Active sensing. Ehud Ahissar
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1 Active sensing Ehud Ahissar 1
2 Active sensing Passive vs active sensing (touch) Comparison across senses Basic coding principles Perceptual loops Sensation-targeted motor control Proprioception Controlled variables Active vibrissal touch: encoding and recoding 2
3 Eye movements during fixation 3
4 sensory encoding: What receptors tell the brain Sensory organs consist of receptor arrays: somatosensation audition vision ~200 µm Finger pad 10 µm cochlea 10 µm retina Spatial organization => Spatial coding ( which receptors are activated ) Movements => Temporal coding ( when are receptors activated ) 4
5 Temporal coding in action 5
6 Coding space by time 1. Spatial frequency 2. Spatial phase 6
7 Touch: Temporal encoding of spatial features Darian-Smith & Oke, J Physiol, 1980 anesth. monkey, MR fibers 7
8 RA fiber Vel - constant f = SF * V dt = dx / V 8
9 SF Vel SA fiber 9
10 SF Vel RA fiber V1 V2 V3 G1 G2 G3 10
11 SF Vel PC fiber 11
12 Coding ranges 12
13 Temporal filtering (by intrinsic factors) eye K P M W X Y whisker Frequency (Hz) Frequency (Hz) finger SA RA PC Frequency (Hz)
14 Coding space by time 1. Spatial frequency 2. Spatial phase 14
15 Vision: Temporal encoding due to eye movement space Veye RF(1) RF(2) space retinal outputs 1 2 time 15
16 Vision: Temporal encoding due to eye movement x space Veye RF(1) RF(2) space retinal outputs 1 2 t time 16
17 Vision: Temporal encoding due to eye movement x space Veye RF(1) RF(2) space retinal outputs 1 2 t time 17
18 Vision: Temporal encoding due to eye movement x space Veye RF(1) RF(2) space retinal outputs 1 2 t time 18
19 Vision: Temporal encoding due to eye movement x space Veye RF(1) RF(2) space retinal outputs 1 2 t time 19
20 Spatial vs temporal coding Spatial Temporal faster better resolution scanning allows sensing in between receptors 20
21 Passive vs Active sensing of stationary objects Passive Active threshold low high accuracy low high Systems involved sensory Sensory + motor coding spatial Spatial + temporal Processing speed fast slow Used in detection Exploration Localization Identification 21
22 Central processing of touch where touch begins? Text book: at the receptors 22
23 Sensory-motor loops of the vibrissal system Primary Sensory Cortex Secondary Cortex Primary Motor Cortex Zona Incerta VPM-dm VPM-vl Thalamus POm - VL Thalamic Nuclei Sensory extralemniscal Cerebellar/Olivary Red Nucleus Pontine Reticular Nucleus Superior Colliculus Brainstem Reticular Formation Motor Trigeminal Nuclei Brainstem Loop + Facial Nucleus Trigeminal Ganglion Vibrissae 23
24 Sensory-motor loops of the vibrissal system Cortex E D C Identification ( what ) Localization ( where ) B Whisking A m a The old view Thalamus WT T W Brainstem WT T W 24
25 Sensory-motor loops of the vibrissal system Primary Sensory Cortex Secondary Cortex Primary Motor Cortex Zona Incerta VPM-dm VPM-vl Thalamus POm - VL Thalamic Nuclei Sensory extralemniscal Cerebellar/Olivary Red Nucleus Pontine Reticular Nucleus Superior Colliculus Brainstem Reticular Formation Motor Trigeminal Nuclei Brainstem Loop + Facial Nucleus Trigeminal Ganglion Vibrissae 25
26 Cortex E D C Identification ( what ) Localization ( where ) B Whisking A m a Thalamus WT T W Brainstem WT T W 26
27 Sensory-motor loops of the vibrissal system E D C B A m a WT T W WT T W 27
28 Central processing of touch where touch begins? Text book: at the receptors Active touch does not begin at the receptors Sensor motion determines the interaction between the receptors and external objects 28
29 Break?
30 Motor control Closed loops Proprioceptive feedback Reflexes tool for probing loop function Controlled variables motor vs sensory 30
31 Motor control Closed loops Proprioceptive feedback Reflexes tool for probing loop function Controlled variables motor vs sensory 31
32 Excitation Contraction Coupling Phase 1: Firing of Motor Neuron Phase 2: Release of Neurotransmitter 32
33 Excitation Contraction Coupling Phase 1: Firing of Motor Neuron Phase 2: Release of Neurotransmitter Phase 3: Muscle contraction 33
34 Open-loop system Information flows in one direction (from neurons to muscles 34
35 Open-loop system Information flows in one direction (from neurons to muscles Closed-loop system Information flows in a closed loop: from neurons to muscles and from muscles to neurons What kind of information? 35
36 Closed-loop system The direct feedback from muscles and joints is mediated by proprioceptive signals Proprioceptive receptor types Name: Muscle spindle receptors Golgi tendon organs Joint receptors Sensitive to: muscle length muscle tension Flexion, extension 36
37 Proprioceptive receptor types Name: Muscle spindle receptors Golgi tendon organs Joint receptors Sensitive to: muscle length muscle tension Flexion, extension Location: Fleshy part of the muscle Between muscle and tendon Joint capsule Parallel to muscle fibers Serial to muscle fibers 37 Between bones
38 Motor control Closed loops Proprioceptive feedback Reflexes tool for probing loop function Controlled variables motor vs sensory 38
39 What proprioceptors encode? 39
40 Proprioceptive receptor types Name: Muscle spindle receptors Golgi tendon organs Joint receptors Sensitive to: muscle length muscle tension Flexion, extension From Arthur Prochazka, University of Alberta 40
41 Proprioceptive receptor types Name: Muscle spindle receptors Golgi tendon organs Joint receptors Sensitive to: muscle length muscle tension Flexion, extension Encode: force f = k 1 F 41
42 Proprioceptive receptor types Name: Muscle spindle receptors Golgi tendon organs Joint receptors Sensitive to: muscle length muscle tension Flexion, extension Encode: Length + velocity f = k 1 L + k 2 V 0.6 force f = k 1 F angle f = k 1 θ 42
43 Proprioceptive receptor types Name: Muscle spindle receptors Golgi tendon organs Joint receptors Sensitive to: muscle length muscle tension Flexion, extension Encode: Length + velocity f = k 1 L + k 2 V 0.6 force f = k 1 F angle f = k 1 θ θ θ θ 43
44 PID control Proportional (to the controlled variable) Integral (of the controlled variable) Derivative (of the controlled variable) Present Past Future θ θ θ 44
45 Negative feedback loop Characteristic: The effect of a perturbation is in the opposite direction Requirement: The cumulative sign along the loop is negative Function: Can keep stable fixed points 45
46 Positive feedback loop Characteristic: The effect of a perturbation is in the same direction Requirement: The cumulative sign along the loop is positive Function: amplifies perturbations + 46
47 Motor control Closed loops Proprioceptive feedback Reflexes tool for probing loop function Controlled variables motor vs sensory 47
48 The stretch reflex probes the control function of muscle spindles 48
49 Is the loop positive or negative? The stroke stretches the muscle As a result the muscle contracts The result opposes the perturbation => negative FB loop 49
50 the anatomical loop Muscle spindle excites the motor neuron Motor neuron excites muscle fibers Muscle contraction suppresses spindle response 50
51 Proprioceptive receptor types Name: Muscle spindle receptors Golgi tendon organs Joint receptors Sensitive to: muscle length muscle tension Flexion, extension Encode: force f = k 1 F Why proprioceptors fire at rest? And why aren t we aware of it? 51
52 What about the flexor muscles? Positive or negative loop? What is the underlying circuit? Take it as homework may appear in the exam 52
53 Pain reflex Positive or negative? What is the underlying 53 circuit? Same
54 Motor control Closed loops Proprioceptive feedback Reflexes tool for probing loop function Controlled variables motor vs sensory 54
55 Break?
56 Sensory-motor loops of the vibrissal system Primary Sensory Cortex Secondary Cortex Primary Motor Cortex Zona Incerta VPM-dm VPM-vl Thalamus POm - VL Thalamic Nuclei Sensory extralemniscal Cerebellar/Olivary Red Nucleus Pontine Reticular Nucleus Superior Colliculus Brainstem Reticular Formation Motor Trigeminal Nuclei Brainstem Loop + Facial Nucleus Trigeminal Ganglion Vibrissae 56
57 Basic principles of closed-loop control 57
58 Set point - + f Vd Vs Vm=f(-Vs) Vs=g(Vm) Vs Vm V s0 V m0 Vm V 58
59 Set point - + f Vd Vs Vm=f(Vd-Vs) Vs=g(Vm) Vs Vm V s0 Vsd V md V m0 Vm V 59
60 Direct control without direct connection - + f Vd Vs Vm=f(Vd-Vs) Vs=g(Vm) Vs Vm V s0 Vsd V md V m0 Vm V 60
61 Nested loops + - f f Vd Vs Vm=f(Vd-Vs) Vs=g(Vm) Vs Vm V s0 Vsd V md V m0 Vm V 61
62 Parallel loops + - f 2 Vs Vm2=f(Vd-Vs)Vs=g(Vm2) - + f Vm2 V s0 Vsd V md V m0 Vm2 Vs Vm1 Vs V s0 Vsd Vm1=f(Vd-Vs)Vs=g(Vm1) V V md V m0 Vm1 62
63 Parallel loops + - f 2 Xs Xm=f(Xd-Xs) Xs=g(Xm) - + f Xm X s0 Xsd X md X m0 Xm Xs Vs Vm Vs Vm=f(Vd-Vs) Vs=g(Vm) V s0 Vsd V V md V m0 Vm 63
64 Closed loops in active sensing The controlled variables can be f 2 + f Motor (Xm) (velocity, amplitude, duration, direction, ) Sensory (Xs) (Intensity, phase, ) Object (via Xm Xs relationships) (location, SF, identity, ) Xs Vs Vm Xm V 64
65 Sensory-motor loops of the vibrissal system Primary Sensory Cortex Secondary Cortex Primary Motor Cortex Zona Incerta VPM-dm VPM-vl Thalamus POm - VL Thalamic Nuclei Sensory extralemniscal Cerebellar/Olivary Red Nucleus Pontine Reticular Nucleus Superior Colliculus Brainstem Reticular Formation Motor Trigeminal Nuclei Brainstem Loop + Facial Nucleus Trigeminal Ganglion Vibrissae 66
66 Sensory-motor loops of the vibrissal system Primary Sensory Cortex Secondary Cortex Primary Motor Cortex Zona Incerta VPM-dm VPM-vl Thalamus POm - VL Thalamic Nuclei Sensory extralemniscal Cerebellar/Olivary Red Nucleus Pontine Reticular Nucleus Superior Colliculus Brainstem Reticular Formation Motor Trigeminal Nuclei Brainstem Loop + Facial Nucleus Trigeminal Ganglion Vibrissae 67
67 Sensory-motor loops of the vibrissal system Primary Sensory Cortex Secondary Cortex Primary Motor Cortex Zona Incerta VPM-dm VPM-vl Thalamus POm - VL Thalamic Nuclei Sensory extralemniscal Cerebellar/Olivary Red Nucleus Pontine Reticular Nucleus Superior Colliculus Brainstem Reticular Formation Motor Trigeminal Nuclei Brainstem Loop + Facial Nucleus Trigeminal Ganglion Vibrissae 68
68 Sensory-motor loops of the vibrissal system Primary Sensory Cortex Secondary Cortex Primary Motor Cortex Zona Incerta VPM-dm VPM-vl Thalamus POm - VL Thalamic Nuclei Sensory extralemniscal Cerebellar/Olivary Red Nucleus Pontine Reticular Nucleus Superior Colliculus Brainstem Reticular Formation Motor Trigeminal Nuclei Brainstem Loop + Facial Nucleus Trigeminal Ganglion Vibrissae 69
69 Active sensing in the vibrissal system 70
70 Sensory signal conduction The vibrissal system 71
71 whisker Sensory signal conduction The vibrissal system Meisner Merkel Ruffini Lanceolate free endings 72
72 73
73 Sensory-motor loops of the vibrissal system Primary Sensory Cortex Secondary Cortex Primary Motor Cortex Zona Incerta VPM-dm VPM-vl Thalamus POm - VL Thalamic Nuclei Sensory extralemniscal Cerebellar/Olivary Red Nucleus Pontine Reticular Nucleus Superior Colliculus Brainstem Reticular Formation Motor Trigeminal Nuclei Brainstem Loop + Facial Nucleus Trigeminal Ganglion Vibrissae 74
74 Motor control of whiskers Intrinsic muscles 75 Dorfl J, 1982, J Anat 135:
75 Follicle as a motor-sensory junction Motor signals move the follicle and whisker Follicle receptors report back details of self motion = proprioception Plus perturbations of this motion caused by the external world Dorfl J, 1985, J Anat 142:
76 Motor control of whiskers Intrinsic muscles 78 Dorfl J, 1982, J Anat 135:
77 Vibrissal proprioception Each follicle contains ~2000 receptors About 20% of them convey pure proprioceptive information 79
78 Vibrissal system Skeletal system Proprioceptive loop Proprioceptive loop 80
79 Whiskers come with different muscle sizes Intrinsic muscles 0.5 mm 81 Dorfl J, 1982, J Anat 135:
80 Whisking behavior reflections of control loops 82
81 Perception of external objects Object localization What signals must the brain process in order to infer a location of an external object in space? Reafferent + exafferent signals 86
82 What the whiskers tell the rat brain Reafference: Their own movement ( Whisking ) Exafference: Touch 88
83 What the whiskers tell the rat brain Whisking space Whisker position vs. time time 89
84 What the whiskers tell the rat brain Whisking space Whisker position vs. time time 90
85 What the whiskers tell the rat brain Whisking space Whisker position vs. time time 91
86 What the whiskers tell the rat brain Whisking space Whisker position vs. time time 92
87 What the whiskers tell the rat brain Whisking space Whisker position vs. time time 93
88 What the whiskers tell the rat brain Whisking space Whisker position vs. time time 94
89 What the whiskers tell the rat brain Whisking space Whisker position vs. time time 95
90 What the whiskers tell the rat brain Whisking space Whisker position vs. time time 96
91 What the whiskers tell the rat brain Whisking space Whisker position vs. time time 97
92 What the whiskers tell the rat brain Whisking space Whisker position vs. time time 98
93 What the whiskers tell the rat brain Whisking space Whisker position vs. time time 99
94 What the whiskers tell the rat brain Reafference: Their own movement ( Whisking ) Exafference: Touch 100
95 What the whiskers tell the rat brain Touch space Whisker position vs. time time 101
96 What the whiskers tell the rat brain Touch space Whisker position vs. time time 102
97 What the whiskers tell the rat brain Touch space Whisker position vs. time time 103
98 What the whiskers tell the rat brain Touch space Whisker position vs. time time 104
99 What the whiskers tell the rat brain Touch space Whisker position vs. time time 105
100 What the whiskers tell the rat brain Touch space Whisker position vs. time time 106
101 What the whiskers tell the rat brain Touch space Whisker position vs. time time 107
102 What the whiskers tell the rat brain Touch space Whisker position vs. time time 108
103 What the whiskers tell the rat brain Touch space Whisker position vs. time time 109
104 What the whiskers tell the rat brain Touch space Whisker position vs. time time 110
105 What the whiskers tell the rat brain Touch space Whisker position vs. time time 111
106 Whisking: What the whiskers tell the rat brain How can the brain use this information? space Whisker position vs. time time Touch: contact with object space Whisker position vs. time time 112
107 Whisking: What the whiskers tell the rat brain How can the brain use this information? space? Whisker position vs. time time Touch: contact with object space? Whisker position vs. time time 113
108 How can the brain extract the location of the object Whisking: space Whisker position vs. time time Touch: contact with object 114
109 How can the brain extract the location of the object Whisking: space Whisker position vs. time time Touch: contact with object 115
110 sensory encoding: What receptors tell the brain Sensory organs consist of receptor arrays: somatosensation audition vision ~200 µm Finger pad 10 µm cochlea 10 µm retina Spatial organization => Spatial coding ( which receptors are activated ) Movements => Temporal coding ( when are receptors activated ) 116
111 Orthogonal coding of object location Vertical object position is encoded by space Horizontal object position is encoded by time Radial object position is encoded by rate 117
112 Active sensing The End 118
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