scholarly journals Actin filaments, stereocilia, and hair cells of the bird cochlea. I. Length, number, width, and distribution of stereocilia of each hair cell are related to the position of the hair cell on the cochlea.

1983 ◽  
Vol 96 (3) ◽  
pp. 807-821 ◽  
Author(s):  
L G Tilney ◽  
J C Saunders
Keyword(s):  
Hair Cell ◽  
Surface Area ◽  
Hair Cells ◽  
Actin Filaments ◽  
Hexagonal Lattice ◽  
Frequency Discrimination ◽  
Sensory Epithelium ◽  
Scanning Microscopy ◽  
Apical Surface ◽  
Length Width

Located on the sensory epithelium of the sickle-shaped cochlea of a 7- to 10-d-old chick are approximately 5,000 hair cells. When the apical surface of these cell is examined by scanning microscopy, we find that the length, number, width, and distribution of the stereocilia on each hair cell are predetermined. Thus, a hair cell located at the distal end of the cochlea has 50 stereocilia, the longest of which are 5.5 microns in length and 0.12 microns in width, while those at the proximal end number 300 and are maximally 1.5 microns in length and 0.2 micron in width. In fact, if we travel along the cochlea from its distal to proximal end, we see that the stereocilia on successive hair cells gradually increase in number and width, yet decrease in length. Also, if we look transversely across the cochlea where adjacent hair cells have the same length and number of stereocilia (they are the same distance from the distal end of the cochlea), we find that the stereocilia of successive hair cells become thinner and that the apical surface area of the hair cell proper, not including the stereocilia, decreases from a maximum of 80 microns2 to 15 microns2. Thus, if we are told the length of the longest stereocilium on a hair cell and the width of that stereocilium, we can pinpoint the position of that hair cell on the cochlea in two axes. Likewise, if we are told the number of stereocilia and the apical surface of a hair cell, we can pinpoint the location of that cell in two axes. The distribution of the stereocilia on the apical surface of the cell is also precisely determined. More specifically, the stereocilia are hexagonally packed and this hexagonal lattice is precisely positioned relative to the kinocilium. Because of the precision with which individual hair cells regulate the length, width, number, and distribution of their cell extensions, we have a magnificent object with which to ask questions about how actin filaments that are present within the cell are regulated. Equally interesting is that the gradient in stereociliary length, number, width, and distribution may play an important role in frequency discrimination in the cochlea. This conclusion is amplified by the information presented in the accompanying paper (Tilney, L.G., E.H. Egelman, D.J. DeRosier, and J.C. Saunders, 1983, J. Cell Biol., 96:822-834) on the packing of actin filaments in this stereocilia.

scholarly journals The actin filament content of hair cells of the bird cochlea is nearly constant even though the length, width, and number of stereocilia vary depending on the hair cell location.

1988 ◽  
Vol 107 (6) ◽  
pp. 2563-2574 ◽  
Author(s):  
L G Tilney ◽  
M S Tilney
Keyword(s):  
Hair Cell ◽  
Actin Filament ◽  
Cross Sections ◽  
Hair Cells ◽  
Actin Filaments ◽  
Membrane Surface ◽  
Scanning Electron Micrographs ◽  
Length Width ◽  
Filament Content ◽  
Filament Bundle

By direct counts off scanning electron micrographs, we determined the number of stereocilia per hair cell of the chicken cochlea as a function of the position of the hair cell on the cochlea. Micrographs of thin cross sections of stereociliary bundles located at known positions on the cochlea were enlarged and the total number of actin filaments per stereocilium was counted and recorded. By comparing the counts of filament number with measurements of actin filament bundle width of the same stereocilium, we were able to relate actin filament bundle width to filament number with an error margin (r2) of 16%. Combining this data with data already published or in the process of publication from our laboratory on the length and width of stereocilia, we were able to calculate the total length of actin filaments present in stereociliary bundles of hair cells located at a variety of positions on the cochlea. We found that stereociliary bundles of hair cells contain 80,000-98,000 micron of actin filament, i.e., the concentration of actin is constant in all hair cells with a range of values that is less than our error in measurement and/or biological variation, the greatest variation being in relating the diameters of the stereocilia to filament number. We also calculated the membrane surface needed to cover the stereocilia of hair cells located throughout the cochlea. The values (172-192 micron 2) are also constant. The implications of our observation that the total amount of actin is constant even though the length, width, and number of stereocilia per hair cell vary are discussed.

Actin filaments, stereocilia and hair cells of the bird cochlea. VI. How the number and arrangement of stereocilia are determined

Development ◽  
1992 ◽  
Vol 116 (1) ◽  
pp. 213-226 ◽  
Author(s):  
L.G. Tilney ◽  
D.A. Cotanche ◽  
M.S. Tilney
Keyword(s):  
Hair Cell ◽  
Surface Area ◽  
Hair Cells ◽  
Apical Cell ◽  
Apical Surface ◽  
Cell Cytoplasm ◽  
Cross Sectional ◽  
Cross Sectional Profile ◽  
Sectional Profile ◽  
Round Cross

Beginning in 8-day embryos, stereocilia sprout from the apical surface of hair cells apparently at random. As the embryo continues to develop, the number of stereocilia increases. By 10 1/2 days the number is approximately the same as that encountered extending from mature hair cells at the same relative positions in the adult cochlea. Surprisingly, over the next 2–3 days the number of stereocilia continues to increase so that hair cells in a 12-day embryo have 1 1/2 to 2 times as many stereocilia as in adult hair cells. In short, there is an overshoot in stereociliary number. During the same period in which stereocilia are formed (9-12 days) the apical surface of each hair cell is filled with closely packed stereocilia; thus the surface area is proportional to the number of stereocilia present per hair cell, as if these features were coupled. The staircase begins to form in a 10-day embryo, with what will be the tallest row beginning to elongate first and gradually row after row begins to elongate by incorporation of stereocilia at the foot of the staircase. Extracellular connections or tip linkages appear as the stereocilia become incorporated into the staircase. After a diminutive staircase has formed, eg. in a 12-day embryo, the remaining stereocilia located at the foot of the staircase begin to be reabsorbed, a process that occurs during the next few days. We conclude that the hair cell determines the number of stereocilia to form by filling up the available apical surface area with stereocilia and then, by cropping back those that are not stabilized by extracellular linkages, arrives at the appropriate number. Furthermore, the stereociliary pattern, which changes from having a round cross-sectional profile to a rectangular one, is generated by these same linkages which lock the stereocilia into a precise pattern. As this pattern is established, we envision that the stereocilia flow over the apical surface until frozen in place by the formation of the cuticular plate in the apical cell cytoplasm.

The Organization of Actin Filaments in the Stereocilia of the Bird Cochlea

1980 ◽  
Vol 38 ◽  
pp. 554-555
Author(s):  
E.J. Battles ◽  
D. DeRosier ◽  
J.C. Saunders ◽  
L.G. Tilney
Keyword(s):  
Hair Cell ◽  
Diffraction Pattern ◽  
Sea Urchin ◽  
Actin Filaments ◽  
Optical Diffraction ◽  
Dense Material ◽  
Apical Surface ◽  
Structural Rigidity ◽  
High Degree ◽  
Degree Of Order

Extending from the apical surface of each hair cell of the chick cochlea are from 75 to 200 microvilli or stereocllia and one true cllium, the kinocilium. The stereocllia are arranged in rows of progressively increasing length (Fig. 1). Within each tapering sterocilium is a bundle of actin filaments with over 900 filaments near the tip yet only approximately 25 at the base where filaments are enmeshed in a dense material (Fig. 1); from here some of the filaments enter the apical surface of the cell (cuticular plate) as a rootlet. Examination of longitudinal sections of the stereocilia (Fig. 2) show that the filaments are aligned parallel to each other and show considerable order. Examination of an optical diffraction pattern of this bundle (Fig. 4) reveal that the actin filaments are packed such that the crossover points of adjacent actin filaments are inregister. A prominent reflection at 125Å−1 demonstrates that the filaments are cjossbridged by a macromolecular bridge situated at an average of 125Å−1 intervals (Fig. 4) in transverse sections the filaments appear hexagonally packed although there are regions where the filaments are less ordered (Fig. 3). In images processed in the computer to remove, noise and enhance detail periodic nature of the bridge can be clearly seen (see arrows Fig. 5). This image resembles that of an actin paracrystal formed from sea urchin extract composed of bundles of actin filaments crossbridged by a second protein. Thus the actin filaments in the bird stereocilia by being cross-bridged and packed with a high degree of order and produces a structure with considerable structural rigidity. Embryos were studied at various stages in development in an attempt to determine how the stereocilia form and how does the actin packing develops. These stages will be discussed.

scholarly journals Preliminary biochemical characterization of the stereocilia and cuticular plate of hair cells of the chick cochlea.

1989 ◽  
Vol 109 (4) ◽  
pp. 1711-1723 ◽  
Author(s):  
M S Tilney ◽  
L G Tilney ◽  
R E Stephens ◽  
C Merte ◽  
D Drenckhahn ◽  
...  
Keyword(s):  
Hair Cells ◽  
Actin Filaments ◽  
Biochemical Characterization ◽  
Cell Types ◽  
Sensory Epithelium ◽  
Tectorial Membrane ◽  
Uncharacterized Protein ◽  
Proteolytic Fragment ◽  
Cuticular Plate ◽  
Filament Bundles

The sensory epithelium of the chick cochlea contains only two cell types, hair cells and supporting cells. We developed methods to rapidly dissect out the sensory epithelium and to prepare a detergent-extracted cytoskeleton. High salt treatment of the cytoskeleton leaves a "hair border", containing actin filament bundles of the stereocilia still attached to the cuticular plate. On SDS-PAGE stained with silver the intact epithelium is seen to contain a large number of bands, the most prominent of which are calbindin and actin. Detergent extraction solubilizes most of the proteins including calbindin. On immunoblots antibodies prepared against fimbrin from chicken intestinal epithelial cells cross react with the 57- and 65-kD bands present in the sensory epithelium and the cytoskeleton. It is probable that the 57-kD is a proteolytic fragment of the 65-kD protein. Preparations of stereocilia attached to the overlying tectorial membrane contain the 57- and 65-kD bands. A 400-kD band is present in the cuticular plate. By immunofluorescence, fimbrin is detected in stereocilia but not in the hair borders after salt extraction. The prominent 125 A transverse stripping pattern characteristic of the actin cross-bridges in a bundle is also absent in hair borders suggesting fimbrin as the component that gives rise to the transverse stripes. Because the actin filaments in the stereocilia of hair borders still remain as compact bundles, albeit very disordered, there must be an additional uncharacterized protein besides fimbrin that cross-links the actin filaments together.

scholarly journals A counter gradient of Activin A and follistatin instructs the timing of hair cell differentiation in the murine cochlea

eLife ◽  
2019 ◽  
Vol 8 ◽  
Author(s):  
Meenakshi Prajapati-DiNubila ◽  
Ana Benito-Gonzalez ◽  
Erin Jennifer Golden ◽  
Shuran Zhang ◽  
Angelika Doetzlhofer
Keyword(s):  
Cell Differentiation ◽  
Hair Cell ◽  
Hair Cells ◽  
Activin A ◽  
Sensory Epithelium ◽  
Outer Hair Cells ◽  
Longitudinal Gradient ◽  
Hair Cell Differentiation ◽  
Local Gradient ◽  
Molecular Signals

The mammalian auditory sensory epithelium has one of the most stereotyped cellular patterns known in vertebrates. Mechano-sensory hair cells are arranged in precise rows, with one row of inner and three rows of outer hair cells spanning the length of the spiral-shaped sensory epithelium. Aiding such precise cellular patterning, differentiation of the auditory sensory epithelium is precisely timed and follows a steep longitudinal gradient. The molecular signals that promote auditory sensory differentiation and instruct its graded pattern are largely unknown. Here, we identify Activin A and its antagonist follistatin as key regulators of hair cell differentiation and show, using mouse genetic approaches, that a local gradient of Activin A signaling within the auditory sensory epithelium times the longitudinal gradient of hair cell differentiation. Furthermore, we provide evidence that Activin-type signaling regulates a radial gradient of terminal mitosis within the auditory sensory epithelium, which constitutes a novel mechanism for limiting the number of inner hair cells being produced.

scholarly journals Interactions between actin filaments and between actin filaments and membranes in quick-frozen and deeply etched hair cells of the chick ear.

1982 ◽  
Vol 95 (1) ◽  
pp. 249-261 ◽  
Author(s):  
N Hirokawa ◽  
L G Tilney
Keyword(s):  
Hair Cells ◽  
Actin Filaments ◽  
Apical Cell ◽  
Apical Surface ◽  
Contractile Ring ◽  
Vestibular Organ ◽  
Apical Cell Membrane ◽  
Cuticular Plate ◽  
Quick Freezing ◽  
Two Populations

Replicas of the apical surface of hair cells of the inner ear (vestibular organ) were examined after quick freezing and rotary shadowing. With this technique we illustrate two previously undescribed ways in which the actin filaments in the stereocilia and in the cuticular plate are attached to the plasma membrane. First, in each stereocilium there are threadlike connectors running from the actin filament bundle to the limiting membrane. Second, many of the actin filaments in the cuticular plate are connected to the apical cell membrane by tiny branched connecting units like a "crow's foot." Where these "feet" contact the membrane there is a small swelling. These branched "feet" extend mainly from the ends of the actin filaments but some connect the lateral surfaces of the actin filaments as well. Actin filaments in the cuticular plate are also connected to each other by finer filaments, 3 nm in thickness and 74 +/- 14 nm in length. Interestingly, these 3-nm filaments (which measure 4 nm in replicas) connect actin filaments not only of the same polarity but of opposite polarities as documented by examining replicas of the cuticular plate which had been decorated with subfragment 1 (S1) of myosin. At the apicolateral margins of the cell we find two populations of actin filaments, one just beneath the tight junction as a network, the other at the level of the zonula adherens as a ring. The latter which is quite substantial is composed of actin filaments that run parallel to each other; adjacent filaments often show opposite polarities, as evidenced by S1 decoration. The filaments making up this ring are connected together by the 3-nm connectors. Because of the polarity of the filaments this ring may be a "contractile" ring; the implications of this is discussed.

scholarly journals Actin filaments, stereocilia, and hair cells of the bird cochlea. V. How the staircase pattern of stereociliary lengths is generated

1988 ◽  
Vol 106 (2) ◽  
pp. 355-365 ◽  
Author(s):  
LG Tilney ◽  
MS Tilney ◽  
DA Cotanche
Keyword(s):  
Hair Cell ◽  
Hair Cells ◽  
Actin Filaments ◽  
Developmental Time ◽  
Final Phase ◽  
Set Up

The stereocilia on each hair cell are arranged into rows of ascending height, resulting in what we refer to as a "staircase-like" profile. At the proximal end of the cochlea the length of the tallest row of stereocilia in the staircase is 1.5 micron, with the shortest row only 0.3 micron. As one proceeds towards the distal end of the cochlea the length of the stereocilia progressively increases so that at the extreme distal end the length of the tallest row of the staircase is 5.5 micron and the shortest row is 2 micron. During development hair cells form their staircases in four phases of growth separated from each other by developmental time. First, stereocilia sprout from the apical surfaces of the hair cells (8-10-d embryos). Second (10-12-d embryos), what will be the longest row of the staircase begins to elongate. As the embryo gets older successive rows of stereocilia initiate elongation. Thus the staircase is set up by the sequential initiation of elongation of stereociliary rows located at increased distances from the row that began elongation. Third (12-17-d embryos), all the stereocilia in the newly formed staircase elongate until those located on the first step of the staircase have reached the prescribed length. In the final phase (17-d embryos to hatchlings) there is a progressive cessation of elongation beginning with the shortest step and followed by taller and taller rows with the tallest step stopping last. Thus, to obtain a pattern of stereocilia in rows of increasing height what transpires are progressive go signals followed by a period when all the stereocilia grow and ending with progressive stop signals. We discuss how such a sequence could be controlled.

scholarly journals The Differentiation of Stem/Progenitor Cells Derived from the Cochlear Sensory Epithelium into Hair Cells Requires a Specific Spatial Supporting Structure of p27kip1-Positive Cells

2020 ◽  
Author(s):  
Xianren Wang ◽  
Xuemei Zhang ◽  
Di Jiang ◽  
Danqing Liu ◽  
Hongyan Jiang
Keyword(s):  
Hair Cell ◽  
Progenitor Cells ◽  
Hair Cells ◽  
Hollow Sphere ◽  
Morphological Changes ◽  
Hollow Spheres ◽  
Solid Sphere ◽  
Sensory Epithelium ◽  
Adherent Culture ◽  
Epithelial Sheets

Abstract Background: Cochlear sensory epithelium-derived progenitor cells initially give rise to compact solid/round spheres. These compact solid/round spheres then gradually convert into irregular and partially hollow spheres, which then ultimately transform into large hollow spheres. The purpose of this study was to observe the differentiation of cochlear sensory epithelium-derived progenitor cells into spheres, and determine factors necessary for their development into hair cells.Methods: Cochlear epithelial sheets from postnatal day 1 C57BL/6 mice were dissociated and sphere cells were cultured. The morphological changes of the spheres were observed, and the different types of sphere cells were examined for their ability to differentiate into hair cell-like cells.Results: Solid spheres formed first, and then gradually transformed into hollow spheres over approximately 260 hours. Adherent culture and Transwell culture assays, and immunohistochemistry staining revealed that neither solid nor hollow sphere cells alone could differentiate into mature hair cells. Solid sphere cells, however, were able to differentiate into mature hair cells when co-cultured with p27kip1-positive hollow sphere cells. Direct contact of the cells was necessary for the differentiation of the solid sphere cells into mature hair cell-like cells.Conclusions: Cochlear sensory epithelium-derived progenitor cells require specific conditions to differentiate into mature hair cells.

scholarly journals SYSTEMIC KANAMYCIN TREATMENT HAS NO EFFECT ON CELL NUMBERS IN THE MOUSE UTRICULAR MACULA

2011 ◽  
Vol 24 (2) ◽  
pp. 69 ◽  
Author(s):  
Mette Kirkegaard ◽  
Stig Å Severinsen ◽  
Lise Wogensen ◽  
Jens R Nyengaard
Keyword(s):  
Hair Cell ◽  
Hair Cells ◽  
Caspase 3 ◽  
Cell Loss ◽  
Supporting Cell ◽  
Cell Number ◽  
Sensory Epithelium ◽  
Positive Staining ◽  
Survival Times ◽  
Cell Numbers

The aim of the present study is to estimate the total number of the sensory hair cells (chalice innervated and bouton innervated) and supporting cells in the mouse utricular sensory epithelium at two different time points after systemic kanamycin treatment. Mice were given two daily subcutaneous injections of kanamycin (600 or 900 mg/kg) for 15 consecutive days and allowed to survive either 1 or 3 weeks after end of treatment. Cell numbers were estimated using a physical fractionator. Paraffin-embedded tissue was immunohistochemically stained for active caspase-3 in order to detect apoptosis. There was no change in hair cell or supporting cell number after treatment with kanamycin and the survival time had no effect. Although no positive staining for caspase-3 was seen, hair cells with swollen chalices and dark stained nuclei were observed in the sensory epithelium of the treated animals, indicating some effect of the treatment. In conclusion, the dosing regime and survival times studied here are not sufficient to induce hair cell loss in the mouse utricle.

scholarly journals Conserved and Divergent Principles of Planar Polarity Revealed by Hair Cell Development and Function

2021 ◽  
Vol 15 ◽  
Author(s):  
Michael R. Deans
Keyword(s):  
Hair Cell ◽  
Inner Ear ◽  
Hair Cells ◽  
Model Organism ◽  
Vestibular Function ◽  
Sensory Receptor ◽  
Apical Surface ◽  
Planar Polarity ◽  
Vestibular Hair Cells ◽  
Sensory Epithelia

Planar polarity describes the organization and orientation of polarized cells or cellular structures within the plane of an epithelium. The sensory receptor hair cells of the vertebrate inner ear have been recognized as a preeminent vertebrate model system for studying planar polarity and its development. This is principally because planar polarity in the inner ear is structurally and molecularly apparent and therefore easy to visualize. Inner ear planar polarity is also functionally significant because hair cells are mechanosensors stimulated by sound or motion and planar polarity underlies the mechanosensory mechanism, thereby facilitating the auditory and vestibular functions of the ear. Structurally, hair cell planar polarity is evident in the organization of a polarized bundle of actin-based protrusions from the apical surface called stereocilia that is necessary for mechanosensation and when stereociliary bundle is disrupted auditory and vestibular behavioral deficits emerge. Hair cells are distributed between six sensory epithelia within the inner ear that have evolved unique patterns of planar polarity that facilitate auditory or vestibular function. Thus, specialized adaptations of planar polarity have occurred that distinguish auditory and vestibular hair cells and will be described throughout this review. There are also three levels of planar polarity organization that can be visualized within the vertebrate inner ear. These are the intrinsic polarity of individual hair cells, the planar cell polarity or coordinated orientation of cells within the epithelia, and planar bipolarity; an organization unique to a subset of vestibular hair cells in which the stereociliary bundles are oriented in opposite directions but remain aligned along a common polarity axis. The inner ear with its complement of auditory and vestibular sensory epithelia allows these levels, and the inter-relationships between them, to be studied using a single model organism. The purpose of this review is to introduce the functional significance of planar polarity in the auditory and vestibular systems and our contemporary understanding of the developmental mechanisms associated with organizing planar polarity at these three cellular levels.

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