256 Neuroscience Letters, 124 (1991) 256-259

~) 1991 Elsevier Scientific Publishers Ireland Ltd. 0304-3940/91/$ 03.50

ADONIS 030439409100054Z
NSL 07648

On the functionality of the visually deprived occipital cortex in early blind

persons

F. Uhl, P. Franzen, G. Lindinger, W. L a n g a n d L. Deecke

Neurological University Clinic, Vienna (Austria)
(Received 31 January 1990; Revised version received 27 December 1990; Accepted 7 January 1991)

Key words: Occipital cortex; Visual deprivation; Blind human; Tactile

In early blind mammals, the deprived visual cortex undergoes anatomical and functional alterations. Its functional role was investigated in the
early human blind by using patterns of cortical activation as measured by scalp-recorded event-related slow negative DC potential shifts. The blind
showed higher occipital negativity than did sighted persons both during a tactile reading task and a non-reading tactile control task. Results point
to a possible role for the blind's visual cortex in tactile processes.

The observation of von Senden [15] that blind-born
humans face great difficulties in the use of vision if their
eyes are operated late has stimulated various studies on
the visually deprived occipital cortex of the animal [11].
In sum, both changes in microstructure and in neural
functioning have been demonstrated for visual areas.
For instance, single neurons in the visual association
cortical area 19 of blind-born monkeys exhibited a de-
creased response to visual stimuli, but an increased res-
ponse to somatic stimuli [5]. In contrast, attempts to
study functional alterations in the human have not all
yielded confirmatory results. The occipital metabolic ac-
tivity of early blind human subjects as measured by posi-
tron emission tomography (PET) has been reported as
normal [10] or above normal [16]. However, in order to
establish the functional significance of this cortical area,
evidence is required for a task-dependent modulation of
the activity of this cortical area. Thus, cortical activity
was monitored by means of the scalp-recorded event-
related negative DC potential shift [2] which has been
shown to covary with increased neural activity [1] in the
upper cortical layers and with regional cerebral blood
flow (rCBF, ref. 7). Furthermore, the sensitivity of the
present method exceeded that of rCBF in two joint stu-
dies [4, 7, 12].
Eleven right-handed subjects (median age 26 years)
who became blind within the first year of life (n = 2) or
who were congenitally blind (n = 9) participated. Their
Correspondence: F. Uhl, Neurologische Universit~itsklinik Wien,
Lazarettgasse 14, A- 1090 Wien, Austria.
history and clinical examination were free of cerebral
disease (stroke, tumor, craniotomy). Eight were totally
blind. Three had residual dark/bright perception without
having had any contour or pattern vision. All subjects
started to learn Braille reading at the age of 6 years. In
the experimental condition, tactile reading of sentences
coded in Braille was required using the left index finger
for scanning. In the control condition, subjects were
passing their left index finger over a random dot pattern
in a left to right manner similar to that used while Braille
reading. Prior to each trial, subjects positioned their left
hand at the left border of the stripes of either Braille text
or random dot pattern. Using DC amplifiers and Ag/
AgC1 surface electrodes, cortical potentials referred to
resistor (5 k~) linked earlobes were recorded from left
(F3) and right frontal (F4), left (C3), mid (Cz), and right
central (C4), left (P3) and right parietal (P4), left (T5)
and right posterior temporal (T6) and left (O1) and right
occipital (02) scalp positions along with the diagonal
EOG. To minimize habituation effects, the tasks were
switched after each block of 16 trials, until 64 artifact-
free trials per task were collected. Averaging was per-
formed time-locked to the self-paced initiation of the
task. According to Noebels et al. [9], trials containing
detectable EOG activity were rejected. To quantify DC
negativity accompanying the performance of the task,
the mean amplitude was calculated within a 2000 ms in-
terval from 6 to 8 s after initiation (termed performance-
related negativity, N-P; stippled area in Fig. 1).
For comparison, the same paradigm was applied to
sighted normals familiar with the Braille code. Since
even teachers for the blind use their eyes for reading
 257

Braille, only three of such persons were willing to under- up of 3-6 letters) was presented instead of Braille code
go a tedious training program enabling them to partici- thus allowing a statistically sufficient number (17) of
pate in the present tactile task. Thus, a relief script sighted normals (10 female; median age 27 years) to par-
(embossed upper case letters spelling short words made ticipate. The sighted and the blind subjects both used the

II EarLy Blind 17 Sighted NormaLs 3 Sighfed Teochers

_ ~ ........:~:~.dEOG . . _ ~ ~ . . . . ~ ' dEOG

F3

_ ~ F4 F4 ._~ _ . ~ ~

[4 C4

P3 ;_. ~ P3
/
~...........~
P4 . . . ~ ' : : ' : : : : ' : P4

T5 _ ~ : ......" " ~ " TS

T6 _~
...w.,,"" ...........
.... • o" ....

0
Rar~om d o t - -
BraiLLe........ ReLief........ BrailLe........
Fig. 1. Grand averages. The continuous line waveforms stand for the sensorimotor control task of passing the finger over random dots. The dotted
waveforms represent the tasks of tactile reading (Braille in the blind and teachers, relief letters in the 17 normals). Negativity up. The onset of task
performance is represented by the vertical line (t=0). The hatched window stands for the time interval used for calculating the mean amplitude
N-P.
258
same random dot control pattern. Following testing, the
subjects were questioned as to their use of visual imagery
while scanning the test patterns and they were scored
with zero for no use of imagery, 1 for 'a bit', 2 for
'marked use', and 3 for 'very vivid use of imagery'. The
blind reported to have used virtually no imagery (mean
0.3) as opposed to the sighted subjects (teachers: mean
2.0; 17 sighted normals: mean 2.1).
It was hypothesized that the blind's occipital cortex
would be involved in tactile tasks and thus would exhibit
additional occipital negativity as related to negativity at
C4, reflecting activation of the right central cortex [6]
considered to be primarily involved in a tactuomotor
task by the left hand.
Sustained negative DC potential shifts accompanied
the performance of the tasks (Fig. 1). In the blind, the
increase in DC negativity while scanning Braille sen-
tences as compared to the random dot control task was
maximum at occipital sites. There was no increase at the
right central electrode site (C4) positioned over those
cortical areas considered of primary importance for sen-
sorimotor guidance of the left hand. In sighted normals
and teachers, however, such a task-related increase was
more widely distributed, particularly affecting central
sites.
For statistical assessment, right occipital (02) N-P
was submitted to a repeated-measures two-factorial ana-
lysis of variance with the between subject factor of
'group' (11 early blind vs 17 sighted normals) and the
within-subjects factor of 'task' (random dot vs Braille/
Relief scanning). ANOVA* gave rise to a main effect of
group (df=l,25; F=9.8; P=0.004), indicating an in-
crease in the blind, and of condition (df= 1,25; F = 16.3;
P = 0.001), indicating an increase with tactile reading.
The topographical distribution of the DC potentials
was assessed by use of the centro-occipital gradient,
obtained by subtracting N-P recorded at 0 2 from N-P
recorded at C4. Again, ANOVA yielded a main effect of
'group' (df= 1,24; F=5.7; P=0.025): in the blind, right
occipital negativity was greater than the right central (on
the average, such gradient was - 4 . 5 pV _+ 1.2 S.E.M.
with reading, but - 0 . 9 pV +_ 2.0 with random dots),
while the opposite topographical distribution occurred
in sighted normals (+2.4 pV _+ 1.2 with reading; + 1.2
/tV _+ 0.8 with random dots). The main effect of 'task'
was non-significant (df= 1,24; F = 1.9; P = 0.18), but the
significant 'group-by-task' interaction (df-- 1,24; F = 7.7;
P = 0.011) indicated a further increase in the blinds' occi-
*Explanation for the reduced degree of freedom for sighted subjects:
In subject9, C4 recordingsof both tasks had to be dropped as a conse-
quence of too few artifact-freetrials (i.e. free of tempoalismuscleactiv-
ity). In subject 14, C4 and 02 recordingsin the readingtask were not
evaluateddue to too few artifact-freetrials.
pital preponderance with Braille reading (df=l,10;
F = 5.4; P=0.043), but no such increase for the sighted
(df= 1,14; F = 1.6; P = 0.23). Finally, hemispheric latera-
lization was assessed by use of the occipital hemispheric
difference score 'dO (N-P at O1 subtracted from N-P at
02). The two-factorial ANOVA yielded a main effect of
'group' (dr=l,25; F=4.9; P---0.037) indicating right
occipital lateralization of negativity in the blind (average
dO: - 0 . 4 pV +_0.8 with reading; -2.1 /~V +_0.4 with
random dots) as opposed to left hemispheric lateraliza-
tion in sighted (average dO: +0.5 pV +_0.8 with reading;
+ 1.2 pV +_0.7 with random dots). The main effect of
task was non-significant (df= 1,25; F=2.9; P=0.36)
while the significant 'group-by-task' interaction
(df= 1,25; F=5.7; P=0.025) indicated that the groups
differed more in the random dot control task.
If scalp-recorded negative DC shifts indeed reflect
neural activation of the underlying cortical areas [13, 14],
then the findings suggest that the visually deprived occi-
pital cortex of early blind humans seems not to be inac-
tive, but rather to have retained considerable function.
First, occipital negativity in the blind was greater than
in sighted persons both in conditions of passive (random
dots) and active touch (Braille or relief reading). Second,
these differences cannot be explained by diffuse arousal
effects on the general DC level, since distinct modula-
tions in topography emerged. In sighted normals, perfor-
mance-related negativity was distributed clearly in
favour of the contralateral central cortex considered to
be primarily involved. By contrast, the blind seemed to
exhibit additional occipital negativity which even
increased when tactile reading of Braille was required.
Unlike in sighted normals, this occipital increase cannot
be accounted for by visual imagery [12]. The finding
might suggest some involvement of the deprived occipi-
tal cortex in tactile reading of Braille. These positive re-
sults might encourage further investigations trying to
tease apart the effects of the tactile and of the language-
component involved in tactile reading of Braille. Such
tactile - versus-language - considerations do not ques-
tion the essence of the present results, since the addition-
al occipital negativity was already observed in the non-
reading random dot task.
In sum, results corroborate Doty's observation that
'the visual cortex has non-visual functions as well' [3].
With visual deafferentation such as in blindness, non-
visual modalities apparently increase in gain. Also, the
auditorily elicited mismatch negativity (MMN) evoked
by changes in tone-location revealed a more posterior
scalp distribution in blind than in sighted subjects [8].
The cooperation of the blind volunteers and the direc-
tors of the Vienna 'Bundesblindenerziehungsinstitut'
 259

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