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6: Other Senses Cognitive Neuroscience David Eagleman Jonathan Downar Chapter Outline Detecting Data from the World Hearing The Somatosensory System Chemical Senses The Brain is Multisensory Time Perception 2 Detecting Data from the World We use five senses to interact with the outside world, as traditionally defined. There are many subdivisions of those five, plus sensations from inside the body. For all senses, there are specialized receptors to transduce the stimulus. All senses project to primary sensory cortex, and have some form of mapping. 3 Hearing The Outer and Middle Ear Converting Mechanical Information into Electrical Signals: The Inner Ear The Auditory Nerve and Primary Auditory Cortex The Hierarchy of Sound Processing Sound Localization Balance 4 The Outer and Middle Ear Sounds are vibrations carried through air or water as waves We interpret the frequency (measured in Hz) of the vibration as the pitch of the sound. The amplitude of the vibration is interpreted as the loudness of the sound. 5 The Outer and Middle Ear 6 The Outer and Middle Ear The pinna collects and amplifies certain frequencies of sound and directs that sound down the ear canal. The sound energy strikes the tympanic membrane and causes it to vibrate at the same frequency as the sound wave. 7 The Outer and Middle Ear 8 The Outer and Middle Ear In the middle ear, vibrations of the tympanic membrane cause movement of the three bones of the middle ear. Malleus (Hammer) Incus (Anvil) Stapes (Stirrup) Movement of these bones causes movement of the oval window. 9 Converting Mechanical Information into Electrical Signals The oval window is part of the cochlea, which is the inner ear. The cochlea contains three fluid-filled tubes, wound around a central axis, like a snail shell. Inside the cochlea is the basilar membrane, which vibrates in time with the sound wave. 10 Converting Mechanical Information into Electrical Signals The basilar membrane is tighter at one end (the base) and looser at the other end (the apex). This difference means there is a tonotopic map of frequencies along the basilar membrane High frequencies near the base. Low frequencies near the apex. 11 Converting Mechanical Information into Electrical Signals 12 Converting Mechanical Information into Electrical Signals 13 Converting Mechanical Information into Electrical Signals Inner hair cells along the basilar membrane transduce sound into electrical signals. The vibration of the membrane causes the stereocilia to flex closer together or further apart. 14 Converting Mechanical Information into Electrical Signals Tip links on the stereocilia cause ion channels to be pulled open, depolarizing the cell, when the cilia move one direction. When cilia move the other direction, the channels close and the cell is hyperpolarized. 15 Converting Mechanical Information into Electrical Signals 16 Converting Mechanical Information into Electrical Signals The outer hair cells help to amplify and sharpen the incoming sound. The basilar membrane can break apart a complex sound into the component frequencies. 17 The Auditory Nerve and Primary Auditory Cortex The auditory (cochlear) nerve carries information from the inner hair cells to the cochlear nucleus of the brainstem. Each fiber is a labeled line, carrying information about only one frequency. Information travels from the cochlear nucleus through a number of nuclei to the primary auditory cortex in the temporal lobe. 18 The Auditory Nerve and Primary Auditory Cortex 19 The Hierarchy of Sound Processing Deafness can result from damage to the outer, middle, or inner ear. Damage that occurs to the auditory pathway after the inner ear typically results in damage to the ability to process sounds. Higher auditory areas are involved in the interpretation of sounds. 20 The Hierarchy of Sound Processing 21 Sound Localization We can localize sounds that occur around us based on characteristics of the sound and interaural differences. Interaural timing differences are used to identify the location of a sharp, brief sound. Interaural phase differences are used to localize a continuous sound. 22 Sound Localization 23 Balance The vestibular system, with three semicircular canals and two otolith organs, provides information about orientation. The semicircular canals detect head rotation and angular acceleration. The otolith organs detect linear acceleration. Both systems detect movement by the displacement of hair cells, similar to the auditory system. 24 Balance 25 Balance 26 The Somatosensory System Touch Temperature Pain Proprioception Interoception The Somatosensory Pathway 27 Touch The somatosensory system tells us about the external world, where our limbs are in space, and about our internal world. Receptors are found all over our skin and within our internal organs. 28 Touch Touch receptors are mechanoreceptors, which respond to stretching or bending. Meissner’s corpuscles and Merkel’s disks are located close to the surface of the skin and have small receptive fields. Pacinian corpuscles and Ruffini’s endings are located deeper in the skin and have large receptive fields. 29 Touch 30 Temperature Thermoreceptors are mechanoreceptors that convey temperature information. These receptors carry information about how the stimulus differs from the temperature of the skin. One population carries information about stimuli warmer than the skin and a separate population carries information about stimuli cooler than the skin. 31 Pain The perception of pain by nociceptors is necessary for our survival. There are three types of nociceptors. Mechanical nociceptors are activated by physical damage. Thermal nociceptors are activated by very high or very low temperatures. Chemical nociceptors are activated by particular chemicals. 32 Pain Some nociceptors are responsive to more than one type of stimuli and are called polymodal. Silent nociceptors respond to the body’s own chemical signals are can play a role in the increased sensitivity to stimulation following injury. 33 Pain Nociceptors appear to be free nerve endings. Different nociceptors transmit their signals at different rates. C fibers are small and unmyelinated, carrying the signal slowly. A delta fibers are myelinated and carry mechanical and thermal pain signals quickly. 34 Proprioception Proprioception is the sense of position and movement of our own body. Muscle spindles detect the length of the muscle and the speed of stretching. Golgi tendon organs provide information about muscle tension. 35 Proprioception 36 Interoception Interoception is our ability to perceive the internal state of our body, such as hunger, thirst, and mood. Receptors include stretch receptors and nociceptors. 37 Interoception 38 Interoception Gate control theory describes why we sometimes notice pain and sometimes do not, depending on the situation. If information from the interoceptors arrives at the central nervous system at the same time as nociceptive information, this can overwhelm the CNS, and the pain signals are blocked. 39 The Somatosensory Pathway Information from the head and face is carried by the trigeminal cranial nerve. Information from each dermatome of the body is carried into the dorsal horn of the spinal cord. Different types of somatosensory information follow different pathways to the brain. 40 The Somatosensory Pathway 41 The Somatosensory Pathway Somatosensory information decussates and is relayed by the thalamus to the primary somatosensory cortex (S1) S1 is in the parietal lobe, immediately posterior to the central sulcus. There is a somatotopic map of the body, known as the homunculus. 42 The Somatosensory Pathway 43 Chemical Senses Taste Smell The Sense of Flavor Pheromones 44 Taste For both taste and smell, molecules bind to receptors and trigger the release of neurotransmitters. Taste receptors are found on the tongue, palate, pharynx, epiglottis, and esophagus. Taste cells are grouped into taste buds, which are grouped into papillae. 45 Taste 46 Taste There are currently five basic tastes: Sweet, Sour, Bitter, Salty, Umami Salty and sour trigger ionotropic receptors. Sweet, bitter, and umami mostly trigger metabotropic receptors, but activate some ionotropic receptors as well. 47 Taste Most taste receptors appear to have a preferred type of taste they respond to, but can respond to other tastes in high concentration. Gustatory afferent neurons relay taste information to the brainstem and on to the frontal operculum. 48 Taste 49 Smell The olfactory epithelium is found in the back of the nasal cavity. Odorants dissolve in the mucus covering the epithelium and bind to receptors on the cilia of the olfactory receptor cells. Each olfactory receptor cell expresses only one type of receptor. Receptors are all GPCRs. 50 Smell 51 Smell Olfactory receptor cells project to the glomeruli in the olfactory bulb. 52 Smell Olfactory pathway travels from the olfactory bulb to the primary olfactory cortex in the rhinencephalon. 53 The Sense of Flavor Flavor is a combination of taste, smell, temperature, and texture. Higher-level processing within the brain evokes memories of past experiences. 54 Pheromones Pheromones are chemicals released to transmit information to and influence another member of the same species. Pheromones are detected by the vomeronasal organ. At this time, it is not clear to what extent humans use pheromones. 55 The Brain is Multisensory Synesthesia Combining Sensory Information The Binding Problem The Internal Model of the World 56 Synesthesia The brain needs to bring together information from different senses. Synesthesia is a condition where different senses are integrated inappropriately, such as when a letter is associated with a particular color. It appears to be due to heightened connectivity between brain regions. 57 Synesthesia 58 Combining Sensory Information Many neurons in the brain are more responsive when presented with more than one sensation at a time. In the McGurk effect, what a subject hears can be influenced by what they see at the same time. 59 Combining Sensory Information 60 The Binding Problem We do not perceive a stimulus as having separate visual and auditory components, but as a unified stimulus. How the sensory signals are integrated is known as the binding problem. Recurrent connections between the sensory systems likely are important for solving the binding problem. 61 The Binding Problem 62 The Internal Model of the World Our final perception does not rely only on stimuli from the outside world, but also on expectations from past experiences. In anosognosia, a patient is apparently unaware of their own physical limitations. 63 Time Perception Time involves input from all of the sensory systems. Time perception is a construct of the brain. Time appears to slow down in crucial situations because the amygdala encodes memories more thoroughly. These more dense memories make it seem like the event took longer. 64