Medical Mycology 1999.

37,295-306 Accepted I January 1999

Review Article

Ma/assezia pachydermatis: a review

J. GUILLOT* & R. BONDt
*Unite de Parasitologie-Mycologie, UMR Biologie Mo/eculaire et lmmunologie Parasitaires et Fongiques, Ecole
Nationale Veterinaire d'Aifort, Maisons-Aifort, France and tDepartment of Small Animal Medicine and Surgery. The
Royal Veterinary College, Hatfield, Herts, UK

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Malassezia pachydermatis is of importance in both veterinary and human medicine.
Its taxonomic status and physiological characteristics are now better understood.
Skin disease associated with this lipophilic yeast is now commonly recognized,
especially in dogs. However, further studies are required to elucidate the mechanisms
which allow this yeast to proliferate and induce disease. Skin colonization is common
in pet carnivores which consequently constitute a source of M. pachydermatis for
susceptible humans. In the future, the development of efficient typing systems should
allow better understanding of the transmission mechanisms.
Keywords Lipophilic yeast, Malassezia pachydermatis

Introduction in intensive care units [7,8]. The purpose of this review is

The lipophilic yeast Malassezia pachydermatis is part of
the normal cutaneous microftora of most warm-blooded
vertebrates. The fungus was first described as Pityrospo-
rum pachydermatis in 1925 [1]. Recently, the nomenclatu-
ral revision of the genus Malassezia did not question the
integrity of the only non-lipid-dependent lipophilic spe-
cies [2-4]. M. pachydermatis shows distinct morphologi-
cal and physiological characteristics, allowing thus a
straightforward distinction from any other yeast. In the
two la;t decades, the opportunistic nature of M. pachy-
dermatis has been clearly demonstrated. The normally
commensal yeast may become a pathogen whenever alter-
ation in the skin surface microclimate or host defence
occurs. Otitis externa as well as seborrheic dermatitis in
dogs and cats are frequently associated with a large
number of M: pachydermatis. Similar cases have also
been described in wild animals. Reports concerning hu-
man infections caused by M. pachydermatis have been
fewer than those with lipid-dependent Malassezia yeasts
[5,6]. However, the species may be isolated from neonates
<;:orrespondence: J. Guillot, Unite de Parasitologie-Mycologie,
Ecole Nationale Veterinaire d'Alfort, 7 avenue du General de
Gaulle, 94704 Maisons-Alfort, France. Tel: + 33-1-43-967157; fax:
+ 33-1-43-753507; email: j.guillot@vet-alfort.fr
to present the current status of the knowledge about the
lipophilic yeast M. pachydermatis.
Mycology
Taxonomy and nomenclature
In 1925, Weidman observed bottle-shaped yeast-like cells
in scales from an Indian rhinoceros (Rhinoceros unicor-
nis) with exfoliative dermatitis [1]. This microorganism
was able to grow on ordinary media apparently without
lipid supplementation. Cultural characteristics included
slow growth, dark yellow colour and dryness of the
colonies. Weidman classified this yeast in the genus Pity-
rosporum because of its resemblance to P. ovate ('spore of
Malassez'), the purported agent of human dandruff.
However, the regularity in form and the smaller size of
the yeast recovered from the rhinoceros (2-3 f.Lm against
3-8 f.Lm for P. ovale) led Weidman to describe the new
species as P. pachydermatis. Ciferri and Redaelli [9] ex-
amined a strain from Weidman and chose P. pachyder-
matis as the type of the genus since Weidman's strain was
at that time the best studied Pityrosporum strain. In her
review of anascosporogenous yeasts, Lodder [10] com-
pared a subculture from the Weidman isolate and a strain

© 1999 ISHAM
 Gu1llot & Bor.d

probably originating from Sabouraud's collection that T~!Jie l Composition of the genus ;'v/alasse~ia.

she combined under the unpublished name P. rhinucero-

Species Isolates rRNA sequences
sum. The cultures were considered to be identical but,
unfortunately, were not preserved after 1934. L'd. fiojio· CBS 1878, neo- AF063214
In 1955, Gustafson isolated bottle-shaped yeasts from type culture
otitis externa in dogs (II]. He related his isolates to the !VI. pachyderma- CBS 1879, neo- AF0632!5 (sequence type
tis type culture '!a')
genus Pityrosporum on the basis of their cellular shape
ill/. sympodia!is CBS 7222, type AF064024
and the type of budding, but distinguished them from culture
already described species. In contrast to the other lipid- !VI. globosa CBS 7966, type AF064025
dependent species (P. ovale and the newly described P. culture
Jvf. restncta CBS 7877, type AF064026

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orbiculare, Gordon 1951), Gustafson's isolates did not
culture
require fatty acid supplementation for growth. These ivf. obtusa CBS 7876, type AF064027
isolates grew easily and were kept alive by monthly culture
transfers on common media. Gustafson wrongly con- ivt. slooffiae CBS 7956, type AF064028
cluded from Lodder's description that the yeasts recov- culture
ered from rhinoceros skin by Weidman grew poorly and
were very difficult to maintain. For this reason,
Gustafson created the new species P. canis. Fraser [12] classified into seven sequence types, named sequevars
also studied many strains from both healthy and diseased (I a- Ig) [18] (Fig. I). Some of these variants (Ic, Ie, If)
seemed to be host specific; they included isolates exclu-
ears of dogs. He concluded that Pityrosporum strains
sively recovered from rhinoceros, dogs or ferrets, respec-
from dogs were similar to those described by Weidman
tively. DNA/DNA reassociation experiments displayed
and Lodder. There was subsequently no reason to justify
lower values with strains from different sequevars (i.e.
the establishment of P. canis. In 1970, Slooff assigned all
from different origin) (85-90'%) than with strains of the
strains of Pityrosporum growing on media without lipid
same type (87 -I OO'Y<,). These results do not question the
supplementation to a single species P. pachydermatis [13].
taxonomic unity of J'vf. pachydermatis, but might indicate
Evidence for the synonymy of lv!alasse::.ia (Bail!on 1889) that this species is in the course of differentiation and
and Pityrosporum (Sabouraud 1904) was clearly recog- probably adaptation to specific animal hosts [3, 18].
nized and accepted in 1984 with anterioiity for the
generic Jl;falasse::.ia [14]. P. pachydermatis was then ac-
cepted as !vi. pachydermatis, a combination already pro- <y::: 0
lb

posed by Dodge in 1935 (15] and accepted by Gordon in pe Ic

1979 [!6]. In absence of original material (Weidman's M. pachydermatis
e ld

~
isolate), the strain 238S isolated from otitis externa in the
type le
dog by Gustafson [II] was designated as neotype culture
~~~~-1
(CBS 1879, ATCC 14522). lype lf

Genomic studies recently confirmed the specific status type lg

of non-lipid-dependent lv!alassezia yeasts. Strains recov-

ered from wild animals (including rhinoceros) were
proved to be conspecific with those recovered from dogs. M. obtusa
The nDNA base composition (GC content) was found
around 55·5'% and DNA/DNA reassociation experiments
!.......--~~~-~ M. sympodU!lis
consistently showed percentages over 85% [3, 17]. In con-
trast, lipid-dependent Malassezia yeasts were distributed 1-....~~~~~~~~~~~~~ M. slooffiae

among six distinct taxa (lvf. jurji1r, M. sympodialis and

four new species, lVf. globosa, lvl. obtusa, M. restricta and
J'vl. sloojfiae Table I) on the basis of genomic compari-
sons and ribosomal large subunit (LSU) sequencing [2].
The latter method allowed a further infraspecific separa-
tion with most of the Mafasse::.ia species characterized by
several sequences differing from each other by only a few
nucleotides. Consequently, with partial LSU rRNA se-·
quencing, one hundred i'vf. pachydermatis isolates were
lipid-dependent Malassezia yeasis
Fig. I Phylogenetic tree based on partial large subunit rRNA
sequences of iv!a/asse::ia yeasts (after Guillot & Gw§ho, 1995 [4] and
Gw§ho eta!., 1996 [2]). Seven sequence types were detected for 1'Vf.
pachydermalis. Most of lipid-dependent Afalasse~ia yeasts were also
represented by several closely related sequences (data not shown).
The distance between two species or subspecies, measured in substi-
tutions per nucleotides. is obtained by summing the length of the
connecting branches along the horizontal axis.

© 1999 ISHAM, Medical Mycology, 31. 295-306


Other typing methods have been applied to ;v!. pachy-
dermatis isolates. Karyotyping seemed to provide little
information, since all lvf. pachydermatis isolates revealed
rather uniform banding patterns [19-22]. Using restriction
fragment length polymorphism (RFLP) analysis on whole
cell DNA, Midgley & Schechtman distinguished nine
distinct patterns out of 50 isolates, mostly of canine origin
[23]. The same method, but using a different restriction
enzyme, allowed Anthony et a!. [19] to distinguish 10
patterns among the 32 isolates examined. In the latter
study, Southern blotting of Bglii digests with a poly (GT)
probe was also performed; this method together with PCR
fingerprinting appeared to be the most efficient epidemio-
logical tool [19,22]. M. pachydermatis isolates of various
origin were unequivocally identified, whereas all clinical
isolates recovered in the same intensive care unit were
shown to be indistinguishable [22]. The heterogeneity of
the species was also demonstrated using yeast killer system
(24]. Recently, multilocus enzyme electrophoresis (MLEE)
analysis was proved to be useful for the study of genetic
structuration and biodiversity of M. pachydermatis (25].
Interestingly, both RFLP and l'viLEE results matched very
well with those reported using· LSU rRNA sequencing
[3, 17,25].
Phylogeny
Sexual reproduction has been reported neither for !vi.
pachydermatis nor for the other lipophilic yeasts. However,
high GC content ( > 50'1<,), lamellar nature and biochemi-
cal composition of the cell wall (cf. infra), positive diazo-
nium blue B staining reaction and urea activity clearly
indicate that the genus Malassezia has basidiomycete
affinities (2,3,26-28]. Ribosomal sequence comparisons
confirmed the latter assignment but also demonstrated that
lvfalasse::.ia yeasts could not be clearly related to any other
existing order of heterobasidiomycetes [43]. Cell wall
characteristics (ultrastructure, chitin distribution) confi-
rmed this remote position (3,28-30]. Lipophilic yeasts
fonned a monophyletic group that probably diverged from
other basidiomycetes a long time ago. In fact, the high rate
of nucleotide substitution reported between 1\1alassezia
species largely exceeded the values observed within other
yeast genera [31]. These findings were in accordance with
an early emergence and differentiation. They might also be
incorporated in the hypothesis of an unusually fast 'molec-
ular clock' leading to extensive polymorphism and subse-
quently to overevaluation of phylogenetic distances [3,4].
Morphology
Most 1j,f. pachydermatis strains have short oval to ellip-
© 1999 ISHAM. Medical /'1ycology. 37, 295-306
rV!aiassezJa pachyderrnws: a review
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Fig. 2 Scanning electron micrograph of lvi. pachydermatis. Unipo-
lar budding yeast cells with proeminent collarettcs are easily recog-
nized.
soidal cells (2·0-3·0 x 4·0-5·0 ~tm). Globose cells may also
be present. Slooff (13] distinguished a small-celled type
(2·5-4·8 x 2·6-5·0 J.lm) from a large-celled type (3·8-
6·0 x 4·8- 7·0 ~tm). Kisser a!. [32] also reported microscop-
ical differences between globose (3·8-3·9 x 4·7 -4·9 J.lm)
and a rode-shaped morphotype (1·9-2·1 x 5·1-5·3 ~tm).
As with other lipophilic yeasts, its reproduction proceeds
by enteroblastic unipolar budding. M. pachydermatis
shows a broad budding base with a distinct collarette (Fig.
2). The medium used for cultivation does not seem to
influence the yeast cell micromorphology. Under micro-
aerophilic conditions, a beginning of yeast-mycelial trans-
formation was reported for i\.I. pachydermatis by
Faergemann & Bernander [33]. However, only very short
filaments were observed. Contrary to several lipid-depen-
dent llifalasse::.ia yeasts, lvf. pachydermatis may not, there-
fore, be considered as a species able to filament [2].
On Sabouraud's glucose agar at 32 °C, M. pachyderma-
tis colonies are mat, convex and pasty with a dry and
smooth surface. Initially white to ivory in colour, they
darkened with age from tan to brown. Gordon [16]
observed that there was sometimes a pink tinge in the early
growth, whereas Slooff [13] reported a pink1sh shade
after one month at 20 oc. Similarly, Midgley described
a strain (GM 439 isolated from the skin of a civet) that
formed colonies with a. constant and persistent pinkish
colour [3]. After 7 days incubation at 32 oc. colonies
usually have a diameter from 3 to 5 mm. However,
Huang & Little [34] described differences in the size
of M. pachydernwtis colonies. They distinguished large
and small-colony types among isolates recovered from
 Guillot & Bone
canine ears or skin. These phenotypic differences were
reinforced by variations in fatty acid composition. More
recently, Bond & Anthony [35] examined /II. pachyder-
rnatis strains that apparently behaved like lipid-depen-
dent yeasts; they initially either failed to develop on
regular media or formed very small colonies on
Sabouraud's glucose agar. Electrophoretic karyotypes of
these strains were consistent with illl. pachyderrnatis. Ri-
bosomal RNA sequencing proved that all these atypical
isolates represented a genetically distinct population and
could be related to the sequence type 'Id' composed of
canine strains exclusively [ 18].
Ultrastructure
Transmission electron microscopical observations
demonstrated a thick (up to 0·25 ~Lm) and multilame!lar
cell wall which is typical of basidiomycetous fungi. De-
pending on culture conditions as well as fixation and
staining methods, two or three layers were usually dis-
tinguished for both M. pach.vdermatis and lipid-depen-
dent Malasse::ia yeasts [28.29,36)7]. An additional outer
layer was described for ll;f. pachydermatis by Winiarczyk
[38]. This very thin outer layer could be observed only
in specimens which were prepared by means of methods
leading to a minimal washing out of lipids. A similar
structure was also reported for J1;f. ji1rji1r and identified
in this case as a diffuse microfibrillar capsule [29]. In
fact, the most interesting ultrastructural feature is the
corrugated nature of the inner surface of the cell wall.
This structure, unique in the Kingdom Fungi, corre-·
sponds to the helicoidal invagination of the plasma
membrane around the cell [28,37,39,40]. In the bud, this
characteristic feature becomes progressively apparent as
it matures.
The typical enteroblastic budding of basidiomycetous
yeasts leaves a collarette on the mother cell wall. For iVJ.
pachydermatis, the budding process was precisely de-
scribed by Nishimura et al. [41]; these authors distin-
guished several stages from the protrusion of the
cytoplasm and the dimpling of the mother cell wall to
the complete release of the bud. lvf. pachyderrnatis was
clearly characterized by the width of the bud base (0·9-
1·1 Jlm in diameter). A much narrower base was re-
ported for lipid-dependent yeasts (0· 5-0· 7 Jlm) [42].
Cytoplasmic organelles appeared comparable to those
described for other yeasts [38,41]. The size of the nu-
cleus was the same as that of l'vf. jiojitr (0·8 ~Lm in
diameter) [4!]. Gabal [37] reported a distinctive struc-
ture of the nucleus in k/. pachydemwtis. This result was
not confirmed by other investigators [38.41 ].
Physiolog-;
l1.1. pachydermatis is a lipophilic yeast and the growth is
improved by addition of lipids in the medium. However,
this species differs from all the other lv!alasse::ia yeasts
by the fact that lipid supplementation is not an absolute
requirement for growth. Thus, M. pachydermatis is the
easiest Malassezia species to identify [43]. M. pachyder-
matis has no block in de novo synthesis of myristic acid
and it has been shown that growth does not depend on
the presence of long chain fatty acid precursors in the
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medium. However, differences in lipid requirements
have been reported [35]. Isolates belonging to the se-
quence type 'Id' exhibit a slow rate of growth and a
rapid loss of viability when initially cultivated on
Sabouraud's glucose agar [18]. After several transfers,
these isolates form small colonies, but usually loose their
exacting lipid requirements [3,18].
lvf. pachydermatis does not develop in liquid media
used in the standard assimilation tests [13, 14]. Using
auxanogram plates, Slooff [13] and Saez [44] reported
that growth is possible with glucose o-mannitol and
o-glucitol, and sometimes with glycerol, sorbitol, lactic
and succinic acids. Sanguinetti et al. [45] found that
glucose and o-mannitol were the only carbon sources to
be assimilated. Kiss et a!. [32] distinguished two physio-
logical types on the basis of o-mannitol and sorbitol
assimilation, hydrolysis of Tween 80 and requirement
for nicotinic acid. In usual commercial yeast identifica-
tion systems, such as API-:ZOC (bio-iVIerieux), JVj_ pachy-
dermatis is usually inactive [3,5,32]. However, Marcon &
Powell [5] reported that iVF pachydermatis might be pos-
itive for glucose, glycerol and sorbitol leading to the
misidentification of the yeast as Ccmdida lipolytica.
Sugar fermentation has never been demonstrated
[13,14,32,44]. Potassium nitrate is not assimilated
[13,32]. Urea is hydrolysed and the catalase reaction is
usually positive but with a delayed gas formation [2,32].
Surprisingly, Mathieson et al. [46] reported variations in
urease activity between strains. In fact in most cases,
the reaction was negative on a solid medium supple-
mented with phenol red. Using API-ZYM system (bio-
Merieux, Marcy !'Etoile, France), alkaline and acid
phosphatase, phosphoamidase and two esterase lipase
activities were detected [26,47]. Using esters of lauric,
palmitic and stearic acids, Kiss et a!. [32] confirmed that
the yeast is positive for esterase activity. In the latter
study, peroxidase and lecithinase activities were also de-
tected. but no coagulase or haemagglutinating activity
was observed. The proteolytic activity of lVI. pachyder-
matis was first demonstrated on bovine serum agar [48]
and further confirmed using Sabouraud glucose agar
and Sabouraud glucose liquid medium, supplemented
with casein [46].
© 1999 ISHAM. i'1edJcall"lycology. 31. 295-306

M. pachydermatis is resistant to cycloheximide
(1000 ppm) (2,44,49]. In fact, increased growth has even
been observed with high concentrations of cycloheximide
(2000 ppm) [50]. Some isolates may be sensitive to lauric
acids [3,43]. The latter characteristic was first reported by
Ushijima eta!. [49], who devised selective and differential
media for lipid-dependent Malassezia yeasts. Supplemen-
tation with laurate (750 mg mi- 1) leads to the complete
growth inhibition of M. pachydermatis.
Growth is possible at temperatures from 25 to 41 oc
[2,13,44,45]. Optimum temperature is 37 oc [2,13]. M.
pachydermatis develops equally well in aerobic, mi-
croaerophilic and capneic conditions, while in anaerobic
conditions, growth is poor [33].
M. pachydermatis isolates could be maintained by
monthly transfers on Sabouraud's glucose agar. After
7 days incubation at 32 or 37 °C, yeasts are kept alive at
room temperature. M. pachydermatis seems to be particu-
larly sensitive to the cold [51]. After 3 months at 4 °C,
most isolates were no longer viable, whereas some lipid-
dependent Malassezia yeasts were still alive even after
5 years at the same temperature. Cells survive lyophiliza-
tion [2,51]. Preservation in liquid nitrogen is also efficient
[51 ,52]. Lorenzini & de Bern ardis (53] suggested the use
of the glucose nutrient agar supplemented with 1·5';{,
yeast extract and 1% Tween 80; cultures maintained at
room temperature were kept alive for at least 3 months.
The chemical composition of M. pachydermatis cells
has been investigated by only a few authors. The protein
fractions of 30 isolates from dogs was investigated by
SDS-Page [54]. Numerous protein bands ranging from
15·0-145·7 kDa were detected, but profiles did vary ac-
cording to the origin of the strains (external ear canal or
skin). Breierova et a!. [52] demonstrated that the content
of unsaturated acids (oleic, linoleic, myristoleic and
palmitoleic) was 2·5-7·5 times greater than that of satu-
rated acids (palmitic, stearic and myristic). Great varia-
tions were noticed between strains. The small-colony type
described by Huang & Little [34] contained less linoleic
acid than the large-colony type and oleic acid could not
be detected. Furthermore, the small-colony type con-
tained five unidentified fatty acids. Addition of fatty acids
in the growth medium changed the fatty acid composi-
tion of strains. Saturated fatty acids were incorporated as
such into the cells but unsaturated acids induced complex
changes, including the production of a new fatty acid.
The unexpected effect of unsaturated acids on the bio-
chemical composition of cellular membranes might ac-
count for the mycostatic action of oleic and linoleic acids
in vitro [34].
Hamamoto et a!. [55] demonstrated that M. pachyder-
matis was characterized by the coenzyme Q9 system. The
© 1999 ISHAM, Medical Mycology. 37, 295-306
Malassezia pachydermatis: a review
membrane sterol composition was analysed for isolates
sensitive or resistant to polyene antibiotics, and a greater
amount of ergosterol was found in sensitive strains than
in resistant ones (56]. The cell wall of M. pachydermatis
stains red to violet with diazonium blue B [2,28]. This
feature, together with the absence of cell wall lysis by
13-(1-3)-o-glucanase [26], is consistent with the ba-
sidiomycetous nature of M. pachydermatis.
Ecology
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Skin carriage
Malassezia yeasts are principally isolated from the skin of
a variety of mammals and birds and are seldom recov-
ered from the environment [6,51]. M. pachydermatis is
frequently found on wild and domestic carnivores, in-
cluding dogs, cats, bears, ferrets and foxes [3,57,58]. It
has also been isolated from species as diverse as rhi-
noceros, pigs, primates, pinnipeds, horses and birds
(3,44,57,59-64]. In contrast, M. pachydermatis was not
recovered from rodents and lagomorphs in one recent
survey [57]. Certain host species such as pigs, rhinoceros
and cats, may be colonized by both M. pachydermatis
and lipid-dependent Malasse:::ia spp. [57,62,65-67]. Hu-
man skin is frequently colonized by lipid-dependent
Malassezia yeasts, but seldom by M. pachydermatis
(6,68].
Carriage of M. pachydermatis has been widely studied
in dogs because of its importance as an apparent skin
pathogen in these animals. Gustafson [11] first isolated
this yeast from the external ear canal of both healthy
dogs and dogs with otitis externa. In subsequent investi-
gations, the yeast was recovered from this site in between
2 and 49°!£, of healthy dogs [11,12,69-71]. Hajsig eta!.
(72] sampled various mucosal sites and isolated M.
pachydermatis from 33 out of 57 healthy dogs. The yeast
was isolated from the anus, rectum, anal sacs and vagina
in approximately 40% of dogs, whereas it was infre-
quently recovered from the mouth, nostrils and nasal
planum. Dufait (59] cultured M. pachydermatis from 81
out of 94 hair samples from dogs and found the fre-
quency of isolation from healthy dogs and dogs with skin
disease to be equivalent.
Bond et a!. [73] isolated M. pachydermatis from the
anus of 52·5% and the oral cavity of 27·5% of 40 healthy
dogs of various breeds, whereas it was less often found in
the nose, prepuce and vulva. Skin carriage was frequently
observed in the interdigital area (60%), on haired skin of
the lower lip (75%) and in the external ear canal (32·5%),
but it was rarely recovered from the axilla, groin or
dorsum. Kennis et a!. [74] also found the chin and perio-
ral area to be frequently colonized in healthy dogs. In
basset hounds, a breed predisposed to ;l;Ja/asse::ia der-
matitis [75,76], both the frequencies of isolation and
populations sizes of the yeast in healthy hounds were
Dermatitis and otitis
found to be significantly higher at skin and mucosal sites
when compared with those of healthy dogs of various
other breeds [77]. In a temporal study of ivf. pachyderma-
tis carriage in eight healthy beagles sampled fortnightly
over I 0 weeks, persistent carriage was identified at the
external ear canal in seven dogs and at the anus in three
dogs [78].
Infections associated with M, pa.chydermatis
1n dogs
Gustafson [11] first proposed that ivf. pachydermaris
might have been involved in the pathogenesis of otitis
externa in the dog. He was able to induce otitis externa
by applying Jlvf. pachydermatis to the external ear canal,
whereas a Pityrosporum ovale strain of human origin had
no effect. Fraser [12] isolated M. pachydermatis in com-
parable frequencies from healthy dogs and dogs with
otitis externa, but found that affected dogs tended to

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Adherence to canine corneocytes
Adherence. the specific attachment of microorganisms to
cells and tissues, is thought to play an important role in
the colonization and infection of mammals by pathogenic
fungi. Adherence allows the fungus to resist physical
forces which may otherwise result in its removal from the
host, and may precede other pathogenic events such as
germination and invasion [79.80]. Factors which influ-
ence the adherence of Candida spp. to epithelial cells have
been extensively investigated [81], whereas the adherence
of Malasse::ia spp. has received less attention [82,83].
Suspensions of 111. pachyder;rwtis adhered to canine
corneocytes attached to adhesive tape in a dose (P <
0·001) and time-dependent < 0·01) manner; adherence
was maximal after 2 h [84]. Af. pachydermatis cells were
approximately 10 times more adherent than Saccha-
romyces cerevisiae (P < 0·00 1) cells after 2 h ·incubation.
Stationary-phase cells adhered better < 0·05) than ex-
ponential-phase cells. Pre-treatment of yeasts, or cor-
neocytes, with 0·1 '% trypsin for 30 min reduced (P < 0·0 I)
the adherence of four and two, out of five strains, respec-
tively, suggesting that trypsin-sensitive proteins or glyco-
proteins on the yeast cell wall and on the corneocyte
surface play an important role in the adherence of lvf.
pachydermatis to canine corneocytes in vitro. Incubation
with 300 mM solutions of D( +) mannose, sucrose and
N-acetyl D-glucosamine had no consistent effect.
A panel of four lectins was used to further investigate
the role of carbohydrates in the adherence of i\1. pachy-
demwtis to canine corneocytes in ~·irro [85]. Pre-treatment
of canine corneocytes with concanavalin A (10 f.lg m]- 1)
inhibited (P < 0·01) the adherence of one out of six M.
pachydermatis strains. This effect was abrogated by pre-
incubating concanavalin A with its hapten inhibitor (6%
methyl a-D-mannopyranoside), suggesting that manno-
syl-bearing carbohydrate residues on the epithelial cells
serve as ligands for adhesins expressed by this strain.
Howevec treatment of corneocytes with soybean. wheat
germ and gorse lectin, and pre-treatment of the yeast cells
with either of the four lectins had no reproducible effect
on the adherence of two strains.
have larger numbers of the yeast on smears of cerumen.
Subsequent studies have either confirmed Fraser's find-
ings [86], or have reported an increased prevalence of the
yeast in otitis externa [70. 71]. Studies in which cases of
otitis externa in dogs responded to antifungal therapy
have been described [69,87 -89], and current opinion sug-
gests that the yeast acts as an opportunistic secondary
pathogen within the ear canal, under circumstances
which remain poorly defined. Otitis externa associated
with IV!. pachydermatis is often characterized by a waxy,
moist, brown or yellow exudate with variable erythema
and pruritus [90].
More recently, interest in ivl. pachydermatis has in-·
creased with the recognition that the organism may be
recovered from dogs with more generalized skin disease.
The complete change in opinion on the importance of
'Jv!alassezia dermatitis' in canine dermatology is illus-
trated by a review of the recent editions of the standard
text, 'Small Anima! Dermatology'. Skin disease associ-
ated with ivf. pachydermatis, other than otitis externa. is
not mentioned in the third edition [91]. In the fourth
edition [92], IV/alassezia dermatitis is almost dismissed in
the space of eight lines, whereas five pages are devoted to
it in the fifth edition [93].
Dufait [94] is credited with the first reports of M.
pachydermatis as a cause of more widespread dermatitis
in the dog. He described a series of 50 dogs with pruritic
dermatitis, from which the yeast could be readily recov-
ered by cytology or culture, and which responded to
antifungal therapy. Skin lesions consisted of erythema
and hyperpigmentation which most often affected the
ventral abdomen, although the face, feet and perineal
regions were also commonly affected. Larsson et a!. [95]
described similar findings in 23 cases from Brazil. Scott &
Miller [96] described eight West Highland White terriers
with a severely pruritic seborrhoeic skin disorder from
which M. pachydernzatis could be isolated. Mason &
Evans [97] described II cases in detail and demonstrated
the yeast in the stratum corneum in all nine cases from
which skin biopsies were obtained. Concurrent allergic
diseases were recognized in four dogs. and one dog had
@ 1999 ISHAM. Medical fv!yco!ogy. 31, 295-306

hepatic disease. In each case, antibacterial therapy was
without benefit, but all dogs responded to either systemic
ketoconazole or topical antifungal therapy. This publica-
tion and other presentations by Mason in the early 1990s
[97-99] were instrumental in highlighting widely the po-
tential importance of this yeast to members of the veteri-
nary dermatology community, many of whom were
unfamiliar with, or had dismissed, the earlier reports by
Dufait [94].
Further evidence for a pathogenic role of M. pachyder-
matis was provided by a double-blind study in which the
clinical responses in 33 basset hounds with Malassezia-
associated seborrhoeic dermatitis were related to skin
microbial populations, before and after treatment with
either miconazole-chlorhexidine or selenium sulphide
shampoos [100]. At the first assessment, M. pachyderma-
tis populations in affected skin, determined using a deter-
gent-scrub technique, were typically 100-1 0000-fold
higher than those of healthy dogs. Marked increases in
skin populations of coagulase-positive staphylococci,
principally Staphylococcus intermedius, were also demon-
strated. The miconazole-chlorhexidine treated dogs had
superior clinical responses and lower post-treatment
counts of M. pachydermatis and coagulase-positive
staphylococci, suggesting that skin infection with M.
pachydermatis and/or bacteria, played an important role
in the development of dermatitis in those basset hounds.
However, the evidence for a pathogenic role for the
yeast remains circumstantial. Some of the antifungal
agents used in the studies described previously have other
effects. For example, ketoconazole has been shown to
have anti-inflammatory properties [99, 101]. Concurrent
colonization or infection of lesional skin by S. inter-
medius is common [73,97]. The miconazole-chlorohexi-
dine shampoo has substantial antibacterial activity [1 00]
and it is difficult to determine the relative importance of
the reductions in yeast and bacterial populations. How-
ever, antibacterial therapy alone has previously been re-
ported to have little effect in such cases [97] and there is
some evidence to suggest that the best therapeutic re-
sponse is achieved when both M. pachydermatis and
bacteria are removed by topical therapy [102].
A recent study suggested that increased corneocyte
receptivity for the yeast is not important in the pathogen-
esis of seborrhoeic dermatitis associated with M. pachy-
dermatis in basset hounds [103]. Similar results were
obtained by Bergbrant & Faergemann [82] who found
that M. furfur was better able to adhere to stratum
corneum cells from healthy individuals than to cells from
patients with seborrhoeic dermatitis. However, Schecht-
man et a!. [83] reported that there was no relationship
between the severity of seborrhoeic dermatitis of humans
© 1999 ISHAM. Medical Mycology. 37. 295-306
Malassezia pachydermatis: a review
and the adherence of M. furfur to human keratinocytes in
vitro.
The factors which favour proliferation of M. pachyder-
matis and its transition from a commensal organism to
an apparent pathogen on canine skin are poorly under-
stood, but presumably reflect disturbances of the normal
physical, chemical or immunological mechanisms which
restrict microbial colonization of skin. Breed predilec-
tions have been identified, although geographical varia-
tions are apparent. Basset hounds, dachshunds, cocker
spaniels, West Highland White terriers, poodles and Aus-
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tralian Silky terriers have been reported to be at in-
creased risk [75,76,94,98]. There is no gender predilection
and Malassezia-associated skin diseases may develop in
both young and old dogs. Plant et a!. [76] reported that
recent antibacterial therapy and a clinical presentation
consistent with seborrhoeic dermatitis were associated
with high counts of M. pachydermatis. Concurrent dis-
eases have been recognized in many, but not all, cases of
Malassezia dermatitis in dogs, and correction of the
concurrent disease may prevent, or reduce the frequency
of relapse in some dogs. Some atopic dogs carry large
numbers of M. pachydermatis yeasts in both lesional skin
and in unaffected areas [104,105]. Immediate intradermal
test reactivity has been observed in atopic dogs following
the injection of M. pachydermatis extracts at concentra-
tions which cause no reaction in healthy dogs [106],
suggesting that hypersensitivity responses to yeast aller-
gens may be involved in the pathogenesis in some cases
of atopic disease. These observations are analogous to
the suggestion that IgE-mediated hypersensitivity to
Malassezia-derived allergens may be important in the
pathogenesis of the 'head-neck' form of atopic dermati-
tis in humans [107].
Diagnostic criteria for Malassezia dermatitis in the dog
have not been firmly established. It has been suggested
that a diagnosis of Malasse::ia dermatitis is appropriate,
based on current knowledge, when a dog with elevated
M. pachydermatis populations on lesional skin shows a
good clinical and mycological response to appropriate
antifungal therapy [108]. Cytological, cultural and histo-
pathological techniques can be used to identify the yeast.
Cytological techniques used include impression methods
using glass slides [76,96], cotton swabs [98], skin scrap-
ings [94] and tape-strip preparations [98]. Although cytol-
ogy is one of the most useful methods in practice, it is not
clear which of these te~hniques are most efficient. Con-
tact-plate cultural techniques provide a rapid and conve-
nient method for isolation of the yeast but the results are
not immediately available [104]. This quantitative
method is superior to standard swabbing methods be-
cause population sizes can be determined, and in most
 Guiliot & Bor;d
canine cases, population densities in affected areas
greatly exceed those of healthy skin [73,77,103,104]. Al-
though /vf. pachydermatis is not lipid-dependent, the use
of lipid-supplemented media, especially the modified
Dixon's medium, was shown to be very advantageous for
quantitative culture of the yeasts from canine skin [109].
Both incubation temperatures, 27 and 37 °C, were then
equally suitable. The Sabouraud glucose agar seemed to
be less efficient, especially when incubated at 27 oc. En-
hanced growth on that medium could be obtained with a
37 oc incubation or in a carbondioxide-enriched atmo-
sphere [109-111].
Histopathological examination of skin biopsy speci-
mens typically shows parakeratotic hyperkeratosis, irreg-
ular epidermal hyperplasia, intercellular oedema and
lymphocytic superficial perivascular or interstitial der-
matitis [97, 112]. However, iV!. pachydermatis is not al-
ways visible in the epidermal stratum corneum, even in
cases where large numbers are seen cytologically [78, 1!2];
this probably reflects disruption of this layer during
processing.
Therapeutic options for canine Malasse::.ia dermatitis
include systemic therapy with k€toconazole or itracona-
zole [97,99] or topical therapy with a variety of agents,
especially azole derivatives [113]. A 2'% miconazole-2'Yt,
chlorhexidine product has been shown to have good
efficacy (100], but selenium sulphide, chlorhexidine, enil-
conazole and ketoconazole-containing products are also
helpful in some cases [98,93]. Identification and correc-
tion of underlying causes should be attempted in re!aps-
mg cases.
Dermatitis and otitis in other animals
M. pachydermatis appears to be a relatively infrequent
pathogen in cats, at least when compared with dogs. This
may reflect the less frequent carriage rates seen in healthy
cats [57,65,66, 72,86, 114]. Ceruminous otitis externa re-
sponsive to antifungal ear drops is the commonest clini-
cal presentation of i14alasse.::ia-associated skin diseases in
cats. Occasional cases of localized or generalized
Malasse.::ia dermatitis have been described in cats
[99, 115]. Exfoliative erythroderma, greasy exudation and
varying degrees of pruritus may be seen. Diagnostic and
therapeutic measures are similar to those used in dogs.
At. pachydermatis-associated otitis externa have also
been described in ferrets, fennecs, pigs and dromedaries
[3,57, 116, 117]. An exfoliative Malasse:ia dermatitis was
first reported in 1925 in an Indian rhinoceros from
Philadelphia [1]. More recently similar lesions were ob-
served in a southern white rhinoceros [59]. Jv!. pachyder-
matis was held responsible for a case of focal dermatitis
in a California sea lion (Zalophus cahfornianus). The
animal developed multiple wheals on both flanks and
chest [61]. The excellent response to specific antifungal
therapy suggested that the yeasts were opportunistic
pathogens.
Human infections
A case of canaliculi tis was reported in a 61-year-old man
[118]. The yeasts was also isolated from a cutaneous
wound of a 67-year-old man and from the urine of a
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patient with chronic granulomatous disease [16]. How-
ever, the most commonly described human infection due
to lvl, pachydennatis is an intravascular catheter-acquired
sepsis [5]. Usually, patients are premature infants with
multiple complications of prematurity. They require hos-
pitalization in a neonatal intensive care unit and have
been receiving broad spectrum antibiotics and parenteral
lipid emulsions for several weeks through a central
venous catheter [5,8, 1I 9]. Signs of illness may be absent
[8]. When observed, symptoms are similar to those asso-
ciated with fungemia due to M. fwjitr [8, 119]. Fever,
lethargy, respiratory distress or bradycardia are usually
reported. Multiple anatomic sites can be sources of lvf.
pachydermatis [ 120,121]. Removal of the infected catheter
seems to be the most successful mode of treatment [5].
Genetic typing of clinical isolates clearly demonstrated
the nosocomial nature of epidemics caused by !Vi. pachy-
dermatis in neonatal intensive care units [22]. All isolates
recovered from both patients and incubator surfaces were
genetically comparable. Furthermore, despite regular
cleaning of the incubators, yeasts were shown to persist
on the glass surfaces at least for 2 months. Somerville
[ 122] reported that 16'% of patients with chronic cuta-
neous diseases harboured 1\I. pachydermatis on the skin.
These findings were not confirmed by other investigators
who concluded that, in most cases, the presence of lvf.
pachydermatis on human skin was rare and transient
[6,68, 123, 124]. Mickelsen et a!. [8] suggested that im-
munocompromised patients might coru,titute suitable
hosts for M. pachydermatis. Transmission would be most
likely from animal skin (pets) [7] or environmental
sources [22]. With partial LSU rRNA sequencing, all the
strains recovered from humans were classified in sequevar
'I a', i.e. the most frequent sequence-type of the strains
isolated from dogs [18]. In a recent study concerning a
epidemic caused by iV!. pachydermatis in an intensive care
nursery, isolates from patients, health care workers and
health care workers' pets had identical pattern of RFLP
[7]. The introduction and subsequent nosocomial trans-
mission of pet associated Jvlalasse::;ia yeasts in nursery
could be prevented by careful hand washing by health
care workers.
© 1999 ISHAI'-'1. i"/ed!cJI Mycology, 37. 295 ·- 306

Acknowledgements
The authors gratefully acknowledge the invaluable advice
and constant help provided by Eveline Gueho from the
Pasteur Institute of Paris.
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