Responses in Macaque Visual Area V4 following Inactivation of the
Visual Neuron Responses
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Transcript of Visual Neuron Responses
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Visual Neuron Responses
• This conceptualization of the visual system was “static” - it did not take into account the possibility that visual cells might change their response selectivity over time and it was firmly based in the classical notion of a receptive field
– Logic went like this: if the cell is firing, its preferred line/edge must be present and…
– if the preferred line/edge is present, the cell must be firing
• We will encounter examples in which these don’t apply!
• Representing boundaries and surfaces must be more complicated than simple edge detection! WHY??
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What can a visual neuron “know” about the image?
• If a neuron is only an edge detector and/or only has a small receptive field, it can’t represent information about the relationship between the contents of its receptive field and other receptive fields elsewhere in the display.
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What can a visual neuron “know” about the image?
• If a neuron is only an edge detector and/or only has a small receptive field, it can’t represent information about the relationship between the contents of its receptive field and other receptive fields elsewhere in the display.
• For example the famous 1961 Rosebowl hoax…no single person could know what the big picture showed
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What can a visual neuron “know” about the image?
• If a neuron is only an edge detector and/or only has a small receptive field, it can’t represent information about the relationship between the contents of its receptive field and other receptive fields elsewhere in the display.
• Also the 2004 Harvard – Yale Game:
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Visual Neuron Responses
• Edges are important because they are the boundaries between objects and the background or objects and other objects
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Visual Neuron Responses
• Boundaries between objects can be defined by color rather than brightness
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Visual Neuron Responses
• Boundaries between objects can be defined by texture
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Visual Neuron Responses
• Boundaries between objects can be defined by motion and depth cues
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Visual Neuron Responses
• Boundaries between objects can be defined by motion and depth cues
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Feed-Forward and Feed-Back Processing in the Visual System
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The Feed-Forward Sweep
• What is the feed-forward sweep?
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The Feed-Forward Sweep
• The feed-forward sweep is the initial response of each visual area “in turn” as information is passed to it from a “lower” area
• Characteristics:– a single spike per synapse
– no time for lateral connections
– no time for feedback connections
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The Feed-Forward Sweep
• The feed-forward sweep is the initial response of each visual area “in turn” as information is passed to it from a “lower” area
• What does it mean for an area to be “lower” or “higher”
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The Feed-Forward Sweep
• Hierarchy of visual cortical areas defined anatomically
Dorsal “where”/”how”
Ventral “what”
Notice the direct connection from SC to MT/V5
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The Feed-Forward Sweep
• Hierarchy can be defined more functionaly
• The feed-forward sweep is the initial response of each visual area “in turn” as information is passed to it from a “lower” area
• Consider the latencies of the first responses in various areas
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The Feed-Forward Sweep
• Thus the “hierarchy” of visual areas differs depending on temporal or anatomical features
• aspects of the visual system account for this fact:
– multiple feed-forward sweeps progressing at different rates (I.e. magno and parvo pathways) in parallel
• M pathway is myelinated
• P pathway is not
– signals arrive at cortex via routes other than the Geniculo-striate pathway (LGN to V1)
• Will be important in understanding blindsight
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The Feed-Forward Sweep
• The feed-forward sweep gives rise to the “classical” receptive field properties– tuning properties exhibited in very first spikes
• Orientation tuning in V1
• Optic flow tuning in MST
– think of cortical neurons as “detectors” only during feed-forward sweep
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After the Forward Sweep
• By 150 ms, virtually every visual brain area has responded to the onset of a visual stimulus
• But visual cortex neurons continue to fire for hundreds of milliseconds!
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After the Forward Sweep
• By 150 ms, virtually every visual brain area has responded to the onset of a visual stimulus
• But visual cortex neurons continue to fire for hundreds of milliseconds!
• What are they doing?
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After the Forward Sweep
• By 150 ms, virtually every visual brain area has responded to the onset of a visual stimulus
• But visual cortex neurons continue to fire for hundreds of milliseconds!
• What are they doing?
• with sufficient time (a few tens of ms) neurons begin to reflect aspects of cognition other than “detection”
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Extra-RF Influences
• One thing they seem to be doing is helping each other figure out what aspects of the entire scene are contained within a given receptive field
– That is, the responses of visual neurons begin to change to reflect global rather than local features of the scene
– recurrent signals sent via feedback projections are thought to mediate these later properties
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Extra-RF Influences
Note that these are responses to the same stimulus!
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Extra-RF Influences
• consider texture-defined boundaries
– classical RF tuning properties do not allow neuron to know if RF contains figure or background
– At progressively later latencies, the neuron responds differently depending on whether it is encoding boundaries, surfaces, the background, etc.
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Extra-RF Influences
• Consider this analogy:
– Imagine when each fan puts up a card he or she is told to shake it – so that the entire scene is full of shaking cards
– After some delay, the fans holding up the red cards are told to keep shaking but the fans holding white cards are told to stop…the words will be enhanced
– But the fans can’t each figure that out on their own because they don’t actually know the color of the card they are holding
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Extra-RF Influences
• How do these data contradict the notion of a “classical” receptive field?
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Extra-RF Influences
• How do these data contradict the notion of a “classical” receptive field?
• Remember that for a classical receptive field (i.e. feature detector):
– If the neuron’s preferred stimulus is present in the receptive field, the neuron should fire a stereotypical burst of APs
– If the neuron is firing a burst of APs, its preferred stimulus must be present in the receptive field
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Extra-RF Influences
• How do these data contradict the notion of a “classical” receptive field?
• Remember that for a classical receptive field (i.e. feature detector):
– If the neuron’s preferred stimulus is present in the receptive field, the neuron should fire a stereotypical burst of APs
– If the neuron is firing a burst of APs, its preferred stimulus must be present in the receptive field
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Recurrent Signals in Object Perception
• Can a neuron represent whether or not its receptive field is on part of an attended object?
• What if attention is initially directed to a different part of the object?
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Recurrent Signals in Object Perception
• Can a neuron represent whether or not its receptive field is on part of an attended object?
• What if attention is initially directed to a different part of the object?
Yes, but not during the feed-forward sweep
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Recurrent Signals in Object Perception
• curve tracing
– monkey indicates whether a particular segment is on a particular curve
– requires attention to scan the curve and “select” all segments that belong together
– that is: make a representation of the entire curve
– takes time
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Recurrent Signals in Object Perception
• curve tracing
– neuron begins to respond differently at about 200 ms
– enhanced firing rate if neuron is on the attended curve
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Feedback Signals and the binding problem
• What is the binding problem?
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Feedback Signals and the binding problem
• What is the binding problem?
• curve tracing and the binding problem:
– if all neurons with RFs over the attended curve spike faster/at a specific frequency/in synchrony, this might be the binding signal
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Feedback Signals and the binding problem
• So what’s the connection between Attention and Recurrent Signals?
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Feedback Signals and Attention
• One theory is that attention (attentive processing) entails the establishing of recurrent “loops”
• This explains why attentive processing takes time - feed-forward sweep is insufficient
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Feedback Signals and Attention
• Instruction cues (for example in the Posner Cue-Target paradigm) may cause feedback signal prior to stimulus onset (thus prior to feed-forward sweep)
• think of this as pre-setting the system for the upcoming stimulus
• What does this accomplish?
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Feedback Signals and Attention
• What does this accomplish?
• Preface to attention: Two ways to think about attention– Attention improves perception, acts as a gateway to memory
and consciousness
– Attention is a mechanism that routes information through the brain
• It is the brain actively reconfiguring itself by changing the way signals propagate through networks
• It is a form of very fast, very transient plasticity
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Feedback Signals and Attention• Put another way:
– It may strike you as remarkable that a single visual stimulus should “activate” so many brain areas so rapidly
– In fact it should be puzzling that a visual input doesn’t create a runaway “chain reaction”
• The brain is massively interconnected
• Why shouldn’t every neuron respond to a visual stimulus
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Feedback Signals and Attention
• We’ll consider the role of feedback signals in attention in more detail as we discuss the neuroscience of attention