Photoreceptors II Photoreceptors II • Logarithmic photoreceptor, passive and active • Adaptive photoreceptor • Circuit techniques – Capacitive feedback, resistive “adaptive” elements, cascode, Miller effect, Speedup in transimpedance amplifiers, secondorder system analysis • Imaging arrays • Passive pixel sensor • Active pixel sensor (APS) • Charge-coupled device (CCD) Delbruck, CNS course 28.1.06 T. Delbruck, S.C. Liu Visual Pathway in Brain (Side View) Visual Pathway in Brain (Bottom View) Cross-Section through Human Retina Cross-Section through Human Eye Cones and Rods Outer nuclear layer Outer plexiform layer Horizontal cell Bipolar cell Inner nuclear layer Inner plexiform layer Ganglion cell 1 Two types of Photoreceptors: Rods & Cones Rods (~126 million) • • • • Spectral Sensitivity S R Sensitive in very dim light – lowest 3 decades Saturate at high illumination One light-sensitive pigment in blue-green Not present in fovea L M Cones (~6 million) • Only sensitive to brighter light – upper 9 decades • Three types with pigments with different wavelength sensitivities color vision 3 types of cones: L cones (sensitive to long wavelengths), M cones (sensitive to medium wavelengths), S cones (sensitive to short wavelengths) are only 10 % of the total number of cones, and absent from the center of the fovea. Rods (R) are sensitive to intermediate wavelengths. Rod/Cone Distribution X1000 Rodieck Photoreceptor Response • Outer segments are modified cilia with disks filled with opsin, the molecule that absorbs photons, as well as voltage-gated Na channels. • The protein opsin contains a pigment molecule called retinal. In rods, the opsin+retinal together are called rhodopsin. In cones, there are diferent types of opsins combined with retinal to form pigments called photopsins. Rodieck Photoreceptor Response • When a photon absorbed by a rod/cone visual pigment, the retinal undergoes a photoisomerization to all-trans retinal which changes conformation of opsin. • This conformation change triggers a biochemical cascade that causes hyperpolarization of the cell and a decrease of glutamate release. • Bipolar cells respond to the subsequent decrease in glutamate. Response to flash of light, (Baylor, 1987) 2 Biological phototransduction uses a chain of amplifiers Biological photoreceptors amplify changes more than DC Turtle cone, intracellular recording Mahowald, Mahowald,1992 1992 Normann & Perlman, 1979 Logarithmic Photosensors M1 V out How the source follower receptor works M1 M2 M1 Vb Vout Vout Iph Iph Iph E Diode-connected MOSFET E E D1 D1 D1 Double diode-connected MOSFET Source-follower MOSFET Characteristics of Logarithmic Photosensors Diode-connected MOSFET: Vout Double diode-connected MOSFET: Vout Source-follower MOSFET: Vout Contrast encoding: Logarithmic Photosensor with Feedback Loop Vdd 1 I ph Vdd U T log 2 I0 I ph V g U T log I0 dV out U T dI ph • Photodiode has large capacitance Ib I ph log I0 UT M3 Vb M1 Vout Vs M2 Iph E – Small photocurrent takes a long time to change VS • High-gain feedback loop clamps VS and speeds up response • Output voltage is read at the gate of M1 D1 I ph 3 Adaptive Photoreceptor Current-Voltage Characteristics of Logarithmic Photosensor with Feedback Loop • Logarithmic photosensor covers large range of illumination, but contrast-response is small (~100mV for a b/w transition (one decade of illumination)) • Imaging arrays suffer from fixed-pattern noise (FPN), i.e. different photosensors have different output signals for same illumination, due to spatial variation of fabrication parameters (different I0, 1.85 Iph Vout 1 VS UT log I0 A dIph UT dIph dVout UT A1 Iph Iph 1.8 1.7 1.65 1.6 dVS 1.55 0.01 0.1 1 10 100 – Low-FPN amplification of transient signals required to detect lowcontrast features – Use capacitive divider for amplification and resistive element for adaptation UT dIph UT dIph A1 Iph A Iph 1000 E(mW/m2) Amplification of Response by Speedup of response by A Adaptive Photoreceptor Adaptive photoreceptor from a general feedback perspective Ib C1 M3 R1 Vfb b(s) x(s) + C2 y(s) a(s) Vs M2 Iph + • Transient amplification of logarithmic signal Vfb onto Vout by capacitive gain stage (C1, C2) Vout • Slow adaptation of Vout to Vfb and change of adaptation state via current through resistive element R1 • State of adaptation stored as charge Qfb on Vfb node Vb M1 E D1 Comparing source-follower and adaptive photoreceptor Adaptive Photoreceptor Response Curves Square wave input over 7 decades illumination Adaptive receptor 200mV Vout (V) 1.75 Time 5s Decade changes in background illumination Delbruck and Mead 1993 Source follower receptor Constrast ratio 2:1 Brightest 3W/m2 4 Characteristics of Adaptive Photosensor Resistive Elements I ph Vfb 1 VS UT log I 0 C C2 Transient amplification: AC 1 C2 M2 Adapted signal: M3 Ifb Qfb C1Vdd Vb Iout Vfb Vout Vfb Vout ACVfb Small-signal response: A dIph U dIph dVout ACUT AC T A1 I ph I ph C2 Vfb Expansive element with low common-mode sensitivity Delbruck 1993 Vout Q1 M1 Transient signal: Vout M1 Expansive element with more symmetric characteristics Compressive element Delbruck and Mead 1993 Liu 1998 Expansive Resistive Element Vout > Vfb : diode-connected MOSFET D S Vfb Vout Ifb p n D Iout p+ p+ n+ S ISD B Vfb > Vout : 2-collector bipolar transistor E Vfb Vout Ifb p C2 n E C1 C2 Iout p+ IEC2 n+ p+ C1 IEC1 B IEB Delbruck and Mead 1993 Current-Voltage Characteristics of Expansive Resistive Element Current-Voltage Characteristics of Expansive Resistive Element -2 10 -4 10 -I (V >V ) fb fb out -I (V >V ) out fb out -6 I (A) 10 Ifb(Vout>Vfb) -8 10 -10 10 Iout(Vout>Vfb) -12 10 0 0.5 1 1.5 2 |V -V | (V) out fb Delbruck and Mead 1993 5 Step-Response of Adaptive Photosensor Adaptive Photosensor with Cascode E • Miller effect: Parasitic gateto-drain capacitance of M2 is driven by Vout node and thus effectively amplified by large gain of inverter Shield drain of M2 from large output voltage swing with cascode M4 to speed up response Ib 2.2 C1 2.1 M3 Vb R1 M1 Vfb Vout Vout (V) 2 C2 1.9 M4 Vc 1.8 Vs 1.7 M2 Iph 1.6 E D1 1.5 0 50 100 150 200 Tim e(s) Delbruck 1993 Small signal analysis of active log photoreceptor Measured effect of cascode on adaptive photoreceptor 6 Root locus plot of poles of H(s) of adaptive photoreceptor Delbruck and Mead 1993 Imaging arrays Integrating Photodiode Pixels • 1D or 2D array of photosensors can record optical images projected onto it by lens system • Individual photosensor in an imaging array is called pixel (picture element) • High image resolution requires small pixels • Outputs of a set of pixels have to be multiplexed onto a single signal line small read-out duty cycle • Two strategies for pixel operation: • Continuous-time mode: Conversion of photogenerated charge into steady-state photocurrent • Integration mode: Collection of photogenerated charge on capacitor until readout, followed by reset Vref Vr M1 Vrs M1 Vph Vph E E D1 D1 Photogates for Charge-Coupled Devices (CCDs) depletion p- substrate p- substrate E E Surface charge storage Surface-channel CCD Active pixel Acquisition period: charge packets collected under photogates Readout period: charge packets shifted underneath photogates by clocking gate potentials between two values n diffusion depletion Vro Charge-Packet Transport in CCDs Vg inversion M3 Vout Passive pixel Vg M2 Bulk charge storage Buried-channel CCD F3 F2 F1 F2 F1 t1 t2 t3 t4 t5 t6 t1 t2 t3 t4 3-phase clock 2-phase clock with stepped oxide 7 In biological photoreceptors, the rise time is not proportional to the gain CCD Registers imaging array vertical readout register transfer gate vertical readout register imaging array Time to peak response (s) output horizontal readout register output Gain (arbitrary units) horizontal readout register Frame-transfer CCD Interline-transfer CCD Rise time ~ Gain0.25 The gain is distributed Light a a Signal a log(gain) log(gain) log(bandwidth) log(Illumination) log(frequency) 8