Mechanisms underlying feature selectivity in primary sensory - - PowerPoint PPT Presentation
Mechanisms underlying feature selectivity in primary sensory - - PowerPoint PPT Presentation
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
In other to get access to the speaker’s comments, click on: Comments\show comments list
Kaniza Triangle
0o 90oL 90oR A
AL AR
LGN Optic tract Optic radiation pulvinar pretectum
- Post. commissure
Medial geniculate caudate fornix
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
ON-center and OFF-center LGN cells
Retinotopic organization – “simple” but “distorted”
Anterograde transsynaptic degeneration due to loss of right eye
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
Waking state
The gate is open Thalamic reticular nucleus (RNT)
Sleep
The gate is closed
M-cells
- orientation selective
- direction selective
- stereoscopically sel.
- just like LGN Mcell RFs
To extrastriate cortex (MT) pia white matter
Orientation selectivity of cortical cells
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
Orientation selectivity
Hubel and Wiesel, 1962
IVCalpha IVB
The Feedforward Model of Orientation Selectivity in Primary Visual Cortex
Contrast Invariance of Orientation Tuning
Contrast Invariance of Orientation Tuning
Trial-to-Trial Response Variability and the Origin of Contrast- Invariant Orientation Tuning in Simple Cells
The Biophysical Mechanisms Underlying stimulus dependant changes in spike threshold of V1 Simple Cells
INA ANA
The Biophysical Mechanisms Underlying the Response Properties of V1 Simple Cells
Lateral connectivity! (similar to retina)
Reference RF # Reference RF recorded at the asterisk doesn’t change
Kaniza Triangle a b c
27
Schizophrenia and Visual Discrimination
Spencer et al., Proc Nat Acad Sci. 101:17288-17293, 2004 Normal Schiz
Cat V1 Layer IV simple cell
FF > 1.0
Precise but not reliable Reliable but not precise
Mainen and Sejnowski, 1995
Biophysical Limits?
CNQX + APV + bicuculline
Reinagel and Reid, 2000
Synaptic Limits? Lateral Geniculate Nucleus
Stone et al., 1979
What about visual cortex?
What about visual cortex?
LGN Layer IV
Excitatory Synapse
Bright Stimulus LGN Layer IV
Excitatory Synapse Inhibitory Synapse (via interneuron, not shown)
Bright Stimulus Push-Pull Push only
A better stimulus: contrast modulated Gabor patch
Spatial location Spatial frequency Orientation Spatial Phase Length Width
Jones and Palmer, 1987a,b,c
Optimization:
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).
X = 46 Y = 24 S = 50 Number of events
Key question
- How is the cortex able to maintain the precision of
its principle inputs? The stimulus itself – produces synchronous activation of LGN cells
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
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)
Σ Σ
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.
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
Excite two non-overlapping populations of LGN afferents at different times relative to each other.
Methods A B
Facilitation and suppression of the response Bar A suppresses Bar B
Intracellular recording (sharpies)
Intracellular recording (sharpies)
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)
- 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……