The Other Senses

Total Page:16

File Type:pdf, Size:1020Kb

The Other Senses COGS 17 Fall 2009 The Other Senses Mary ET Boyle, Ph.D. Department of Cognitive Science UCSD Peripheral Vestibular Structure: • Inner ear miniaturized accelerometers inertial guidance devices Continually reporting information about: motions and position of head and body Information goes to: brainstem cerebellum somatic sensory cortices 1 Central Vestibular Structure: • Vestibular Nuclei Directly controls motor neurons controlling: extraocular cervical postural Important for: stabilization of gaze head orientation posture during movements The Vestibular Labyrinth: • Main peripheral component Connected with cochlea Uses same specialized hair cells Transduce physical motion into neural impulses head movements inertial effects due to gravity ground-borne vibrations Vestibular endolymph (like cochlear endolymph) high in K+ and low in Na+ 2 Vestibular navigation • Translational movements are in terms of x, y, z Saccule Utricle Rotational movements roll, pitch, yaw • Roll – tumbling left or right -- move your head from your left to your right shoulder • Pitch – nod your head “yes” • Yaw – shake your head “no” Semicircular canals 3 The Vestibular Labyrinth – otolith organs • Two otolith organs - (vestibular sacs) Utricle – hair cells are located on the floor- horizontal plane Saccule – hair cells are located on the wall – vertical motion •Respond to: Information about the position of the head relative to the body utricle cochlea Vestibulocochlear nerve VIII saccule The Vestibular Labyrinth – semicircular canals • Three semicircular canals- (vestibular sacs) Oriented in three planes Ampullae – located at the base of each of the semicircular canals. •Respond to: Rotational accelerations of the head ampullae cochlea Vestibulocochlear nerve VIII 4 The Vestibular Hair cells Similar to auditory hair cells Mechanically gated transduction Channels located at the tips of the stereocilia Otolithic hair cells Scanning EM of calcium carbonate crystals (otoconia) in the utricular macula of the cat. Each crystal is about 50mm long. Lindeman, 1973 5 Otolithic neurons sense linear accelerations of the head 6 Semicircular neurons sense angular acceleration of the head 7 Adaptation Adaptation is explained in the gating spring model by adjustment of the insertion point of tips links. Movement of the insertion point up or down the shank of the stereocilium, perhaps driven by a Ca2+-dependent protein motor, can continually adjust the resting tension of the tip link. (Hudspeth and Gillespie, 1994.) components • Semicircular canals -head movements -head rotation •Vestibular sacs -position of head relative to the body 8 Vestibular Pathways • Vestibular hair cells convert information about passive head movement and active head rotation into an increase or decrease in neurotransmitter release synapse with bipolar neurons Vestibular Pathways • Cell bodies of bipolar neurons form: vestibular ganglia (receive input from vestibular hair cells) axons of the vestibular ganglia become the vestibu lar ne rve (combin e wi th cochle ar ner ve fibers to form the auditory nerve) 9 Most vestibular nerve fibers synapse with vestibular nuclilei in the mmdlledulla. 10 When vestibu lar nucle i projec t to the spina l cord and cerebe llum, they influence the coordination of balance, changes in body position, and body movement. Vestibulo-Cervical Reflex & Vestibulo-Spinal Reflex • Postural adjustments of the head & body • Descending projections 11 When they project to other areas of the medulla and to the pons, they coordinate head and eye movements (movement of the eyes to compen stsate for hdhead mmmovements) . VOR – Vestibulo-occular Reflex 12 Thalamocortical Pathways Motion Sickness • Feelings of dizziness and nausea; occur when the body is moved passively without motor activity and corresponding feedback to the brain. 13 • The vestibular system detects movements, but motor actions that could have produced the movement have not occurred (e.g., riding in a car, plane, or boat). Inconsistent information • The vestibular system senses movement inconsistent with the information about movement sensed by the eyes (e.g., spinning around with eyes closed and then stopping and opening eyes). 14 The Somatosenses • Somatosense—The skin sensations of touch, pain, temperature, and proprioception. • Proprioception—The somatosense that monitors body position and movement, acts to maintain bodyyp position, and ensures the accuracy of intended movements; located in the muscles, tendons, and joints; essential to the control of movement. Skin Receptors • The functions of the skin include protecting the internal organs from injury; • helping regulate body temperature by producing sweat, which cools the body when it becomes too hot; • and providing a first line of defense against invading microorganisms. 15 The Somatosenses: Receptors • Skin receptors • Pacinian corpuscles • Free nerve endings • Meissner’s corpuscles • Merkel’s disks • Ruffini’s corpuscles • Pacinian corpuscles—The largest of the somatosensory receptors of the skin Approximately 0.5 mm wide by 1.0 mm long Have quite large receptive fields Sensitive to touch stimulation, especially to high-frequency vibrations (200 to 300 Hz) 16 • Free nerve endings—Located just below the surface in both hairy and hairless skin detects temperature change and pain stimuli (both fast pain and slow pain) • Meissner’s corpuscle—A type of skin receptor in hairy skin located in the elevations of the dermis into the epidermis responds to pressure and low- frequency vibrations; small receptive fields 17 • Merkel’s disk—A type of skin receptor in the base of the epidermis near the sweat ducts sensitive to pressure, but not to vibrations small receptive fields • Ruffini’s corpuscle—A type of skin receptor just below the surface detects low-frequency vibrations, but not pressure large receptive fields 18 Somatosensory Pathways • Once information from the skin reaches the CNS, the neural message travels through one of three somatosensory systems: The dorsal column-medial lemniscal system The anterolateral system The spinocerebellar system • Dorsal column-medial lillemniscal system—A somatosensory pathway that begins in the spinal cord and transmits information about touch and proprioception to the primary somatosensory cortex. 19 • Anterolateral system— The stsssomatosensory pathway that begins in the spinal cord and transmits information about temperature and pain to the brain stem, reticular formation, and the primary and secondary somatosensory cortices. • Spinocerebellar system—The somatosensory pathway that begins in the spinal cord and transmits proprioceptive information to the cerebellum. 20 Locating Input on the Somatosensory System • The somatosensory system is topographically organized – adjacent places on the skin activate adjacent neurons in the primary somatosensory cortex, though the cortical organization is upside down. • Not all body parts are equall y represented. The greatest representation is for areas such as the hands, lips, and tongue, which are involved in fine tactile discrimination. 21.
Recommended publications
  • Common Vestibular Function Tests
    Common Vestibular Function Tests Authors: Barbara Susan Robinson, PT, DPT; Lisa Heusel-Gillig PT DPT NCS Fact Sheet The purpose of Vestibular Function Tests (VFTs) is to determine the health of the vestibular portion of the inner ear. These tests are commonly performed by ENTs, Audiologists, and Otolaryngologists Electronystagmography or Videonystagmography Electronystagmography (ENG test) or Videonystagmography (VNG test) evaluate the inner ear. Both record eye movements during a group of tests in light and dark rooms. During the ENG test, small electrodes are placed on the skin near the eyes to record eye movements. For the VNG test, eye movements are recorded by a video camera mounted inside of goggles that are worn during testing. ENG and VNG tests evaluate eye movements while following a visual target (tracking Produced by test) or during body and head position changes (positional test). The caloric test evaluates eye movements in response to cool or warm air (or water) placed in the ear canal. If there is no response to warm or cool air or water, ice water may be used in order to try to produce a response. The caloric test determines the difference between the function of the left and right inner ear. During this test, you may experience dizziness or nausea. You may be asked questions (math questions, city names, alphabet tasks) to distract you in order to get the best results. A Special Interest Group of Contact us: ANPT Other Common Vestibular Function Tests 5841 Cedar Lake Rd S. The rotary chair test is used along with the VNG to confirm the diagnosis and assess Ste 204 compensation of the vestibular system.
    [Show full text]
  • Anatomy of the Ear ANATOMY & Glossary of Terms
    Anatomy of the Ear ANATOMY & Glossary of Terms By Vestibular Disorders Association HEARING & ANATOMY BALANCE The human inner ear contains two divisions: the hearing (auditory) The human ear contains component—the cochlea, and a balance (vestibular) component—the two components: auditory peripheral vestibular system. Peripheral in this context refers to (cochlea) & balance a system that is outside of the central nervous system (brain and (vestibular). brainstem). The peripheral vestibular system sends information to the brain and brainstem. The vestibular system in each ear consists of a complex series of passageways and chambers within the bony skull. Within these ARTICLE passageways are tubes (semicircular canals), and sacs (a utricle and saccule), filled with a fluid called endolymph. Around the outside of the tubes and sacs is a different fluid called perilymph. Both of these fluids are of precise chemical compositions, and they are different. The mechanism that regulates the amount and composition of these fluids is 04 important to the proper functioning of the inner ear. Each of the semicircular canals is located in a different spatial plane. They are located at right angles to each other and to those in the ear on the opposite side of the head. At the base of each canal is a swelling DID THIS ARTICLE (ampulla) and within each ampulla is a sensory receptor (cupula). HELP YOU? MOVEMENT AND BALANCE SUPPORT VEDA @ VESTIBULAR.ORG With head movement in the plane or angle in which a canal is positioned, the endo-lymphatic fluid within that canal, because of inertia, lags behind. When this fluid lags behind, the sensory receptor within the canal is bent.
    [Show full text]
  • Tip-Link Integrity and Mechanical Transduction in Vertebrate Hair Cells
    Neuron, Vol. 7, 985-994, December, 1991, Copyright 0 1991 by Cell Press Tip-link Integrity and Mechanical Transduction in Vertebrate Hair Cells John A. Assad,*+ Gordon M. G. Shepherd,* This suggests that the gating springs are not rigid ele- and David P. Corey*511 ments, but can be slack-that they can pull but not *Department of Neurobiology push on the channels (Corey and Hudspeth, 1983). *Program in Neuroscience The structural correlate of this process has not been Harvard Medical School well established. A simple model has evolved from Boston, Massachusetts 02115 several independent observations. First, measure- SDepartment of Neurology ment of current flow near moving bundles indicated Massachusetts General Hospital thatthetransductionchannelsareatornearthetipsof Boston, Massachusetts 02114 the stereocilia (Hudspeth, 1982). While this has been IINeuroscience Group challenged by measurements with a Ca*+ indicator Howard Hughes Medical Institute dye (Ohmori, 1988), two additional experirnents have corroborated the localization of the channels at the tips (Huang and Corey, 1990, Biophys. Sot., abstract; Summary Jaramillo and Hudspeth, 1991). Second, the discovery of fine filaments between the tips of adjacent ster- An attractive hypothesis for hair-cell transduction is that eocilia led to the suggestion that these “tip links”were fine, filamentous “tip links” pull directly on mechani- the actual mechanical linkages to the channels (Pick- cally sensitive ion channels located at the tips of the les et al., 1984). The geometry of the bundle is such stereocilia. We tested the involvement of tip links in the that excitatory displacements would stretch the tip transduction process by treating bundles with a BAPTA- links and apply tension to the channels; inhibitory buffered, low-Ca*+ saline (1O-v M).
    [Show full text]
  • Balance and Equilibrium, I: the Vestibule and Semicircular Canals
    Anatomic Moment Balance and Equilibrium, I: The Vestibule and Semicircular Canals Joel D. Swartz, David L. Daniels, H. Ric Harnsberger, Katherine A. Shaffer, and Leighton Mark In this, our second temporal bone installment, The endolymphatic duct arises from the en- we will emphasize the vestibular portion of the dolymphatic sinus and passes through the ves- labyrinth, that relating to balance and equilib- tibular aqueduct of the osseous labyrinth to rium. Before proceeding, we must again remind emerge from an aperture along the posterior the reader of the basic structure of the labyrinth: surface of the petrous pyramid as the endolym- an inner membranous labyrinth (endolym- phatic sac. phatic) surrounded by an outer osseous laby- The utricle and saccule are together referred rinth with an interposed supportive perilym- to as the static labyrinth, because their function phatic labyrinth. We recommend perusal of the is to detect the position of the head relative to first installment before continuing if there are gravity (5–7). They each have a focal concen- any uncertainties in this regard. tration of sensory receptors (maculae) located The vestibule, the largest labyrinthine cavity, at right angles to each other and consisting of measures 4 to 6 mm maximal diameter (1–3) ciliated hair cells and tiny crystals of calcium (Figs 1–3). The medial wall of the vestibule is carbonate (otoliths) embedded in a gelatinous unique in that it contains two distinct depres- mass. These otoliths respond to gravitational sions (Fig 4). Posterosuperiorly lies the elliptical pull; therefore, changes in head position distort recess, where the utricle is anchored.
    [Show full text]
  • Pejvakin, a Candidate Stereociliary Rootlet Protein, Regulates Hair Cell Function in a Cell-Autonomous Manner
    The Journal of Neuroscience, March 29, 2017 • 37(13):3447–3464 • 3447 Neurobiology of Disease Pejvakin, a Candidate Stereociliary Rootlet Protein, Regulates Hair Cell Function in a Cell-Autonomous Manner X Marcin Kazmierczak,1* Piotr Kazmierczak,2* XAnthony W. Peng,3,4 Suzan L. Harris,1 Prahar Shah,1 Jean-Luc Puel,2 Marc Lenoir,2 XSantos J. Franco,5 and Martin Schwander1 1Department of Cell Biology and Neuroscience, Rutgers the State University of New Jersey, Piscataway, New Jersey 08854, 2Inserm U1051, Institute for Neurosciences of Montpellier, 34091, Montpellier cedex 5, France, 3Department of Otolaryngology, Head and Neck Surgery, Stanford University, Stanford, California 94305, 4Department of Physiology and Biophysics, University of Colorado School of Medicine, Aurora, Colorado 80045, and 5Department of Pediatrics, University of Colorado School of Medicine, Aurora, Colorado 80045 Mutations in the Pejvakin (PJVK) gene are thought to cause auditory neuropathy and hearing loss of cochlear origin by affecting noise-inducedperoxisomeproliferationinauditoryhaircellsandneurons.Herewedemonstratethatlossofpejvakininhaircells,butnot in neurons, causes profound hearing loss and outer hair cell degeneration in mice. Pejvakin binds to and colocalizes with the rootlet component TRIOBP at the base of stereocilia in injectoporated hair cells, a pattern that is disrupted by deafness-associated PJVK mutations. Hair cells of pejvakin-deficient mice develop normal rootlets, but hair bundle morphology and mechanotransduction are affected before the onset of hearing. Some mechanotransducing shorter row stereocilia are missing, whereas the remaining ones exhibit overextended tips and a greater variability in height and width. Unlike previous studies of Pjvk alleles with neuronal dysfunction, our findings reveal a cell-autonomous role of pejvakin in maintaining stereocilia architecture that is critical for hair cell function.
    [Show full text]
  • Acoustically Responsive Fibers in the Vestibular Nerve of the Cat
    The Journal of Neuroscience, October 1994, 74(10): 6056-6070 Acoustically Responsive Fibers in the Vestibular Nerve of the Cat Michael P. McCue1v2*a and John J. Guinan, Jr.r.2.3-4 ‘Eaton-Peabody Laboratory of Auditory Physiology, Department of Otolaryngology, Massachusetts Eye and Ear Infirmary, Boston, Massachusetts 02114, 2Harvard-MIT Division of Health Science and Technology and Research Laboratory of Electronics, and 3Department of Electrical Engineering and Computer Science, Massachusetts Institute of Technology, Cambridge, Massachusetts 02139, and 4Department of Otology and Laryngology, Harvard Medical School, Boston, Massachusetts 02115 Recordings were made from single afferent fibers in the and levels within the normal range of human hearing. We inferior vestibular nerve, which innervates the saccule and suggest a number of auditory roles that these fibers may posterior semicircular canal. A substantial portion of the fi- play in the everyday life of mammals. bers with irregular background activity increased their firing [Key words: saccule, otoliths, auditory system, mamma- in response to moderately intense clicks and tones. lian sound reception, middle-ear muscles, cochlear nucleus] In responsive fibers, acoustic clicks evoked action poten- tials with minimum latencies of I 1 .O msec. Fibers fell into The vertebrate inner ear contains several senseorgans involved two classes, with the shortest latency either to condensation in the maintenance of equilibrium and the detection of vibra- clicks (PUSH fibers) or to rarefaction clicks (PULL fibers). tion. The precise sensory role assumedby homologous organs Low-frequency (800 Hz) tone bursts at moderately high sound varies among species.For example, the sacculeis thought to act levels (>80 dB SPL) caused synchronization of spikes to asa linear accelerometerin mammals(Fernindez and Goldberg, preferred phases of the tone cycle.
    [Show full text]
  • VESTIBULAR SYSTEM (Balance/Equilibrium) the Vestibular Stimulus Is Provided by Earth's Gravity, and Head/Body Movement. Locate
    VESTIBULAR SYSTEM (Balance/Equilibrium) The vestibular stimulus is provided by Earth’s gravity, and head/body movement. Located in the labyrinths of the inner ear, in two components: 1. Vestibular sacs - gravity & head direction 2. Semicircular canals - angular acceleration (changes in the rotation of the head, not steady rotation) 1. Vestibular sacs (Otolith organs) - made of: a) Utricle (“little pouch”) b) Saccule (“little sac”) Signaling mechanism of Vestibular sacs Receptive organ located on the “floor” of Utricle and on “wall” of Saccule when head is in upright position - crystals move within gelatinous mass upon head movement; - crystals slightly bend cilia of hair cells also located within gelatinous mass; - this increases or decreases rate of action potentials in bipolar vestibular sensory neurons. Otoconia: Calcium carbonate crystals Gelatinous mass Cilia Hair cells Vestibular nerve Vestibular ganglion 2. Semicircular canals: 3 ring structures; each filled with fluid, separated by a membrane. Signaling mechanism of Semicircular canals -head movement induces movement of endolymph, but inertial resistance of endolymph slightly bends cupula (endolymph movement is initially slower than head mvmt); - cupula bending slightly moves the cilia of hair cells; - this bending changes rate of action potentials in bipolar vestibular sensory neurons; - when head movement stops: endolymph movement continues for slightly longer, again bending the cupula but in reverse direction on hair cells which changes rate of APs; - detects “acceleration”
    [Show full text]
  • Anatomic Moment
    Anatomic Moment Hearing, I: The Cochlea David L. Daniels, Joel D. Swartz, H. Ric Harnsberger, John L. Ulmer, Katherine A. Shaffer, and Leighton Mark The purpose of the ear is to transform me- cochlear recess, which lies on the medial wall of chanical energy (sound) into electric energy. the vestibule (Fig 3). As these sound waves The external ear collects and directs the sound. enter the perilymph of the scala vestibuli, they The middle ear converts the sound to fluid mo- are transmitted through the vestibular mem- tion. The inner ear, specifically the cochlea, brane into the endolymph of the cochlear duct, transforms fluid motion into electric energy. causing displacement of the basilar membrane, The cochlea is a coiled structure consisting of which stimulates the hair cell receptors of the two and three quarter turns (Figs 1 and 2). If it organ of Corti (Figs 4–7) (4, 5). It is the move- were elongated, the cochlea would be approxi- ment of hair cells that generates the electric mately 30 mm in length. The fluid-filled spaces potentials that are converted into action poten- of the cochlea are comprised of three parallel tials in the auditory nerve fibers. The basilar canals: an outer scala vestibuli (ascending spi- membrane varies in width and tension from ral), an inner scala tympani (descending spi- base to apex. As a result, different portions of ral), and the central cochlear duct (scala media) the membrane respond to different auditory fre- (1–7). The scala vestibuli and scala tympani quencies (2, 5). These perilymphatic waves are contain perilymph, a substance similar in com- transmitted via the apex of the cochlea (helico- position to cerebrospinal fluid.
    [Show full text]
  • Molecular Characterization of Hair Cell Metabolism and Tip Link Damage
    Molecular characterization of hair cell metabolism and tip link damage by Kateri J. Spinelli A dissertation in partial fulfillment of the requirements for the degree of Doctor of Philosophy Presented to the Neuroscience Graduate Program Oregon Health & Science University School of Medicine February 2012 Table of Contents List of Figures ………………………………………………………………………………………………………..…… iv AcKnowledgments ……………………………………………………………………………………………………… vii Abstract ……………………………………………………………………………………………………………………… ix Chapter 1 – Introduction ............................................................................................. 1 Chapter 2 – Distinct energy metabolism of auditory and vestibular sensory epithelia revealed by quantitative mass spectrometry using MS2 intensities ........................... 13 Abstract ........................................................................................................................ 14 Introduction .................................................................................................................. 15 Results .......................................................................................................................... 17 Label-free quantitation with MS2 intensities ............................................... 17 Quantitation of proteins in chicK auditory and vestibular epithelia ............. 18 Verifying mass spectrometry quantitation ................................................... 21 Comparing protein and mRNA abundance ..................................................
    [Show full text]
  • Physiology of the Inner Ear Balance
    § Te xt § Important Lecture § Formulas No.15 § Numbers § Doctor notes “Life Is Like Riding A § Notes and explanation Bicycle. To Keep Your Balance, You Must Keep Moving” 1 Physiology of the inner ear balance Objectives: 1. Understand the sensory apparatus of the inner ear that helps the body maintain its postural equilibrium. 2. The mechanism of the vestibular system for coordinating the position of the head and the movement of the eyes. 3. The function of semicircular canals (rotational movements, angular acceleration). 4. The function of the utricle and saccule within the vestibule (respond to changes in the position of the head with respect to gravity (linear acceleration). 5. The connection between the vestibular system and other structure (eye, cerebellum, brain stem). 2 Control of equilibrium } Equilibrium: Reflexes maintain body position at rest & movement through receptors of postural reflexes: 1. Proprioceptive system (Cutaneous sensations). 2. Visual (retinal) system. 3. Vestibular system (Non auditory membranous labyrinth1). 4. Cutaneous sensation. } Cooperating with vestibular system wich is present in the semicircular canals in the inner ear. 3 1: the explanation in the next slide. • Ampulla or crista ampullaris: are the dilations at the end of the semicircular canals and they affect the balance. • The dilations connect the semicircular canals to the cochlea utricle Labyrinth and saccule: contain the vestibular apparatus (maculla). Bony labyrinth • bony cochlea, vestibule & 3 bony semicircular canals. • Enclose the membranous labyrinth. Labyrinth a. Auditory (cochlea for hearing). b. Non-auditory for equilibrium (Vestibular apparatus). composed of two parts: • Vestibule: (Utricle and Saccule). • Semicircular canals “SCC”. Membranous labyrinth • Membranous labyrinth has sensory receptors for hearing and equilibrium • Vestibular apparatus is responsible for equilibrium 4 Macula (otolith organs) of utricle and saccule } Hair cell synapse with endings of the vestibular nerve.
    [Show full text]
  • Stiffness and Tension Gradients of the Hair Cell's Tip-Link Complex In
    RESEARCH ARTICLE Stiffness and tension gradients of the hair cell’s tip-link complex in the mammalian cochlea Me´ lanie Tobin1,2, Atitheb Chaiyasitdhi1,2†, Vincent Michel2,3,4†, Nicolas Michalski2,3,4, Pascal Martin1,2* 1Laboratoire Physico-Chimie Curie, Institut Curie, PSL Research University, CNRS UMR168, Paris, France; 2Sorbonne Universite´, Paris, France; 3Laboratoire de Ge´ne´tique et Physiologie de l’Audition, Institut Pasteur, Paris, France; 4UMRS 1120, Institut National de la Sante´ et de la Recherche Me´dicale (INSERM), Paris, France Abstract Sound analysis by the cochlea relies on frequency tuning of mechanosensory hair cells along a tonotopic axis. To clarify the underlying biophysical mechanism, we have investigated the micromechanical properties of the hair cell’s mechanoreceptive hair bundle within the apical half of the rat cochlea. We studied both inner and outer hair cells, which send nervous signals to the brain and amplify cochlear vibrations, respectively. We find that tonotopy is associated with gradients of stiffness and resting mechanical tension, with steeper gradients for outer hair cells, emphasizing the division of labor between the two hair-cell types. We demonstrate that tension in the tip links that convey force to the mechano-electrical transduction channels increases at reduced Ca2+. Finally, we reveal gradients in stiffness and tension at the level of a single tip link. We conclude that mechanical gradients of the tip-link complex may help specify the characteristic frequency of the hair cell. DOI: https://doi.org/10.7554/eLife.43473.001 *For correspondence: [email protected] †These authors contributed equally to this work Introduction The cochlea—the auditory organ of the inner ear—is endowed with a few thousands of mechanosen- Competing interests: The sory hair cells that are each tuned to detect a characteristic sound frequency (Fettiplace and Kim, authors declare that no 2014).
    [Show full text]
  • Development and Evolution of the Vestibular Sensory Apparatus of the Mammalian Ear
    Journal of Vestibular Research 15 (2005) 225–241 225 IOS Press Development and evolution of the vestibular sensory apparatus of the mammalian ear Kirk W. Beisel, Yesha Wang-Lundberg, Adel Maklad and Bernd Fritzsch ∗ Creighton University, Omaha, NE and BTNRH, Omaha, NE, USA Received 21 June 2005 Accepted 3 November 2005 Abstract. Herein, we will review molecular aspects of vestibular ear development and present them in the context of evolutionary changes and hair cell regeneration. Several genes guide the development of anterior and posterior canals. Although some of these genes are also important for horizontal canal development, this canal strongly depends on a single gene, Otx1. Otx1 also governs the segregation of saccule and utricle. Several genes are essential for otoconia and cupula formation, but protein interactions necessary to form and maintain otoconia or a cupula are not yet understood. Nerve fiber guidance to specific vestibular end- organs is predominantly mediated by diffusible neurotrophic factors that work even in the absence of differentiated hair cells. Neurotrophins, in particular Bdnf, are the most crucial attractive factor released by hair cells. If Bdnf is misexpressed, fibers can be redirected away from hair cells. Hair cell differentiation is mediated by Atoh1. However, Atoh1 may not initiate hair cell precursor formation. Resolving the role of Atoh1 in postmitotic hair cell precursors is crucial for future attempts in hair cell regeneration. Additional analyses are needed before gene therapy can help regenerate hair cells, restore otoconia, and reconnect sensory epithelia to the brain. Keywords: Ear, development, sensory epithelia, sensory neurons, otoconia, cupula 1. Introduction c) Sound reaches the cochlea where the basilar membrane motion and hair cell stimulation con- verts specific frequencies into tonotopic informa- The mammalian ear contains three sensory systems: tion that encodes place of cochlear stimulation a) Angular acceleration perception is accomplished and intensity.
    [Show full text]