Sensing Tactile Kinesthetic (position / force) B R A I L L E Manipulation HAPTICS
Slide courtesy of Prof. Roberta L. Klatzky, CMU, 2005
Sensing
Tactile Kinesthetic(position / force)
B R A I L L E
Manipulation
HAPTICS
Slide courtesy of Prof. Roberta L. Klatzky, CMU, 2005
Haptic Perception &Human Response to VibrationsHaptic Perception &Human Response to Vibrations
Outline:
1. Neural Coding of Touch Primitives
2. Functions of Peripheral Receptors
Slide courtesy of Prof. Roberta L. Klatzky, CMU, 2005
1. Neural Coding of Touch Primitives1. Neural Coding of Touch Primitives
• Touch Receptors– Mechanoreceptors and their Function
– Other skin receptors: thermal, pain
– Kinesthetic receptors
• Pathways from receptors to brain
Slide courtesy of Prof. Roberta L. Klatzky, CMU, 2005
Touch ReceptorsTouch Receptors
• Touch sensations are mediated by receptors that respond to pressure, vibration, and heat flow.
• The receptors are found in two regions:
-- Within skin: cutaneous sensing pressure, temperature, pain
-- Beneath skin in muscles, tendons, joints:
kinesthetic sensinglimb position and movement
Slide courtesy of Prof. Roberta L. Klatzky, CMU, 2005
Classes of ReceptorsClasses of Receptors
• Receptors that respond to pressure and vibration are called mechanoreceptors
• Mechanoreceptors are found in skin (cutaneous) and muscles, tendons, and joints (kinesthesis)
• The skin also includes other receptors – that signal skin warming and cooling (thermo-receptors)
– that signal pain (nociceptors)
Slide courtesy of Prof. Roberta L. Klatzky, CMU, 2005
Skin Mechanoreceptors have specialized endingsSkin Mechanoreceptors have specialized endings
Johannson & Valbo, 1983
Epidermis
Dermis
Subcutis
Meissner
Merkel
Ruffini
Pacinian
Slide courtesy of Prof. Roberta L. Klatzky, CMU, 2005
Functional characteristics of Skin Mechanoreceptors:Receptive field size (I = small, II = large)and adaptation rate (FA = fast adapting, SA = slow adapting)
Functional characteristics of Skin Mechanoreceptors:Receptive field size (I = small, II = large)and adaptation rate (FA = fast adapting, SA = slow adapting)
Kandel et. al., 2000
Meissner’s Merkel Pacinian RuffiniCorpuscle Cell Complex Corpuscle Ending
Receptors
Receptive
Field
Intensity and Time Course of Neural Signal (adaptation)
NeuralSpike train
Stimulus
FA I SA I FA II SA II
Slide courtesy of Prof. Roberta L. Klatzky, CMU, 2005
Distribution of Mechanoreceptors on the Palm: Receptors with small receptive fields are densely packed on fingertips.
Distribution of Mechanoreceptors on the Palm: Receptors with small receptive fields are densely packed on fingertips.
Result: ~.5 mm 2-point discrimination on fingertip
Slide courtesy of Prof. Roberta L. Klatzky, CMU, 2005
Frequency Sensitivity: One-channel-per-mechanoreceptor modelFrequency Sensitivity: One-channel-per-mechanoreceptor model
Amplitude required for threshold response at given frequency(dB relative to 1 μm)
Bolanowski et al., 1988
SA I SA II
FA I
FA II (PC)
Average data
Slide courtesy of Prof. Roberta L. Klatzky, CMU, 2005
Thermo-ReceptorsThermo-Receptors
Thermo-receptors lack specialized endings; respond to temperature change.
Two populations of nerve fibers: warm and cold
Kenshalo, 1976
NormalBody temp
Cold fibers
Warmfibers
Slide courtesy of Prof. Roberta L. Klatzky, CMU, 2005
The brain: Primary and Secondary Somatosensory CortexThe brain: Primary and Secondary Somatosensory Cortex
SI includes Brodmann areas:3a: Muscle3b: Skin (SA and FA)1: Skin (FA)2: Pressure, joints
SI projects to SII, 5, 7
SI
SII
Central Sulcus SI Areas 5 & 7
SIILateral Sulcus
Slide courtesy of Prof. Roberta L. Klatzky, CMU, 2005
Somatosensory “Homunculus” in SIresults from somatotopic mapping*Somatosensory “Homunculus” in SIresults from somatotopic mapping*
*Adjacent on skin⇒Adjacent in SI
There are severalsuch maps in SI and SII.
Penfield & Rasmussan, 1950
Slide courtesy of Prof. Roberta L. Klatzky, CMU, 2005
Somatosensory areas in brain are plastic: Re-assignment of receptive fields after amputation of a digit
Somatosensory areas in brain are plastic: Re-assignment of receptive fields after amputation of a digit
Areas in SI that once responded to 3rd fingertip are now activated by finger 2 and 4, plus base of 3
Merzenich et al., 1984
Amputation point
Slide courtesy of Prof. Roberta L. Klatzky, CMU, 2005
Cortical plasticity causes phantom limb painCortical plasticity causes phantom limb pain
ThumbIndex finger5th digit
Stimulating face and arm activates cortical areas previously responsive to fingers, now taken over by face and arm.
face finge
rs
arm
Slide courtesy of Prof. Roberta L. Klatzky, CMU, 2005
2. Functions of peripheral receptors2. Functions of peripheral receptors
• Slowly adapting cutaneous mechanoreceptors provide array (tactile) sensing.
• Fast adapting cutaneous receptors signal pressure changes on the skin.
• Kinesthetic mechanoreceptors provide a sense of limb position.
• Therefore, the mechanoreceptor populations support different human abilities.
Slide courtesy of Prof. Roberta L. Klatzky, CMU, 2005
Grasping is supported by FAI mechanoreceptors that detect incipient slipGrasping is supported by FAI mechanoreceptors that detect incipient slip
FAIs fire under near-slipGrasp, lift… Return
Load Force N
Grip Force N
Position mm
Grip: Load ratio
Time
Load Force N
Grip Force N
Position mm
Force Ratio
FA ISA IFA IISA II
Slip threshold
Johansson & Westling, 1984; Westling, 1986
Slide courtesy of Prof. Roberta L. Klatzky, CMU, 2005
Tactile Pattern Perception is based on SAI mechanoreceptorsTactile Pattern Perception is based on SAI mechanoreceptors
Input pattern
Response
Phillips, Johansson & Johnson, 1990
Spatial plot of the response of a skin mechanoreceptor to a Braille pattern swept through its receptive field: Each tick mark is a neural impulse given contact from that stimulus location.
PatternpreservationSA I
FA I
SA II
FA II
Slide courtesy of Prof. Roberta L. Klatzky, CMU, 2005
Curvature perception also reflects SA I responsesCurvature perception also reflects SA I responses
-4 -2 0 2 4Distance of receptive field from center of indentation (mm)
MeanSA IResponse
Data are shown for 7 curves, ranging from radius zero to radius 1.44 mm
Goodwin et al., 1995
Slide courtesy of Prof. Roberta L. Klatzky, CMU, 2005
PC receptors (FA II) signal causes of deep vibrationsPC receptors (FA II) signal causes of deep vibrations• Transient vibrations from contact•Micron-element textural variations, as in this stimulus
Only Pacinian Corpuscles respond continuously to the textured portion of a half-textured plate with ht. 1 μm
Srinivasan, Whitehouse, & LaMotte, 1990
Stimulus plate Receptor Responses Therefore, PCs underlie roughness percept at micro-scale.
Slide courtesy of Prof. Roberta L. Klatzky, CMU, 2005
Impact on functions of cutaneous receptors when skin is covered with a rigid sheath
Impact on functions of cutaneous receptors when skin is covered with a rigid sheath
Decrements in spatial resolution, differential force sensing, pattern perception --but vibration (sensed by deeper PCs) remains essentially intact
Lederman & Klatzky, 1999
This simulates some force-feedback environments.