Neural Plasticity and Consciousness (Figures Are Missing in This Version) Susan Hurley

NEURAL PLASTICITY AND CONSCIOUSNESS1
[figures are missing in this version]
Susan Hurley
< s.l.hurley@warwick.ac.uk >
Alva Noë
< anoe@cats.ucsc.edu>
Submitted to Biology and Philosophy for the special issue on Philosophy and the
Neurosciences edited by Kathleen Akins and Philip Gerrans
Why does neural activity in a particular area of cortex give rise to experience of
red, say, rather than green, or to visual experience rather than auditory? Why, for that
matter, does it have any conscious qualitative expression at all? These familiar questions
point to the explanatory gap between neural activity and qualities of conscious
experience.
In fact, these questions indicate that there are three different types of explanatory
gap for consciousness.2 There’s the absolute gap: Why should neural processes be
‘accompanied’ by any conscious experience at all? And there are two comparative gaps.
First, there’s the intermodal comparative gap: Why does certain neural activity give rise
to visual rather than auditory experience, say? Second, there’s the intramodal
comparative gap: Why does certain neural activity give rise to experience as of red, say,
rather than experience as of green?
It seems natural to adopt an inward focus in response to such explanatory gap

questions, to assume that they must be answered in terms of the intrinsic properties of the
neural correlates of consciousness. But there are well-known grounds for skepticism
1
For helpful discussions and comments, we are grateful to David Chalmers, Alan Cowey, Jeffrey
Gray, Mark Greenberg, Robert Hanna, Colin McGinn, Philip Pettit, Kim Plunkett, Nicholas
Rawlins, Evan Thompson, Michael Tooley, and Larry Weiskrantz. We are also grateful to
participants in the 2002 NEH Summer Institute on Consciousness and Intentionality in Santa
Cruz, California,. A.N. gratefully acknowledges the support of faculty research funds
granted by the University of California, Santa Cruz as well as that of a UC President's
Fellowship in the Humanities.
2
Compare Chalmers 1996, p. 5, who in effect distinguishes the absolute and comparative gaps,
but not the intermodal and intramodal comparative gaps.
Neural plasticity and consciousness:
about this strategy of response. Neural properties are qualitatively inscrutable. 3 If you
were to land in the visual system as a microscopic alien, you couldn’t tell, by looking
around at the local fireworks, whether experience was happening, or whether, if it was, it
was visual experience, or whether, given that it was visual, it was visual experience as of
something red. Mueller’s theory (1838) of “specific nerve energies” recognized this. On
his view, it is not the intrinsic character of the neural activity that makes it visual. Rather,
it is the fact that the neural activity is set up by stimulation of the retina, and not, say, the
cochlea. But this view still leaves the explanatory gap unbridged: why should
differences in the peripheral sources of input, leading to differences in the cortical
locations of the neural activity, make for the difference between what it is like to see and
what it is like to hear?
We suggest that an inward focus in response to explanatory gap worries can be

misleading. To find explanations of the qualitative character of experience, our gaze
should be extended outward, to the dynamic relations between brain, body, and world. In
this paper we apply this general strategy to the comparative explanatory gaps, both
intermodal and intramodal. We set aside the absolute gap, dividing in hopes of
conquering. We believe we can make progress by concentrating on the comparative
gaps; whether our approach will help to bridge the absolute gap is a further question.
We take our start from consideration of neural plasticity. This phenomenon

deserves serious attention from philosophers concerned with explanatory gaps, since it
reveals that neural activity in a given area can change its function and its qualitative
expression. We introduce a distinction between cortical dominance and cortical
deference4, and apply it to various examples in which input is rerouted either
intermodally or intramodally to nonstandard cortical targets. In some cases of rerouting
but not others, cortical activity ‘defers’ to the nonstandard sources of input and takes on
the qualitative expression typical of the new source.
This distinction is puzzling, and raises closely related empirical and philosophical
issues. What explains why qualitative character defers to nonstandard inputs in some
cases but not others? How does explanation of this difference address the comparative
explanatory gaps?5 After laying out the dominance/deference distinction, with both
intermodal and intramodal illustrations, we consider and criticize some possible
explanations of it. We then put forward a dynamic sensorimotor account of the
distinction. This promising hypothesis has the potential, if correct, to bridge the
comparative explanatory gaps.
3
“How pulses of water in pipes might give rise to toothaches is indeed entirely incomprehensible,
but no less so than how electro-chemical impulses along neurons can”. Maudlin 1989, 413.
4
Compare related discussions in von Melcher et al 2000; Merzenich 2000; Pallas 2000.
5
The distinction raises further philosophical issues as well, about how supervenience claims
relate to explanatory gaps. We address these in another article, in progress. In particular, we
claim that our dynamic sensorimotor account is compatible with neural supervenience claims, but
does more to address explanatory issues.
2
Whether or not our dynamic sensorimotor proposal turns out to be correct, our
main claim here is that the dominance/deference distinction is important and worthy of
further study, both empirical and philosophical. Explaining this distinction will help us
to understand how qualities of consciousness are related to the rest of the natural world.
1. The distinction introduced: cortical dominance vs. cortical deference
What happens when areas of cortex receive input from sensory sources that would not
normally project to those areas? When an area of cortex is activated by a new source,
what is it like for the subject? Is the qualitative character of the subject’s experience
determined by the area of cortex that is active, or by the source of input to it?
Empirical work on neural plasticity shows that it can go either way. In cases of
cortical dominance, cortical activation from a new peripheral input source gives rise to
experience with a qualitative character normally or previously associated with cortical
activity in that area. In such cases, we can say that cortical activity in a particular region
dominates, that is, it retains its ‘natural sign’ or normal qualitative expression. In cases of
cortical deference, in contrast, cortical activity in a given area appears to take its
qualitative expression from the character of its nonstandard or new input source. In these
cases, the qualitative expression of cortical activity in that area changes, deferring to the
new input source.
Cortical dominance is illustrated by phantom limb cases in which there appears to

be no change in the normal qualitative expression of activation of a given area of cortex,
despite change in the source of input. Normally, tactile inputs from arm and face map
onto adjacent cortical areas. After amputation of part of an arm, tactile inputs from the
face appear to invade deafferented cortex whose normal qualitative expression is a
feeling as of an arm being touched. When this area of cortex is activated from its new
source, the face, it retains its normal qualitative expression, the touch-to-arm feeling.
Thus, stroking the face is felt as the stroking of a phantom arm, as well as a stroking of
the face (Ramachandran and Hirstein 1998; see also Ramachandran and Blakeslee 1998,
28, 38).
Cortical deference is illustrated when congenitally blind persons read Braille.

During Braille reading, visual cortex is active. Moreover, stimulation of visual cortex via
transcranial magnetic stimulation (TMS), which in normal subjects distorts visual but not
tactile perceptions, in such blind subjects distorts tactile perceptions (Sadato et al, 1998,
Buchel et al, 1998, Cohen et al, 1997a, Sadato et al, 1996). In these subjects, visual
cortex seems not only to perform a tactile perceptual function, but also to have tactile
qualitative expression. Visual cortex defers qualitatively to its nonstandard tactile inputs.
We describe this and further illustrations of cortical deference below.
It may be natural to expect cortical dominance to be the norm. Perceptual scientists

may assume that for every type of experience, there is a locus in the brain such that
3
experience of that type supervenes on neural activity at that locus. Activity at such a
neural locus may be held to be necessary and/or sufficient to produce experience of the
relevant type however that activity is produced, whether by normal perceptual processes,
by direct stimulation, or by stimulation from a nonstandard source. Such a locus is
sometimes called a neural correlate of consciousness (NCC), or a bridge locus.6 It may
seem that if there is such a bridge locus for a given type of experience, then cortex should
dominate in the event of rerouting.
There are two points to note about this assumption of dominance as the norm.
First, an empirical point: cortex does not in fact always dominate, as we have noted. It
is important to recognize that cortical deference occurs as well. Second, a philosophical
point: the supervenience of types of experience on neural properties at bridge loci does
not entail dominance, but is equally compatible with deference, since neural activity at a
given locus can have different neural properties. We argue this point elsewhere, where
we claim that our account of the dominance/deference distinction, though compatible
with neural supervenience, addresses explanatory gaps in a way that neural supervenience
does not. To avoid distraction for purposes of this article, it is helpful to keep in mind
that we do not regard cortical deference, or our account of the dominance/deference
distinction, as threatening to the neural supervenience of experience.
Why does cortex defer in some cases but dominate in others? How could an
explanation of this difference contribute to bridging the comparative explanatory gaps?
This article makes an initial approach to these questions. We lay the groundwork in the
next two sections by giving a general schematization of the distinction and applying it to
various examples.
2. The distinction schematized
We schematize the distinction in terms of the relations between two mappings, one from
peripheral sources of input to cortical target areas, the other from cortical areas to
qualitative expressions.
Suppose there are two different peripheral (i.e. proximal) sources of afference or
input, A and B. A and B can be specified broadly to give an intermodal comparison, as in
6
Teller and Pugh (1983) introduced the term “bridge locus”. They write: “Most visual scientists
probably believe that there exists a set of neurons with visual system input, whose activities form
the immediate substrate of visual perception. We single out this one particular neural stage, with a
name: the bridge locus. The occurrence of a particular activity pattern in these bridge locus
neurons is necessary for the occurrence of a particular perceptual state; neural activity elsewhere
in the visual system is not necessary. The physical location of these neurons in the brain is of
course unknown. However, we feel that most visual scientists would agree that they are certainly
not in the retina. For if one could set up conditions for properly stimulating them in the absence
of the retina, the correlated perceptual state would presumably occur” (Teller and Pugh, 1983, p.
581). For discussion of this conception, see Noë and Thompson, forthcoming.)
4
visual stimulation (a pattern of light hitting the eye) vs. tactile stimulation (a pattern of
touch to the skin). Or they can be specified more narrowly to give an intramodal
comparison, such as tactile simulation to the face vs. tactile stimulation to the arm.
Suppose also there are two different target areas of cortex to which afference of kinds A
and B normally project, respectively. A normally projects to area 1, and B to area 2.
These areas are identified anatomically. The normal qualitative expression associated
with area 1 is the A-feeling and that associated with area 2 is the B-feeling.7
Now we hypothesize a rerouting of afference. First, suppose that area 2, to which B

normally projects, is deafferented so that the projection from B to 2 is eliminated. It may
be removed through surgical intervention (severance of the afferent channel, or by
removal of the limb or organ which is the peripheral source of the afference), or accident,
or it may be congenitally missing. Second, suppose that afference from source A now
somehow comes to project to area 2. A general question then arises.
Will activation of area 2 by afference with source A give rise to experiences or

feelings of the same type as activation of area 2 by afference with source B? That is, will
area 2 retain is normal qualitative expression, the B-feeling, when it is activated by the
new, nonstandard source? If so, then cortex dominates input source in the determination
of ‘what it is like’ for the subject.
Or, will activation of area 2 by afference with source A give rise instead to
sensation of the same type as activation of area 1 by afference with source A? That is,
will the qualitative expression of area 2 change from the B-feeling to the A-feeling,
reflecting the new source of afference? If so, then the qualitative character of the
subject’s experience depends in some way on the character of the source of input, rather
than just on whether cortical area 1 or 2 is active: cortex defers, apparently to the source
of input, in the determination of ‘what it is like’ for the subject.
FIGURE 1
Notice that cortical dominance thus schematized involves illusory

experience, while cortical deference involves veridical experience. If A and B are
stimulations of different sensory organs, such as tactile and visual stimulations, then we
can speak of intermodal dominance or deference. If A and B are stimulations within one
modality, such as touch to the face and to the arm, we can speak of intramodal
dominance or deference.
3. The distinction at work
7
It will sometimes be convenient to refer to the A-feeling as ‘A’, as in the looks-yellow-feeling
and ‘looks yellow’, in the discussion of synaesthesia in section 3 below.
5
We now explain how the distinction applies to the examples mentioned above and to
further examples.
Phantom limb cases provide examples of intramodal dominance. Suppose A and B

are touches to face and arm, respectively. Area 1 of somatosensory cortex normally
receives tactile afference from the face, while the adjacent area 2 of somatosensory cortex
normally receives tactile afference from the arm. The normal qualitative expression of
area 1 is the feeling of the face being touched, while that of area 2 is the feeling of the
arm being touched.
The arm is then lost, so there is no longer any afference from the arm reaching area
2. Instead, area 2 appears to be invaded by afference from the face, which now projects to
area 2 as well as the adjacent area 1. Touches to the face now activate both cortical areas
1 and 2. The question arises: will touches to the face feel only like touches to the face, or
will they also feel like touches to the missing ‘phantom’ arm? The answer is: in many
cases they feel also like touches to the missing arm (Ramachandran and Hirstein 1998,
Ramachandran and Blakeslee1998, 29, 33, 45).8 This suggests that activation of area 2
retains its normal qualitative expression, touch-to-arm feeling. Cortex dominates the new
source in determining what it is like for the subject.
Synaesthesia may provide examples of intermodal dominance. Color-graphemic

synaesthetes experience vivid sensations of color when reading or hearing words, letters,
or digits. Particular colors can be associated with particular letters or digits. There is
evidence that synaesthetic experience is automatic and truly perceptual, rather than
merely a matter of metaphorical association. For example, in a variant on the usual
Ishihara tests for color-blindness, synaesthetes who see numerals as colored were
presented with a collection of 2s and 5s such that the 5s were mirror images of the 2s.
The numerals were arranged so that the 5s made a pattern. Normal subjects could not see
the pattern. But it simply ‘popped out’ for the synaesthetes, since they saw the 2s and the
5s in different colors (Ramachandran and Hubbard 2001).
It is unclear whether synaesthesia results from nonstandard neural projections.

However, recent imaging work (Nunn et al 2002) has found that when synaesthetes with
colored hearing listen to spoken words, there is clear activation in an area of visual cortex
that has been identified as a color-experiencing area (Hadjikhani et al 1998; cf. Zeki et al.
1998; Tootell and Hadjikhani 1998). Activation of this area under the same conditions is
8
Such referral of touches to the face to the phantom arm can occur less than one day after
amputation, suggesting that the referral may be due to the unmasking of ordinarily silent inputs
rather than the sprouting of new axon terminals (Borsook et al 1998; Ramachadran and Rogers-
Ramachandran 1996, 385). However, the precise topographic mapping from facial stimulation to
phantom arm can become extremely disorganized and may be highly unstable over time,
suggesting that the relevant alterations in sensory processing may not be hardwired but rather be
mediated by an extensive and interconnected neural network with fluctuating synaptic strengths
(Knecht et al 1998; see also Halligan et al 1994).
6
not found in normal subjects. This suggests that language inputs get routed in
synaesthetes not just to their normal destinations but also to this area of visual cortex,
where they elicit experiences as of color. Cortical activation dominates over the source of
stimulation.
To spell out this suggestion about synaesthesia in terms of our schematized

distinction: Suppose that A is stimulation of auditory channels generated by the spoken
word “Wednesday” and B is a pattern of light entering the eye from a yellow visual
stimulus. Input from A activates area 1, whose normal qualitative expression is ‘sounds
like “Wednesday”’. Input from B activates area 2, whose normal qualitative expression is
‘looks yellow’. Here, there is no disconnection of area 2 from input B. Rather, there are
additional nonstandard neural projections: input from A also activates area 2, perhaps via
area 1, again eliciting area 2’s normal qualitative expression ‘looks yellow’. Since area 2
retains its normal qualitative expression ‘looks yellow’ even when activated by an input
from a different modality, synaesthesia thus interpreted would count as a case of
intermodal dominance.
Visual to auditory rerouting in ferrets provides an example of intermodal

deference. In newborn ferrets, nerves from the retina that would normally project to
visual thalamus and visual cortex have been surgically rerouted to project instead to
auditory thalamus and auditory cortex. Auditory areas are thus deprived of their normal
auditory inputs, and provided instead with inputs the source of which is visual
stimulation. Here, A and B are retinal and auditory inputs, and areas 1 and 2 are visual
and auditory cortex, respectively.
As a result of this rerouting, 2-dimensional retinotopic maps (similar to those

normally found in visual area V1) form in auditory cortex (Roe et al 1990, 1992). Some
single cells in auditory cortex develop orientation and direction selectivity normally
found in cells in visual cortex. Groups of cells in auditory cortex form orientation
modules and acquire some visual field properties (Roe et al 1990, 1992; Pallas and Sur
1993; Sur et al 1999; Sharma et al 2000; Elman et al 1996, 273ff. See also and compare
Rauschecker on auditory to visual rewiring in the cat). However, auditory cortex with
visual input did not make ectopic connections with visual cortex, but maintained its
connections with other auditory cortical areas (Pallas and Sur 1993).
Moreover, the visual information thus carried in rewired auditory cortex can be
made to mediate visual behavior. Unilaterally rewired ferrets are trained to respond
differently to light stimuli and sound stimuli presented to the non-rewired hemisphere.
Then, when light stimuli are presented to the rewired hemisphere, in the portion of the
visual field that is ‘seen’ only by this induced projection to auditory cortex, the rewired
ferrets respond as though they perceive the stimuli to be visual rather than auditory. The
researchers suggest that the functional specification and perceptual modality of a given
cortical area can be instructed to a significant extent by its extrinsic inputs; as a result, “...
The animals ‘see’ with what was their auditory cortex” (Mezernich 2000, 821; cf.
Carman et al 1992; von Melcher et al 2000). It is argued that the different characteristics
of input activity from specific sources (visual vs. auditory) generate not just
7
representational structure specific to that source but also source-specific sensory and
perceptual qualities. To put the point in our terminology, this recognition of cortical
deference is seen as a striking departure from the traditional and widely held assumption
cortical dominance (Mezernich 2000).
However, this work on ferrets, striking though it is, still leaves room for skepticism
about whether there has really been an alteration in the qualitative expression, as opposed
to representational and functional roles, of a given area of cortex (expressed to us, for
example, by Ned Block). Other work, on human subjects, leaves no such room for
skeptical manoeuver.
Early blind readers of Braille provide examples of intermodal deference in human

subjects. Brain imaging work on congenitally and early blind subjects reveals activation
in visual cortex during tactile tasks, including Braille reading, whereas normal controls
show deactivation (measured by PET scans) of visual cortex during tactile tasks (Sadato
et al 1996, 1998; see also Buchel 1998, Buchel et al 1998). 9 The researchers suggest that
the neuronal mechanisms of cross-modal plasticity include unmasking of normally silent
inputs (here, projections from tactile input to visual cortex), stabilization of normally
transient connections, and axonal sprouting. Referring back to our schematism, A and B
are here peripheral tactile and visual stimulations, and areas 1 and 2 are somatosensory
cortex and visual cortex, respectively.
The question arises how the early blind experience such activation of visual cortex:
what is its qualitative expression? This question is directly addressed by work that uses
TMS to produce transient interference with visual cortex activity during Braille reading.
In the early blind subjects, TMS applied to visual cortex produced both errors in Braille
reading and reports of tactile illusions (“missing dots”, “extra dots”, and “dots don’t
make sense”) (Cohen et al 1997a).10 By contrast, in normal subjects TMS to visual cortex
had no effect on tactile tasks or sensations, whereas similar stimulation is known to
disrupt the visual performance of normal subjects. In our terms, the qualitative expression
of area 1 in normal subjects is visual experience, while in these early blind subjects it is
tactile experience: the qualitative expression of activation of visual cortex here defers,
apparently to the source of input. The researchers view their work as supporting “the idea
that perceptions are dynamically determined by the characteristics of the sensory inputs
rather than only by the brain region that receives these inputs” (Cohen et al 1997a, 182;
cf. Maudlin 1989, 408).11
9
Other imaging work has shown that visual cortex of blind subjects is activated by sound
changes, when the task is to detect these changes (Kujala et al 2000).
10
Speech was unaffected by TMS, and subjects given a chance to correct their reports after TMS
had ended did not do so, suggesting that errors were not due to interference with speech output.
11
The perception of signing by the congenitally deaf provides another example suggestive of
intermodal deference. Auditory cortex lights up when some congenitally deaf persons receive
inputs from the visual periphery (where manual signing is perceived by those fluent at sign
8
4. How can the distinction be explained?
The fact that both dominance and deference occur needs explanation. Why do some cases
of neural rerouting result in dominance while others result in deference? What explains
whether qualitative expression goes one way or the other in particular cases? What
explains why activity in a certain cortical area is experienced as like this rather than like
that? To take one of our intramodal examples, why is cortical activity in a certain area
expressed as a touch-to-arm feeling rather than merely as a touch-to-face feeling? And in
the intermodal examples, why is cortical activity in a certain area expressed as tactile
rather than visual feeling, or as visual rather than auditory? These questions express
comparative explanatory gap issues, but they are open to empirical answer.
An initial hypothesis might be that we find dominance in cases that involve

intramodal plasticity and deference in intermodal cases. For example, the experience of
touch to the face as touch to the phantom limb is a case of dominance, and involves only
the sense of touch. By contrast, the experience of tactile distortion as a result of TMS
applied to visual cortex is a case of deference. This is a case of cross-modal plasticity in
which tactile inputs find a nonstandard target in visual cortex.
But the intermodal deference/intramodal dominance hypothesis is not satisfactory,

for at least two reasons.
First, it does not accommodate all the cases we’ve considered, even so far. We have
seen that some intermodal cases are plausibly regarded as examples of dominance, such
as synaesthesia. Moreover, there is evidence that the referral of sensations to phantom
limbs may be highly unstable over time, so even here they may be departures from strict
dominance. However, the hypothesis could be reformulated, so that intermodal rerouting
is necessary but not sufficient for deference.
But second, even if this reformulated hypothesis is correct, we’ll still want to know
why. Indeed, even if it were to turn out, on further reflection, that the
intermodal/intramodal distinction does coincide with the deference/dominance
distinction, we would still want to know why. The intermodal deference/intramodal
dominance hypothesis, if correct, would still not be explanatory, would be too close to
mere redescription of data (though it might provide clues to a more explanatory account).
A quite different suggestion turns on whether damage or rerouting has occurred

early or late. The hypothesis is that deference to nonstandard sources of input tends to
result from early rerouting of inputs to nonstandard targets, while dominance results from
late rerouting. The intuition behind this early deference/late dominance hypothesis is that
language, who focus on faces; Elman et al 1996, 299). It would be interesting to discover
whether in these cases TMS to auditory cortex produces visual distortions.
9
dominance is the norm for a mature brain with established qualitative expressions, while
deference results from early rerouting, before the brain has settled into a quality space.
However, consider the fact that patients born without arms may nevertheless have
phantom arms (Ramachandran and Blakeslee 1998, 40-42). The early deference/late
dominance hypothesis would predict that such patients should not experience the kind of
referred sensation experienced by amputees with phantoms, such as in the case of
dominance we described above in which a touch to the face is felt also as a touch to the
phantom arm. We do not know if referred sensation is found in congenital phantoms as
well as in late-acquired phantoms. Again however, if it were, the hypothesis could be
reformulated, so that early rerouting is necessary but not sufficient for deference. This
reformulation would also be prompted by synaesthesia, an apparent example of
dominance that starts very early in development (from as far back as synaesthetes can
remember).12
The reformulated prediction would then be that late rerouting should give rise to
cortical dominance. Here, the evidence at present seems less than decisive. Sadato et al
(1998) studied 8 blind subjects, 4 of whom were blind at birth and 4 of whom became
blind later (on average, at 8.5 years). “...[T]he critical point is that the primary visual
cortices of both early and later blind groups are activated during Braille reading....”,
irrespective of the onset of time of blindness (Sadato et al 1998, 1215; see and compare
Buchel 1998; Buchel et al 1998).
However, this study did not directly address the question of how the later blind
subjects experienced this activation by applying TMS to visual cortex, as did Cohen et al
(1997a). It would be interesting to know whether Sadato’s later blind subjects would
experience tactile distortions from TMS to visual cortex. Even if visual illusions also
resulted, tactile distortions from TMS to visual cortex in these subjects would show
cortical deference resulting from relatively late rerouting, contrary to the present
prediction.13
Cohen et al (1997b, 1999) studied blind subjects who lost their sight later in life
still (14-15 years) than Sadato’s subjects. But in these subjects, activation was not found
in visual cortex during Braille reading. Moreover, TMS to visual cortex did not disturb
Braille reading. However, since there is no imaging evidence of tactile to visual rerouting
in these subjects in the first place, the issue of dominance vs. deference is not raised by
Cohen’s late blind studies.
However, if the prediction that late rerouting leads to cortical dominance holds up,
further explanation would still be needed. We’d still want to know why: what is it about
early but not late rerouting that permits deference? We’d want an explanation at a deeper
12
Thanks here to Jeffrey Gray.
13
See also Kujala et al 2000 for further evidence on cross-modal cortical reorganization in the
mature brain.
10
level, one that sheds light on why qualitative expression can come to reflect the source of
input in early but not late rerouting.
Thus, both the intermodal deference/intramodal dominance and the early

deference/late dominance hypotheses are explanatorily shallow, even if they turn out to
contain elements of truth. What could give us a deeper level of explanation?
5. Intermodal plasticity without neural rerouting: adaptation to TVSS.
In order to move toward a deeper explanation, let’s consider some examples of

plasticity that do not involve neural rerouting. These examples are not captured by the
dominance/deference distinction as we have schematized it so far, because they involve
external rather than neural rerouting. What is altered in these cases is the external
relation between the objects of perception—the distal sources of input—and the
perceiver’s sensory organs—the peripheral source of inputs—, rather than the internal
relation between peripheral sources of input and cortical targets. Even so, these examples
illustrate a distinctive feature of cases of cortical deference, namely, changes in
qualitative expression as a result of rerouting. We’ll consider both intermodal and
intramodal examples of such external rerouting leading to deference. These examples
lead us to extend the dominance/deference distinction and motivate a dynamic
sensorimotor account of the distinction.
Consider first perceptual adaptation to a tactile-visual substitution system, which

involves an intermodal external rerouting. In a well-known series of studies by Bach-y-
Rita (1972, 1984, 1996; Sampaio et al. 2001), blind patients are outfitted with a tactile-
vision substitution system (TVSS).14 Vibrators or electrodes on the back, thigh or tongue
receive inputs from a camera fitted on the subject’s head or shoulder. Visual input to the
camera produces tactile stimulation of the skin, which in turn gives rise to activity in
somatosensory cortex, and to tactile experience.
After a period of adaptation (as short as a few minutes), subjects report perceptual
experiences that are distinctively non-tactile and quasi-visual. For example, objects are
reported to be perceived as arrayed at a distance from the body in space and as standing
in perceptible spatial relations such as “in front of” or “partially blocking the view of,”
etc. However, Bach-y-Rita emphasizes that the transition to quasi-visual perception
depends on the subject’s exercising active control of the camera (1984, 149). If the
camera is stationary, or if someone else controls it while the subject passively receives
tactile inputs from the camera, subjects report only tactile sensation.
In our schematism, A is here peripheral tactile input (patterns of vibration and

pressure on the skin by mechanical fingers) and B is peripheral visual input (patterns of
14
See also Arho et al 1999 on the possibility of an auditory-visual substitution system, as well as
Meijer (1992,) and also http://ourworld.compuserve.com/homepages/Peter_Meijer/).
11
light falling on the eye). Call cortical target area 1 somatosensory cortex and cortical
target area 2 visual cortex. Note that TVSS involves no rerouting from peripheral source
of input to cortical areas, either before or after perceptual adaptation. Peripheral tactile
input continues to stimulate cortical activity in somatosensory cortex throughout.
What has been rerouted is the external relationship between distal sources of visual
input, objects in space, and peripheral sources of tactile input. So, we can add to our
schematism a new lowest level of distal sources of inputs, A’ and B’. Let A’ be a distal
source of tactile input, an object that is touching the skin, and B’ be a distal source of
visual input, namely, an array of objects in space. TVSS effects a new external
intermodal mapping from distal sources of visual input to peripheral tactile inputs and on
to somatosensory cortex. As a result, the qualitative expression of somatosensory cortex
after adaptation appears to change intermodally, to take on the visual character of normal
qualitative expressions of visual cortex. Such a change in qualitative expression involves
no neural rerouting and so does not fit our original characterization of cortical deference.
That is, in contrast to the cases of deference we considered earlier, here there is no
apparent deference to a nonstandard source of peripheral input, for there is no change in
peripheral sources of input. However, because there is still a change in qualitative
expression of activity in given area of cortex, this case prompts us to extend our
characterization of deference to include cases of external rerouting.
FIGURE 2
Someone might be skeptical that there is genuinely an intermodal change in

qualitative expression of somatosensory cortex. However, there are good reasons to think
that, at the very least, the new qualitative expression of somatosensory cortex after
adaptation to TVSS is like vision (Noë 2002; O’Regan and Noë 2001a, b, c;). There are
structural respects in which tactile-vision is more like vision than it is like touch. In
vision, and in tactile-vision, we make perceptual contact with objects arrayed out before
us at a distance in space. Neither vision nor tactile-vision requires immediate physical
contact with perceptual objects. In contrast, touch is a perceptual mode that proceeds by
bringing a touched object into direct contact with the surfaces of the body. When a person
is first outfitted with TVSS, she feels tactile sensations on her back (say). When
adaptation is complete, she may still continue to feel sensations on the back (at least if
she were to attend to them), but she now also “feels” the presence of objects in space
around her.
There are other similarities as well. When you see an object, you make perceptual
contact only with the facing side or aspect of the object. You only see what is in view.
This fact has important dynamic sensorimotor implications. To bring more of an object
into view, it often suffices to move in relation to the object. This pattern of dependence of
what you perceive on what you do holds for tactile-vision in much the same way that it
holds for vision. In a similar vein, both vision and TVSS are governed by laws of
occlusion for which there is no analog in touch. You see, or TVSS-perceive, objects
around you only if they are not blocked “from view” by other opaque objects.
12
What makes tactile-vision visual? Here we cannot appeal to the proximal source of
stimulation, or to the fact that visual areas of the brain are activated. For TVSS-
perception is visual despite the fact that eyes and visual cortex are not directly activated.
The visual character of tactile vision stems from the way perceivers can acquire and use
practical knowledge of the common laws of sensorimotor contingency that vision and
TVSS share (O’Regan and Noë 2001a). For example, as you move closer to an object, its
apparent tactile-visual size increases, just as it would if you were seeing it. As you turn
to the left, objects in “view” swing to the right in your tactile-visual field, just as they
would if you were seeing them. As you move around an object, hidden portions of its
surface come into tactile-visual view, just as they would if you were seeing them. What it
is like to see is similar to what it is like to perceive by TVSS because seeing and tactile-
vision are similar ways of exploring the environment: they are governed by similar
sensorimotor constraints, draw on similar sensorimotor skills, and are directed toward
similar visual properties, including perspectivally available occlusion properties such as
apparent size and shape.
Recall that if the camera is stationary, or if someone else controls it, TVSS subjects
do not adapt to achieve vision-like experience of objects in space, but continue to report
only tactile sensation. Adaptation to TVSS does not occur unless the subject actively
controls the camera. This is what a dynamic sensorimotor approach would predict: active
movement is required in order for the subject to acquire practical knowledge of the
change from sensorimotor contingencies characteristic of touch to those characteristic of
vision and the ability to exploit this change skillfully.15
In TVSS, somatosensory cortex defers: but to what? Our original cases of

deference make it clear that qualitative expression cannot be explained just in terms of
the area of cortex activated. Extended deference in TVSS shows that it is not enough to
appeal in addition to the character of peripheral sources of input. In TVSS, there is an
intermodal change in the qualitative expression of somatosensory cortex, yet there is no
rerouting of peripheral inputs to somatosensory cortex. Rather, external rerouting
between distal and peripheral input sources induces the change in qualitative expression.
However, we do not suggest that this external rerouting in itself explains the
change in qualitative expression. Rather, the external rerouting effects a change in the
dynamic pattern of sensorimotor contingencies in which peripheral tactile inputs
participate, a change from the pattern characteristic of touch to the pattern characteristic
of vision. This change makes distinctively visual know-how and skills newly available to
the active subject. In TVSS, somatosensory cortex defers to distinctively visual qualities
of distal objects, but this deference is mediated by the perceiver’s new sensorimotor
skills. It is the perceiver’s practical knowledge of distinctively visual patterns of
sensorimotor contingency that give TVSS visual objects.
15
This is also argued in Hurley 1998, ch. 9, and O’Regan and Noë (2001, a, b, c; Noë 2002).
13
6. A dynamic sensorimotor hypothesis
These insights about TVSS can be generalized. According to our dynamic

sensorimotor hypothesis, changes in qualitative expression are to be explained not just in
terms of the properties of sensory inputs and of the brain region that receives them, but in
terms of dynamic patterns of interdependence between sensory stimulation and embodied
activity. What drives changes in qualitative expression of a given area of cortex, and
hence what explains the difference between dominance and deference, is not simply a
remapping from the sources of input, whether internal or external, to that area of cortex,
but rather higher-order changes, in relations between mappings from various different
sources of input to different areas of cortex and from cortex back out to effects on those
sources of input, which are in turn fed back to various areas of cortex. Note that there is
an essential and inextricable motor element to our account; intramodal, intermodal, and
sensorimotor relations are all potentially relevant. Qualitative adaptation depends on a
process of sensorimotor integration.
A general account of intermodal differences in qualitative expression is thus

suggested by a dynamic sensorimotor approach.16 Different sensory modalities are
governed by different, rich and systematic patterns of dynamic interdependence between
sensory stimulation and active movement. For example, to see something is to interact
with it in a way governed by the dynamic patterns of sensorimotor contingency
characteristic of vision, while to hear something is to interact with it in a different way,
governed by the different patterns of sensorimotor contingency characteristic of
audition.17 Your visual impressions are affected by eye movements and blinks in
specific, lawlike ways, while eye movements and blinks are irrelevant to the character of
your auditory impressions. Again, as you approach an object, visual field flow expands,
while as you withdraw visual field flow contracts. By contrast, as you approach the
source of a sound slowly, the amplitude of the auditory stimulus increases, while as you
withdraw the amplitude decreases. Perceivers are familiar with these distinctively
different dynamic patterns of sensorimotor contingency and know how to exploit them to
explore and negotiate their environments. According to the dynamic sensorimotor view,
perceptual experience is a skillful activity, in part constituted by such practical know-
how.
If the dynamic, sensorimotor view is to provide a bridge across comparative

explanatory gaps, it should not be explanatorily shallow. It must be explanatorily
satisfying. And it is.18 When it is brought to our attention that certain sensorimotor
16
Here we draw on Noë and O’Regan (2002), O’Regan and Noë (2001a, b, c) and on Hurley
1998a, especially chapter 9.
17
A more satisfyingly detailed characterization of the distinctive sensorimotor patterns charcteristic of
different modalities is given in O’Regan Noë 2001a, c.)
18
We do not accept the constraint that explanatory gaps concerning consciousness can only be
bridged by conceptual truths. See and cf. Levine 1993; van Gulick 1993, 146; Chalmers 1996.
14
contingencies are characteristic of vision, others of hearing, others of touch, there is an

‘aha!’ response. What we have learned does not have the character of a brute fact.
Rather, it is intelligible why it is like seeing rather than hearing to perceive in a way
governed by the sensorimotor contingencies characteristic of vision rather than those
characteristic of audition. It is not intuitively tempting to respond: “Yes, that correlation
of sensorimotor contingencies with vision may well hold, but why does it hold? Why do
those patterns of interdependence go with what it is like to see, rather than to hear or to
touch?” By contrast, if it is brought to our attention that activity in a certain brain area is
correlated with vision, we do indeed still want to ask: “ But why does brain activity there
go with what it is like to see, rather than to hear or touch?” Sensorimotor contingencies
are more promising in respect of intermodal comparative explanatory gaps than neural
correlates of consciousness.
Readers should keep in mind, when consulting their own intuitions for the ‘aha!’
response, that we are here addressing comparative rather than absolute explanatory gaps.
For present purposes the relevant question is not: when we describe certain sensorimotor
patterns, do we make it compelling that there is something it is like for the agent at all? It
is rather: assuming that there is something it is like for the agent, when we describe the
sensorimotor patterns characteristic of, say, vision by contrast with those characteristic of
audition, does our description make it compelling that it is like seeing rather than like
hearing for the agent?
Because TVSS effects a change from patterns of sensorimotor contingencies

characteristic of touch to patterns characteristic of vision, a dynamic sensorimotor view
predicts deference and an intermodal change in qualitative expression of somatosensory
cortex in this case. But can a dynamic sensorimotor approach be extended to intramodal
differences of qualitative expression?
It might be suggested that the dynamic sensorimotor hypothesis is on stronger

ground in predicting intermodal deference than intramodal deference, since intermodal
rerouting results in larger-scale, more global changes in sensorimotor contingencies than
does intramodal rerouting. For example, changes in the sensorimotor contingencies
between touching the face and touching the arm, or between looking at something red
and looking at something green, are relatively minor, restricted and subtle, compared
with those between looking at something and listening to it, or between looking at
something and touching it. If this suggestion were to hold up, then perhaps the dynamic
sensorimotor hypothesis could provide a deeper level of explanation for the intermodal
deference/intramodal dominance hypothesis considered above, which we said was
explanatorily shallow.
However, it does not hold up. In principle, the dynamic sensorimotor account
applies to intramodal as well as intermodal differences in qualitative character. Perhaps
For a different kind of attempt to bridge the explanatory gaps in sensorimotor terms, see
O’Regan, Myin and Noë, submitted and O’Regan and Noë 2001c).
15
sensorimotor contingencies change in subtler, less global ways within modalities than
between modalities than within modalities. But at the qualitative level also, there is a
subtler difference between seeing red and seeing green than there is between seeing and
touching, or between seeing and hearing. There are nevertheless significant differences
in sensorimotor contingencies between qualities within one modality.19 So the dynamic
sensorimotor hypothesis does not necessarily predict intramodal dominance. And this is
all to the good, since we find striking intramodal deference when we consider adaptation
to goggles, to which we now turn.
7. Intramodal plasticity without neural rerouting: reversing goggles
Consider the results of experiments on the long-term effects of wearing left-right

reversing goggles (Taylor 1962; see also Stratton 1897; Kohler 1951). Here again there is
an external rather than a neural rerouting. And here it results in intramodal deference.
The initial effect of left-right reversing goggles is to produce a left-right reversal

in perceptual content. The goggle wearer’s right hand looks as if it is on the left and vice
versa. The explanation is straightforward. Normally, without goggles, a rightward distal
object would produce certain peripheral visual inputs that would in turn project to a
certain area of visual cortex, which we’ll call right visual cortex, or RV-cortex. 20 The
normal qualitative expression of RV-cortex is ‘looks rightward’. Similarly, a leftward
distal object would produce different peripheral visual inputs that would project to left
visual cortex, or LV-cortex, the normal qualitative expression of which is ‘looks
leftward’. The goggles effect an external intramodal rerouting: now a rightward distal
object produces peripheral visual inputs that project to LV-cortex, and a leftward distal
object produces peripheral visual inputs that project to RV-cortex. So the goggles
initially make the right hand look as if it is on the left and vice versa. Notice that again
there is no internal neural rerouting of the projections from peripheral inputs to cortical
targets; these are unchanged.
FIGURE 3
However, putting on the goggles initially disrupts vision dramatically. Movements

of eyes and head and body give rise to surprising, unanticipated, confusing sensory
19
This may hold even for differences in color (O’Regan and Noë 2001a; O’Regan, Myin & Noë
submited; Noë, forthcoming; see also Pettit in press). Also relevant is the fact that there are
asymmetries in color relationships that prevent inversions from working smoothly. SeeThompson
1995; Myin 2001; van Gulick 1993, 144-145.
20
Note that this terminology can mislead. RV-cortex is the part of visual cortex that normally subserves the
experience as some something being visually on the right. It is not the anatomically right part of the visual
cortex.
16
effects. For example, when you rotate your head, the world, dizzyingly, seems to move
around you. It used to be that you had to move your head leftward to bring leftward
objects more into view, but that no longer works. When you try to move your right hand
rightward, it feels as if your right hand is moving rightward, but it looks as if your left
hand is moving leftward. Moving your right hand still activates right proprioceptive
cortex, the qualitative expression of which remains ‘feels right’, even though it looks
leftward. So there is a conflict between vision and proprioception when they are co-
stimulated by the same right hand movement. The external intramodal visual rerouting
effected by the goggles results in an intermodal conflict between vision and
proprioception, such that proprioception is veridical and vision is not.
FIGURE 4
Thus, movements of your eyes, head, limbs, and whole body quickly demonstrate
that the old patterns of sensorimotor contingency no longer apply. While the mapping
from distal objects, such as your moving hand, to visual cortex has altered, the mapping
from your moving hand to proprioceptive cortex has not altered. Nor has the mapping
from motor cortex to your moving hand. Since the mapping to visual cortex has changed,
but not the mappings to proprioceptive cortex or from motor cortex, the higher-order
relations between these mappings, or sensorimotor contingencies, have changed.
According to our view, for you to experience something as visually leftward is for
it to present itself to you as occupying a certain position in a space of sensorimotor
possibilities, through which you have the skills to navigate. For example, when
something looks to you as though it is on the left, you know how to move your eyes, or
turn your head, to bring the thing more into view, you know how to raise your arm and
hand in order to block it from view, or to rearrange things that block your view of it so
that they no longer do, and so on. When you put this goggles on, you initially lose this
know-how.
If perceptual experience depends on the perceiver’s sensorimotor know-how, then

one way to interfere with perceptual experience would be to alter the dynamic patterns of
sensorimotor contingency his know-how exploits. As we’ve seen, this is just what putting
on left-right reversing goggles does. However, as the goggle-wearer learns how to
navigate through the new space of sensorimotor contingencies, our view predicts that his
perceptual experience should adapt accordingly.
This prediction is born out by Taylor’s (1962) description of the experience of a

left-right reversing goggle subject, who moved freely through and interacted with his
environment over a long period. At first his vision was disrupted as described above.
But over time, in addition to seeing a leftward object on the right, he began to see a
ghostly version of the same object on the left in its true position. According to Taylor,
his visual experience eventually adapted, so that leftward objects came once more to look
as if they were on the left and not on the right at all. After adaptation, vision was again
veridical, and did not conflict with proprioception.
17
FIGURE 5
Taylor emphasizes that this result was achieved as a result of a rigorous and
extended training program during which the goggle wearer engaged in intensive
sensorimotor interactions with his environment. Visual adaptation was modular, and
reflected specific practice sessions. For example, after lots of bike riding with goggles,
buildings on the left came to look leftward, while the writing on signs on those buildings
was still left-right reversed. After subsequent practice at reading with goggles, the
writing appeared normal as well. Just as intermodal adaptation to TVSS depends on the
active control and sensorimotor interactions, so intramodal visual adaptation to left-right
reversing goggles depends on the subject’s activity. 21
Like TVSS, Taylor’s goggle-adaptation result fits our extended characterization of

deference: the qualitative expression of activity in RV-cortex changes as a result of
external rerouting. In this case, by contrast with TVSS, the change is intramodal: from
‘looking rightward’ when the goggles are first put on to ‘looking leftward’ after
adaptation.
Again, we ask: deference to what? The change in qualitative expression cannot be

explained in terms of a change in the peripheral source of visual inputs, as there has been
no rerouting between peripheral inputs and their cortical targets and so there is no change
in peripheral input source. Rather, the change in qualitative expression is induced by
external rerouting between distal and peripheral input sources.
Our response is again that the external rerouting effects a change in the dynamic
pattern of sensorimotor contingencies in which peripheral inputs participate. As a result,
the subject temporarily loses the know-how on which his visual experience depends.
But with practice he regains it, having learned a new way of navigating skillfully through
a restructured space of sensorimotor contingencies with which he eventually has become
familiar. Leftward objects once more present themselves to him as in terms of a certain
characteristic trajectory in this multidimensional space, as related to changing
proprioceptive and motor mappings in characteristic ways. Now, when he moves his
head leftward, it does bring objects that look (and are) leftward more into view. When he
tries to move his right hand rightward, it feels as if his right hand is moving rightward
and it also looks as if his right hand is moving rightward. Vision here defers to the true
position of distal objects, but this deference is mediated by the perceiver’s reconfigured
21
Taylor reports that one of his long-term subjects experienced no aftereffect when removing or
reinstating the goggles (while riding a bicycle!). This suggests a very striking variability in
qualitative expression. With goggles on, left arm stimulates RV-cortex, and looks leftward. With
goggles off, right arm stimulates RV-cortex, and looks rightward. Here, the qualitative
expression of activity in RV-cortex would vary between ‘looks rightward and ‘looks leftward’.
The subject has acquired know-how in relation to both sets of patterns of sensorimotor
contingencies, with and without goggles, and switches between them seamlessly. See Hurley
1998a, ch. 9, for further discussion.
18
dynamic patterns of sensorimotor skills in relation to such objects. The perceiver’s

practical knowledge of the dynamic patterns of sensorimotor contingencies characteristic
of seeing leftward objects, within the larger space of sensorimotor contingencies that
characterize vision generally, is what makes them look leftward.
8. A challenge.
Charles Harris (1965, 1980) questions whether there is genuinely visual adaptation
to reversing goggles of the kind claimed by Taylor. Some long-term goggle wearers may
judge that their visual experience has veridicalized, that leftward objects eventually come
to look leftward even while wearing goggles. But Harris suggests that any such
judgements are mistaken. He explains two aspects of adaptation to reversing goggles.
He argues, first, that adaptation to the goggles is the result not of vision righting
itself, but rather of the adaptation of proprioception to reversed vision. The right hand,
which looks as if it is on the left, now comes to feel (proprioceptively) as if it is on the
left too. Behavioral dispositions are also reversed and so brought into accord with
reversed visual experience, thus eliminating the intermodal discord and confusion
induced initially by putting on the goggles. Harris says that
…so many visual judgments and visually guided behaviors are affected [by
processes of adaptation] that one could talk about a modification of visual
perception, as long as one bears in mind that here too what is actually modified is
the interpretation of nonvisual information about positions of body parts” (1980,
113).
In his view, if perceivers interpret the adaptive change in their experience as a change in
visual experience, their judgments about their own experience are wrong; nonvisual
experience rather than visual experience has really adapted.
Secondly, Harris argues that a process of familiarization can explain why even
though proprioception, not vision, has really adapted, it nevertheless can seem to subjects
that vision has reverted to normal. Just as with practice mirror-writing can come to seem
“normal,” so, with practice, reversed vision can come to seem normal and familiar. But
Harris suggests that visual experience really remains left-right reversed even though it
comes to seem normal. Harris’ hypothesis is a kind of left-right positional version of an
inverted spectrum hypothesis.
If Harris is right, then proprioception adapts intramodally, not vision. We don’t get
a change in the visual qualitative expression of RV-cortex, as Taylor’s work suggests.
Rather, thanks to the power of vision to influence proprioception, we get a change in the
proprioceptive qualitative expression of neural activity in what we can call right-
proprioceptive cortex (RP-cortex). RP-cortex changes its qualitative expression from
‘feels rightward’ to ‘feels leftward’. Note that here again there has been no rerouting
19
between sources of proprioceptive inputs and their targets in proprioceptive cortex. The
right hand projects to RP-cortex both before and after adaptation.
So, if Harris is right, we still have a kind of deference. RP-cortex defers; but to
what? Here again it defers not to a new source of proprioceptive input (there is no new
source), but to a new co-stimulation relationship with visual cortex. Actions are intended,
felt, and seen, in relation to the environment, giving rise to patterns of neural activity.
The goggles alter the way neural patterns are coordinated by action on and in the world.
An intramodal external rerouting in vision generates an intermodal conflict between
vision and proprioception, which in turn induces an intramodal change in proprioceptive
qualitative expression that resolves the conflict, relative to the power of vision.22 The
illusion is thus compounded, as proprioception inherits the illusion the goggles have
perpetrated on vision. If Harris is right, deference here cannot be deference to the true
position of distal objects. But nevertheless, what would drive the change in qualitative
expression of LP-cortex, and could eventually give rise to the secondary illusion that
visual experience is re-inverted, would still be a change in the dynamic patterns of
sensorimotor contingency among intentional movement, proprioception and vision.
We are skeptical about Harris’ view. We will consider his view as a rival to
Taylor’s in order to spell out our misgivings about it. Consider why proprioception
might bend to vision, as Harris argues his experiments show it to do? Why might the new
co-stimulation relation induce illusory proprioceptive deference rather than veridical
visual deference? Suppose, for the sake of argument, that there are two distinct
qualitative possibilities, veridical Taylor-type visual adaptation and illusory Harris-type
proprioceptive adaptation with the secondary adaptation such that vision again seems
normal. Then it is an empirical question which form of adaptation in fact occurs.
Perhaps it is even possible that adaptation occurs one way under certain conditions and
the other way under other conditions. Either way, cortical activity somewhere changes its
qualitative expression intramodally, driven by the changes in dynamic patterns of
22
Visual capture is a widespread phenomenon. It is exhibited in such phenomena as the
ventriloguist effect, in which we hear the voice coming from the mouth of the dummy because we
see things that way, and in the McGurk effect, in which visual information about lip-movements
trumps accoustic information in determing heard phonemes. An interesting case has recently been
reported (Botvinik and Cohen 1998). Subjects were "seated with their left arm resting upon a
small table. A standing screen was positioned beside the arm to hide it from the subject's view
and a life-sized rubber model of a left hand and arm was placed on the table directly in front of
the subject. The subject sat with eyes fixed on the artificial hand while we used two small
paintbrushes to stroke the rubber hand and the subject's hidden hand, synchronising the timing of
the brushing as closely as possible" (Botvinik and Cohen 1998, 756). After a short interval
subjects have the distinct and unmistakable feeling that they sense the stroking and tapping in the
visible rubber hand and not in the hand which is in fact being touched. Further tests show that if
you ask subjects with eyes closed now to point to the left hand with the hidden hand, their
pointings, after experience of the illusion, are displaced toward the rubber hand.
20
sensorimotor contingency. So either way, adaptation to reversing goggles illustrates

intramodal deference.
It would be nice to stop there. But we have yet to pin down, in sensorimotor
terms, which intramodal change, in vision or in proprioception, has occurred in response
to changes in dynamic patterns of sensorimotor contingency. What does a dynamic
sensorimotor approach predict about which modality will, as it were, host the adaptive
change? Does it predict the kind of deference illustrated by Taylor’s view or that
illustrated by Harris’ view?
As we saw in the previous section, the dynamic sensorimotor approach gives a

plausible characterization of intermodal differences. Vision and hearing and touch each
have distinctive sensorimotor relations to bodily movement and to each other. But for
one animal, the dynamic patterns of sensorimotor contingency governing different
simultaneously operative modalities are interpenetrating, superposed on the same
organism and its neural system operating in a given environment. We propose that
qualities of experience reflect practical knowledge of higher-order sensorimotor patterns.
But when a change occurs in the higher-order sensorimotor patterns in which several
modalities participate and the animal comes to acquire know-how in relation to the new
pattern, how does a dynamic sensorimotor approach attribute the corresponding
qualitative adaptation to one modality or another? For example, when a change occurs in
the normal relations between visual and proprioceptive inputs and motor outputs, on what
basis does a dynamic sensorimotor approach predict that visual or proprioceptive
experience will adapt with renormalization?
To answer this question, we need to consider what is needed for sensorimotor

know-how to be regained by someone wearing left-right reversing goggles. First, you
need, presumably, to be in intermodal sensorimotor harmony, so that vision, hearing,
touch, proprioception, and motor intentions are integrated and not in conflict. Intermodal
harmony may demand intramodal adaptation, but it may be possible to satisfy this
demand in more than one way, as the difference between Taylor’s and Harris’s views
suggests. It can be satisfied so that intermodal harmony is veridical, as in Taylor’s
view—your right hand looks and feels rightward--or is illusory, as in Harris’ view—you
right hand both looks and feels leftward.
Second, however, sensorimotor know-how also requires you to be able to negotiate

your public environment successfully. Now recall the two parts of Harris’ view. The
primary aspect of adaptation of proprioception to vision means that your right hand
comes at the same time to feel as well as look leftward. But there is also a secondary
aspect, in that universal mirror reversal comes at the same time to look normal and
familiar. If adaptation only had the primary aspect, your environmental know-how would
be compromised by the primary illusion. Here, for example, is how things might go
wrong with only the primary adaptation: When your wedding ring is on your left hand, it
doesn’t look right. So you put your wedding ring on your right hand, which looks and
feels leftward. When someone you don’t like at all indicates his romantic interest in you,
21
you hold your right hand, stylishly, near to his face, hoping to stop him in his tracks. His
response is not at all what you intended.
Can the secondary aspect of adaptation save you from such blunders? While it is
not completely clear how Harris understands this secondary aspect, it could be interpreted
as a kind of higher-order illusion: because everything rightward seems leftward, you
come to interpret seeming leftward, wrongly, as seeming rightward. Whether or not this
is the correct interpretation of Harris, it is worth spelling out why we are skeptical of this
dual-illusion hypothesis.
The dual illusion hypothesis under scrutiny is that while rightward things really
look and feel leftward to you, they come to seem to look and feel rightward. So the true
qualities of your experience are no longer self-evident to you. Both the primary and
secondary adaptations generate illusions. By canceling out the primary illusion, such a
secondary illusion would at least restore your know-how in relation to your environment.
It would, for example, save you from the kind of wedding ring blunder just described, as
your right hand would come with ‘normalization’ to seem to you to look and feel
rightward, even though it still really looks and feels leftward. So your wedding ring
would stay where it belongs.
FIGURE 6
The primary aspect of adaptation postulated by Harris without such a secondary

illusion would not restore dynamic sensorimotor know-how; it would leave you open to
blunders. So a dynamic sensorimotor view will not predict the primary illusion without
the secondary illusion to cancel it out for practical purposes. But both the doubly illusory
adaptation and Taylor’s veridical adaptation will equally restore intermodal and
environmental know-how in the context of the new pattern of contingencies imposed by
the goggles. Does our account favor one or the other?
Note that the dual-illusion hypothesis conflicts with the claim that qualities of
experience are self-evident. In effect, the subject loses knowledge of the qualities of his
own experience, in regaining sensorimotor know-how. So any independent arguments
for self-evidence would favor Taylor’s hypothesis over the dual- illusion hypothesis.23
23
See Hurley 1998, ch. 4; cf. Chalmers 1996, ch. 5, on the paradox of phenomenal judgment.
While we cannot pursue the details here, we note that two distinctions are critical in assessing the
relevance of Chalmers’ arguments to present concerns.
First, self-evidence should be explicitly distinguished from incorrigibility: if experience
must be self-evident, there is an entailment from the character of experience to judgment. But the
reverse entailment does not follow: someone’s judgment the character of his experience need not
be incorrigible, even if his experience is self-evident. Hurley 1998 denies incorrigibility, but
defends a version of self-evidence. Self-evidence is all we need to rule out Harris’ dual-illusion
adaptation, not incorrigibility. Chalmers also denies incorrigibility, insisting that our judgements
about our own consciousness do not entail the truths of such judgments; a zombie could judge
himself to be conscious. His position on self-evidence is more complex; see pp. 97, 205ff.
22
Our view leads to the same conclusion by a different route. It predicts that the two
postulated illusions would cancel one another out qualitatively as well as practically.
The dual- illusion hypothesis relies on the bare idea that this is what it is really like
qualitatively for the subject, even though it does not seem to him that this is what is really
like. But our view predicts that if there were no difference in sensorimotor know-how
between the dual- illusion view and Taylor’s view, then there would be no qualitative
difference either.
It may be natural to presuppose that experience is either visual or not visual

intrinsically, or at least in virtue of something other than dynamic sensorimotor know-
how. But to do so in effect begs what is an empirical question against our account of
intermodal differences in qualities of experience. Why is this presupposition natural?
Perhaps because it is natural to assume intramodal cortical dominance: if RV-cortex is
still active, then visual experience must still be of something rightward. But certainly
Harris would not be entitled to this assumption, by his own lights, since if his view is
correct then proprioceptive cortex defers intramodally, even if visual cortex dominates, as
explained above. Moreover, we have argued that cortical dominance cannot in general be
assumed. It is an empirical question whether cortex dominates or defers, and the
evidence from cases of neural plasticity suggests it can do either.
To sum up our response to Harris’ position: If adaptation involves illusory

proprioception, as Harris suggests, then a canceling secondary adaptation or illusion is
required to restore full know-how. If there is a qualitative difference between adaptation
on the dual illusion interpretation and adaptation on Taylor’s veridical interpretation, then
the former implies intramodal cortical deference just as much as the latter does.
Nevertheless, we suggest that adaptation on the dual illusion view would not be
qualitatively different from adaptation on Taylor’s view, since there is no difference in
sensorimotor know-how. The two illusions would cancel out qualitatively as well as
practically.
Note that this response is not put forward on a priori verificationist grounds, along
the lines of denying a priori the qualitative possibility of an inverted spectrum despite
complete practical adaptation. Rather, it is an empirical prediction based on a theory
supported by evidence in other cases (see also Cole 1990). Indeed, it could prove
difficult to verify this particular prediction after adaptation, although the dynamic
sensorimotor theory is in general open to empirical assessment, as we have explained. 24
Second, we should distinguish absolute from comparative issues: self-evidence and

incorrigibility may be more or less plausible for absolute or comparative issues, and our concern
here is with comparative issues, not zombies.
24
Moreover, it is difficult to see how Harris’s view could explain for the visual doubling that
Taylor’s subject reported before adaptation was complete: “the simultaneous perception of an
object and its mirror image, although…the chair on the right [its true position] was rather ghost-
like” (1962, 202, 206).
23
9. The dominance/deference distinction explained.
We will now review in general terms how the cases we have considered constrain
explanation of the dominance/deference distinction and move us toward a dynamic
sensorimotor account.
Deference resulting from neural rerouting, in our original cases, shows that
qualitative expression cannot be explained just in terms of the area of cortex activated.
Deference resulting from external rather than neural rerouting, as with adaptation to
TVSS and reversing goggles, shows that it is not enough to appeal in addition to the
character of peripheral sources of input, since there is no rerouting from peripheral inputs
to cortex in these cases. Rather, rerouting, whether internal or external, changes the
pattern of dynamic sensorimotor contingencies in which given areas of cortex participate.
Recall the way changes in qualitative expression in TVSS and goggle adaptation depend
on the agents’ active control and exploratory movement in their environment. As a result
of rerouting plus a subject’s activity, a global dynamic pattern characteristic of a specific
modality, or a more local pattern characteristic of a specific quality within a modality,
may be newly established or relocated to new neural pathways. And a given area of
cortex may find itself newly participating in and integrated into such dynamic patterns of
sensorimotor contingency. Such characteristic patterns govern agents’ skillful
perceptual activities in their environments, their perceptual know-how. Changes in the
neural paths of such characteristic sensorimotor patterns after rerouting can disrupt
agents’ perceptual know-how and with it the qualitative character of experience, but with
practice such know-how can be reacquired. Deference reflects agents’ know-how in
relation to dynamic patterns of sensorimotor contingencies that are characteristic of
specific modalities or qualities, but which use nonstandard neural paths that include areas
of cortex that would normally participate in different sensorimotor patterns. We suggest
that the dominance/deference distinction can be explained in terms of such skill-
governing sensorimotor patterns in both intermodal and intramodal cases.
Our account predicts deference where two general conditions are met: first, where
perceptual experience of the kinds in question normally arises out of distinct patterns of
sensorimotor contingency, which are systematically transformed by rerouting, and
second, where the agent is able to explore and learn the new operative contingencies and
their relations to the old ones. Thus, dominance should result when the second condition
is not met because the agent is relatively passive, or where the first condition is not met
because particular kinds of rerouted input give rise to ‘dangling’ cortical activity, not
substantially tied into a dynamic pattern of sensorimotor contingencies, of cross-modal
and feedback relationships. Such dangling activity latter could nevertheless generate a
limiting case of perceptual experience. There may be a spectrum of degrees of richness
and complexity in patterns of sensorimotor contingency, with a kind of nearly-null case at
one extreme whereby a source of input stimulates only one cortical area and is unaffected
by motor activity. These two conditions for dominance are connected: inactivity by the
24
subject may leave a new input dangling, until activity ties it in to the network of
sensorimotor contingencies through co-stimulation and feedback.
Mappings from different input sources to cortex are affected in different ways and
to different degrees by motor activity, and the way and degree to which they are affected
can be altered by rerouting. Rerouting of inputs that are affected by motor activity should
generally produce changes in dynamic patterns of sensorimotor interdependence. Thus,
on this approach, such rerouting should generally produce deference. Deference is the
norm, and dominance is the exception that needs to be explained, as a kind of limiting
case.
How do these predictions play back onto our original cases? Intermodal deference
in the ferret and Braille cases are straightforwardly predicted, since neither condition
predicting dominance is met: the agents are active, and the rerouted input does not dangle
but is tied into a network of sensorimotor contingencies. It would be interesting to know
what would happen if TMS were applied to visual cortex when no Braille reading activity
by the subject is taking place: would it still generate a tactile sensation?
By contrast, dominance could be predicted in the phantom referral case on the basis
that the rerouted input from face-stroking to cortex that used to signal touch to arm is
dangling as a result of inactivity in relation to this specific input. Why? Because the
experimenter, not the subject himself, does the face stroking, so no feedback is set up. If
the subject were to stroke his or her own face, while also watching in a mirror, rather than
having the experimenter stroke it, the dynamic sensorimotor hypothesis would predict
that qualitative expression would defer. Such self-stroking would make available a new
set of relevant contingencies.
This prediction is in line with Ramachandran’s well-known mirror-box results

(Ramachandran and Blakeslee, 1998, 47ff; Ramachandran and Rogers-Ramachandran
1996; Ramachandran et al 1995). Ramachandran’s patient had an immobilized phantom
hand, paralyzed in a painful position for ten years since he had lost his limb.
Ramachandran used a box in which mirrors had been positioned to create an illusion of
the patient’s intact hand in the felt position of his phantom hand. The patient was asked
to try to move both his hands simultaneously. When he moved his intact hand and saw it
move in the mirrors, in the felt position of his phantom hand, he felt his phantom hand
move as well. Moreover, the movement in his phantom relieved the pain in his phantom.
The mirrors created illusory visual feedback of phantom movement, harking back to the
sensorimotor contingencies familiar to the subject from before the loss of the limb.
Experience changed accordingly. Ramachandran suggests that when the brain sends out
motor commands for movement, and copies of these commands, but gets no
corresponding feedback of actual arm movement because the arm is missing, it learns that
the arm does not move, that it is paralyzed. The illusory feedback created by the mirror
box allows it temporarily to unlearn paralysis.
How then, Ramachandran asks, can we understand the persisting experience of

phantom limb movements in congenital phantoms? He suggests that a normal adult has a
25
lifetime of practical familiarity with what in our terminology are dynamic patterns of
sensorimotor contingency. These are missing after amputation; the brain’s normal
‘expectations’ of sensorimotor feedback are ‘disappointed’, so adaptation is needed. As a
result the phantom may freeze or even disappear over time. But movement in a
congenital phantom may persist indefinitely because the congenital absence of a limb to
provide co-stimulation and feedback relationships between various modalities and motor
activity means that there are no normal expectations of sensorimotor feedback from such
a limb to be disappointed. So no adaptation is called for. In effect, the part of the innate
body image corresponding to the phantom is never overwritten but dangles, disconnected
from the network of sensorimotor contingencies into which it would normally be
integrated (See Ramachandran and Blakeslee 1998, 57).
A challenge for the dynamic sensorimotor view is to explain apparent intermodal

dominance in synaesthesia. We do not have an account of this worked out. But here it is
interesting to note that while synaesthetes are like normal subjects in displaying cross-
modal priming effects for consciously perceived colors, they do not display the covert
cross-modal priming effects shown by normal subjects. This may provide a clue of use to
the dynamic sensorimotor hypothesis, suggesting a degree of ‘dangle’, or disconnection
of synaesthetic color perceptions from the usual network of cross modal contingencies. 25
25
In particular, colored graphemic synaesthetic perception does not have all the properties of
normal color perception. Synaesthetically induced colors give rise to cross-modal priming
effects, as do colors perceived by normal subjects. However, synaesthetically induced colors do
not give rise to covert cross-modal priming effects, while colors perceived by normal subjects do
(Mattingly et al 2001).
Here is what this means, operationally. Normal subjects asked to name the color of ink in
which a word is written show longer reaction times when the word spells the name of an
incongruent color: for example, if the word ‘red’ is printed in blue ink, they will take longer to
say ‘blue’ than they will if the word ‘blue’ is printed in blue ink (the Stroop effect). Synaesthetes
display a synaesthetic version of a Stroop effect: if they are asked to judge the physical color of a
letter that induces a different synaesthetic color, their reaction times are slowed (whereas that of
normal subjects are not). If they are shown a letter prime that induces experience of a certain
synaesthetic color, and then shown a color patch and asked to name its color, they are slower
when the induced synaesthetic color of the letter prime is different from the color of the color
patch. Syneasthetic and normal colors are thus similar in respect of Stroop priming effects, where
subjects are conscious of the colors in question.
However, normal subjects also display covert priming effects: a letter is briefly presented
and masked so that subjects are not consciously aware of having seen it, and then asked to name a
subsequently presented letter. Normal subjects are slower to name the subsequent letter when
the masked prime was a different letter, even though they are not consciously aware of the
masked prime. And synaesthetes show this same covert priming effect on letter recognition,
displaying unconscious processing of the letter prime. Note that this task does not involve
synaesthetic perceptions.
However, if the synaesthetes are asked to name not a letter but the color of a color-patch,
preceded by a masked letter prime which they are not aware of having seen, there is no covert
priming effect: reaction times are no longer when the masked letter prime would, if unmasked,
26
To the extent color perceptions resulting from synaesthetic rerouting do dangle, the
dynamic sensorimotor approach would predict dominance.
10. Conclusion
Our main aims in this article have been to draw the dominance/deference
distinction, to indicate its relationship to the comparative explanatory gaps, and to raise
the question of how the distinction should be explained. In our view, the NCC
perspective does not by itself have the right kind resources to explain the
dominance/deference distinction. We have proposed a dynamic sensorimotor hypothesis
as a way of explaining the dominance/deference distinction and have suggested further
experiments.26 If our dynamic sensorimotor hypothesis is successful---and we emphasize
that its success or otherwise turns among other things on empirical issues---the approach
should by the same token go some way to bridge the comparative explanatory gaps.
We are thus suggesting that an empirical account can in principle scratch an

explanatory gap itch-—in particular, the comparative gap itches. Understanding the way
certain dynamic patterns of sensorimotor contingency are characteristic of particular
modalities and qualities provides a kind of insight and intuitive illumination into what
they are like, which does not leave us asking at once, “OK, but why does that
characteristic sensorimotor pattern go with what it is like to see?” When the
sensorimotor pattern characteristic of vision is explained, we have an “aha!” reaction; we
see through the dynamic pattern of sensorimotor contingency to what vision in particular
is like.27 Again, keep in mind that these claims relate to the comparative explanatory
gaps, and not the absolute gap.
By contrast, understanding that activity in certain brain areas is characteristic of

particular modalities and qualities leaves us wanting to rephrase the question
induce a different synaesthetic color from the color of the presented patch. Thus, synaesthetically
induced colors in particular appear to generate distinctively synaesthetic intermodal priming
effects only when they are consciously perceived, even though synaesthetes show normal covert
intramodal priming effects for letter recognition.
This suggests that synaesthetic color perception lacks some of the links that normal color
perception has. Mattingley et al 2001 suggests that synaesthetic interactions occur after initial
processes of recognition in the inducing modality are complete. See also Rich and Mattingley
2002.
26
Namely, we have suggested investigating whether applying TMS to auditory cortex of the cogenitally
deaf produces visual (or other sensory) distortions; whether Sadato’s later-blind subject would experience
tactile distortions when TMS is applied to visual cortex; whether TMS produces tactile illusions when no
Braille reading is in progress when applied to visual cortex in the congitally and early blind; whether
referred sensations are found in congenital phantoms; and whether referred sensations are found in late-
acquired phantoms who stroke themselves while watching in a mirror.
27
Again, more detailed sensorimotor characterizations of the modalities are given in O’Regan and Noë
2001a, c. We recommend attention to these for the most robust ‘aha!’ reaction.
27
immediately. Finding neural correlates of consciousness, or NCCs, is a splendid thing to

do, but it does not by itself scratch explanatory gap itches. We still want to know, for
example, why the qualitative expression of activity in a given brain area is like seeing
instead of hearing, or like one quality rather than another. Viewed out of the contexts of
the dynamic patterns of sensorimotor contingency in which they function, NCCs are
qualitatively inscrutable; we do not see through them to what their qualitative expressions
are like.
This contrast is a matter of degree. Characteristic patterns of sensorimotor

contingency may be qualitatively translucent rather than transparent. This may stem in
part from differences in the degree to which patterns of sensorimotor contingency are
accessible at the personal level, a thought we hope to pursue in further work. But even if
sensorimotor patterns are not always completely qualitatively transparent, they are a lot
more qualitatively scrutable than NCCs.
The itch-scratching/scrutability contrast just drawn between dynamic sensorimotor

and NCC approaches is at a psychological level. But meta-questions arise about this
contrast itself. If the expanded gaze strategy has the potential to provide more satisfying
answers than the inward neural scrutiny strategy, why is the latter so prevalent? What
assumptions orient us inwardly this way? This may be related to the question of the
logical relationship of our account to the claim that qualitative character supervenes on
neural properties. It may seem that our account is incompatible with a claim of neural
supervenience, but we deny this. Both are empirical claims, and they are logically
compatible. As an empirical matter, both claims may be true. Qualitative character may
supervene on neural properties even if the qualitative expression of neural activity is
determined, as we have argued, by dynamic sensorimotor considerations. This is so
because the rerouting (whether neural or extraneural) that changes patterns of
sensorimotor contingency may well also induce changes in neural properties at a given
locus. In work in progress we explain the compatibility claim further. But if both claims
are true, we hold that our account is explanatory in a way that the neural supervenience
claim is not. The neural supervenience claim may be true, but may nevertheless
encourage us to look in the wrong place for an explanation of qualitative character. The
explanatory promise of our approach, we suggest, is closely connected to the fact that it
expands our scrutiny both spatially and temporally, to the dynamic relations between
brain, body, and world.
28
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NEURAL PLASTICITY AND CONSCIOUSNESS1

[figures are missing in this version]

It seems natural to adopt an inward focus in response to such explanatory gap

We suggest that an inward focus in response to explanatory gap worries can be

We take our start from consideration of neural plasticity. This phenomenon

1. The distinction introduced: cortical dominance vs. cortical deference

Cortical dominance is illustrated by phantom limb cases in which there appears to

Cortical deference is illustrated when congenitally blind persons read Braille.

It may be natural to expect cortical dominance to be the norm. Perceptual scientists

2. The distinction schematized

Now we hypothesize a rerouting of afference. First, suppose that area 2, to which B

Will activation of area 2 by afference with source A give rise to experiences or

Notice that cortical dominance thus schematized involves illusory

3. The distinction at work

Phantom limb cases provide examples of intramodal dominance. Suppose A and B

Synaesthesia may provide examples of intermodal dominance. Color-graphemic

It is unclear whether synaesthesia results from nonstandard neural projections.

To spell out this suggestion about synaesthesia in terms of our schematized

Visual to auditory rerouting in ferrets provides an example of intermodal

As a result of this rerouting, 2-dimensional retinotopic maps (similar to those

Early blind readers of Braille provide examples of intermodal deference in human

4. How can the distinction be explained?

An initial hypothesis might be that we find dominance in cases that involve

But the intermodal deference/intramodal dominance hypothesis is not satisfactory,

A quite different suggestion turns on whether damage or rerouting has occurred

Thus, both the intermodal deference/intramodal dominance and the early

5. Intermodal plasticity without neural rerouting: adaptation to TVSS.

In order to move toward a deeper explanation, let’s consider some examples of

Consider first perceptual adaptation to a tactile-visual substitution system, which

In our schematism, A is here peripheral tactile input (patterns of vibration and

Someone might be skeptical that there is genuinely an intermodal change in

In TVSS, somatosensory cortex defers: but to what? Our original cases of

6. A dynamic sensorimotor hypothesis

These insights about TVSS can be generalized. According to our dynamic

A general account of intermodal differences in qualitative expression is thus

If the dynamic, sensorimotor view is to provide a bridge across comparative

contingencies are characteristic of vision, others of hearing, others of touch, there is an

Because TVSS effects a change from patterns of sensorimotor contingencies

It might be suggested that the dynamic sensorimotor hypothesis is on stronger

7. Intramodal plasticity without neural rerouting: reversing goggles

Consider the results of experiments on the long-term effects of wearing left-right

The initial effect of left-right reversing goggles is to produce a left-right reversal

However, putting on the goggles initially disrupts vision dramatically. Movements

If perceptual experience depends on the perceiver’s sensorimotor know-how, then

This prediction is born out by Taylor’s (1962) description of the experience of a

Like TVSS, Taylor’s goggle-adaptation result fits our extended characterization of

Again, we ask: deference to what? The change in qualitative expression cannot be

dynamic patterns of sensorimotor skills in relation to such objects. The perceiver’s

sensorimotor contingency. So either way, adaptation to reversing goggles illustrates

As we saw in the previous section, the dynamic sensorimotor approach gives a

To answer this question, we need to consider what is needed for sensorimotor

Second, however, sensorimotor know-how also requires you to be able to negotiate

The primary aspect of adaptation postulated by Harris without such a secondary

It may be natural to presuppose that experience is either visual or not visual

To sum up our response to Harris’ position: If adaptation involves illusory

Second, we should distinguish absolute from comparative issues: self-evidence and

9. The dominance/deference distinction explained.

This prediction is in line with Ramachandran’s well-known mirror-box results

How then, Ramachandran asks, can we understand the persisting experience of