Mechanisms underlying feature selectivity in primary sensory - - PowerPoint PPT Presentation

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Mechanisms underlying feature selectivity in primary sensory cortices Thomas Bessah In other to get access to the speakers comments, click on: Comments\show comments list Kaniza Triangle 0 o A 90 o L 90 o R A L A R pulvinar pretectum


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Mechanisms underlying feature selectivity in primary sensory cortices

Thomas Bessaïh

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In other to get access to the speaker’s comments, click on: Comments\show comments list

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Kaniza Triangle

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0o 90oL 90oR A

AL AR

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LGN Optic tract Optic radiation pulvinar pretectum

  • Post. commissure

Medial geniculate caudate fornix

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1,2 = magnocellular (M); 3,4,5,6 = parvocellular (P) 1,4,6 = contralateral; 2,3,5 = ipsilateral ON and OFF cells separated in P layers (not M layers) Six layers Why? Optic tract

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ON-center and OFF-center LGN cells

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Retinotopic organization – “simple” but “distorted”

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Anterograde transsynaptic degeneration due to loss of right eye

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LGN receptive fields: same as in retina M cells (10%)– large, motion sensitive, no color selectivity P cells (80%) – small, center-surround, color selective

In foveal representation the pathway is one to one: RGC LGN Layer IVC of area 17

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Waking state

The gate is open Thalamic reticular nucleus (RNT)

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Sleep

The gate is closed

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M-cells

  • orientation selective
  • direction selective
  • stereoscopically sel.
  • just like LGN Mcell RFs

To extrastriate cortex (MT) pia white matter

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Orientation selectivity of cortical cells

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On-center RF of cell in IVB Off-center RF of cell in IVB RGC LGN IVCα IVB

Retina thalamus striate cortex

M-cell pathway

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Orientation selectivity

Hubel and Wiesel, 1962

IVCalpha IVB

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The Feedforward Model of Orientation Selectivity in Primary Visual Cortex

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Contrast Invariance of Orientation Tuning

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Contrast Invariance of Orientation Tuning

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Trial-to-Trial Response Variability and the Origin of Contrast- Invariant Orientation Tuning in Simple Cells

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The Biophysical Mechanisms Underlying stimulus dependant changes in spike threshold of V1 Simple Cells

INA ANA

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The Biophysical Mechanisms Underlying the Response Properties of V1 Simple Cells

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Lateral connectivity! (similar to retina)

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Reference RF # Reference RF recorded at the asterisk doesn’t change

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Kaniza Triangle a b c

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27

Schizophrenia and Visual Discrimination

Spencer et al., Proc Nat Acad Sci. 101:17288-17293, 2004 Normal Schiz

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Cat V1 Layer IV simple cell

FF > 1.0

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Precise but not reliable Reliable but not precise

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Mainen and Sejnowski, 1995

Biophysical Limits?

CNQX + APV + bicuculline

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Reinagel and Reid, 2000

Synaptic Limits? Lateral Geniculate Nucleus

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Stone et al., 1979

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What about visual cortex?

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What about visual cortex?

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LGN Layer IV

Excitatory Synapse

Bright Stimulus LGN Layer IV

Excitatory Synapse Inhibitory Synapse (via interneuron, not shown)

Bright Stimulus Push-Pull Push only

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A better stimulus: contrast modulated Gabor patch

Spatial location Spatial frequency Orientation Spatial Phase Length Width

Jones and Palmer, 1987a,b,c

Optimization:

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Methods

  • Extracellular recording in vivo, barbiturate anesthetized cat
  • LGN X- and Y-cells, Layer IV Simple cells in Area 17 (V1)
  • Contrast modulated Gabor patch optomized for each cell
  • Contrast drawn from Gaussian distribution (mean = 0, SD = 31%) at 125 Hz
  • Spike timing is acquired with a time resolution of 0.1 ms and spike times are

rebinned using 1 ms bins (typically ISImin > 2 ms, therefore only 1 spike/bin).

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X = 46 Y = 24 S = 50 Number of events

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Key question

  • How is the cortex able to maintain the precision of

its principle inputs? The stimulus itself – produces synchronous activation of LGN cells

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69% 77% 75% 69%

LGN neurons of the same cell type respond similarly given the same Gabor patch sequence.

Suggests: All LGN neurons located within a lobe of simple cell’s receptive field, respond similarly.

Similarity of LGN input across cells and cats

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The responses of on-centered, LGN neurons are similar to the responses of off-centered, LGN neurons to inverted version of the same stimulus. Similarity of LGN ON-NS/OFF-IS inputs to Cortex

Suggests: Our stimulus synchronously activates all the LGN afferents of Layer IV simple cells.

70% 75%

Time (s) Time (s) Time (s)

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Σ Σ

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Summed ON-ns and OFF-is LGN Response Simple Cell

Simple cells are as precise as any one of their inputs and more precise than the sum of their inputs.

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Key questions

  • How is the cortex able to maintain the precision of

its principle inputs?

Cortical cells are sensitive to highly synchronous

input from LGN afferents

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Excite two non-overlapping populations of LGN afferents at different times relative to each other.

Methods A B

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Facilitation and suppression of the response Bar A suppresses Bar B

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Intracellular recording (sharpies)

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Intracellular recording (sharpies)

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Supralinear for spikes but sublinear for EPSPs

Can be explained by the accelerating non-linearity in the spike generating mechanism (Reid, et al., 1987; DeAngelis et al., 1993a,b; Anzai et al., 1999; Carandini & Ferster, 2000; Nykamp & Ringach,

2002)

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  • LGN X and Y cells and cortical simple cells have highly stereotyped responses to

stimuli containing rapid transients

  • Simple cells are as precise as any one of their LGN X-cell inputs and more precise

than their putative pooled input

  • Membrane potential in simple cells follows the pooled LGN input closely
  • Biophysical mechanisms related largely to the spike generation in simple cells render

the response to synchronized thalamic input larger and more precise than expected

  • The stimulus…
  • Coincidence detection……