff high temporal frequency decreases (Ferry-Porter law) peak contrast sensitivity occurs at lower temporal frequency the low temporal frequency rolloff disappears 50 100 Retinal Illuminance (Td) 9300 850 77 7.1 0.65 0.06 The temporal contrast sensitivity function B a Is ec m ea ba n m or e om es su re of t em nt ra s co k pe a a as d. .. po r.. ... ta . be t.. nd ar y H 4. 0% 0% 0% 0% bo u 3. th e 2. Is the boundary between contrasts you can see and ones you cannot see Has a peak contrast at around 1 Hz at high mean luminance Is a measure of temporal acuity Becomes more bandpass as the mean luminance is decreased Is 1. The center-surround interactions of retinal neurons may account for a low frequency roll-off in temporal CSF of individual neurons Actually, there is a mid-temporal frequency enhancement of sensitivity The delayed arrival of the surround signal, relative to the center signal can cause the surround to add with the center at some temporal frequencies The delayed arrival of the surround signal, relative to the center signal can cause the surround to add with the center at mid-range temporal frequencies Temporal CSFs have several features in common with spatial CSFs: band pass shape, cutoff high frequency indicating the acuity limit, and a low frequency rolloff. Threshold Contrast Contrast Sensitivity 200 0.005 100 0.01 50 0.02 20 0.05 10 0.1 5 0.2 2 0.5 1 1 2 5 10 20 Frequency (Hz) This is like Figure 6.9 in the spatial domain 50 100 Retinal Illuminance (Td) 9300 850 77 7.1 0.65 0.06 The temporal CSF is a useful measure for diagnosing retinal disorders 1. Artificially increased IOP produces reduced temporal CSF (but no effect on CFF) 2. Temporal CSF is reduced with glaucoma and ocular hypertension • Glaucoma - Frequency-doubling perimeter measures contrast threshold for 0.25 c/deg grating flickering at 25 Hz (mediated by MY [nonlinear magno] cells?) 3. Eyes at risk for exudative (wet) AMD show reduced sensitivity at 5 - 40 Hz (5 Hz & 10 Hz alone discriminate from healthy eyes) Importance? Early diagnosis can lead to earlier treatments The low temporal frequency rolloff of the temporal CSF Is re l cr at ed ea t to f.. . to e th e M ac h cu b. .. ... pr om m or e el ps ec B H a om es “m id -te m po r .. 0% 0% 0% 0% lly 3. 4. re a 2. Is really a “mid-temporal frequency enhancement produced by the longer latency of the receptive field surround Becomes more prominent at low mean luminance levels Helps create Mach bands Is related to the cutoff high temporal frequency Is 1. Five Parts to this Chapter Temporal Acuity (critical flicker frequency [CFF]) The Temporal Contrast Sensitivity Function Temporal Summation (Bloch’s Law & Broca-Sulzer) Masking Motion Detection (Real and Apparent) Log Threshold Luminance (quanta/s/deg2) Fig. 2.5 Log Background Intensity 7.83 5.94 4.96 3.65 No Background 9 8 7 6 5 4 Stimulus area = 0.011 deg2 0.001 0.01 0.1 1 Flash Duration (s) 10 100 Bloch’s Law holds for durations shorter than the critical duration Lxt=C Eq. 7.7 where L is the threshold luminance of the flash, t is its duration, and C is a constant Remember: luminance (L) is directly proportional to the number of quanta (Q) in a flash and inversely proportional to the duration (t) and area (A) of the flash, or L=Q/txA quanta x duration C duration x area Eq. 2.6 There is a constant # of quanta in a threshold flash as L decreases Part A – threshold measures Temporal Summation and Bloch's Law When a brief flash is used to determine the threshold intensity, the visual system does not distinguish the “temporal shape” of the flash if the flash duration is less than the “critical duration” A B Number of Quanta Critical Duration Time Critical Duration Time Two ways to show Bloch’s Law: L x t = C Log Threshold Luminance Bloch’s Law holds Log Threshold Luminance x Time Bloch’s Law holds 1 10 100 1 10 100 Flash Duration (msec) “Holds” means that Bloch’s Law accounts for the threshold values Bloch's law is a consequence of the temporal filtering properties of vision. But I will not hold you responsible for this section Bottom of 198 & top of 199 Bloch's law is a consequence of the temporal filtering properties of vision. Fourier Synthesis: can construct complex waveforms by adding together simple ones A Luminance 0.0 0.2 0.4 0.6 0.8 1.0 Horizontal Position (deg) 11F F+3F+5F+7F+9F+11F 10 00 B Relative 1 00 Contrast 10 1 9F 7F 0.1 F+3F+5F+7F+9F F+3F+5F+7F 1 3 5 7 11 17 25 Spatial Frequency (cycles/deg) 10 C Relative Sensitivity 1 0.1 0.01 5F 3F F+3F+5F F+3F 1 3 5 7 11 17 25 Spatial Frequency (cycles/deg) D Relative Contrast 10 00 1 00 10 1 F 0.1 F 1 3 5 7 11 17 25 Spatial Frequency (cycles/deg) E Brightness 0.0 0.2 0.4 0.6 0.8 1.0 Horizontal Position (deg) Flashes of various durations shorter than the critical duration all have the same temporal frequency spectrum. Flashes longer than the critical duration contain less contrast at intermediate temporal frequencies, after filtering through the temporal CSF and are therefore less visible. Thus, more quanta are need to be added to bring them up to threshold. The critical duration for a brief flash against a background decreases as the luminance of a background light or area of the flash increases Log Threshold Retinal Illuminance (Td) 4 fovea 1o 3 Background Luminance (Td) 2 2500 3400 456 115 21 9.5 1.9 0 0.43 1 0 -1 -2 0 1 2 Log Flash Duration (msec) 3 Critical duration also depends on stimulus area. As the area of the flash is increased, the critical duration decreases. When the stimulus diameter is small (1.5 - 2 min arc), Bloch's Law holds for flash durations up to around 0.10sec (100 msec). If the test flash diameter increases to approximately 5 deg., Bloch's Law only holds for flashes up to about 30 msec in duration. For flash durations less than the critical duration, Bloch’s Law holds and 1. The flash cannot be seen when it is above threshold 2. The number of quanta in a threshold flash is the same for different flash durations 3. L x C = t 4. None of the above Five Parts to this Chapter Temporal Acuity (critical flicker frequency [CFF]) The Temporal Contrast Sensitivity Function Temporal Summation (Bloch’s Law & Broca-Sulzer) Masking Motion Detection (Real and Apparent) Part B – above-threshold brightness Supra-threshold flashes of a certain brief duration appear brighter than longer and shorter flashes of the same physical intensity (Broca-Sulzer effect) 600 500 400 300 170 lux Broca 126 lux 200 170 126 64.5 lux 100 64.5 32.4 16.2 Flash Duration (sec) 0.5 0.25 0.2 0.125 0.1 0 0.037 0.046 0.062 32.4 lux 16.2 lux 0.01 Comparative Brightness 700 Neural Explanation •Intense stimuli produce photoreceptor overshoot •This produces (via the bipolar cells) an initial burst of action potentials in the ganglion cells •Brightness is related to the firing rate of the cells (spikes/second) •For long flashes, the firing rate after the initial burst signals the brightness •For brief flashes, only the initial burst occurs, so the only information the neurons in central structures can use is a high firing rate, which makes the flash appear brighter than when it is long. Neural explanation of the Broca-Sulzer effect Note: the photoreceptor membrane potentials are upside down (negative is up on the graph) to demonstrate the similarity in shape to the BrocaSulzer effect. Neural Explanation •Intense stimuli produce photoreceptor overshoot •This produces (via the bipolar cells) an initial burst of action potentials in the ganglion cells •Brightness is related to the firing rate of the cells (spikes/second) •For long flashes, the firing rate after the initial burst signals the brightness •For brief flashes, only the initial burst occurs, so the only information the neurons in central structures can use is a high firing rate, which makes the flash appear brighter than when it is long. Five Parts to this Chapter Temporal Acuity (critical flicker frequency [CFF]) The Temporal Contrast Sensitivity Function Temporal Summation Masking (Temporal interactions between visual stimuli) Motion Detection (Real and Apparent) Temporal Interactions between Visual Stimuli Masking is any situation in which the detection of a visual stimulus is reduced by another stimulus presented before, during, or after the target stimulus. 1) masking of light by light, 2) masking of a pattern by light, and 3) masking of a pattern by a pattern. The effects of a masking stimulus may continue forward, after its cessation, and backwards, before its onset Log Threshold Test Field Energy Masking Stimulus (1.38o) 0 Test Stimulus (0.36o) Backward Masking might remind you of Early Dark Adaptation -1 -2 -3 -4 Mask On Mask Off Test On Test Off Forward masking Backward Masking masking 0 250 500 750 Test Field Onset Time (msec) 0 Test flash Masking flash Log Threshold Test Field Energy Masking Stimulus (1.38o) 0 Test Stimulus (0.36o) Backward Masking might remind you of Early Dark Adaptation -1 -2 -3 -4 Mask On Mask Off Test On Test Off Forward masking Backward Masking masking 0 250 500 750 Test Field Onset Time (msec) 0 Simultaneous and forward masking are signal detection problems 0 Mask follows target by 300 milliseconds 0 60 0 Mask follows target by 100 milliseconds 0 1000 Time (milliseconds) 2000 Spikes per second 60 Mask pulse alone 60 Spikes per second Spikes per second 0 60 Spikes per second Target pulse alone 60 Spikes per second 60 Spikes per second Spikes per second Backward masking may be explained by the response latency and duration of the test flash 60 Mask follows target by 50 milliseconds 0 Mask follows target by 20 milliseconds 0 0 Mask follows target by 10 milliseconds 0 1000 Time (milliseconds) 2000 Masking effects may occur when the test and mask are spatially separated metacontrast (backwards) and paracontrast (forwards) are masking in which the test flash and masking flash do not overlap spatially on the retina Masking effects do not require spatial coincidence of test and masking stimuli; they may occur when the test and mask are spatially separated by as much as 3 degrees This suggests that the same cells must be stimulated by the edges of both stimuli to obtain metacontrast When the gap between the stimuli becomes large enough, different populations of retinal neurons are stimulated by the test and masking flashes. Any masking has to occur upstream in the visual pathway, where receptive fields get larger Masking • Masking is any situation in which the detection of a visual stimulus is reduced by another stimulus presented before, during, or after the target stimulus. • Metacontrast (backwards masking with physicallyseparated stimuli) and paracontrast (forward masking with physically separated stimuli) • Dichoptic masking – masking where the two stimuli are presented to different eyes Dichoptic masking - A masking stimulus presented to one eye affects vision of a test stimulus at a corresponding retinal location in the other eye Cannot occur until inputs from the two eyes meet at a binocular cell in V1 or later Saccadic suppression Saccadic suppression is defined as a reduction in sensitivity to visual stimuli that occurs before, during and after a saccade (Look at your eyes in a mirror and try to see them move when you make a saccade) Decreased sensitivity (increased threshold) to visual stimuli occurring before, during, and after saccadic eye movements 100 100 Eyes moving 80 Visual suppression 60 60 40 40 Pupil suppression 20 20 0 0 -120 -80 -40 0 40 80 120 Pupil Response (percent) Visual Response (percent) 80 160 Time of Flash (msec) But you can see the strobe lights atop Red Mountain if you time your saccade just right Masking includes 1. 2. 3. 4. any situation in which the detection of a visual stimulus is reduced by another stimulus presented before, during, or after the target stimulus Paracontrast Dichoptic masking All of the above Five Parts to this Chapter Temporal Acuity (critical flicker frequency [CFF]) The Temporal Contrast Sensitivity Function Temporal Summation Masking Motion Detection (Real and Apparent) Motion is a continuous change in an object’s location as a function of time Three reasons motion detection is important: •detect moving objects against a background (see edges) •detect own motion through the environment •determine 3-D shape (crudely) Demo – shape from motion (If you can’t see the edges, you can’t see the object) http://www.biomotionlab.ca/Demos/BMLwalker.html Real Motion: Motion involve an image changing its location on the retina Contrast with smooth pursuit (moving the eyes smoothly). This prevents the image from changing its location on the retina. We are not studying smooth pursuit. There is an upper limit to our ability to see motion – stimuli can be moving “too fast to see” It turns out that the reason is that rapidly moving images have a temporal frequency that is too high for our visual system to detect (frequency is above the temporal highfrequency cutoff). To understand this – need to look at movement from the point of view of an individual retinal neuron. From the viewpoint of any one cell in the retina, motion is a change in luminance that occurs at a rate that depends on the speed with which the object moves and on the spatial frequency composition of the object •A high spatial frequency grating moving at constant velocity (degrees per second) has a faster temporal frequency than a lower spatial frequency moving at the same velocity This grating moved one full cycle Motion involves interactions of both spatial frequency and temporal frequency •Now, a lower spatial frequency (about half of the first one) moving at the same velocity (degrees per second). It has lower temporal frequency (cycles per second) at a given spot This grating moved about ½ cycle. Measured at the green dot (symbolizing a receptive field), it has a temporal frequency (flicker rate) about half of the higher spatial frequency Can determine the temporal frequency of a drifting grating by multiplying its spatial frequency times its velocity in degrees per second A 3 cycle/deg grating moving 10 deg/sec has a temporal frequency of 30 Hz; 30 cycle/deg =300 Hz As object velocity increases, spatial CSF shifts to lower spatial frequencies; temporal CSF remains constant Contrast Sensitivity A B Velocity (deg/s) 0 1 10 100 800 1000 100 10 1 0.01 0.1 1 10 Spatial Frequency (cycles/deg) 25 50 0.01 0.1 1 10 Temporal Frequency (cycles/s) 25 50 How fast a velocity can you see moving? The limiting factor in motion detection is the temporal resolution of the visual system. If you present a very, very low spatial frequency (and high contrast) can see motion of several thousand degrees per second The ability to see rapidly-moving (high velocity) objects 1. Is limited by the temporal frequency 2. Occurs only in the visual cortex 3. Is set by the velocity of the objects 4. Cannot be measured Apparent Motion: Apparent motion is the perception of real motion that can be produced when a stimulus is presented discontinuously. Phi phenomenon http://www.yorku.ca/eye/balls.htm Apparent Motion: Apparent motion is the perception of real motion that can be produced when a stimulus is presented discontinuously. The “rules” for producing apparent motion are the same as for real motion: the optimal stimulus duration and spacing is the same as would occur if a real object moved. Real vs. Apparent Motion Motion sampled stroboscopically appears like real motion due to the Optimal Distance (arc min) insensitivity of vision to high 200 temporal and spatial frequencies 100 50 20 10 5 2 0.2 0.5 1 2 Optimal Time (msec) 5 10 20 50 100 Burr and Ross, 1982 Van Deenna and Kimurama, 1982 Nakayama and Silverman, 1984 Kelly, 1979 200 100 50 20 0.2 0.5 1 2 5 10 Velocity (deg/sec) 20 50 100 To produce optimal apparent motion of 10 degrees per second, need each spot to be about 25’ apart and be on for about 35 msec. A real object, traveling at 10 degrees per second would move 21’ in the same time In order to make apparent motion look like real motion 1. You have to “fool” some of the neurons all of the time 2. You need a string of lights 3. You need real motion 4. You need to present the stimuli with the same separation and duration as would occur with real motion Detection of motion and sensitivity to direction of motion is achieved in hierarchic fashion in Areas V1 of the striate and middle temporal region of the cortex Newsome and colleagues sampled the activity of neurons in area MT Each cell has a receptive field that responded to motion in some location in the visual field (some retinal location). Each neuron was direction selective; it had an optimal direction (most spikes per second) and a null direction (fewer spikes per second). Stimuli with a range of correlation of the motion of the spots were used to determine threshold amount of correlation for the monkey, and also the threshold for neurons in the monkey’s area MT (in a twoalternative, forced-choice situation). A Number of Trials 20 Correlation - 12.8% 20 Correlation - 3.2% Correlation - 0.8% 20 Non-preferred direction Preferred direction 0 B Spikes per Trial 100 Percent Correct 1.0 0.9 0.8 0.7 0.6 0.5 Psychometric Function Neurometric Function 0.4 0.1 1 10 Correlation (%) 100 Using signal detection theory, a “neurometric” function could be produced for each neuron and compared with the monkey’s psychometric function Frequency of Occurence 7 Mean of Noise A 6 5 Maintained Discharge (Noise) Distribution 4 3 2 1 0 Overlap: Possible Confusion Mean of Noise + Signal 7 B 6 5 Maintained Discharge (Noise) + Response to Flash (Signal) Distribution 4 3 2 1 0 0 1 2 3 4 5 6 7 8 9 10 11 12 Number of Action Potentials in 50 msec Period 13 14 15 d'=1.5 A d'=1.0 B d'=0.5 C Srimulus Absent Stimulus Present ROC Curve A Number of Trials 20 Correlation - 12.8% 20 Correlation - 3.2% Correlation - 0.8% 20 Non-preferred direction Preferred direction 0 B Spikes per Trial 100 Percent Correct 1.0 0.9 0.8 0.7 0.6 0.5 Psychometric Function Neurometric Function 0.4 0.1 1 10 Correlation (%) 100 Using signal detection theory, a “neurometric” function could be produced for each neuron and compared with the monkey’s psychometric function The psychometric function for the monkey was matched well by directionselective neurons in area MT. Number of Neurons 20 15 10 5 0 0.1 Neuron more sensitive than the monkey 1 10 Monkey more sensitive than the neuron Threshold Ratio (neuron/behavior) Real & apparent motion seem to be detected by neurons in the parietal (MT) “stream” The monkeys’ “neurometric function” 1. Did not match the psychometric function 2. Could not be accurately estimated 3. Closely matched the psychometric function 4. None of the above Adapting to one direction of motion can produce a motion aftereffect when the movements stops (the “waterfall illusion”) May be due to neurons in MT Waterfall Illusion http://www.yorku.ca/eye/mae.htm Five Parts to this Chapter Temporal Acuity (critical flicker frequency [CFF]) The Temporal Contrast Sensitivity Function Temporal Summation Masking Motion Detection (Real and Apparent)