IMPLICATIONS OF BEHAVIORAL SENSITIVITY
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2 IMPLICATIONS OF BEHAVIORAL SENSITIVITY rod phototransduction - single photons reliably transduced rod! bipolar! AII! amacrine! cone cone! bipolar! ganglion! synaptic transmission - reliable transmission of single photon responses neural coding - absorption of a few photons produces change in optic nerve activity
3 IMPLICATIONS OF BEHAVIORAL SENSITIVITY rod! bipolar! rod cone cone! bipolar! phototransduction - single photons reliably transduced - reproducible responses to each absorbed photon How do you make an accurate single molecule timer? synaptic transmission - reliable transmission of single photon responses AII! amacrine! ganglion! neural coding - absorption of a few photons produces change in optic nerve activity
4 IMPLICATIONS OF BEHAVIORAL SENSITIVITY phototransduction - single photons reliably transduced - reproducible responses to each absorbed photon How do you make an accurate single molecule timer? synaptic transmission - reliable transmission of single photon responses - separation of sparse signal from noise What is optimal readout for array of noisy sensors when tiny fraction active? neural coding - absorption of a few photons produces change in optic nerve activity
5 REPRODUCIBILITY OF ROD RESPONSES TO SINGLE PHOTONS Rh* 2 pa 0 photocurrent flash monitor 0 10 sec 20 30
6 REPRODUCIBILITY OF ROD RESPONSES TO SINGLE PHOTONS 2 pa 0.4 Rh* 2 pa singles photocurrent flash monitor failures 0 10 sec CVarea = standard deviation / mean = 0.34 ± 0.01 (n=30) much less then other signals produced by single molecules! sec
7 EXPECTATION FOR FIRST-ORDER PROCESS if Rhodopsin behaves like typical single molecule Rh* activity Time CV =1
8 MULTIPLE SHUTOFF STEPS COULD DECREASE VARIABILITY Rieke and Baylor, 1998 Field and Rieke, 2002 if Rhodopsin behaves like typical single molecule proposed model for rhodopsin inactivation Rh* activity Rh* activity Time CV =1 Time CV =1/ N
9 MULTIPLE SHUTOFF STEPS COULD DECREASE VARIABILITY Rieke and Baylor, 1998 Field and Rieke, 2002 if Rhodopsin behaves like typical single molecule proposed model for rhodopsin inactivation Rh* activity Rh* activity Time CV =1 Time CV =1/ N phosphorylation via kinase quenching via arrestin R*... R*-P n R i -P n -arrestin Key Prediction: variability should change in graded and systematic manner as # phosphorylation sites decreased
10 FEWER PHOSPHORYLATION SITES = MORE VARIABILITY 0 sites 1 site 2 sites 3 sites 5 sites 6 sites (control) 2 pa 10 s 10 s 10 s 0.5 s 0.5 s 0.5 s increasing # phosphorylation sites decreasing response variability Thuy Doan Tony Azevedo
11 VARIABILITY HAS GRADED AND SYSTEMATIC DEPENDENCE ON # SITES CV area 0.6 n=15 n=16 n=17 predicted 1/!# sites n=20 n=27 n=29 n= Number of phosphorylation sites 2 pa 10 s 10 s 10 s 0.5 s 0.5 s 0.5 s Thuy Doan Tony Azevedo
12 VARIABILITY HAS GRADED AND SYSTEMATIC DEPENDENCE ON # SITES CV area 0.6 n=15 n=16 n=17 predicted 1/!# sites CV area 0.4 n=27 n=20 n=29 n= Number of phosphorylation sites # sites 6 2 pa 10 s 10 s 10 s 0.5 s 0.5 s 0.5 s Thuy Doan Tony Azevedo
13 IMPLICATIONS OF BEHAVIORAL SENSITIVITY phototransduction - single photons reliably transduced - reproducible responses to each absorbed photon How do you make an accurate single molecule timer? synaptic transmission - reliable transmission of single photon responses - separation of sparse signal from noise What is optimal readout for array of noisy sensors when tiny fraction active? neural coding - absorption of a few photons produces change in optic nerve activity
14 CONVERGENCE AND SPARSE SIGNALING IN MAMMALIAN RETINA At visual threshold photons < 0.1% of the rods contribute signals while all rods generate noise Under these conditions averaging is a disaster General problem in nervous system ~0.001 Rh*/rod Petri Ala-Laurila Greg Field A. P. Sampath 200 ms
15 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 General problem in nervous system ~0.001 Rh*/rod Petri Ala-Laurila Greg Field A. P. Sampath explaining ganglion sensitivity requires that circuit effectively separates signals from noise 200 ms
16 LOOKING FORWARD... How do you make an accurate single molecule timer? rod 2 pa singles rod! bipolar! AII! amacrine! cone cone! bipolar! ganglion! sec failures functional significance: permits photon counting, but... timing information? need tools to explore stimulus space What is optimal readout for array of noisy sensors when tiny fraction active?
17 LOOKING FORWARD... How do you make an accurate single molecule timer? rod What is optimal readout for array of noisy sensors when tiny fraction active? cone rod! bipolar! AII! amacrine! cone! bipolar! ganglion! general theoretical argument: multiple stages time-varying input dependence on sensor signal/noise properties
18 Current lab members Petri Ala-Laurila Juan Angueyra Tony Azevedo Jon Cafaro Will Grimes Michael Rudd Greg Schwartz Past lab members Thuy Doan (Stanford) Felice Dunn (UW) Greg Field (Salk) Cecilia Gold (Penn) Gabe Murphy (Janelia) Philipp Khuc-Trong (Max Planck) Kerry Kim (FHL) A.P. Sampath (USC) Fred Soo (Princeton) Barry Wark (Physion) Collaborators EJ Chichilnisky Valerie Uzzell Peter Detwiler Jeannie Chen Ana Mendez Felice Dunn
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