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SOKENDAI physiology lecture course 2013
"Neuroscience of Cognition and Motor control" #4
"Attention and Motor Control"
May 24, Friday 10:00-12:00, Myodaiji Staff Hall 2F Meeting room
Masatoshi Yoshida (NIPS and SOKENDAI, Assistant professor)
1. Introduction
1-1. What is attention? -- Let’s start from
examples
• We cannot consciously perceive a big change
in a scene when we do not pay attention to it.
• Attention has a strong influence on what we
see.
1-2. Definition of attention
• William James (Principles of Psychology
(1890)): "Everyone knows what attention is. It
is the taking possession by the mind in clear
and vivid form, of one out of what seem
several simultaneously possible objects...It
implies withdrawal from some things in order to
deal effectively with others..."
1.3 Taxonomy of attention
• Two kinds of attention
• Selective attention: ability to focus on
positions or objects
• Sustained attention: alertness, ability to
concentrate
• What drives selective attention?
• Bottom-up: stimulus-driven (pre-attentive,
pop-out)
• Top-down: goal-directed
• What is selected in selective attention?
• Position: spatial attention
• Object (feature): object (feature)-based
attention
• Bottom-up vs. top-down attention in pre-cue
task
• Bottom-up attention1
• Top-down attention
2. Attention and eye movements
2-1. Neural network for attention and
saccadic eye movements
• Brain regions involved in bottom-up attention
• Brain regions involved in top-down attention2
• Brain regions involved in saccadic eye
movements
2-2. Overt attention and covert attention in
human
• Overt attention and Covert attention3
• Overt selection of visual features contained
within a portrait as revealed by the scanning
eye movements.
• The female monkey on the left is looking
straight ahead. However, it is apparent that
she is fully attending to her neighbor.
• Overt attention and covert attention activate
the same brain regions. 4,5
2-3. Overt attention and covert attention in
monkey
• Saccade control system
• FEF: frontal eye field (premotor cortex for
eye)
• SC: superior colliculus (one of the midbrain
structures)
• FEF is involved in covert attention6
• Electrical stimulation to FEF directs covert
attention without evoking saccades.
• SC is involved in covert attention7
• Inactivation of SC leads to deficits in covert
attention but does not affect MT and MST.
• The effect of inactivation is not an indirect
effect on MT/MST but a direct effect on SC
(or FEF).
• Cortex is not necessary for attention!
3. Dorsal and ventral attention network
3-1. Dorsal and ventral attention network in
human
• When top-down attention is manipulated by
pre-cue, both dorsal and ventral attention
systems are activated.8
• Note these are NOT the dorsal and ventral
visual pathway
• Resting state BOLD activity8
• Strong and significant positive temporal
correlation of spontaneous activity at rest.
• The dorsal network is anti-correlated with the
default network.
3-2. Spatial hemineglect as a disorder of
attentional network
• What is 'spatial hemineglect'?
• The inability to report, respond, or orient to
stimuli in the contralesional space.
• The deficit must not be fully attributable to
primary sensory deficits (e.g., hemianopia) or
motor disturbance (e.g., hemiparesis).
• Neglect occurs more commonly in those with
brain injury affecting the right cortical
hemisphere.
• Where is the key lesion site?
• Hemineglect as a disorder of network9
• Activity of MFG and STS shows a sign of
recovery in the chronic period.
• However, functional connectivity did not
recover in the chronic period.
3-3. Ventral attention network in monkey?
• The ventral attention system (TPJ-IFG) is not
identified in monkeys.
• Monkeys have no BA40 (TPJ/SMG).
• (Putative BA40 is part of BA7).
• The dorsal network may be mediated by SLFII.
• SLF(the superior longitudinal fasciculus)
• SLFII: spatial attention and spatial
awareness
• The ventral network may be mediated by AF
(the arcuate fasciculus).10
• AF connection is very sparse in macaque11
4. Saliency computational model
4-1. What is saliency map?
• What is saliency?
• salient (adj): “very easy to notice”
• visual salience (or visual saliency): the
distinct subjective perceptual quality which
makes some items in the world stand out
from their neighbors and immediately grab
our attention.(Scholarpedia by Laurent Itti)
• What grabs our attention?
• Where is the object that is different from the
rest?
• Feature integration theory12
• Original computational model13
• A unique saliency map, independent of
features
• Winner-take-all rule (WTA)
• Selection was made on the saliency map
• What is saliency map?
• An explicit 2D map that encodes the saliency
of objects in the visual environment
• A computational concept
• Saliency computational model14
• Iterative calculation of center-surround
differentiation and normalization
• Intracortical lateral inhibition is
mathematically equivalent to second
derivative (=laplacian). It is used for edge
detection.
4-2. Applications of the saliency model
• Application #1: Modeling visual search
• Application #2: Advertisement
• AD people collect eye-tracking data to
evaluate the effect of advertisement.
• But it is expensive and time-consuming.
• You do not need to collect eye-tracing data if
you use the saliency model instead.
• Now it is commercially available.
4-3. Saliency map in the brain
• Brain network involved in saliency
4-4: Saliency map in the brain - Parietal
cortex
• Temporal saliency drives LIP neurons15
• Visual information (stimuli in RF) is
sufficient?
• Or saliency (abrupt onset of stimuli) is
necessary?
• Stimuli with low saliency (stable stimuli) did
not drive a LIP neuron.
• Visual information (stimuli in RF) is not
enough!
• Stimuli with high salience (recently flashed
stimuli) drive a LIP neuron.
• LIP represents visual salience.
• Decoding saliency from the brain16
• The difference of saliency in four quadrants
can be decoded from pIPS and early visual
cortex.
4-5: Saliency map in the brain - Is V1
necessary?
• Computational model predicts monkey’s gaze
• Salient stimuli attract gazes of blindsight
monkeys17
• The gaze positions have higher saliency,
than expected from random eye movement.
• Revised view of saliency computation
4-6: Clinical application 1 - ADHD
• Classification of neurological disorders18
• Eye-tracking (20min) is sufficient to classify
subjects, if combined with the saliency
model.
4-7: Clinical application 2 - autism
• Autistic people do not look at eyes. But why?
• Visual saliency of eyes and mouth matters.19
• The authors generated many images with
different saliency for eyes and mouth.
• The autistic people see the mouth even
when the saliency is low.
• The autistic people do not avoid the eyes.
Rather, they preferentially look at the mouth.
4-8: Clinical application 3 - schizophrenia
• Aberrant salience hypothesis of psychosis20
5. Summary
• Attention and eye movements share the same
brain circuitry.
• Dorsal and ventral attentional networks are
identified in humans (but not yet in monkeys).
• Saliency computational model is able to
predict gaze of humans and monkeys.
References
1. Ikeda, T., Yoshida, M. & Isa, T. Lesion of primary
visual cortex in monkey impairs the inhibitory but
not the facilitatory cueing effect on saccade.
Journal of Cognitive Neuroscience 23, 1160–1169
(2011).
2. Baluch, F. & Itti, L. Mechanisms of top-down
attention. Trends in Neurosciences 34, 210–224
(2011).
3. Moore, T., Armstrong, K. M. & Fallah, M.
Visuomotor origins of covert spatial attention.
Neuron 40, 671–683 (2003).
4. Beauchamp, M. S., Petit, L., Ellmore, T. M.,
Ingeholm, J. & Haxby, J. V. A parametric fMRI
study of overt and covert shifts of visuospatial
attention. NeuroImage 14, 310–321 (2001).
5. de Haan, B., Morgan, P. S. & Rorden, C. Covert
orienting of attention and overt eye movements
activate identical brain regions. Brain Research
1204, 102–111 (2008).
6. Moore, T. & Fallah, M. Control of eye movements
and spatial attention. Proc Natl Acad Sci USA 98,
1273–1276 (2001).
7. Zénon, A. & Krauzlis, R. J. Attention deficits without
cortical neuronal deficits. Nature 489, 434–437
(2012).
8. Corbetta, M. & Shulman, G. L. Spatial neglect and
attention networks. Annu. Rev. Neurosci. 34,
569–599 (2011).
9. He, B. J. et al. Breakdown of Functional
Connectivity in Frontoparietal Networks Underlies
Behavioral Deficits in Spatial Neglect. Neuron 53,
905–918 (2007).
10. Petrides, M. & Pandya, D. N. Distinct Parietal and
Temporal Pathways to the Homologues of Broca's
Area in the Monkey. Plos Biol 7, e1000170 (2009).
11. Rilling, J. K. et al. The evolution of the arcuate
fasciculus revealed with comparative DTI. Nat
Neurosci 11, 426–428 (2008).
12. Treisman, A. Feature binding, attention and object
perception. Philosophical Transactions of the Royal
Society B: Biological Sciences 353, 1295–1306
(1998).
13. Koch, C. & Ullman, S. Shifts in selective visual
attention: towards the underlying neural circuitry.
Hum Neurobiol 4, 219–227 (1984).
14. Itti, L. & Koch, C. Computational modelling of visual
attention. Nat Rev Neurosci 2, 194–203 (2001).
15. Gottlieb, J. P., KUSUNOKI, M. & Goldberg, M. E.
The representation of visual salience in monkey
parietal cortex. Nature 391, 481–484 (1998).
16. Bogler, C., Bode, S. & Haynes, J.-D. Decoding
successive computational stages of saliency
processing. Curr. Biol. 21, 1667–1671 (2011).
17. Yoshida, M. et al. Residual attention guidance in
blindsight monkeys watching complex natural
scenes. Current Biology 22, 1429–1434 (2012).
18. Tseng, P.-H. et al. High-throughput classification of
clinical populations from natural viewing eye
movements. J Neurol 1–10 (2012).
doi:10.1007/s00415-012-6631-2
19. Neumann, D., Spezio, M. L., Piven, J. & Adolphs, R.
Looking you in the mouth: abnormal gaze in autism
resulting from impaired top-down modulation of
visual attention. Social Cognitive and Affective
Neuroscience 1, 194–202 (2006).
20. Kapur, S. How antipsychotics become
anti-‘psychotic’--from dopamine to salience to
psychosis. Trends Pharmacol. Sci. 25, 402–406
(2004).

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総研大講義 「#4 注意と運動」レジメ

  • 1. SOKENDAI physiology lecture course 2013 "Neuroscience of Cognition and Motor control" #4 "Attention and Motor Control" May 24, Friday 10:00-12:00, Myodaiji Staff Hall 2F Meeting room Masatoshi Yoshida (NIPS and SOKENDAI, Assistant professor) 1. Introduction 1-1. What is attention? -- Let’s start from examples • We cannot consciously perceive a big change in a scene when we do not pay attention to it. • Attention has a strong influence on what we see. 1-2. Definition of attention • William James (Principles of Psychology (1890)): "Everyone knows what attention is. It is the taking possession by the mind in clear and vivid form, of one out of what seem several simultaneously possible objects...It implies withdrawal from some things in order to deal effectively with others..." 1.3 Taxonomy of attention • Two kinds of attention • Selective attention: ability to focus on positions or objects • Sustained attention: alertness, ability to concentrate • What drives selective attention? • Bottom-up: stimulus-driven (pre-attentive, pop-out) • Top-down: goal-directed • What is selected in selective attention? • Position: spatial attention • Object (feature): object (feature)-based attention • Bottom-up vs. top-down attention in pre-cue task • Bottom-up attention1 • Top-down attention 2. Attention and eye movements 2-1. Neural network for attention and saccadic eye movements • Brain regions involved in bottom-up attention • Brain regions involved in top-down attention2 • Brain regions involved in saccadic eye movements 2-2. Overt attention and covert attention in human • Overt attention and Covert attention3 • Overt selection of visual features contained within a portrait as revealed by the scanning eye movements. • The female monkey on the left is looking straight ahead. However, it is apparent that she is fully attending to her neighbor. • Overt attention and covert attention activate the same brain regions. 4,5 2-3. Overt attention and covert attention in monkey • Saccade control system • FEF: frontal eye field (premotor cortex for eye) • SC: superior colliculus (one of the midbrain structures) • FEF is involved in covert attention6 • Electrical stimulation to FEF directs covert attention without evoking saccades. • SC is involved in covert attention7 • Inactivation of SC leads to deficits in covert attention but does not affect MT and MST. • The effect of inactivation is not an indirect
  • 2. effect on MT/MST but a direct effect on SC (or FEF). • Cortex is not necessary for attention! 3. Dorsal and ventral attention network 3-1. Dorsal and ventral attention network in human • When top-down attention is manipulated by pre-cue, both dorsal and ventral attention systems are activated.8 • Note these are NOT the dorsal and ventral visual pathway • Resting state BOLD activity8 • Strong and significant positive temporal correlation of spontaneous activity at rest. • The dorsal network is anti-correlated with the default network. 3-2. Spatial hemineglect as a disorder of attentional network • What is 'spatial hemineglect'? • The inability to report, respond, or orient to stimuli in the contralesional space. • The deficit must not be fully attributable to primary sensory deficits (e.g., hemianopia) or motor disturbance (e.g., hemiparesis). • Neglect occurs more commonly in those with brain injury affecting the right cortical hemisphere. • Where is the key lesion site? • Hemineglect as a disorder of network9 • Activity of MFG and STS shows a sign of recovery in the chronic period. • However, functional connectivity did not recover in the chronic period. 3-3. Ventral attention network in monkey? • The ventral attention system (TPJ-IFG) is not identified in monkeys. • Monkeys have no BA40 (TPJ/SMG). • (Putative BA40 is part of BA7). • The dorsal network may be mediated by SLFII. • SLF(the superior longitudinal fasciculus) • SLFII: spatial attention and spatial awareness • The ventral network may be mediated by AF (the arcuate fasciculus).10 • AF connection is very sparse in macaque11 4. Saliency computational model 4-1. What is saliency map? • What is saliency? • salient (adj): “very easy to notice” • visual salience (or visual saliency): the distinct subjective perceptual quality which makes some items in the world stand out from their neighbors and immediately grab our attention.(Scholarpedia by Laurent Itti) • What grabs our attention? • Where is the object that is different from the rest? • Feature integration theory12 • Original computational model13 • A unique saliency map, independent of features • Winner-take-all rule (WTA) • Selection was made on the saliency map • What is saliency map? • An explicit 2D map that encodes the saliency of objects in the visual environment • A computational concept • Saliency computational model14 • Iterative calculation of center-surround differentiation and normalization • Intracortical lateral inhibition is mathematically equivalent to second derivative (=laplacian). It is used for edge detection. 4-2. Applications of the saliency model • Application #1: Modeling visual search • Application #2: Advertisement • AD people collect eye-tracking data to evaluate the effect of advertisement. • But it is expensive and time-consuming. • You do not need to collect eye-tracing data if
  • 3. you use the saliency model instead. • Now it is commercially available. 4-3. Saliency map in the brain • Brain network involved in saliency 4-4: Saliency map in the brain - Parietal cortex • Temporal saliency drives LIP neurons15 • Visual information (stimuli in RF) is sufficient? • Or saliency (abrupt onset of stimuli) is necessary? • Stimuli with low saliency (stable stimuli) did not drive a LIP neuron. • Visual information (stimuli in RF) is not enough! • Stimuli with high salience (recently flashed stimuli) drive a LIP neuron. • LIP represents visual salience. • Decoding saliency from the brain16 • The difference of saliency in four quadrants can be decoded from pIPS and early visual cortex. 4-5: Saliency map in the brain - Is V1 necessary? • Computational model predicts monkey’s gaze • Salient stimuli attract gazes of blindsight monkeys17 • The gaze positions have higher saliency, than expected from random eye movement. • Revised view of saliency computation 4-6: Clinical application 1 - ADHD • Classification of neurological disorders18 • Eye-tracking (20min) is sufficient to classify subjects, if combined with the saliency model. 4-7: Clinical application 2 - autism • Autistic people do not look at eyes. But why? • Visual saliency of eyes and mouth matters.19 • The authors generated many images with different saliency for eyes and mouth. • The autistic people see the mouth even when the saliency is low. • The autistic people do not avoid the eyes. Rather, they preferentially look at the mouth. 4-8: Clinical application 3 - schizophrenia • Aberrant salience hypothesis of psychosis20 5. Summary • Attention and eye movements share the same brain circuitry. • Dorsal and ventral attentional networks are identified in humans (but not yet in monkeys). • Saliency computational model is able to predict gaze of humans and monkeys. References 1. Ikeda, T., Yoshida, M. & Isa, T. Lesion of primary visual cortex in monkey impairs the inhibitory but not the facilitatory cueing effect on saccade. Journal of Cognitive Neuroscience 23, 1160–1169 (2011). 2. Baluch, F. & Itti, L. Mechanisms of top-down attention. Trends in Neurosciences 34, 210–224 (2011). 3. Moore, T., Armstrong, K. M. & Fallah, M. Visuomotor origins of covert spatial attention. Neuron 40, 671–683 (2003). 4. Beauchamp, M. S., Petit, L., Ellmore, T. M., Ingeholm, J. & Haxby, J. V. A parametric fMRI study of overt and covert shifts of visuospatial attention. NeuroImage 14, 310–321 (2001). 5. de Haan, B., Morgan, P. S. & Rorden, C. Covert orienting of attention and overt eye movements activate identical brain regions. Brain Research 1204, 102–111 (2008). 6. Moore, T. & Fallah, M. Control of eye movements and spatial attention. Proc Natl Acad Sci USA 98, 1273–1276 (2001). 7. Zénon, A. & Krauzlis, R. J. Attention deficits without cortical neuronal deficits. Nature 489, 434–437 (2012). 8. Corbetta, M. & Shulman, G. L. Spatial neglect and attention networks. Annu. Rev. Neurosci. 34, 569–599 (2011). 9. He, B. J. et al. Breakdown of Functional Connectivity in Frontoparietal Networks Underlies Behavioral Deficits in Spatial Neglect. Neuron 53, 905–918 (2007). 10. Petrides, M. & Pandya, D. N. Distinct Parietal and Temporal Pathways to the Homologues of Broca's Area in the Monkey. Plos Biol 7, e1000170 (2009).
  • 4. 11. Rilling, J. K. et al. The evolution of the arcuate fasciculus revealed with comparative DTI. Nat Neurosci 11, 426–428 (2008). 12. Treisman, A. Feature binding, attention and object perception. Philosophical Transactions of the Royal Society B: Biological Sciences 353, 1295–1306 (1998). 13. Koch, C. & Ullman, S. Shifts in selective visual attention: towards the underlying neural circuitry. Hum Neurobiol 4, 219–227 (1984). 14. Itti, L. & Koch, C. Computational modelling of visual attention. Nat Rev Neurosci 2, 194–203 (2001). 15. Gottlieb, J. P., KUSUNOKI, M. & Goldberg, M. E. The representation of visual salience in monkey parietal cortex. Nature 391, 481–484 (1998). 16. Bogler, C., Bode, S. & Haynes, J.-D. Decoding successive computational stages of saliency processing. Curr. Biol. 21, 1667–1671 (2011). 17. Yoshida, M. et al. Residual attention guidance in blindsight monkeys watching complex natural scenes. Current Biology 22, 1429–1434 (2012). 18. Tseng, P.-H. et al. High-throughput classification of clinical populations from natural viewing eye movements. J Neurol 1–10 (2012). doi:10.1007/s00415-012-6631-2 19. Neumann, D., Spezio, M. L., Piven, J. & Adolphs, R. Looking you in the mouth: abnormal gaze in autism resulting from impaired top-down modulation of visual attention. Social Cognitive and Affective Neuroscience 1, 194–202 (2006). 20. Kapur, S. How antipsychotics become anti-‘psychotic’--from dopamine to salience to psychosis. Trends Pharmacol. Sci. 25, 402–406 (2004).