Devel.l)meplml Bra#l Re.search. 25 (1986) 71 81
Elscvicr
71
BRD 5O34O
Effects of Monocular Exposure to Oriented Lines on Monkey Striate Cortex*
MARY CARLSON, DAVID H. HUBEL and TORSTEN N. WIESEL**
Department of Neurobiology. Harvard ~,ledical School. Boston. MA 02115 ( U. 5'. ,4. J
(Accepted August 6th. 1985)
Key aords." monocular exposure - - visual experience - - monkcy striate cortex - - orientation selectivity
This study examines the extent to which the restriction of visual experience to lines of a single orientation influences the organization of the striatc cortex in infant monkeys (Macaca mulatta}. Previous studies of kittens raised with monocular exposure to a single
line orientation have consistently shown the response preference of cells driven by that eye to be biased towards the experienced
orientation. Studies of binocular exposure to restricted orientations have been equivocal. In the infant monkcv cortex responses to
oricntcd lines have virtually all the specificity of responses seen in the adult animal. In an effort to clarity the phenomenon and the
mechanism by which orientation bias might be obtained, we examined the effects of monocular exposure to a restricted orientation in
infant macaques. Three monkeys were used. Each monkey was raised with one open eye exposed to lines of a single orientation and
one cvc occluded by lid suture. As in other cases of monocular deprivation in either cat or monkey, fcw binocularly driven cclls wcrc
recorded and the majority of cells were dominated by the open eye. Cells driven bv the open eye had normal rcprcscntation of all
oricntation preferences and there was no overall increase in the number of cells preferring the orientation to which the eye had been
exposed. The cells dominatcd by the occluded eye, howcver, showed a lack of cells responding to orientations to which thc open cyc
had bccn cxposcd. These tindings suggest that a competitive mechanism operates between the two eyes to providc an orientationselective adx antagc to the open cyc.
INTRODU('TION
er t h e c h a n g e s reflect shifts in o r i e n t a t i o n a l l e g i a n c e
of i n d i v i d u a l cells, o r r e p r e s e n t a m e r e r e d u c t i o n in
T h e o r i g i n a l e x p e r i m e n t s e x a m i n i n g t h e effects of
the selectivity o r r e s p o n s i v e n e s s of s o m e cells, or a
early m o n o c u l a r lid s u t u r e o n t h e a n a t o m y a n d physi-
b i a s i n g of t h e fate of cells t h a t at b i r t h w e r e u n c o m -
ology of t h e cat visual p a t h w a y I~.> h a v e since b e e n
m i t t e d . B e h i n d m o s t of t h e s e e x p e r i m e n t s h a s l u r k e d
e x t e n d e d to i n c l u d e a n u m b e r of s p e c i e s a n d d i f f e r e n t
t h e old c o n t r o v e r s y o v e r t h e d e g r e e to w h i c h cortical
k i n d s of d e p r i v a t i o n s . O n e of t h e m o s t i n t e r e s t i n g
n e u r o n a l c o n n e c t i o n s a r e d e t e r m i n e d as a r e s u l t of
v a r i a t i o n s h a d its o r i g i n w i t h e x p e r i m e n t s b v H i r s c h
e x p e r i e n c e , as o p p o s e d to g e n e t i c s .
a n d Spinelli; a n d B l a k e m o r e a n d C o o p e r I, in w h i c h
C o m p a r e d with t h e cat, t h e r h e s u s m o n k e y at b i r t h
k i t t e n s h a d vision r e s t r i c t e d to s t r i p e s of a single
has a far m o r e m a t u r e visual s y s t e m . Cells in t h e
o r i e n t a t i o n . Since t h e n , t h e s e e x p e r i m e n t s h a v e b e e n
striate c o r t e x s h o w a d e g r e e of r e s p o n s e specificity
repeated
with a w i d e r a n g e o f m o d i f i c a t i o n s a n d
t h a t in m a n y r e s p e c t s ( i n c l u d i n g line o r i e n t a t i o n pref-
widely d i v e r g i n g r e s u l t s , f r o m f a i l u r e to r e p l i c a t e t h e
e r e n c e ) a p p r o a c h e s t h a t of t h e a d u l t a n d t h e s y s t e m
origimtl ( a n d very d r a m a t i c ) f i n d i n g s to c o n f i r m a t i o n
of o r i e n t a t i o n c o l u m n s is a l r e a d y established-'-'. Be-
a n d e x t e n s i o n of t h e m (for r e v i e w s see refs. 5 a n d
c a u s e of t h e s e d i f f e r e n c e s it s e e m e d w o r t h w h i l e to re-
13). T o d a y t h e r e s e e m s to be little d o u b t t h a t by re-
p e a t s o m e of t h e s t r i p e - r e a r i n g e x p e r i m e n t s in m o n -
stricting e a r l y vision to s t r i p e s of a single o r i e n t a t i o n
keys. W e w e r e also m o t i v a t e d by s t u d i e s t h a t s h o w
o n e can c h a n g e t h e r e l a t i v e n u m b e r s o f c o r t i c a l cells
t h a t in m a n e a r l y a s t i g m a t i s m gives rise to a l o w e r e d
t h a t r e s p o n d to t h a t o r i e n t a t i o n . It is less c l e a r w h e t h -
visual acuity r e s t r i c t e d to t h e o r i e n t a t i o n s t h a t w e r e
* These results were first presented at the annual meeting of the Society for Neuroscience in St. Louis, MO, 1978.
** Present address: The Rockefeller University, New York, NY 10021, U.S.A.
Correspondence: M. Carlson. Present address: Washington University School of Medicine, Box g07(I, St. Louis, MO 631 ll), U.S.A.
01~'~5-38116:86/$03.5II© 1986 Elsevier Science Publishers B.V. (Biomedical Division)
2
out of focus 12. In the present experiments we occluded one eye by lid suture and exposed the other
eye to stripes of a single orientation. Monocular closure was used because it is known to produce, in infant monkeys, dramatic physiological and anatomical
changes in eye dominance, even after brief periods of
deprivation 2`~,~s. We assumed that combining lid
closure with orientation deprivation would provide
us with two measures of the effects of the rearing procedures.
MATERIALS AND METHODS
Subjects
Three infant macaques were raised with the left
eye exposed only to lines of a single orientation and
the right eye sutured for the duration of the exposure. Table I presents the age, species, and the duration and orientation of exposure.
Rearing apparatus and procedures
The newborn monkeys, all of normal gestational
age and birth weight, were removed from the mother
the day after birth. Prior to surgery for right eye closure they were housed in ordinary incubators in the
Primate Center nursery unit which was lighted about
12 h/day. After right-eye suture the animals were
transferred to a completely darkened, light-tight
room which contained the rearing apparatus.
The animal was housed in a plastic cage (30.5 x
30.5 x 30.5 cm) with a thermostatically controlled
electric heating pad on the floor. A molded plastic
face mask with eye holes was mounted over a 7.6-cm
opening in the front of the cage. When the monkey's
head was properly positioned inside the mask the entire visual field was filled by a 33.0-cm translucent,
mushroom-shaped glass globe. Two electrodes were
mounted in the brow area of the mask which, when
contacted simultaneously, activated a Kodak carousel projector. The projector, in turn, illuminated the
globe and displayed stripe patterns of a single orientation for the duration of contact. When the animal
remained in contact with the electrodes for a specified duration, it received a drop of milk (SMA human
formula) from a nipple mounted in the mouth area of
the mask. Since the animal was in total darkness except when in contact with the face mask and electrodes, this arrangement assured a fixed orientation
of the monkey's head relative to the stripe pattern
stimulus.
On the first day in the apparatus the animal's head
was placed in the mask and held against the nipple'
and electrodes to assure that it learned the correct
procedure to obtain reinforcement. After repeated
placement in the mask on the first day, the animal
learned the location of the mask and spent increasing
lengths of time with its head in the mask. Over the
next few days it spent increasing lengths of time with
its head in the mask without assistance. During the
first week it received a single drop of milk for each accumulated 2-s electrode contact and exposure to the
stripe pattern. Over the next few weeks a variable
duration schedule was introduced into the programming system, which required different durations
of contact to obtain a single milk delivery (variable
durations of 2 s-2 rain, for an average 30-s duration).
The hours of exposure obtained in the apparatus tor
the 3 animals are shown in Fig. 1. Regular observations of the animal during exposure confirmed that
the head was upright during the projection of the
stripe patterns. Through a fiber optic viewer mounted
in the eye area of the mask, it was possible to verify
that the left eye was open during exposure periods.
The animals were taken directly from the rearing apparatus in a light-tight box to the laboratory for physiological recordings.
The projector contained 80 slides of 4 different
patterns of 6-8 lines which changed automatically
each minute. The lines were composed of strips of
stippled transparencies which diminished in density
TABLE 1
Age and condition of rearing and physiological procedures
Macaque
Age at right
eye suture
Orientation
exposure
Age at onset
of exposure
Duration of
exposure
Age and date at
physiological recordings
No. 1 M. Arctoides
No. 2 M. Mulatta
No. 3 M. Arctoides
23 days
12 days
6days
vertical
vertical
horizontal
41 days
13 days
7 days
15 h (37 days)
57 h (42 days)
107 h (57 days)
78 days (April 23, 1975)
55 days (June 24, 1975)
64 days (November 13, 1975)
73
Hours
No. 3
100
-
75-
50
25
No I
0
Birth
]0
20
30
40
I
50
~0
I
70
I
80
!
90
I
]00
the end of the recording sessions. Two or 3 penetrations were made in each animal and small electrolytic
lesions ( 2 ~ A for 2 s) were made at transitional points
through the p e n e t r a t i o n (e.g. right eye to left eye
transition, layer 1 V c - V border). A t the termination
of the recording sessions animals were perfused with
saline followed by formol saline. The brains were
blocked and cut in the parasagittal plane, and the 25um frozen sections were stained with cresyl violet. Finally, histological reconstructions were made of electrode tracks, all of which were well within area 17.
Days
Fig. 1. The graph shows accumulated monocular exposure (in
hours) to stripe patterns plotted relative to age (in days) for the
3 infimt monkeys. The initial point of each curve indicates that
day the animal was placed under the fully restricted routine in
the rearing apparatus.
and width at the ends to eliminate an obvious edge.
At their widest dimension the lines were 5 - 2 0 ° in
width and spaced at 1-16 ° intervals, representing a
variety of spatial frequencies. Half the slides consisted of light lines on a dark b a c k g r o u n d and half of
dark lines on a light background. The illumination
levels were in the photopic range, dark areas measured 8 cd/m 2 and bright areas 300 cd/m 2.
Physiological and anatomical methods
The recording m e t h o d s used in these studies have
been described previously'~, 10. A n i m a l s were kept
continuously anesthetized with sodium p e n t o b a r b i tal, paralyzed with a mixture of Flaxedil and D-tubocurarine and artificially ventilated during the course
of the recording session. Body t e m p e r a t u r e and expirated C O , were m o n i t o r e d and kept within normal
physiological range. The sutured lids were o p e n e d
and the eyes refracted by means of contact lenses
upon a tangent screen 1.45 m away.
Unit activity was recorded through tungsten electrodes moved in steps of 2 5 - 5 0 u m in extensive tangential penetrations in area ~0,17. The size, orientation
specificity and binocular interactions of receptive
fields of cells were m a p p e d on the tangent screen
with a hand-held projector. The location of the receptive fields was d e t e r m i n e d relative to the projected locations of the optic discs and foveas for each
eye. Two of the authors (Hubel and Wiesel) did not
know the rearing orientation for the 3 animals until
RESULTS
The 3 experimental animals were raised under the
same conditions of monocular exposure to lines of a
single orientation, but the age of lid suture, onset and
duration of exposure differed as summarized in
Table |. The rate and total accumulation of restricted
visual stimulation in the rearing apparatus are illustrated in Fig. 1.
M o n k e y No. 1 had normal visual experience until
23 days of age, when the right eye was closed. Over
the next two weeks the animal was put into the rearing apparatus and exposed to the vertical stripe patterns for several hours a day but was otherwise kept
in a normal l i g h t - d a r k environment. The fully restricted rearing routines were not begun until 41 days
of age. During the next 37 days the m o n k e y exposed
its left eye to a total of 15 h of vertical stripe stimulation: the rest of the time it spent in complete darkness.
Some effects of both monocular deprivation and of
restricted viewing of vertical orientations can be seen
in this animal although it has a history of 23 days of
binocular and 18 days of monocular visual exposure
in the nursery before the orientation restriction procedure was begun. All the cells recorded (excluding
non-oriented cells recorded in layer IVc) r e s p o n d e d
normally to oriented lines. There was, however, a
decline in the n u m b e r of binocular cells and the ocular dominance distribution was skewed in favor of
the exposed eye (Fig. 2A). The histogram is not unlike that found in animals raised with artificial strabismus s.2,~ or after binocular closure 21. Table II
shows that of the 64 o r i e n t e d cells recorded, the left
(exposed) eye was d o m i n a n t for 599} (groups 1-3),
whereas the right (closed) eye d o m i n a t e d in only
74
A
30--
25~
U
20-
15"-
10-
Z
5-
I
ii
1
I
1
2
3
Exposed
I
I
4
eye
I
5
6
Closed
7
II
eye
OCULAR DOMINANCE
Hot
B
Or.
Cells
Vert.
+45)
20-
C
Or
Cells
(~45)
20
15
10
10
5-
I
1
1
I
2
3
I
4
I
5
6
7
3
i
Exposed eye
OCULAR
Closed
DOMINANCE
eye
Exposed eye
OCULAR
I t-'r4
I
5
I
6
CIo~*ed
7
I
it
eye
DOMINANCE
Fig. 2. Monkey No. 1 experienced a normal visual environment for the first 23 days at which time its right eye was sutured, It received
15 h of restricted orientation exposure over the next 37 days. Recordings were begun at 78 days of age. A: ocular dominance histogram
for all oriented cells recorded. B: ocular dominance histogram for cells with + 45 ° horizontal orientation preference. C: ocular dominance histogram for cells with _+ 45 ° to vertical orientation preference. (Definition of ocular dominance groups: cells of group 1 were
driven only by the contralateral eye; for cells of group 2 there was marked dominance of the contralateral eye: for group 3, slight dominance. For cells in group 4 there was no obvious difference between the two eyes. In group 5 the ipsilateral eye dominated slightly: in
group O. markedly, and in group 7 the cells were driven only bv the ipsilateral eye. )
75
TABLE l[
Orientation pr~ference and ocular dominance
RE, right eye; LE, left eye; H-OR, ceils with _+45° horizontal orientation preference; V-OR, cells with _+ vertical orientation preference: LE-DOM, cells in groups 1-3 bins in ocular dominance histograms; RE-DOM, cells in groups 5-7 bins in ocular dominance histograms; GRP 4, cells in group 4 bin, equally dominated by RE and LE; T. total.
Macaque Oriented
cells
H-OR
cells
(c;~fT)
V-OR
cells
(C'(oJT)
L E - D O M RE-DOM Group 4
cells
cells
cells
(~ofT)
(~'(ofT) ( % o f T )
H-OR Cells
LE
( ~ of H)
RE
(¢'( of H)
Group 4
('~; of H)
LE
RE
No. 1
64
No. 2
76
96
39
(61)*
28
(37)
54
(56)*
38
(59)*
49
(64)*
66
(69)*
13
(52)*
28
(58)*
31
(74) *,~*
11
(44)
20
(42)
11
(26)
1
(4)
-
No. 3
25
(39)
48
(63)*
42
(44)
25
(64) ×.~
21
(75)* ~
35
(65)*
14
(36)
7
(25)
19
(35)
25
(39)
27
(36)
30
(31)
1
(2)
-
V-OR Cells
-
* Highest percent for single orientation.
** Highest percent of LE dominance for either orientation.
39% (groups 5 - 7 ) . W h e n the cells were divided
into two groups according to orientation preference
(cells favoring horizontal _+ 45 ° and those favoring
vertical _+ 45°), the horizontal group showed no preference of one eye over the other, while in the vertical
group, cells d o m i n a t e d by the exposed eye exceeded
those d o m i n a t e d by the closed eye by nearly 2:1 (Fig.
2C; see also Table II).
The second m o n k e y was raised under similar conditions having the right eye occluded by lid suture
and the left exposed to vertical stripes. The main differences in this animal were that the deprivation (the
monocular occlusion and stripe exposure) began at
12 and 13 days of age and that the total accumulation
of visual stimulation was 57 h c o m p a r e d to 15 h in the
first instance.
The activity of cells in the striate cortex a p p e a r e d
normal also in this animal, as in the first one, with the
usual distribution of cells responding selectively to
different orientations. The cells r e c o r d e d in a single,
4-ram long, tangential p e n e t r a t i o n through the superficial layers is shown in Fig. 3. T h r o u g h o u t the penetration the orientation preference shifted in small
regular steps first counterclockwise, then clockwise,
just as seen in normal m o n k e y s 1° and in newborn
monkeys with little or no visual experience 22. In particular there was no suggestion of any compression or
expansion of areas preferring horizontal or vertical
orientations, as might be expected if the rearing procedure had led to a distortion of orientation columns.
There was, however, a clear decrease in the repre-
sentation of the occluded eye and an increase in the
areas d o m i n a t e d by the exposed eye, especially in regions of vertical orientation preference.
This differential distribution of orientation preferences for individual cells is shown in Fig. 4. Fig. 4A
consists of cells d o m i n a t e d by the occluded eye and
Fig. 4B consists of the cells preferring the exposed
eye. There was a striking paucity of cells which prefer
vertical orientation in the occluded eye and a rather
uniform distribution of orientation preferences in the
eye exposed to vertical stripes. The few binocular
cells recorded had the same orientation preference in
the two eyes and those with orientations a r o u n d the
vertical ( m a r k e d by numbers 186, 188 and 190 in Fig.
4B) were d o m i n a t e d by the exposed eye.
A n examination of the ocular d o m i n a n c e histogram of the cells recorded in the second m o n k e y also
revealed this asymmetry. As in the first m o n k e y
there was a decline in the n u m b e r of binocular cells
and an overall dominance of the exposed eye (Fig.
5A and Table II). This dominance was slight for cells
preferring horizontal orientations (Fig. 5B), but
more m a r k e d for cells with orientation preference
around the vertical (Fig. 5¢').
In the third case the animal was lid-sutured at 6
days of age and entered the rearing apparatus on the
following day. The m o n k e y was exposed to horizontal stripes over a p e r i o d of 57 days and accumulated a
total of 107 h of exposure. Thus this m o n k e y had
more stimulation to a single orientation than the others and the exposures were confined to the period of
76
s
80-
-
60-
/
40-
t
20-
I
0-
-20DQg
\
-40-
"
-60"
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00
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o
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2.2
2.4
2.6
2.8
3.0
3.2
3.4
3.6
3.8
4.0
TRACK
DISTANCE
-
turn
Fig. 3. Monkey No. 2 had normal visual experience until 12 days of age at which time its right eye was sutured. On the following day it
was placed continuously in the rearing apparatus where it received a total of 57 h of exposure to vertical lines over the next 42 days.
Recordings were begun at 55 days of age. Graph of orientation vs tract distance for cells driven by the open eye (closed circles) and for
cells driven by the closed eye (open circles) recorded in a 4-ram oblique penetration in Monkey No. 2.
high susceptibility to visual deprivationJt. Of the 3
monkeys, this one showed the most marked decline
in the number of binocular cells and the most striking
asymmetry, again in favor of the open eye (Fig. 6A).
To our surprise, however, there was little difference
in this asymmetry between cells favoring orientations
around the horizontal axis and those favoring the vertical axis (Fig. 6B, C). The asymmetry was slightly
g r e a t e r for h o r i z o n t a l , but not so great that o n e was
able to guess w h a t o r i e n t a t i o n had b e e n used for the
rearing ( T a b l e ll).
T o s u m m a r i z e , the m o n o c u l a r s t r i p e - r e a r i n g p r o c e d u r e in t h e s e 3 m o n k e y s resulted in a decline in the
n u m b e r of b i n o c u l a r cells and a decline in the influe n c e of the closed eye. T h e decline in the effectiveness of the closed eye, seen mainly for cells r e s p o n d -
77
A
B
.....
i
Fig. 4. Monkev No. 2. Orientation of cells recorded in the oblique penetration through striate cortex illustrated in Fig. 3. ('ontinuous
lines indicate monocular cells or cells preferring the designated eye. Interrupted lines indicate binocular cells or cells prcterring thc
other eve. Numbers refer to micrometer depth readings (in mm × 100). A: orientation of cells driven by the closcd cvc. B: oricntation
of cells driven by the exposed c\c.
ing to orientations close to the one used in the rearing
procedure, was statistically significant (P = 0.02,
two-sample/-tests on 3 animals combined). The difference in the effectiveness of the two eyes for cells
responding to orthogonal orientations was not significant (see Table ll).
DIS('LISSION
The results from these monkeys make it clear that
preferential exposure of one eye early in life to
stripes of a single orientation can change the ocular
dominance distribution of cortical cells. In till 3 animals, the population of cells as a whole came to favor
the open eye, and the earlier the deprivation and the
longer its duration, the greater was the shift in eye
preference (Figs. 2, 3.6, parts A). Not unexpectedly,
the shift was greatest for the cells having orientations
at and around the orientation of the stripes used in
the exposure. That this difference was least striking
- - and perhaps not even significant - - in the third
monkey, which had the earliest and longest deprivation, was a surprisc to us and we have no ready explanation.
In the case of the first two monkeys, it might at first
glance seem strange that after exposure of one eve to
stripes of one orientation the distribution of cells
according to optimal orientation through that eye
should remain unchanged (e.g. Fig. 4B) and that
meanwhile the other eye should be robbed of its control over cells having the orientation used in rearhtg
(Fig. 4C). This result, however, is ,just what is predicted from the earlier simpler procedures, in both
cats and monkeys, of rearing in a normal visual enviro n m e n t with one eye closed, as opposed to having
both eyes closed. The monocular closure is by far the
more drastic procedure, and there is now an abundance of evidence that the difference is due to competitive effects between the two eyes <t~,25. In the
present experiments where, say, only vertical stripes
were presented to one eye, the procedure a m o u n t e d
to a binocular deprivation for horizontal and monocular deprivation for vertical. In cats very similar results have been obtained in kittens in which the rearing procedures took advantage of this competition in
a similar way, with selective exposure of one eye to a
single orientation, the other eye being either open or
closed 15. Similar results were likewise obtained when
the selective depriwition was achieved by means of
artificial astigmatism produced by a cylindrical lens
over one eye 4. In facL studies in which competition
has played a part have consistently produced striking
78
A
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30-
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gB
20-
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10-
!
I
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!
3
2
4
5
eye
7
6
i
Closed eye
OCULAR D O M I N A N C E
B
Or,
Hot.
30-
C e l l s (+_45)
Vert.
C
u~
Or.
Cells
(~45)
20-
25
u~
~J
u~
20
1510
~0Z
Z
5-
5"
I
I
2
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3
eye
OCULAR
I r-v-4
5
6
Closed eye
DOMINANCE
I
7
I
2
Exposed
3
eye
OCULAR
r--J
I
4
I
5
6
7
Closed eye
DOMINANCE
Fig. 5. M o n k e y No. 2. A: ocular dominance histogram for all oriented cells recorded. B: ocular dominance histogram for cells with _+
45 ° horizontal preference. C: ocular dominance histogram for cells with _+ 45 ° vertical preference.
79
A
60-
u~
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30-
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t0-
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I!
1
I
2
3
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;
4
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5
6
!
I
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OCULAR DOMINANCE
Hor. Or.
B
Cells
Vert.
(+_45)
G
4o-
Or.
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u')
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u.,i
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2
J
3
4
5
i
Exposed eye
OCULAR
6
i
7
lJ
Closed eye
DOMINANCE
i0-
q
1
2
3
Exposed eye
OCULAR
4
5
6
7
ii
Closed eye
DOMINANCE
Fig. 6. Monkey No. 3 was given normal visual experience until 6 days of age when its right eye was sutured. It was placed in the rearing
apparatus on the following day. Over the next 57 days a total of 107 h of restricted exposure to horizontal lines was obtained. Recordings were begun at 64 days of age. A: ocular dominance histograms for all oriented cells recorded. B: ocular dominance histogram for
cells with +_ 45 ° horizontal preference. C: ocular dominance histogram for cells with _+ 45 ° vertical preference,
8(1
abnormalities in the cortical physiology, beginning
the orientation orthogonal to the rearing orientation.
with the original work of Hirsch and SpinelliT, in
Furthermore. in sequences of cells recorded in long
which goggles produced stimuli of one orientation to
one eye and the orthogonal to the other, up to recent
tangential penetrations we saw regular shifts m
confirmation in a more strictly controlled study by
Stryker et al. z:. The results of kitten experiments in
which both eyes were exposed to a single orientation
have been much more varied, from the profound effects seen originally by Blakemore and Cooper~ to
the lack of any effects seen bv Stryker and Sherk L~.
Rauschecker and Singer i4 raised kittens with cylin-
orientation, similar to those seen in newborn or normally reared m o n k e y s 1<22. This indicates that the
orientation columns present at birth remained intact
despite orientation deprivation in one eye and pattern deprivation in the sutured eye during the first
two months of life. Thus these results give no support
to the suggestion that orientation preferences are induced by visual experience.
drical lenses and concluded from their study that both
"selective and instructive processes" may be operating in the development of orientation preference.
The wide discrepancies between all these studies remain unexplained but are perhaps not surprising in
view of the variations in results that can be seen in
ACKNOWLEDGEMENTS
The monkeys used in these experiments were
raised at the New England Regional Primate Center:
we wish to express our appreciation for the use of
a single set of experiments - - for example Figs. 2 and
5 vs. Fig. 6 of the present paper!
Despite the changes reported here, the present ex-
these excellent facilities and for the generous help
given by Dr. P.K. Sehgal, Ellen Newhouse and many
periments in macaque monkeys provided no support
for the notion that it is possible to modify innately determined orientation preference of striate cortex
cells by rearing animals viewing only horizontal or
others. We also wish to thank Dr. F e r n a n d o Gonzalez who programmed the schedules and Dr. Jerome
Blue and Mary Russell for assistance in rearing the
animals. Bea Storai and Karen Larsen did the histo-
vertical lines. Cells of all orientations were recorded
logy of the brain tissue and Cathy Cross and Peter
Peirce the photography, for which we are grateful.
in the exposed eve of these monkeys and there was
no obvious preponderance of cells preferring the
orientation to which the eye has been exposed. In
The work was supported by NIH Grants EY0605 and
EY0606 to Drs. D . H . H . and T . N . W . M . C . was sup-
fact, in monkeys No. 2 and No. 3 the majority of cells
(63c+ and 56C+ respectively) showed preferences for
ported by NS14261 during the preparation of the
manuscript.
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