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1 Structure and functions of skin


and coat
The skin and its adnexae make up the most important organ of the body and form an essential
anatomical and physiological harrier between the external environment and the internal organism. Knowledge of
the anatomy and histology of the various cutaneous structures, some of which are specitic to cats, allows better
understanding of all the functions of the skin in the cat. The skin also acts as a mirror which reflects the health and
comct functioning of the organism.

Structure of the skin and its adnexae


The skin of cats is thinner than that of dogs, 0.4 to 2 mm thick on average. It is thicker on the dorsal regions
and proximal limbs, and thinner on the ventrum and distal limbs '.

Epidermis
The epidermis is a squamous keratinised epithelium made up of 3 to 5 cellular layers (excluding the horny
layer), 25 pm thick ' (Diagram 1 : 1) (Fig. 1 : 1). In the non-hairy areas (nasal planurn, footpads), it is
thicker (up to 900 p)and lined by a more developed horny layer (Fig. 1 : 2). The thickness of the horny
layer is between 3 and 20 p in the hairy areas and between 15 and 35 pm on the footpads '. The
epidermis does not contain any blood vessels, its nutrients being provided by diffusion fmm the dermal
hlocd supply.
Keratinocytes make up the majority (85-90%) of the epidermal cells. Other epidermal cells, such as
Langerhans' cells (3-8%), melanocytes (2-5%) and Merkel cells, appear as clear cells under the light
microscope. Lymphocytes and occasionally mast cells can also be found ' (Fig. 1 : 3).
Keratinocytes undergo keratinisation or comification, a differentiation and maturation process which
progressively transforms the small, round, basal layer cells into large, flat, anuclear, polyhedral
corneocytes (Diagram 1 : 1). The corneocytes are shed by desquamation. The replacement rate of
keratinocytes has not yet been determined in cats. In dogs (Beagles), it takes 22 days for a basal cell to
reach the horny layer '.

Diagram 1 : 1 : Schematic diagmm of the epidermis

Horny layer

Granular layel

Spinous (Malpighian) layer

Basal layer
Basement membrane
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Melanocytes are dendritic cells which synthesise melanin pigments in specific structures called
melanosomes, situated in the epidermal basal layer or in the hair follicle matrix. They generally appear
as clear cells, without specific colouration, under the light microscope. In black or grey cats, these
melanocytes are easily recognisahle due to their intracytoplasmic black p u l e s . The melanocytes, via
their dendrites, are in contact with several keratinocytes to which they transfer their pigment granules.
There are up to 25 keratinocytes per melanocyte in the epidermis.
Langerhans' cells are antigen-presenting cells which phagocytose antigens at the skin surface, and
migrate into the drainmg lymph node where they present the captured antigen to T lymphocytes. They are
responsible for triggering specific immune reactions. Langerhans' cells contain specific intracytoplasmic
organelles, shaped like rackets, called Birbeck p u l e s 4. Their immunophenotypes are CDlat, M H C P
and C D ~ +In. cats, the density of Langerhans' cells in the skin is higher on the p m a e and middle of the
back, and lower on the abdomen I.
Merkel cells are mechanoreceptor cells situated in the basal layer, in contact with nerve fibres.

Basement membrane
The keratinocytes of the basal layer sit on the basement membrane which separates the epidermis from
the dermis. They are attached to it by hemidesmosomes and other adhesion molecules. The basement
membrane acts as mechanical supporl for the epidermis and regulates metabolic transfer between the
dermis and epidermis. It is around 40 nm thick. It can be seen under the light microscope usimg special
stains such as Periodic Acid-Schiff (PAS). Ultrastructural studies have enabled three zones to be
disimguished: the lamina lucida, in contact with cells of the basal layer, the lamina densa and the
sublamina densa which permits adhesion to dermal collagen.

Dermis
The dermis is a rich network of fibres, intercellular ground substance, blood and lymphatic vessels,
nerves, muscles and cells.
Collagenfibres produced by fibroblasts are made up of 90% collagen, a filamentous protein with great
tensile strength.
Elastinfibres make up around 4% of dermal fibres and form a network in the dermis beneath the pilo-
sebaceous units.
Reticulinfibres are very h e fibres which form a loose network around collagen fibres and other dermal
structures.
Dermal ground substance is made up of proteoglycans, glycoproteins and large quantities of water. It
appears as an amorphous gel, produced by fibroblasts, and has an important barrier function with regard
to micro-organisms and large molecules passing between the epidermis and subcutaneous tissue.
Three interconnected vascularplen'are found in the dermis. The uppermost sub-epidermal plexus brings
nutrients to the epidermis and hair follicle infundibula. The middle plexus, situated at the level of the
sebaceous glands, provides a blood supply to the glands, muscles and follicular isthmus. The deep plexus
below the hair follicles, supplies the dermal papilla, apocrine glands and the two other plexi.
Arteriovenous shunts, situated mainly in the distal limbs, are associated with thermoregulation.
Lymph vessels in the deep dermis enable drainage of cutaneous fluids and maintain homeostasis.
Nervefibres follow the blood capillaries and are organised in three plexi. Free nerve endings reach the
epidermis or form more complex structures such as Pacinian corpuscles..
Dermal cells are numerous. Fibroblasts responsible for collagen synthesis and production of dermal
ground substance also produce enzymes, notably a collagenase and a gelatinase, capable of catabolising
ground substance and fibres. The fibroblasts are therefore responsible for maintaining the dynamic
homeostasis of the dermis. They also play an important role in inflammation and healing. The numerous
mast cells in cat skin are situated mainly in the perivascular region and synthesise a range of enzymes
(chymase, tryptase) (Fig. 1 : 3). Other cells found in the dermis include macrophages, lymphocytes,
neutrophils, eosinophils and plasma cells.
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1 : Strucw and functions of sldn and coat

Figure I :I : Normal cot epidermis: note thinness of the epidermis Figure 1 :2: Normalfoofpadepidermis: note thickness of the horny layer
(HE,x 402) (HE,x l w

Figure I :3 :Intraepidermal mast cells, (+) (Tolu~dlneblue, x 100) Figure 1 :4 : I n ~ u l w ofn a normal hairfoNicle (HE,x 460)

Pigave 1 :5 :Hairfoll~clebulb in nnagenphose(HE,x400) Fkure I : 6 : Tr~ctwgmmof central pnmmy, lateral primnry and
secondary h r s (x 100)
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Subcutaneous connective tissue


Subcutaneous tissue is composed of a layer of connective tissue, rich in adipose cells. It has many roles,
the most important of which are storage of lipids and fat-soluble substances, thermal insulation and
protection from physical trauma.

Hair follicles
Hair follicles are epidermal invagimations into the dermis, which synthesise and provide support for the
hair. Hair follicles are divided into three regions (Diagram 1 :2): the infundibulum (Fig. 1 :4), the isthmus
and the bulb (Fig. 1 : 5).
In the adult cat, each follicle has its own bulb and isthmus but several follicles share the same
infundibulum. These are called compound hair follicles. Ten to twenty hairs are present per infundibulum.
The density of the coat is around 250 hairs/cmi. The coat is denser on the abdomen than on the back.
Newly-born kittens up to the age of 3-4 months have simple hair follicles.
Types of hair
Hair follicles generally contain one central primary hair, up to five lateral primary hairs and up to twenty
secondaq hairs. The clusters of follicles on the back have larger primary hairs and fewer secondaq hairs
than those on the abdomen.
Central primary hairs (40 to 80 p in diameter) have a large medulla and thin cortex ' (Fig. 1 : 6). They
are rigid, cover the whole skin surface, provide protection against the rain and are responsible for coat
colour. The primary hair follicles are the largest and longest in the follicular cluster, extending into the
deep dermis. They are associated with sebaceous glands, apocrine sweat glands and an m c t o r pili
muscle.
Lateralprimary hairs (25 to 40 pm in diameter) are moderately supple '. They have a different orientation
to the undercoat, therefore providing better thermal insulation. They are characterised by a sub-apical
bulge at the base of the hair.
Secondary hairs (10 to 20 pm in diameter) are fine and supple with a relatively thin medulla and a thick
cortex (Fig, 1 : 6). They form the undercoat responsible for maintaining temperature. Secondary
follicles can sometimes have a single sebaceous gland, but never apocrine sweat glands nor m c t o r
muscles.
Hair shrcture
The hair consists of a colunm of comified, vety adherent, stratified cells, arranged in a cuticle, a cortex
and a medulla (Diagram 1 : 2).
The cuticle is made up of a single layer of cuboidal epithelial cells which dierentiate into anuclear, flat
and adherent comeocytes. These comeocytes line and protect the hair, like tiles on a roof, and are
orientated towards the isthmus. The cuticle cells are oriented in the opposite direction to those of the inner
follicular root sheath and are therefore overlapping. This protects the hair follicle and ensures support for
the hair during its growth phase in the deep p a s of the hair follicle. In cats, the cuticle cells are thin and
arranged in a very smooth flattened manner, which is why cat hair feels softer to the touch than dog hair.
The cortex consists of elongated comified cells arranged parallel to the axis of the hair. In primary hairs,
the cortex makes up a sixth of the diameter of the hair 3. Secondaq hairs have a relatively thicker cortex
than the primary hairs which have a much larger medulla.
The medulla is the internal part of the hair. It is produced by the hair matrix and generally contains air,
glycogen vacuoles or pigments. The medullary cells of cats are flatter than those of dogs and arranged at
a 90" angle to the axis of the hair 9.
Structure of the hair follicle
The base of the hair follicle is made up of a bulb matrix, a dermal papilla and a hair bulb. Above the bulb
are the inner and outer epithelial root sheaths which are surrounded and supported by a layer of dense
connective tissue (Diagram 1 : 2).
The bulb matrix is formed from small, round, basophilic epithelial cells which divide actively at the centre
to create the hair, and at the outside to create the inner root sheath. Melanocytes ate responsible for
pigmentation by transfer of melanosomes to the cells of the hair's cortex and medulla. The bulb matrix is
nourished by the dermal papilla, made up of connective tissue rich in blood vessels and nemes within a
mucopolysaccharide matrix.
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1:Smcture and functlons of skin and coat

F w e I :7 :Xar fdllirle In anugen p b e (HE,&Oil] P i g m 1 :8 :Holrfollicie inmagenirelogenphme (HE,x 400)

F i g m I :9 : Sebaewur $ I d s ( H E , 4WJ

Fipre 1 :11 :Pac~nrancorpuscle (PAS, x IW) Figure I :12 :Whrsker note the blood slnur a m m i the h l r (HE,x 100)
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Diagram 1:2 : Schematic diagram of a hair follicle


Hair
Cuticle
Medulla
Isthmus

Inner root sheath

Outer root sheath

Apocrine sweat gland \


Dermal papilla

At the base of the follicle above the bulb matrix, the hair is surrounded by two concentric sheaths. The
outer root sheath represents an invagimation of the superficial epidermis into the dermis. In the deeper part
of the isthmus and in the bulbal zone, the outer root sheath is not cornifid. In the isthmus and upper
bulbal zone, the cells of the outer root sheath appear larger and clearer due to high levels of glycogen. In
the deeper parts of the bulb, they appear small and basophilic, like the mahix cells. The inner root sheath
swounds and supports the hair, from the matrix where it is produced, up to the isthmus where it is
destroyed.
Follicular cycle
Hair growth consists of diierent phases: a growth (anagen) phase (Fig. 1 : 7), an intermediate (catagen)
phase (Fig. 1.8) and a resting (telogen) phase. Duration of the anagen phase depends on breed and region
of skin. Long-haired cats have a longer anagen phase than short-haired cats. Dorsal hair has a longer
anagen phase than nasal hair. Hairs grow from 0.04 to 1 mm each day and long hairs grow faster than
short hairs.
The genetics of different coat types
Hair length is influenced by various genes, a dominant, L "short hair" gene (average 4.5 cm) and a
recessive, I "long hair" gene (up to 13 cm). Other lengths (very long as in the Persian, mid-length as in
the Norwegian or the Maine Coon) are produced by polygenes or modifier genes.
Genes of the series "I", "h" and " W engender important modifications to the texture and structure of
hairs. The "I" series is seen in "Rex" cats which do not have central primary hairs, and whose lateral
primary and secondary hairs and whiskers are abnormal. There are different types of Rex cats (Cornish,
Devon, Oregon, Dutch and Sekirk) depending on recessive or dominant mutations of this gene. The "h"
series is expressed in the Sphinx, (no relation to the Rex), which has no primary hairs. However,
secondary hairs are present on the extremities (nose, pinnae, limbs, tail). The dominant "Wh" gene is
responsible for the development of very curly hair. In this case, the three types of hair are entwined.
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1 : Structure and functions of skin and coat

Genetics of hairpigmentation
The colour of the skin and coat depends on the quantity, type and location of melanin pigments in the
skin. There are two types of melanin, eumelanin @lack/brown)and phaeomelanin (red/yellow). The large
number of colours seen in the various breeds of cats depends on the prevalence, combination and
distribution of these two types of melanin in the skin. White colouration occurs through an absence of
pigmentation resulting from lack of melanisation of melanocytes. This is called albinism when complete
( i s is very light blue or red).
The most important gene involved in feline hair pigmentation is the dominantA "agouti" gene responsible
for the o r i g ' i , wild, banded pigmentation of primary hairs. Cats carrying this gene have dark-tipped
hairs with one or more bands of colour. Those which express the recessive a, "non-agouti" gene, have
uniformly pigmented hair. Wid cats also carry the T "tabby" gene, responsible for dark markings on a
lighter, generally yellow or grey, base. There are different variants of the "tabby" gene which produce
variable distribution of stripes or spots on the body.
Colours seen in domestic cats are divided into phaeomelanic colours (orange / g'mger (red) and cream
(diluted red)) and eumelanic colom (black, grey (generally called blue, diluted black), chocolate
(chestnut), lilac (diluted brown), cinnamon (light chestnut) and fawn (diluted cinnamon). As the genes
responsible for hair colour are situated on the X chromosome, cats with concurrent red and black/brown
colours are generally females which can carry the two red and black/brown genes on each of the two X
chromosomes.
Colour dilution is caused by the recessive d gene. In colour-dilute breeds, l i e the C h W u x , hairs contain
irregularly distributed melanosomes grouped in melanin clusters in the melanocytes and hair cortex. A
foUicular dysplasia of colour-dilute cats has been described in the Cornish Rex.
Silver coats are coded by the I "colour inhibitionn gene. Colour is lost in the proximal hair which grows
out white. There are different types of silver coat, depending on the extent of hair depigmentation.
The recessive variants of one pdcular gene, the C gene, are responsible for "colour point" coats, typical
of the Siamese. Tnis gene codes for an enzyme whose activity is temperature-dependent. This causes
excessive colouration in colder regions such as the extremities. The ca gene is responsible for blue eye
albinism, the c gene for red eye albinism.
White spots are due to the expression of a different gene, the S "white spotting" gene, whereas generalised
white colouration of the coat is encoded by the dominant W gene. This latter gene is often associated with
deafness because it causes degeneration df the cochlea and atrophy of the organ of Corti.

3utaneous glands
Sebaceous glands
Sebaceous glands are simple alveolar holocrine glands, associated with hair follicles in groups of two or
thee (Fig. 1 : 9). Their excretorv ducts open into the follicular isthmus. Sebum is the product of sebaceous
gland cell destruction in the Addle of the gland. It combines with sweat, produied by the apocrine
glands, to form a lipid emulsion (the surface hydrolipid film) which protects the skin surface. The
sebaceous glands are large and numerous in the skin around the lips and on the chin, and play a role in
territorial marking when the cat rubs its face against objects and human beings. The perianal glands and
supracaudal organ are modified sebaceous glands which are larger than the other glands. As development
is hornonally dependant, they may be particularly large in entire male cats. Excessive accumulation of
glandular secretion in this region is known as stud tail.
Sweat glands
Apocrine sweat glands are present over the whole surface of hairy skin and their ducts open above the
sebaceous gland ducts into the follicular isthmus. Apocrine glands are simple tubular glands with a
straight duct and a convolutedsecretory part surrounded by myoepithelial cells (Fig. 1 : 10).Tney produce
an aqueous secretion which forms an emulsion with sebum at the skin surface (the surface hydrolipid
film).
Eccrine sweat glands have the same structure as apocrine sweat glands but are only found in non-hairy
areas of skin l i e footpads. Their ducts open directly at the skin surface. These sweat glands are situated
close to blood vessels and are controlled by blood adrenaline and noradrenaline. Hence, frightened cats
sweat excessively from their footpads.
1~;
~.4hctical
~~ ~

GuidetoFehe Dermatology
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Modified sweat glands make up the mammary glands, ceruminal glands and the glands of the anal sacs
There are no sweat glands on the nasal planum '.

Functions of the skin and coat


The skin and coat have many important functions including protection from the external environment,
thermoregulation and maintenance of biochemical homeostasis. They also have metabolic, sensory, immunological
and social functions.

Mechanical protection
The coat is the first barrier to mechanical trauma. It is often thick because of an undercoat covering the
whole skin surface. Dermal collagen fibres impart a high tensile strength and prevent skin tearing.

Protection against water


Thanks to this thick coat and the surface hydrolipid film,water does not readily reach the skin surface.
The orientation of primary hairs ensures that water droplets are quickly shed. Constant grooming keeps
primary hairs clean and ensures optimal protection from water. Certain breeds, such as the Norwegian
Forest, exhibit a very hydrophobic coat. This characteristic is often tested by show judges who let several
drops of water fall on the hairs to see if the drops slide off the coat without wetting the animal. If water
does reach the skin surface, it cannot pass through it because intercellular spaces in the horny layer
contain lipids impermeable to water and water-soluble substances.

Protection from light


The coat is an excellent barrier to visible light and ultra-violet (UV) rays. In the areas where the coat is
sparse or absent, pigments (mainly melanins), keratin, proteins and blood absorb the UV rays and prevent
certain skin lesions. Nevertheless, in white or light-coated cats with little pigment, solar keratotic lesions
and epidermal carcinomas can appear in sparsely-haired areas such as the pinnae, nose, eyelids and the
a m behind the ears, following prolonged or frequent exposure to the sun.

Thermoregnlation
Temperature regulation is also an important function of the coat and skin. The thick coat and the layer of
subcutaneous connective tissue, rich in adipose cells, protect the cat when the temperature is cold. An
insulating air cushion exists between the hairs. The size of the air cushion can be increased or decreased
by the action of arrector muscles. The undercoat is shed in the spring and re-grows in the autumn in breeds
which experience large seasonal temperature variations. This is_particularlytrue in the k k i s h Van breed,
origimating in central Anatolia, where the temperature variation between summer and winter can be as
much as 50°C. The highly developed dermal vascular system plays an important role in thermoregulation.
Vasodilatation produces heat loss and vasoconstriction causes closure of arteriovenous shunts, which
prevents excessiveheat loss from the circulation. The cutaneous vascular system stores large quantities of
blood and peripheral vasoddatation or vasoconstriction can also have an effect on central blood pressure.
Cats have a particular way of reducing their body temperature as, unlike in humans, sweat glands are not
involved in thermoregulation. Cats constantly wet their coats by licking. This produces a cooling effect
as saliva evaporates from the skin surface.

Biochemical homeostasis
Many substances are stored in the dermis and subcutaneous comective tissue. Large quantities of water
and electrolytes are associated with proteoglycans and other molecules in the ground substance, while
lipids and fat-soluble molecules are stored in subcutaneous adipose tissue.

Metabolic and immunological functions


For a long One, the skin was considered to be an organ with only a passive protective role, not one that
involved the individual's general metabolism or immunological defence mechanisms, with the exception
of vitamin D synthesis. Various metabolic pathways prodicing systemic effects (for example, effect of
androgens and oestrogens on smell) have recently been demonstrated in the skin. In addition, an
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1 : Structure and functions of skin and coat

immunological system has been identified, capable of specitic reactions against micro-organisms,
parasites or foreign antigens present on skin.

Sensory functions
Various different sensations such as pain, pruritus, heat, cold, pressure and touch are felt at the cutaneous
level. Free nerve endimgs in the epidermis are responsible for sensations of pruritus, pain, heat and cold.
Specialised structures, l i e Pacinian corpuscles (Fig. 1 : 1l), tylohich pads, whiskers " (Fig. 1 : 12), hair
follicles, type C mechanoreceptors and Merkel cells all have mechanoreceptor functions 12.

Social functions
The colour of wild cats l i e the Lynx, enables the animal to hide easily in woods and forests (mimicry).
Domesticated breeds have lost this characteristic, perhaps with the exception of black cats used in the
Middle Ages, to chase mice and rats from the holds of ships. Specialised glands, like the hepatoid
circumanal glands, the anal sacs, the supracaudal organ and the sebaceous glands of the lips and chin,
produce pheromones. Pheromones are used in territorial marking, facilitate recognition between
individuals and influence sexual attraction. Various fractions (F3, F4) have been isolated recently. Cats in
dangerous situations raise the hairs on their back and tail and present themselves sideways-on to their
adversary in order to appear larger.

Cutaneous microbi9logy
-- and barrier functions
Horny layer
' The compact horny layer represents the f h t physical banier to infections and parasitic infestations. In
normal conditions, water, water-soluble molecules and micro-organisms cannot cross it. Constant
shedding of the most supeficial cells reduces excessive bacterial colonisation which could otherwise
predispose to infection.

Dermis
If a micro-organism or parasite enters the dennis, it must pass through a very dense network of collagen
fibres and intercellular ground substance molecules, before fmdiig itself in contact with a very active
immune system.

Hydrolipid surface film


The sebum and sweat form an emulsion on the skin surface which acts both as a physical banier,
preventing the passage of water and water-soluble substances, and a chemical banier, made possible by
the presence of certain substances involved in skin defence mechanisms. These include Ransferrin which
l i i t s bacterial proliferation, and free fatty acids such as linoleic acid, produced from surface higlycerides
by bacterial lipases, which prevent skin colonisation by pathogenic micro-organisms. Other lipids such as
glycosphingolipids arising from the decomposition of the horny layer might play an even greater
antibacterial role. Specific factors, such as complement and immunoglobulins, can attach themselves to
the skin surface and prevent microbial adherence and proliferation. Skin pH in cats varies between 5.73
and 6.01, with the exception of the nose and footpads (areas of eccrine secretion) where it is less acidic,
between 6.81 and 7.97". A low pH has bacteriostatic and bactericidal effects.

Resident microbial flora


The resident surface flora, which occupies micro-ecological niches, prevents colonisation by pathogenic
micro-organisms by producing antibiotics, enzymes or other toxic substances. They utilise available
nutrients. Resident bacteria on cat skin include Micrococcus spp., coagulase-negative staphylococci
(especially Staphylococcus simulans), a-haemolytic streptococci and Acinetobacter spp. 12. It is possible
to isolate more bacteria from cats in close contact with humans than froiii cats living in catteries, which
suggests that humans may be involved in transmitting bacteria to cats ". Transient bacteria, isolated from
cat skin, include R-haemolyhc streptococci, Escherischia coli, Proteus mirabilis, Pseudomonas spp.,
Alcaligenes spp., Bacillus spp., and staphylococci 12. These hacteria can occasionally become pathogenic
if conditions are favourable for multiplication.
Yeasts of the genus Malassezia are commensal micro-organisms found on the skin of cats, especially in
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humid regions such as ears, penanal and inguinal regions Id. Malassezia pachydermatis, Malassezia
sympodialis and Malassezia globosa have been isolated from cat skin 14.
Van'ous saprophytic fungi can be isolated from the skin and coat of normal cats. These include Alternaria
spp., Aspergillus spp., Cladosporium spp., Mucor spp., Penicillium spp. and Rhodotorula spp. ",''.
Microsporum canis isolated from normal cats must always be considered a pathogen 16.

REFERENCES
1. Strickland, J. H. & Lois Calhoun, M. Amer J Vet. Res. 24, 1018-1029 (1963).
2. Scott, D. W Vet.Dermatol. 1,6569 (1990).
3. Baker, B. B. Amer. J. Vet. Res. 3,93 (1973).
4. TsagaraIds, C., Marchd, T., Magnol, J. P., Fournel, C., Dezutter-Dambuyant, C. & Schmitt, D. Research in Virology 145,245-249 (1994).
I 5. Saint-Andr6 Marchal, I., Dezutter-Dambuyant. C. & Mutin, I. P. and others Br. J. Dermarol. 136,961-965 (1997).
6. Badleston, D. L., Roosje, P. & Goldschmidt, M. H. J. Vet. Allergy Clin. Immunol. 5,54-58 (1997).
7. Blazej, A. Galatik, A. Galatik, I. Krul, Z. & Mladek, M. Atlas of Microscopic Structures of Fur Skins (Elsevier, Amsterdam, 1989).
8. Lochte, m.Atlas der Menschlichen und TierischenHaare (Paul Schps Verlag, Leipzig, 1938).
9. Creed, R. E S. Vet.Rec. 70, 171-175 (1958).
10. Kumamoto, K., Takei, M., Kinoshita, M., Ebara, S. & Matsuura, T. J. Anat. 182,23-28 (1993).
11. Ikeda, M. & Okada, S. Okajimas Folia Anaromia Japanensis 67,365-369 (1990).
12. Scott, D. W Miller Jr, W. H. & Griffin, C. E. Muller & Kirk's Small Animal Dermatology,5th edih'on (Saunders, W.B., Philadelphia, 1995).
13. Meyer, W. & Neurand, K. Arch. Dermatol. Res. 283.16-18 (1991).
14. Bond, R., Howell, S.A.,Haywood, P. I. & Lloyd, D. H. Vet. Rec. 141,200-201 (1997).
15. Moriello, K. A. & DeBoer, D. J. Amer J. Vet. Res. 52,602-606 (1991).
16. Moriello, K. A,, Kunkle, G. A. & DeBoer, D. J. Vet. Dermatol. 5,57-62 (1994).

AKNOWLEDGEMENTS
Diagrams 1 : 1 and 1 : 2 have been reprinted with kind permission fmm Peters S.: "Haut und Haarkleid kim Hund", Ferdinand EnkeVerlag, 1997,Pages 4 and 21.