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31 Olfactory Impairement in Disease and Aging
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Electron Microscopy andtheNose
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Core Messages
• Electron microscopy is a very important tool
for evaluating the ultrastructural features of
the nose and helping diagnose diseases related
to nasal structures.
• Biopsies should be small (less than 2mm) and
xed in seconds to prevent autolysis and to
obtain optimum diffusion of the xatives.
• Cilia are hairlike extensions of the apical
plasma membrane containing microtubules.
The basal body is a microtubule-organizing
center located in the apical region of the ciliated cell. The existence of the characteristic
“9+2” organization of the axonemes of the
cilia and the presence of the basal bodies are
important for the normal function of the cilia.
• The goblet cell is common in the airway epithelium. The parasympathetic nervous system
does not control release from goblet cells.
Rather, these cells respond to physical and
chemical irritants; however, mediators have
not yet been clearly identied.
• Basal cells are stem cells from which other
cell types arise. They lie on the basement
membrane and do not reach the lumen.
• The olfactory segment is the region at the roof of
the nasal cavity. In humans, the olfactory region
M. Hayran (*)
Department of Anatomy, Ekonomi University Faculty
of Medicine, İzmir, Turkey
e-mail: mtuncel@hacettepe.edu.tr
is a small area formed by a modied pseudostratied epithelium. The olfactory epithelium is
composed of olfactory receptor cells, supporting
or sustentacular cells, basal cells, and brush cells.
In contrast with the other regions of the nasal cavity, there are no goblet cells in this area.
32.1 Electron Microscopy
andtheNose
32.1.1 The Electron Microscope
The electron microscope is a type of microscope
that uses an electron beam accelerated under high
vacuum instead of light source to create an image
of the specimen. In studying the microscopic
anatomy of the nose, it is essential to understand
the requirements and capabilities of the electron
microscope and which tissues and cells should be
clearly observed by electron microscopy.
The electron microscope uses an accelerated
electron beam, emitted by a cathode and controlled by a series of electrostatic and electromagnetic lenses [1, 2]. Components of the
electron microscope are (Fig.32.1):
1. Electron optical column.
2. Electron gun that consists of an electron
source to produce electrons, such as a tungsten lament.
3. Magnetic lenses to demagnify the beam.
© The Author(s), under exclusive license to Springer Nature Switzerland AG 2023
Ö. Ö. Celebi, T. M. Önerci (eds.), Nasal Physiology and Pathophysiology of Nasal Disorders,
https://doi.org/10.1007/978-3-031-12386-3_32
419
420
TEM SEM
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M. Hayran
1
2
6
7
Fig. 32.1 Diagram of standard transmission electron
microscopy (TEM) and scanning electron microscopy
(SEM), showing major components: 1 electron optical
4. Magnetic coils to control and modify the
beam.
5. Apertures to dene the beam and prevent
electron spray.
6. Detectors to collect, detect, and display the
signal.
7. Digital imaging systems that produce an
image from the signal.
There are also vacuum systems consisting of vacuum pumps and a vacuum chamber.
There are two basic types of electron microscopy, transmission electron microscopy (TEM)
and scanning electron microscopy (SEM).
TEM uses an electron beam that transmits
through ultrathin (60–90 nm) sections that are
glutaraldehyde-xed and usually double-stained.
TEM produces two-dimensional images on a uo-
rescent screen, photographic lm, or CCD (charge-
column, 2 electron gun, 3 magnetic lenses, 4 magnetic
coil, 5 apertures, 6 detector, and 7 digital imaging
systems
coupled device) camera. TEM can detect structures
by the transmission of the electron beam and discriminate details of 0.2nm. However, the quality
of the obtained image mainly depends on the preparation of the biological sample [1].
SEM obtains topographic, three-dimensional
images with a resolution of about 2nm. The lens
system of SEM produces a small focused spot of
electrons that are then scanned over the specimen
surface by a deection coil. SEM is able to produce an image by detecting secondary electrons
and backscattered electrons generated from the
specimen. A secondary electron detector in the
SEM builds the image by mapping the signals of
a nely focused electron beam that is scanned on
the sample surface [1, 3, 4].
Both TEM and SEM can provide only blackand- white images. Although the original image
is monochrome, micrographs can be colored
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digitally to emphasize details. In recent years,
digital imaging systems provide incredible
opportunities to obtain high-quality electron
micrographs.
Biological materials usually require processing before being viewed by electron microscopy
[1, 3]. The tissue preparation technique varies
depending on the type of microscope and the type
of specimen. Low-vacuum SEMs and the environmental scanning electron microscope (ESEM)
overcome both these limitations [5].
The stages for tissue preparation for TEM are
as follows
(a) Fixation: can be achieved by perfusion and
microinjection or immersion using various
xatives including aldehydes.
(b) Post xation: performed in OsO
4.
(c) Dehydration: done with a graded series of
alcohol.
(d) Epoxy resin block preparation: treat with
propylene and embed in epoxy resin.
(e) Semi-thin sectioning: one- to two-
micrometer- thick semi-thin sections obtained
from the epoxy resin blocks should be stained
with methylene blue or azure for light
microscopy.
(f) Ultrathin sectioning: ultramicrotome, an
instrument for cutting extremely thin sections, is used for ultrathin sectioning of tissue. Ultrathin sections are obtained from
selected areas and then double-stained with
uranyl acetate/lead citrate.
During SEM sample preparation, after the xation step, the specimens must be dried. Electron
microscopists prefer to use Critical Point Drying.
By removing carbon dioxide after the transition
from the liquid to the gas phase at the critical point,
the specimen can be dried without structural damage. Specimens must be mounted onto a holder
that can be inserted into the scanning electron
microscope. The last step prior to sample imaging
is coating the samples. The objective of this coating is to increase its conductivity in the scanning
electron microscope and to prevent the buildup of
high-voltage charges on the specimen. Typically,
specimens are coated with a thin layer of gold,
gold-palladium, or platinum [3].
In addition to visual inspection, a grading system can be used to quantitatively evaluate the
samples to compare different experimental conditions. The data can then be analyzed statistically to make further evaluations. This grading
system was established based on similar principles of methods used for evaluating different tissue samples [6–10].
It is important to pay close attention during
the sample preparation to get small-sized (less
than 2mm) biopsy materials and to x them in
seconds to prevent autolysis and obtain optimum
diffusion of the xatives [3].
32.1.2 Microscopic Anatomy
oftheNose
The nose humidies, lters, and warms the air we
breathe as well as provides the sense of olfaction.
The nose is considered to have two parts: the
external nose and the nasal cavity.
The external nose consists of the skin and a
framework of compact bone and hyaline cartilage
that forms a projection covered by skin. Electron
microscopic observation of this part does not
show any regional specications. The skin consists of two main layers. The outer layer is the
epidermis, which is composed of a keratinized
and stratied squamous epithelium (Figs. 32.2
and 32.3) (see Sect. 32.1.2.1). The inner layer is
the dermis, which is dense connective tissue
including epithelial derivatives of the skin such
as hair follicles and sweat and sebaceous glands
[11]. The supporting framework is composed of
nasal bones, the frontal process of the maxillae,
and the nasal part of the frontal bone and septum,
as well as major and minor alar cartilages. Bone
is also a connective tissue characterized by a mineralized extracellular matrix containing mainly
type I collagen along with other non-collagenous
matrix proteins [1]. Bones of the external nose
consist of layers of relatively thick compact bone
with a layer of spongy bone covered by periosteum, which is a sheath of dense brous connective tissue containing osteoprogenitor cells.
The type of cartilage that contributes to the
framework of the nose is hyaline cartilage
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Fig. 32.2 The epidermis, which is composed of a keratinized stratied squamous epithelium (E), and the dermis
(D) (scale bar: 25μm). (a) Light micrograph of the epidermis of the external nose (parafn block, stain: H&E). (b)
Light micrograph of the epidermis of the external nose
(araldite block, stain: methylene blue). The specimens
were obtained from a fresh frozen cadaver from a microscopic anatomy lab at Hacettepe University, Faculty of
Medicine, Department of Anatomy. H&E hematoxylin
and eosin
(Fig.32.4). The matrix of the hyaline cartilage
consists of collagen, predominantly type II brils
and other cartilage-specic collagen molecules
[1]. The chondrocytes are either rounded or ellipsoidal (Figs.32.4 and 32.5). The plasma membrane is folded into a moderate number of
microvilli. Numerous cytoplasmic laments and
coarse granules of glycogen are prominently
present in the cytoplasm. The Golgi complex is
Fig. 32.3 Electron micrograph (TEM) keratinized stratied squamous epithelium of the epidermis of the external
nose (scale bar: 5μm) (Araldite block, stain: uranyl acetate/lead citrate). The specimen was obtained from a fresh
frozen cadaver from a microscopic anatomy lab at
Hacettepe University, Faculty of Medicine, Department of
Anatomy. TEM transmission electron microscopy, 1 stratum corneum, 2 stratum spinosum, 3 stratum granulosum,
and 4 stratum basale
also prominent and frequently contains dilated
vesicles enclosing small dense particles. A small
amount of rough endoplasmic reticulum is present, while unattached ribosomes are not numerous. There are a few lipid droplets in the
cytoplasm. The nuclei are ovoid and usually contain a single large nucleolus (Fig. 32.5).
Mitochondria are small and not very numerous.
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Fig. 32.4 The hyaline cartilage, rounded or ellipsoidal
chondrocytes (Ch), and broblast-like cells of the perichondrium (P). (a) Light micrograph (parafn block,
Fig. 32.5 Electron micrograph (TEM) of the hyaline cartilage, rounded or ellipsoidal chondrocytes (Ch) (scale
bar: 2 μm) (Araldite block, stain: uranyl acetate/lead
citrate). The specimen was obtained from a fresh frozen
cadaver at a microscopic anatomy lab at Hacettepe
University, Faculty of Medicine, Department of Anatomy.
TEM transmission electron microscopy, Arrows cytoplasmic laments, Gly coarse granules of glycogen
The matrix is composed mostly of collagen brils
and matrix granules. Frequently, granules appear
to be linked together by extremely ne intergranular brils, usually less than 50Å thick, which
connect the projections of adjacent granules.
Infrequently, clusters of membrane-bounded
matrix vesicles are observed between collagen
brils of the matrix [12].
stain: H&E) (scale bar: 100μm). (b) Light micrograph
(Araldite block, stain: methylene blue) (scale bar:
100μm). H&E hematoxylin and eosin
The perichondrium, a rmly attached dense
connective tissue composed of broblast-like
cells, surrounds the hyaline cartilage (Fig.32.4).
The nasal cavity is divided into paired chambers separated by a bony and cartilaginous septum. Each chamber is divided into three regions:
(a) Vestibule of the nasal cavity.
(b) Respiratory region.
(c) Olfactory region.
32.1.2.1 Vestibule oftheNasal Cavity
The nasal cavity extends from the nares anteriorly
to the choanae posteriorly. Just behind the nares,
the nasal cavity widens and forms the vestibule
[13]. It is lined with keratinized stratied squamous
epithelium and the dermis (Fig.32.6) that contains
connective tissue elements, many hair follicles
(hairs in this region are called vibrissae) (Figs.32.7
and 32.8), and sebaceous glands and sweat glands
(Figs.32.9 and 32.10) (see Sect. 32.1.2).
The stratied squamous epithelium consists of
several layers of cells. The attened cells form its
outer layer and the deepest cells are columnar.
The epidermis is composed of four distinct layers. These are the stratum corneum, stratum spinosum, stratum granulosum, and stratum basale
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Fig. 32.6 (a) Light micrograph of the epidermis of the
vestibule (scale bar: 5μm) (Araldite block, stain: methylene blue). (b) Electron micrograph (TEM) of the epidermis
of the vestibule (scale bar: 5μm) (Araldite block, stain: ura-
nyl acetate/lead citrate). Both specimens were obtained
from fresh frozen cadavers at a microscopic anatomy lab at
Hacettepe University, Faculty of Medicine, Department of
Anatomy. TEM transmission electron microscopy, E epidermis, D dermis, 1 stratum corneum, 2 stratum spinosum,
3 stratum granulosum and 4 stratum basale
a b
Fig. 32.7 Light micrograph from the dermis of the vestibule (scale bar: 100μm). (a) Hair follicle and vibrissae,
longitudinal section (parafn block, stain: H&E). (b) Hair
follicle and vibrissae, cross (HC) and longitudinal section
(HL) (Araldite block, stain: methylene blue) (scale bar:
100μm). The specimens were obtained from fresh frozen
cadavers at a microscopic anatomy lab at Hacettepe
University, Faculty of Medicine, Department of Anatomy.
H&E hematoxylin and eosin, E epithelial cells
ab
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Fig. 32.8 Electron micrographs (TEM) from the dermis
of the vestibule. (a) Cross and (b) longitudinal sections of
the vibrissae (scale bar: 5μm) (Araldite block, stain: uranyl acetate/lead citrate). The specimen was obtained from
a fresh frozen cadaver at a microscopic anatomy lab at
Hacettepe University, Faculty of Medicine, Department of
Anatomy. TEM transmission electron microscopy, E epithelial cells
b
a
c
Fig. 32.9 The sebaceous glands (arrows) from the dermis of the vestibule (scale bar: 5μm). (a) Light micrograph (parafn block, stain: H&E). (b) Light micrograph
(Araldite block, stain: methylene blue). (c) Electron
micrographs (TEM) (Araldite block, stain: uranyl acetate/
lead citrate). H&E hematoxylin and eosin, TEM transmission electron microscopy
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M. Hayran
a
Fig. 32.10 The sweat glands and their myoepithelial
cells (arrows) from the dermis of the vestibule. (a) Light
micrograph (parafn block, stain: H&E) (scale bar: 5μm).
(b) Electron micrographs (TEM) (Araldite block, stain:
b
uranyl acetate/lead citrate) (scale bar: 100 μm). H&E
hematoxylin and eosin, TEM transmission electron
microscopy
(stratum germinativum) (Fig.32.6). The cells of
the stratum corneum are anucleate corneal cells
(squamous), called corneocytes or cornied cells.
The corneocytes are attened cells that lack
nuclei and cytoplasmic organelles. The cells contain aggregated keratin laments. The upper spinous layer and granular cell layer also contain
smaller lamellate granules called lamellar,
membrane- coating granules (MCGs or Odland
bodies). These are numerous within the upper
spinous layer. They play an important role in providing the barrier and intercellular cohesion
functions of the stratum corneum. They release
their lipid components into the intercellular
space. The basal and spinous cells together are
called the Malpighian layer, which includes cells
such as melanocytes, Langerhans cells, and
Merkel cells. When outer cells become damaged,
cell division occurs within the basal layer. The
cells move outwards to the stratum corneum,
passing through the stratum spinosum. The characteristics of these cells then transdifferentiate to
become the cells of the stratum corneum. There
are biochemical and signaling interactions
between the epithelial cells, including desmosomes, adherens junctions, gap junctions, and
tight junctions [14].
Posteriorly, where the vestibule ends, the
stratied squamous epithelium becomes thinner
and undergoes a transition to the pseudostratied
epithelium that characterizes the respiratory
region. At this site the sebaceous glands end [15].
At the level of the limen nasi, the lining of the
nasal cavity gradually changes from squamous
epithelium to non-ciliated cuboidal or columnar
epithelium. At the level of the inferior turbinate,
the epithelium continues as pseudostratied cili-
ated columnar epithelium [16].
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32.1.2.2 Respiratory Region
oftheNasal Cavity
The mucosa of the respiratory region warms,
moistens, and lters inspired air. The lamina propria of the respiratory region has a rich, vascular
network that includes a complex set of capillary
loops. The nasal mucosa microvasculature is
composed of arterioles, venules, capillaries, and
cavernous sinuses. Both arterioles and venules
run parallel to the long axis of the nasal concha.
The capillaries and cavernous sinuses are particularly abundant and interconnect with numerous
short anastomoses to form a rich dense network
[17]. The arrangement of the vessels allows the
inhaled air to be warmed by blood ow through
the part of the capillaries closest to the surface.
These capillaries provide a mechanical heat
exchange system. Submucosal capillaries and
venules have fenestrated endothelial linings and
relatively porous basement membranes, facilitating the transit of uid and white blood cells to the
mucosal surface. The lamina propria then
becomes distended with uid, resulting in a
marked swelling of the mucous membrane and
consequent obstruction of the air passage. This
makes breathing difcult. The mucosa contains
large venous-like spaces known as swell bodies,
which may become congested during allergic
reactions or infections. The morphological view
of fenestrated endothelia might change in
response to alterations in the physiological conditions. It is essential to dene the different
shapes and courses of the muscle cells responsible for constriction and dilatation of nasal swell
bodies for proper clinical diagnoses [18].
The nasal vasculature is controlled by dense
innervations. Myelinated nerve bundles and
small axons are found in the arterial wall located
in the adventitia. Veins also have nerve structures,
but they are fewer and are found in the muscle
layer. Therefore, no axons are present in capillaries. The differences in the density of axons indicate that these vessels are controlled by neural
structures and play an important role in the swelling of the nasal mucosa [19].
Seromucous glands are one of the main components of the human nasal mucosa. Their secretion contributes to the moistening function of the
goblet cells in the respiratory epithelium. The
terminal segments of the glands are surrounded
by contractile myoepithelial cells in a basketlike
fashion. These cells, in particular, show a high
number of mitochondria [20]. This innervation
pattern is important in understanding the control
of different physiological glandular functions.
Unmyelinated nerve bers have typical neuronal
components such as neurolaments, neurotubules, and mitochondria in their cytoplasm [21].
Inhaled agents contact the nasal mucosa and
cause a local immune response. Because of the
nature of these local immune responses, nasal
mucosal antibody production is best achieved via
direct stimulation of IgA-committed, nasalassociated lymphoid tissue-derived B cells [22].
The respiratory region constitutes most of the
volume of the nasal cavities. The medial wall of
the respiratory region, the nasal septum, is
smooth, but the lateral walls contain three shelflike, bony projections called turbinates or conchae. The turbinates increase surface area to
more efciently warm inspired air. This air is also
ltered by the mucus-covered walls of the nasal
cavity. Particles trapped in this layer of mucus are
transported to the pharynx by means of coordinated sweeping movements of cilia and are subsequently swallowed. Therefore, these motile
cilia play a critical role in mucociliary clearance.
This segment is lined by a ciliated, pseudostratied columnar epithelium. The pseudostratied
respiratory epithelium actually consists of one
layer of cells, but their nuclei frequently lie at different levels, and some cells do not reach the epithelial surface. Hence, the epithelium looks
stratied even if all the cells rest on a basement
membrane located between the epithelial cells
and the loose lamina propria. Basal cells, situated
close to the basement membrane, replace the ciliated cells or the goblet cells when needed [1, 23].
The lamina propria is attached to the periosteum
of the adjacent bone. The submucosa contains
blood vessels, venous plexus, glandular elements,
sensory nerves, and immune system cells.
The ciliated, pseudostratied columnar epithelium of the respiratory region is composed of
ve cell types: ciliated columnar cells, nonciliated columnar cells, goblet cells, basal cells,
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