Virtue Ethics
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C. Swanton
Vision for Action: Neural Mechanisms
Sensory input received by the visual receptors from an
object in space results in a complex pattern of
excitation from which object properties relevant to
different functions have to be extracted. This selection
is the basis for a specific mechanism called visuomotor
transformation, the shaping of the motor effectors
into configurations precisely adapted to capturing,
manipulating, using, and transforming visual objects.
In primates, the remarkable evolution of the hand into
a grasping apparatus represents an ultimate acqui-
sition for skilled interaction with objects and use of
tools. This article will focus on anatomical and
functional analysis of neural mechanisms involved in
visuomotor transformation.
Beginning in the 1970s, the anatomical description
of cortical visual pathways in the monkey has led to
the formulation of the two visual systems model
(Ungerleider and Mishkin 1982). According to this
model, the primary visual cortex in the occipital lobe
projects along two main streams that reach, respect-
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Figure 1
Lateral view of the left hemisphere of a monkey brain.
Two of the main sulci have been opened to show the
buried areas: the intraparietal sulcus (IPs) and the
lateral sulcus (Ls, which separates the parietal lobe
from the temporal lobe). The other main sulci are the
central sulcus (Cs) which separates the parietal lobe
from the frontal lobe, the arcuate sulcus (As) and the
principal sulcus (Ps) within the frontal lobe, and the
superior temporal sulcus (STs) within the temporal
lobe. The anterior part of the parietal lobe, which
includes somatosensory areas SI and SII (in the upper
bank of the lateral sulcus), is not concerned with
visuomotor transformation. The posterior part of the
parietal lobe includes areas 7 and 5 on the cortical
surface, and the intraparietal areas (medial, lateral,
ventral, anterior intraparietal areas, respectively). The
motor strip of the frontal lobe includes the primary
motor cortex (MI, the command area), and the dorsal
and ventral premotor cortex (PMd, PMv, respectively).
ively, the posterior parietal lobe (the dorsal pathway)
and the inferotemporal cortex (the ventral pathway).
Although the two pathways are densely intercon-
nected with each other, they are now viewed as
relatively independent functional entities.
The dorsal pathway, which will be under scrutiny in
this article, was once considered as a space channel.
Indeed, its experimental destruction in the monkey
produced spatial disorientation of the animal. Later
on, a more complete description of the anatomical
organization of the posterior parietal cortex, which is
outlined in Fig. 1, led to a new conception of its
functional role. First, a number a specialized areas,
many of them located in the intraparietal sulcus (e.g.,
LIP, MIP, AIP), were added to the classical cyto-
architectonic areas (e.g., areas 7 and 5). Second, these
newly described areas were found to be connected with
motor regions of the frontal cortex. These connections
appear to be selectively organized: parietal area MIP is
preferentially connected with the upper part of the
premotor cortex (PMd in Fig. 1), whereas AIP is
preferentially connected with its lower part (PMv).

Both PMd and PMv project to the motor cortex itself,
where the commands to the relevant muscles are
generated. Thus, the dorsal pathway, whereby visual
information is transferred from the primary visual
cortex to the premotor and motor cortex, can better be
conceived as a pathway specialized for visuomotor
transformation.
1. A Mechanism for Reaching Objects in Visual
Extrapersonal Space
Reaching for a visual object requires that the set of
coordinates in which the object-oriented movement is
computed must have its origin on the agents body
(egocentric coordinates). This constraint stems from
the fact that object position in space must be recon-
structed by adding signals from eye, head, and body
positions for the arm to be ultimately directed at the
proper location. Determining object position in space
by computing its retinal position alone would be
inappropriate, because this locus in space would
correspond to different retinal positions according to
respective eye, head, and body positions (Jeannerod
1988). The following paragraphs describe some of the
mechanisms which have been identified in recent
experiments in monkeys and which could account for
transforming retinal (eye centered) coordinates into
(body centered) egocentric coordinates. These mech-
anisms are primarily located in the intraparietal sulcus.
An influential hypothesis proposes that the basic
information about where to reach for an object is the
motor command sent to the eyes to move in its
direction (the eye position signal). Neurons in area 7a
have been shown to encode visual stimuli for a specific
position of the eye in the orbit, i.e., they fire prefer-
entially when the object stimulates a given location on
the retina (the neurons receptive field), and when the
gaze is fixating in that direction (Andersen and
Mountcastle 1983). These neurons would provide a
representation of visual space that could subsequently
be projected to the premotor cortex for guiding the
arm reach. This mechanism is apparently compatible
with the organization of visuomotor behavior: the
gaze systematically moves first to the location of the
desired stimulus, followed by the arm movement
within about 150 ms. A closer look, however, shows
that, because the motor command to the arm muscles
anticipates arm displacement by about the same
amount, the motor commands to the eye and to the
arm are in fact more or less synchronous (Biguer et al.
1982). Duhamel and his colleagues found the solution
to that enigma. They discovered in area LIP neurons
which fire in response to a stimulus briefly presented,
not within their receptive field, but within the area of
space where their receptive field will project after an
eye movement is made. This effect begins some 80 ms
before the eye movement starts (Duhamel et al. 1992).
The retinal position of receptive fields is thus modified
Vision for Action: Neural Mechanisms
prior to the occurrence of an eye movement. This
remapping of the representation of space in the
parietal cortex accounts for perceptual stability of
the visual scene during eye movements. In addition, the
existence of this mechanism demonstrates that signals
related to the intended eye position are available to the
motor cortex. In area VIP, neuron discharges testify to
a different form of remapping. Their receptive fields
remain in the same spatial position irrespective of eye
position in the orbit. In this case, therefore, the
receptive fields appear to be anchored, not to the eye,
but to the head, i.e., the receptive fields move across
the retina when the eyes move (Duhamel et al.
1997).
Eye and head position signals, however, should not
be sufficient in themselves for generating a directional
signal for guiding the arm. Besides the position of the
stimulus on the retina and the position of the eye in the
orbit, the position of the arm with respect to the body
must also be available to the arm motor command.
This function is achieved by neurons presenting
receptive fields in two sensory modalities. Neurons in
area MIP, for example, respond to both a localized
visual stimulus and a cutaneous stimulation applied to
one of their hands. These bimodal neurons are strongly
activated when the monkey reaches with that hand to
the visual target within their receptive field (see
Andersen et al. 1997, Colby 1998).
Neurons located in motor areas outside the parietal
cortex also present bimodal activation. Graziano and
his colleagues found bimodal neurons in the ventral
premotor cortex: these neurons, which in addition
encode the direction of arm movements, have tactile
receptive fields and visual receptive fields that extend
from the tactile area into adjacent space. For those
neurons which have their tactile receptive field on the
arm, the visual receptive field moves when the arm is
displaced (Graziano et al. 1994). These visual receptive
fields anchored to the arm encode visual stimulus
location in body (arm) coordinates and thus may
provide a solution for directional coding of reaching.
In what way does this premotor mechanism relate to
parietal mechanisms? What is the role of connections
between parietal areas and the premotor cortex? These
points remain unclear. It is likely that, instead of a
single directional signal used for all purposes, multiple
transformations and multiple spatial maps are gener-
ated and adapted to each purpose: guiding the eye,
guiding the arm, stabilizing the visual world, etc.
Data from single cell recordings are supported by
experimental data in human studies which clearly
point to the parietal lobe as the site of visuomotor
transformation for reaching. Although human data
are less direct than monkey data, they provide deeper
insight into the function due to the unique human
ability to execute tasks without having to learn them.
Functions like visually guided reaching can be
examined in more naturalistic situations. Observation
of patients with focal posterior parietal lesions in one
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Vision for Action: Neural Mechanisms
hemisphere show deep alterations of reaching move-
ments, usually limited to the hand contralateral to the
lesion: kinematics are altered, and large reaching
errors are observed. This misreaching is more marked
in the condition where the visual target is presented
outside the fixation point and increases when the
target distance from the fixation point increases (e.g.,
Milner et al. 1999). These clinical findings are cor-
roborated by brain imaging studies (based on changes
in localized cerebral blood flow) in normal subjects
during pointing at visual targets. The activated area, in
the parietal lobe contralateral to the pointing hand,
corresponds closely to the location of lesions that
produce misreaching (Faillenot et al. 1997). Finally,
the involvement of the posterior parietal cortex in
visuomotor control is confirmed by transcranial
magnetic stimulation (TMS). This technique allows
the application, via magnetic induction, of very brief
single shocks to a given cortical zone. When TMS was
applied to posterior parietal cortex shortly after onset
of a reaching movement directed to a visual target, it
prevented online corrections needed to reach the
target. TMS only affected the arm contralateral to the
stimulated cortex, which excludes the possibility that
the stimulation perturbed visual processing of target
location: only the adaptation of the motor command
to the visual position of the target was perturbed
(Desmurget et al. 1999).
2. The Case of Hand and Finger Moements
Grasping is often the ultimate achievement of
reaching. For this reason, the two functions can hardly
be dissociated. Grasping, however, relies on a quite
different system than reaching: it involves distal
segments, multiple degrees of freedom, and a high
degree of reliance on tactile function. It is an ensemble
of visuomotor transformations which preshape the
hand while it is transported to the object location.
Preshaping deals with object features which are not
relevant to reaching, like object size, shape, or orien-
tation (Jeannerod et al. 1995).
Indeed, anatomical data suggest that the cortical
pathway connecting posterior parietal and premotor
finger areas and controlling grasping is separate from
the pathway controlling reaching. Hand movement-
related neurons were disclosed by Sakata and his
colleagues (1995) in a small zone within the anterior
part of intraparietal sulcus, corresponding to an area
(AIP) closely connected with the ventral part of the
premotor cortex (PMv in Fig. 1). Neurons from AIP
were recorded in monkeys trained to manipulate
various types of objects, which elicited from the animal
different motor configurations of the hand: most of
them were selective for grasping objects of a given
shape. Task-related neurons were classified as motor
dominant or visual dominant according to whether becomes apparent when measuring the orientation of
their discharge related more to the action of grasping
16226
or to the inspection of the object. Most importantly,
neuron activity was not influenced by changing the
position of the object in space, which suggests that
they were related to distal hand and finger movements
(for which spatial position is irrelevant) rather than to
proximal (reaching) movements of the arm (Sakata et
al. 1995). Finally, a distinct population of neurons
sensitive to the 3-D orientation of the longitudinal axis
of visual stimuli was found in the caudal part of the
intraparietal sulcus. It is therefore likely that these
neurons process the 3-D characteristics of the object
and that the output of this processing is sent to AIP.
Transient inactivation of the AIP region, by locally
injecting a GABA agonist (muscimol), produces a
subtle change in the performance of visually guided
movements during grasping. Lack of preshaping and
grasping errors are observed in tasks requiring a
precision grip. In addition, there is a clear-cut dis-
sociation of the muscimol effects on grasping, which is
altered, and reaching, which is preserved. These results
(Gallese et al. 1994) support the view that these parietal
neurons play a specific role in the visual guidance of
grasping.
In humans, clinical observations also stress the role
of the posterior parietal cortex as a structure con-
trolling the visuomotor action of grasping. During
prehension of objects, parietal lesioned patients show
little or no preshaping with their contralateral hand;
they tend to open their finger grip too wide with
respect to object size and to close it only when their
fingers come into contact with the object; the
opposition axis (the axis which passes through the two
opposing fingertips and the center of gravity of the
object) is improperly oriented, which results in mis-
placement of fingertips on the object surface
(Jeannerod et al. 1994). In normal subjects, the above
brain imaging study (Faillenot et al. 1997) revealed an
area specifically involved in grasping, at the ventral
extremity of the intraparietal sulcus, also including
part of the somatosensory area S II. It is likely that this
cortical zone includes the human homologue of the
monkey AIP area.
An intriguing question thus remains to be discussed:
in which system of coordinates are finger movements
during grasping coded? The main aspect of grasping,
grip formation, has been described using parameters
such as grip size, orientation of opposition axis,
number of fingers involved, etc., which, arguably,
should not depend on the object position in space.
This is also the case for another aspect of grasping, the
encoding of grasp and lift forces: these parameters
depend on visual cues for object weight, for which
object location is irrelevant. There are limits to this
reasoning, however. Reaching and grasping cannot be
considered in isolation, as they represent two aspects
of the same action, reaching to grasp an object. The
need for coordination between these two aspects
the opposition axis during grasping. The position of

the fingertips on the object surface should only be
determined by object shape and 3-D characteristics in
order to achieve a stable grasp. Paulignan et al. (1997),
however, using upright cylinders as targets (e.g.,
objects affording multiple possible positions for the
fingertips) showed that orientation of the opposition
axis changed as a function of the cylinder position in
the workspace. This result indicates that the action of
grasping must encode not only finger movements but
also movements of the proximal joints: due to bio-
mechanical limitations of the arm, the configuration of
the upper limb cannot be the same for grasping the
same object at two different locations.
Even though reaching and grasping cannot be
considered as functionally independent (in spite of
segregation of their anatomical pathways), they pro-
cess different types of information and operate at
different rates. Experiments using visual perturba-
tions synchronized with movement onset clearly
demonstrate this difference (Paulignan et al. 1991a,
1991b). If, for example, the target object to be reached
by the subject jumps (by means of an optical device) by
10 degrees to the right as the subject starts reaching in
its direction, a corrective movement is produced so
that the new target is adequately reached: the first
kinematic evidence for correction occurs around
100 ms following the perturbation. If, in another
experiment, the object keeps the same location, but
changes size (e.g., increases) as the subject starts to
move, it takes at least 300 ms for the finger grip to
adjust to the changing object size. This sharp contrast
between the timing of the two types of corrections
accounts for the functional differences between the
two components of the action. Although fast execution
during reaching has an obvious adaptive value (e.g.,
for localizing and catching moving targets), this is not
the case for grasping, the function of which is to
prepare fine manipulation of the object once it has
been caught.
3. Conclusion
Posterior parietal cortex thus appears as a critical zone
for organizing object-oriented action, whether move-
ments are executed by the proximal or the distal
channels. The specialization of the dorsal visual path-
way for visuomotor transformation is complementary
with that of other brain regions, mostly located within
the ventral pathway, and specialized for object identifi-
cation and recognition. This suggests the existence of a
dual mode of processing for object properties like
spatial location, orientation, shape, or size. Perception
requires that object properties are processed within
an allocentric frame of reference, that is, independent
from the perceiver and where the relationships be-
tween objects within the visual array are preserved.
Action, by contrast, as has been shown in this article,
requires that information is processed within an ego-
centric frame of reference where everything in the
Vision for Action: Neural Mechanisms
visual array that is not the target for the movement is
ignored.
See also: Attention and Action; Eye Movements
and Perception; Motion Perception Models; Object
Recognition: Theories; Perception and Action; Per-
ceptual Organization
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Vision, High-level Theory of
The concept of high-level vision is used to denote
aspects of vision that are influenced by information
stored in memory such as an objects identity, its
name, or the context in which it belongs. It is usually
distinguished from low-level visual processing, which
is driven primarily by the visual input. High-level
vision involves functions such as object recognition,
face perception, mental imagery, picture naming,
visual categorization, or scene perception.
1. Background
In hierarchical models of vision (e.g., Marr 1982, see
also Marr, Daid (194580)), higher levels of visual
processing operate on the building blocks delivered by
more primitive visual mechanisms. In Marrs ap-
proach to perception, each stage has its own algo-
rithms and its own format of representing processing
output. The primal sketch represents the basic features
and their spatial relations (low-level vision) and the
2,5-D sketch represents surfaces and shapes from the
observers viewpoint (intermediate-level vision).
Recognition, that is, matching of object descriptions
stored in visual memory with visual input, Marr
believed only possible when the full 3-D structure of
an object and its components is represented in a so-
called 3-D object model (high-level vision).
Marrs approach has been highly influential as a
framework to integrate disparate work on the pro-
cessing of, among others, luminance, texture, shad-
ing, color, stereo, or motion (see Palmer 1999).
However, his assumption of strictly bottom-up pro-
cessing in sequential stages has gradually lost attrac-
tion. One reason for this theoretical evolution has
been the increased popularity in the 1980s of so-called
interactive activation models in which different levels
of processing interact with one another (see also
Connectionist Approaches; Artificial Neural Networks:
Neurocomputation): Partial outputs from lower-level
processes initiate higher-level processes and the out-
puts of higher-level processes feed back into the lower-
level processes.
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Copyright # 2001 Elsevier Science Ltd. All rights reserved.
International Encyclopedia of the Social & Behavioral Sciences
When different levels are distinguished in more
recent theories, great care is taken to avoid the
assumption of strict sequential processing. However,
the issue of where early vision ends, where high-level
vision starts, and how to demarcate the two has
remained important in current theorizing. Pylyshyn
(1999) has proposed the notion of cognitive impene-
trability as the major criterion for separating early (or
low-level) vision from later (or higher-level) vision.
Early visual processes run autonomously; they cannot
be influenced by what we know about the world or
expect in a specific situation. Most aspects of visual
processing which seem cognitive (e.g., recovering
unique 3-D percepts from ambiguous 2-D inputs) are
attributed to embodied natural constraints derived
from principles of optics and projective geometry,
internal to the visual mechanisms themselves. Only
effects of object-specific beliefs on earlier visual pro-
cessing would count as cognitive penetration. A large
number of so-called top-down effects of higher-level
processes on early vision are then explained as effects
of attention (preceding early vision) or as post-
perceptual effects on the decision stage based on the per-
ceptual output. For example, perceptual learning,
perceptual expertise and effects of categorization on
discrimination might be attributed to learning to focus
attention to the important features or attributes of a
stimulus. Semantic effects of the context of the whole
scene (e.g., a kitchen) on the identification of the
objects embedded within the scene (e.g., bread) have
been shown to result from response bias rather than
from genuine perceptual effects (e.g., Henderson and
Hollingworth 1999).
Implicit in Pylyshyns (1999) effort to delineate early
vision from cognitive influences is his beliefperhaps
shared by many vision scientiststhat real scientific
progress is possible only in early vision. Arguably,
however, in recent years considerable progress has
been made in the area of high-level vision as well (e.g.,
Edelman 1999, Ullman 1996). Most of this progress
has been made in object recognition, and, thus, this
topic will be covered predominantly (see also Binocular
Space Perception Models).
2. Object Recognition
2.1 Biederman (1987)
Object recognition concerns the identification of an
object as a specific entity (i.e., semantic recognition) or
the ability to tell that one has seen the object before
(i.e., episodic recognition). Interest in object recog-
nition is at least partly caused by the development of a
new theory of human object recognition by Biederman
(1987). Building on Marrs recognition ideas, where
perceptually derived 3-D volumetric descriptions are
matched to those stored in memory, Biederman in
addition incorporated Gestalt principles of perceptual
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