VISION, CENTRAL
Magnetically induced phosphenes in sighted,
blind and blindsighted observers
Alan CoweyCA and Vincent Walsh
University of Oxford, Department of Experimental Psychology, South Parks Road, Oxford OX1 3UD, UK
CA
Corresponding Author
Received 13 June 2000; accepted 31 July 2000
Direct stimulation of visual cortex can produce illusory ¯ashes
of light, called phosphenes. Here we describe the spatial and
motion properties of phosphenes produced by transcranial
magnetic stimulation in normal subjects and in two subjects
with peripheral or cortical blindness. The totally retinally blind
subject experienced normal phosphenes, apart from their
concentration in the centre of the visual ®eld, whereas the
Key words: Areas V1 and V5; Awareness; Blindsight; Phosphenes; Transcranial magnetic stimulation
INTRODUCTION
Direct stimulation of the visual cortex by transcranial
magnetic stimulation (TMS) can reveal several aspects of
visual processing. Amassian et al. [1] were the ®rst to use
TMS as a `virtual lesion' in studies of vision by brie¯y
inactivating visual cortex, and this method has since been
used to examine visual detection and discrimination [2±6],
awareness [7], visual attention [8], plasticity [9,10] and
cross-modal interactions [11]. As predicted by Pen®eld and
Rasmussen [12], albeit with respect to direct electrical
stimulation rather than magnetic stimulation, the effects
caused by posterior occipital stimulation, involving early
cortical visual areas, arise from interference with normal
visual processing in the retinotopically corresponding part
of the visual ®eld [3], irrespective of whether the stimula-
tion produces a visual image, i.e. a phosphene.
Phosphenes can also be used to probe the functional
sparing and residual properties of damaged visual cortex
as well as to brie¯y impair vision. We therefore attempted
to induce phosphenes by TMS to assess the integrity of
visual cortex in a totally retinally blind subject and to
compare the results with those obtained by stimulating the
same regions in normally sighted individuals and in a
well-documented hemianopic subject who shows blind-
sight in his impaired ®eld. The purpose of this was to test
the hypothesis that in the absence of area V1, the retinal
projection to visual region MT‡/V5 alone, via the mid-
brain and the pulvinar nucleus, is suf®ciently effective to
give rise to conscious visual impressions, especially of
moving stimuli [13±15].
In normal vision, the perceived angular size of an after-
image produced by intense retinal stimulation is constant
over different imagined viewing distances, i.e. the after-
0959-4965 & Lippincott Williams & Wilkins
NEUROREPORT
hemianopic subject, lacking area V1, did not experience
phosphenes when his surviving extrastriate visual areas were
stimulated. In the absence of V1, magnetically induced activity
was unable to generate a conscious visual percept in the ®eld
defect. NeuroReport 11:3269±3273 & 2000 Lippincott Williams
& Wilkins.
image looks bigger the further away it is perceived. This is
known as Emmert's law. Whether this size/distance scal-
ing has a cortical basis in any particular cortical area or
areas is unknown. Phosphenes are like after-images but are
produced cortically as opposed to retinally. We induced
phosphenes by stimulating occipital cortex (areas V1/V2
and perhaps V3) and region MT‡/V5 and asked subjects
to indicate their size on a surface at three different viewing
distances in order to see whether Emmert's law still holds.
Extensive and intensive stimulation of the damaged left
hemisphere in the blindsight subject did not yield phos-
phenes, with one notable exception, even when applied to
area MT‡/V5 on that side. The phosphenes reported by
the peripherally blind subject were as easily elicited and as
reproducible as those experienced by other subjects but
their spatial distribution suggests that retinotopic mapping
in this subject's visual cortex is degraded. Despite the
coarser spatial mapping the blind subject showed other-
wise normal phosphenes including perception of move-
ment when MT‡/V5 was stimulated.
MATERIALS AND METHODS
Peripherally blind subject PS, now aged 61, suffered
transection of one optic nerve and compressive destruction
of the other in a road traf®c incident and has been totally
blind for almost 8 years. He cannot even tell when the
lights in a room are turned on and off and has no pupillary
response to a bright light when measured with a pupill-
ometer. Hemianopic patient GY suffered almost complete
destruction of left striate cortex (V1), with some additional
damage to extrastriate areas V2 and V3, in a traf®c accident
at the age of eight [16]. His area MT‡/V5 has been
localized in both hemispheres in functional neuroimages
Vol 11 No 14 28 September 2000 3269
NEUROREPORT
co-registered with structural images [15±17]. We used
these published structural and functional localisations,
together with our own structural MR images of his brain
and skull, to locate MT‡/V5. In his damaged hemisphere
MT‡/V5 is more or less mirror symmetrical with its
position in the normal hemisphere. He is now in his early
forties and has been a subject in many experiments on
blindsight (see [18] for review). The remaining six subjects
were neurologically normal and included both experimen-
ters. All subjects were tested with the approval of the
Departmental Ethical Committee and in accordance with
the declaration of Helsinki and gave their informed con-
sent for the experiments.
For the ®rst experiment, to plot the spatial localization of
phosphenes, the subjects ®xated the centre of a tilted
drawing board placed 57 cm in front of them, and de-
scribed and drew any visual sensations produced by the
TMS. Magnetic stimulation was applied over the occipital
scalp with the centre of a 70 mm ®gure-of-eight coil
positioned at various points between 1 and 4 cm dorsal to
the inion and 1±3 cm laterally, for occipital stimulation
above the region of V1, V2 and V3 until we found sites at
which the subjects consistently reported visual phos-
phenes. The coil was held tangential to the skull with the
handle pointing downwards at 458 to the axis of the
spinal cord. For direct excitation of region MT‡/V5 we
stimulated points on the scalp centered 3 cm above the
inion and 5 cm lateral, as described previously [19,20]. The
subject wore an elasticated swimming cap, secured with
surgical tape to the temples, forehead and nape of the
neck. The inion and the midline were marked on the cap.
Stimulating points were similarly marked by adhesive tabs
at 1 cm intervals, but stimulation was often applied be-
tween marked points.
Pulses were delivered using a MagStim Super-Rapid
system with four boosters giving a maximum output of
2 T. After each pulse or pulse train the subjects were
asked whether they saw anything. If the answer was yes
they were asked `where was it, did it have a colour or a
shape, and did it move?'. Soon, subjects responded without
being asked. When a phosphene site was located we
systematically moved the coil to sites 1±2 cm away from
this site and the stimulation was repeated. At every site,
stimulation was delivered three times, >8 s apart, but often
longer. Between triplets of stimulation the interval was
> 20 s while the subject summarized the results of each
triplet and the experimenters recorded the observations
and selected the next stimulating point. The subject drew
the outline of each phosphene on the drawing board.
In the ®nal procedure subjects were seated 50, 100 or
200 cm from a white wall bearing a ®xation point and used
a laser pen to trace the outlines of phosphenes, which were
then faintly repeated in pencil by one of the experimenters.
All testing took place in a dimly lit room while subjects
®xated a just-visible point.
RESULTS
All six fully sighted subjects drew their phosphenes in
different locations according to the site of stimulation.
When the stimulating coil was moved forwards and
dorsally from the inion the phosphenes became progres-
sively more eccentric in the contralateral lower visual
3270 Vol 11 No 14 28 September 2000
A. COWEY AND V. WALSH
quadrant. When the coil was moved laterally, away from
the vertex, the phosphenes moved away from the midline
of the visual ®eld, as would be expected if the TMS is
stimulating the upper bank of the calcarine sulcus, in
which the horizontal retinal meridian is represented later-
ally in the fundus of the sulcus [16,21,22]. Figure 1a shows
an example, which was characteristic of all six normal
subjects. In all of them the phosphenes produced by TMS
near the mid-line were often coloured, albeit faintly, but
were rarely described as moving.
Stimulation over the region of the occipital pole of the
peripherally blind subject also produced phosphenes (to
his surprise and pleasure) but in three separate testing
sessions several months apart there was less systematic
spatial organization to his perceptions (Fig. 1b) and he
drew clustered overlapping phosphenes around the centre
of ®xation, which was his ®nger tip placed on a raised
mark on the centre of the drawing board. We could not
determine whether his anatomical visual axis was aligned
with his ®nger tip but his phosphenes were reliably
reproduced with a scatter of only a few degrees. However,
there could well be a constant error in their position, for
example if his anatomical visual axis was consistently
above his ®ngertip. Like normal subjects he frequently
described them as coloured, most often using the descrip-
tions red, green, blue, silver, gold, orange, or brown. In
contrast, blindsight subject GY showed normal spatial
organization in the ®eld ipsilateral to his damaged striate
cortex (Fig. 1c) but when TMS was applied over his
damaged left occipital region it was not possible to elicit
any phosphenes, even with repetitive pulse TMS (rTMS) of
up to 10 Hz, 0.5 s and maximal stimulator output.
We next explored regional specialization in TMS-
induced phosphenes. We have shown elsewhere [19,20]
that rTMS applied over visual area MT‡/V5 can produce
moving phosphenes. Our six sighted subjects reliably
perceived movement in the visual ®eld contralateral to the
hemisphere of stimulation (Fig. 2a). The illusory movement
in some subjects was a single elongated blob which jumped
in one direction but in one subject it was like a detuned TV
screen. In response to questions about their colour, these
moving phosphenes were usually described as silvery,
cloudy, sparkling, greyish, like pepper and salt, or a snow-
storm, and only rarely having any colour, which was
always faint. In all subjects other than retinally blind PS,
the movement on each trial tended to have an overall
centrifugal or centripetal direction. PS also reported move-
ment but his reports were consistent with the disruption of
topographical representation revealed by stimulation at
and near the midline (Fig. 2b). Whereas sighted subjects
often reported discrete and unidirectional perceptions of
movement, phosphenes reported by PS drifted or swirled
or shimmered, and changed direction and position from
trial to trial at the same scalp location. Blindsighted subject
GY also perceived moving phosphenes but only when
TMS was applied over MT‡/V5 of his intact right hemi-
sphere (Fig. 2c). To ensure that we had not simply missed
his motion area in the damaged hemisphere we located it
from functional and anatomical PET and MRI scans of his
brain [15±17] and applied TMS at and around this region
at 100% of stimulator output for 1 s at 10 Hz, within a grid
of 4 3 4 cm. Such dense sampling and intense stimulation
MAGNETICALLY INDUCED PHOSPHENES NEUROREPORT

(a) (a)
V1 phosphenes VW Single pulse; 70% of output V5 phosphenes
Midline and lateral above inion
VW 10 Hz, 0.5 s; 70% of output
A: 2.5, 0 Midline and lateral above inion
B: 4, 0 A A: 3, 5 right hemisphere
C: 5, 0
B: 3, 5 left hemisphere
D: 4, 1 (right) B
1 2 3 4 5 6
E: 4, 1 (left) 7 6 54 3 2 1
A

B E
D
C

(b) PS5 Hz, 0.5 s; 80% of output

Right hemisphere 0 Hz, 0.5 s; 80% of
Midline and lateral above inion output
B PS
A: 2, 0 (b) Midline and lateral
B: 2, 1 B above inion
A
C: 2, 2 A: 3, 5
1 2 3 4 5 6 7 D: 3, 0 B B: 3, 5
E: 3, 1 B D: 4, 4
B 1 2 34 5 6 7
F: 3, 2
G: 3, 3
D A
H: 4, 0 A
I: 4, 1
B
J: 4, 2

(c) GY 8 Hz, 0.5 s; 70% of output

Right hemisphere
Midline and lateral above inion GY 80% of output
A: 2, 1 Right hemisphere
B: 2, 2 (c) Midline and lateral above inion
D C: 4, 1 A: 3, 6 4 Hz, 0.5 sec
B A
D: 3, 1 C B: 3, 6 12 Hz, 0.5 sec
4 8 121620 24
E: 2.5, 2 B 2 4 6 8 1012 C: 4, 5 8 Hz, 0.5 sec
E A

Fig. 1. Examples of retinotopic mapping of TMS induced phosphenes:
(a) in a normally sighted observer (b) in retinally blind subject PS and (c)
in hemianopic patient GY. The coordinates give the site of stimulation in
dorsal-lateral order. For example, 2,1 indicates that the coil was centered
2 cm above the inion and 1 cm lateral. Note that as the coil is moved
dorsally and forward, away from the inion, the phosphenes migrate
inferiorly (cf A and C in GY, part c), and that as the coil is moved
laterally from the midline the phosphenes migrate further from the
vertical meridian, in the contralateral visual ®eld (cf A and B in GY, part
c). In retinally blind subject PS (b) the phosphenes were clustered in the
central 38 of the visual ®eld despite stimulation being delivered between
2 and 4 cm above the inion and up to 3 cm lateral. Where phosphenes
were elicited beyond the central 28 they are in the opposite direction to
that predicted by normal retinotopic mapping i.e. stimulating more
dorsally yielded more superior rather than inferior phosphenes. PS drew
all his phosphenes as small smudges, about 18 across. As they overlapped
extensively, only their centres are marked. The arrowheads on the
medial outlines of phosphenes D and E in (c) indicate that the phosphene
extended much further into the periphery to a border that could be
indicated only vaguely by the subject.
Fig. 2. Moving phosphenes: (a) in a normally sighted observer (b) in
peripherally blind subject PS and (c) in the intact ®eld of hemianopic
patient GY. All three subjects reported moving phosphenes. In GY the
centripetal direction was particularly strong, as indicated by the arrows.
In PS stimulation at the same scalp position a few seconds apart yielded
phosphenes with different positions. Such extensive variability was not
seen in six sighted subjects.
around this area makes it unlikely that we failed to
stimulate MT‡/V5. Intriguingly, on three trials (out of
. 100) in which rTMS was applied to left hemisphere
MT‡/V5, he reported faint perceptions in the normal
ipsilateral visual ®eld, which suggests that we weakly
activated neurons in the normal hemisphere via callosal
connections, an interpretation consistent with a recent
functional imaging study [16]. These faint ipsilateral phos-
phenes did not move and we could not replicate them on a

Vol 11 No 14 28 September 2000 3271

NEUROREPORT A. COWEY AND V. WALSH

subsequent testing session. It is not disputed that area (a) 110

MT‡/V5 is essential for normal perception of visual
motion but it is clear that shorn of its major input from V1,
activity in MT‡/V5 alone, when induced by TMS, was
insuf®cient to generate a visual percept of movement in
the contralateral ®eld defect.
The results of stimulating occipital cortex in the retinally
blind subject PS (Fig. 1b) show that the cortex remains
excitable but suggest that spatial representation is de-
graded. The latter ®nding apparently contradicts the re-
sults of Brindley and Lewin [23], who observed clear
retinotopic organization with direct electrical stimulation
via electrodes implanted on the medial surface of the
occipital lobe in a peripherally blind subject. A possible
explanation is that the more diffuse nature of TMS may
make precise localization impossible if the cortical repre-
sentation has become coarsened as a result of changes in
sensitivity and perhaps in intracortical connexions. Thus
the visual cortex of the retinally blind may retain retinoto-
pic speci®city when conditions are optimized, as they are
with direct and focal electrical stimulation.
The last experiment investigated the similarity between
phosphenes reported by blindsight subject GY and by
sighted subjects with respect to Emmert's law of size
scaling. The retinally blind subject was excluded from this
comparison because he could only draw things with
reference to his ®ngertip on the drawing board within
reach. Nor was it clear that he could imagine ®xating at
different distances and projecting his phosphenes appro-
priately. Figure 3 shows the phosphenes perceived by a
sighted subject and by blindsighted GY when magnetically
stimulated over striate cortex (Fig. 3a) or area MT‡/V5
(Fig. 3b) when seated 50, 100 or 200 cm away from the
®xation spot. Clear but imperfect size scaling is present
with phosphenes from both stimulated regions.
DISCUSSION
It was straightforward to elicit phosphenes from stimula-
tion of occipital sites in all subjects, except from the
damaged hemisphere of the blindsighted subject. Not all
investigators have found it easy to induce phosphenes but
several have had a success rate comparable to that reported
here [3, 24±27].
The retinotopic speci®city of the phosphenes seen by
sighted and hemianopic subjects, together with the func-
tional specialization present in all subjects, including retin-
ally blind PS, indicates that the locus of phosphene
induction must be cortical, not only because the experience
of phosphenes induced from areas V1/V2/V3 and MT‡/
V5 is so similar but also because it is unlikely that
subcortical activation could produce movement speci®c
phosphenes from a coil positioned remotely over MT‡/V5
but not over V1/V2/V3. As we noted above, stimulation
with the high magnetic ®eld intensities used here would
lead to activation in visual areas surrounding V1. It is
therefore noteworthy that just as stimulation of left MT‡/
V5 failed to elicit a perception of visual motion in the
blindsighted subject GY, so too did the probable stimula-
tion of areas V3 and V3a, adjacent to the lesion in V1 and
V2, even though they can be activated in functional
neuroimaging experiments [16,17]. These results can not be
attributed to inadequate sampling. For example, in a single
F
E D
15
0 Size
(cm)
A
B 25
C
210
30 25 20 15 10 5
(b)
110
B
C A
15
F
D
E Size
0
(cm)
25
210
30 25 20 15 10 5
Distance from fixation (arbitrary units)
Fig. 3. Phosphenes and size-distance scaling. (a) Occipital stimulation.
Phosphenes A, B and C were obtained in a normally sighted subject with
TMS applied 4 cm above the inion and 1 cm to the right. The viewing
distances were 50 (A), 100 (B) and 200 (C) cm from the ®xated surface.
Phosphenes D, E and F were obtained in the normal hemi®eld of
blindsight subject GY with TMS applied 4 cm above and 1 cm to the right
of the inion. (b) Stimulation of MT‡/V5. Phosphenes A,B and C were
obtained in subject AC with TMS applied 3 cm above and 5 cm to the
right of the inion. Phosphenes D,E and F were obtained in subject GY
with TMS applied 3 cm above and 5.5 cm to the right of the inion. The
phosphenes are drawn according to a size scale in cm shown on the
ordinate. Their relative lateral displacement from ®xation is shown in
arbitrary units on the abscissa because of the different eccentricities of
phosphenes in the two subjects and the two visual areas.
testing session (out of three sessions) when MT‡/V5 was
being explored in the blindsighted subject GY, TMS
applied at 36 different points over a region of 5 3 5 cm
centred on the expected position of this area in the left
hemisphere did not yield a single phosphene. On the other,
normal hemisphere, in a smaller 3 3 3 cm grid centred on
MT‡/V5, there were 33 phosphene sites and only six sites,
at the periphery of the grid, where there was no phos-
phene. This difference between the hemispheres was
statistically signi®cant (÷2 ˆ 54.4; df ˆ 1; p , 0.001). The
phosphenes elicited in the peripherally blind subject,
despite being described as bright, like lightning, were
unusual in being con®ned to the central few degrees of the
contralateral visual hemi®eld. Too little is known about the
excitability of the visual cortex following almost instanta-
neous total and long-term peripheral blindness in adult-
hood to provide a convincing reason for the abnormality

3272 Vol 11 No 14 28 September 2000

MAGNETICALLY INDUCED PHOSPHENES
but EEG measures might indicate whether activity in
visual cortex is depressed. It should even be possible to
combine single-pulse TMS with high density multi-
electrode EEG monitoring to determine whether TMS is
only effective in eliciting activity in the cortex representing
the central retina, which has a high magni®cation factor
[28,29]. If the results on PS are con®rmed in other patients
it is tempting to think that in congenitally blind subjects
one possible answer to Molyneux's question [30] about the
nature of restored sight is that it would allow recognition
of at most only one object at a time because multiple
objects might not readily be spatially separable within the
compromised retinotopic map.
Whether area V1 is necessary for phenomenal visual
consciousness is contested [15,18,31]. For example, it has
been reported that high-contrast swiftly moving stimuli
can elicit qualia of visual motion within the otherwise
dense visual ®eld defects of patients with damage to striate
cortex [15]. However, in no instance where the damage to
V1 is veri®ably complete is it clear whether the patient has
experienced a phenomenal visual percept or is instead
aware of an event produced by a visual stimulus e.g. an
impression or feeling produced by, for example, a re¯exive
eye movement. The latter has been called blindsight type 2
by Weiskrantz [31]. The results of stimulating extrastriate
cortex with TMS in patient GY suggest that he really is
phenomenologically blind, i.e. no visual qualia, in the
hemisphere that lacks area V1 and when it is only MT‡/
V5 that is directly activated. It will be important to
investigate this in other patients whose ®eld defects and
blindsight arise from different cortical pathology.
CONCLUSION
Phosphenes were readily induced by TMS applied to
occipital cortex in normal subjects and in a peripherally
blind subject. Their absence when TMS was similarly
applied to extrastriate visual regions of a patient with
hemianopia caused by destruction of V1 and parts of V2
suggests that TMS can only induce conscious visual per-
cepts when V1/V2 are intact.
Acknowledgements: This work was supported by MRC grants G971/397/B and G9711247, by the Royal Society and by the Dr
Hadwen Trust. We are very grateful to PS and GY for their enthusiastic co-operation and to Dr Amanda Ellison for her help in
preparing the ®gures.
NEUROREPORT
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