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The Auditory System Sound is created by pressure waves in air; these waves are induced by vibrating membranes such as vocal cords. Because the membranes usually vibrate in a regular manner, the pressure waves have a fixed spacing. Sound has two main characteristics: A. the spacing between waves or period. This can also be thought of in terms of how many wave cycles pass by in one secondthat is, the sound’s frequency in Hz=cycles/second. B. The intensity of amplitude of the sound measured in decibels. Bear et al. The Auditory Periphery Bear et al. Kandel et al. The outer ear and canal guide and filter sound. The tympanic membrane and ossicles transmit the vibrations to the cochlea itself; the vibrations enter the cochlea via the round window and exit via the round window. As they pass through the endolymph of the scala vestibuli and tympani, sound waves cause the basilar membrane to vibrate. This is the key to auditory function. The Cochlea Sound waves cause the basilar membrane to vibrate. The basilar membrane is stiff at its base and loose at its apex. Just like a guitar string, this causes it to resonate to high frequencies at its base and low frequencies at its apex. There is therefore a tonotopic map- a location code- formed on the cochlea.This is a fundamental feature of auditory coding and is preserved right up to cortex. Note that this is a separate code from synchronization. Bear et al. Bear et al. Hair Cells Hair cells transduce vibrations into depolarization. This in turn leads to vesicular release that excites auditory afferent fibers and causes them to discharge. Hair cells are firmly attached to the basilar membrane and therefore move up and down with it as it vibrates. The “hairs” or cilia of these cells are attached to a tectorial membrane; this membrane is fixed- it does not vibrate in response to sound. So, as you can imagine, when the basilar membrane moves upward, the cilia will be bent. This is the first step in the transduction process. A scanning electron microscopic view of the beautiful organization of hair cell cilia in the cochlea. Kandel et al. Hair Cells 2 When the cilia bend in one direction it causes the hair cell to hyperpolarize; bending in the opposite direction causes depolarization. Bear et al. This effect is due to the mechanical coupling of the cilia to K+ channels at their tips. The depolarization causes Ca2+ entry and the fusion of vesicles and release of glutamate from hair cells. This cause excitation and spiking of the auditory afferent fibers. Auditory Afferent fibers Each auditory afferent fiber is tuned to a specific frequency. The tuning is simply due to the location of its hair cell along the cochlea. This tonotopic mapping is preserved in the projection of cochlear afferents to the cochlear nuclei in the medulla. As I mentioned earlier, cochlear afferents are also phase locked to sound (especially low frequency sounds). So there are two ways to code for sound. A firing rate “place” code (tonotopy) and a temporal code. The central auditory system uses both codes for various purposes. This is a general principle. Sensory systems are flexible and can use multiple coding strategies. Bear et al. Central Auditory Pathways Auditory reach the cochlear nuclei of the medulla (DCN, VCN). From these nuclei a direct pathway goes to the inferior colliculus, then the thalamic medial geniculate nucleus (MGN) and then onto the auditory primary cortex. Note that this pathway is bilateral unlike the contralateral somatosensory pathway. This makes sense since sound always reaches both ears. On the way up to cortex axons from VCN also terminate in nuclei of the superior olivary complex. Neurons in this cell group use relative sound timing and intensity in the two ears to estimate the spatial location of a sound source. Bear et al. Sound Localization Sounds coming from the right arrive at the right ear a little earlier than the at the left ear. This small time difference is used by the superior olive. Bear et al. A cell in the superior olive responds with an increase in firing rate to a time difference in the arrival of sound to the ears. This cue to the sounds location is conveyed up to the inferior colliculus and onto cortex. Relative sound intensity is also used as a cue in different neurons in the superior olive region. The neural mechanisms involved are becoming understood but are beyond the scope of an introductory course. Cortical Processing of Sounds Bear et al. The MGN projects up to the primary auditory cortex (there are secondary auditory areas as welll). Notice that the cortex still has tonotopy. However auditory cortex neurons also respond to complex features of sound such as modulations of amplitude or frequency. If you pay attention to speech or music you will hear many examples of both kinds of modulation. Auditory cortex projects to numerous secondary cortical areas including multisensory areas (allow us to recognize animals or other humans by both sound and sight) and to regions specifically involved in communication. Communication and environmental sounds are separated after the AC. It is also noteworthy that the AC and MGN project to the amygdala; as we’ll see later this permits sounds to be linked with dangerous stimuli (fear conditioning for conditioned avoidance). Processing of Natural Sounds Nelken, 2004 Acoustic input typically comes from many different sources; they have different combinations of frequencies, their amplitudes change independently and they have different locations. But the sound waves coming into the ears are just the sum of all these different acoustic signals. The auditory system then separates these acoustic inputs to generate the “sounds” we hearthe “auditory scene”. This takes extensive learning early in life but the mechanisms are not understood. One major point is that the both the signal fine structure (the frequencies present) and the envelope (amplitude modulation of the individual frequencies) must be extracted and connected with their location. This process occurs in cortex.