Handbook of Clinical Neurology, Vol.

151 (3rd series)

The Parietal Lobe
G. Vallar and H.B. Coslett, Editors
https://doi.org/10.1016/B978-0-444-63622-5.00023-1
Copyright © 2018 Elsevier B.V. All rights reserved

Chapter 23

The dorsal “action” pathway

JASON P. GALLIVAN1,2,3 AND MELVYN A. GOODALE4,5*
1
Department of Psychology, Queen’s University, Kingston, Ontario, Canada
2
Department of Biomedical and Molecular Sciences, Queen’s University, Kingston, Ontario, Canada
3
Centre for Neuroscience Studies, Queen’s University, Kingston, Ontario, Canada
4
Department of Psychology, University of Western Ontario, London, Ontario, Canada
5
Brain and Mind Institute, University of Western Ontario, London, Ontario, Canada

Abstract
In 1992, Goodale and Milner proposed a division of labor in the visual pathways of the primate cerebral
cortex. According to their account, the ventral pathway, which projects to occipitotemporal cortex, con-
structs our visual percepts, while the dorsal pathway, which projects to posterior parietal cortex, mediates
the visual control of action. Although the framing of the two-visual-system hypothesis has not been with-
out controversy, it is clear that vision for action and vision for perception have distinct computational
requirements, and significant support for the proposed neuroanatomic division has continued to emerge
over the last two decades from human neuropsychology, neuroimaging, behavioral psychophysics, and
monkey neurophysiology. In this chapter, we review much of this evidence, with a particular focus on
recent findings from human neuroimaging and monkey neurophysiology, demonstrating a specialized role
for parietal cortex in visually guided behavior. But even though the available evidence suggests that ded-
icated circuits mediate action and perception, in order to produce adaptive goal-directed behavior there
must be a close coupling and seamless integration of information processing across these two systems.
We discuss such ventral–dorsal-stream interactions and argue that the two pathways play different, yet
complementary, roles in the production of skilled behavior.

INTRODUCTION1 close – and for centuries, scientists have been trying to

understand how exactly vision manages to achieve this
Of all our senses, vision has fascinated us more than any
incredible feat. Even today, the study of visual perception
other. The reason for this is clear: our reflections on the
and its neural substrates is a flourishing – and
past, awareness of the present, and imaginings of the
expanding – enterprise in laboratories around the globe.
future are so intimately tied to, and in many ways insep- It is important to remember, however, that, in addition
arable from, our visual experience that it is virtually
to allowing us to see and interpret our world, vision
impossible to envision our mental events being governed
enables exquisite control of the movements we make in
by any other sense. The prominence of vision lies, in part,
that world, from the simple act of picking up a glass of
in its unique capacity to provide incredibly rich and
wine at dinner to the skilled thrust of a foil in a fencing
detailed information about the world far beyond the sur-
match. Yet, with the notable exception of eye movements,
face of our skin. No other sensory system comes even
which could be regarded as an information-seeking

1
Abbreviations used in the chapter are listed at the end of the chapter before References section.

*Correspondence to: Melvyn A. Goodale, Department of Psychology, Brain and Mind Institute, University of Western Ontario,
London, Ontario, Canada N6A 5B7. Tel: +1-519-661-2070, E-mail: mgoodale@uwo.ca
450 J.P. GALLIVAN AND M.A. GOODALE
adjunct to visual perception, it is only relatively recently
in the august history of vision science that researchers
have turned their attention to the role of vision in the pro-
gramming and control of our actions, particularly the
movements of our hands and limbs. One reason why
the visual control of action has been ignored for so long
is the prevalent belief that the same visual representation
that enables us to perceive our environment also guides
our actions in that environment. Indeed, visual percep-
tion is so directly linked to our conscious experience that
it is difficult to imagine that a separate visual system
might underlie our interactions with our surroundings.
As we will outline in this chapter, however, evidence
from a broad range of studies shows that vision is not
a unitary system. Rather, the control of skilled actions
depends on visual processes and neural mechanisms that
are quite separate from those mediating perception and
awareness.
DIFFERENCES IN THE
COMPUTATIONAL REQUIREMENTS
FOR VISUOMOTOR CONTROL AND
VISUAL PERCEPTION
To understand why the visual mechanisms mediating the
control of action are different from those mediating per-
ception, it is first necessary to consider differences in
their computational requirements, i.e., differences in
the way in which visual input is transformed for perform-
ing an action versus the way in which it is transformed for
constructing a percept. The visual information required
for the guidance of skilled actions must reflect the real
metrics of the world and be effector-specific. In the case
of grasping, for example, the location and disposition of
the goal object must be computed with respect to the
hand, or, at the very least, be selected from learned
“look-up tables” that link neurons coding a particular
set of visual inputs with neurons that code the desired
state of the grasping hand (for discussion of these issues,
see Thaler and Goodale, 2010; Goodale, 2011). More-
over, the time at which these computations are performed
is equally critical. Actors and goal objects rarely stay in a
static relationship with one another and, as a conse-
quence, the location and disposition of a target object
with respect to the limb can change radically from
moment to moment. In order to deal with these constantly
changing situations, “just-in-time” computations are
required: the required coordinates for action need to be
computed at the very moment the movements are per-
formed and dynamically updated in an “online” fashion
as the movement unfolds.
In contrast to visuomotor control, visual perception
does not need to deal with the absolute size of objects
or their egocentric locations (i.e., their locations with
respect to the particular effectors being used to perform
the action). In fact, very often such computations would
be counterproductive because our viewpoint with respect
to objects, as noted above, rarely remains the same from
one moment to the next. Instead, it changes constantly as
we move through our environment, even though our per-
ceptual representations of those objects typically show
constancy; i.e., we continue to see them as having the
same size, shape, and location even though our view-
point might change dramatically.
Indeed, for the purposes of perception, one can argue
that it would be better to encode the size, orientation, and
location of objects relative to others in the scene, even
though the projection of the images of those objects on
our retinas will vary as we move around. Such a
scene-based frame of reference permits a perceptual rep-
resentation of objects that transcends particular view-
points, and at the same time preserves information
about their spatial relationships (as well as their relative
sizes and orientations) as the observer moves around. To
do this, some of the brain networks underlying percep-
tion might use an array of viewer-centered representa-
tions of the same object; others might use canonical
representations; still others might be truly “object-
centered” (Riesenhuber and Poggio, 2000; for a discus-
sion of these issues, see Biederman, 2000; Hummel,
2013). But whatever the particular coding might be, it
is the identity of the object, not its disposition with
respect to the observer, that is of primary concern to
the perceptual system. The products of perception, like
object recognition and identity, also need to be available
over a much longer timescale than the visual information
used in the control of action. Thus, by working with per-
ceptual representations that are object- or scene-based,
we are able to maintain the constancies of size, shape,
color, lightness, and relative location, over time and
across different viewing conditions.
TWO VISUAL SYSTEMS
In 1992 Goodale and Milner suggested that these differ-
ences in the required computations for visual perception
and the visual control of action led to the emergence of
separate visual pathways in the primate cerebral cortex:
a ventral visual stream projecting from early visual areas
to inferotemporal cortex that mediates our perception of
the world, and a dorsal visual stream projecting to the
posterior parietal cortex (PPC), that mediates the visual
control of action (Fig. 23.1). According to their two-
visual-systems account of visual function, the identifica-
tion and selection of goal objects and an appropriate
course of action depend on the perceptual machinery
of the ventral stream and associated cognitive modules
in the temporal and frontal lobes, whereas the execution
 THE DORSAL “ACTION” PATHWAY
Fig. 23.1. Schematic representation of the two streams of
visual processing in the human cortex. Visual information
landing on the retina is projected to the lateral geniculate
nucleus of the thalamus (LGNd), which then projects to the pri-
mary visual cortex (V1). Within cortex, the information from
early visual areas (V1+) then projects along two separate path-
ways: a ventral stream (red) that projects to occipitotemporal
cortex and a dorsal stream (green) that projects to posterior
parietal cortex. The posterior parietal cortex also receives
visual input by the superior colliculus (SC) route, relayed
through the pulvinar. On the right, approximate locations of
the pathways are shown on a partially inflated human cortical
surface (PALS-B12 template: van Essen, 2005).
of subsequent goal-directed actions is mediated by ded-
icated online control systems in the dorsal visual stream
and associated motor and supplementary motor areas.
The two-visual-systems model recognizes that, in
order to have evolved, both the dorsal and the ventral
streams must contribute to the production of adaptive
actions and behavior. After all, natural selection can
operate only on the products of an organism’s actions
on the world. Indeed, a system that simply generates
internal percepts in isolation from outward, observable
actions could never have evolved. Rather, as was already
described, the percepts generated by the ventral stream
enable the organism to identify goals and select the
appropriate course of action to achieve those goals.
But, as Milner and Goodale have pointed out (2008),
the contribution of the ventral stream to the production of
adaptive actions does not stop there. By mediating the
learning of associations between the visual appearance
of an object and other properties such as its mass, density,
compliance, and surface friction, the ventral stream also
provides critical information for the successful manipu-
lation of the object once contact has been made. More-
over, many objects, such as tools, for example, have
functional properties and must be grasped and manipu-
lated in particular ways. In such cases, the ventral stream
contributes to the selection of the appropriate hand pos-
ture even though it is the dorsal stream that deals with the
metric requirements of the programming and control of
the eventual action (for a discussion of this issue, see
Milner and Goodale, 2006).
In this chapter, we review the evidence for these
claims, particularly evidence from recent neuroimaging
studies in humans. We will focus largely on studies of
451
the role of the dorsal stream in the visual control of
action, but will also touch on some of the contributions
of the ventral stream. But first, we briefly review the early
work on neurologic patients that drove the initial concep-
tion of the two-visual-systems model.
BRIEF REVIEW OF THE
NEUROPSYCHOLOGIC EVIDENCE
Some of the most compelling evidence for the division of
labor proposed by Goodale and Milner (1992) has come
from studies of patterns of spared visual abilities and
visual deficits in patients with damage to either the dorsal
or ventral stream. Patients with lesions in the dorsal
stream, particularly lesions of the superior regions of
the PPC that invade the territory of the intraparietal sul-
cus and/or the parieto-occipital sulcus, typically have
problems in using vision to direct a grasp or aiming
movement towards the correct location of a visual target
placed in different positions in the visual field, particu-
larly in the peripheral visual field. This deficit is typically
referred to as optic ataxia (following Balint, 1909; see
Chapter 11). But the failure to locate an object with the
hand should not be construed as a general problem in spa-
tial vision. Many of these patients, for example, can
describe the relative position of an object in space quite
accurately, even though they cannot accurately direct
their hand towards it (Perenin and Vighetto, 1988).
Moreover, sometimes the deficit will be seen in one hand
but not in the other (Balint, 1909). In some patients with
damage to the PPC, the deficit in acquiring targets with
the hand encompasses somatosensory-defined targets,
such as parts of the body that are touched by the exper-
imenter (see Chapter 11). But in other cases, the deficit
is restricted to visually defined targets (Balint, 1909).
Some individuals with damage to the dorsal stream are
unable to use visual information to rotate their hand,
scale their grip, or configure their fingers properly when
reaching out to pick up an object, even though they have
no difficulty describing the orientation, size, or shape of
objects in that part of the visual field (Fig. 23.2). More-
over, these deficits in grasping can even be seen in central
vision in patients with bilateral dorsal-stream lesions
(Jakobson et al., 1991). This pattern of deficits suggests
that the PPC plays a critical role in the visual control of
skilled actions (for a more detailed discussion of optic
ataxia and related sensorimotor deficits, see Milner and
Goodale, 2006, and Chapter 11).
The opposite pattern of deficits and spared abilities
has been described in patients with visual agnosia. Take
the case of patient DF (see Chapter 13), who developed a
profound visual form agnosia following carbon monox-
ide poisoning (Goodale et al., 1991). Although magnetic
resonance imaging (MRI) showed evidence of diffuse
452 J.P. GALLIVAN AND M.A. GOODALE

Fig. 23.2. Difference in brain damage and preserved function between an optic ataxic and visual form agnosic patient. Brain
images (top) highlight lesion differences between patient R.V., an individual with optic ataxia (left; from Goodale MA,
Meenan JP, Bulthoff HH, et al. (1994) Separate neural pathways for the visual analysis of object shape in perception and prehen-
sion. Curr Biol 4: 604–610), and patient D.F., an individual with visual form agnosia (right; from James TW, Culham J, Humphrey
GK, et al. (2003) Ventral occipital lesions impair object recognition but not object-directed grasping: an fMRI study. Brain 126:
2463–2475, with permission from Oxford University Press). Pink coloring highlights the lesioned areas in each individual. The
graphs (bottom) show the size of the aperture between the index finger and thumb during object-directed grasping and manual
estimates of object width for R.V. (left) and D.F. (right). The left panel shows that R.V. was able to indicate the size of the objects
reasonably well (individual trials marked as open diamonds), but her maximum grip aperture in flight did not show appropriate
object scaling. She simply opened her hand as wide as possible on every trial. In contrast, the right panel shows that D.F. exhibited
excellent grip scaling, opening her hand wider for the 50-mm-wide object than for the 25-mm-wide object. D.F.’s manual estimates
of the width of the two objects, however, were grossly inaccurate and showed enormous variability from trial to trial.

damage consistent with hypoxia, most of the damage
early on was evident in ventrolateral regions of the occip-
ital cortex, with V1 remaining largely spared (Fig. 23.2).
More recent high-resolution MRI scans confirmed
this damage, but revealed that her ventral-stream lesions
are focused in a region of the lateral occipital cortex (area
LO), that we now know is involved in the visual recog-
nition of objects, particularly their geometric structure
(James et al., 2003). Even though DF’s “low-level”
visual abilities are reasonably intact, she can no longer
recognize everyday objects or the faces of her friends
and relatives; nor can she identify even the simplest of
geometric shapes. (If an object is placed in her hand,
of course, she has no trouble identifying it by touch.)
Remarkably, however, DF shows strikingly accurate
guidance of her hand movements when she attempts to
pick up the very objects she cannot identify. Thus, when
she reaches out to grasp objects of different widths, her
hand opens wider midflight for wider objects than it does
for narrower ones, just like it does in people with normal
vision (Fig. 23.2). Similarly, she rotates her hand and
wrist appropriately when she reaches out to grasp objects
in different orientations, and she places her fingers cor-
rectly on the surface of objects with different shapes.
At the same time, she is quite unable to distinguish
between any of these objects when they are presented
to her in simple discrimination tests. She even fails in
manual matching tasks in which she is asked to show
how wide an object is by opening her index finger and
thumb a corresponding amount.
DF’s spared visuomotor skills are not limited to grasp-
ing. She can step over obstacles during locomotion as well
as control subjects, even though her perceptual judgments
about the height of those obstacles are far from normal.
Contrary to what would be predicted from the “what vs.
where” hypothesis (Ungerleider and Mishkin, 1982),
 THE DORSAL “ACTION” PATHWAY
a profound loss of form perception coexists in DF with a
preserved ability to use form in guiding a broad range of
actions. Such a dissociation, of course, is consistent with
the idea that there are separate neural pathways for trans-
forming incoming visual information for the perceptual
representation of the world versus for the control of
action. Presumably it is the former and not the latter that
is compromised in DF (for more details, see Goodale
and Milner, 2004; Milner and Goodale, 2006). It appears
that the selective damage to area LO has disrupted DF’s
ability to perceive the form of objects. These lesions have
not interfered with her ability to use visual information
about form to shape her hand when she reaches out and
grasps objects – presumably because the visuomotor net-
works in her dorsal stream that mediate grasping are largely
spared, with damage restricted to the most posterior part
of the parieto-occipital cortex (Whitwell et al., 2014).
Since the original work on DF, other patients with
ventral-stream damage have been identified who show
strikingly similar dissociations between vision for per-
ception and vision for action. Patient SB, who suffered
severe bilateral damage to his ventral stream early in life,
shows remarkably preserved visuomotor skills (he plays
table tennis and can ride a motorcycle) despite having
profound deficits in his ability to identify objects, faces,
colors, visual texture, and words (Le et al., 2002;
Dijkerman et al., 2004).
Recently, another patient, who sustained bilateral
damage to the ventral stream following a stroke, was
tested on several of the same tests that were given to
DF more than a decade ago. Remarkably, this new patient
(JS) behaved almost identically to DF. In other words,
despite his inability to perceive the shape and orientation
of objects, he was able to use these same object features
to program and control grasping movements directed
towards those objects (Karnath et al., 2009).
Finally, it is worth noting that if one reads the early
clinical reports of patients with visual form agnosia,
one can find a number of examples of what appears to
be spared visuomotor skills in the face of massive deficits
in form perception. Campion (1987), for example,
reports that patient RC, who showed a profound visual
form agnosia after carbon monoxide poisoning, “could
negotiate obstacles in the room, reach out to shake hands
and manipulate objects or [pick up] a cup of coffee.”
Taken together, the pattern of visual deficits and
spared abilities in DF (and in SB, JS, and other patients
with visual form agnosia) is in many ways the mirror
image of that observed in the optic ataxia patients
described earlier. Indeed, DF, who has damage in her
ventral stream, can reach out and grasp objects whose
form and orientation she does not perceive, whereas
patients with optic ataxia, who have damage in their dor-
sal stream, are unable to use vision to guide their reaching
453
and/or grasping movements to objects whose form and
orientation they can perceive. This double dissociation
is consistent with the division of labor between percep-
tion and action first proposed by Goodale and Milner
(1992). While the aforementioned patient work does well
in describing the consequences of damage to the separate
pathways for perception and action, some of the most
convincing evidence for the perception–action proposal
has come from neurophysiologic studies of the dorsal
and ventral streams in nonhuman primates and functional
MRI (fMRI) studies in the human. In the following sec-
tions, we briefly review the functional organization of the
visuomotor control networks in the primate brain, focus-
ing largely on the dorsal stream.
FUNCTIONAL ORGANIZATION OF THE
DORSAL PATHWAY IN HUMANS AND
NONHUMAN PRIMATES
Although there has been much debate about the precise
nature of the underlying computations, the roles of pri-
mary and secondary motor areas in frontal cortex in
the generation of movement have been well established
for more than a century (Fritsch and Hitzig, 1870; Ferrier,
1876, 1890; Sherrington, 1906; Penfield and Boldrey,
1937; see Graziano, 2009, for a detailed review). By
comparison, the contributions of the dorsal stream to
movement control, and those of the PPC in particular,
have been a topic of investigation only relatively recently
(Mountcastle et al., 1975; Mountcastle, 1976). This
was in part due to the historic assessment of the PPC
as a purely sensory association area, a designation
grounded in its anatomic location at the boundaries of
the primary visual, auditory, and somatosensory cortical
areas (Critchley, 1953; see Chapter 7). Consistent with this
idea, the PPC has been well characterized by its multi-
sensory functional properties. For example, the anterior
regions of the parietal cortex respond to tactile stimulation
and exhibit somatosensory responses (Mountcastle, 1957;
see Chapter 4) whereas the posterior regions show a vari-
ety of neural response properties linked to visuospatial
attention (Ungerleider and Mishkin, 1982; Colby and
Goldberg, 1999; Husain and Nachev, 2007). However,
it is now widely accepted, based on neurophysiologic
findings in nonhuman primates, patient work in humans,
and more recently, human fMRI studies, that the PPC
is indeed selectively involved in processes related to
movement planning and control (Mountcastle et al.,
1975; Andersen et al., 1997, 2014; Kalaska et al., 1997;
Culham and Valyear, 2006; Gallivan and Culham, 2015).
The PPC in both humans and nonhuman primates
consists of a mosaic of specialized subregions, each of
which has been shown to be involved in the planning
and control of specific bodily effectors (Culham et al.,
454 J.P. GALLIVAN AND M.A. GOODALE
2006; see Chapter 8, Fig. 8.8). First in awake, behaving
monkeys with neurophysiologic recording techniques
and then later in humans with neuroimaging methods,
these areas were identified by their activation selectivity
for movements of the hand (anterior intraparietal area
(AIP) in monkeys and AIP sulcus (aIPS) in humans),
arm (visual area 6 (V6A) and medial intraparietal area
(MIP) in monkeys and superior parieto-occipital cortex
(SPOC) and anterior precuneus (aPCu) in humans), eyes
(lateral intraparietal area (LIP) in monkeys and parietal
eye fields (PEF) in humans), and head (ventral intrapar-
ietal area (VIP) in monkeys, and VIP sulcus (VIPs) in
humans) (Gallivan and Culham, 2015; reviewed in
Culham and Valyear, 2006).
Mirroring these putative subdivisions in the PPC,
there are similar functional specializations in the motor
areas of the frontal cortex – an organization that reflects
reciprocal interconnections between the parietal and
frontal cortex (Johnson et al., 1996; Shipp et al., 1998;
Matelli and Luppino, 2000; Andersen and Buneo,
2002; Tanne-Gariepy et al., 2002). Although not an
exhaustive list, these frontal areas include the ventral pre-
motor cortex (PMv), involved in hand movements, the
dorsal premotor cortex (PMd), involved in arm move-
ments, and the frontal eye fields (FEF) involved in eye
movements (Weinrich and Wise, 1982; Bruce and
Goldberg, 1985; Rizzolatti et al., 1988). In addition,
the medial surface of frontal cortex contains secondary
motor areas, such as the supplementary motor area
(SMA), the pre-supplementary motor area (pre-SMA),
and the cingulate motor zones (discussed elsewhere;
see Dum and Strick, 2002; Amiez and Petrides, 2014).
All of these frontal areas, in addition to receiving projec-
tions from the PPC and exhibiting connectivity with one
another, send direct projections to the primary motor cor-
tex (M1), the main source of descending motor com-
mands to the spinal cord (Porter and Lemon, 1993), as
well as to the spinal cord directly (Dum and Strick,
1991, 1996, 2002; Boudrias et al., 2009).
Importantly, the PPC, in contrast to regions in the
frontal lobe, has considerably fewer direct anatomic pro-
jections to the spinal structures (less than one-fifth the
number of connections; see Murray and Coulter, 1981;
Toyoshima and Sakai, 1982), suggesting that the dorsal
stream largely exerts its influence over motor output
through its projections to frontal cortical regions. This
pattern of connections suggests that the PPC is more
likely to be involved in the earlier stages of sensorimotor
transformations for action, at a level that is more
abstractly removed from the precise muscle commands
needed to achieve the desired actions (Fig. 23.3).
Based on patterns of anatomic connectivity in mon-
keys and the response properties of neurons in the differ-
ent areas involved (Johnson et al., 1996; Matelli and
Luppino, 2000; Tanne-Gariepy et al., 2002), the PPC
has been putatively subdivided into two parallel proces-
sing streams for prehension: (1) a dorsolateral circuit
involving interconnections between CIP, AIP, and
PMv, thought to be specialized for object grasping and
manipulation; and (2) a dorsomedial circuit involving
interconnections between V6A, MIP, and PMd, thought
to be specialized for reaching and online corrections
during movement (Galletti et al., 2003; Johnson and
Grafton, 2003; Rizzolatti and Matelli, 2003; Grafton,
2010; Fig. 23.4). (A separate, third pathway, involving
interconnections between LIP and FEF, has been shown
to be involved in the preparation of eye movements.)
Whereas the anatomic divisions proposed by this dorsal
parallel pathway view are not in dispute, the extent to
which they actually support separable functional circuits
for grasping and reaching is still a matter of some debate.
A recent wealth of evidence demonstrates that macaque
V6A, an area presumed to be primarily involved in reach-
ing, also exhibits a wide range of grasp-selective neuronal
properties (Fattori et al., 2009, 2010, 2012), and neur-
oimaging experiments in humans have bolstered these
observations (Cavina-Pratesi et al., 2010; Gallivan et al.,
2011b; Monaco et al., 2011; Gutteling et al., 2015).
Nevertheless, the connectivity of the dorsolateral cir-
cuit does appear to indicate a more specialized role in
object grasping. Area AIP, for example, exhibits promi-
nent interconnections with inferotemporal cortex (Borra
et al., 2008). This presumably provides a direct access
route through which perceptual information about object
properties (e.g., surface texture, compliance) can be
shared with the dorsal-stream brain structures that use
this information for visually guided grasping, particu-
larly for the final phase when the required grip and load
forces are critical.
Furthermore, the dorsolateral pathway lies directly
adjacent to, and is interconnected with, the inferior pari-
etal lobule (IPL: Borra et al., 2008), a brain region
that, in monkeys, has been shown to represent the
higher-level goals of motor acts in which grasping is
embedded (Fogassi et al., 2005; Rozzi et al., 2008;
Bonini et al., 2011) and, in humans, it is an area that
has undergone considerable cortical expansion through
evolution (Van Essen and Dierker, 2007). It is perhaps
not surprising then that the IPL has been heavily impli-
cated in higher-order human-related behaviors such as
hand gestures and tool use (Johnson-Frey, 2004; see
Chapters 17 and 25), language processing (Davis and
Johnsrude, 2003), and action understanding (see
Chapter 28). In the next section, we provide a more
detailed discussion of each of the purported functional
subdivisions of human and monkey PPC, and attempt to
link each to a specialized role in visuospatial processing
for action.
 THE DORSAL “ACTION” PATHWAY 455

Fig. 23.3. Schematic representation of action-related areas of parietal and premotor cortex shown on the cortical surface of the
human (left) and macaque monkey (right) brain. The cortical surfaces (dorsal, lateral, and medial views) have been partially
inflated to reveal regions within the sulci while preserving a sense of curvature, and are based on the PALS-B12 template human
brain (left: Van Essen, 2005) and the F99 template macaque brain (right: Van Essen, 2004). Brain areas are color-coded according
to the legend at the bottom and are an approximation based on general consensus from human neuroimaging (left) and macaque
neurophysiology (right). For legibility, functionally defined regions have been grouped into the most well-established regions
implicated in hand and eye movements (note that the schematic is not intended to veridically show the extent and overlap of acti-
vation). Areas coded with the same color in parietal and premotor cortex within each species indicate direct connectivity and sim-
ilar functional specializations. Areas coded with the same color across species suggest possible functional homologies. White
dashed lines indicate major sulci. Sulci acronyms: AS, arcuate sulcus; Ces, central sulcus; CiS, cingulate sulcus; IFS, inferior fron-
tal sulcus; IPS, intraparietal sulcus; LuS, lunate sulcus; PoCes, postcentral sulcus; POS, parieto-occipital sulcus; PrCes, precentral
sulcus; PS, principal sulcus; SFS, superior frontal sulcus. Areal acronyms: AIP, anterior intraparietal area; aIPS, anterior intrapar-
ietal sulcus; APCu, anterior precuneus; CIP, caudal intraparietal area; FEF, frontal eye fields; LIP, lateral intraparietal area; MIP,
medial intraparietal area; mIPS, medial intraparietal sulcus; PEF, parietal eye fields; pIPS/cIPS, posterior or caudal intraparietal
sulcus; PMd, dorsal premotor cortex; PMv, ventral premotor cortex; SPOC, superior parieto-occipital cortex; VIP, ventral intra-
parietal area; vIPS, ventral intraparietal sulcus.

REGIONS IN POSTERIOR PARIETAL

Fig. 23.4. Subdivisions of the dorsal visual pathway. The two
parallel pathways for prehension within the dorsal stream are
shown according to Rizzolatti and Matelli (2003) and Galletti
et al. (2003). Pathways are projected on to the F99 template
macaque brain (Van Essen, 2004).
CORTEX
One brain region that appears to be critically involved in
both reaching and grasping actions in humans is the
SPOC, a putative human homolog of area V6A in the mon-
key (Pitzalis et al., 2006), located medially and slightly
anterior to the parieto-occipital sulcus. In monkeys, neu-
rons in V6A exhibit spatial tuning for parameters linked
to reaching, such as direction and amplitude, and more
recently have been shown also to encode parameters rele-
vant for object grasping, such as grip formation and wrist
orientation (for review, see Fattori et al., 2017).
456 J.P. GALLIVAN AND M.A. GOODALE
Consistent with these functional roles, experimentally
induced lesions to this area in monkeys lead to both
misreaching and misgrasping (e.g., a failure to correctly
reach towards, and appropriately preshape the fingers
of the hand for the target object; see Battaglini et al.,
2002). In line with these observations, convergent find-
ings from human fMRI (Prado et al., 2005; Bernier and
Grafton, 2010; Cavina-Pratesi et al., 2010; Gallivan
et al., 2011a, b; Tosoni et al., 2015), transcranial magnetic
stimulation (Vesia et al., 2010; Ciavarro et al., 2013) and
patient studies (Perenin and Vighetto, 1988; Karnath and
Perenin, 2005) all implicate SPOC in the preparation and
execution of reaching and reach-to-grasp actions. For
example, recent neuroimaging studies using neural decod-
ing techniques have shown that signals from SPOC reli-
ably predict different upcoming object-directed grasping
actions to be performed by an individual (Gallivan
et al., 2011b) as well as the spatial direction of their reach-
ing movements (Gallivan et al., 2011a).
Just anterior to SPOC in humans is the aPCu, an area
that has also been implicated in reaching (and pointing)
actions (Filimon et al., 2009; Bernier and Grafton, 2010).
Although both SPOC and aPCu are frequently
co-activated in sensorimotor tasks, they are likely to be
functionally distinct (Filimon et al., 2009; Bernier and
Grafton, 2010; Konen et al., 2013), a notion supported
by differences in their underlying cytoarchitecture
(Scheperjans et al., 2008a, b) and the observation that
they exhibit distinct patterns of whole-brain functional
connectivity (Margulies et al., 2009; Hutchison et al.,
2015). Notably, whereas several arguments have been
made for functional equivalence between human SPOC
and macaque V6A (Mars et al., 2011; Hutchison et al.,
2015; Fattori et al., 2017), the functional equivalent of
aPCu in the macaque is not yet clear. Nevertheless, in
the monkey, both of these neuroanatomical regions have
connections with the MIP and PMd (Matelli and
Luppino, 2000), regions that are both heavily implicated
in sensorimotor transformations for reaching (Weinrich
and Wise, 1982; Weinrich et al., 1984; Caminiti et al.,
1990; Snyder et al., 1997; Pesaran et al., 2006).
The intraparietal sulcus (IPS), which divides the
superior parietal lobule medially from the IPL laterally,
is perhaps one of the best-studied yet poorly understood
regions in parietal cortex. The area houses a highly
complex cytoarchitecture (Grefkes and Fink, 2005;
Gregoriou et al., 2006), exhibits a highly distributed
pattern of corticocortical connectivity (Mars et al.,
2011), and the structures contained within it have been
implicated in a dizzying array of high-level sensorimotor
and cognitive functions. These include, but are not lim-
ited to, roles in action planning and coordinate transfor-
mations (Andersen and Buneo, 2002), decision making
and value estimation (Gold and Shadlen, 2007), forward
state estimation and prediction (Wolpert et al., 1998;
Shadmehr and Wise, 2005; Mulliken et al., 2008), goal
encoding (Musallam et al., 2004; Andersen et al.,
2010), cognitive set (Stoet and Snyder, 2004), categori-
zation (Freedman and Assad, 2006), shape recognition
(Sereno and Maunsell, 1998), timing (Walsh, 2003;
Jazayeri and Shadlen, 2010, 2015), attention (Colby
and Goldberg, 1999; Bisley and Goldberg, 2003), learn-
ing (Clower et al., 1996; Rossetti et al., 1998; Linden
et al., 1999), number and magnitude processing
(Walsh, 2003; Hubbard et al., 2005; Ansari, 2008;
Nieder and Dehaene, 2009), and working memory
(Xu and Chun, 2006, 2009).
Located at the very posterior extent of the IPS, situ-
ated early in the hierarchy of the dorsal visual stream,
is the CIP. Neurons in macaque CIP exhibit tuning for
both two- and three-dimensional object visual surfaces
(Sakata et al., 1998; Tsutsui et al., 2002; Tsao et al.,
2003; Katsuyama et al., 2010; Rosenberg et al., 2013),
features that must be extracted and updated during move-
ment planning and control. The exact role of CIP in trans-
forming this visual information for the purposes of
action, however, is unclear, as reversible inactivation
of area CIP does not produce a grasping deficit, but rather
only a perceptual deficit in the discrimination of object
tilt/slant (Tsutsui et al., 2001). In humans, the posterior
extent of the IPS (pIPS) is activated in fMRI tasks requir-
ing object visual selection and attention (Wojciulik and
Kanwisher, 1999; Szczepanski et al., 2010) and it has
been shown to encode both target- and effector-related
information for movement (Beurze et al., 2007, 2009;
Gallivan et al., 2011a, 2013). With respect to the latter,
recent neuroimaging evidence in both macaques and
humans suggests a role for CIP and pIPS, respectively,
in eye- rather than arm-related movements (Leone
et al., 2014; Premereur et al., 2015; Saber et al., 2015),
although additional evidence is needed to verify this
effector selectivity.
Moving further anteriorly, the middle segment of the
IPS has been shown, in both the human and the monkey,
to be reliably activated by both eye and arm movements.
The LIP area, located on the lateral bank of the middle
IPS in monkeys, contains neurons more selective for
eye movements than for reaching movements (Snyder
et al., 1997; Cui and Andersen, 2007). Consistent with
this, electric stimulation of LIP produces fixed-vector
saccades (Thier and Andersen, 1996, 1998; Constantin
et al., 2007), and oculomotor behavior can be reliably
predicted from LIP population responses (Graf and
Andersen, 2014, 2015). Interestingly, it has been shown
that LIP activity represents upcoming decisions related to
not just eye (Gold and Shadlen, 2007) but also hand
movements (de Lafuente et al., 2015), suggesting a more
general role for the area in perceptual decision making.
 THE DORSAL “ACTION” PATHWAY
By contrast, area MIP, located on the medial bank of the
middle IPS in monkeys, contains neurons that are selec-
tive for reach movements rather than eye movements
(Snyder et al., 1997; Calton et al., 2002; Cui and
Andersen, 2007) and, in the context of decision making,
it only represents upcoming decisions related to arm
movements (de Lafuente et al., 2015). This latter obser-
vation is consistent with recent findings showing that
MIP inactivation impacts reach but not saccade choices
(Christopoulos et al., 2015) and execution (Hwang
et al., 2012).
Whereas in the monkey the functional organization of
the middle segment of the IPS makes it relatively easy to
dissociate eye- and arm movement-related neural sig-
nals, experimental investigations in the human have
painted a considerably fuzzier picture of such effector
selectivity. Indeed, the majority of human neuroimaging
studies exploring this topic have revealed largely over-
lapping activations for the two effectors in PPC
(Astafiev et al., 2003; Connolly et al., 2003; Hagler
et al., 2007; Levy et al., 2007; Beurze et al., 2009;
Filimon et al., 2009).
Such species-specific differences may be due to sev-
eral nonmutually exclusive factors. First, the neuroana-
tomic locations of LIP and MIP in the monkey lie
directly adjacent to each other (on opposing sulcal sur-
faces) and, in comparison to single neuron recordings,
fMRI has far poorer spatial resolution to allow for similar
levels of detection and separability. Second, there may
actually be less effector-specificity in human compared
to monkey PPC (Heed et al., 2011). Third, the human
middle IPS may not actually house the functional homo-
logs of LIP and MIP and the circuits may be organized in
a different fashion (Leone et al., 2014). Lastly, the signals
being measured by fMRI (blood oxygenation response)
do not reflect a one-to-one relationship with the action
potentials recorded via neural electrodes in monkeys
(Logothetis et al., 2001).
Nevertheless, setting these factors aside, fMRI work
has shown that activation patterns from middle IPS can
be used to decode prepared arm movements and saccadic
eye movements, as well as their spatial directions, to be
performed moments later (Gallivan et al., 2011a). Thus,
even if effector representations in the human are not as
compartmentalized as in the macaque, distributed activ-
ity patterns in the middle IPS still appear to exhibit sim-
ilar kinds of information. No doubt the close proximity of
arm- and eye-related representations in the IPS (as in
frontal cortex) is not by chance but rather an organiza-
tional feature that facilitates the close coupling and coor-
dination of the two effectors in everyday behavior
(Andersen and Buneo, 2002; Andersen and Cui, 2009).
The most anterior section of the IPS, area AIP in
the monkey, contains neurons selectively involved in
457
three-dimensional shape coding (Srivastava et al., 2009;
Theys et al., 2012), and hand preshaping for grasping
(Taira et al., 1990; Sakata et al., 1997; Murata et al.,
2000; Baumann et al., 2009). Consistent with a role in
visual transformations for object grasping, muscimol
(g-aminobutyric acid agonist) injections into AIP disrupt
visually guided object-directed hand preshaping, while
leaving reach behavior relatively intact (Gallese et al.,
1994; see Janssen and Scherberger, 2015, for review).
In agreement with this functional organization, the human
aIPS also shows selectivity for grasp planning and execu-
tion with fMRI (Culham et al., 2003; Frey et al., 2005;
Begliomini et al., 2007; Cavina-Pratesi et al., 2010).
Moreover, neurologic patients with damage to this area
show deficits in grasping but little disturbance in reaching
(Binkofski et al., 1998), and transcranial magnetic
stimulation to the region selectively interferes with the
grasp component of reach-to-grasp actions (Tunik et al.,
2005; Rice et al., 2006).
As emphasized above, neurophysiologic evidence
across human and nonhuman primates points to the dor-
sal stream as being critically involved in the visual con-
trol of action. As noted at the outset, however, the dorsal
stream does not typically act in isolation, but rather
receives considerable input from the ventral stream,
which identifies goal objects in the environment and aids
in the selection of an appropriate action based on the
current situation and context. In the following section,
we turn to a discussion of some of the important ways
in which the two visual streams may come to interact
in the service of goal-oriented behavior.
INTERACTIONS BETWEEN THE
DORSAL AND VENTRAL STREAMS
The two-visual-systems framework provides a convinc-
ing account for the role of the dorsal pathway in mediat-
ing reaching and grasping movements directed towards
objects, where the relevant object metrics (e.g., location,
size, shape) can be computed directly from visual input.
But, as discussed earlier, reaching out and grasping an
object is most often the prelude to actually manipulating
and interacting with objects (i.e., lifting and moving them)
in our environment. Preparing such actions requires
knowledge of an object’s mechanical properties (e.g.,
weight and the object’s associated dynamics, as well as
its compliance, surface friction coefficients, and even its
temperature) – properties that cannot be reliably derived
from retinal inputs alone and must instead be estimated
based on stored knowledge linking visual information
about the object (e.g., size or material) to its mechanical
properties.
To lift an object efficiently, for example, an accurate
prediction of object weight is critical (Wolpert and
458 J.P. GALLIVAN AND M.A. GOODALE
Flanagan, 2001; Flanagan et al., 2006; Johansson and
Flanagan, 2009). Individuals will smoothly increase their
lifting forces (i.e., vertical load force) to a level that just
exceeds the estimated object weight and, when this pre-
diction is correct, the object will be lifted efficiently.
When lifting an object for the first time, people rely on
well-learned correlations, or priors, relating material
and size to weight (Gordon et al., 1991a, b, 1993;
Flanagan and Beltzner, 2000; Buckingham et al., 2009;
Baugh et al., 2012). For example, we learn from an early
age that large objects tend to be heavier than smaller
objects and, likewise, that metal objects tend to be
heavier than wood objects, and readily exploit this infor-
mation to appropriately scale the fingertip forces used
when lifting objects.
Undoubtedly, the use of visual cues, such as size and
material, in the initial calibration of lifting forces indi-
cates a role for the ventral visual pathway in the control
of action. Indeed, at the level of brain mechanisms, a
wealth of evidence points to regions in occipitotemporal
cortex (OTC) as being critically involved in the coding of
visual object properties, such as size and material (Grill-
Spector and Malach, 2004; Cant and Goodale, 2007,
2011), and behavioral research suggests that size infor-
mation, used to predict weight when lifting, is actually
processed in the ventral stream (Brenner and Smeets,
1996; Jackson and Shaw, 2000). Thus, one possibility
is that structures in OTC provide the dorsal pathway,
via projections to parietal cortex (Borra et al., 2008), with
visual information about object properties relevant for
computing weight used when lifting. In which case,
we might be expected to find an encoding of object
weight information only in the dorsal visual pathway,
consistent with its known role in computing the real-
world metrics of objects relevant for sensorimotor
control.
Another possibility, however, consistent with evi-
dence showing that activity in OTC is sensitive to previ-
ous sensorimotor experience (Weisberg et al., 2007),
real-world object properties (Konkle and Oliva, 2012),
and a variety of sensory inputs linked to object identity
(e.g., tactile, auditory; see Amedi et al., 2001; James
et al., 2011), is that object weight is actually encoded
at the level of object-processing regions in the ventral
visual pathway.
Interestingly, consistent with this latter prediction,
Gallivan and colleagues (2014) showed, using human
fMRI and neural decoding methods, that object-
processing areas in OTC, in addition to areas in primary
and secondary motor cortex, are involved in representing
object weight. The encoding of a motor-related object
feature in OTC, though perhaps unexpected in the con-
text of strict interpretations of the dual-pathways view
(McIntosh and Schenk, 2009; Schenk and McIntosh,
2010), makes sense in light of the fact that object weight
cannot be directly appreciated from vision. Rather, it
requires the extraction of lasting and detailed information
about objects, via learning and memory associations
(Milner and Goodale, 2006).
While weight predictions based on size-weight and
material-weight priors are often accurate, they can, at
times, be erroneous (e.g., when one braces to lift a large
bag of luggage only to quickly realize, at lift-off, that it is
in fact empty). Not surprisingly, in such circumstances
individuals are capable of using knowledge obtained
from previous lifts, known as sensorimotor memory, to
adapt their lifting force in subsequent lifts of the same
object (Johansson and Westling, 1988; Cole, 2008;
Baugh et al., 2012). Nevertheless, these well-learned
priors are robust and extremely difficult to override, as
revealed through the size weight and material weight illu-
sions (SWI and MWI, respectively).
The SWI refers to the observation that individuals,
when lifting two objects of different volumes but equal
weight, will almost without fail judge the smaller object
to be heavier (Charpentier, 1891; Murray et al., 1999).
This illusion is thought to arise from a simple mismatch
between what one expects the object to weigh and what
it actually weighs (Flanagan et al., 2008). That is, because
the smaller object is heavier than expected, it is judged
to be heavier (and vice versa with the larger object).
Similarly, the MWI refers to the observation that individ-
uals will often report, when lifting two objects of different
materials but of the same volume and weight, the object
with the lighter-looking material (e.g., expanded poly-
styrene) as feeling heavier than the one made from
heavier-looking materials (e.g., metal: Harshfield and
DeHardt, 1970; Ellis and Lederman, 1999; Buckingham
et al., 2009). Interestingly, neither of these illusions is
weakened when the actor is informed that the objects
are of equal weight (Flourney, 1894; Nyssen and
Bourdon, 1955) or lessened with repeated lifting of the
objects (Wolfe, 1898; Seashore, 1899; Buckingham
et al., 2009). This is despite the fact that individuals will cor-
rectly adapt their lifting forces to the actual weights of the
objects in subsequent lifts (Flanagan and Beltzner, 2000;
Buckingham et al., 2009) (Fig. 23.5).
Interestingly, it has recently been shown that, by alter-
ing the expectation that weight increases with size
(through having participants practice lifting objects over
repeated daily sessions in which small objects are always
heavier than large ones), the SWI can be attenuated and
eventually even inverted, such that the bigger of two
equally weighted objects will actually be judged as
heavier (Flanagan et al., 2008). Despite this significant
change in perceptual report, the sensorimotor system
nevertheless quickly learns to predict the weights of
the inverted-weight objects accurately, as revealed
 THE DORSAL “ACTION” PATHWAY 459

Fig. 23.5. Demonstration of the size–weight illusion (SWI). (A) Size-weight stimuli. Subjects lift the large and small objects
(equal in weight) by grasping a handle using a precision grip (between thumb and index finger). The handle is equipped with
two force-torque sensors that can be quickly moved from object to object. A light-sensitive diode, embedded into the lifting plat-
form, records object lift-off. (B) Fingertip force records for a representative individual. Grip force (in newtons), load force (LF),
grip and load force rates, and light-sensitive diode recorded in the initial (left) and later (right) lifts of each object. This subject lifted
the large object (blue traces) and then the small object (red traces), in an alternating fashion. In all trials, subjects grasped the object
and increased grip and load force together until lift-off occurred (signaled by the light diode). In the initial trials (left), peak grip and
load force rates were scaled to object size whereas in the later trials (right), the peak force rates were statistically indistinguishable
for the two objects (i.e., appropriately scaled to the equally weighted objects). Vertical lines indicate object lift-off. (C) Lift param-
eters, averaged across subjects. Grip force rate and load force rate are plotted as a function of trial number. Each dot represents an
average of 20 subjects (vertical lines are standard errors). Curves were fitted for each object to provide a visual impression of the
force adaptation over trials. (D) Heaviness rating, plotted as a ratio of the small-to-heavy object. Participants indicated, on a
10-point scale, the perceived heaviness of the two objects throughout the duration of testing. A positive score indicates the smaller
object as being perceived heavier. (Data kindly provided by Randy Flanagan and modified from Flanagan and Beltzner (2000),
with permission from Nature Publishing Group.)
460 J.P. GALLIVAN AND M.A. GOODALE
through the fingertip forces used when lifting. This dem-
onstrates that the well-learned priors serving the sensori-
motor system are not hard-wired or crystallized during
development, but can be gradually modified through
the experience of lifting objects (Flanagan et al., 2008).
Although the brain mechanisms that support the dis-
sociation between force estimation (for lifting) and per-
ceived heaviness remain poorly understood, fMRI work
has suggested the left PMv as playing a role in estimating
object density based on size and weight (Chouinard et al.,
2009), a factor thought to be important in driving the SWI
(Grandy and Westwood, 2006; Buckingham, 2014).
Taken together, this elegant work on object lifting pro-
vides support for the dual, but not mutually exclusive,
proposition that: (1) the control of object manipulation
requires access to ventral visual pathway resources (such
as computations about size and material); and (2) like
other action–perception dissociations, the sensorimotor
system can often operate independently of the cogni-
tive/perceptual system.
CONCLUDING REMARKS
When Goodale and Milner (1992) first proposed the idea
of a separate vision-for-action system in the dorsal
stream, the emphasis was on the independence of this
system from vision for perception in the ventral stream.
But, even in the original conception, it was understood
that the two systems would have to work closely together
in the generation of purposive (and adaptive) behavior.
Indeed, as noted earlier, in light of the pressures of natural
selection, if the computations of the ventral stream did
not in some way assist in the selection of our actions
(and help the organism survive) then it is unlikely to have
ever evolved.
One way to think about the interaction between the
two streams (an interaction that capitalizes on the comple-
mentary differences in their computational constraints) is
in terms of a tele-assistance metaphor borrowed from
robotics engineering (Goodale and Humphrey, 1998).
In tele-assistance, a human operator, who has identified
a goal object and decided what to do with it, communi-
cates with a semiautonomous robot that actually performs
the required motor act on the flagged goal object (Pook
and Ballard, 1996). In terms of this tele-assistance meta-
phor, the perceptual-cognitive system in the ventral
stream, with its rich and detailed representations of the
visual scene (and links with cognitive systems), would
be the human operator. Processes in the ventral stream
participate in the identification of a particular goal and
then flag the relevant object in the scene, perhaps by
means of an attention-like process. Once a particular goal
object, such as a wine glass on the dinner table, has been
flagged, dedicated visuomotor networks in the dorsal
stream (in conjunction with related circuits in premotor
cortex, basal ganglia, cerebellum, and brainstem) are then
activated to transform the visual information about the
cup into the appropriate coordinates and motor commands
for the desired action (i.e., in the metaphor above, the
dorsal stream would be the semiautonomous robot). This
means that in many instances a flagged object in the scene
will be processed in parallel by both ventral and dorsal
stream mechanisms – each transforming the visual infor-
mation in the array for different purposes. In other situa-
tions, however, where the visual stimuli are particularly
salient, visuomotor mechanisms in the dorsal stream
can operate without any immediate supervision by the
ventral stream’s perceptual mechanisms.
Of course, the tele-assistance metaphor does not cap-
ture the entire interaction between the dorsal and ventral
streams that must occur in a natural reach-to-grasp
movement. Although the dorsal stream might be able
to provide most of the metric computations required
to guide the hand towards the goal object, the ventral
stream clearly plays a pivotal role in selecting the
appropriate hand posture (in the case of tools, for exam-
ple) and in determining the required grip and load
forces for interacting with the object once contact is
made. In short, the two streams play different but com-
plementary roles in the production of skilled and adap-
tive actions.
ACKNOWLEDGMENTS
J.P.G. was supported by a Canadian Institutes of Health
Research (CIHR) fellowship and M.A.G. was supported
by CIHR, the Natural Sciences and Engineering
Research Council of Canada, and the Canada Research
Chairs program.
ABBREVIATIONS
AIP, anterior intraparietal area; aIPS, anterior intraparie-
tal sulcus; aPCu, anterior precuneus; CIP, caudal
intraparietal area; FEF, frontal eye fields; fMRI,
functional magnetic resonance imaging; IPL, inferior
parietal lobule; IPS, intraparietal sulcus; LIP, lateral
intraparietal area; LO, lateral occipital area; M1,
primary motor cortex; MIP, medial intraparietal area;
MRI, magnetic resonance imaging; MWI, material
weight illusion; OTC, occipitotemporal cortex; PEF,
parietal eye fields; pIPS, posterior intraparietal sulcus;
PMd, dorsal premotor cortex; PMv, ventral premotor
cortex; PPC, posterior parietal cortex; pre-SMA, pre-
supplementary motor area; SPOC, superior parieto-
occipital sulcus; SMA, supplementary motor area;
SWI, size weight illusion; +V6A, visual area 6; VIP,
ventral intraparietal area; VIPs, ventral intraparietal
sulcus.
 THE DORSAL “ACTION” PATHWAY
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