Lovell et al.

, Anat Physiol 2012, 2:3

Anatomy & Physiology http://dx.doi.org/10.4172/2161-0940.1000106

Research Article Open Access

Scanning and Transmission Electron Microscope Examination of Cochlea

Hair and Pillar Cells from the Ear of the Mongolian Gerbil (Meriones
unguiculatus)
JM Lovell1,3*, M Brosch1, E Budinger1, J Goldschmidt1, H Scheich1, S Tschorn1, U Wendt2 and W Zuschratter1
1
Leibniz Institute for Neurobiology, Magdeburg, Germany
2
Institute of Materials and Joining Technology, Otto von Guericke University, Magdeburg, Germany
3
Deutsches Zentrum fur Neurodegenerative Erkrankungen (DZNE), Bonn, Germany

Abstract
The rodent inner ear and especially from the Mongolian Gerbil (Meriones unguiculatus) has received considerable
attention in recent years for hearing research purposes, with models employed, for example, comprehension of age
and noise related hearing loss, genetic disorders in auditory processing and the effects of ototoxic drugs. To aid both
histological and physiological investigations, topographic details of the hair cells along the organ of Corti are imaged
using the Scanning Electron Microscope (SEM) to reveal the overall length of the cochlea and to indicate the number
of hair cells present. This information is of use in, for example, surgical planning for the implantation of electrodes
and other interfaces and for pharmacological research purposes where the effects of certain drugs and other agents
on the ear can be determined pathologically.

Keywords: Cochlea; Ear; Hair cell; Mongolian Gerbil; Meriones
unguiculatus; Pillar cell
Abbreviations: am: auditory meatus; bl: bony spiral lamina; bm:
basilar membrane; bma: basilar membrane zona arcuata; bmp: basilar
membrane zona pectinata; bv: bronx waltzer mouse mutation; cb: cell
body; c: cochlea; Cx26: Connexin 26; Cx30 Connexin 30; cp: cuticular
plate; dc. Deiters’ cells; fn: foramina nervosa; Hz: hertz; IHC: Inner Hair
Cell; inner pillar cell; J: junction; kHz: kilohertz; kV: kilovolt; MRM:
Magnetic Resonance Microscopy; m: malleus; µm: micrometre; mm:
millimetre; mv: microvilli; mV: millivolt; nm: nanometre; n: nucleus;
OHC: Outer Hair Cell; opc: outer pillar cell; PC: Personal Computer;
pH: power of hydrogen; pnf: peripheral auditory nerve fibres; rm:
Reissner’s membrane; SD: Standard Deviation; SEM: Scanning Electron
Microscope; sg: Spiral ganglion; slm: spiral limbus; spo: supporting
cells for the spiral organic; st: stereocilia; sv: synaptic vesicles; TEM:
Transmission Electron Microscope; Tl+: thallium; tm: tectorial
membrane; tym. tympanic membrane; VIII: peripheral auditory
nerve; VASP: Vasodilator-stimulated phosphoprotein;
Introduction
Whilst the inner ear from M. unguiculatus has been examined
using a variety of histological and physiological methodologies, none
describe the fine structural organization of hair cells and pillar cells of
the cochlea. The inner ear is situated in the osseous labyrinth of the
auditory periotic in the tympanic part of the temporal bone, lying
below the squama and in front of the mastoid process. A series of
interconnected membranous pouches, ducts and canals are contained
within the osseous labyrinth which consists of three main parts; the
vestibule, the semicircular canals and cochlea. The cochlea is divided
into three longitudinal compartments; the scala tympani, scala media
(containing the organ of Corti) and scala vestibuli [1,2], coiled around
the modiolus (a hollow bone pillar containing the cochlea nerve).
In general, the hair cells along the organ of Corti are divided into a
single row of Inner Hair Cells (IHCs) and three rows of Outer Hair
Cells (OHCs) [2]. The position of the cells is specified during early
development and possibly follows the layout of the supporting pillar
cells [3-6]. The structure of the pillar cells is formed from actin and
myosin filaments, which are especially present in the apical region
forming part of the cuticular plate [7,8] that increase in number with
location around the basilar membrane and reticular lamina [9-11].
In addition, supporting cells have bundled microtubules containing
tubulin that provides structural support to the organ of Corti [12].
Waldeyer [13] first introduced the hypothesis of radial self-
movements within the pillar cells, and Rhode & Geisler [14] came to
a similar conclusion when mathematically modeling the displacement
between opposing points on the tectorial membrane and reticular
lamina. Schick et al. [15] found Vasodilator-stimulated phosphoprotein
(VASP) and the protein Zyxin co-localised with pan-actin, suggesting
actin based dynamics in pillar cells was possible. VASP is a crucial
factor in the regulation of actin dynamics and the formation of actin
filaments [15,16]. VASP has been found to be associated with motility
in the Outer Hair Cells (OHCs), and its expression along with F-actin
in pillar cells coincides with the onset of hearing [16-18]. The bronx
waltzer (bv) mouse mutation is an autosomal recessive mutation that
affects both the inner hair cells (IHCs) and pillar cells in the organ of
Corti and vestibular system; severer mutations lead to deafness and
vestibular dysfunction [19,20]. Until recently, pillar cells were assumed
to form a highly rigid bridge between the IHCs and OHCs [11], though
this rigidity is contrary to the findings of Gummer et al. [21], Hu et
al. [22] and Hemmert et al. [23]. In light of this, it appears that pillar
cells may be more dynamically involved in controlling the protracted
mechanical properties of the cochlea than is generally recognised [15].
During an acoustical event, the stapes in the middle ear transmits
*Corresponding author: JM Lovell, Leibniz Institute for Neurobiology,
Magdeburg, Deutsches Zentrum fur Neurodegenerative Erkrankungen (DZNE),
Bonn, Germany, Tel: +49-391-6263-32; E-mail: j.lovell@ifn-magdeburg.de
Received March 21, 2012; Accepted June 23, 2012; Published June 25, 2012
Citation: Lovell JM, Brosch M, Budinger E, Goldschmidt J, Scheich H, et al. (2012)
Scanning and Transmission Electron Microscope Examination of Cochlea Hair and
Pillar Cells from the Ear of the Mongolian Gerbil (Meriones unguiculatus). Anat
Physiol 2:106. doi:10.4172/2161-0940.1000106
Copyright: © 2012 Lovell JM, et al. This is an open-access article distributed under
the terms of the Creative Commons Attribution License, which permits unrestricted
use, distribution, and reproduction in any medium, provided the original author and
source are credited.

Anat Physiol
ISSN:2161-0940 Physiol, an open access journal Volume 2 • Issue 3 • 1000106
Citation: Lovell JM, Brosch M, Budinger E, Goldschmidt J, Scheich H, et al. (2012) Scanning and Transmission Electron Microscope Examination
of Cochlea Hair and Pillar Cells from the Ear of the Mongolian Gerbil (Meriones unguiculatus). Anat Physiol 2:106. doi:10.4172/2161-
0940.1000106
the vibrational movement of the tympanic membrane to the perilymph
in the scala vestibuli via the fenestra ovalis. The motion of perilymph
in turn vibrates the endolymph in the scala media, the perilymph in
the scala tympani, the structure of the basilar membrane and organ of
Corti, thus causing movement of the stereocilia located at the tips of the
hair cells. The kinetic deflection of the stereocilia opens mechanically
gated ion channels through the cilia membrane and generates receptor
potentials, the strength of which depend on the orientation of the
stereocilia array relative to the direction of fluid particle velocity.
The endolymph in the scala media is rich in potassium salts and has
a high positive charge from 80 to 120 mV [24] whilst the hair cells
have a negative potential of around - 50 mV. This electrochemical
gradient between the hair cells and endolymph encourages an influx
of potassium ions into the negatively charged cell body. Once the cell
membrane becomes sufficiently depolarised, the cell generates a brief
electrical signal or non-spiking receptor potential [25]. Both inner
and outer hair cells respond to sound by generating potentials [24,26].
Cochlea hair cells are also known to possess a specialised synapse
and a reserve of readily releasable vesicles [27] that allows for a rapid
and long lasting response of the cells when appropriately stimulated.
However, OHCs are generally considered to be an effector or motor cell
[28], as they receive little if any afferent innervation [29]. OHCs are also
known to display electromotile properties evoked by depolarization of
the cell [25,30] as part of a feedback system that contributes to both
frequency selectivity and hearing sensitivity, by amplifying basilar
membrane motion [28,31]. The middle ear of the Mongolian gerbil
(M. unguiculatus) is voluminous, with relatively thin bone covering
the cochlea and brain [32], facilitating studies of both the ear and
auditory cortex [33]. Interestingly, the range of frequencies audible
to M. unguiculatus is comparatively wide, and covers a bandwidth of
between 0.1 kHz to 60.0 kHz, with low frequencies being comparable
to man [34]
Various techniques have been used to reveal internal and external
hair cell structures; Spoendlin [29] and Sato et al. [35] used a light
microscope to examine the innervating nerve distributions and hair
cells in the cochlea of the cat, and Wever et al. [36] used a similar
methodology for examining the cochlea from the bottlenose dolphin
(Tursiops truncates). Richter et al. [37] imaged soft tissue in the gerbil
organ of Corti to micrometer resolution using hard X-rays. In addition,
Thorne et al. [38] used Magnetic Resonance Microscopy (MRM) to
generate three dimensional reconstructions of the endolymphatic and
perilymphatic fluid spaces in the human, guinea pig, bat, rat, mouse,
and gerbil. Lenoir et al. [39] and Ma et al. [40] used the Scanning
Electron Microscope (SEM) to study maturation of the rat cochlea,
whilst Kopke et al. [41] and Lovell et al. [42] used the SEM to study
surface detail of the hair cells in the non mammalian vertebrate
vestibule. Internal cellular structures were studied using Transmission
Electron Microscopy (TEM), in accordance with Tucker et al. [6] and
Ma et al. [40] who used the TEM to study the hair cells in the cochlea
of the mouse. Tolomeo and Holley [7] used the TEM to study the pillar
cells in the guinea pig and Souter et al. [43] used both SEM and TEM to
study postnatal maturation of the organ of Corti in gerbils. Plassmann
et al. [44] studied the cochlea of 5 different gerbil species using cochlear
microphonic recordings, serial sections and computerized quantitative
reconstructions of the cochleae and their specific morphological
structures. Tange et al. [45] used the Scanning Electron Microscope
(SEM) to quantify vancomycin-induced cochlear damage in M.
unguiculatus.
Anat Physiol
ISSN:2161-0940 Physiol, an open access journal
Page 2 of 6
Materials and Methods
SEM preparation methodology
All experiments were approved by the animal ethics committee of
the Land Sachsen-Anhalt (No. 42502-2-825 IfN MD), in accordance
with the regulations of the German Federal Law on Care and Use of
Laboratory Animals, and with the European Communities Council
Directive of 24 November 1986 (86/609/EEC). The heads from 6 mature
gerbils were removed immediately following an overdose of ketamine,
then divided medially and a section of the brainstem, auditory nerve
and associated arterial system were excised from the cranium, together
with the periotic bone containing the inner ear and placed in chilled
fixative (3.5% glutaraldehyde in Sodium Cacodylate buffer (0.1 M pH
7.2)) within around 4 minutes. After two hours in the fixative, the ears
were washed in buffer and the periotic mounted in a clear Perspex
clamp placed in Petri dish, then filled with buffer until the cochlea
was completely covered to a depth of 1mm. The Petri dish was fixed
to a rotating vice and x,y slide and mounted on a metal table bearing a
binocular microscope and Dremmel drill with a stand and micro burr
tool (tip diameter 300 µm). Encapsulating bone was carefully removed
from the continuously submerged cochlea using the micro burr, fine
forceps and a needle probe. On completion, the cochlea was dehydrated
through a graded ethanol series ranging from 35% through 50%, 70%
and 90% to absolute ethanol, prior to desiccation using the critical point
drying method used by Lovell et al. [46]. Fully desiccated end organs
were subsequently mounted on a specimen stub using either a carbon
tab or silver dag, and coated with c. 8 nm of gold in an Emitech K 550
sputter coater (working at approximately 2 x 10-6 Torr). The processed
specimens were investigated and photographed using a FEI XL30
scanning electron microscope operated at 25 kV or 2 kV, respectively
and a 15mm working distance and the images digitally stored on a PC.
TEM preparation methodology
The TEM preparation methodology employed in this study was
based on techniques used by Lovell & Harper [46]. As with the SEM
methodology, the heads of four mature male gerbils were divided
medially and the periotic bone containing the inner ears from the left
and right sides were removed and placed into chilled fixative for two
hours. Following this, the periotic bone was trimmed and placed in a
transparent micro-vice in a small dish filled with chilled buffer solution
until it covered the cochlea to a depth of approximately 1mm. Removal
of the bone encapsulating the cochlea was achieved manually under
a binocular microscope using the aforementioned SEM methodology.
Bone was removed a turn at a time and the canal cut into sections at
each turn, then freed from the modiolus by running a micro-scalpel
around the inner edge of the spiral lamina (the bony shelf projecting
partially into the cochlea canal that supports the basilar membrane).
After cutting, the canal was lifted from the periotic and placed into
chilled Sodium Cacodylate buffer. The canal was then secondary
fixed in a 1% Osmium Tetroxide in Sodium Cacodylate buffer for
one hour for the first set of samples, and omitted for the second. The
tissue was then rinsed twice in buffer, then dehydrated in an ethanol
series. Once in 100% ethanol, the tissue was placed in increasing
concentrations of Durcupan epoxy resin until it was fully infiltrated
to 100%, in accordance with Glauert [47]. The resin and samples were
then placed in a small plastic dish filled to a depth of approximately
2mm, with polymerisation achieved by placing the dish in a 70°C oven
for 48 hours. The resulting resin block was divided into smaller blocks
each containing a section of cochlea, then sectioned using a Reichert-
Jung Ultracut and a Diatome diamond knife. Ultra thin sections were
Volume 2 • Issue 3 • 1000106
Citation: Lovell JM, Brosch M, Budinger E, Goldschmidt J, Scheich H, et al. (2012) Scanning and Transmission Electron Microscope Examination
of Cochlea Hair and Pillar Cells from the Ear of the Mongolian Gerbil (Meriones unguiculatus). Anat Physiol 2:106. doi:10.4172/2161-
0940.1000106
picked up using 200 mesh thin bar copper grids or coated slot grids,
and stained with a saturated ethanol solution of Uranyl Acetate. The
fully processed images were taken using a LEO EM 912 transmission
electron microscope and analysed using the ImageJ software.
Results
General dissection
Partial removal of the bone covering the inner ear from M.
unguiculatus reveals the cochlea encased in the dense auditory
periotic bone, along with the eardrum and malleus. Further removal
of the remaining bone covering the posterior part of the inner ear
reveals the membranous labyrinth of the vestibule, which contains the
saccule, utricle and semi-circular canals; each filled with endolymph.
Six complete gerbil cochlea were imaged using the Scanning Electron
Microscope and measured along the organ of Corti from the lower
basal end to the upper tip (approximately 2.75 turns). The innervated
organ of Corti had a mean length of 11.67 mm (min 11.44mm, max
11.91 mm) with a standard deviation of 0.208.
Hair cell morphology
The rodent cochlea (in common with other mammals) has two
types of hair cell, Inner Hair Cells (IHCs) and Outer Hair Cells (OHCs),
orientated in a highly ordered arrangement toward the outer wall of the
cochlea. Inner hair cells occur in a single row around the inner edge of
the lamina, whilst the outer hair cells are arranged in a band of three
rows. Each of the OHCs bears up to 60 stereocilia arranged in a W
formed from three to four consecutively shorter rows. The approximate
hair cell count along the entire length of the organ of Corti was
calculated to be 1260 inner hair cells to 5000 outer hair cells, configured
in four highly ordered rows with a ratio of approximately 1 inner hair
Figure 1: 1a. Low magnification SEM micrograph of the cochlea at the second turn. cb. cell body, n. nucleus, slm. spiral limbus. 1b: SEM micrograph of the
stereocilia from the single row of IHCs. 1c: SEM micrograph of the stereocilia from an OHC from the second turn of the cochlea. 1d: SEM micrograph of an IHC
showing rows of stereocilia.
Anat Physiol
ISSN:2161-0940 Physiol, an open access journal
Page 3 of 6
cell to between 3 and 4 outer hair cells. Figure 1a to Figure 1d present
micrographs of the cochlea at the second and basal turns.
The IHCs in the cochlea have a cell body at the apex of the
helicotrema of approximately 37.5 µm (SD 1.284, min 35.6, max 38.8)
in length and approximately 9.1 µm (SD 1.005, min 7.6, max 10.0) in
width, and containing a nucleus with a mean diameter of 3.5 µm (SD
0.438, min 2.96, max 4.1). The cell body lies at almost 90° from the
OHCs, which are orientated almost vertically; though this angle can
vary by as much as 45° from vertical. The mean length of the cuticular
plate was 5.4 µm (SD 0.485, min 4.7, max 6.0) and positioned on the
side of the inner hair cell, so the stereocilia protrude upward to a mean
height of 10.0 µm (SD 0.642, min 8.8, max 10.5) and at a similar angle
to the cilia from the outer hair cells. IHCs from the first turn of the
cochlea are approximately 22.0 µm (SD 0.790, min 21.1, max 23.3 µm)
in length and approximately 8.3 µm (SD 0.742, min 7.311, max 9.229
µm) in width, and containing a nucleus with a mean diameter of 3.8 µm
(SD 0.316, min 3.327, max 4.207 µm). The mean length of the cuticular
plate was 4.9 µm (SD 0.689, min 4.2, max 5.8 µm), with stereocilia
protruding upward to a mean height of 5.6 µm (SD 0.407, min 5.0, max
6.2 µm). At the basal end of the second turn of the cochlea the IHCs are
approximately 20.5 µm (SD 1.374, min 18.4, max 21.8) in length and
approximately 8.2 µm (SD 1.201, min 6.1, max 9.3 µm) in width, and
containing a nucleus with a mean diameter of 2.5 µm (SD 0.383, min
2.0, max 3.0 µm). However, it should be noted that these data may not
include the maximum nucleus width at this location, as other sections
from a similar sample had a mean width of 4.9 µm (SD 0.669, min 3.4,
max 5.5 µm). The mean length of the cuticular plate was 4.2 µm (SD
0.161, min 4.0, max 4.4 µm), and stereocilia had a mean height of 2.7
µm (SD 0.283, min 2.5, max 3.2 µm). The body of the outer hair cells
(OHCs) are slightly smaller than the IHCs, being approximately 31.8
µm (SD 3.082, min 26.5, max 35.5 µm) in length and approximately 6.9
Volume 2 • Issue 3 • 1000106
Citation: Lovell JM, Brosch M, Budinger E, Goldschmidt J, Scheich H, et al. (2012) Scanning and Transmission Electron Microscope Examination
of Cochlea Hair and Pillar Cells from the Ear of the Mongolian Gerbil (Meriones unguiculatus). Anat Physiol 2:106. doi:10.4172/2161-
0940.1000106

Page 4 of 6

Figure 2: 2a TEM micrographs of the outer hair cell tip and connecting pillar cell from the second turn of the cochlea. Note; the striated myofibril like structures
below the OHC tip are the pillar cells projecting from the two proximal hair cells. Bar = 2 µm. 2b: Micrograph of the junction between the tip of an outer hair cell and
the connecting pillar cell. Bar = 0.5 µm. 2c: Micrograph of a transverse section through a pillar cell. Bar = 1 µm. 2d: Micrograph of an Outer Hair Cell from the first
half turn of the cochlea. Bar = 10 µm. 2e: Micrograph of the basal inner pillar cell (pc.) and bony spiral lamina (bl.) containing peripheral fibres of the auditory nerve
(pnf.), entering the foramina nervosa (fn.) located 10 to 15 µm below the distal end of the inner hair cell. cp. cuticular plate, J. junction, n. nucleus, st. stereocilia,
sv. synaptic vesicles.

µm (SD 0.818, min 5.9, max 8.3 µm) in width, and containing a nucleus
with a mean diameter of 4.7 µm (SD 0.746, min 3.4, max 5.6 µm) at the
apex. The uppermost portion of the cell contains the cuticular plate;
in the outer hair cells it is approximately 4.8 µm across, supporting
around 60 stereocilia and is in close proximity to the outer pillar cell.
The base of the OHCs are connected to Deiters’ cells below which lies
the terminal end of the outer pillar cells, which extend upward to the
IHCs and roof of the tunnel of Corti. Each OHC is separated from its
neighbour by a region void of any obvious structure approximately 10
µm wide, containing perilymph from the scala tympani.
Pillar Cells
Figure 2a through Figure 2e present ultra thin sections of the pillar
cells, viewed using the Transmission Electron Microscope (TEM). The
thinnest part of the pillar cell body is approximately 2 µm in thickness
and projects from the cuticular plate at the top of the OHCs (Figure 2a
and Figure 2b), running just below the epithelial surface toward the
single row of IHCs positioned approximately 25 µm from the outer hair
cells (Figure 2d).
Transverse sectioning of the pillar cell (Figure 2c) revealing thin
diameter actin and thicker diameter myosin filaments producing a
cross striated architecture (lower structure in Figure 2a). As Figure 2d
shows, the pillar cell thickens as it passes below the tip of the IHC, then
becomes thinner again as it descends to the bony spiral lamina (Figure
2e). The lamina contains peripheral fibres from the auditory nerve,
which are located between 10 to 15 µm below the distal end of the inner
hair cell. Synaptic vesicles can be clearly seen above the peripheral
fibres, which project toward the vesicles through the foramina nervosa.
The terminal end or foot of the inner pillar cell can be seen in close
proximity to the VIII nerve fibres and synaptic vesicles.
Discussion
All mammalian cochleae appear to function according to the same
basic principles; however, the effective frequency range differs between
species [48]. For example, the range of audible frequencies is about 20
Hz to 16 kHz in the human cochlea and about 100 Hz to 60 kHz in
M. unguiculatus [34]. In humans, the cochlea is coiled approximately
two and one-half turns around the modiolus, and in M. unguiculatus
it is coiled two and three-quarter turns. The diversity in the number
of spiral turns in the cochlea and variation in the basilar membrane
length between rodents can be seen in Table 1 [46]. The number of
cochlea turns and the basilar membrane length from M. unguiculatus,
(identified in this study) have also been included, along with the hearing
frequency range as defined by Ryan [34].
In this work, the structure of pillar cells is investigated, and

Anat Physiol
ISSN:2161-0940 Physiol, an open access journal Volume 2 • Issue 3 • 1000106
Citation: Lovell JM, Brosch M, Budinger E, Goldschmidt J, Scheich H, et al. (2012) Scanning and Transmission Electron Microscope Examination
of Cochlea Hair and Pillar Cells from the Ear of the Mongolian Gerbil (Meriones unguiculatus). Anat Physiol 2:106. doi:10.4172/2161-
0940.1000106

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Order and species Spiral Basilar membrane Lower and upper 2. Retzius G (1884) Das Gehörorgan der Wirbelthiere. Samson & Wallin Vols 1 &
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Guinea pig 4.25 18 19 0.045 49 Three microtubule-organizing centres collaborate in a mouse cochlear epithelial
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Chinchilla 3 18.5 0.05 33 Sci 108: 37-50.
Chinchilla langer 4. Manley GA, Köppl C (1998) Phylogenetic development of the cochlea and its
Laboratory rat 2.25 9.7 0.39 72 innervation. Curr Opin Neurobiol 8: 468-474.
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5. Tucker JB, Mogensen MM, Henderson CG, Doxsey S J, Wright M, et al. (1998)
Laboratory mouse 2 7 0.9 79
Nucleation and capture of large cell surface-associated microtubule arrays that
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6. Tucker JB, Mackie JB, Bussoli TJ, Steel KP (1999) Cytoskeletal integration in a
Table 1: The spiral turns of the cochlea and basilar membrane length with the
highly ordered sensory epithelium in the organ of Corti: response to loss of cell
range of audible frequencies from a selection of rodents (adapted from Lovell and
partners in the Bronx waltzer mouse. J Neurocytol 28: 1017-1034.
Harper [46]) and includes data for M. unguiculatus.
7. Tolomeo JA, Holley MC (1997) Mechanics of microtubule bundles in pillar cells
from the inner ear. J Biophys 73: 2241-2247.

8. Haas KF, Woodruff E 3rd, Broadie K (2007) Proteasome function is required to

maintain muscle cellular architecture. Biol Cell 99: 615-626.

9. lurato S (1961) Submicroscopic structure of the membranous labyrinth. Z Zell

Mikrosk Anat 52: 105-128.

10. Angleborg C, Engstrom H (1972) Supporting elements in the organ of Corti I.

Fibrillar structures in the supporting cells of the organ of corti of mammals. Acta
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11. Kikuchi T, Takasaka T, Tonosaki A, Katori Y, Shinkawa H (1991) Microtubules of

guinea pig cochlear epithelial cells. Acta Otolaryngol 111: 286-290.

12. Saha S, Slepecky NB (2000) Age-related changes in microtubules in the guinea

Figure 3: Schematic showing the relationship between the inner and outer
hair cells and the pillar cells from M. unguiculatus. The arrows labelled A, B
and C indicates the possible direction of movement caused by the motion of
the pillar cells. bm. basilar membrane, dc. outer phalangeal (Deiters) cells,
Hc. Hensen cells, IHC. Inner Hair Cell, ipc. inner pillar cell, iphal. inner
phalangeal cells, OHC. Outer Hair Cell, opc. outer pillar cell, pha. phalanx of
the outer phalangeal cells, sv. synaptic vesicles, VIII. auditory nerve fibres.
Note: the apical region of the outer hair cells is in contact with the and the
outer hair cells from the first row do not directly contact the outer pillar cell.
striated banding within the cell body can be clearly seen in Figure 2a.
Transverse sectioning shows this architecture has a repeating pattern of
alternating thin diameter actin and thicker diameter myosin filaments
[7,8]. The number of actin and myosin filaments within the pillar
cell increase progressively with location in the cochlea [9-11]. Outer
pillar cells express Cx26 while the inner pillar cells express Cx30; it is
considered that these proteins form pathways between cells [37], and
can be imaged using bright field, fluorescent and confocal fluorescent
techniques. Genetic mutations in Connexin 26 (Cx26) and Connexin
30 (Cx30) have been linked to prelingual deafness in humans.
It is therefore proposed here, that it is not only the outer hair cells
that are motile [25,30] but the pillar cells themselves have a structure
that may responds sympathetically with the movement of the outer
hair cells; thus pillar cell motion may resemble the movement of cilia
or flagella. Rapid movement of pillar cells would further increase hair
cell motility; in line with the findings of Waldeyer, Rhode & Geisler
and Schick et al. [13-15]. The schematic diagram in Figure 3 has been
annotated with arrows that suggest possible directions of motion
during an acoustical event.
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Anat Physiol
ISSN:2161-0940 Physiol, an open access journal Volume 2 • Issue 3 • 1000106