FREQUENCY OF SEEING EXPERIMENTS (Hecht, Shlaer and Pirenne, 1942) PROBLEM: No way to account for false positives (noise)
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2 FREQUENCY OF SEEING EXPERIMENTS (Hecht, Shlaer and Pirenne, 1942) 1 target 8 fixation point target covers ~5 rods % seen Θ Θ = 2 Θ = 7 = photons at cornea CONCLUSION: Θ = 5-7 photons absorbed spread over 5 rods PROBLEM: No way to account for false positives (noise)
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4 IMPLICATIONS OF BEHAVIORAL SENSITIVITY rod phototransduction - single photons reliably transduced rod! bipolar! cell AII! amacrine! cell cone cone! bipolar! cell ganglion! cell synaptic transmission - reliable transmission of single photon responses neural coding - absorption of a few photons produces change in optic nerve activity
5 CONVERGENCE AND SPARSE SIGNALING IN MAMMALIAN RETINA At visual threshold photons <.1% of the rods contribute signals while all rods generate noise Under these conditions averaging is a disaster General problem in nervous system
6 ROD-ROD BIPOLAR SIGNAL TRANSFER AND ABSOLUTE VISUAL SENSITIVITY keep discard Rod-rod bipolar signal transfer is nonlinear. Nonlinear signal transfer eliminates or severely attenuates majority of rodʼs single photon responses. Rejection of noise more than compensates loss of signal - thus rod bipolars provide near-optimal readout of rod signals near visual threshold.
7 SUCTION ELECTRODE RECORDING (Baylor et al., 1979) photocurrent (pa) time (sec)
8 DARK NOISE IN MAMMALIAN RODS (Baylor et al., 1984) 1 pa thermal event ave single Dark + Instrumental Noise 1 pa continuous thermal.2 sec Instrumental Noise 1 pa 2 time (sec) 4 6
9 SEPARATION OF ROD SIGNAL AND NOISE BY THRESHOLDING NONLINEARITY photocurrent 1 pa keep throw away flash times time (sec) 14 (modified from Wilson, 22)
10 keep discard Mouse rod-rod bipolar signal transfer is nonlinear. - dependence of response on flash strength - discreteness of dim flash response Nonlinear signal transfer eliminates or severely attenuates majority of rodʼs single photon responses. Rejection of noise more than compensates loss of signal - thus rod bipolars provide near-optimal readout of rod signals near visual threshold.
11 DIM FLASH RESPONSES OF RODS GROW LINEARLY WITH FLASH STRENGTH pa -5-1 log(r/r max ) -1 rods sec log(rh*) rod bipolar why log-log?
12 RESPONSES OF ROD BIPOLARS BUT NOT RODS GROW SUPRALINEARLY WITH FLASH STRENGTH pa -5-1 log(r/r max ) -1 rods sec log(rh*) rod bipolar pa log(r/r max ) time (sec) log(rh*/rod)
13 photocurrent 1 pa flash times time (sec) 14 keep throw away Does rod-rod bipolar signal transfer separate rod signal and noise?
14 MOUSE ROD SINGLE PHOTON RESPONSES ARE PARTIALLY OBSCURED BY NOISE 2 pa # responses Rh* rods..2 sec rod bipolar
15 ROD BIPOLARS GENERATE DISCRETE RESPONSES TO DIM FLASHES 2 pa # responses Rh* rods..2 sec rod bipolar 2 pa # responses Rh*/rod..2 sec
16 keep discard Mouse rod-rod bipolar signal transfer is nonlinear. Nonlinear signal transfer eliminates or severely attenuates majority of rodʼs single photon responses. Rejection of noise more than compensates loss of signal - thus rod bipolars provide near-optimal readout of rod signals near visual threshold.
17 MODEL FOR ROD-ROD BIPOLAR SIGNAL TRANSFER probability rods nonlinearity gain Σ sum predict nonlinearity and discreteness of rod bipolar response flash strength # responses
18 signal and noise distributions and discrimination
19 keep discard Mouse rod-rod bipolar signal transfer is nonlinear. Nonlinear signal transfer eliminates or severely attenuates majority of rodʼs single photon responses. Rejection of noise more than compensates loss of signal - thus rod bipolars provide near-optimal readout of rod signals near visual threshold.
20 DISTRIBUTION OF ROD RESPONSES AT VISUAL THRESHOLD Rod experiments (~1 Rh*) Visual threshold (.1 Rh*) signal noise nonlinearity probability nonlinearity gain
21 ROD BIPOLAR PROVIDES NEAR OPTIMAL READOUT OF ROD SIGNALS AT VISUAL THRESHOLD (.1 Rh*/rod/integration time) Σ sum rods estimate signal-to-noise of output nonlinearity gain SNR/SNR linear probability all responses signal noise 1.2 observed nonlinearity nonlinearity midpoint 2
22 NONLINEAR SIGNAL TRANSFER LIMITS SENSITIVITY WELL ABOVE VISUAL THRESHOLD (.1 Rh*/rod/integration time) Σ sum rods estimate signal-to-noise of output nonlinearity gain probability SNR/SNR linear all responses signal noise nonlinearity midpoint 2 observed nonlinearity
23 olfactory bulb (CNS) glomerulus olfactory epithelium (receptors) Mombaerts et al, 1996
FREQUENCY OF SEEING EXPERIMENTS (Hecht, Shlaer and Pirenne, 1942) PROBLEM: No way to account for false positives (noise)
FREQUENCY OF SEEING EXPERIMENTS (Hecht, Shlaer and Pirenne, 1942) 100 target 80 fixation point target covers ~500 rods % seen 60 40 20 Θ Θ = 2 Θ = 7 = 12 0 10 100 photons at cornea CONCLUSION: Θ = 5-7
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