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Whats a good experiment in cognitive brain imaging?

A personal view
C Frith, 13th March 2002

1) Mapping

What is the function of the various Brodmann areas?

This question cannot be answered with one experiment.

Mapping the known: retinotopic areas in the visual system (Sereno et al., 1995).

Mapping the partially known: biological motion the need for a localiser task (Grossman
et al., 2000)
Mapping the unknown:
a) classifying cognitive functions
Stroop (Barch et al. 2001), Working memory (Petit et al., 1998)
b) contrasting specific cognitive functions (Rowe et al., 2000)

2) Mechanisms

Bottom-up & top-down (Corbetta et al., 1991 features, OCraven et al., 1999, objects +
Site & Source: Macaluso et al., 2000, multi-modal, Wojcuilik et al., 1999, many task one
top-down source, Kastner et al., 1999, activity before stimulus, de Fockert et al., working
memory and attention.

3) Connectivity

Attention: modulating connectivity. Bchel & Friston, 1997


Borders Of Multiple Visual Areas In Humans Revealed By Functional Magnetic-


Sereno Mi, Dale Am, Reppas Jb, Kwong Kk, Belliveau Jw, Brady Tj, Rosen Br, Tootell

Science, 268 (5212): 889-893 May 12 1995

The borders of human visual areas V1, V2, VP, V3, and V4 were precisely and
noninvasively determined. Functional magnetic resonance images were recorded during
phase-encoded retinal stimulation. This volume data set was then sampled with a cortical
surface reconstruction, making it possible to calculate the local visual field sign (mirror
image Versus non-mirror image representation). This method automatically and
objectively outlines area borders because adjacent areas often have the opposite field
sign. Cortical magnification factor curves for striate and extrastriate cortical areas were
determined, which showed that human visual areas have a greater emphasis on the center-
of-gaze than their counterparts in monkeys. Retinotopically organized visual areas in
humans extend anteriorly to overlap several areas previously shown to be activated by
written words.

Brain areas involved in perception of biological motion

Grossman E, Donnelly M, Price R, Pickens D, Morgan V, Neighbor G, Blake R

Journal Of Cognitive Neuroscience, 12 (5): 711-720 SEP 2000

These experiments use functional magnetic resonance imaging (fMRI) to reveal neural
activity uniquely associated with perception of biological motion. We isolated brain areas
activated during the viewing of point-light figures, then compared those areas to regions
known to be involved in coherent-motion perception and kinetic-boundary perception.
Coherent motion activated a region matching previous reports of human MT/MST
complex located on the temporo-parietooccipital junction. Kinetic boundaries activated a
region posterior and adjacent to human MT previously identified as the kinetic-occipital
(KO) region or the lateral-occipital (LO) complex. The pattern of activation during
viewing of biological motion was located within a small region on the ventral bank of the
occipital extent. of the superior-temporal sulcus (STS). This region is located lateral and
anterior to human MT/MST, and anterior to KO. Among our observers, we localized this
region more frequently in the right hemisphere than in the left. This was true regardless
of whether the point-light figures were presented in the right or left hemifield. A small
region in the medial cerebellum was also active when observers viewed biological-
motion sequences. Consistent with earlier neuroimaging and single-unit studies, this
pattern of results points to the existence of neural mechanisms specialized for analysis of
the kinematics defining biological motion.
Anterior cingulate cortex and response conflict: Effects of response modality and
processing domain

Barch DM, Braver TS, Akbudak E, Conturo T, Ollinger J, Snyder A

Cerebral Cortex 11 (9): 837-848 SEP 2001

Studies of a variety of higher cognitive functions consistently activate a region of anterior

cingulate cortex (ACC), situated posterior to the genu and superior to the corpus
callosum. However, it is not clear whether the same ACC region is activated for all
response modalities (e.g. vocal and manual) and/or all processing domains (e.g. verbal
and spatial). To explore this question, we used rapid event-related functional magnetic
resonance imaging and a spatial Stroop task with conditions tapping both verbal and
spatial processing. We also employed novel methods that allowed us to acquire the
accuracy and reaction times of both manual and vocal responses. We found one large
ACC region that demonstrated significant response conflict effects with both vocal and
manual responses, and three ACC regions that demonstrated significant response conflict
effects with both spatial and verbal processing. We did not find any ACC regions that
demonstrated activity selective to either a specific response modality or processing
domain. Thus, our results suggest that the same regions of ACC are responsive to conflict
arising with both manual and vocal output and with both spatial and verbal processing.

Sustained activity in the medial wall during working memory delays

Petit L, Courtney SM, Ungerleider LG, Haxby JV

Journal Of Neuroscience 18 (22): 9429-9437 NOV 15 1998

We have taken advantage of the temporal resolution afforded by functional magnetic

resonance imaging (fMRI) to investigate the role played by medial wall areas in humans
during working memory tasks. We demarcated the medial motor areas activated during
simple manual movement, namely the supplementary motor area (SMA) and the
cingulate motor area (CMA), and those activated during visually guided saccadic eye
movements, namely the supplementary eye field (SEF). We determined the location of
sustained activity over working memory delays in the medial wall in relation to these
functional landmarks during both spatial and face working memory tasks. We identified
two distinct areas, namely the pre-SMA and the caudal part of the anterior cingulate
cortex (caudal-AC), that showed similar sustained activity during both spatial and face
working memory delays. These areas were distinct from and anterior to the SMA, CMA,
and SEF. Both the pre-SMA and caudal-AC activation were identified by a contrast
between sustained activity during working memory delays as compared with sustained
activity during control delays in which subjects were waiting for a cue to make a simple
manual motor response. Thus, the present findings suggest that sustained activity during
working memory delays in both the pre-SMA and caudal-AC does not reflect simple
motor preparation but rather a state of preparedness for selecting a motor response based
on the information held on-line.

The prefrontal cortex: Response selection or maintenance within working memory?

Rowe JB, Toni I, Josephs O, Frackowiak RSJ, Passingham RE

Science 288 (5471): 1656-1660 JUN 2 2000

It is controversial whether the dorsolateral prefrontal cortex is involved in the

maintenance of items in working memory or in the selection of responses. We used event-
related functional magnetic resonance imaging to study the performance of a spatial
working memory task by humans, We distinguished the maintenance of spatial items
from the selection of an item from memory to guide a response. Selection, but not
maintenance, was associated with activation of prefrontal area 46 of the dorsal lateral
prefrontal cortex, In contrast, maintenance was associated with activation of prefrontal
area 8 and the intraparietal cortex The results support a role for the dorsal prefrontal
cortex in the selection of representations. ntis accounts for the fact that this area is
activated both when subjects select between items on working memory tasks and when
they freely select between movements on tasks of willed action.

Selective And Divided Attention During Visual Discriminations Of Shape, Color, And
Speed - Functional-Anatomy By Positron Emission Tomography

Corbetta M, Miezin Fm, Dobmeyer S, Shulman Gl, Petersen Se

Journal Of Neuroscience 11 (8): 2383-2402 AUG 1991

Positron emission tomography (PET) was used to identify the neural systems involved in
discriminating the shape, color, and speed of a visual stimulus under conditions of
selective and divided attention. Psychophysical evidence indicated that the sensitivity for
discriminating subtle stimulus changes in a same-different matching task was higher
when subjects selectively attended to one attribute than when they divided attention
among the attributes.

PET measurements of brain activity indicated that modulations of extrastriate visual

activity were primarily produced by task conditions of selective attention. Attention to
speed activated a region in the left inferior parietal lobule. Attention to color activated a
region in the collateral sulcus and dorsolateral occipital cortex, while attention to shape
activated collateral sulcus (similarly to color), fusiform and parahippocampal gyri, and
temporal cortex along the superior temporal sulcus.

Outside the visual system, selective and divided attention activated nonoverlapping sets
of brain regions. Selective conditions activated globus pallidus, caudate nucleus, lateral
orbitofrontal cortex, posterior thalamus/colliculus, and insular-premotor regions, while
the divided condition activated the anterior cingulate and dorsolateral prefrontal cortex.

The results in the visual system demonstrate that selective attention to different features
modulates activity in distinct regions of extrastriate cortex that appear to be specialized
for processing the selected feature. The disjoint pattern of activations in extravisual brain
regions during selective- and divided-attention conditions also suggests that perceptual
judgments involve different neural systems, depending on attentional strategies.

fMRI evidence for objects as the units of attentional selection

O'Craven KM, Downing PE, Kanwisher N

NATURE 401 (6753): 584-587 OCT 7 1999

Contrasting theories of visual attention emphasize selection by spatial location(1), visual

features (such as motion or colour)(2-4) or whole objects(5,6). Here we used functional
magnetic resonance imaging (fMRI) to test key predictions of the object-based theory,
which proposes that pre-attentive mechanisms segment the visual array into discrete
objects, groups, or surfaces, which serve as targets for visual attention(5-9). Subjects
viewed stimuli consisting of a face transparently superimposed on a house, with one
moving and the other stationary. In different conditions, subjects attended to the face, the
house or the motion. The magnetic resonance signal from each subject's fusiform face
area(10), parahippocampal place area(11) and area MT/MST12 provided a measure of the
processing of faces, houses and visual motion, respectively. Although all three attributes
occupied the same location, attending to one attribute of an object (such as the motion of
a moving face) enhanced the neural representation not only of that attribute but also of
the other attribute of the same object (for example, the face), compared with attributes of
the other object (for example, the house). These results cannot be explained by models in
which attention selects locations or features, and provide physiological evidence that
whole objects are selected even when only one visual attribute is relevant.

Modulation of human visual cortex by crossmodal spatial attention

Macaluso E, Frith CD, Driver J

SCIENCE 289 (5482): 1206-1208 AUG 18 2000

A sudden touch on one hand can improve vision near that hand, revealing crossmodal
Links in spatial attention. It is often assumed that such Links involve only multimodal
neural structures, but unimodal brain areas may also be affected. We tested the effect of
simultaneous visuo-tactile stimulation on the activity of the human visual cortex. Tactile
stimulation enhanced activity in the visual cortex, but only when it was on the same side
as a visual target. Analysis of effective connectivity between brain areas suggests that
touch influences unimodal visual cortex via back-projections from multimodal parietal
areas. This provides a neural explanation for crossmodal links in spatial attention.

The generality of parietal involvement in visual attention

Wojciulik E, Kanwisher N

NEURON 23 (4): 747-764 AUG 1999

Functional magnetic resonance imaging (fMRI) was used to determine whether different
kinds of visual attention rely on a common neural substrate. Within one session, subjects
performed three different attention experiments (each comparing an attentionally
demanding task with an easier task using identical stimuli): (1) peripheral shifting, (2)
object matching, and (3) a nonspatial conjunction task. Two areas were activated in all
three experiments: one at the junction of intraparietal and transverse occipital sulci
(IPTO), and another in the anterior intraparietal sulcus (AIPS). These regions are not
simply involved in any effortful task, because they were not activated in a fourth
experiment comparing a difficult language task with an easier control task. Thus, activity
in IPTO and AIPS generalizes across a wide variety of attention-requiring tasks,
supporting the existence of a common neural substrate underlying multiple modes of
visual selection.

Increased activity in human visual cortex during directed attention in the absence of
visual stimulation

Kastner S, Pinsk MA, De Weerd P, Desimone R, Ungerleider LG

NEURON 22 (4): 751-761 APR 1999

When subjects direct attention to a particular location in a visual scene, responses in the
visual cortex to stimuli presented at that location are enhanced, and the suppressive
influences of nearby distracters are reduced. What is the top-down signal that modulates
the response to an attended versus an unattended stimulus? Here, we demonstrate
increased activity related to attention in the absence of visual stimulation in extrastriate
cortex when subjects covertly directed attention to a peripheral location expecting the
onset of visual stimuli. Frontal and parietal areas showed a stronger signal increase
during this expectation than did visual areas. The increased activity in visual cortex in the
absence of visual stimulation may reflect a top-down bias of neural signals in favor of the
attended location, which derives from a fronto-parietal network.

The role of working memory in visual selective attention

de Fockert JW, Rees G, Frith CD, Lavie N

SCIENCE 291 (5509): 1803-1806 MAR 2 2001

The hypothesis that working memory is crucial for reducing distraction by maintaining
the prioritization of relevant information was tested in neuroimaging and psychological
experiments with humans. Participants performed a selective attention task that required
them to ignore distracter faces while holding in working memory a sequence of digits that
were in the same order (Low memory Load) or a different order thigh memory Load) on
every trial. Higher memory Load, associated with increased prefrontal activity, resulted in
greater interference effects on behavioral performance from the distracter faces, plus
increased face-related activity in the visual cortex. These findings confirm a major role
for working memory in the control of visual selective attention.

Modulation of connectivity in visual pathways by attention: Cortical interactions

evaluated with structural equation modelling and fMRI

Buchel C, Friston KJ

CEREBRAL CORTEX 7 (8): 768-778 DEC 1997

Electrophysiological and neuroimaging studies have shown that attention to visual

motion can increase the responsiveness of the motion-selective cortical area V5 and the
posterior parietal cortex (PP). Increased or decreased activation in a cortical area is often
attributed to attentional modulation of the cortical projections to that area. This leads to
the notion that attention is associated with changes in connectivity. We have addressed
attentional modulation of effective connectivity using functional magnetic resonance
imaging (fMRI). Three subjects were scanned under identical stimulus conditions (visual
motion) while varying only the attentional component of the task. Haemodynamic
responses defined an occipito-parieto-frontal network, including the, primary visual
cortex (V1), V5 and PP. A structural equation model of the interactions among these
dorsal visual pathway areas revealed increased connectivity between V5 and PP related to
attention. On the basis of our analysis and the neuroanatomical pattern of projections
from the prefrontal cortex to PR we attributed the source of modulatory influences, on the
posterior visual pathway, to the prefrontal cortex (PFC). To test this hypothesis we
included the PFC in our model as a 'modulator' of the pathway between V5 and PR using
interaction terms in the structural equation model. This analysis revealed a significant
modulatory effect of prefrontal regions on V5 afferents to posterior parietal cortex.