Tatiana Pasternak1-3 and William H.
Merigan"
Motion Perception following
Lesions of the Superior Temporal
Sulcus in the Monkey
We examined the effect of bilateral ibotenic acid lesions,
aimed at areas MT/MST in three macaques, on their
perception of motion. The medial boundary of the le-
sions in the three monkeys was near the dorsal end of
the STS, but the lesions extended different lengths ven-
tralry along the STS. The lesions extended the shortest
distance ventrally in monkeys 1 and 2, covering most
of MST but possibly sparing a portion of lateral MT.
That in monkey 3 damaged all of MT and MST bilaterally
and extended through most of FST. All three lesions
caused a temporary disruption, followed by at least
partial recovery, of most motion thresholds. Permanent
effects of the lesions on visual sensitivity were graded
with lesion extent Contrast sensitivity for detecting
low-spatial-frequency (1 cycle/degree) drifting gratings
over a wide range of drift rates, as well as for identifying
their direction of motion, was slightly affected only in
monkey 3. Only monkeys 2 and 3 showed a deficit in
discriminating stimulus speed, and the size of the loss
was two- to fourfold. Discrimination of opposite direc-
tions of dot pattern motion, which required integration
of local motion signals, was mildly affected in monkeys
2 and 3, and not affected in monkey 1. However, ad-
dition of directional noise to this discrimination caused
the performance of all monkeys to be permanently dis-
rupted, especially that of monkeys 2 and 3. Finally,
direction difference thresholds were elevated by a factor
of 2-4 after the lesions in all three monkeys. Many of
these deficits were more pronounced during the first 2
months of testing following the lesion.
Thus, our results demonstrate that areas within dor-
sal STS make an important contribution to the perfor-
mance of various motion perception tasks including the
discrimination of smaD differences in direction and speed,
and the perception of global motion in the presence of
directional noise. The residual motion perception, even
in the monkey with virtually complete removal of areas
MT/MST, may suggest either that these tasks are nor-
mally mediated in part by cortical areas outside of areas
MT and MST, or that the disrupted functions were par-
tially assumed by other cortical areas after lesions.
Departments of "Neurobiology and Anatomy, and
2
Ophthalmology, and 'Center for Visual Science,
University of Rochester, Rochester, New York 14642
Physiological properties of neurons in cortical areas
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MT and MST of the macaque monkey suggest an im-
portant role of these areas in the processing of visual
motion signals. Neurons in area MT have large re-
ceptive fields, many are directionally selective, and
some respond to an integrated direction of motion
when stimulated with the motion of multiple, ori-
ented moving patterns (Zeki, 1974; Maunsell and Van
Essen, 1983; Albright, 1984; Movshon et al., 1985;
Rodman and Albright, 1989). In addition, recent phys-
iological evidence (Newsome et al., 1989; Britten et
al., 1992) indicates that the sensitivity of MT neurons
to visual motion is similar to that of the psychophys-
ically tested monkey. Additional evidence that MT
may be important for motion perception is the dem-
onstration that microstimulation of small numbers of
MT neurons can alter the visual choices of a monkey
toward the direction to which the stimulated neurons
are most sensitive (Salzman et al., 1992). Similarly,
recordings from the adjacent area, MST, also suggest
a role in motion processing, specifically in the analysis
of object motion and optic flow (Duffy and Wurtz,
1991; Tanaka et al., 1993) and in pursuit eye move-
ments (Komatsu and Wurtz, 1988).
Lesions of MT and MST affect motion perception
in a way that suggests a central role for these areas.
Lesions of areas MT (Newsome et al., 1985) and MST
(Dursteller and Wurtz, 1988) resulted in deficits in
smooth pursuit eye movements that suggested im-
paired perception of the velocity of motion, although
these effects were transient and recovered within a
week (Yamasaki and Wurtz, 1991). Newsome and Pare
(1988) used dynamic random dots to examine the
discrimination of motion direction. They reported def-
icits in thresholds for discriminating opposite direc-
tions of motion that were severe on the day of the
lesion, but largely recovered after a week. When the
lesion was larger, and included both MT and a large
portion of MST, there was a twofold threshold ele-
vation, which appeared to be permanent.
While the above work suggests some role for MT/
MST in motion perception, the limited and largely
transient deficits following lesions of these areas
question how essential they are. The basis for this
rapid recovery of function is an important and as yet
unresolved question. Related questions that formed
the basis of the present study are the possibilities (1)
that other motion-related capacities might be sub-
stantially and permanently affected by MT lesions and
Cerebral Cortex May/June 1994;4.247-259; 1O47-3211/94/M.OO
(2) that extensive lesions involving both MT and MST
would have even more dramatic effects on some mo-
tion-related thresholds.
Materials and Methods
Subjects
Three adult monkeys were used. On weekdays, water
was restricted for a period of 23 hr before testing and
they received their daily water ration, in the form of
fruit juice, during behavioral testing. On weekends,
the monkeys were not tested behaviorally and had
unrestricted access to water. Food was continually
available in the home cage, and their body weights
were monitored daily. The three monkeys (with orig-
inal numbers in parentheses) were labeled 1 (047),
2 (854), and 3 (9103) in order of the completeness
of their MT/MST lesions to facilitate clarity of de-
scription of their behavioral deficits.
Stimuli and Behavioral Procedures
Grating Stimuli
Stimuli. Stimuli and behavioral procedures were sim-
ilar to those described in Pasternak and Leinen (1986).
The stimuli were sinusoidal, moving or stationary ver-
tical gratings that were generated with an Innisfree
stimulus generator on a Hewlett-Packard 1332a os-
cilloscope screen (P-31 phosphor). The gratings ap-
peared behind two 6° diameter circular openings in
a white surround of the same mean luminance (75
cd/m 2 ). All gratings had a slow, smoothed onset (200
msec raised cosine), and they remained on until the
first response after 2 sec. Contrast sensitivity functions
for detecting gratings and discriminating their direc-
tion of motion were measured for 1 cycle/degree grat-
ings at temporal frequencies from 2.24 Hz to 19 Hz.
Difference thresholds for speed were measured over
a range of stimulus contrasts, with grating contrast
defined as ( £ m - £„,„)/(£„„ + Z ^ ) , where L^ and
Zmta are the maximum and minimum luminance in the
grating. The mean contrast values were 1.5%, 3%, 6%,
12%, 24%, and 48%. To eliminate cues due to differ-
ences in apparent contrast between the standard and
comparison gratings, contrast was randomly varied
±15% around the mean for each trial, except at the
highest contrast level (48%), where a single contrast
value was used. Finally, spatial frequency difference
thresholds were measured with the same stimuli. All
measurements were performed at a distance of 35 cm.
The grating stimuli usually remained on until the
response was made.
Contrast sensitivity and speed discrimination. The
monkeys were trained to perform three different tasks:
to detect the presence of a moving grating, to dis-
criminate its direction of motion, and to discriminate
small differences in the speed of moving gratings. The
behavioral procedure was identical for all three tasks.
The monkeys viewed the comparison stimuli binoc-
ularly, and were rewarded for pushing one of two
buttons that was located on the same side as the cor-
248 Effects of MT/MST Lesions on Motion Thresholds • Pasternak and Merigan
rect stimulus. Incorrect responses resulted in a 6 sec
tone and no juice reward. To eliminate position bias,
a correction procedure was used: three consecutive
incorrect responses on the same side resulted in a
trial being repeated until the animal made a correct
response. Data from correction trials were discarded.
Responses were effective only after the stimulus was
present for 2 sec. In the detection task, the monkeys
viewed a uniform field and a moving grating of the
same mean luminance, and were rewarded for choos-
ing the target containing the grating. In the direction
discrimination task, they viewed two identical grat-
ings moving at the same speed in opposite direction,
and chose the grating that moved rightward. In the
speed discrimination task, the monkeys were pre-
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sented with two gratings of the same spatial frequency
moving in the same direction at two different rates.
They were rewarded for choosing the target that moved
faster.
Initially the monkeys were trained to detect and
to discriminate the direction of motion of a high-
contrast, 1 cycle/degree grating, drifting at 2.24 Hz.
After they acquired both tasks (criterion: three con-
secutive 200 trial sessions >90% correct, or four con-
secutive sessions >80% correct), they were trained
to discriminate differences in speed. Thresholds were
measured with a staircase procedure in which three
consecutive correct responses resulted in a decrease
of contrast (or for speed thresholds, a decrease in the
speed of the comparison stimulus), while each in-
correct response resulted in an increase of contrast
or the difference in speed between the two compar-
ison stimuli. A psychometric function was constructed
from the results of all trials, and contrast and speed
thresholds were calculated by interpolation at the
stimulus value consistent with 75% correct. Contrast
thresholds were converted into contrast sensitivity by:
I/contrast threshold x 100, while speed thresholds
were converted into Weber fractions: increment in
speed/base speed (AV/V). Each testing session con-
sisted of 200 trials, and the monkeys were tested in
two consecutive sessions. Thus, contrast sensitivities
for detection and direction discrimination for a given
stimulus were usually measured on the same day.
Spatial frequency discrimination. This function
was measured using stimuli identical to those used
in other discriminations to see if the effects of MT/
MST lesions were specific to motion tasks. This could
be determined by comparing the smallest discrimin-
able difference in spatial frequency measured here to
the smallest discriminable difference in speed mea-
sured in the motion tasks. In this task the monkey
discriminated between two stationary gratings of
identical contrast, that differed only in spatial fre-
quency. A base spatial frequency of 1 cycle/degree
was always compared with a higher spatial frequency,
and the monkey was rewarded for responding to the
higher frequency. Temporal onset of the gratings fol-
lowed a cosine envelope of 200 msec rise time. A
staircase procedure, identical to that described above
for speed discrimination, was used and threshold was
taken at a value corresponding to 75% correct per-
formance. Measurements were performed at two con-
trast levels: 12% and 50%.
Dynamic Random Dot Stimuli
Stimuli. Dynamic random dot targets, used in the
present study, were identical to those used recently
by Pasternak et al. (1990). The stimuli, first intro-
duced by Williams and Sekuler (1984), consisted of
dots repeatedly displaced with a direction of motion
chosen randomly from a uniform distribution of di-
rections. The speed with which the dots were dis-
placed was determined by the step size (Ax) and the
temporal interval between steps (At), and was either
5 or 15 degrees/sec. When the range of the distri-
bution of directions was 360°, there was only local
random motion of individual dots, and no coherent
motion could be seen. However, when the distribu-
tion was less than about 320-340°, the dots appeared
to flow coherently in the direction of the mean of the
distribution. The saliency of the stimulus was manip-
ulated by varying the range of the direction distri-
bution (direction range) and/or the proportion of the
dots moving in nonrandom directions (% motion sig-
nal).
The dot stimuli were displayed for 2 sec on two
adjacent regions of the screen of a Tektronix 606 os-
cilloscope (P-31 phosphor), placed at a distance of
35 cm from the observer, behind two immediately
adjacent 7° diameter circular openings in a white sur-
round. Each stimulus contained 50 dots, the dot den-
sity in all experiments was 0.94 dots/degree 2 , and the
mean luminance of the display was 0.1 cd/m 2 . Each
dot was 0.08° in diameter and its luminance was set
3.5 log units above detection threshold for human
observers. Under these conditions there was no in-
dication of streaks produced by moving dots.
Tasks. The monkeys were trained on two tasks in-
volving random dots: discrimination of opposite di-
rections, and discrimination of small differences in
direction. Thefirsttask was identical to that used with
gratings. The monkeys viewed two stimuli consisting
of rightward and leftward moving dots, and were re-
warded for choosing the rightward motion. The
broadest range of directions in the stimulus that al-
lowed the animals to perform the discrimination at
75% correct was determined at several levels of mo-
tion signal: 100%, 50%, 25%, 12%, and 6%. In addition,
a threshold for the limit of motion signal was mea-
sured with direction range held at 0°. In the second
task, one stimulus moved to the right and the other
downward but less than 90° from rightward, the mon-
keys were rewarded for choosing rightward motion,
and the minimal discriminable difference in direction
was measured. Direction difference thresholds were
measured for the following direction ranges: 0°, 90°,
180°, and 270°. Other procedural details were iden-
tical to those described above for gratings.
Preoperative Testing
Preoperative testing was performed over a period of
3-6 months for different monkeys. Since during the
initial data collection improvement in thresholds with
continued testing is common, a large number of
threshold determinations was performed until there
was no indication of further improvement. To reach
stable performance it was often necessary to repeat
the testing 4-10 times. At least four threshold deter-
minations contributed to the mean used as a baseline
threshold estimate at each stimulus parameter. In all
tasks, the sequence of testing at each contrast level
(speed discrimination), temporal frequency (contrast
thresholds), range, step size, and so on, was irregular,
with interleaving of different conditions.
Postoperative Testing
Since the present studies were designed to examine
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the permanent effects of lesions, rather than transient
deficits that occur immediately after the lesions (e.g.,
Newsome and Pare, 1988), postoperative testing be-
gan between 3 and 7 d after the lesions and continued
over a period of 6 months. Nonetheless, in all three
animals we were able to assess the extent of early
deficits on some of the tasks. The sequence of testing
following the lesions was as follows. In monkey 1,
single threshold determinations were performed for
all tasks during the first 2 weeks of the postoperative
behavioral testing. After that, systematic measure-
ments of each visual function resumed. In all mon-
keys, we made at least three to five measurements of
speed, contrast, direction, and range thresholds. In
cases when animals were unable to perform discrim-
inations, attempts to measure thresholds were re-
peated several times. When there was any sign of im-
provement, the monkeys received additional testing
to determine at what level performance would sta-
bilize. In monkey 2, range thresholds were measured
first, followed by contrast sensitivity testing at a single
temporal frequency (9 Hz), followed by direction dif-
ference threshold testing and speed discriminations.
After these initial measurements, the monkey re-
ceived extensive systematic testing on all tasks. In
monkey 3, contrast thresholds for detecting and dis-
criminating direction over a range of temporal fre-
quencies were measured first. This was followed by
measurements of speed discriminations, and finally
by range and direction thresholds for random dots.
The performance on all tasks stabilized after the first
2 months of testing. Thresholds for all tasks were
rechecked during the last month of the experiment,
6 months after the lesion.
Ibotenic Acid Lesions
Lesions were made by injecting ibotenic acid (10 /ig/
lil) at multiple sites along the superior temporal sul-
cus (STS) in both hemispheres. Injections began near
the dorsal end of the STS and continued along the
STS for about 8 mm. Individual injections were made
with about 2.5 mm separation anterior and posterior
to the lumen of the STS along its dorsalmost 8 mm.
Injections were made with a Hamilton syringe and
their location based on direct visualization of the STS.
In monkey 1, 106 and 118 injections of 1.5 MI (total,
3360 ng) were placed in the left and right hemi-
spheres, respectively. In monkey 2, 73 and 74 injec-
Cerebral Conex Mayflune 1994, V 4 N 3 249

Rgore 1 . Representative Kstotogical section from which the intenic acid lesions were reconstructed. This is a photograph of a 40-jim-thick horizontal section of the brain of
monkey 1 stained for CO to illustrate the lesion location. In tfw photograph up is lateral and leh is anterior. The locations of three suld are indicated: lunate [LS]. superior
temporal [STS), and mtraparieal [IP). The lesion a located along both banks of the STS. Scale bar, 1 mm.
tions of 1.5 Ml (total, 2205 Mg) were made in the left
and right hemispheres, and in monkey 3, 96 and 112
injections of 1.5 MI (total, 3120 tig). Due to the length
of the injection procedure, the lesions in each hemi-
sphere were made in separate surgeries a week apart,
during which time visual thresholds were measured
for monkeys 1 and 3 to determine the effect of the
surgery itself (plus the unilateral lesion) on thresh-
olds.
Histology
At the conclusion of behavioral testing, each animal
was killed with an overdose of barbiturate and per-
fused with a phosphate buffer rinse followed by 4%
paraformaldehyde fixative. A second rinse was used
to removed excess fixative. The brain was removed,
blocked, and sectioned at a thickness of 32 or 40 pm.
Triplets of adjacent sections at intervals of 0.25 or 0.50
mm were stained for cytochrome oxidase (Wong-Rlley,
1979), with cresyl violet, and for myelinated fibers
(Gallyas, 1979).
Results
Lesion Reconstruction
A representative histological section through one le-
sion is shown in Figure 1. Lesions usually produced
a very uniform loss of neurons with an abrupt tran-
2(0 Effects of MT/MST Lesions on Motion Thresholds • PasternaV and Merigan
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sition to normal-appearing neuropil at the lesion edg-
es as shown in Figure 1. This lesion was complete
along both anterior and posterior banks of the STS,
except for a small region in which deeper layers of
cortex survived. In instances when the lesion termi-
nated with a sulcus, we often found a small region of
damage to only superficial layers bordering the full
thickness lesion.
The location and extent of complete and partial
lesions are illustrated on flattened 2-D maps of the
cortical surface in Figure 2. Lesions were reconstruct-
ed primarily from the cytochrome oxidase (CO) sec-
tions, which clearly showed the extent of neuron loss.
Comparison with cresyl violet and myelin stains
showed thinning of the cortical surface at lesion sites,
with substantial gliosis but no apparent damage to
fibers. The lesions are shown, as in previous studies
(e.g., Newsome and Pare, 1988), in relation to the
borders and fundus of the STS. Also shown is the
secondary fundus of the STS, which is a reliable in-
dicator of the location of lateral (central) MT (Van
Essen et al., 1981). Small regions along the lateral
border of the lesions in monkeys 1 and 2 showed the
myeloarchitecture that is characteristic of area MT in
unlesioned monkeys (Van Essen et al., 1981; Unger-
leider and Desimone, 1986), and these areas are shown
by solid lines in Figure 2.
Lesions in all three monkeys began near the medial
end of the STS but had a variable anterior-posterior Monkey
extent. The lesion in monkey 1 appears to have dam- 1
aged all of dorsal MT and MST, but to have partially
spared approximately one-fourth of ventral MT (cor-
responding to approximately the central 4° of the vi-
sual 6eld), as well as a portion of ventral MST. As
indicated by the shading, damage to superficial layers
extended to the estimated border of MT. The lesion
in monkey 2 again damaged all of dorsal MT and MST,
and extended slightly farther ventral than the lesion
in monkey 1, as can be seen from the length of the
secondary fundus of the STS (dotted line to the left
of the primary fundus of the STS) that was spared. A
small portion of the most ventral edge of MT was
partially spared (deeper layers) on both sides in this

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monkey, as well as part of ventral MST. This monkey
also had small lesions in the left hemisphere near the
anectant gyrus and in the interparietal sulcus. In mon-
key 3, the lesions in both hemispheres appear to cover
all of MT and MST, as well as a substantial part of FST.
There are also small lesions near the anectant gyrus
and in ventral cortex in the inferior occipital sulcus.
Thus, all three lesions involved large and slightly dif-
ferent portions of MT and MST.

Contrast Sensitivity: Detection and

Direction Discrimination
Preoperative (open circles) and stable postoperative
(solid circles and triangles) contrast sensitivity for
drifting gratings as a function of temporal frequency
is shown in Figure 3. Data collected shortly after the
lesion ("early") are shown by dashed-dotted lines
and will be described at the end of Results. The stable
postoperative data were collected during a period of
about 4 weeks, several months after the lesion. Mon-
keys 1 and 2 showed no permanent deficit on either
task. The slightly higher postlesion sensitivity for some
conditions was most likely due to a gradual increase
in sensitivity during the course of testing. Monkey 3
showed a small sensitivity loss (less than twofold) for
detection, and a somewhat larger loss for direction
discrimination at all temporal frequencies.
In addition to the measures for drifting gratings
shown here, in monkey 3 we also measured contrast
sensitivity for a 9 Hz counterphasemodulated grating
of 1 cycle/degree spatial frequency. For this stimulus,
postlesion contrast sensitivity remained 0.28 log units
below that for a drifting grating. A similar greater
sensitivity for drifting than flickering gratings is com-
monly found in normal observers (Levinson and Sek-
uler, 1975; Pasternak, 1986).
Speed Discrimination
Pre- and postoperative speed discrimination are shown
as Weber fractions (speed increment/base speed) in
Figure 4. Preoperatively, the monkeys could discrim-
inate 5-10% differences in speed over a broad range
of grating contrasts. Their performance was less ac-
curate only at the lowest grating contrast (1.5%). Mon-
key 1, which had the most substantial sparing of por-
tions of MT and MST, showed no permanent elevation
of speed threshold after the lesions, while the thresh-
Figure 2. Reconstnjctkm of lesions on a flattened representatm of visual cortex
(Van Essen and Maunseil, 1980). an approach that allows 3 f l information about
cortex, obtained from histtriogcat seaiore. to be displayed on a 2-0 map. The senrnds
shown to the left of each conical map shows the location of the foveal portion of
V I . which contacts the rest of cortex at atong the borders, when are shown separated
m this figure only IH*TITP of Qeometncal constrains. The rtoht hemisphere is shown
as h would appear in the brain, with righi bang approximately anterior. The left
hemuphere is shown mirror reversed for ease of comparison. Dotted lines show the
location of the fundus of suta, and three of these, the inferior occtprtal [IDS), the
superior temporal [STS), and the tunate [LS], are labeled. The secondary fundus of
the STS is slightly posterior (left) of the primary fundus, and its ventral end can be
seen in monkeys 1 and 2. Area MT a located on the posterior bank of the STS, with
its center close to the dorsal limit of the secondary sulcus of the STS. Regions with
the characteristic rrryeioarchitecture of MT that extended beyond complete lesions were
present only in monkeys 1 and 2 and are shown by a thm fm. Regions in which aO
layers of cortex were damaged are shown in A t a t white areas with the loss of only
superficial cortical layers are shown by a tnken tkK, The plane ol section was
horoontal for rmnkeys 1 and 2 and sagittal for monkey 3.
olds of the two monkeys with more extensive damage
to these areas were elevated about two- to fourfold,
although thresholds remained independent of con-
trast. While these two monkeys showed a reliable el-
evation of thresholds, they were still able to discrim-
inate differences in speed of less than 40%.
In these experiments variation in stimulus speed
is confounded with variation in temporal frequency,
making it unclear which stimulus feature was dis-
criminated. To assess the possibility that monkey 1
discriminated temporal frequency rather than speed,
we compared the above discrimination to the dis-

Cerebral C o n e i May/June 1994, V 4 N 3 Ml

 Detection Direction Discrimination
:
1 c/deg j ^ Jarion 1 c/deg
300 300

100 100

30 ^ 'early 30

10 *»" 10 .^early

3 3
Monkey 1 •*•
Monkey 1

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300 . lesion 300 lesion
•—"?
C/3 100 • intact 100
^ ^
early >j
a 30 30 A early
00
.4—>
10 • 10
8C 3 3
Monkey 2 Monkey 2
O
u
300
300

100 100

30 30

10 • 10
-early /'
3 3
Monkey 3 Monkey 3
10 20 10 20

Temporal Frequency (Hz)

Figure 3. Effects of MT/MST lestons on contrast sensitivity functions for detection [left) and direction discrimination [right). Preoperative (moo. open symbols) and postoperative
(teen) pufoimance is shown as a function of temporal frequency (drift rate) for the three monkeys. Thresholds measured 3-6 months after the bilateral lesions are shown with
solid syntols and dashed fines. Each date point e based on at least three threshold determinations. Error bars are ± 1 SEM. Where not w i d e , SEMs are smaSer than symbols.
Open tnatfes indicate single thresholds measured after the l i t e r a l injection of iboienit add. DaslHloned tines show the performance measured shortly (3-7 d) after the lesion
was placed.

crimination of temporal frequency using a counter-
phase flickering grating of the same spatial and tem-
poral frequency. These results are shown in Figure 5.
Both before and after the lesion this monkey was less
accurate in discriminating differences in flicker rate
than differences in speed. Thus, it is unlikely that the
speed discrimination was performed on the basis of
apparent flicker rate.
Spatial Frequency Discrimination
Thresholds for discriminating differences in spatial
frequency were examined with stationary gratings in
monkey 1 to determine if deficits resulting from MT/
MST lesions were limited to motion thresholds. Re-
sults in Figure 6 show that the lesion had no reliable
effect on spatial frequency discrimination at either of
the tested contrasts. Thresholds before and after the
lesion were on the order of 6% irrespective of grating
contrast. Thus, although this monkey showed a three-
fold loss in accuracy in the discrimination of small
differences in direction (see below), it showed no
deficit in the discrimination of spatial frequency dif-
ferences.
Dynamic Random Dots: Discrimination of
Opposite Directions
Preoperative and postoperative range thresholds, that
is, the broadest range of directions in a dot display

262 Effects of MT/MST Lesions on Motion Thresholds • Pasternak and Merigan

 Monkey 1
0.6
lc/dcg Intact Lesion
2.24 Hz
8"
3- 03
0.4
1"
I"
0.2 10 50 2 10 50

Grating Contrast (%)

intact R g u r a 6 . A companion of difference thresholds for speed and temporal frequency

Monkey 1 for monkey 1. Speed dierence thresholds were measured relative to a 1 cycle/degree
grating dnfting at 2.24 H2 (2.24 degrees/sec), whde temporal frequency difference
thresholds were measured in comparison wnh a 1 cycle/degree grating flickering in

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> U.8
counterphase at 2 24 H L Prenperatrve performance [ m a . open symbols) a shown
on the left while postoperative performance {lesion, solid symbols) is shown on the
<
right Both before and after the taston the monkey was less accurate m discnmnating

II 0.6

leskm
tempers! frequency differences than in discriminating differences m speed.

0.4
discriminate motion direction in the presence of sub-
Q stantial stimulus noise. However, in the absence of
0.2 noise (larger motion signal), the residual perfor-
"8 Monkey 2
mance on the direction task was only slightly below
normal, suggesting that the ability to integrate local
CO
motion signals was largely preserved after the lesion.
0.6 r

Dynamic Random Dots: Discrimination of

Small Differences in Direction
0.4 Direction difference thresholds were measured at two
stimulus speeds, 5 and 15 degrees/sec, as a function
of the range of directions in the stimulus. Preopera-
tively, the accuracy of direction discrimination de-
0.2 creased with an increasing range of directions, but
increased with stimulus speed. For example, at 5 de-
intact
Monkey 3
1 10 50
Spatial Frequency Discrimination

Contrast (%) 0.08 I c/deg

C stationary
R g u r a 4 . Effects of MT/MST lesions on speed discriminations. Preoperative [open (/a
symbols) and postoperative [solid symbols) speed discnmmatcn thresholds are shown 2
as a function of paling contrast It? the three monkeys. Ossh-dotted lines for monkeys
1 and 3 and a solid trimgfe for monkey 2 show the performance measured 7 or 30 -O 0.06
d after the bilateral lesion. o
VI

2 0.04
for which opposite directions of global motion could
s:
be discriminated, are shown in Figure 7. When all the 0)
dots in the distribution were directionally biased c
(100% signal), range thresholds reached 330-340°. 0.02
The thresholds decreased with percentage of signal
a
.0
and the monkeys were unable to perform the task
when the motion signal dropped below about 5%
(i.e., over 95% of dots moved in random directions).
The postoperative performance of all monkeys was
affected by the lesion, and the effect was greatest at
Grating Contrast (%)
low motion signal. While this effect was most pro- 6. The effect of MT/MST lesions on spatial frequency (fiscrrminatjon. Pre-
nounced in monkeys 2 and 3, there was a small deficit operative [light tars, BUsct) and postoperative (flEsDr bffs, lesion) difference thresholds
are shown for the spatial frequency of a stationary grating. Base spatial frequency
even in monkey 1, which may have had partial sparing was 1 cycle/degree. Each 4 M point represents at least three threshold detaminaiions.
of the central representation of MT. Thus, lesions of Error bars are ± 1 SEM. The thresholds were measured at two contrail revets. 5 1 %
areas MT and MST permanently affected the ability to and 1256-

Cerebral Cortei May/June 1994, V 4 N 3

 360 ' 5 deg/sec 5 deg/sec

270
V
intactyj^
180 00
• early u
2-
90 / Aesion .
o Monkey 1 s>e 15 deg/sec

0 d I L 90

Monkey 1
10
360

Downloaded from http://cercor.oxfordjournals.org/ at University of Bath Library & Learning Centre on July 7, 2015
270 Monkey 2 Monkey 2
intact

180

90 10

0
Monkey 2 Monkey 3 Monkey 3

90 180 270 90 180 270

360 Direction Range (deg)

intact
270
180
90 /'early
0
Monkey 3
10 20 50 100
Motion Signal (%)
Rgnra 7 . Effects of MT/MST lesons on the abihy to integrate local directional
signals. Preopsratrve [imsa, open symbols) and postoperative [lesion, solid symbols)
range thresholds as a function of motion signal are shown for the three monkeys. The
data labeled lesion were collected 3-6 months after placing the brtatera) STS lesion
8Rd are means of ai teas three threshold determinations. Open trusses m the data
for monkeys 1 and 3 show single thresholds measured within days of placement of
the unilateral lesion. The data shown by the doutashed a m (labeled esrty] are
thresholds measured within the first 7 d (monkey 2). 14 d (monkey 1), and 45 d
(monkey 3) following the lesion. Each data poim on these curves represents a single
threshold determination. Speed, 5 degrees/sec (Ar = 30 msec to •=• 0.151; task,
discrimination between rignrward and leftward motion (see inset). Prims at 0° range
were measured by varying the proportion of randomly moving dots while keeping the
"s^jnaJ" dots at 0° range.
grees/sec monkey 2 discriminated at 10° difference
in direction, while at 15 degrees/sec, thresholds im-
proved to about 5° (see Fig. 8). For all monkeys le-
sions of MT/MST permanently reduced the precision
with which they judged target direction by about a
Direction difference thresholds as a ftnction of the range of directors in
the stimulus. The direction range ts 0° when all dots in the display move in parallel
in a smgte direction. The direction range is 180° when the dens are allowed to move
in directions ± 9 0 ° from rightward motion. Error bars are ± 1 SEM. Each data poim
includes at least three threshold deterrrinauons. The data labeled exiy for monkey 1
and connected by a dohdashed line are thresholds measured 14 d after the bilateral
lesions were placed. The open triangle indicates the direction threshold measured
within days of placement of the unilateral lesion. Speeds. 5 degrees/sec (Ar - 30
msec; to - 0.15°) or 15 degrees/sec (Ar = 15 msec, to - 0225°); 100%
signal, task, discrimination of small direction differences (see inset).
factor of 2-3, and this effect appeared to be largely
independent of the range of directions in the target,
except for monkey 1, which showed no deficit for the
stimulus containing the broadest range of directions
(270°).
The above data were obtained with stimuli in which
dot displacement was 0.15°. Since the maximal dis-
placement that allows the perception of motion, Dnal,
has been used as a measure of the spatial scale of
motion mechanisms (Cleary and Braddick, 1990; Mor-
gan, 1992), we examined the effect of the size of
spatial displacement on motion thresholds in two
monkeys (Fig. 9). The temporal interval between dot
displacements was adjusted to maintain a constant
speed of 5 degrees/sec. Before the lesion, direction
thresholds were greater for larger step sizes, and for
steps greater than 0.6-0.8° the monkey was unable to
perform the task. After the lesion, the variation of
range threshold with step size was less pronounced,
resulting in relatively normal performance at larger
step sizes. The maximal step size for which a direction
of 90° could be discriminated was 0.67-0.72°, which
was at least as large as the prelesion maximum.

IU Effects of MT/MST Lesions on Motion Thresholds • Pasternak and Merigan

Transient Effects
Dot-dashed lines in all of the above figures show
thresholds measured relatively shortly after the le-
sions. These data do not have the same reliability as
the stable postlesion data, since the experiments were
not designed to examine short-term effects, and be-
cause early threshold measures were usually not re-
peatedly tested until they were stable. Moreover, the
early threshold measures had to be limited to a small
number of tests, and they were not performed in all
three animals. Nonetheless, these data, especially the
more complete early data available for monkey 1, give
a clear impression of dramatic disruptions in thresh-
olds shortly after the lesions, with a gradual recovery
of at least partial prelesion sensitivity.
Contrast Sensitivity
In all three animals, measurements of sensitivity for
detection and direction discrimination revealed some
transient depression of sensitivity. The most complete
data were obtained from monkey 1, for whom we were
able to measure complete functions for detection and
discrimination during the first 6 d after the lesions by
making determinations of detection and direction
thresholds each day. Loss was more pronounced for
direction discrimination than detection, suggesting
that these early transient effects were specific either
to motion mechanisms or to the greater demands of
discrimination than detection tests. Interestingly, the
pronounced effects at lower temporal frequencies de-
creased within 3 d, with an eventual return to normal
preoperative levels. A pronounced early loss in di-
rection contrast sensitivity was also seen in monkey
3, and again the deficit was most marked for slowly
moving gratings. The loss for monkey 3 at 7 d was
greater than that in monkey 1 at 6 d, and this differ-
ence was consistent with the final thresholds, which
showed loss only for monkey 3.
This early sensitivity loss was unlikely to be a sim-
ple consequence of surgical trauma, since for two
monkeys (monkeys 1 and 3) we tested sensitivity dur-
ing the week between placement of the two unilateral
lesions, and found little, if any, loss in contrast thresh-
olds (see open triangles in Fig. 3). Thus, even though
the animals had received a unilateral MT/MST lesion,
there was little effect.
Dynamic Random Dot Tests
Transient losses on tests involving discrimination of
opposite directions usually exceeded permanent def-
icits by about a factor of 2, with the most consistent
early effects across monkeys seen on range thresholds.
Monkeys 2 and 3 were unable to discriminate direc-
tion above chance levels even 4-6 d after the lesions.
Early data, collected during the first few weeks fol-
lowing the lesion, show a dramatic drop in range
thresholds in all monkeys even when all the dots in
the display were directionally biased (100% signal).
Before the lesion both monkeys could discriminate
direction when the breadth of the direction distri-
bution was about 330°. However, after the lesions,
range thresholds in monkeys 2 and 3 decreased to
-a
5deg/sec
1 *>
30
8
I
10
S
§
•a
Monkey 1
0.2 0.4
,§©,
0.6 0.8 1
Monkey 3
0.2 0.4 0.6 0.8
Q Step Size (deg)
Figure 9. Pro- snd postoperative direction difference thresholds as a function of the
A x (step sizel of individual dots for two of the monkeys. The direction thresholds
Downloaded from http://cercor.oxfordjournals.org/ at University of Bath Library & Learning Centre on July 7, 2015
were measured at 5 degrees/sec, whrii was kept constam by adjusting the temporal
nterval. The largest step sue on each graph is the maximal spatial displacement
( 0 _ ) , when was determined by varying the step see (and temporal interval) end
asking the monkeys to discnrraraie between nghtward and downward moving stimuli
|90° difference m direction) The earr> postoperative data were collected 14 d after
the bilateral festoa Direction range, &. 100% signal.
less than 100°. The deficit in monkey 1 was less pro-
nounced, with range thresholds reduced to about 270°.
With lower-percentage signal, early losses were even
more pronounced, and all monkeys were unable to
perform the task with less than 20% motion signal.
With repeated testing (or the passage of time), per-
formance at high levels of motion signal (50% and
100% signal) improved nearly to preoperative levels,
although performance with low motion signal re-
mained poor.
Data obtained in the interval between the first and
second lesions in monkeys 1 and 3 (Fig. 7, open tri-
angles) indicate that even unilateral lesions substan-
tially affected these thresholds. In both monkeys the
effect of the unilateral lesion was comparable to the
transient effect of the complete lesions. Since the uni-
lateral lesion caused little or no effect on contrast
sensitivity for detection and direction discrimination
in these same animals (open triangles in Fig. 3), these
deficits in global motion thresholds could reflect ei-
ther a greater sensitivity of these measures to simple
surgical trauma or a dependence of these thresholds
on a contribution from both visual fields.
Discussion
Our results show that large lesions of areas MT and
MST produce permanent deficits in some tasks in-
volving motion discriminations. These effects includ-
ed a reduction in the accuracy of speed and direction
discrimination, and reduced ability to integrate local
directional signals in the presence of directional noise.
However, the most striking feature of the present re-
sults was not the observed deficits, but rather the sub-
stantial preservation of motion discrimination. It was
particularly impressive that the lesioned monkeys dis-
criminated direction of grating motion at essentially
contrast threshold, and when stimuli contained no
directional noise, they showed near-normal perfor-
mance on discriminations that require the integration
of local motion vectors.
Cerebral Cortex May/June 1994, V 4 N 3 2W
Lesion Location and Extent
Interpretation of the lesion effects requires both a
knowledge of which visual areas were damaged and
which portions of the visual field were affected in each
of the involved areas. These issues are particularly
difficult for the conical areas studied in this experi-
ment. While area MT is retinotopically organized, its
relatively large receptive fields make it difficult to
interpret partial lesions. Area MST shows only mini-
mal retinotopy and it has very large receptive fields
(Saito et al., 1986), making interpretation of partial
lesions even more difficult. In this study we attempted
to make complete lesions of areas MT and MST, but
lesion reconstruction indicates sparing of deep but
not superficial layers of a portion of central MT in
monkeys 1 and 2. On the other hand, the lesion in
monkey 3 appeared complete, and the pattern of re-
sults was qualitatively similar in all three monkeys.
This similarity suggests that the regions of central MT
in which there was sparing of only deep layers in
monkeys 1 and 2 may not have been functional. Al-
ternatively, these regions of central MT may not have
been sufficient to prevent permanent loss of some
motion sensitivity. In any case, it does not appear
possible that the substantial survival of motion per-
ception seen in this study was due to undamaged
regions of MT or MST.
Detection and Direction Contrast Sensitivity
Intact sensitivity for detecting stationary and drifting
gratings has previously been reported for monkeys
after MT lesions (Newsome and Pare, 1988), and hu-
mans with damage to parieto-occipital cortex (Hess
et al., 1989; Plant et al., 1990). A similar preservation
of contrast sensitivity for detection has also been ob-
served in cats with lesions of the lateral suprasylvian
cortex (Pasternak et al., 1989), an area often compared
to area MT in the monkey (Payne, 1993). In these
studies, observers detected gratings at contrast levels
close to, or only slightly above, normal thresholds. In
the present study, monkeys 1 and 2 showed no per-
manent loss of sensitivity, while monkey 3 showed a
small but persistent decrease in sensitivity for both
thresholds. The small depression in sensitivity in
monkey 3 may have been due to the larger extent of
its lesion, which appeared to include all of MT and
MST as well as a portion of area FST. This decreased
sensitivity for drifting gratings was accompanied by a
parallel decrease in sensitivity for counterphase-mod-
ulated gratings flickered in counterphase, such that
the nearly twofold greater sensitivity for drifting grat-
ings was maintained after the lesion. Superior sensi-
tivity for drifting gratings suggests, as does other ev-
idence discussed below, that grating detection at
contrast threshold was mediated by directionally se-
lective mechanisms (Watson et al., 1980; Pasternak,
1986).
The largely preserved sensitivity for the direction
of image motion is similar to that found by Plant et
al. (1990) in a human patient with extrastriate lesions,
who was otherwise severely deficient in discriminat-
ing speed differences. It is also consistent with the
158 Effects of MT/MST Lesions on Motion Thresholds • Pasternak and Merigan
results of Pasternak et al. (1989), who measured con-
trast sensitivity for detection and direction discrimi-
nation in cats after ibotenic acid lesions of a cortical
area possibly involved in motion analysis, the lateral
suprasylvian (LS) cortex. As in these monkeys, no loss
of sensitivity was found for either detection or direc-
tion discrimination. They concluded that direction
signals present in undamaged cortical areas may me-
diate this behavior.
A more severe deficit in contrast sensitivity for di-
rection discrimination was found in a human patient
with an occipito-parietal cortical lesion (Hess et al.,
1989). This loss in sensitivity was more reminiscent
of that found in cats developmentally deprived of mo-
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tion perception, which have a widespread loss of cor-
tical directionality (Pasternak et al., 1985; Pasternak
and Leinen, 1986), than in cats with LS lesions or
monkeys with lesions of MT and MST. One implica-
tion of this analysis is that the human patient of Hess
et al. (1989) probably has a more widespread lesion
than either the cats with LS lesions or the monkeys
with MT/MST lesions.
Speed Discrimination
Two of the monkeys showed consistent and perma-
nent two- to fourfold losses in the accuracy of speed
discrimination over a wide range of contrasts. How-
ever, postoperative performance remained indepen-
dent of stimulus contrast, suggesting that, at least in
some respects, mechanisms underlying the perfor-
mance retained their preoperative properties.
Since differences in grating speed are accompa-
nied by differences in temporal frequency, it was con-
ceivable that the residual performance on speed dis-
crimination was based on flicker cues, rather than on
speed per se. If this were the case, we would expect
that the discrimination of flicker differences would
be at least as accurate as the discrimination of speeds.
However, previous reports (McKee et al., 1986; Pas-
ternak, 1987), and the data from one of the monkeys
in the present study, show that discrimination thresh-
olds for temporal frequency are about a factor of 2-3
higher then speed thresholds. Moreover, it has been
demonstrated that the discrimination of flicker rates
cannot be performed in the absence of directionally
selective velocity mechanisms (Pasternak, 1987). Thus,
the residual performance of the monkeys appears to
be based on differences in target speed.
Although the lesions produced some loss in the
accuracy of speed discrimination, the monkeys re-
tained substantial ability to judge speed differences.
Preservation of these speed discrimination abilities
after destruction of areas MT/MST, suggests that neu-
rons outside the STS can perform some analysis of
stimulus speed, and that MT neurons do not make a
unique contribution to the analysis of speed.
Our results on speed discrimination address the
issue of a special role for MT in the perception of
velocity. The first evidence that MT might be involved
in the processing of target velocity came from the
study of Newsome et al. (1985), who showed that
ibotenic acid lesions of MT produced transient defi-
cits in smooth pursuit eye movements, a result that
was interpreted as reflecting a loss of velocity infor-
mation. Effects of MST lesions on smooth pursuit were
similar in that the monkey was unable to match eye
speed to target speed (Dursteller and Wurtz, 1988).
Subsequently, deficits in speed discrimination (like
those shown here) were reported in patients with
lesions of posterior occipitoparietal cortex (Hess et
al., 1989; Vaina, 1989; Plant et al., 1990). The impli-
cations of all of these findings remain somewhat un-
clear. First, the eye movement deficits of Newsome
and colleagues and Dursteler et al. appeared more
transient than the speed deficits measured in this study.
This difference could be due in pan to the variation
in lesion size and location in the different studies.
Comparison of speed difference effects in the present
macaque and previous human studies is even more
difficult given the uncertain extent to the lesions in
studies of humans, and the unclear relationship of
macaque MT to human occipitoparietal cortex.
Global Motion
To assess the role of MT in the integration of local
motion signals, we used dynamic random dots. MT
neurons have several properties that make them par-
ticularly well suited for integrating dot motion, in-
cluding their large, directionally selective receptive
fields and the directionally integrative response of
many of its neurons to multiple directions of motion
(Movshon et al., 1985). Like Newsome and Pare
(1988), we found that the integration of local direc-
tional signals (the range threshold for dot motion)
was transiently disrupted by lesions. A similar inabil-
ity to perform motion tasks in the presence of noise
has been reported for a "motion blind" patient by
Baker et al. (1991). Permanent range threshold def-
icits at low levels of motion signal suggest either that
these areas play a particularly important role in some
processing that enhances signal-to-noise ratios (e.g.,
summation) or simply that lesions may exacerbate
noise levels in motion-sensitive areas. The possibility
that MT neurons may play some role in extracting
motion signals from noise is supported by studies of
Britten and Newsome (Newsome et al., 1989; Britten
et al., 1992), who have recently shown that cells in
MT respond to the direction of motion in noisy targets
at signal-to-noise levels near psychophysical thresh-
old. Moreover, recent microstimulation experiments
(Salzman et al., 1992) have shown that it is possible
to influence the monkey's ability to extract correlated
motion by stimulating individual neurons in MT.
In addition to deficits in the discrimination of op-
posite directions for dots and gratings, deficits in the
discrimination of small direction differences were
found in all three monkeys. Since such discrimina-
tions are likely based on a comparison of the response
of local motion detectors, involving a pooling of re-
sponse differences across directions (Wilson and Gelb,
1984; Watamaniuk et al., 1989), the direction deficit
could result from disruption of the response of local
directional detectors or the pooling of directional out-
puts. It is unlikely that the MT/MST lesion affected
directionally selective neurons in areas most likely
involved in local analysis of direction (e.g., VI). In-
deed, our results showed very limited effects of MT/
MST lesions on contrast sensitivity for direction, a
function that may reflect local directionally selective
responses (Pasternak and Maunsell, 1992). It seems
more likely that the pooling of local directional re-
sponses was affected by the lesion, although the near-
ly intact range thresholds we found in the absence of
directional noise suggest that at least some aspects of
pooling remained intact after MT/MST lesions. Com-
putational approaches to directional difference
thresholds suggest shallower directional tuning func-
tions or increased variability as likely causes of in-
Downloaded from http://cercor.oxfordjournals.org/ at University of Bath Library & Learning Centre on July 7, 2015
creased direction thresholds (Wilson and Gelb, 1984).
These considerations would be consistent with either
an alteration in these properties in neurons mediating
direction thresholds, or assumption of these functions
by other neurons with less optimal directional tuning
and variability.
Preoperative values of D m measured in this study
(0.7-0.8° for a target moving at 5 degrees/sec) are
close to the upper limit for dot displacement in di-
rectionally selective neurons in area MT (Mikami et
al., 1986). Likewise, psychophysical measures of D^
as a function of eccentricity closely match the maxi-
mal spatial displacement for MT neurons at different
eccentricities (Baker and Braddick, 1985). These par-
allels between the properties of MT neurons and the
psychophysical performance of humans have sug-
gested that receptive fields of MT neurons could be
the substrate of the maximal spatial displacement for
direction discrimination. In this study, postoperative-
ly reduced accuracy of directional judgments was less
pronounced at larger spatial displacements, and there
was no detectable deficit at all at the largest step sizes.
In fact, the maximal spatial displacement (£>„„) for
direction discrimination was somewhat larger after
the lesion. This increase in £>„„ suggests a postop-
erative increase in the spatial scale of mechanisms
mediating the perception of direction (Baker and
Braddick, 1985).
Specificity of the Motion Deficit
All three monkeys showed initial and often perma-
nent deficits on a variety of discriminations of the
speed and direction of moving targets. On the other
hand, we found almost no deficits on the detection
task, which did not require the use of motion cues.
It is possible that this difference simply reflects a
greater vulnerability of discrimination than detection
tasks, although this explanation is not supported by
the intact ability of monkey 1 on a nonmotion dis-
crimination: the spatial frequency of stationary grat-
ings. Thus, it seems likely that the effects of MT/MST
lesions were probably limited to the domain of image
motion. On the other hand, the effect of the MT/MST
lesion on the spatial frequency discrimination was
examined in only a single monkey. Since the lesion
in monkey 1 was less extensive and the accompanying
deficit in motion thresholds was less pronounced than
that in the other two monkeys, the question of the
Cerebral Cortex May/June 1994, V 4 N 3 M7
specificity of MT/MST lesion effects needs to be fur-
ther explored.
7*e Meaning of Transient Deficits
The nature of transient effects is of great importance
in understanding the contribution of MT and MST to
motion perception, since it is not clear from previous
studies whether the more profound earlier effects of
MT lesions are qualitatively different from permanent
effects. Unfortunately, it is impossible to study the
detailed profile of MT effects at the same time as the
detailed time course of transient effects, since such
effects often last only a day or two. Thus, the present
study was designed primarily to examine permanent
effects of MT/MST lesions on a large number of visual
functions, to set the stage for subsequent studies of
the detailed time course of interesting effects.
However, some early effects of lesions were ex-
amined in this study, and as has been previously ob-
served, they were more profound than permanent ef-
fects. At present we can only speculate about the
relation of transient and permanent effects of MT/
MST lesions to the functional role of these areas. One
possibility is that these areas alone are central to mo-
tion processing in the normal monkey and that rapid
recovery simply reflects the postlesion (plastic) as-
sumption of motion processing by other cortical areas
(Newsome and Pare, 1988). A second possibility is
that motion processing normally involves a wide net-
work of many cortical areas, and that recovery time
represents an adaptation of those not damaged. Fi-
nally, it could be that many motion perception ca-
pacities are not critically dependent on MT or MST,
and that measured disruption simply represents an
increase in noise throughout all cortical areas. Clearer
evidence that deficits were highly specific to motion
processing would argue against this possibility (see
above). These issues may best be approached by a
careful comparison of the time course and nature of
both neural and behavioral disruptions.
Conclusions
Permanent deficits in the detection of motion in the
presence of directional noise and the discrimination
of small differences in direction and speed provide
strong support for a role of areas MT/MST in motion
processing. On the other hand, since many aspects
of motion perception survive damage to MT/MST, it
is likely that intact areas outside ventral STS, perhaps
at lower cortical levels, can perform many of the op-
erations required by the motion tasks of the present
study. It remains to be seen whether cortical reor-
ganization following the lesion contributes to the dis-
tribution of these capacities.
Notes
This work was supported by grants from the National Eye
Institute, EY06175, EY05911, and EY01319 (Core Grant to
the Center for Visual Science). John Maunsell participated
in early stages of the experiment We thank Dawn Ferris,
Walter Polashenski, Judy Tompkins, and Peter Vamvakias for
excellent technical assistance, and Emil Rainero for devel-
258 Effects of MT/MST Lesions on Motion Thresholds • Pasternak and Merigan
oping testing software. We also thank William Newsome and
John Maunsell for comments on the manuscript.
Correspondence should be addressed to Tatiana Paster-
nak, Center for Visual Science, University of Rochester,
Rochester, NY 14627.
References
Albright TD (1984) Direction and orientation selectivity
of neurons in visual area MT of the macaque. J Neurosci
42:1106-1130.
Baker CL.BraddickOJ (1985) Eccentricity-dependent scal-
ing of the limits for short range motion perception. Vision
Res 25:803-812.
Britten KH, Shadlen MN, Newsome WT, MovshonJA (1992)
The analysis of visual motion: a comparison of neuronal
and psychophysical performance. J Neurosci 12:4745-
4765.
Downloaded from http://cercor.oxfordjournals.org/ at University of Bath Library & Learning Centre on July 7, 2015
Cleary R, Braddick OJ (1990) Direction discrimination for
band-pass filtered random dot kinematograms. Vision Res
30:303-316.
Duffy C.WurtzR (1991) Sensitivity of MST neurons to optic
flow stimuli. I. A continuum of response selectivity to
large-field stimuli. J Neurophysiol 65:1329-1345.
Dursteller MR, Wurtz RH (1988) Pursuit and optokinetic
deficits following lesions of cortical areas MT and MST.
60:940-965.
Gallyas F (1979) Silver staining of myelin by means of
physical development. Neurol Res 1:203-209.
Hess RF, Baker CL, Zihl J (1989) The "motion-blind" pa-
tient: low-level spatial and temporal filters J Neurosci
9:1628-1640.
Komatsu H, Wurtz RH (1988) Relation of cortical areas MT
and MST to pursuit eye movements. I. Localization and
visual properties of neurons. J Neurophysiol 60:580-603.
Levinson E, Sekuler R (1975) The independence of chan-
nels in human vision selective for direction of movement.
J Physiol (Lond) 250:347-366.
Maunsell JHR, Van Essen DC (1983) Functional properties
of neurons in the middle temporal area of the macaque
monkey. I. Selectivity for stimulus direction, speed and
orientations. J Neurophysiol 49:1148-1167.
McKee S, Nakayama K, Silverman GH (1986) Precise ve-
locity discrimination despite variations in temporal fre-
quency and contrast. Vision Res 26:609-619.
Mikami A, Newsome WT, Wurtz RH (1986) Motion selec-
tivity in macaque visual cortex- spatio-temporal range of
directional Interactions in MTandVl.J Neurophysiol 55.
1328-1339.
Morgan MJ (1992) Spatial filtering precedes motion de-
tection. Nature 355:344-346.
Movshon JA, Adelson EH, Gizzi MS, Newsome WT (1985)
The analysis of moving visual patterns. In: Pattern rec-
ognition mechanisms (Chagas C, Gattas R, Gross CG,
eds), pp 117-151. Vatican City: Ponticifica Academia
Scientiarum.
Newsome WT, Pare EB (1988) A selective impairment of
motion perception following lesions of the middle tem-
poral visual area (MT). J Neurosci 8:2201-2211.
Newsome WT, Wurtz RH, Dursteler MR, Mikami A (1985)
Deficits in visual motion processing following Ibotenic
acid lesions of the middle temporal visual area of the
macaque monkey. J Neurosci 5:825-840.
Newsome WT, Britten KH, Movshon JA, Shadlen M (1989)
Single neurons and the perception of visual motion. The
Woodlands, TX- Portfolio.
Pasternak T (1986) The role of cortical directional selec-
tivity in detection of motion and flicker. Vision Res 26:
1187-1194.
Pasternak T (1987) Discrimination of differences in speed
and flicker rate depends on directionally selective mech-
anisms. Vision Res 27:1881-1890
Pasternak T, Leinen LJ (1986) Pattern and motion vision
in cats with selective loss of cortical directional selectiv-
ity. J Neurosci 6:938-945.
Pasternak T, Maunsell JHR (1992) Spatiotemporal sensi-
 tivity following lesions of area 18 in the cat. J Neurosci
12.4521-4529.
Pasternak T, Horn KM, Maunsell JHR (1989) Deficits in
speed discrimination following lesions of the lateral su-
prasylvian cortex in the cat. Visual Neurosci 3:365-375.
Pasternak T, Albano JE, Harvftt D (1990) The role of di-
rectionally selective neurons in the perception of global
motion. J Neurosci 10:3079-3086.
Pasternak T, Schumer RA, Gizzi MS, Movshon JA (1985)
Abolition of cortical directional selectivity impairs visual
behavior in cats. Exp Brain Res 61-214-217.
Payne BR (1993) Evidence for visual cortical area homo-
logs in cat and macaque monkey. Cereb Cortex 3:1-25
Plant G, Laxer KD, Barbaro NM, Nakayama K (1990) Im-
paired speed discrimination restricted to the hemifield
contralateral to an occipitoparietal surgical resection.
Soc Neurosci Abstr 16:5.

Downloaded from http://cercor.oxfordjournals.org/ at University of Bath Library & Learning Centre on July 7, 2015
Rodman HR, Albright TD (1989) Single-unit analysis of
pattern-motion selective properties in the middle tem-
poral visual area (MT). Exp Brain Res 75:53-64.
Saito HA, Yukie M, Tanaka K, Hikosaka K, Fukada Y, Iwai E
(1986) Integration of direction signals of image motion
in the superior temporal sulcus of the macaque monkey.
J Neurosci 6:145-157.
Salzman CD, Mirasugi CM, Britten KH, Newsome WT (1992)
Microstimulation of visual area MT: effects of direction
discrimination performance. J Neurosci 12:2331-2356.
Tanaka K, Sugita Y, Moriya M, Saito HA (1993) Analysis
of object motion in the ventral part of the medial superior
temporal area of the macaque visual cortex. J Neurophy-
siol 69:128-142.
Vaina LM (1989) Selective impairment of visual motion
interpretation following lesions of the right occipito-pa-
rietal area in humans Biol Cybern 61-347-359.
Van Essen D, Maunsell JHR (1980) Two-dimensional maps
of the cerebral cortex. J Comp Neurol 191:255-281.
Van Essen D, Maunsell JHR, BigsbyJL (1981) The middle
temporal visual area in the macaque: myeloarchitecrure,
connections, functional properties, and topographic or-
ganization. J Comp Neurol 199:293-326.
WatamaniukSNJ.SekulerR, Williams DW (1989) Direction
perception in complex dynamic displays, the integration
of direction information. Vision Res 29-47-60.
Watson AB, Thompson PG, Murphy BJ, NachmiasJ (1980)
Summation and discrimination of gratings moving in op-
posite directions. Vision Res 20:341-348.
Williams DW, Sekuler R (1984) Coherent global motion
perception from stochastic local motions. Vision Res 24.
55-62.
Wilson HR, Gelb DJ (1984) Modified line-element theory
for spatial-frequency and width discrimination. J Opt Soc
Am 1:124-131.
Wong-RileyM (1979) Changes in the visual system of mon-
ocularly sutured or enucleated cats demonstrable with
cytochrome oxidase histochemlstry. Brain Res 171:11-
28.
Yamasaki DS, Wurtz RH (1991) Recovery of function after
lesions In the superior temporal sulcus in the monkey.
J Neurophysiol 66:651-673.
Zeki SM (1974) Functional organization of a visual area in
the posterior bank of the superior temporal sulcus of the
rhesus monkey. J Physiol (Lond) 236:549-573.

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