| Volume 3, Number 10, Article 2, Pages 599-609 |
doi:10.1167/3.10.2 |
http://journalofvision.org/3/10/2/ |
ISSN 1534-7362 |
Discrimination and identification of luminance contrast stimuli
Emily S. Kachinsky |
Visual Science Laboratories, University of Chicago, Chicago, IL USA |
|
Vivianne C. Smith |
Visual Science Laboratories, University of Chicago, Chicago, IL USA |
|
Joel Pokorny |
Visual Science Laboratories, University of Chicago, Chicago, IL USA |
|
Abstract
The goal of this study was to compare luminance contrast discrimination and polarity identification in the inferred Parvocellular (PC-) and Magnocellular (MC-) pathways. The position identification task tested ability to locate a contrast change within a stimulus. The polarity identification task tested ability to classify the contrast change as either brighter or darker. Three paradigms were employed to find these thresholds for the inferred MC- and PC-pathways: Pulsed-Pedestal, Steady-Pedestal and Pedestal-Δ-Pedestal. Position and polarity identification thresholds were the same for the Pulsed-Pedestal stimuli (inferred PC-pathway). The position identification thresholds were lower than the polarity identification thresholds for Steady-Pedestal stimuli (inferred MC-pathway). The position identification thresholds were lower than the polarity identification thresholds for Pedestal-Δ-Pedestal stimuli when the Δ-Pedestal contrast steps were sub-threshold or close to threshold. When the contrast steps were larger, position identification and polarity identification thresholds were similar for the Pedestal-Δ-Pedestal stimuli. Comparisons were also made between position identification and polarity identification thresholds for a short and a long stimulus presentation. There were no systematic differences dependent on presentation duration.
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|
History
Received February 26, 2003; published October 28, 2003
Citation
Kachinsky, E. S., Smith, V. C., & Pokorny, J. (2003). Discrimination and identification of luminance contrast stimuli.
Journal of Vision, 3(10):2, 599-609,
http://journalofvision.org/3/10/2/,
doi:10.1167/3.10.2.
Keywords
luminance contrast discrimination, luminance contrast identification, PC-pathway, MC-pathway
for related articles by these authors
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Thomas (1985)
provided a conceptual framework relating detection (the ability to distinguish a
change within a stimulus) and identification (the ability to classify a change
within a stimulus as one of a number of alternatives), postulating two
sequential stages. The first stage is the encoding stage; the physical stimulus
activates tuned pathways and is represented within the visual system according
to the pathway responses. This stage should be the same for both detection and
identification because the tasks use the same sensory information in the form of
the encoded signal. The second stage is the decision stage, during which a
judgment is made based on the combination of the pathway responses with respect
to a decision rule. This stage should be different for the tasks because
different decision rules are applied to the encoded signal ( Thomas, 1985; Watson & Robson, 1981). Noise limits the
accuracy of detection and identification due to random fluctuations inherent in
the pathways.
The identification task is often described in terms of
"labeled lines". These are tuned pathways that signal that a specific property
of interest is present in a stimulus, e.g. spatial frequency or contrast. On-
and Off-pathways are examples of labeled lines. Retinal ganglion cells in
On-pathways increment their firing rate to increments in contrast; cells in
Off-pathways increment their firing rate to decrements in contrast. For the
detection task, we may apply an inclusive-or rule, which means that detection
occurs if at least one pathway is activated ( King-Smith & Carden, 1976; Sachs, Nachmias & Robson, 1971). For example,
if the task is to detect a change in luminance from the steady state, activity
in either the On- or the Off-pathway (or both) may signal detection. However,
for the identification task, we may apply a simple maximum rule to the labeled
line information. This information is used to classify the stimulus property of
interest and to base the judgment according to which line or channel is most
active ( Thomas, 1985). In the example of a
contrast change, if the On-pathway has the higher amount of activity, the
stimulus will be identified as an increment in luminance. In this case, we are
comparing the relative amounts of activity within different labeled lines and
using that information to make a judgment about what happened.
In the current experiment, we were interested in
discrimination and identification of luminance contrast signals. Previous
studies showed that chromatic stimuli were identified at the detection
threshold, e.g. a "red-ward" pulse from white is seen as “reddish”
and a "green-ward" pulse from white is seen as “greenish” ( Gille, 1984; Mullen
& Kulikowski, 1990; Smith, Pokorny
& Sun, 2000). The result for chromatic stimuli is not surprising and can
be explained by a model of chromatic processing in the PC pathway ( Smith et al., 2000). Further, the information
carried in the PC-pathway is graded. The observer can report the polarity of a
chromatic pulse at the chromatic contrast discrimination threshold e.g. a
red-ward increment on a “pinkish” pulse is seen as “more
pink” and a decrement toward white is seen as “less pink”. The
PC-pathways comprise four parallel sub-channels carrying information about color
and luminance contrast ( Derrington, Krauskopf
& Lennie, 1984). A color contrast step from an equiluminant background
causes firing in two of these sub-channels. It is an open question if labeled,
graded responses are carried for both chromaticity and luminance in the
PC-pathway.
Achromatic contrast signals also activate the MC
pathways. Tolhurst & Dealy (1975)
found a small, 0.05 log unit difference between detection and identification
thresholds for achromatic contrast. The need for greater contrast for
identification than detection in Tolhurst & Dealy’s experiment may
reflect the fact that the pulse activates both On- and Off-pathways in multiple
channels of matched spatial frequency. In other experiments using achromatic
contrast, discrimination of different spatial frequencies was accurate at
detection threshold ( Furchner, Thomas &
Campbell, 1977; Nachmias & Weber,
1975) when the ratio of the frequencies was greater than three. Presumably,
the use of a spatial frequency variable selectively activated different spatial
frequency channels. This result might suggest that activation of an appropriate
labeled line is required for achromatic contrast identification in the
MC-pathway.
The rationale for the current study lies in our ability
to segregate achromatic processing in MC- and PC-pathways psychophysically ( Pokorny & Smith, 1997; Smith & Pokorny, 2003; Smith et al., 2000). In these experiments a
four square array replaced a constant luminance background at a fixed increment
or decrement in retinal illuminance. The contrast of one of the squares, chosen
randomly, was varied to elicit an increment or decrement threshold. We used two
paradigms that differed only in the stimulus pattern viewed during the
interstimulus interval. In the Pulsed-Pedestal Paradigm, the observer adapted to
the constant background and the entire four square array was pulsed on during
the trial. In the Steady-Pedestal Paradigm, the observer adapted to the four
square array and only one square varied during the trial. In the Pulsed-Pedestal
Paradigm, for large contrast steps
( Δ
I/I > +/- 0.1), discrimination was ascribed to activity in
PC-processing channels. In the Steady-Pedestal Paradigm, discrimination was
ascribed to activity in MC-processing channels. In a third paradigm the
Pedestal-Δ-Pedestal Paradigm, contrast discrimination at low contrast steps
(Δ I/I < +/- 0.1) was ascribed to the MC-pathway. The goal of the
present experiment was to assess contrast discrimination and polarity
identification in these inferred PC- and MC-pathways.
The test stimulus consisted of an array of four
1o x 1o squares with a 0.07o separations
between squares. The surround was 8o x 8o, filled the
separations, and had a luminance that was held constant at 12 candelas per meter
square (cd/m2;115 effective trolands (td)). The surround and test
chromaticities were metameric to the equal energy spectrum. The monitor was
viewed binocularly at a distance of 1 m. A chin rest was used for head
stabilization.
The stimuli were generated by a Macintosh Power PC
G4/350 computer with a 10-bit Radius Thunder Power 30/1600 video card and were
presented on a 17” NEC Multisync FE750 color monitor. The resolution of
the display was 832 x 624 pixels and the refresh rate was 75 Hz. An Optronics
OL754 spectroradiometer was used to measure the spectral power distributions of
the phosphors. The luminance of each phosphor was measured for 1024 levels of
input integer values, by a Minolta LS-100 luminance meter. Look-up tables were
constructed for each phosphor to convert the voltage integer value to luminance.
During the adaptation period, a spatially homogenous
surround was presented continuously. A four square array (pedestal), briefly
pulsed during the test period, replaced the surround at a fixed increment or
decrement in retinal illuminance. One of the squares (the test square) either
appeared at a higher or lower retinal illuminance than the other three.
During the adaptation period, the four square array
appeared continuously in the surround, with the array at a fixed increment or
decrement in retinal illuminance. Only the test square changed to a higher or
lower retinal illuminance during the test period.
Pedestal-∆-Pedestal Paradigm
The four square array was presented at a steady
pedestal retinal illuminance different from the surround during the adaptation
period. During the test period, the entire four square array changed in retinal
illuminance (the ∆-Pedestal) with the test square incremented or
decremented from the other three by a different amount.
Figure 1 is a movie
illustrating increment and decrement stimuli for the three
paradigms.
Figure 1. Illustrations of the stimuli of the three paradigms. Double-click on each image to activate the movie.
The observer’s task was to determine the position
of a luminance difference in the test array (position identification) and to
report the polarity (increment or decrement) of the luminance difference
(polarity identification). To eliminate external factors and ensure responses
were given under similar conditions, both questions were answered on every trial
(the 2X2 design). A position judgment is made concerning the location of the
test in the four square array; then a polarity judgment requires the
identification of the brightness of the test square relative to the remaining
squares. The test square could be an increment or decrement relative to the
array and four separate interleaved staircases were run, two based on correct
position, two based on correct polarity identification.
First, the observer adapted for two minutes to the
surround. There was an additional one-minute adaptation to the surround and
pedestal in the Steady-Pedestal and the Pedestal-∆-Pedestal Paradigms. A
small cross that served as a fixation aid disappeared at the beginning of the
trial period and reappeared following the trial period. The observer pressed one
of four buttons on a Gravis USB Gamepad controller to indicate the spatial
location of the test square and then pressed one of two buttons to indicate
whether the luminance difference was an increment or a decrement. No feedback
was given. The duration of the test was 26.7 ms given by two screen refreshes at
the 75-Hz monitor refresh rate. Additional data were collected on two observers
at 267 ms (twenty screen refreshes).
Trials consisted of four random alternating staircases.
There were two staircases for position identification, one for test square
increment thresholds and the other for decrement thresholds. There were also two
staircases for polarity identification, one for correct identification of
increments and one for correct identification of decrements. All staircases
began with an easily discriminable test contrast. Subsequently, the step size
decreased by half following reversals, until reaching a criterion step size
of 1% contrast after which there was no further change in the step size and staircases continued according to a reversal rule. For the position staircases, test contrast decreased after two correct responses and increased after one incorrect response. For the identification staircases, two correct position responses with correct identification led to a decrease in test contrast and one incorrect response for either position or identification or both led to an increase in test contrast. Thresholds for each staircase were estimated from the average of ten reversals per staircase. Each session lasted 20 to 30 minutes allowing the measurement of the four staircase thresholds for two or three of the starting pedestal retinal illuminances. For the Pulsed-Pedestal and Steady-Pedestal Paradigms and for each stimulus duration, two or three sessions were necessary to gather a complete set of thresholds. For the Pedestal-∆-Pedestal Paradigm, six sessions were
necessary to gather a complete set of thresholds. Thresholds for each condition
were calculated from the average of the three or four repetitions.
The five observers (EK, female age 25; AL, female age
30; ER, female age 21; LJ, female age 19; YS, male age 26) were all normal
trichromats as assessed with the Ishihara pseudoisochromatic plates and the
Neitz OT anomaloscope; all had Farnsworth 100-hue error scores of 32 or fewer.
Observers EK (an author) and AL were familiar with the experimental design and
psychophysical procedures. Observers ER, LJ and YS were untrained observers
recruited for the experiment and were naïve to the purpose and design of
the experiment. Observers EK, AL and YS participated in Experiment 1, EK and AL
performed all conditions and observer YS provided data at 26.7 ms confirming the
main results. Observers LJ and ER were recruited for Experiment 2. To provide
training and for comparison purposes we first collected baseline data of
Experiment 1. Observers EK, LJ and ER participated in Experiment 2. Written
informed consent was obtained from all observers.
In Experiment 1, we replicated the previous study
comparing Pulsed- and Steady-Pedestal Paradigms ( Pokorny & Smith, 1997), but used the 2X2 design to include the polarity task. The pedestal had one of five retinal illuminances: two dimmer (75 and 91 td) than the surround, two brighter (145 and 182 td) and one equal (115 td) to the surround. No systematic differences were found between increment and decrement thresholds in pilot data and these thresholds were averaged.
Figure 2 shows results for four observers (clockwise panels AL, EK, YS, ER) for the 26.67 ms stimulus; data for LJ are presented in the Appendix. Data are shown for both the Pulsed- and Steady Pedestal Paradigms (open and closed symbols respectively). The delta log retinal illuminance (threshold from the pedestal) is plotted as a function of log retinal illuminance of the pedestal expressed in trolands. The panels show the position (circles) and polarity identification (squares) thresholds (±2 SE). Figure 2. Position (circles) and polarity
identification (squares) for the Pulsed-Pedestal paradigm (open symbols) and the
Steady-Pedestal paradigm (closed symbols). The log D retinal illuminance is plotted as a function of the log retinal illuminance of the pedestal. The arrow in the graph indicates surround retinal illuminance. V-shapes are fits to Equation 1 in the text; lines are fits to Equation 2 in the text. Error bars are ±2 SE. Panels show data for four observers, AL, EK, ER, and YS.
For the Pulsed-Pedestal Paradigm, the thresholds for
position and polarity identification increased similarly as the pedestal
contrast increased with respect to the surround, forming a V-shape. The minimum
threshold values occurred when the pedestal was at the same retinal illuminance
as the surround (indicated by arrow on graphs). When the pedestal has equal
retinal illuminance to the surround, the two paradigms are replications. This
data point is not included in the analysis of the Pulsed-Pedestal Paradigm. Pokorny and Smith (1997) described the data as
reflecting the contrast response of the PC-pathway to achromatic contrast. Their
equation, revised in Smith, Sun & Pokorny,
(2001) is expressed as
logΔ I:
| logΔI
=
log{(Csat
+ |C|)2/
[Csat
–
Kc(Csat
+ C)]} + log(KPIs) | (1) |
where
Kc
represents a positive criterion ( Pokorny &
Smith,1997) and
KP
accounts for threshold sensitivity and gain. For the Pulsed Pedestal Paradigm,
the data did not approach saturation and
Kc
can be set at 0.001. The dashed lines show fits obtained by varying
Csat
and
KP.
The fits are good, with similar parameters for the three observers. Values of
Csat
were 0.76 - 1.5, consistent with previous work ( Pokorny & Smith, 1997), showing that
position judgments are always contrast
discriminations. For the Steady-Pedestal Paradigm,
the thresholds increased monotonically as pedestal retinal illuminance increased
for increment pedestals. Pokorny & Smith
(1997) showed that the data could be fit by a unit slope, implying
Weber’s Law. For pedestal decrements, a few thresholds departed from
monotonicity. This phenomenon was noted in earlier studies (Pokorny & Smith, 1997) and was ascribed
to stray light since the effect was increased with reduction in size of the
four-square array (Smith, Sun & Pokorny,
2001) and decreased with reduction in surround width. The solid lines are
linear fits to the data; obtained by varying
KM: | logΔI
= log(I) +
log(KMIs) | (2) |
The observers showed little variation in the
one free parameter. This result confirms previous data and suggests that for the
Steady Pedestal Paradigm, the position thresholds were always detections from
the pedestal luminance. Thresholds for polarity
identification were consistently higher
(~0.14 log unit) than position
thresholds for all observers. The figures additionally showed that the polarity
identification thresholds for the Steady-Pedestal Paradigm were below the
contrast discrimination threshold for the Pulsed-Pedestal Paradigm. This result
suggests that polarity identification for the Steady-Pedestal Paradigm was
mediated in the presumed MC-pathway.
We calculated the difference thresholds between
position and polarity identification for the two paradigms, for each observer.
For the Pulsed-Pedestal Paradigm, the difference threshold at the surround
luminance was combined with the Steady-Pedestal Paradigm data. The difference
thresholds were evaluated by ANOVA. For the Pulsed-Pedestal Paradigm, the
difference thresholds ranged from –0.011 to 0.051 among observers and were
not significant. For the Steady-Pedestal Paradigm, the difference thresholds
ranged from 0.085 to 0.189 among observers and were significant. For two
observers we collected data using a 267 ms duration. Thresholds showed an
increase in sensitivity and the data were parallel to the 26.7 ms data. Polarity
identification for long duration stimuli in the Steady-Pedestal Paradigm did not
improve due to temporal separation of the pulse onset and offset.
Our average separation between position and polarity
discrimination on the Steady-Pedestal Paradigm was larger than noted by Tolhurst & Dealy (1975). For the zero
pedestal, the four observers showed greater sensitivity than predicted for
detection and the separation between detection and polarity identification was
less, more similar to Tolhurst & Dealy
(1975).
Experiment 1 showed that polarity identification
thresholds were slightly higher than position thresholds in the Steady-Pedestal
Paradigm and we have argued that both thresholds were mediated in the
MC-pathways. Two possibilities suggest themselves. First, we have argued that
the MC-pathway adapts to the Steady-Pedestal. The pulse thus excites both On-
and Off-pathways as suggested in the introduction for the zero pedestal
condition. Polarity identification might depend on pathway isolation. Previous
work by Tolhurst (1975) revealed that at low
spatial frequencies responses to the onset and offset of a pulse were about
equal when they were independently detectable events spaced in time. We used the
long duration pulse in an attempt to separate the responses of the On- or the
Off-pathway by separating their onset and offset in time. We thought that the
observer might be able to attend separately to the onset and offset. However,
use of the longer duration did not improve polarity identification. A second
explanation might be that the contrast pedestal does favor the correct pathway
but the signal is noisy. Polarity identification could require a larger signal
in order to maintain accuracy. We can isolate the MC-pathway On-and Off-pathways
by using pulsed pedestals of very low contrast ( Pokorny & Smith, 1997). However there is
only a small range to observe the MC-pathway contrast response since there is a
transition to presumed PC-pathway function for contrast steps > 0.15 ( Pokorny & Smith, 1997). By placing the
entire array on a pedestal, we can extend the available range between the
presumed PC- and MC-pathways; this is the Pedestal-Δ-Pedestal Paradigm. The
Pedestal-Δ-Pedestal Paradigm revealed the contrast discrimination behavior
of the presumed MC-pathway with a steep V-shape ( Pokorny & Smith, 1997). Further, a study
of the time course of recovery from the Δ-Pedestal showed that increment
and decrement thresholds showed different time courses ( Pokorny, Sun & Smith, 2003). The
data suggested that increment discriminations at Δ-Pedestal onset were
mediated in On-pathways and decrement discriminations were mediated in
Off-pathways. The increment and decrement staircases should thus be averaged
separately although this separation was not performed in the initial study.
Further, the smallest contrast steps may fall in the range where threshold
summation has been reported ( Foley & Legge,
1981), although this also was not observed in previous work ( Pokorny & Smith, 1997; Pokorny et al., 2003).
In Experiment 2, the steady pedestal retinal
illuminance was 182 td and there were 12 values of the ∆-Pedestal varying
between 91 and 363 td. These included eight small luminance steps
(155,162,170,174, 191, 195, 205, and 214 td) adjacent to the steady pedestal and
four large luminance steps (91, 115, 289, and 364 td). The stimulus duration was
26.7 ms. Increment and decrement thresholds for both position and polarity
identification were averaged separately.
The data of Experiment 2 are displayed in Figures 3-5 for observers EK, LJ and ER,
respectively. Each figure compares results for position (circles) and polarity
identification (squares) in separate panels for the increment (upper panel) and
decrement (lower panel) staircases. The dashed lines are predictions based the
steady Pedestal Paradigm of Experiment 1. The dashed V-shape is from the
PC-pathway prediction used for the Pulsed Pedestal paradigm of Experiment 1,
adjusted to predict the conditions of the Pedestal-Δ-Pedestal Paradigm.
There are two separate regions of the graphs to consider: the outlying regions
where the Δ-Pedestal contrasts are high and the region near the steady
pedestal where the Δ-Pedestal contrasts are low. When the Δ-Pedestal
contrasts are > +/–0.35, the thresholds (the four outlying points)
correspond to the dashed line prediction of the PC-pathway. We interpret these
thresholds at high Δ-Pedestal contrasts to reflect the PC-pathway contrast
response function. Position and polarity identification are equivalent for both
increment and decrement thresholds, consistent with the Pulsed Pedestal data of
Figure 2. Where the Δ-Pedestal contrasts
are < +/–0.15, thresholds rise sharply from the steady pedestal
threshold. This sharp rise shows the rapidly saturating contrast response of the
MC-pathway. The V-shape indicates that these are contrast discriminations
mediated in isolated On- and Off-pathways. Contrast discrimination and polarity
identification are
equivalent.
Figure 3. Comparison of position
(circles) and polarity identification (squares) for the
Pedestal-∆-Pedestal Paradigm. The upper panel shows increment data and
the lower panel shows decrement data. The dashed lines are the fits to the
Steady-Pedestal data of Experiment 1 and the V-shape is the predicted PC-pathway
function from the Pulsed-Pedestal Paradigm adjusted to the
Pedestal-∆-Pedestal Paradigm. The solid V-shapes are fits to the combined
data for both panels based on Equation 3 in the text. These fits are described further in the text. The data are for observer EK.
Figure 4. Comparison of position and
polarity identification for the Pedestal-∆-Pedestal Paradigm. The plot
format is as for Figure 3; the observer is
LJ.
Figure 5. Comparison of position and
polarity identification for the Pedestal-∆-Pedestal Paradigm. The plot
format is as for Figure 3; the observer is
ER .
Within the region of rapid threshold rise, the
increment and decrement thresholds show an asymmetry about the steady pedestal
luminance. Increment thresholds are lower than decrements on increment
Δ-Pedestals and decrement thresholds are lower than increment thresholds on
decrement Δ-Pedestals. The results are consistent across observers and for
both discrimination and polarity identification tasks. A similar result was
previously reported by Pokorny, (Pokorny, Sun
& Smith, 2003). For increment staircases on increment Δ-Pedestals
and for decrement staircases on decrement Δ-Pedestals, the effective
contrast is reduced by a factor (0<κ<1). Equation (1) was thus rewritten to reflect the
MC-pathway
response:
| logΔI
=
log{(Csat
+ |C
κ|)2/[Csat
–
Kc(Csat
+ |C κ |)]} + log(KMIs) | (3) |
The solid lines are fits to the combined position and
polarity identification data using Equations
(1) and (3) with individual adjustment of
Csat,
κ, and
KM.
The value of
Kc
was set at 0.01. Compared with the fits of equation (1) to Pulsed Pedestal data,
the value
Csat
was at a lower value (usually near 0.1–0.15) consistent with the steep
MC-pathway saturation ( Pokorny & Smith,
1997). For observer LJ, we allowed an additional constant added to the
polarity identification for decrements (see Appendix). The fits were good with
little variation in the parameters among the observers. There were
over-estimations near the steady pedestal luminance where the Δ-Pedestal
contrast ( ~0.05) is
sub-threshold for our observers. At pedestal contrast near 0.05, measured
thresholds were more sensitive than for the zero pedestal for increments
measured on the increment pedestal (upper panels) and for decrements measured on
the decrement pedestals (lower panels). These data may reflect the effect of
sub-threshold summation, when the staircase and the Δ-Pedestal are in the
same direction.
Subthreshold effects have been noted for simple spatial
displays in the achromatic ( Bowen, 1995) and in
the chromatic domains ( Cole Stromeyer &
Kronauer, 1990). Although sub-threshold summation was originally attributed
simply to physical addition of light, this explanation was discredited with the
arrival of signal detection accounts of discrimination ( Nachmias & Kocher, 1970). The current
explanations for dipper effects include a non-linear transducer ( Chen, Foley & Brainard, 2000; Graham, 1989; Nachmias & Kocher, 1970) and uncertainty
reduction ( Pelli, 1985). Neither approach
explains the bumper effect ( Bowen, 1995). For
our simple displays, the physical addition of light offers the simplest account
of both phenomena. We assume that with sub-threshold pedestals the observers
revert to detecting the odd square against the background rather than
discriminating among four squares. This conclusion for simple color stimuli was
also reached by Eskew (1999).
These threshold interactions do not occur uniformly among observers. We reviewed
data collected in our laboratory for the Pedestal-Δ-Pedestal paradigm ( Pokorny & Smith, 1997), but did not find
consistent evidence of sub-threshold effects. In this previous work,
sub-threshold pedestals were not included since assessment of threshold
interactions was not the goal of the study. Additionally, some of the variation
may reflect observer variability. Our three observers in Experiment 2 were not
as highly trained as in the previous work, and showed higher thresholds
overall.
The predictions describe both contrast discrimination
and polarity identification above detection threshold. In this region, the
appropriate pathway is isolated and contrast discrimination and polarity
identification are the same.
In the Pulsed-Pedestal Paradigm, thresholds for
contrast detection and polarity identification were the same for all non-zero
pedestal contrasts. Previous studies found no differences between chromatic
contrast detection ( Gille, 1984; Mullen & Kulikowski, 1990) or chromatic
contrast discrimination ( Smith, Pokorny &
Sun, 2000) and identification. These results suggest that the PC-pathway
does indeed have labeled lines, which signal polarity identification for both
chromaticity and luminance at detection threshold. The results suggest that the
four retinal pathways, +L+M, +L-M, -L+M and –L-M, combine at higher
cortical areas in different combinations to separate luminance and chromatic
information. This interpretation has been called demultiplexing ( Ingling & Martinez-Uriegas, 1983; Lennie & D'Zmura, 1988). However,
physiological evidence of demultiplexing has not yet been found ( Lennie, Krauskopf & Sclar, 1990). Of note
is that both for color and for luminance the cortex extracts a continuous graded
signal.
The situation in the MC-pathway is different. At steady state, both On- and Off-pathways are active and contribute to detection. Polarity identification for our display required about 0.14 log unit more contrast. We thought that observers might have better identification performance with long pulses for which they could separate attention to the onset and offset of the Pulsed Pedestal. However, they apparently did not use this strategy. With the Pedestal-∆-Pedestal paradigm, increment contrast discrimination was
determined in On-pathways and decrement contrast discrimination was determined
in Off-pathways. When the pathway was isolated, polarity identification was at
contrast discrimination threshold except in the region where the
∆-Pedestals were sub-threshold. This result supports the hypothesis that
polarity identification requires isolation of an appropriate pathway.
The data of Figures 3 -
5 showed very high thresholds for decrement staircases measured with
increment pedestals and increment staircases measured with decrement pedestals.
The thresholds were in fact higher than the predicted PC-pathway thresholds.
This phenomenon might reflect a cognitive component, e.g. the observer is
monitoring only the MC-pathway until it saturates and the ∆-Pedestals are
above PC-pathway threshold. Alternatively, perhaps the close alignment of the
squares interferes with the observers' judgment. We thought it would be
instructive to look at the actual illuminances at the threshold ΔI. Figures 6 - 8 show this calculation for the three
observers. This plot is in linear units of retinal illuminance. The abscissa
shows the ∆-Pedestal illuminance; the ordinate shows the threshold
illuminance at the discrimination step for position (open symbols) and for
polarity (closed symbols). The vertical and horizontal lines dividing the plot
frame into quadrants are placed at the steady pedestal illuminance. Increment
∆-Pedestals occur in the right hemi field; decrement ∆-Pedestals
occur in the left hemi field. Increment thresholds with increment pedestals fall
in the upper right quadrant above the diagonal (shown as a dashed line) and
decrement thresholds with decrement pedestals fall in the lower left quadrant
below the diagonal. These results were as expected. The unexpected results shown
by the plot concern the decrement staircases with increment ∆-Pedestals.
These thresholds fell in the lower right quadrant indicating that discrimination
and polarity identification of a decrement required a decrement from the steady
pedestal luminance. A parallel result occurred for the increment staircases with
decrement ∆-Pedestals. These thresholds fell in the upper left quadrant
indicating that both discrimination and polarity identification required an
increment from the steady pedestal luminance. It can be seen that this untoward
event occurred only for the ∆-Pedestals that were ascribed to the
MC-pathway. At higher contrasts (stippled area), where we have suggested that
the MC-pathway saturates and the PC-pathway mediates achromatic discrimination,
the thresholds are distributed in the appropriate quadrants: increments above
the diagonal, decrements below the diagonal. These results are consistent with
the continuous graded achromatic response signaled by the PC-pathway. The
results suggest that the MC-pathways are labeled by pathway only and do not
signal a graded response. Under our stimulus conditions, the response
“brighter” is signaled only by On-pathways while the response
“dimmer” is signaled only by
Off-pathways. Figure 6. The test square illuminance at
the position and the polarity identification thresholds plotted as a function of
the ∆-Pedestal illuminance in trolands. Open symbols are for position
identification; closed symbols are for polarity identification. Increment
thresholds are shown by circles and decrement thresholds are shown by squares.
The solid horizontal and vertical lines indicate the retinal illuminance (183
td) of the steady pedestal. The dashed line indicates the illuminance of the
∆-Pedestal. Icons across the top correspond to (from left to right): a decremental pedestal (decrement test), a pedestal equal to the surround (decrement test) and an incremental pedestal (increment test). The observer is EK.
Figure 7. The test square illuminance at
the position and the polarity identification thresholds plotted as a function of
the ∆-Pedestal luminance in trolands. The plot format is as for Figure 6; the observer is LJ.
Figure 8. The test square illuminance at
the position and the polarity identification thresholds plotted as a function of
the ∆-Pedestal luminance in trolands. The plot format is as for Figure 6; the observer is ER.
In summary, we suggest that graded polarity signals are
generated in the PC-pathway for both chromatic and achromatic contrast. For the
MC-pathway, isolation of On- and Off-pathways can lead to correct polarity
identification but this information is not graded.
Response bias: In previous work with contrast
discrimination paradigms, we have noted no differences in increment and
decrement staircases using the Pulsed- and Steady-Pedestal Paradigm. It was on
this basis that we averaged data for increment and decrement thresholds. In
fact, no differences occurred in the present work for the discrimination task.
Further, no differences in the increment and decrement staircases were found for
the position task for the Pulsed-Pedestal Paradigm.
Our method does allow for the possibility of response
bias in polarity identification in the Steady-Pedestal paradigm where many
trials are detected but not identified. Since there are only two choices for
identification, "brighter" or "dimmer", response bias could occur if the
observer tended to favor one response on trials where the position was correctly
identified but the polarity identification was below threshold. Since staircases
are driven by the correct responses, response bias could preferentially drive
the polarity staircase. Suppose the observer tends to respond "dimmer". The
number of correct identifications on the decrement staircase would be above
chance. Decrement staircases would approach the detection threshold. In
examining the data of Experiment 1, two observers showed no evidence of response
bias for polarity identification. One observer (AL) showed a bias to "dimmer".
For the 26.7ms pulses her average (Polarity – Detection) difference of
0.107 reflected a difference of 0.1625 on the increment staircase and 0.05 on
the decrement staircase. Two new observers were recruited for Experiment 2. The
data of ER (shown in Figure 5) were similar to
those of AL, with a similar mild bias to the "dimmer" response. ER showed an
average polarity-detection difference of 0.124 composed of 0.05 for decrements
and 0.24 for increments. For LJ however, a very pronounced response bias to
"brighter" was noted in Experiment 1 and shown in Figure 9. For the Pulsed-Pedestal Paradigm,
increment and decrement thresholds were the same and were averaged. Contrast
discrimination and polarity identification were also the same. All the data
followed the two rising arms of the PC-pathway contrast response. For the
Steady-Pedestal Paradigm, the position judgments were the same for increment and
decrements and were averaged. However, the polarity identification staircases
were different for increments and decrements. The increment polarity
identification was the same as the detection. However, the decrement polarity
identification thresholds fell on the data for the Pulsed-Pedestal Paradigm. In
the Pedestal-Δ-Pedestal Paradigm, the decrement contrast discrimination was
in the Off-pathway but there was still a deficit in polarity identification.
These data were consistent with a pure response bias. The observer told us that
she was trying to guess when she could not tell the polarity, however her
behavior did not match her
words. Figure 9. Comparison of the
Pulsed-Pedestal (open symbols) and Steady-Pedestal (closed symbols) paradigm for
observer LJ. Position (circles) and polarity (squares) identification is shown
for the Pulsed-Pedestal. Polarity identification for the Steady-Pedestal is
separated by staircase with increments shown by upright triangles and decrements
shown by inverted triangles.
NEI Research Grant EY00901 and NIMH Training Grant T32 MH20029 supported this research. We thank our observers for their time and attention, and Linda Glennie for assistance with programming. We thank Patrick Monnier and Barry Lee for useful discussions of the results. Publication supported in part by an unrestricted grant to the Department of Ophthalmology and Visual Science from Research to Prevent Blindness.
Commercial relationships: none.
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