Crowe et al’s data are consistent with Ungerleider and Mishkin’s
(1982) classical delineation between a cortical ventral stream that is
engaged in identifying and recognizing objects and a dorsal stream that is
concerned with computing spatial relationships. Milner and Goodale (1995)
have challenged this conceptualization of the functional specialization of
the dorsal and ventral streams. In doing so these authors question not
only the forms of spatial encoding undertaken by the dorsal stream but,
more critically, question the purpose for which that spatial encoding is
employed. Within Ungerleider and Mishkin’s framework the dorsal stream
underlies our conscious perception of spatial relationships. Milner and
Goodale have argued that not only is scene-based spatial cognition the
preserve of the ventral stream (Carey, Dijkerman, Murphy, Goodale, Milner,
2006) but that spatial coding in the dorsal stream does not serve
perception per se. Rather, in conjunction with certain types of object-
form analysis, the dorsal stream functions to supply information regarding
‘how’ to act upon objects in the world.
At first glance Crowe and colleagues’ results appear not only to
support Ungerleider & Mishkin’s (1982) proposed division, but also
seem to contradict Milner and Goodale’s model. Indeed, the task utilized
by Crowe et al. is quite cunningly designed to avoid any need to program
spatial relationships for making an action. The monkey does not, for
example, have to respond by pointing to the left or right, but rather has
to press abutton at the right point in time. Such a behavioral output,
however, does not guarantee that the dorsal stream will not compute
information to guide an action. In fact, several studies have shown that
the dorsal stream performs computations related to performing a given
action even if the participant does not intend to act on that information
(Grezes, Tucker, Armony, Ellis and Passingham, 2003). Furthermore many
area-7 neurones are clearly modulated by the execution of manual actions:
whilst they might encode the properties of a stimulus viewed passively
they will respond much more vigorously if the animal performs an action on
that stimulus (Mountcastle et al., 1975).
Moreover, the form of object-centered spatial coding identified by
Crowe and colleagues could play an important role in ‘vision for action’.
When picking up a cup, for example, it is important to encode the location
of the handle with respect to the overall object. There are, however, more
complicated forms of spatial representation that would prove more
difficult for the ‘vision for action’ model of the dorsal stream. Milner
and Goodale argue that allocentric encoding (between different objects)
requires the ventral stream (see Carey et al., 2006). Distinguishing
between truly allocentric encoding and object-based encoding (between
elements of a single object), of course, raises the issue of defining what
is perceived as a single object. The inverted T-stimulus used by Crowe and
colleagues intuitively groups to form a single object. It would therefore
be theoretically important to extend the Crowe et al. paradigm to explore
whether there are neurons in area 7a that encode not just the relationship
of a target element to a single object but the relationship between
objects, for example by locating them at distances at which they would
clearly not be grouped into a single object. If activity in area 7a
neurones still predicted the relationship between these objects, it would
follow that the dorsal stream could encode complex spatial relationships
that play no obvious role in action and provide a serious challenge to the
vision-for-action model of the dorsal stream.
Finally, it is worth drawing attention to the interesting coincidence
between the localization of the proposed viewer- to object-centered
transformation and the location of 7a within the anatomical hierarchy that
embodies the dorsal and ventral streams. Andersen et al. (1990) point out
that although there are other connections between early levels of
processing in the two streams, 7a provides the only direct link between
the endpoints of processing within the two streams. Andersen and
colleagues therefore suggest that area 7a may play a theoretically
significant role in communicating information in the two streams. Milner
and Goodale themselves argue that communication between the two streams is
often required for coordinated behavior. For instance, the dorsal stream
might program the exact kinematics of how to act upon a given object while
the ventral stream selects which objects to act upon (Milner and Goodale,
1995).
In the context of communicating between different egocentric (i.e.
hand-, head-, or retina-based) frames of reference purely within the
parietal lobe, Cohen and Andersen (2002) have argued that the
establishment of a common frame of reference could prove useful in
coordinating the action of multiple effectors. The creation of an object-
centered frame of reference in an area that just happens to provide the
most direct connection between the higher levels of processing between the
two streams might have an important role to play in establishing a common
reference frame within which the two streams can communicate.
References:
Andersen RA, Asanuma C, Essick G, Siegel RM (1990) Corticocortical
Connections of Anatomically and Physiologically Defined Subdivisions
within the Inferior Parietal Lobule. J Comp Neurol, 296(1), 65-113.
Carey DP, Dijkerman HC, Murphy KJ, Goodale MA, Milner AD (2006)
Pointing to places and spaces in a patient with visual form agnosia.
Neuropsychologia 44(9): 1584-1594.
Cohen YE, Andersen RA (2002) A common reference frame for movement
plans in the posterior parietal cortex. Nat Rev Neurosci 3(7):553-62
Grezes J, Tucker M, Armony JL, Ellis R, Passingham RE (2003) Objects
automatically potentiate action: an fMRI study of implicit processing.
Eur J Neurosci 17: 2735-2740.
Milner AD, Goodale MA, (1995) The Visual Brain in Action. Oxford:
Oxford University Press.
Mountcastle VB, Lynch JC, Georgopoulos A, Sakata H, Acuna C (1975)
Posterior Parietal Association Cortex of Monkey - Command Functions for
Operations within Extrapersonal Space. J Neurophysiol 38(4): 871-908.
Ungerleider LG, Mishkin M, (1982) Two cortical visual systems. In:
Analysis of Visual Behavior (Ingle DJ, Goodale MA, Mansfield RJW, ed),
pp549-586. Cambridge MA: MIT Press.