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Chapter Six: Vision Assorted Materials from Modules 1-3 Chapter Seven: Audition Module 1 Visual Coding and Retinal Receptors Reception- absorption of physical energy (electromagnetic waves) by receptors Transduction-the conversion of physical energy to an electrochemical pattern in the neurons Coding- one-to-one correspondence between some aspect of the physical stimulus and some aspect of the nervous system activity Visual Coding and Retinal Receptors From Neuronal Activity to Perception coding of visual information in the brain does not duplicate the stimulus being viewed General Principles of Sensory Coding Muller and the law of specific energies-any activity by a particular nerve always conveys the same kind of information to the brain Now close your lid and poke your eye… do you see light (fosphenes)(sp?) Qualifications of the Law of Specific Energies the rate of firing or pattern of firing may signal independent stimuli timing of action potentials may signal important information indicating such things as movement the meaning of one neuron depends on what other neurons are active at the same time Visual Coding and Retinal Receptors“Look into My Eyes!” The Eye and Its Connections to the Brain Pupil-opening in the center of the eye that allows light to pass through Lens-focuses the light on the retina Retina-back surface of the eye that contains the photoreceptors The Fovea-point of central focus on the retina The Route Within the Retina photoreceptors-rods and cones bipolar cells-receive input from rods and cones ganglion cells-receive input from bipolar cells optic nerve-made up of axons of ganglion cells blind spot-the point where the optic nerve leaves the eye Figure 6.2 Cross section of the vertebrate eye Note how an object in the visual field produces an inverted image on the retina. Figure 6.4 Visual path within the eyeball The receptors send their messages to bipolar and horizontal cells, which in turn send messages to the amacrine and ganglion cells. The axons of the ganglion cells loop together to exit the eye at the blind spot. They form the optic nerve, which continues to the brain. Receptive Field- the part of the visual field to which any one neuron responds Key Point: Fields have both excitatory and inhibitory regions •The receptive field of a receptor is simply the area of the visual field from which light strikes that receptor. •The receptive field for any other neurons in the visual system is determined by which receptors connect to the cell in question. Figure 6.6 Two demonstrations of the blind spot of the retina Close your left eye and focus your right eye on the o in the top part. Move the page toward you and away, noticing what happens to the x. At a distance of about 25 cm (10 inches), the x disappears. Now repeat this procedure with the bottom part. At that same distance what do you see? Visual Receptors: Rods and Cones, Reception & Transduction continued Rods -abundant in the periphery of the retina -best for low light conditions -see black/white and shades of gray Cones abundant around fovea best for bright light conditions see color Table 6.1 is very good Transduction Both Rods and Cones contain photopigments (chemicals that release energy when struck by light) 11-cis-retinal is transformed into all-trans-retinal in light conditions this results in hyperpolarization of the photoreceptor the normal message from the photoreceptor is inhibitory Light inhibits the inhibitory photoreceptors and results in depolarization of bipolar and ganglion cells Theories Color Vision The Trichromatic (Young-Helmholtz) Theory KEY POINT: We perceive color through the relative rates of response by three kind of cones, each kind maximally sensitive to a different set of wavelengths, but receptors are not equally distributed across retina. (exercises) Bowmaker & Dartnall (1980) projected a known amount of light directly through the outer segments of photoreceptors and measured how much light was absorbed by the photopigment molecules. They found four classes of photopigments. The wavelength of maximum absorbance is indicated at the top of each curve. The 420 curve is for the short wavelength cones (blue), the 498 curve is for the rods, and the 534 and 564 curves are for the middle (green) and long wavelength (red) sensitive cones respectively. More Theories of Color Vision The Opponent-Process Theory we perceive color in terms of paired opposites RED vs. GREEN; YELLOW vs. BLUE; BLACK vs. WHITE one color the result of excitation, the other the result of inhibition of bipolar cells Can explain negative color after-image effects The Retinex Theory When information from various parts of the retina reaches the cortex, the cortex compares each of the inputs to determine the brightness and color perception for each area. can explain color constancy only works when entire view has been tinted Figure 6.12 Possible wiring for one bipolar cell Short-wavelength light (which we see as blue) excites the bipolar cell and (by way of the intermediate horizontal cell) also inhibits it. However, the excitation predominates, so blue light produces net excitation. Red, green, or yellow light inhibit this bipolar cell because they produce inhibition (through the horizontal cell) without any excitation. The strongest inhibition is from yellow light, which stimulates both the long- and medium-wavelength cones. Therefore we can describe this bipolar cell as excited by blue and inhibited by yellow. White light produces as much inhibition as excitation and therefore no net effect. (Actually, receptors excite by decreasing their usual inhibitory messages. Here we translate that double negative into excitation for simplicity.) Neural Basis of Visual Perception An Overview of the Mammalian Visual System Rods and Cones synapse to amacrine cells and bipolar cells Bipolar cells synapse to horizontal cells and ganglion cells Axons of the ganglion cells leave the back of the eye The inside half of the axons of each eye cross over in the optic chiasm Pass through the lateral geniculate nucleus Transferred to visual areas of cerebral cortex Neural Basis of Visual Perception Concurrent Pathways in the Visual System In the Retina and Lateral Geniculate Two categories of Ganglion cells – Parvocellular-smaller cell bodies and small receptive fields, located near fovea; detect visual details, color – Magnocellular-larger cell bodies and receptive fields, distributed fairly evenly throughout retina; respond to moving stimuli and patterns In the Cerebral Cortex V1-Primary Visual Cortex-responsible for first stage visual processing V2-Secondary Visual Cortex-conducts a second stage of visual processing and transmits the information to additional areas Ventral stream-visual paths in the temporal cortex Dorsal stream-visual path in the parietal cortex Figure 6.20 Three visual pathways in the cerebral cortex (a) A pathway originating mainly from magnocellular neurons. (b) A mixed magnocellular/parvocellular pathway. (c) A mainly parvocellular pathway. Neurons are heavily connected with other neurons in their own pathway but only sparsely connected with neurons of other pathways. Area V1 gets its primary input from the lateral geniculate nucleus of the thalamus; the other areas get some input from the thalamus but most from cortical areas. (Sources: Based on DeYoe, Felleman, Van Essen, & McClendon, 1994; Ts’o & Roe, 1995; Van Essen & DeYoe, 1995) Neural Basis of Visual Perception- The Cerebral Cortex: The Shape Pathway Hubel and Wiesel’s Cell Types in the Primary Visual Cortex Simple Cells has fixed excitatory and inhibitory zones in its receptive field Complex Cells receptive fields cannot be mapped into fixed excitatory and inhibitory zones Respond to a pattern of light in a particular orientation Figure: The receptive field of a complex cell in the visual cortex: -It is like a simple cell in that its response depends on a bar of light’s angle of orientation. -It is unlike a simple cell in that its response is the same for a bar in any position within the receptive field. Neural Basis of Visual Perception- The Columnar Organization of the Visual Cortex Columns are grouped together by function – Ex: cell within a given column respond best to lines of a single orientation Are Visual Cortex Cells Feature Detectors? Feature Detectors-neurons whose responses indicate the presence of a particular feature Shape Analysis Beyond Areas V1 and V2 Inferior Temporal Cortex (V3)-detailed information about stimulus shape (V4)-Color Constancy; Visual Attention (V5)-Speed and Direction of Movement Neural Basis of Perceptual Disorder Disorders of Object Recognition Visual Agnosia-Inability to Recognize Objects Prosopagnosia-Inability to recognize faces Color Vision Deficiencies Complete and Partial Color Blindness-inability to perceive color differences Generally results from people lacking different subsets of cones genetic contributions- same photopigment made on medium and longwave wavelength receptors Neural Basis of Visual PerceptionThe Cerebral Cortex The Cerebral Cortex: The Color Pathway Parvocellular to V1 (blobs) to V2, V4, and Posterior Inferior Temporal Cortex The Cerebral Cortex: The Motion and Depth Pathways Structures Important for Motion Perception Middle-temporal cortex-V5-speed and direction of movement Motion Blindness-Inability to detect objects are moving Experience and Visual Development Early Lack of Stimulation of One Eye-blindness occurs in that one eye Early Lack of Stimulation of Both Eyes-if this occurs over a long period of time, loss of sharp receptive fields is noted Restoration of Response and Early Deprivation of Vision-deprive stimulation of the previously active eye and new connections will be made with the inactive eye Uncorrelated Stimulation in Both Eyes-each cortical neuron becomes responsive to the axons from just one eye and not the other Experience and Visual Development Early Exposure to a Limited Array of Patterns—most of the neurons in the cortex become responsive only to the patterns that the subject has been exposed to Lack of Seeing Objects in Motion-become permanently disable at perceiving motion Effects of Blindness on the Cortex-parts of the visual cortex become more responsive to auditory and tactile stimulation Chapter Seven The Nonvisual Sensory Systems- Auditory System Module One Audition Sound and the Ear Physical and Psychological Dimensions of Sound Amplitude=intensity of wave=loudness frequency=number of waves/second=pitch Figure 7.1 Four sound waves The time between the peaks determines the frequency of the sound, which we experience as pitch. Here the top line represents five sound waves in 0.1 second, or 50 Hz—a very low-frequency sound that we experience as a very low pitch. The other three lines represent 100 Hz. The vertical extent of each line represents its amplitude or intensity, which we experience as loudness. Anatomy of the Ear Structures of the Ear Pinna-cartilage attached to the side of the head Tympanic Membrane-eardrum middle ear boneshammer/anvil/stirrup oval window-membrane leading to inner ear cochlea-three fluid-filled tunnels scala vestibuli scala media scala tympani basilar membrane-flexible membrane tectorial membrane-rigid membrane hair cells-auditory receptors Figure 7.2 Structures of the ear When sound waves strike the tympanic membrane in (a), they cause it to vibrate three tiny bones—the hammer, anvil, and stirrup—that convert the sound waves into stronger vibrations in the fluid-filled cochlea (b). Those vibrations displace the hair cells along the basilar membrane in the cochlea. (c) A cross section through the cochlea. The array of hair cells in the cochlea is known as the organ of Corti. (d) A closeup of the hair cells. Pitch Perception Theories of Pitch Perception Frequency theory the basilar membrane vibrates in synchrony with a sound, causing auditory nerve axons to produce action potentials at the same frequency Place theory the basilar membrane resembles the strings of a piano in that each area along the membrane is tuned to a specific frequency and vibrates whenever that frequency is present Volley principle the auditory nerve as a whole can have volleys of impulses up to about 5,000 per second, even though no individual axon can approach that frequency by itself Figure 7.4 The basilar membrane of the human cochlea High-frequency sounds produce their maximum displacement near the base. Low-frequency sounds produce their maximum displacement near the apex. Figure 7.5 Traveling waves in the basilar membrane set up by different frequencies of sound Note that the peak displacement is closer to the base of the cochlea for high frequencies and is toward the apex for lower frequencies. In reality the peak of each wave is much narrower than shown here. Pitch Perception in the Cerebral Cortex Primary auditory cortex Each cell responds best to one tone Cells preferring a given tone cluster together Secondary auditory cortex Each cell responds to a complex combination of sounds Figure 7.6 Route of auditory impulses from the receptors in the ear to the auditory cortex The cochlear nucleus receives input from the ipsilateral ear only (the one on the same side of the head). All later stages have input originating from both ears. Hearing Loss Conductive Deafness bones of the middle ear fail caused by tumors, infection, disease usually corrected by surgery or hearing aids Nerve Deafness damage to cochlea, hair cells or auditory nerve usually treated with hearing aids caused by genetics, disease, ototoxic drugs, etc. Localization of Sound Sound Shadowloudest in nearest ear Time of arrivalarrives at one ear soonest Phase differencesounds arrive out of phase dependent on frequency Figure 7.10 Phase differences between the ears as a cue for sound localization Note that a low-frequency tone (a) arrives at the ears slightly out of phase. The ear for which the receptors fire first (here the person’s left ear) is interpreted as being closer to the sound. If the difference in phase between the ears is small, then the sound source is close to the center of the body. However, with a high-frequency sound (b) the phase differences become ambiguous. The person cannot tell which sound wave in the left ear corresponds to which sound wave in the right ear.