Cutaneous & Chemical Senses Flashcards
What’s an auditory space?
- Perception of where sounds are located in space
- Auditory space extends around a listener’s head in all directions, existing wherever there is a sound
What’s auditory localization?
The perception of the location of a sound source in auditory space
Why is it easier to locate visual stimuli in the environment over auditory stimuli?
- Ex: consider a tweeting bird and a meowing cat near a tree
- Visual information for the relative locations of the bird and the cat is contained in the images of the bird and the cat on the surface of the retina
- The bird’s “tweet, tweet” and the cat’s “meow” on the other hand stimulate the cochlea based on their sound frequencies, and these frequencies cause patterns of nerve firing that result in our perception of a tone’s pitch and timbre
- Activation of nerve fibers in the cochlea is based on the tones’ frequency components and not on where the tones are coming from
- Meaning that 2 tones with the same frequency that originate in different locations will activate the same hair cells and nerve fibers in the cochlea
- The auditory system must therefore use information other than the place on the cochlea to determine location -> location cues
What are location cues?
- In hearing, characteristics of the sound reaching the listener that provide information regarding the location of a sound source
- These are created by the way sound interacts with the listener’s head and ears
What are the 2 types of location cues?
- Binaural cues
- Spectral cues
What do binaural cues locations cues depend on?
They depend on both ears
What do spectral cues location cues depend on?
They depend on both ears
Researchers studying the different sound location cues have determined how well people can utilize these cues to locate the position of a sound in what 3 dimensions?
- Azimuth
- Elevation
- Distance
What’s the azimuth dimension?
In hearing, specifies locations that vary from left to right relative to the listener
What’s the elevation dimension?
In hearing, sound locations that are up and down relative to the listener
What’s the distance dimension?
- How far a stimulus is from the observer
- In hearing, the distance coordinate specifies how far the sound source is from the listener
- Localization in distance is much less accurate than azimuth or elevation localization, working best when the sound source is familiar, or when cues are available from room reflections
What are binaural cues?
- Sound localization cue that involves both ears
- They use information reaching both ears to determine the azimuth (left–right position) of sounds
- Sounds that are off to the side are more intense at one ear than the other and reach one ear before the other
What are the primary binaural cues?
- Interaural time difference
- Interaural level difference
- Both are based on a comparison of the sound signals reaching the left and right ears
What’s the interaural level difference (ILD)?
- It is based on the difference in the sound pressure level (or just “level”) of the sound reaching the 2 ears (left & right)
- A difference in level between the 2 ears occurs because the head is a barrier that creates an acoustic shadow, reducing the intensity of sounds that reach the far ear
- This reduction of intensity at the far ear occurs for high-frequency sounds (greater than about 3,000Hz for humans), but not for low-frequency sounds
- The ILD provides a cue for sound localization for high-frequency sounds
Stimulating receptors on the tip of the finger, and other outer body areas, creates perceptions of what?
- Touch
- Pressure
- Temperature
People with a rare condition that results in losing the ability to feel sensations through the skin often suffer with what?
Constant bruises, burns, and broken bones in the absence of the warnings provided by touch and pain
What could losing the sense of touch result in?
- Increase the chance of injury
- Makes it difficult to interact with the environment because of the loss of feedback from the skin that accompanies many actions
What have experiments in which participants have had their hands temporarily anesthetized shown?
They’ve shown that the resulting loss of feeling causes them to apply much more force than necessary when carrying out tasks with their fingers and hands
Describe the case of Ian Waterman
- One of the most extreme examples of the effect of losing the ability to sense with the skin
- As a result of an autoimmune reaction that destroyed most of the neurons that transmitted signals from his skin, joints, tendons, and muscles to his brain, he lost the ability to feel skin sensations so he couldn’t feel his body when lying in bed, and he often used inappropriate force when grasping objects—sometimes gripping too tightly, and sometimes dropping objects because he hadn’t gripped tightly enough
- Destruction of the nerves from his muscles, tendons, and joints eliminated Ian’s ability to sense the position of his arms, legs, and body, so the only way he could carry out movements was by visually monitoring the positions of his limbs and body
- Ian’s problems were caused by a breakdown of his somatosensory system
What’s the somatosensory system?
The system that includes the cutaneous senses (senses involving the skin), proprioception (the sense of position of the limbs), kinesthesis (sense of movement of the limbs), haptic perception, and the vestibular system
What are the cutaneous senses?
The ability to perceive sensations, such as touch and pain, that are based on the stimulation of receptors in the skin
What’s proprioception?
The ability to sense the position of the body and limbs
What’s kinesthesis?
The ability to sense the movement of the body and limbs
What are the cutaneous senses important for?
- They’re important not only for activities like grasping objects and protecting against damage to the skin, but also for motivating sexual activity
- They also create good feelings -> social touch
- They’re important both for day-to-day functioning and survival
List the various feelings involving the cutaneous senses in the skin
- Touch
- Pain
- Pressure
- Vibration
- Tickle
- Temperature
- Pleasure
What does the sense of touch enable us to perceive?
It enables us to perceive properties of surfaces and objects such as details, vibrations, texture, and shape
What did M. Comèl (1953) call the skin?
The “monumental facade of the human body”
What’s the heaviest organ in the human body?
- The skin
- It’s also the most obvious, especially in humans, whose skin isn’t obscured by fur or large amounts of hair
What are the functions of skin?
- Warning function
- Prevents body fluids from escaping and at the same time protects us by keeping bacteria, chemical agents, and dirt from penetrating our bodies
- Skin maintains the integrity of what’s inside and protects us from what’s outside, but it also provides us with information about the various stimuli that contact it
- The sun’s rays heat our skin, and we feel warmth; a pinprick is painful; and when someone touches us, we experience pressure or other sensations
What’s the epidermis?
The outer layers of the skin, including a layer of tough dead skin cells
What’s the dermis?
The layer of skin below the epidermis
What are the different layers of the skin?
- Epidermis
- Dermis
What are mechanoreceptors?
- Receptors that responds to mechanical stimulation of the skin, such as pressure, stretching, or vibration
- Located in the epidermis and the dermis
- The Merkel receptor (SA1) and Meissner corpuscle (RA1) are mechanoreceptors close to the surface of the skin, near the epidermis
What’s glabrous skin?
Skin without hairs or projections
Describe the Merkel receptor (SA1)
- A disk-shaped receptor in the skin associated with slowly adapting fibers
- Mechanoreceptor near the surface of the skin
- Fires to continuous pressure
- Perceives fine details and texture
- Perceives shapes
- Because it’s located close to the surface, it has small receptive fields
- Because the nerve fiber associated with the slowly adapting Merkel receptor fires continuously, as long as the stimulus is on, it is called a slowly adapting (SA1) fiber
Describe the Meissner corpuscle (RA1)
- A receptor in the skin, associated with RA1 mechanoreceptors
- It has been proposed that the Meissner corpuscle is important for perceiving tactile slip and for controlling the force needed to grip objects
- Mechanoreceptor near the surface of the skin
- Fires to “on” and “off”
- Perceives handgrip control
- Perceives motion across skin
- Because it’s located close to the surface, it has small receptive fields
- Because the nerve fiber associated with the rapidly adapting Meissner corpuscle fires only when the stimulus is first applied and when it is removed, it is called a rapidly adapting (RA1) fiber
What’s a cutaneous receptive field?
Area of skin that, when stimulated, influences the firing of a neuron
The rapidly adapting (RA1) fiber is associated with which skin receptor?
Meissner corpuscle
The slowly adapting (SA1) fiber is associated with which skin receptor?
Merkel receptor
What are the types of perception associated with the Merkel receptor/SA1 fiber?
- Details
- Shape
- Texture
What are the types of perception associated with Meissner corpuscle/RA1 fiber?
- Controlling handgrip
- Perceiving motion across the skin
What’s the Ruffini cylinder?
- A receptor structure in the skin associated with slowly adapting (SA2) fibers, which responds continuously to stimulation (fires to continuous pressure)
- It has been proposed that the Ruffini cylinder is involved in perceiving “stretching”
What’s the Pacinian corpuscule?
- A receptor with a distinctive elliptical shape associated with RA2 mechanoreceptors
- It’s a rapidly adapting fiber (RA2) which responds when the stimulus is applied or removed (fires to “on” and “off”)
- It transmits pressure to the nerve fiber inside it only at the beginning or end of a pressure stimulus and is responsible for our perception of vibration and fine textures when moving the fingers over a surface
Both the Ruffini cylinder and Pacinian corpuscle are located where?
Deep in the skin, so they have larger receptive fields
The Ruffini cylinder is associated with what kind of perception?
Perceiving stretching of the skin
The Pacinian corpuscle is associated with what kind of perception?
With sensing rapid vibrations and fine texture
The perception of texture often involves what?
The coordinated activity of different types of neurons working together
Where are cutaneous receptors in the skin located?
They’re distributed over the whole body
Why do we sometimes refer to the detection of stimulation of the skin as the “journey of the long-distance nerve impulses”?
- Because of the wide distribution of the cutaneous receptors in the skin, plus the fact that signals must reach the brain before stimulation of the skin can be perceived
- This reference is especially true for signals that must travel from the fingertips or toes to the brain
Signals from all over the body are conducted from the skin to the what?
To the spinal cord
How many segments does the spinal cord have?
Consists of 31 segments, each of which receives signals through a bundle of fibers called the dorsal root
What happens after signals from the stimulation of the cutaneous receptors enter the spinal cord?
Nerve fibers transmit them to the brain along 2 major pathways: the medial lemniscal pathway and the spinothalamic pathway
What are the 2 major pathways from the spinal cord to the brain?
- The medial lemniscal pathway
- The spinothalamic pathway
- Fibers from both pathways cross over to the other side of the body during their upward journey and synapse in the thalamus and then send signals to the somatosensory cortex in the parietal lobe
- Most of the fibers in the cutaneous system synapse in the ventrolateral nucleus of the thalamus
- Signals originating from the left side of the body reach the thalamus in the right hemisphere of the brain, and signals from the right side of the body reach the left hemisphere
- The case of Ian Waterman illustrates the separation in function between these 2 pathways, because although he lost the ability to feel touch and to sense the positions of his limbs (lemniscal pathway), he was still able to sense pain and temperature (spinothalamic pathway)
What’s the medial lemniscal pathway?
- A pathway in the spinal cord that transmits signals from the skin toward the thalamus
- It has large fibers that carry signals related to sensing the positions of the limbs (proprioception) and perceiving touch
- These large fibers transmit signals at high speed, which is important for perceiving and for reacting to touch
What’s the spinothalamic tract/pathway?
- One of the nerve pathways in the spinal cord that conducts nerve impulses from the skin to the somatosensory area of the thalamus
- The spinothalamic pathway consists of smaller fibers that transmit signals related to temperature and pain
What supports the idea that different pathways serve different sensations?
The idea of 2 pathways conducting cutaneous signals to the thalamus and then to the somatosensory cortex
Name some of the brain structures associated with the cutaneous system
- Primary somatosensory cortex (S1)
- Secondary somatosensory cortex (S2)
- Anterior cingulate cortex (ACC)
- The insula
What are 2 of the areas that receive signals from the thalamus?
- The primary somatosensory cortex (S1) in the parietal lobe
- The secondary somatosensory cortex (S2)
- Signals also travel between S1 and S2 and to a network of other areas in the brain in additional pathways
What’s the secondary somatosensory cortex (S2)?
The area in the parietal lobe next to the primary somatosensory area (S1) that processes neural signals related to touch, temperature, and pain
What is the insula important for?
Sensing light touch
What is the anterior cingulate cortex (ACC) involved in?
Pain
How is the somatosensory cortex organized and how was this discovered?
- It’s organized into maps that correspond to locations on the body
- The story behind the discovery of these maps -> when British neurologist Hughlings Jackson observed that in some cases of epilepsy, his patients’ seizures progressed over the body in an orderly way, with a seizure in one body part being followed by a seizure in a neighboring body part
- This sequence, known as “the Jacksonian march,” suggested that the seizures reflected the spread of neural activity across maps in the motor area of the brain
Describe Wilder Penfield and Edwin Boldrey (1937) study of the somatosensory map
- They measured the map of the somatosensory cortex by stimulating points on the brain of awake patients who were having brain surgery to relieve symptoms of epilepsy
- When Penfield stimulated points on the primary somatosensory cortex (S1) and asked patients to report what they perceived, they reported sensations such as tingling and touch on various parts of their body
- Penfield found that stimulating the ventral part of S1 (lower on the parietal lobe) caused sensations on the lips and face, stimulating higher on S1 caused sensations in the hands and fingers, and stimulating the dorsal S1 caused sensations in the legs and feet
- The resulting body map is called the homunculus
What’s the homunculus?
- Latin for “little man”
- The homunculus shows that adjacent areas of the skin project to adjacent areas in the brain, and that some areas on the skin are represented by a disproportionately large area of the brain
- Ex: the area devoted to the thumb is as large as the area devoted to the entire forearm
- This result is analogous to the cortical magnification factor in vision
- Similarly, parts of the body such as the fingers, with the highest tactile acuity are represented by larger areas on the cortex
- A similar body map also occurs in the secondary somatosensory cortex (S2)
Where is the somatosensory cortex located?
In the parietal lobe
Where does the primary somatosensory area (S1) receive inputs from?
From the ventrolateral nucleus of the thalamus
Where is the secondary somatosensory area (S2) located?
It’s partially hidden behind the temporal lobe
Recent research has shown that the primary somatosensory area (S1) is divided into how many interconnected areas?
Recent research has shown that S1 is divided into 4 interconnected areas, each with its own body map and its own functions
The area in S1 involved in perceiving touch is connected to what other area?
It’s connected to another area that is involved in haptics (exploring objects with the hand)
TRUE OR FALSE: The cutaneous system involves numerous areas of the brain which communicate with each other over many pathways
TRUE
What’s one of the most impressive examples of perceiving details with the skin?
- Braille
- The system of raised dots that enables blind people to read with their fingertips
- A Braille character consists of a cell made up of 1-6 dots
- The Braille alphabet consists of raised dots in a 2x3 matrix
- Different arrangements of dots and blank spaces represent letters of the alphabet
- Additional characters represent numbers, punctuation marks, and common speech sounds and words
- Experienced Braille readers can read at a rate of about 100 words/min, slower than the rate for visual reading, which averages about 250-300 words/min
- A Braille reader transforms an array of raised dots into information that goes far beyond simply feeling sensations on the skin
- The ability of Braille readers to identify patterns of small raised dots based on the sense of touch depends on tactile detail perception
What’s the first step in describing research on tactile detail perception?
To consider how researchers have measured tactile acuity
What’s tactile acuity?
- The capacity to detect details of stimuli presented to the skin
- The smallest details that can be detected on the skin
Describe the methods for measuring tactile acuity
- Just as there are a number of different kinds of eye charts for determining a person’s visual acuity, there are a number of ways to measure a person’s tactile acuity
- The classic method of measuring tactile acuity is the two-point threshold
- The two-point threshold is measured by gently touching the skin with two points, such as the points of a drawing compass, and having the person indicate whether he or she feels one point or two
- Another more recent method is the grating acuity method
- Grating acuity is measured by pressing a grooved stimulus onto the skin and asking the person to indicate the orientation of the grating
- Acuity is measured by determining the narrowest spacing for which orientation can be accurately judged
- Finally, acuity can also be measured by pushing raised patterns such as letters onto the skin and determining the smallest sized pattern or letter that can be identified
What’s the two-point threshold?
- A measure of acuity on the skin
- The smallest/minimum separation between 2 points on the skin that when stimulated is perceived as 2 points
- The two-point threshold was the main measure of acuity in most of the early research on touch
What does the firing of the Merkel receptor’s fiber signal?
- It signals details, and more specifically, details of patterns that are pushed onto the skin
- The firing of the fiber reflects the pattern of the stimuli pushed onto the skin
- Demonstrates tactile acuity
What’s one of the things that determine what we experience when the skin is stimulated?
The properties of the receptors
There’s a high density of what kind of receptors in the fingertips?
- Merkel receptors
- Explains why the fingertips are the parts of the body that are most sensitive to details
How has the relationship between locations on the body and sensitivity to detail been studied?
It has been studied psychophysically by measuring the two-point threshold on different parts of the body
What kind of receptors correspond with tactile acuity?
Merkel receptors
A comparison of grating acuity on different parts of the hand shows that better acuity is associated with what?
It’s associated with less spacing between Merkel receptors
Describe the correlation between density of Merkel receptors and tactile acuity
- As the SA1 receptor spacing (mm) increases, so does the distance needed for tactile acuity (mm)
- Fingertips have tactile acuity for the smallest amount of distance and have the smallest receptor spacing
- Then the base of the finger
- Finally, the hand has bigger SA1 receptor spacing with a larger distance needed for tactile acuity
How are regions of high acuity, like the fingers and lips, represented in the cortex?
- They’re represented by larger areas on the cortex
- Ex: the homunculus -> “magnification” of the representation on the brain of parts of the body such as the fingertips
- The map of the body on the brain is enlarged to provide the extra neural processing that enables us to accurately sense fine details with our fingers and other parts of the body
- Also, cortical neurons representing parts of the body with better acuity, such as the fingers, have smaller receptive fields
- Meaning that 2 points that are close together on the fingers might fall on receptive fields that don’t overlap and so would cause neurons that are separated in the cortex to fire
- 2 points with the same separation when applied to the arm are likely to fall on receptive fields that overlap and so could cause neurons that are not separated in the cortex to fire
- The small receptive fields of neurons receiving signals from the fingers translates into more separation on the cortex, which enhances the ability to feel 2 close-together points on the skin as 2 separate points
What are the Two-Point Thresholds (in mm) on Different Parts of the Male Body?
- Fingers (4 mm)
- Upper lip (8 mm)
- Big toe (9 mm)
- Back (42 mm)
- Thigh (44 mm)
- Upper arm (46 mm)
What’s the mechanoreceptor that’s primarily responsible for sensing vibration?
- The Pacinian corpuscle
- One piece of evidence linking the Pacinian corpuscle to vibration is that recording from fibers associated with the corpuscle shows that these fibers respond poorly to slow or constant pushing but respond well to high rates of vibration
Why do the Pacinian corpuscle fibers respond well to rapid vibration?
- The presence of the corpuscle surrounding the nerve fiber determines which pressure stimuli actually reach the fiber
- The corpuscle, which consists of a series of layers, like an onion, with fluid between each layer, transmits rapidly repeated pressure, like vibration, to the nerve fiber, but doesn’t transmit continuous pressure
- Thus, the corpuscle causes the fiber to receive rapid changes in pressure, but not to receive continuous pressure
Because the Pacinian corpuscle doesn’t transmit continuous pressure to the fiber, what should presenting continuous pressure to the corpuscle cause?
It should cause no response in the fiber
Describe Werner Lowenstein (1960) study on the Pacinian corpuscle
- He showed that when pressure was applied to the corpuscle, the fiber responded when the pressure was first applied and when it was removed, but it did not respond to continuous pressure
- But when Lowenstein dissected away the corpuscle and applied pressure directly to the fiber, the fiber fired to the continuous pressure
- Lowenstein concluded from this result that properties of the corpuscle cause the fiber to respond poorly to continuous stimulation, such as sustained pressure, but to respond well to changes in stimulation that occur at the beginning and end of a pressure stimulus or when stimulation is changing rapidly, as occurs in vibration
Vibration plays a role in perceiving what?
Fine textures
What’s surface texture?
The visual and tactile quality of a physical surface created by peaks and valleys
Why is visual inspection a poor way of determining surface texture?
- Because seeing texture depends on the light–dark pattern determined by the angle of illumination
- Thus, although the visually perceived texture of the 2 sides of something could look very different, moving the fingers across the 2 surfaces could reveal that their texture is the same
What’s the duplex theory of texture perception?
- The idea that texture perception is determined by both spatial and temporal cues that are detected by 2 types of receptors
- Originally proposed by David Katz and now called the “duplex theory”
- Although Katz proposed that texture perception is determined by both spatial and temporal cues, research on texture perception has, until recently, focused on spatial cues
What are the spatial cues from the duplex theory of texture perception?
- In tactile perception, information about the texture of a surface that is determined by the size, shape, and distribution of surface elements such as bumps and grooves
- Spatial cues are provided by relatively large surface elements, such as bumps and grooves, that can be felt both when the skin moves across the surface elements and when it is pressed onto the elements
- These cues result in feeling different shapes, sizes, and distributions of these surface elements
- Ex: perceiving a coarse texture such as Braille dots or the texture you feel when you touch the teeth of a comb
What are the temporal cues from the duplex theory of texture perception?
- In tactile perception, information about the texture of a surface that is provided by the rate of vibrations that occur as we move our fingers across the surface
- Temporal cues occur when the skin moves across a textured surface like fine sandpaper
- This type of cue provides information in the form of vibrations that occur as a result of the movement over the surface
- Temporal cues are responsible for our perception of fine texture that cannot be detected unless the fingers are moving across the surface
- Additional evidence for the role of temporal cues in perceiving texture has been provided by research that shows that vibrations are important for perceiving textures not only when people explore a surface directly with their fingers, but also when they make contact with a surface indirectly, through the use of tools
Describe how Mark Hollins and Ryan Risner (2000) presented evidence for the role of temporal cues
- By showing that when participants touched surfaces without moving their fingers and judge “roughness” using the procedure of magnitude estimation, they sensed little difference between 2 fine textures (particle sizes of 10 µm and 100 µm)
- However, when participants were allowed to move their fingers across the surface, they could detect the difference between the fine textures
- Thus, movement, which generates vibration as the skin scans a surface, makes it possible to sense the roughness of fine surfaces
- These results and the results of other behavioral experiments support the duplex theory of perception—that the perception of coarse textures is determined by spatial cues and of fine textures by temporal (vibration) cues
TRUE OR FALSE: You can detect differences in texture by running a tool over a surface
TRUE
- Your ability to detect differences in texture by running a pen (or some other “tool,” such as a stick) over a surface is determined by vibrations transmitted through the tool to your skin
- The most remarkable thing about perceiving texture with a tool is that what you perceive is not the vibrations but the texture of the surface, even though you are feeling the surface remotely, with the tip of the tool
Describe how Justin Lieber and Sliman Bensmaia (2019) studied how textures are represented in the brain and their findings
- By training monkeys to place their fingers on a rotating drum
- Textures ranged from very fine (microsuede) to coarse (dots spaced 5 mm apart)
- They studied the responses of 5 neurons in the somatosensory cortex to 6 different textures
- These patterns showed that
1. Different textures caused different firing patterns in an individual neuron
2. Different neurons responded differently to the same texture - These results showed that texture is represented in the cortex by the pattern of firing of many neurons
- Lieber and Bensmaia found that cortical neurons that fired best to coarse textures received input from SA1 neurons in the skin (Merkel receptors) and neurons that fired best to fine textures received input from PC receptors (Pacinian corpuscles)
What’s active touch?
- Touch in which the observer plays an active role in touching and exploring an object, usually with his or her hands and fingers
- Involves haptic perception
What’s passive touch?
A situation in which a person passively receives tactile stimulation on the skin, as when 2 points are pushed onto the skin to determine the two-point threshold
What’s the difference between active touch and passive touch?
With active touch, where you move your fingers across the object, you’re much more involved in the process than with passive touch, and you have more control over what parts of the objects to which you’re exposed
What’s haptic perception?
- The perception of three-dimensional objects by touch
- An example of a situation in which a number of different systems are interacting with each other
- These processes working together create an experience of active touch that is quite different from the experience of passive touch
What 3 distinct systems are used to arrive at the goal of identifying objects?
- Sensory system -> involved in detecting cutaneous sensations such as touch, temperature, and texture and the movements and positions of your fingers and hands
- Motor system -> involved in moving your fingers and hands
- Cognitive system -> involved in thinking about the information provided by the sensory and motor systems
Why is haptic perception an extremely complex process?
- Because the sensory, motor, and cognitive systems must all work together
- Ex: the motor system’s control of finger and hand movements is guided by cutaneous feelings in the fingers and the hands, by your sense of the positions of the fingers and hands, and by thought processes that determine what information is needed about the object in order to identify it
How did J. J. Gibson (1962) compare the experience of active and passive touch?
- By noting that we tend to relate passive touch to the sensation experienced in the skin, whereas we relate active touch to the object being touched
- Ex: if someone pushes a pointed object into your skin, you might say, “I feel a pricking sensation on my skin”; if, however, you push on the tip of the pointed object yourself, you might say, “I feel a pointed object”
- Thus, for passive touch you experience stimulation of the skin, and for active touch you experience the objects you are touching
According to psychophysical research, how quickly can people accurately identify most common objects using active touch?
Within 1 or 2 seconds
Describe Susan Lederman and Roberta Klatzky (1987, 1990) findings when they observed participants’ hand movements as they identified objects
They found that people use a number of distinctive movements, which they called exploratory procedures (EPs), and that the types of EPs used depend on the object qualities the participants are asked to judge
What are exploratory procedures (EPs)?
- People’s movements of their hands and fingers while they are identifying three-dimensional objects by touch
- People tend to use just 1 or 2 EPs to determine a particular quality
- Ex: people use mainly lateral motion and contour following to judge texture, and they use enclosure and contour following to judge exact shape
What are 4 of the EPs observed by Lederman and Klatzky?
- Lateral motion
- Pressure
- Enclosure
- Contour following
Describe the process that occurs between exploring objects with our fingers to activating specialized neurons
Exploring objects with our fingers and hands activates mechanoreceptors that send signals toward the cortex. When these signals reach the cortex, they eventually activate specialized neurons
What kind of receptive fields do neurons in the ventral posterior nucleus have?
- Center-surround receptive fields
- Similar to the center-surround receptive fields in the lateral geniculate nucleus, which is the visual area of the thalamus
What’s the tactile area of the thalamus?
The ventral posterior nucleus
What kind of neurons responding to touch do we see in the somatosensory cortex?
- In the cortex, we find some neurons with center-surround receptive fields and others that respond to more specialized stimulation of the skin
- There are neurons that respond to specific orientations and neurons that respond to movement across the skin in a specified direction
- There are also neurons in the monkey’s somatosensory cortex that respond when the monkey grasps a specific object -> in a monkey’s parietal cortex
Cortical neurons involved in tactile perception are affected not only by the properties of an object but also by what?
By whether the perceiver is paying attention
Describe Steven Hsiao and coworkers (1993, 1996) study on the effect of attention on cortical neuron responding to tactile information
- They recorded the response of neurons in areas S1 and S2 to raised letters that were scanned across a monkey’s finger
- In the tactile-attention condition, the monkey had to perform a task that required focusing its attention on the letters being presented to its fingers
- In the visual-attention condition, the monkey had to focus its attention on an unrelated visual stimulus
- The results show that even though the monkey is receiving exactly the same stimulation on its fingertips in both conditions, the response is larger for the tactile-attention condition
- Thus, stimulation of the receptors may trigger a response, but the size of the response can be affected by processes such as attention, thinking, and other actions of the perceiver
How does a person’s active participation influence perception?
- It influences what stimuli stimulate the receptors
- It influences the processing that occurs once the receptors are stimulated
What’s interpersonal touching/social touch?
One person touching another person
What kind of nerve fibers are found in hairy skin?
CT afferents
What are CT afferents?
- Unmyelinated nerve fibers found in hairy skin, which have been shown to be involved in social touch
- CT stands for C-tactile
- These fibers are unmyelinated, which means that they aren’t covered by the myelin sheath that covers the fibers associated with the receptors in glabrous skin
- The activity of these slow-conducting CT fibers was first recorded using a technique called microneurography
What’s the difference between myelinated and unmyelinated nerve fibers?
Unmyelinated fibers conduct nerve impulses much more slowly than myelinated fibers, a property which is reflected in the type of stimuli they respond to
What’s microneurography?
- Technique used to record neural signals that involves inserting a metal electrode with a very fine tip just under the skin
- When the skin on the forearm is stroked, the electrodes pick up signals as they are transmitted in nerve fibers conducting signals toward the brain
Describe patient G.L.
- 54-year-old woman who contracted a disease that destroyed all of her myelinated fibers, which she reported had caused her to lose her sensation of touch
- Careful testing revealed that she could detect light brush strokes presented to the hairy part of her forearm, which contains CT afferents
- Her sensations of light touch were accompanied by activation of the insula, which receives signals from CT afferents
- These results led Olausson to propose that CT afferents are involved in “caress-like skin to skin contact between individuals” -> the type of stimulation that came to be called social touch
What’s the social touch hypothesis?
- Research that followed the study of G.L. led to the social touch hypothesis
- Hypothesis that CT afferents and their central projections are responsible for social touch
- This was recognized as a new touch system that is different from the systems which sense the discriminative functions of touch
- The CT system, in contrast, is the basis of the affective function of touch
What are the discriminative functions of touch?
Functions of the touch system such as sensing details, texture, vibration, and objects
What are the affective functions of touch?
Sensing pleasure and therefore often eliciting positive emotions
Describe Line Loken and coworkers (2009) study on the affective functions of social touch
- They focused on the pleasant aspect of social touch by using microneurography to record how fibers in the skin responded to stroking the skin with a soft brush
- Loken found that the stroking caused firing in CT afferents and also in the SA1 (associated with Merkel receptors) and SA2 myelinated fibers associated with the discriminative functions, but with an important difference
- Whereas the response of SA1 and SA2 fibers continued to increase as stroking velocity increased all the way to 30 cm per second, the response of the CT afferents peaked at 3–10 cm per second and then decreased
- CT afferents are therefore specialized for slow stroking
- Loken also had participants rate the pleasantness of the sensation caused by this slow stroking and found a relationship between pleasantness and CT afferent firing (higher firing rates for CT afferents are associated with higher pleasantness ratings)
- Further research showed that maximum pleasantness ratings occurred at stroking speeds associated with optimal CT firing
What’s the main area of the brain that receives input from the CT afferents being stimulated by social touch?
- The insula
- Known to be involved in positive emotions
Describe Monika Davidovic and coworkers (2019) study on the role of the insula in the detection of pleasant touch
- They determined the functional connectivity between different part of the insula caused by pleasant touch and found that slow stroking creates connections between the back of the insula, which receives sensory information, and the front of the insula, which is connected to emotional areas of the brain
- Apparently, this connection to emotional areas helps create the pleasurable response to social touch
What kind of stroking of the arm (and other parts of the body) is pleasant?
Slow stroking
The pleasant effects of slow stroking can be influenced by what other factors in addition to the location and rate of stroking?
Knowledge of who is doing the stroking can determine whether the stroking is perceived as pleasant or unpleasant
Describe how Dan-Mikael Ellingsen and coworkers (2016) demonstrated the effect of knowledge on the perception of whether slow stroking is perceived as pleasant or unpleasant
- They demonstrated this effect by having heterosexual male participants rate the pleasantness on a scale of 1 (very unpleasant) to 20 (very pleasant) of a sensual caress to their arm
- They were led to believe that the caress was delivered by a female or a male
- Although the stroking was the same in both cases, the pleasantness rating was 9.2 if they thought they were being stroked by a male, and 14.2 if they thought they were being stroked by a female
- Results such as this demonstrate that although slow stroking is often pleasant, evaluation of the situation can turn a pleasant interaction into a less pleasant interaction or even a negative one
What’s an example of how top-down processing (AKA knowledge-based processing) can influence the perception of social touch?
The fact that people’s thoughts about who is touching them can influence their perception of pleasantness
What are the functions of pain?
Pain functions to warn us of potentially damaging situations and therefore helps us avoid or deal with cuts, burns, and broken bones
What is pain?
An unpleasant sensory and emotional experience associated with actual or potential tissue damage, or described in terms of such damage
What are the 3 different types of pain determined by Joachim Scholz and Clifford Woolf (2002)?
- Inflammatory pain
- Neuropathic pain
- Nociceptive pain
What’s inflammatory pain?
- Pain caused by damage to tissues, inflammation of joints, or tumor cells
- This damage releases chemicals that create an “inflammatory soup” that activates nociceptors
What’s neuropathic pain?
- Pain caused by lesions or other damage to the nervous system
- Ex: Carpal tunnel syndrome, which is caused by repetitive tasks such as typing; spinal cord injury; and brain damage due to stroke
What’s nociceptive pain?
Type of pain that serves as a warning of impending damage to the skin and is caused by activation of receptors in the skin called nociceptors
What are nociceptors?
- A fiber that responds to stimuli that are damaging or potentially damaging to the skin
- A number of different kinds of nociceptors respond to different stimuli -> heat, chemical, severe pressure, and cold
- Signals from the nociceptors are transmitted to the spinal cord and then up the spinal cord in pathways that lead to the brain
In the 1950s and early 1960s, how was pain explained?
Pain was explained by the direct pathway model of pain
What’s the direct pathway model of pain?
- The idea that pain occurs when nociceptor receptors in the skin are stimulated and send their signals directly from the skin to the brain
- This model doesn’t account for the fact that pain can be affected by other factors in addition to stimulation of the skin
- Examples such as not perceiving the pain from serious wounds or perceiving pain when no signals are being sent to the brain couldn’t be explained by the direct pathway model
Describe the report by Beecher (1959) on American soldiers wounded at the Anzio beachhead in World War II
- He reported that most American soldiers wounded at the Anzio beachhead in World War II “entirely denied pain from their extensive wounds or had so little that they didn’t want any medication to relieve it”
- One reason for this was that the soldiers’ wounds had a positive aspect: they provided escape from a hazardous battlefield to the safety of a behind-the-lines hospital
What’s the phantom limb phenomena?
- A person’s continued perception of a limb, such as an arm or a leg, even though the limb has been amputated
- Example in which pain occurs without any transmission from receptor to brain
- This perception is so convincing that amputees have been known to try stepping off a bed onto phantom feet or legs, or to attempt to lift a cup with a phantom hand
- For many, the limb moves with the body, swinging while walking
- It’s not uncommon for amputees to experience pain in the phantom limb
What causes pain in the phantom limb?
- One idea is that signals are sent from the part of the limb that remains after amputation
- However, researchers noted that cutting the nerves that used to transmit signals from the limb to the brain doesn’t eliminate either the phantom limb or the pain and concluded that the pain must originate not in the skin but in the brain
- This led Ronald Melzak and Patrick Wall (1965, 1983, 1988) to propose the gate control model of pain
What’s the gate control model of pain?
- Melzack and Wall’s idea that perception of pain is controlled by a neural circuit that takes into account the relative amount of activity in nociceptors, mechanoreceptors, and central signals
- This model has been used to explain how pain can be influenced by factors other than stimulation of receptors in the skin
- This model begins with the idea that pain signals enter the spinal cord from the body and are then transmitted from the spinal cord to the brain
- The model proposes that there are additional pathways that influence the signals sent from the spinal cord to the brain
- The central idea behind the theory is that signals from these additional pathways can act to open or close a gate, located in the spinal cord, which determines the strength of the signal leaving the spinal cord
- The gate control system consists of cells in the dorsal horn of the spinal cord
- According to this model, the perception of pain is determined by a balance between input from nociceptors in the skin and non-nociceptive activity from the skin and the brain
Along what 3 pathways does input to the gate control system occur?
- Nociceptors
- Mechanoreceptors
- Central control
Describe how input to the gate control system occurs along the nociceptor pathway
- Fibers from nociceptors activate a circuit consisting entirely of excitatory synapses, and therefore send excitatory signals to the transmission cells
- Excitatory signals from the (+) neurons in the dorsal horn “open the gate” and increase the firing of the transmission cells
- Increased activity in the transmission cells results in more pain
Describe how input to the gate control system occurs along the mechanoreceptor pathway
- Fibers from mechanoreceptors carry information about non-painful tactile stimulation
- An example of this type of stimulus would be signals sent from rubbing the skin
- When activity in the mechanoreceptors reaches the (–) neurons in the dorsal horn, inhibitory signals sent to the transmission cells “close the gate” and decrease the firing of the transmission cells
- This decrease in firing decreases the intensity of pain
What are transmission cells?
- According to gate control theory, it’s the cell that receives + and − inputs from cells in the dorsal horn
- Transmission cell activity determines the perception of pain
Describe how input to the gate control system occurs along the central control pathway
- These fibers, which contain information related to cognitive functions such as expectation, attention, and distraction, carry signals down from the cortex
- As with the mechanoreceptors, activity coming down from the brain also closes the gate, decreases transmission cell activity, and decreases pain
How can top-down processes affect pain perception?
Modern research has shown that pain can be influenced by what a person expects, how the person directs his or her attention, the type of distracting stimuli that are present, and suggestions made under hypnosis
Describe the hospital study on the effect of expectations on pain perception
In a hospital study in which surgical patients were told what to expect and were instructed to relax to alleviate their pain, the patients requested fewer painkillers following surgery and were sent home 2.7 days earlier than patients who were not provided with this information
How does the placebo effect relate to the effect of expectations on pain perception?
- Studies have shown that a significant proportion of patients with pathological pain get relief from taking a placebo, a pill that they believe contains painkillers but that, in fact, contains no active ingredients (placebo effect)
- The key to the placebo effect is that the patient believes that the substance is an effective therapy
- This belief leads the patient to expect a reduction in pain, and this reduction does, in fact, occur
- Many experiments have shown that expectation is one of the more powerful determinants of the placebo effect
What’s a placebo?
A substance that a person believes will relieve symptoms such as pain but that contains no chemicals that actually act on these symptoms
What’s the placebo effect?
A relief from symptoms resulting from a substance that has no pharmacological effect
Describe Ulrike Bingel and coworkers (2011) study on the effect of expectation on painful heat stimulation
- They demonstrated the effect of expectation on painful heat stimulation presented by an electrode on the calf of a person’s leg
- The heat was adjusted so the participant reported a pain rating of 70, where 0 corresponds to “no pain,” and 100 to “unbearable pain”
- Participants then rated the pain in a condition in which a saline solution was presented by infusion (baseline) and 3 conditions in which the analgesic drug remifentanil was presented, but the participants were told:
1. That they were still receiving the saline solution (no expectation)
2. That the drug was being presented (positive expectation)
3. That the drug was going to be discontinued in order to investigate the possible increase in pain that would occur (negative expectation) - The results indicate that pain was reduced slightly, from 66 to 65, in the no expectation condition when the drug infusion began, but dropped to 39 in the positive expectation condition, then increased to 64 in the negative expectation condition
- After the saline baseline condition, the participant was continuously receiving the same dose of the drug
- What was being changed was their expectation, and this change in expectation changed their experience of pain
- The decrease in pain experienced in the positive expectation condition is a placebo effect, in which the positive expectation instructions function as the placebo
- The negative effect caused by the negative expectation instructions is called a nocebo effect
Describe Ulrike Bingel and coworkers (2011) findings on the participants’ brain activity during the study on the effect of expectation on painful heat stimulation
- They found that the placebo effect was associated with increases in a network of areas associated with pain perception, and the nocebo effect was associated with increases in activity in the hippocampus
- A person’s expectation therefore affects both perception and physiological responding
What’s the nocebo effect?
A negative placebo effect, characterized by a negative response to negative expectations