CHAPTER 4: SENSATION

25% of Americans are supertasters, while 50% are normal taters. The remaining 25% are nontasters

PASSIVE PERCIEVERS Empiricism argued that our senses are passive

John Locke (emper.) argued that the human mind is much like a blank tableta tabula rasa, on which experience leaves its mark.

Distal stimulus: an object or event in the outside world.

Proximal stimulus: the energies from the outside world that directly reach our sense organs. e.g. sound waves are the proximal stimulus for hearing.

George Berkeley

Empiricist believed that we can perceive and move around in the three dimensional world because our experience has taught us how to interpret the two dimensional proximal stimulus.

Depth cuesthese cues include whats called visual perspectivea cue used to convey depth in many paintings.

ACTIVE PERCIEVERS

Immanuel Kant argued that perception is possible only because the mind organizes the sensory information into certain preexisting categories. Kant claimed that we have an innate grasp of certain special relationships and an innate grasp of temporal relationships (what it means for one event to occur before another).

PHSYCOPHYSICS

Definition: The area of research that links psychological experience to physical stimuli, is called psychophysics.

We are trying to understand the relationship between the world as it actually is and the world as we perceive it to be.

Sensory threshold

Absolute threshold: the smallest quantity of an input that can be detected.

Difference threshold: the smallest change in an input that can be detected.

When a stimulus is changed by this minimal amount, psychophysicists call it a just noticeable difference or jnd.

Difference thresholds show a consistent property they depend on proportional differences, and not on absolute differences.

Webers Law: The observation that the size of the difference threshold is proportional to the intensity of the standard stimulus

c=I/I

I is the intensity of the standard stimulus, the one to which comparisons are being made, delta I is the amount that must be added to this intensity to produce a just noticeable increase; c is a constant. The right hand side of the equation is known as the Weber fraction Fechners Law: describes the relationship between the physical intensity of a stimulus and the psychological intensity of the experience produced by that experience.

The observation that the strength of a sensation is proportional to the logarithm of physical stimulus intensity.

The law states that the strength of a sensation increases logarithmically with the intensity of the stimulus. S=k*log(I) S stands for psychological magnitude; I is the physical intensity of the stimulus; and k is a constant whose value depends on the value of the Weber fraction. In truth Fechners Law does not hold for all circumstances

A Survey of the Senses Sensory Coding The physical stimulus must be converted into neural signal; this is the step of transduction. Sensory Coding: The process through which the nervous system represents the qualities of the incoming stimulus whether auditory or visual, for example, or whether a red light or a green one, a sour taste or a sweet taste. One aspect of sensory coding involves psychological intensity the difference between the bright light and the dim one, or a subtle scent of cinnamon in contrast to a dense cloud of the smell. The second part of sensory coding is sensory quality how the nervous system represents the difference between, say, vision and hearing; or within a modality, how it represents the difference between, for example, a high pitched note or a low one, or the difference between a sweet taste and a bitter one. Specific theory: This proposal suggest that different sensory qualities (sweet vs. sour, red vs. green) are signaled by different neurons, just as the different sense modalities (vision vs. pressure) are signaled by different nerves. Pattern Theory: the proposal that what allows us to identify the input is the overall pattern of activation which neurons are firing more and which are firing less at a given moment. Sensory Adaptation Our sensory responses are also influenced by change. Sensory Adaptation: This term refers to the way our sensory apparatus registers a strong response to a stimulus when it first arrives, but then gradually decreases that response if the stimulus is unchanged. Vestibular Sense Humans have more than five senses. Kinesthesis: The name for the sensation coming from various receptors in the muscles, tendons, and joints that informs us of our skeletal motion. Vestibular Sense: The sensation generated by receptors in the semicircular canals of the inner ear that inform us about the head s orientation and movement. As we move through the world, our heads move continuously. To compensate for the endless rocking, our eyes have to move accordingly. This adjustment is accomplished by a reflex system

that s coordinated by the cerebellum but initiated by messages from the vestibular sense. In this way, the visual system is effectively stable and operates as if it s resting on a solid tripod. The Skin Senses Skin Senses: The group of senses, including pressure, warmth, and pain, through which we gain information about our immediate surroundings. Skin senses may be the best example of specific coding (or labeled lines ). Pain Pain usually begins with activity in the nociceptors receptors in the skin that have bare nerve endings and that respond to various forms of tissue damage as well as to temperature extremes. These receptors come in two types: 1. A-delta fibers: allow for rapid transmission of information and are responsible for the pain you experience when you re first injured. 2. C-fibers: (in contrast) are unmyelinated and are therefore slower in the transmission; they re the source of the dull ache that remains long after the injury occurs. Natural internal painkillers, called endorphins, help us in many circumstances, to continue functioning despite a seemingly painful injury. Pain can also be blocked via a different mechanism: According to the gate control theory, pain sensation must pass through a neural gate to reach the brain and can be blocked at the gate by neurons that inhibit signals from the nociceptors, so that these signals are never transmitted to the brain. The experience of pain is influenced by someone s beliefs and emotions. Smell Complex molecules drift through the air, and some are drawn up the nose to a mucous membrane called the olfactory epithelium it s located at the top of the nasal cavity and contains the olfactory receptor neurons that respond to airborne molecules called odorants. Olfactory bulb, a brain structure above the nasal cavity and just beneath the frontal lobe. Within the olfactory bulb, the axons converge at sites called glomeruli. The nose contains roughly 1,000 types of receptors. Each glomerulus receives input from one of these receptor types. We ve already mentioned 1,000 types of receptors and therefore 1,000 types of glomeruli but in fact humans can distinguish roughly 10,000 different odors. Right away, these numbers tell us that our sense of smell does not rely on one-glomerulus-per-scent coding we don t rely on

labeled lines that use one neural pathway for each scent. It must be that each odor we can distinguish produces some unique pattern of activation in various glomeruli. Smell-based communication within a species usually involves specialized chemicals called pheromones biologically produced odorants that convey information to other members of the species. What causes menstrual synchrony? The answer turns out to be scent. Taste The proximal stimuli are molecules of particular shape that react with receptor cells. The molecules called tastants are carried by fluid, not by air (odorants). The taste receptors are located primarily on the tongue, although many are found on the roof of the mouth and the upper throat. The tastant molecules are carried to the papillae that cover our tongue. Each papillae contains hundreds of taste buds, and each taste bud contains a hundred or so receptor cells. There are just five sense types evenly distributed over the whole tongue: 1. 2. 3. 4. 5. Salt Sour Bitter Sweet Umami ( savory )

Each receptor type responds, at least to some degree, to all tastants. Whatever their physical basis, though, it s clear that tastes often have a special biological role. For example, many nutritive substances contain some form of sugar, and so natural selection would have favored organisms with a preference for sweets. On the other hand, many toxic substances are bitter, and so natural selection would likely have favored organisms that avoided bitter tastes. Supertasters: These are people who seem enormously sensitive to certain taste, probably because they literally have more papilla than other people do. We can identify supertasters by placing on their tongue a bit of paper impregnated with the chemical compound propylthiouracil(PROP). For supertasters, this chemical has an extremely bitter taste. Other taste preferences are influenced by learning. Conditioned taste aversion: when an organism, due to specific experience, comes to associate a specific taste with illness and from then on seems to find that flavor repulsive.

In other cases the learning may simply involve a matter of familiarity. Familiarity plays a key role in defining a cultures cuisine. Hearing Sound waves are a series of pressure variations in the air that vary in amplitude and wavelength. When these sound waves hit our ears, they initiate a set of further changes that ultimately triggers the auditory receptors. The amplitude (the height of the wave crest used as a measure of sound intensity) is the amount of pressure exerted by each air particle on the next. Frequency: The number of wave peaks per second. In sound, frequency governs the perceived pitch of sound. **Psychologist as well as physicist, audio engineers, and many others routinely use Fourier analysis to describe in detail the sound we encounter in our day-to-day experience. From Sound Waves to Hearing Gathering the Sound Waves

Mammals have their receptors for hearing deep within the ear, in a snail-shaped structure called the cochlea. The outer ear collects the sound waves from the air and directs them toward the eardrum, a taut membrane at the end of the auditory canal. The sound waves make the eardrum vibrate, and these vibrations are transmitted to the oval window, the membrane that seperates the middle ear from the inner ear. This transmission is accomplished by a trio of tiny bones called the auditory ossicles the smallest bones in the human body. The vibration of

the eardrum move the first ossicle (the malleus), which then moves the second (the incus), which in turns moves the third (the stapes). The stapes completes the chain by sending the vibration pattern on to the oval window, which the stapes is attached to. The movement of the oval window then gives rise to waves in the fluid that fills the cochlea, causing (at last) a response to the receptors. In order for us to hear, the changes in air pressure must cause changes in fluid pressure and this is a problem, because fluid is harder to set in motion than air is. To solve this problem, the pressure waves have to be amplified as they move toward the receptors. This is the reason why there s such a long chain of events to stimulate the auditory receptors. Transduction in the Cochlea Basilar membrane: A membrane running the length of the cochlea; sound waves cause a deformation of this membrane, bending the hair cells in the cochlea and thus stimulating the auditory receptors. The actual auditory receptors the 15,000 hair cells in each ear (in the cochlea) are lodged between the basilar membrane and other membranes above it. Place Theory: A proposal about pitch perception stating that regions of the basil membrane respond to particular sound frequencies and the nervous system interprets the excitation from different basil regions as different pitches. Other means of sensing pitch is tied to the firing rate of cells in the auditory nerve. The ear has two ways of encoding pitch: based on the location of maximum movement of movement on the basilar membrane, and based on firing rate of cells in the auditory nerve. For higher-pitched sounds, the location-based mechanism plays a larger role; for lower-pitched sounds, the frequency of firing is more important. Further Processing of Auditory Information Neurons carry the auditory signals from the cochlea to the midbrain. From there, the signals travel to the geniculate nucleus in the thalamus. Other neurons than carry the signals to the primary projection are for hearing, in the context of the temporal lobe. The auditory signal must be analyzed for its timbre sound quality that helps us distinguish a clarinet from an oboe or one persons voice from another s. It turns out that each neuron along the auditory pathway responds to a wide range of pitches, but even so, each has a preferred pitch a frequency of sound to which that neuron fires more vigorously than it fires to any other frequency.

Neurons with similar preferred pitches tend to be located close to each other on the cortex. This arrangement creates what s known as a tonotopic map a map organized on the basis of tone. Vision Visual information overrules the evidence you receive from your ears. The Stimulus: Light Like sounds waves, these light waves can be described in terms of two measurements. First, light waves can vary in amplitude, which is the major determinant of perceived brightness. Second, light waves vary in frequency how many times per second the wave reaches its maximum amplitude. It s more convenient to describe light waves using the inverse of frequency wavelength, the distance between the crest of two successive waves. Wavelengths are measured in nanometers and are a major determinant of perceived color. Visible Spectrum Ranges from 350-750 nm Gathering the Stimulus: The Eye The actual detection of light is done by cells called photoreceptors. These cells are located in the retina and convert light energy into neural impulses. Retinal Image: The image of an object that is projected on the retina. Its size increase with the size of that object and decreases with the object s distance from the eye. The iris is a smooth, circular muscle surrounding the papillary opening. It causes the pupil to dilate (grow large) or contract, thus allowing considerable control over how much light reaches the retina. The cornea and the lens focus the incoming light just like a camera lens does.

The human eye: Light enters the eye through the cornea, and the cornea and lens refract the light rays to produce a sharply focused image on the retina. The iris can open or close to control the amount of light that reaches the retina. The retina is made up of three main layers: the rods and cones, which are photoreceptors; the bipolar cells; and the ganglion cells, whose axons make up the optic nerve. The Visual Receptors It is at the retina that the physical stimulus energy is transduced into a neural impulse. The retina contains two kinds of receptor cells, the rods and the cones. The cones are plentiful in the fovea, a small, roughly circular region at the center of the retinal; but they become less and less prevalent at the outer edges of the retina. The opposite is true of the rods; they re completely absent from the fovea, but more numerous at the retina s edge. The rods and the cones do not report to the brain directly.

The receptor cells stimulate the bipolar cells, and these in turn excite the ganglion cells. The ganglion cells collect information from all over the retina, and the axons of these cells then converge to form a bundle of fiber that we call the optic nerve. Leaving the eyeball the optic nerve carries info first to the lateral geniculate nucleus in the thalamus and then to the cortex. The blind spot: no cones or rods at that location. Rods and cones differ in their structure, number and placement on the retina; they also differ in their function. The rods are the receptors for night vision; they operate at low light intensities and lead to achromatic (colorless) sensations. The cones serve day vision; they respond at much higher levels of illumination and are responsible for sensation of color. So we have two separate receptor systems, one for vision in dim light and the other for vision in bright light. Acuity the ability to perceive detail is much greater in the cones. The fovea is where the cones are more closely packed and visual acuity is greatest. By looking slightly away from the star, you can ensure that the star s image falls outside of the fovea and onto a region of the retina that s dense with the more light sensitive rods. Photopigment: a chemical in the photoreceptor that changes its form in response to light, producing an electrical change that signals to the nervous system that light is present. Rods and cones contain different photopigments. The rods contain rhodopsin, a pigment that breaks down more readily in response to light than the cone pigments do. Consequently, rods can function at lower light levels. There are three different cone photopigments. The differences among the three photopigments are crucial for the cones ability to discriminate color. Each of us is nearly color blind at the visual periphery. Contrast Effect Visual system is also sensitive to spatial contrast A change in brightness typically marks a visual boundary, a point where one object begins and another stop. And, of course these boundaries are immensely important for the visual system because they define the object s shape and shape in turn is generally the info we use to identify an object. Visual system amplifies brightness boundaries by a process called edge enhancement which relies on brightness contrast and allows us to see the edge between objects more clearly.

When a light region borders a dark region, the contrast between the two makes the light region look even lighter and makes the dark regions look darker. The contrast highlights the edge where the two regions meet. Lateral Inhibition is the event in the nervous system that leads to brightness contrast. Lateral inhibition is a pattern of interaction among neurons in which activity in one neuron decreases the response in adjacent neurons. Recordings from single cells in the retina confirm that activity in one cell actually causes the immediately adjacent cell to fire less than they otherwise would. All the cells detecting the edge of a bright surface end up producing a stronger response than that of the cells detecting the middle of the surface. This pattern will then lead to an exaggerated response along the surface s edges, making these edges easier to detect. The reverse happens for cells being stimulated by a patch that s not as bright. Look at pg. 167 for clarification

Color A person with normal color vision can distinguish over 7 million shades of color. But fortunately, this staggering number of colors can be classified in terms of just three dimensions. 1. Hue: This dimension corresponds closely to the way we use color. Varies in wavelength. 2. Brightness: is the dimension that differentiates black from white and the various shades of gray in between. 3. Saturation: the purity of a color The Neural Basis of Color Vision: Color Receptors Human color vision is trichromatic based on three elements, each tied to one type of cone. Each of the three types of cones contains a different photopigment. One pigment is most sensitive to wavelengths in the short-wave region of the spectrum. And so on Note that all three types of cones respond to most of the wavelengths in the visible spectrum. If the input contains A, B, and C the short-preferring cones response to this stimulus will simply be the total of those cones response to A when it s present alone, plus B when present alone, plus their response to C.

All that matters is the sum. And this is crucial, because it s almost always possible to find different mixes of wavelengths that will produce the same three totals. Complementary Hues There is three cones, each with its photopigment. Suppose that you stare at a green patch for a while and then look at a white wall. You ll see a negative afterimage of the patch in this case a reddish spot. In the same way staring at a red patch will produce a green afterimage; staring at something blue will produce a yellow afterimage; and staring at something yellow will produce a blue afterimage. In all cases, the afterimage has the complementary hue of the original stimulus. It appears that colors are paired, such that each color has an opposite that cancels it. The Opponent Process Theory Opponent Process Theory: This theory begins with the undeniable fact that we have three cone types, but it argues that the output from these cones is then processed by another layer of neural mechanism that recode the signal on the basis of three pairs of colors red versus green, blue versus yellow, and black versus white. Many of the neurons in the visual system behave exactly as the theory proposes. Color Blindness Many genes, on at least 19 different chromosomes, can contribute to color blindness. One of the genetic causes involves a gene mutation on the x chromosome. Color blindness is therefore more common among men than it is among women. Men have an xy genetic pattern. If there x chromosome contains the mutated gene, men have no backup gene on another chromosome and color blindness is the result. Perceiving Shape Featured Detectors Results in the lab show that the cells in the visual system all have a preferred target, a certain type of stimulus that s especially effective in causing that cell to fire. David Hubel and Torsten Wiesel These investigators confirmed that each cell in the visual cortex responds to stimuli in only a limited region of space or, equivalently, each cell in the retina responds to stimuli on only a limited region of the retina. This region defines that cells receptive field (the pattern of retinal stimulation that causes the cell in the visual system to fire).

Other cells fire only in response to a line or edge of a particular orientation. Cells like these serve as feature detectors, detecting certain elements within the visual pattern.