Sensations 1 2 3 Organic Sensations General Sensations Special Sensations Sensations important for life. Somatic or Visceral sensations Special sense is one that is localized into a specific organ. OPEN OPEN OPEN The sense of hearing Definition of hearing Reading only Hearing is the neural perception of sound energy, it is a mechanoreceptive sense. Functioning parts of the ear Each ear has three functioning parts : -External ear. -Middle ear. - Cochlea of inner ear. Physical nature of sound • Any device capable of producing disturbance of air molecules in form of regions of compression alternating with regions of rarefaction is a source of sound . The sense of hearing Character of sound Reading only ❶ Pitch ( Frequency )(number of cycles /sec) : The audible range extends in humans from (20 - 20.000 Hz) ( ز - the greatest sensitivity is from( 1000 to 4000 Hz) The ear can discriminate sounds ( تمي وتفهم الصوت ال انت سامعه - discrimination decreases above 4000 Hz. Low Frequency Sound صوت غليظ High Frequency Sound صوت حاد The sense of hearing Character of sound Reading only ❷Intensity (loudness or volume): The intensity, or loudness, of a sound depends on the amplitude of the sound waves. Within the range, the greater the amplitude, the louder the sound . - Unit of measurement : decibels (dB). - minimal sound pressure detected by the human ear (the threshold value = 0.000204 dynes/squ.cm = 0 dB ) - Sounds of intensity greater than 130 dB can permanently damage the sensitive sensory apparatus . 𝐼𝑛𝑡𝑒𝑛𝑠𝑖𝑡𝑦 𝑜𝑓 𝑠𝑜𝑢𝑛𝑑 𝑆𝑜𝑢𝑛𝑑 𝑖𝑛𝑡𝑒𝑛𝑠𝑖𝑡𝑦 𝑑𝐵 = 10 𝑙𝑜𝑔 𝐼𝑛𝑡𝑒𝑛𝑠𝑖𝑡𝑦 𝑜𝑓 𝑠𝑡𝑎𝑛𝑑𝑎𝑟𝑑 𝑠𝑜𝑢𝑛𝑑 The sense of hearing Character of sound Reading only ❸ Timbre ( quality ) of sound : - depends on its overtones : additional frequencies superimposed on the fundamental pitch or tone. - It is analyzed by high cortical centers . ◼Types of sound transmission: • Air conduction. • Bone conduction. The sense of hearing The sense of hearing Function of external ear The pinna Tympanic membrane a sound collector 1 2 The external auditory canal a. tortuous shape and secretes wax → protection against physical trauma and entry of foreign materials. 3 b. The effect of resonance (vibration) it vibrates when wave length of sound is 4 times the length of the tube. The external ear resonance adds 15-20db to the sound pressure at the tympanic membrane 3 The sense of hearing b. The effect of resonance (vibration) it vibrates when wave length of sound is 4 times the length of the tube. The external ear resonance adds 15-20db to the sound pressure at the tympanic membrane The sense of hearing Function of external ear 4 Sound localization by changes in pressure on the tympanic membrane. The pinna Tympanic membrane a sound collector 1 2 The external auditory canal a. tortuous shape and secretes wax → protection against physical trauma and entry of foreign materials. 3 b. The effect of resonance (vibration) it vibrates when wave length of sound is 4 times the length of the tube. The external ear resonance adds 15-20db to the sound pressure at the tympanic membrane The sense of hearing Function of tympanic membrane - The anatomical surface area is 62 mm2. - The physiological surface area is 55 mm2. - The tympanic membrane moves in and out in response to pressure changes produced by sound waves on its external surface so it acts as a resonator. The sense of hearing Function of middle ear The sense of hearing Function of middle ear 1 Conduction of sound from the tympanic membrane to the cochlea a- Amplification of sound intensity Two mechanical devices are built in the middle ear that increase or amplify sound pressure that arrives at the oval window Lever action of the malleus and incus b- Impedance matching by the ossicular system C- Sound protection of the round window The areal ratio of the tympanic membrane and oval window The sense of hearing More distance Less force Less distance More force The sense of hearing The sense of hearing 𝑀𝑎𝑙𝑙𝑒𝑢𝑠 𝐼𝑛𝑐𝑢𝑠 = 𝟏. 𝟑 𝟏 = 1.3 The sense of hearing Function of middle ear 1 Conduction of sound from the tympanic membrane to the cochlea a- Amplification of sound intensity Two mechanical devices are built in the middle ear that increase or amplify sound pressure that arrives at the oval window Lever action of the malleus and incus b- Impedance matching by the ossicular system C- Sound protection of the round window The areal ratio of the tympanic membrane and oval window The sense of hearing Function of middle ear The areal ratio of the tympanic membrane and oval window - The force exerted over the large tympanic membrane (55sq.mm) is concentrated on the tiny footplate of the stapes (3.2 sq.mm) with a gain in pressure equals the surface area of the larger divided by the smaller (55/ 3.2) = 17. The sense of hearing This 17 fold difference times the 1.3 fold ratio of the lever system causes about 22 times as much total force to be exerted on the fluid of the cochlea which has greater inertia than air. The sense of hearing Function of middle ear 1 Conduction of sound from the tympanic membrane to the cochlea a- Amplification of sound intensity b- Impedance matching by the ossicular system ➢ Definition: The function of the middle ear is to transfer sound energy from a gas to a liquid medium without great loss of energy. ➢ Therefore, the tympanic membrane and ossicular system provide Impedance matching between the sound waves in the air and the sound vibrations in the fluid of the cochlear fluid; impedance matching allows utilization of most of the energy of the incoming sound waves. C- Sound protection of the round window The sense of hearing Function of middle ear 1 Conduction of sound from the tympanic membrane to the cochlea a- Amplification of sound intensity b- Impedance matching by the ossicular system C- Sound protection of the round window ➢ Bulging the round window towards the middle ear due to pressure energy of sound in Scala tympani ➢ It serves as a pressure relief hole that prevents overlap between the successive waves, so it allows discrimination of sound waves inside the cochlea The sense of hearing Function of middle ear 2 Functions of middle ear muscles (Tympanic muscle reflexes, attenuation reflex): receptor: the cochlea stimulus: ▪ sound of high intensity (80 db) ▪ low frequency (< 1 KHz = 1000cyc/s). CN Efferent: through motor nuclei of the 5th and 7th cranial nerves afferent: through the cochlear nuclei → superior olivary nucleus SO IC Center: inferior colliculucs The sense of hearing Function of middle ear 2 Functions of middle ear muscles (Tympanic muscle reflexes, attenuation reflex): ➢ Effectors: ✓ the tensor tympani and stapedius muscles respectively. ➢ Effect: ✓ The tensor tympani muscle → pulls the handle of the malleus inward. ✓ The stapedius muscle → pulls the stapes outward. ✓ These two forces → cause the entire ossicular system to develop a high degree of rigidity → greatly reducing the ossicular transmission of loud sounds. This is called attenuation reflux arc. Note ➢ The reflex is concensual i.e. bilateral regardless of whether the sound stimulus is applied to one or both ears. ➢ the reflex occurs after a latent period 40-80 milliseconds. Function of this reflex It partially protects cochlea from damaging vibrations caused by very loud sound, by providing partial reduction of only 40 db. It decreases a person's hearing sensitivity to his own speech. (N.B. collateral signals are transmitted to these muscles during speech). It minimizes transmission of low frequency sounds in a loud environment. ▪ This usually removes a major share of background noise and allows the person to concentrate on sounds > 1000 Hz where most of the pertinent information in voice communication is transmitted. ▪ Also efferent fibers from olivary neurons (which are sensitive to low frequency sounds) passes to cochlea can decrease cochlear response to background noise by triggering hyperpolarization of outer hair cells through an acetylcholine mediated pathway (making effect). This effect raises auditory threshold The sense of hearing The sense of hearing The inner ear Organ of corti stria vascularis The basilar membrane The sense of hearing The sense of hearing The organ of corti (the cochlear mechano-receptive organ) ➢ Located on the basilar membrane. ➢ It contains the mechanically sensitive hair cells, which are the auditory receptors (there are 20,000 outer hair cell and 3500 inner hair cells in the human cochlea). ➢ this organ extends from the base of the cochlea and consequently has a spiral shape. The sense of hearing The organ of corti (the cochlear mechano-receptive organ) ➢ Covering the rows of hair cells is tectorial membrane in which the tips of the hairs (stereocilia) of hair cells are embedded. The sense of hearing The organ of corti (the cochlear mechano-receptive organ) ➢ Neurons of the spiral ganglion within the modiolus send its processes that arborize the bases of hair cells, these fibers transmit electrical changes generated by hair cells to the afferent ascending auditory nerve. The sense of hearing The Steria vascularis - It is a highly vascular structure surrounding the Scala media The sense of hearing The Steria vascularis ➢ Endocochlear potential: ✓ Cells of the stria vascularis have a high concentration of Na+-K+ ATPase. It continuously secretes K+ in the endolymph. ✓ This account for the fact that the Scala media is electrically positive by 80 mV relative to the Scala vestibuli and Scala tympani at rest. ✓ This difference is called endocochlear potential. It serves as a major driving force for signal transduction. The sense of hearing The Steria vascularis ▪ - Intracellular recording reveals that the hair cells have a resting potential of approximately - 40 to -60 mv. ▪ Therefore, the net potential difference across the hair cells apical membrane is 120 to 140 mv. The high electrical potential at the tips of the stereocilia increases their ability to respond to the slightest sound, they are highly excitable. The sense of hearing The basilar membrane The sense of hearing The basilar membrane contains about 25000 basilar fibers: ➢ The length of the fibers increases progressively from the base (oval and round windows) towards the apex of the cochlea. ➢ The diameter of the fibers, decreases from the base towards the apex. ➢ The thick, short fibers near the oval window will vibrate at a high frequency. ➢ The long thin fibers near the apex will vibrate at a low frequency. The sense of hearing The basilar membrane 01 Transmission of sound waves from the oval window to the organ of Corti. 02 Transduction: changing sound pressure into an action potential. 03 Frequency (Pitch) analysis: each group of fibers in the basilar membrane responds to specific frequency. 04 Intensity analysis: by degree of displacement of basilar membrane which is directly proportional to amplitude of action potential. The sense of hearing Transmission of sound waves in the cochlea (Traveling waves) The sense of hearing Transmission of sound waves in the cochlea (Traveling waves) ➢ The movement of the footplate of stapes set up a series of traveling waves in the perilymph of the Scala vestibuli. As the wave moves up the cochlea, its height increases to a maximum and then drops off rapidly. The distance from the stapes to this point of maximum height varies with the frequency of the vibrations initiating the wave: ✓ High pitched sounds: generate waves that reach maximum height near the base of the cochlea ✓ low pitched sounds: generate waves that peak near the apex. The sense of hearing Transmission of sound waves in the cochlea (Traveling waves) Mechanism of transmission of sound waves: ➢ Bony walls of the Scala vestibuli are rigid, but the Reissner's membrane is flexible. ➢ Displacements of the fluid in Scala tympani are dissipated into air at the round window. ➢ Therefore, sound produces distortion of the basilar membrane and the site at which this distortion is maximal is determined by the frequency of the sound waves. The sense of hearing Transduction of sound waves into action potential Definition The hair cells transform the mechanical forces of sound (cochlear fluid vibration) into the electrical impulses of hearing (action potentials propagating auditory messages to the cerebral cortex) خالص قربنا نخلص The sense of hearing The Auditory pathway 1 The auditory nerve is composed of 30,000 axons in humans; about 95% are myelinated and carry signals from inner hair cells. The remaining %5 are unmyelinated and transmit information from outer hair cells. These different innervation patterns suggest that inner and outer hair cells transmit different kinds of information. The large myelinated axons carry signals from a single inner hair cell so it has a private line to the CNS and arrive earlier by 1 to 2 seconds than that of outer hair cells. The unmyelinated axon transmits signals from a relatively large number of outer hair cells and shows considerable convergence. The sense of hearing 2 3 Auditory nerve (myelinated & unmyelinated fibers) courses through the internal auditory meatus to enter the brainstem. Fibers enter the dorsal and ventral cochlear nuclei (the second order neurons in upper part of the medulla) where all fibers synapse . 4 Most of the second order neurons fibers pass to the opposite side to reach the superior olivary nucleus. A few second order fibers pass to the superior olivary nucleus of the same side. The sense of hearing 5 6 From the superior olivary nucleus fibers constitute the lateral lemniscus which ends in the inferior colliculus of the midbrain. 7 The fibers then reach the medial geniculate body of the thalamus. They terminate in the primary auditory cortex (Brodmann's areas 41 and 42) in the superior gyrus of the temporal lobe The sense of hearing The sense of hearing The Auditory pathway ➢ From the primary auditory cortex the fibers then pass to the auditory association area (area 22) then to Wernicke’s area. ➢ Impulse from either ear are transmitted through the auditory pathway of both sides of the brainstem with more projection along the contralateral pathway. ➢ Many collateral branches are given off to the reticular activating system of the brainstem. ➢ The tonotopic organization present in the organ of Corti is preserved within the cochlear nuclei, the inferior colliculus and the primary auditory area The sense of hearing Function of specific parts of The Auditory pathway 1 Dorsal cochlear nucleus ➢ its neurons analyze sound frequencies (lateral neurons receive low frequency sound waves, medial neurons for high frequency waves) as arranged on basilar membrane. ➢ This spatial organization continues all the way up to the cerebral cortex. ➢ Cochlear nuclei of both sides are interconnected. 2 superior olivary nucleus Its function is determination of sound direction in space through difference in amplitude and time lag of signals arriving to the two ears + Masking effect + Sends connections to the reticular formation leads to brain arousal . The sense of hearing Function of specific parts of The Auditory pathway 3 The efferent olivary-cochlear pathway ➢ Connection to the reticular formation leads to brain arousal. ➢ Fibers descend from olivary nucleus through auditory nerve to innervate and decrease the cochlear amplifier (masking effect) by triggering hyperpolarization of the outer hair cells: ✓ this decreases the auditory response to background noises that partially prevents acoustic trauma. Also this pathway can discriminate transient sounds from background noise 4 inferior colliculus It is a center for auditory reflexes (audio-spinal reflexes) lt processes information received from previous nuclei then sends fibers to the geniculate body of thalamus. The sense of hearing Function of specific parts of The Auditory pathway 5 medial geniculate body it plays a role in processing of complex vocal communications as human speech. 6 primary auditory cortex ➢ Pitch discrimination by presence of different tonotopic maps similar to those of (cochlear nuclei, the low frequencies are presented laterally while the high frequencies are presented medially. ➢ Integration and processing of complex auditory signals. ➢ Localization of sound direction in space (by time lag neurons) ➢ Discrimination of timbre The sense of hearing Function of specific parts of The Auditory pathway 7 auditory association cortex (secondary auditory cortex) ❖ Interpretation of the meaning of the sound heard: ➢ it plays an important role in speech perception due to its connection with the language reception center; Wernicke’s area. ➢ It associates sound information with information coming from other cortical sensory areas (as visual cortex) ➢ Auditory information from subcortical structures also projects to the amygdala which is a part of limbic system so emotional responses could be evoked by sounds. The sense of hearing Auditory code 1 The place principle of pitch analysis It states that: ➢ Vibration of fibers of the basilar membrane depends upon their length and diameter, ➢ i.e. the short thick fibers at the base are maximally activated by the high frequency sounds ➢ The long thin fibers at the apex are maximally activated by the low frequency sounds and the ➢ Fibers between the apex and base respond to intermediate frequencies so each pitch (or frequency) would cause vibration of its own particular "place" on the basilar membrane. The sense of hearing ➢ There is spatial organization of the cochlear nerve fibers from the cochlea to the cochlear nuclei in the brainstem i.e. fibers from each respective area of the basilar membrane terminate in a corresponding area of cochlear nuclei. ➢ This spatial arrangement continues all the way up the brainstem to the cerebral cortex. So specific neurons are activated by specific sound frequencies i.e. there is no mixing of fibers, each area has its own fibers in the nerve pathway up to the auditory cortex The sense of hearing 2 Determination of loudness of sound ➢ It is determined by the degree of displacement of the basilar membrane. ➢ Thus increasing the amplitude stimulates the nerve endings at more rapid rates. More and more of the receptors (hair cells) become stimulated. 3 Sound localization ➢ In the horizontal plane (Require bilateral hearing). ➢ It is a function of the superior olivary nucleus and the primary auditory cortex. ➢ They detect the time lag between the entry of sound into one ear and its entry into opposite ear through the presence of time lag neurons. They also detect the difference in the intensity of sound between the two ears. Questions MCQs 1 Amplification of the sound by the middle ear serves to overcome: 01 Hyperpolariz ation of the hair cells 02 03 04 Inertia of fluid in the cochlea Inertia of air in the middle ear Hyperpolarizati on in the cochlear nerve MCQs 2 Which is TRUE about inner hair cells of organ of Corti ? 01 They are considered as true neurons. 02 03 The Hydrodynamic Calcium influx causes glutamate forces of basilar membrane have release at the no effect on hair synaptic area cells 04 Sodium influx is the first event in transduction after bending of stereocilia MCQs 3 What is the range of frequencies that the human ear is most sensitive to? 01 50 Hz to 500 Hz 02 12,000 Hz to 20,000 Hz 03 04 20 Hz to 20,000 Hz 1000 Hz to 4000 Hz MCQs 4 Sound waves are conducted from the air outside the ear to the inner ear by the processes of: 01 02 03 04 Absorption, transmission and refraction. Reflection, transmission and scattering Resonance, leverage and amplification. Resonance, diffraction and refraction. MCQs 5 Which membrane lies over the hair cells found in the organ of Corti? 01 02 03 04 Basilar Vestibular Tectorial Cochlear MCQs 6 Which of the following is the center of auditory reflexes ? 01 Medial geniculate body 02 03 04 Inferior colliculus superior olivary nucleus Dorsal cochlear nucleus MCQs 7 The thick, short fibers near the oval window will vibrate at a low frequency. 01 True 02 False MCQs 8 The physiological surface area of tympanic membrane is 55 cm2. 01 True 02 False