Documenti di Didattica
Documenti di Professioni
Documenti di Cultura
W. A. Phillips
University of Stirling, Scotland
and
Frankfurt Institute of Advanced Studies, Germany
Neural systems must be reliable but flexible. The contrast between these two
requirements is reflected by two frequently opposed perspectives that have arisen from
the neuroscience of the last century. First, there is the Hubel and Wiesel tradition that
sees sensory features and semantic attributes as being signalled by single cells, or
small local populations of cells. These codes are highly reliable. They do not change
from moment to moment, and do not depend upon what is going on elsewhere. Within
this conception feature detection and object recognition are achieved through a fixed
or slowly adapting feedforward projection through a hierarchy of cortical areas, and
this provides the basis for many higher cognitive capabilities.
In contrast, the second perspective emphasizes flexibility. From the early 1980s
onwards, there has been plenty of evidence that, even in sensory systems, activity is
influenced by high-level cognitive state variables such as attention, and by an ever-
changing stimulus context that reaches far beyond the classical receptive field. This
has led many to conclude that the simple Hubel and Wiesel tradition is no longer
viable, and that information is conveyed only by the rich non-linear dynamics of very
large and ever-changing populations of cells.
Our work on cognitive coordination combines these two views. It emphasizes
dynamic contextual interactions, but claims that, instead of robbing the local signals of
their meanings, they make those local signals even more reliable and relevant. Its
central hypothesis is that there are two classes of synaptic interaction: those that
specify the semantics of the signals transmitted, and those that coordinate these
computations so as to achieve current goals in current circumstances. They do this
through the two fundamental processes of contextual disambiguation and dynamic
grouping. These two processes amplify activity relevant to the current task and
stimulus context, group activity into coherent subsets, and combat noise by context-
sensitive redundancy. They are crucial to Gestalt perception, selective attention,
working memory, and strategic coordination.
Contextual disambiguation and dynamic grouping require many locally specific
coordinating interactions between all the detailed processes that compute the cognitive
contents. This implies that coordinating interactions must occur within and between
cortical regions, because it is only they that know the detailed cognitive contents. Our
working assumption is that there is a special class of synaptic interactions that
selectively amplifies and synchronises relevant activities. They are predominantly
mediated by long-range lateral and descending connections and influence post-
synaptic activity via a combination of NMDA receptors and GABA-ergic
interneurons. They do not themselves provide primary drive to post-synaptic cells, but
modulate the effects of those that do. We call them coordinating interactions to
distinguish them from the diffuse effects of the classical neuromodulators.
2
These broad claims of close relations between particular synaptic interactions and
particular cognitive functions are based upon many studies from many labs. They
show detailed relations within and between psychophysical functions, neurobiology,
and psychopathology. There is no time to review that evidence here, but much of it is
reviewed in Phillips and Singer, Behavioral and Brain Sciences 1997, 20:657-722 and
Phillips and Silverstein, Behavioral and Brain Sciences, 2003, 26:65-137. Here I
outline some recent studies from Stirling and Frankfurt.
This work builds upon Steve Silverstein’s original insight of a close relation
between cognitive coordination and cognitive disorganization in schizophrenia. His
argument for this leapt to my attention because, in addition to noting that the cognitive
functions that are impaired are exactly those for which coordination is most crucial, he
also noted that a new view of the pathophysiology of schizophrenia implicated exactly
those mechanisms postulated to be crucial to coordination, i.e. synaptic interactions
mediated by NMDA receptors. Evidence for that view continues to get even stronger,
and I expect a lot more to come, including a better understanding of the consequences
for neuronal dynamics and cognition of the various receptor sub-types that are now
know to occur in cerebral cortex, such as the 2A and 2B sub-types of NMDA receptor.
The first study that I will report here uses a new version of the visual contour
integration paradigm to measure sensitivity of Gestalt perception to temporal cues.
Several earlier studies suggest that temporal resolution is reduced in schizophrenia.
This may reflect a core underlying deficit with various cognitive consequences,
including impairments to Gestalt perception, attention, and working memory. There is
now growing evidence that dynamic aspects of early vision may provide an
endophenotype for cognitive disorganisation in disorders such as schizophrenia. If so,
a better understanding of those dynamic visual processes and their pathologies will
greatly facilitate the search for the distal genetic origins of cognitive disorganisation,
and aid the design of better pharmacotherapies. At Stirling we have therefore
developed a simple, sensitive, and specific psychophysical test of sensitivity to
temporal cues. It has been used extensively to study individual differences within and
between groups of students and schizophrenia patients. EEG and MEG studies using
this paradigm are now just beginning in Frankfurt.
In this paradigm two pseudo-random arrays of Gabor patches are displayed; one
to the left and one to the right of fixation. Within one array a sub-set of elements form
a figure, such as a continuous contour, that can only be reliably detected when their
onset is not synchronized with that of the background elements. For earlier work of
this sort see Hancock and Phillips, Vision Research 2004:2285-99. Using our new
version of the paradigm we found clear evidence on several issues. First, for most
subjects, segregation required an onset asynchrony of 20 – 40 ms. Second, detection
was no better when the figure was presented first, and thus by itself, than when the
background elements were presented first, even though in the latter case the figure
could not be detected in either of the two successive displays alone. This finding is
counter-intuitive. It is evidence against the hypothesis that salience is signalled by the
latency of cortical responses, and contrasts with inferences drawn from studies of
phase relations in rapidly cycling displays. Third, asynchrony segregated subsets of
randomly oriented elements as effectively as those aligned with the underlying
contour. Fourth, asynchronous onsets aligned with the contour could be discriminated
from those lying on the contour but not aligned with it. Fifth, though figure-ground
segregation depended upon asynchrony of the transient neural responses to abrupt
onsets, transient and sustained responses were not processed independently. Finally,
there were wide individual differences in sensitivity to these temporal cues, with
2
3
3
4
regions that modulate spike rate and timing. Local circuits using NMDA receptors and
GABA-ergic interneurons do this. 3. Their consequences include process of dynamic
grouping and contextual disambiguation that are relevant to all domains and levels of
cognition. 4. Their malfunctions can produce underlying impairments in both temporal
resolution and context-sensitivity which can lead to cognitive disorganization as seen
in several psychopathologies.