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) 100 target 80 fixation point target covers ~500 rods % seen Θ Θ = 2 Θ = 7 = photons at cornea CONCLUSION: Θ = 5-7 photons absorbed spread over 500 rods PROBLEM: No way to account for false positives (noise)
3 Signal and noise distributions and false positives (on board)
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5 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 single photon alters optic nerve activity But the task of the retina is in fact more challenging...
6 CONVERGENCE AND SPARSE SIGNALING IN MAMMALIAN RETINA keep discard At visual threshold photons < 0.1% of the rods contribute signals while all rods generate noise Under these conditions averaging is a disaster - instead requires separation of signal from noise General problem in nervous system What is optimal readout of array of detectors when small fraction active? Baylor et al., 1984 van Rossum and Smith, 1998 Field and Rieke, 2002
7 CONVERGENCE AND SPARSE SIGNALING IN MAMMALIAN RETINA keep discard At visual threshold photons < 0.1% of the rods contribute signals while all rods generate noise Under these conditions averaging is a disaster - instead requires separation of signal from noise General problem in nervous system Sizeable behavioral consequences for getting this right! Baylor et al., 1984 van Rossum and Smith, 1998 Field and Rieke, 2002
8 RODS GENERATE PHOTON-LIKE NOISE EVENTS AND CONTINUOUS NOISE continuous noise photon-like noise event ave photon-like noise event ave single photon response 0.2 sec 1 pa 1 pa 0 20 time (sec) Baylor et al., 1984 Greg Field
9 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 rod s 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.
10 THE ROD BIPOLAR PATHWAY rod cone rod bipolar cell AII amacrine cell cone bipolar cell ganglion cell
11 DIM FLASH RESPONSES OF RODS GROW LINEARLY WITH FLASH STRENGTH 0 pa log(r/r max ) -1 rods sec log(rh*) rod bipolar why log-log? Field and Rieke, 2002
12 RESPONSES OF ROD BIPOLARS BUT NOT RODS GROW SUPRALINEARLY WITH FLASH STRENGTH 0 pa log(r/r max ) -1 rods sec log(rh*) rod bipolar pa log(r/r max ) Field and Rieke, 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 sec amplitude 4 rod bipolar Field and Rieke, 2002
15 ROD BIPOLARS GENERATE DISCRETE RESPONSES TO DIM FLASHES 2 pa # responses Rh* rods sec amplitude 4 rod bipolar 20 pa # responses Rh*/rod Field and Rieke, sec amplitude 4
16 keep discard Mouse rod-rod bipolar signal transfer is nonlinear. Nonlinear signal transfer eliminates or severely attenuates majority of rod s 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 amplitude Σ sum predict nonlinearity and discreteness of rod bipolar response amplitude flash strength # responses amplitude
18 Signal and noise distributions and discrimination (on board)
19 keep discard Mouse rod-rod bipolar signal transfer is nonlinear. Nonlinear signal transfer eliminates or severely attenuates majority of rod s 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 ( Rh*) signal noise nonlinearity probability nonlinearity gain amplitude amplitude Field and Rieke, 2002
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) 1 target 8 fixation point target covers ~5 rods % seen 6 4 2 Θ Θ = 2 Θ = 7 = 12 1 1 photons at cornea CONCLUSION: Θ = 5-7 photons absorbed
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