Sensory Systems/Auditory System
The sensory system for the sense of hearing is the auditory system. This wikibook covers the physiology of the auditory system, and its application to the most successful neurosensory prosthesis - cochlear implants. The physics and engineering of acoustics are covered in a separate wikibook, Acoustics. An excellent source of images and animations is "Journey into the world of hearing" .
The ability to hear is not found as widely in the animal kingdom as other senses like touch, taste and smell. It is restricted mainly to vertebrates and insects. Within these, mammals and birds have the most highly developed sense of hearing. The table below shows frequency ranges of humans and some selected animals:
The organ that detects sound is the ear. It acts as receiver in the process of collecting acoustic information and passing it through the nervous system into the brain. The ear includes structures for both the sense of hearing and the sense of balance. It does not only play an important role as part of the auditory system in order to receive sound but also in the sense of balance and body position.
Humans have a pair of ears placed symmetrically on both sides of the head which makes it possible to localize sound sources. The brain extracts and processes different forms of data in order to localize sound, such as:
- the shape of the sound spectrum at the tympanic membrane (eardrum)
- the difference in sound intensity between the left and the right ear
- the difference in time-of-arrival between the left and the right ear
- the difference in time-of-arrival between reflections of the ear itself (this means in other words: the shape of the pinna (pattern of folds and ridges) captures sound-waves in a way that helps localizing the sound source, especially on the vertical axis.
Healthy, young humans are able to hear sounds over a frequency range from 20 Hz to 20 kHz. We are most sensitive to frequencies between 2000 to 4000 Hz which is the frequency range of spoken words. The frequency resolution is 0.2% which means that one can distinguish between a tone of 1000 Hz and 1002 Hz. A sound at 1 kHz can be detected if it deflects the tympanic membrane (eardrum) by less than 1 Angstrom, which is less than the diameter of a hydrogen atom. This extreme sensitivity of the ear may explain why it contains the smallest bone that exists inside a human body: the stapes (stirrup). It is 0.25 to 0.33 cm long and weighs between 1.9 and 4.3 mg.
Anatomy of the Auditory System
The aim of this section is to explain the anatomy of the auditory system of humans. The chapter illustrates the composition of auditory organs in the sequence that acoustic information proceeds during sound perception.
Please note that the core information for “Sensory Organ Components” can also be found on the Wikipedia page “Auditory system”, excluding some changes like extensions and specifications made in this article. (see also: Wikipedia Auditory system)
The auditory system senses sound waves, that are changes in air pressure, and converts these changes into electrical signals. These signals can then be processed, analyzed and interpreted by the brain. For the moment, let's focus on the structure and components of the auditory system. The auditory system consists mainly of two parts:
- the ear and
- the auditory nervous system (central auditory system)
The ear is the organ where the first processing of sound occurs and where the sensory receptors are located. It consists of three parts:
- outer ear
- middle ear
- inner ear
Function: Gathering sound energy and amplification of sound pressure.
The folds of cartilage surrounding the ear canal (external auditory meatus, external acoustic meatus) are called the pinna. It is the visible part of the ear. Sound waves are reflected and attenuated when they hit the pinna, and these changes provide additional information that will help the brain determine the direction from which the sounds came. The sound waves enter the auditory canal, a deceptively simple tube. The ear canal amplifies sounds that are between 3 and 12 kHz. At the far end of the ear canal is the tympanic membrane (eardrum), which marks the beginning of the middle ear.
Function: Transmission of acoustic energy from air to the cochlea.
Sound waves traveling through the ear canal will hit the tympanic membrane (tympanum, eardrum). This wave information travels across the air-filled tympanic cavity (middle ear cavity) via a series of bones: the malleus (hammer), incus (anvil) and stapes (stirrup). These ossicles act as a lever and a teletype, converting the lower-pressure eardrum sound vibrations into higher-pressure sound vibrations at another, smaller membrane called the oval (or elliptical) window, which is one of two openings into the cochlea of the inner ear. The second opening is called round window. It allows the fluid in the cochlea to move.
The malleus articulates with the tympanic membrane via the manubrium, whereas the stapes articulates with the oval window via its footplate. Higher pressure is necessary because the inner ear beyond the oval window contains liquid rather than air. The sound is not amplified uniformly across the ossicular chain. The stapedius reflex of the middle ear muscles helps protect the inner ear from damage.
The middle ear still contains the sound information in wave form; it is converted to nerve impulses in the cochlea.
|Structural diagram of the cochlea||Cross section of the cochlea|
Function: Transformation of mechanical waves (sound) into electric signals (neural signals).
The inner ear consists of the cochlea and several non-auditory structures. The cochlea is a snail-shaped part of the inner ear. It has three fluid-filled sections: scala tympani (lower gallery), scala media (middle gallery, cochlear duct) and scala vestibuli (upper gallery). The cochlea supports a fluid wave driven by pressure across the basilar membrane separating two of the sections (scala tympani and scala media). The basilar membrane is about 3 cm long and between 0.5 to 0.04 mm wide. Reissner’s membrane (vestibular membrane) separates scala media and scala vestibuli.
Strikingly, one section, the scala media, contains an extracellular fluid similar in composition to endolymph, which is usually found inside of cells. The organ of Corti is located in this duct, and transforms mechanical waves to electric signals in neurons. The other two sections, scala tympani and scala vestibuli, are located within the bony labyrinth which is filled with fluid called perilymph. The chemical difference between the two fluids endolymph (in scala media) and perilymph (in scala tympani and scala vestibuli) is important for the function of the inner ear.
Organ of Corti
The organ of Corti forms a ribbon of sensory epithelium which runs lengthwise down the entire cochlea. The hair cells of the organ of Corti transform the fluid waves into nerve signals. The journey of a billion nerves begins with this first step; from here further processing leads to a series of auditory reactions and sensations.
Transition from ear to auditory nervous system
Hair cells are columnar cells, each with a bundle of 100-200 specialized cilia at the top, for which they are named. These cilia are the mechanosensors for hearing. The shorter ones are called stereocilia, and the longest one at the end of each haircell bundle kinocilium. The location of the kinocilium determines the on-direction, i.e. the direction of deflection inducing the maximum hair cell excitation. Lightly resting atop the longest cilia is the tectorial membrane, which moves back and forth with each cycle of sound, tilting the cilia and allowing electric current into the hair cell.
The function of hair cells is not fully established up to now. Currently, the knowledge of the function of hair cells allows to replace the cells by cochlear implants in case of hearing lost. However, more research into the function of the hair cells may someday even make it possible for the cells to be repaired. The current model is that cilia are attached to one another by “tip links”, structures which link the tips of one cilium to another. Stretching and compressing, the tip links then open an ion channel and produce the receptor potential in the hair cell. Note that a deflection of 100 nanometers already elicits 90% of the full receptor potential.
The nervous system distinguishes between nerve fibres carrying information towards the central nervous system and nerve fibres carrying the information away from it:
- Afferent neurons (also sensory or receptor neurons) carry nerve impulses from receptors (sense organs) towards the central nervous system
- Efferent neurons (also motor or effector neurons) carry nerve impulses away from the central nervous system to effectors such as muscles or glands (and also the ciliated cells of the inner ear)
Afferent neurons innervate cochlear inner hair cells, at synapses where the neurotransmitter glutamate communicates signals from the hair cells to the dendrites of the primary auditory neurons.
There are far fewer inner hair cells in the cochlea than afferent nerve fibers. The neural dendrites belong to neurons of the auditory nerve, which in turn joins the vestibular nerve to form the vestibulocochlear nerve, or cranial nerve number VIII'
Efferent projections from the brain to the cochlea also play a role in the perception of sound. Efferent synapses occur on outer hair cells and on afferent (towards the brain) dendrites under inner hair cells.
Auditory nervous system
The sound information, now re-encoded in form of electric signals, travels down the auditory nerve (acoustic nerve, vestibulocochlear nerve, VIIIth cranial nerve), through intermediate stations such as the cochlear nuclei and superior olivary complex of the brainstem and the inferior colliculus of the midbrain, being further processed at each waypoint. The information eventually reaches the thalamus, and from there it is relayed to the cortex. In the human brain, the primary auditory cortex is located in the temporal lobe.
Primary auditory cortex
The primary auditory cortex is the first region of cerebral cortex to receive auditory input.
Perception of sound is associated with the right posterior superior temporal gyrus (STG). The superior temporal gyrus contains several important structures of the brain, including Brodmann areas 41 and 42, marking the location of the primary auditory cortex, the cortical region responsible for the sensation of basic characteristics of sound such as pitch and rhythm.
The auditory association area is located within the temporal lobe of the brain, in an area called the Wernicke's area, or area 22. This area, near the lateral cerebral sulcus, is an important region for the processing of acoustic signals so that they can be distinguished as speech, music, or noise.
Auditory Signal Processing
Now that the anatomy of the auditory system has been sketched out, this topic goes deeper into the physiological processes which take place while perceiving acoustic information and converting this information into data that can be handled by the brain. Hearing starts with pressure waves hitting the auditory canal and is finally perceived by the brain. This section details the process transforming vibrations into perception.
Effect of the head
Sound waves with a wavelength shorter than the head produce a sound shadow on the ear further away from the sound source. When the wavelength is shorter than the head, diffraction of the sound leads to approximately equal sound intensities on both ears.
Sound reception at the pinna
The pinna collects sound waves in air affecting sound coming from behind and the front differently with its corrugated shape. The sound waves are reflected and attenuated or amplified. These changes will later help sound localization.
In the external auditory canal, sounds between 3 and 12 kHz - a range crucial for human communication - are amplified. It acts as resonator amplifying the incoming frequencies.
Sound conduction to the cochlea
Sound that entered the pinna in form of waves travels along the auditory canal until it reaches the beginning of the middle ear marked by the tympanic membrane (eardrum). Since the inner ear is filled with fluid, the middle ear is kind of an impedance matching device in order to solve the problem of sound energy reflection on the transition from air to the fluid. As an example, on the transition from air to water 99.9% of the incoming sound energy is reflected. This can be calculated using:
with Ir the intensity of the reflected sound, Ii the intensity of the incoming sound and Zk the wave resistance of the two media ( Zair = 414 kg m-2 s-1 and Zwater = 1.48*106 kg m-2 s-1). Three factors that contribute the impedance matching are:
- the relative size difference between tympanum and oval window
- the lever effect of the middle ear ossicles and
- the shape of the tympanum.
The longitudinal changes in air pressure of the sound-wave cause the tympanic membrane to vibrate which, in turn, makes the three chained ossicles malleus, incus and stirrup oscillate synchronously. These bones vibrate as a unit, elevating the energy from the tympanic membrane to the oval window. In addition, the energy of sound is further enhanced by the areal difference between the membrane and the stapes footplate. The middle ear acts as an impedance transformer by changing the sound energy collected by the tympanic membrane into greater force and less excursion. This mechanism facilitates transmission of sound-waves in air into vibrations of the fluid in the cochlea. The transformation results from the pistonlike in- and out-motion by the footplate of the stapes which is located in the oval window. This movement performed by the footplate sets the fluid in the cochlea into motion.
Through the stapedius muscle, the smallest muscle in the human body, the middle ear has a gating function: contracting this muscle changes the impedance of the middle ear, thus protecting the inner ear from damage through loud sounds.
Frequency analysis in the cochlea
The three fluid-filled compartements of the cochlea (scala vestibuli, scala media, scala tympani) are separated by the basilar membrane and the Reissner’s membrane. The function of the cochlea is to separate sounds according to their spectrum and transform it into a neural code. When the footplate of the stapes pushes into the perilymph of the scala vestibuli, as a consequence the membrane of Reissner bends into the scala media. This elongation of Reissner’s membrane causes the endolymph to move within the scala media and induces a displacement of the basilar membrane. The separation of the sound frequencies in the cochlea is due to the special properties of the basilar membrane. The fluid in the cochlea vibrates (due to in- and out-motion of the stapes footplate) setting the membrane in motion like a traveling wave. The wave starts at the base and progresses towards the apex of the cochlea. The transversal waves in the basilar membrane propagate with
with μ the shear modulus and ρ the density of the material. Since width and tension of the basilar membrane change, the speed of the waves propagating along the membrane changes from about 100 m/s near the oval window to 10 m/s near the apex.
There is a point along the basilar membrane where the amplitude of the wave decreases abruptly. At this point, the sound wave in the cochlear fluid produces the maximal displacement (peak amplitude) of the basilar membrane. The distance the wave travels before getting to that characteristic point depends on the frequency of the incoming sound. Therefore each point of the basilar membrane corresponds to a specific value of the stimulating frequency. A low-frequency sound travels a longer distance than a high-frequency sound before it reaches its characteristic point. Frequencies are scaled along the basilar membrane with high frequencies at the base and low frequencies at the apex of the cochlea.
Sensory transduction in the cochlea
Most everyday sounds are composed of multiple frequencies. The brain processes the distinct frequencies, not the complete sounds. Due to its inhomogeneous properties, the basilar membrane is performing an approximation to a Fourier transform. The sound is thereby split into its different frequencies, and each hair cell on the membrane corresponds to a certain frequency. The loudness of the frequencies is encoded by the firing rate of the corresponding afferent fiber. This is due to the amplitude of the traveling wave on the basilar membrane, which depends on the loudness of the incoming sound.
The sensory cells of the auditory system, known as hair cells, are located along the basilar membrane within the organ of Corti. Each organ of Corti contains about 16’000 such cells, innervated by about 30'000 afferent nerve fibers. There are two anatomically and functionally distinct types of hair cells: the inner and the outer hair cells. Along the basilar membrane these two types are arranged in one row of inner cells and three to five rows of outer cells. Most of the afferent innervation comes from the inner hair cells while most of the efferent innervation goes to the outer hair cells. The inner hair cells influence the discharge rate of the individual auditory nerve fibers that connect to these hair cells. Therefore inner hair cells transfer sound information to higher auditory nervous centers. The outer hair cells, in contrast, amplify the movement of the basilar membrane by injecting energy into the motion of the membrane and reducing frictional losses but do not contribute in transmitting sound information. The motion of the basilar membrane deflects the stereocilias (hairs on the hair cells) and causes the intracellular potentials of the hair cells to decrease (depolarization) or increase (hyperpolarization), depending on the direction of the deflection. When the stereocilias are in a resting position, there is a steady state current flowing through the channels of the cells. The movement of the stereocilias therefore modulates the current flow around that steady state current.
Lets look at the modes of action of the two different hair cell types separately:
- Inner hair cells:
The deflection of the hair-cell stereocilia opens mechanically gated ion channels that allow small, positively charged potassium ions (K+) to enter the cell and causing it to depolarize. Unlike many other electrically active cells, the hair cell itself does not fire an action potential. Instead, the influx of positive ions from the endolymph in scala media depolarizes the cell, resulting in a receptor potential. This receptor potential opens voltage gated calcium channels; calcium ions (Ca2+) then enter the cell and trigger the release of neurotransmitters at the basal end of the cell. The neurotransmitters diffuse across the narrow space between the hair cell and a nerve terminal, where they then bind to receptors and thus trigger action potentials in the nerve. In this way, neurotransmitter increases the firing rate in the VIIIth cranial nerve and the mechanical sound signal is converted into an electrical nerve signal.
The repolarization in the hair cell is done in a special manner. The perilymph in Scala tympani has a very low concentration of positive ions. The electrochemical gradient makes the positive ions flow through channels to the perilymph. (see also: Wikipedia Hair cell)
- Outer hair cells:
In humans outer hair cells, the receptor potential triggers active vibrations of the cell body. This mechanical response to electrical signals is termed somatic electromotility and drives oscillations in the cell’s length, which occur at the frequency of the incoming sound and provide mechanical feedback amplification. Outer hair cells have evolved only in mammals. Without functioning outer hair cells the sensitivity decreases by approximately 50 dB (due to greater frictional losses in the basilar membrane which would damp the motion of the membrane). They have also improved frequency selectivity (frequency discrimination), which is of particular benefit for humans, because it enables sophisticated speech and music. (see also: Wikipedia Hair cell)
With no external stimulation, auditory nerve fibres discharge action potentials in a random time sequence. This random time firing is called spontaneous activity. The spontaneous discharge rates of the fibers vary from very slow rates to rates of up to 100 per second. Fibers are placed into three groups depending on whether they fire spontaneously at high, medium or low rates. Fibers with high spontaneous rates (> 18 per second) tend to be more sensitive to sound stimulation than other fibers.
Auditory pathway of nerve impulses
So in the inner hair cells the mechanical sound signal is finally converted into electrical nerve signals. The inner hair cells are connected to auditory nerve fibres whose nuclei form the spiral ganglion. In the spiral ganglion the electrical signals (electrical spikes, action potentials) are generated and transmitted along the cochlear branch of the auditory nerve (VIIIth cranial nerve) to the cochlear nucleus in the brainstem.
From there, the auditory information is divided into at least two streams:
- Ventral Cochlear Nucleus:
One stream is the ventral cochlear nucleus which is split further into the posteroventral cochlear nucleus (PVCN) and the anteroventral cochlear nucleus (AVCN). The ventral cochlear nucleus cells project to a collection of nuclei called the superior olivary complex.
Superior olivary complex: Sound localization
The superior olivary complex - a small mass of gray substance - is believed to be involved in the localization of sounds in the azimuthal plane (i.e. their degree to the left or the right). There are two major cues to sound localization: Interaural level differences (ILD) and interaural time differences (ITD). The ILD measures differences in sound intensity between the ears. This works for high frequencies (over 1.6 kHz), where the wavelength is shorter than the distance between the ears, causing a head shadow - which means that high frequency sounds hit the averted ear with lower intensity. Lower frequency sounds don't cast a shadow, since they wrap around the head. However, due to the wavelength being larger than the distance between the ears, there is a phase difference between the sound waves entering the ears - the timing difference measured by the ITD. This works very precisely for frequencies below 800 Hz, where the ear distance is smaller than half of the wavelength. Sound localization in the median plane (front, above, back, below) is helped through the outer ear, which forms direction-selective filters.
There, the differences in time and loudness of the sound information in each ear are compared. Differences in sound intensity are processed in cells of the lateral superior olivary complexm and timing differences (runtime delays) in the medial superior olivary complex. Humans can detect timing differences between the left and right ear down to 10 μs, corresponding to a difference in sound location of about 1 deg. This comparison of sound information from both ears allows the determination of the direction where the sound came from. The superior olive is the first node where signals from both ears come together and can be compared. As a next step, the superior olivary complex sends information up to the inferior colliculus via a tract of axons called lateral lemniscus. The function of the inferior colliculus is to integrate information before sending it to the thalamus and the auditory cortex. It is interesting to know that the superior colliculus close by shows an interaction of auditory and visual stimuli.
- Dorsal Cochlear Nucleus:
The dorsal cochlear nucleus (DCN) analyzes the quality of sound and projects directly via the lateral lemnisucs to the inferior colliculus.
From the inferior colliculus the auditory information from ventral as well as dorsal cochlear nucleus proceeds to the auditory nucleus of the thalamus which is the medial geniculate nucleus. The medial geniculate nucleus further transfers information to the primary auditory cortex, the region of the human brain that is responsible for processing of auditory information, located on the temporal lobe. The primary auditory cortex is the first relay involved in the conscious perception of sound.
Primary auditory cortex and higher order auditory areas
Sound information that reaches the primary auditory cortex (Brodmann areas 41 and 42). The primary auditory cortex is the first relay involved in the conscious perception of sound. It is known to be tonotopically organized and performs the basics of hearing: pitch and volume. Depending on the nature of the sound (speech, music, noise), is further passed to higher order auditory areas. Sounds that are words are processed by Wernicke’s area (Brodmann area 22). This area is involved in understanding written and spoken language (verbal understanding). The production of sound (verbal expression) is linked to Broca’s area (Brodmann areas 44 and 45). The muscles to produce the required sound when speaking are contracted by the facial area of motor cortex which are regions of the cerebral cortex that are involved in planning, controlling and executing voluntary motor functions.