Dermatophytes from cases of skin disease in cats and dogs in Turin, Italy : Dermatophyten als Erreger von Hautkrankheiten bei Katzen und Hunden in Turin, Italien


V. Filipello, Marchisio; M. G. Gallo; V. Tullio; S. Nepote; A. Piscozzi; C. Cassinelli

Mycoses 38(5-6): 239-244

1995


MYCOSES
38,
239-244
(1995)
ACCEPTED:
JANUARY
2,
1995
Dermatophytes
from
cases
of
skin
disease
in
cats
and
dogs
in
Turin,
Italy
Dermatophyten
als
Erreger
von
Hautkrankheiten
bei
Katzen
und
Hunden
in
Turin,
Italien
V.
Filipello
Marchisio
1
,
M.
G.
Gallo
2
,
V.
Tullio',
S.
Nepote
l
,
A.
Piscozzi'
and
C.
Cassinelli'
Key
words.
Microsporum
canis,
dermatophytes,
dermatophytosis,
cats,
dogs,
Italy.
Schliisselworter.
Microsporum
canis,
Dermatophyten,
Dermatophytosen,
Katze,
Hund,
Italien.
Summary.
In
urban
environments,
pet
animals
such
as
dogs
and
cats
may
be
largely
responsible
for
the
progressive
increase
of
zoophilic
over
anthropophilic
dermatophytes
in
the
aetiology
of
human
dermatophytoses.
Mapping,
in
each
city,
of
the
natural
foci
of
zoophilic
dermatophytes
may
thus
be
important
for
understanding
the
epidemi-
ology
of
human
dermatophytoses,
and
for
planning
preventive
measures.
We
have
surveyed
dogs
and
cats
suspected
of
carrying
dermatophytic
lesions
in
the
Turin
area.
Hairs,
skin
scrapings
and
the
contents
of
pustules
or
vesicles
were
inoculated
on
Dermasel
agar
supplemented
with
chloramphen-
icol
and
cycloheximide.
The
mating
type
of
the
isolates
was
checked,
testing
their
compatibility
with
the
two
mating
types
of
Arthroderma
simii
Stockdale,
Mackenzie
and
Austwick.
We
isolated
dermatophytes
from
about
40%
of
the
cases
exam-
ined.
Microsporum
canis
Bodin
was
the
only
dermato-
phyte
found
in
the
lesions.
In
the
cats
the
lesions
were
more
frequent,
in
the
dogs
more
severe.
Animals
less
than
a
year
old
and
male
dogs
were
most
often
affected.
Some
cases
of
transmission
of
the
infection
between
animals
and
from
animals
to
man
are
described.
All
the
isolates
of
M.
canis,
tested
for
mating
behaviour,
were
non-reactive.
Zusammenfassung.
In
stadtischer
Umgebung
konnten
Haustiere
wie
Hunde
und
Katzen
weitge-
'Dipartimento
di
Biologia
vegetale
dell'Universita
di
Torino,
2
Dipartimento
di
Produzioni
Animali,
Epidemiologia
ed
Ecologia
dell'Universita
di
Torino,
and
'Istituto
di
Microbiologia
dell'Universita
di
Torino,
Italy.
Correspondence:
Prof.
Dr
V.
Filipello
Marchisio,
Dipartimento
di
Biologia
vegetale
dell'Universita,
viale
P
A
Mattioli
25,
1-10125
Turin,
Italy.
hend
fur
die
zunehmende
Uberhandnahme
von
zoophilen
gegenuber
antropophilen
Dermatophy-
ten
in
der
Atiologie
menschlicher
Dermatophyto-
sen
verantwortlich
sein.
Die
Kartierung
der
Haufigkeitsschwerpunkte
zoophiler
Dermatophy-
ten
in
jeder
Stadt
konnte
daher
fur
das
Verstandnis
der
Epidemiologie
der
Dermatophytosen
beim
Menschen
und
fur
das
Planen
praventiver
MaBnahmen
wichtig
sein.
Haare,
Hautabschabsel
und
Inhalte
von
Pusteln
oder
Blaschen
wurden
auf
Dermasel-Agar
mit
Chloramphenicol
und
Cycloheximid
angelegt.
Der
Paarungstyp
der
Iso-
late
wurde
bestimmt,
indem
ihre
Kompatibilitat
mit
den
zwei
Paarungstypen
von
Arthroderma
simii
Stockdale,
Mackenzie
und
Austwick
getestet
wurde.
Wir
isolierten
Dermatophyten
von
unge-
fahr
40%
der
untersuchten
Proben.
Microsporum
canis
war
der
einzige
in
den
Lasionen
gefundene
Dermatophyt.
Bei
Katzen
waren
Lasionen
haufi-
ger,
bei
Hunden
waren
diese
schwerer.
Tiere,
die
junger
als
eM
Jahr
alt,
und
Mannchen
bei
den
Hunden
waren
am
meisten
betroffen.
Einige
Falle
von
Ubertragung
der
Infektion
von
Tier
zu
Tier
und
von
Tier
zu
Mensch
werden
beschrieben.
Alle
Isolate
von
M.
canis
waren
in
den
Paarungstesten
nicht
reaktiv.
Introduction
In
both
evolutionary
and
developmental
senses,
a
complex
epidemiological
chain
connects
geophilic,
zoophilic
and
anthropophilic
dermatophytes.
Species
distributions
and
disease
profiles
can
change
over
the
years
and
in
different
countries
as
a
result
of
the
interaction
of
different
factors.
240
V.
FILIPELLO
MARCHISIO
ET
AL.
Population
shifts
and
changes
in
health
habits,
living
standards,
travel
and
clothing
fashions
all
affect
the
epidemiology
and
distribution
of
der-
matophytes
[1].
The
last
several
decades
have
seen,
except
in
the
United
States,
a
worldwide
progressive
increase
in
zoophilic
over
anthropo-
philic
dermatophytes
in
the
aetiology
of
human
dermatophytoses
[1,
2].
In
urban
environments,
pet
animals
such
as
dogs
and
cats
may
be
largely
responsible
for
this
phenomenon,
as
overt
or
symptomless
carriers.
During
the
evolution
of
soil
keratinolytic
species,
and
those
that
specialize
in
the
invasion
and
colonization
of
animal
cornified
tissue,
and
even
confine
themselves
to
one
host
[3,
4],
there
may
have
been
zoophilic
species
able
to
attack
a
wide
spectrum
of
animal
hosts.
Microsporum
canis,
for
example,
can
parasitize
many
wild
and
domestic
animals
[5]
and,
increasingly,
man.
It
seems
that
at
present
tinea
capitis
and
tinea
corporis
in
urban
areas
are
almost
exclusively
due
to
M
canis
[1].
The
organism
has
its
main
reservoir
in
cats
and
dogs,
and
often
causes
lifelong
chronic
infection
in
these
hosts
[1].
It
is
firmly
established
that
the
increased
incidence
of
M.
canis
infections
in
man
is
the
result
of
persistence
in
animal
reservoirs.
In
fact,
in
the
absence
of
such
reservoirs,
it
seems
that
the
fungus
cannot
spread
widely
in
human
populations;
it
loses
its
virulence
after
about
four
passages
between
humans,
and
the
infection
therefore
requires
renewed
contact
with
an
animal
host
to
maintain
virulence
[1].
Mapping,
in
each
city,
of
the
natural
foci
of
zoophilic
dermatophytes
may
thus
be
important
for
understanding
the
epidemiology
of
human
dermatophytoses,
and
for
planning
preventive
measures.
In
this
research
we
have
surveyed
dogs
and
cats
suspected
of
carrying
dermatophytic
lesions
in
the
Turin
area.
This
type
of
survey
has
not
previously
been
done
in
Turin.
Materials
and
methods
The
work
was
done
in
collaboration
with
the
Medical
Clinic
of
the
Veterinary
Faculty,
University
of
Turin,
and
with
the
help
of
several
private
veterinary
clinics.
Over
a
period
of
1
year
203
animals
(98
dogs
and
105
cats),
with
suspected
dermatophyte
lesions
were
examined.
The
modest
number
of
cases
that
we
have
been
able
to
examine
during
a
year
may
be
because
of
the
common
tendency
to
underesti-
mate
the
importance
of
the
lesions
and
failure
to
consult
a
vet.
Samples
taken
from
lesions
(hairs,
skin
scrap-
ings,
contents
of
pustules
or
vesicles)
were
inocu-
lated
on
9-cm
Petri
dishes
containing
Dermasel
(Oxoid)
agar
supplemented
with
50
mg
1
-1
chlor-
amphenicol
and
0.4
g
1
-I
cycloheximide
(Oxoid
antibiotic
supplement
for
Dermasel
agar,
SR
75),
to
limit
the
growth
of
bacteria
and
saprophytic
fungi.
If
the
lesions
had
been
treated
with
antimy-
cotics
or
were
covered
in
pus
or
other
material,
they
were
first
carefully
washed
with
soap
and
water.
Three
inoculations
were
made
from
each
lesion
in
the
same
Petri
dish,
even
when
an
animal
had
several
lesions
on
different
parts
of
the
body.
The
dishes
were
then
maintained
at
24
°C
for
15
days,
followed
by
a
further
15
days
at
room
temperature,
during
which
time
they
were
observed.
Each
developing
colony
was
isolated
in
pure
culture
on
the
same
medium.
To
induce
fructifi-
cation,
Sabouraud
glucose
agar
was
used,
with
the
addition
of
yeast
extract
(5
g
1
-1
),
or
Borelli
Lactritmel
agar
[6].
The
age,
sex,
breed,
place
of
origin,
previous
history,
habits
and
the
type
of
lesion
were
recorded
for
each
animal.
The
presence
of
fungal
infections
among
members
of
the
family
and
other
animals
possibly
present
in
the
household
was
also
noted.
These
data
were
entered
on
forms
specially
pre-
pared
for
the
vets.
However,
as
not
all
samples
were
taken
in
the
presence
of
one
of
the
authors,
a
complete
set
of
data
was
not
available
in
some
cases.
This
disappointing
level
of
collaboration
rendered
difficult
the
development
of
certain
potentially
useful
epidemiological
correlations,
and
may
have
compromised
the
validity
of
others.
For
this
reason,
some
of
the
results
should
be
taken
only
as
indicative.
Because
of
its
epidemiological
importance,
we
also
wished
to
establish
the
mating
type
of
the
isolates,
testing
their
compatibility
with
the
two
mating
types
of
Arthroderma
simii
Stockdale,
Mackenzie
and
Austwick,
using
the
method
of
Stockdale
[7].
The
A.
simii
strains
were
from
the
Centraalbureau
voor
Schimmelcultures,
Baarn,
The
Netherlands.
The
mycelial
colours
are
described
according
to
Rayner
[8].
Results
We
isolated
dermatophytes
from
about
40%
of
the
cases
examined
(Table
1).
This
percentage
might
be
slightly
influenced
by
the
fact
that
1.8%
of
the
animals
had
been
treated
topically
or
orally
with
antimycotics.
Microsporum
canis
Bodin
was
the
only
dermato-
phyte
found
in
the
lesions.
It
developed
in
pure
culture
in
80%
of
the
dishes.
M.
canis
is
an
mycoses
38,
239-244
(1995)
DERMATOPHYTES
IN
CATS
AND
DOGS
IN
TURIN
241
Table
1.
Incidence
of
dermatophytic
lesions
in
the
cats
and
dogs
examined
Samples
examined
Positives
n
%
Cats
105
53
Dogs
98
29
Total
203
82
64.6*
35.4*
40.4f
*
Calculated
on
the
total
of
positives.
f
Calculated
on
the
total
of
samples
examined.
organism
that
appears
to
be
in
an
active
state
of
evolution,
and
strains
associated
with
particular
animal
species
are
known
[1].
We
therefore
note
that
we
found
two
colony
variants
during
this
work,
although
these
did
not
correlate
with
the
type
of
host.
One
variant
possessed
a
prostrate
aerial
mycelium,
adherent
to
the
medium,
with
a
matted
appearance,
and
was
white
to
straw
col-
oured
(colour
46
of
Rayner)
or
cinnamon
(colour
62);
the
underside
was
orange
(colour
7),
tending
to
umber
(colour
9).
The
other
variant
produced
abundant
cottony
erect
aerial
mycelium
coloured
white
to
buff
(colour
45)
or
ochreous
(colour
44);
the
underside
varied
from
colourless
to
straw
(colour
46)
or
orange
(colour
7).
Microscopic-
ally,
the
two
variants
were
indistinguishable.
The
macroconidia
were
variable
in
size:
8.4-19.6
x
50.4-84
gm.
They
were
cigar-shaped,
fusiform
or
ovate,
sometimes
with
one
or
two
curved
beaks.
Lactritmel
agar
was
able
to
induce
macroconidium
formation
in
two-thirds
of
the
sterile
samples.
Microconidia
were
formed
in
only
34%
of
the
isolates;
their
production
was
increased
on
Sabouraud
glucose
agar
plus
yeast
extract.
Arthroconidia
were
produced
more
often
(53%
of
isolates)
and
were
very
variable
in
size
and
shape.
Tables
1-5
show
the
incidence
of
dermatophytic
lesions
in
the
cats
and
dogs,
in
relation
to
age
and
sex.
In
the
cats,
the
lesions
were
more
frequent
(64.6%)
(Table
1).
Animals
less
than
a
year
old
were
most
affected
(Table
2).
In
cats,
no
important
differences
appeared
to
be
correlated
with
sex
(Table
4).
Among
the
dogs,
however,
males
were
more
affected
(Table
5),
and,
as
before,
the
puppies
(Table
3).
In
cats,
the
extent
of
the
lesions
was
generally
modest
and
they
sometimes
went
unnoticed.
Lesions
mostly
appeared
as
simple
dandruff
des-
quamations
with
little
or
no
hair
loss.
They
were
normally
limited
to
the
paws,
the
tail
and
the
head,
as
a
result
of
the
habit
cats
have
of
sleeping
curled
up.
Only
in
some
4-month-old
abandoned
kittens
in
generally
poor
condition
were
the
lesions
widespread
over
the
whole
body.
Table
2.
Relation
between
incidence
and
age
in
cats
Samples
examined
Positives
n
%
Young*
40
31
58.5f
Adults
38
14
26.4f
Unknown
27
8
15.1f
Total
105
53
50.5:
*
Under
1
year
of
age.
t
Calculated
on
the
total
of
positives.
:.
Calculated
on
the
total
of
samples
examined.
Table
3.
Relation
between
incidence
and
age
in
dogs
Samples
examined
Positives
n
%
Young*
27
16
55.2f
Adults
44
10
34.5f
Unknown
27
3
10.3f
Total
98
29
29.6:
*
Under
1
year
of
age.
t
Calculated
on
the
total
of
positives.
1:
Calculated
on
the
total
of
samples
examined.
Table
4.
Distribution
of
lesions
in
cats
between
females
and
males
Samples
examined
Positives
n
%
Females
42
23
43.4*
Males
41
24
45.3*
Unknown
22
6
11.3*
Total
105
53
50.5f
*
Calculated
on
the
total
of
positives.
f
Calculated
on
the
total
of
samples
examined.
Table
5.
Distribution
of
lesions
in
dogs
between
females
and
males
Samples
examined
Positives
n
%
Females
19
3
10.3*
Males
52
18
62.1*
Unknown
27
8
27.6*
Total
98
29
29.6f
*
Calculated
on
the
total
of
positives.
f
Calculated
on
the
total
of
samples
examined.
mycoses
38,
239-244
(1995)
242
V.
FILIPELLO
MARCHISIO
ET
AL.
In
dogs
the
lesions
were
more
severe,
and
showed
more
or
less
extensive
loss
of
hair;
in
puppies,
the
lesions
were
often
suppurating.
The
site
of
the
lesions
was
variable,
but
the
areas
most
often
affected
were
the
head,
followed,
in
order,
by
the
paws,
the
ribs
and
inguinal
areas.
However,
one
15-day-old
Yorkshire
terrier
puppy
had
lesions
over
the
whole
body.
The
infection
was
trans-
mitted
from
the
mother.
The
most
severe
infection
was
seen
in
one
puppy
of
5
months,
bought
in
a
specialist
shop.
At
3
months
the
puppy
had
suffered
from
bronchitis
and
lack
of
vitamin A.
The
first
fungal
lesions
appeared
a
month
later,
with
a
favic
aspect.
Mycological
tests
always
revealed
M.
canis
in
association
with
Chaetomium
globosum
Kunze
ex
Steudel.
Our
data
demonstrated
transmission
of
infection
between
animals
in
only
one
case:
from
an
adopted
stray
cat
to
a
healthy
family
cat.
This
occurred
about
20
days
after
adoption
of
the
stray,
which
had
lesions
on
its
ears.
M.
canis
with
identical
microscopic
characteristics
was
isolated
from
both
cats.
We
should
add
that
most
of
the
cats
that
tested
positive
lived
with
other
cats
or
were
allowed
out
and
could
come
into
contact
with
other
ani-
mals.
Sanitary
inspection
of
these
animals
was
not
possible.
A
further
and
important
source
of
infec-
tion
is
grooming
in
specialized
shops.
This
is
a
frequent
cause
of
infection,
especially
in
cats
with
long
fur,
such
as
Persians.
Transmission
from
animals
to
man,
clearly
occurred
only
in
the
case
of
the
4-month-old
stray
kittens
that
had
lesions
all
over
their
bodies.
The
infection
was
rapidly
transmitted
to
the
mother
of
the
family,
to
her
children
and
to
their
friends,
who
tested
positive
for
M.
canis
(examination
car-
ried
out
at
the
Dermatology
Clinic,
University
of
Turin).
In
13
other
cases,
we
noted
the
simul-
taneous
occurrence
of
lesions
on
domestic
animals
(11
cats
and
two
dogs)
and
in
certain
members
of
the
family,
although
we
could
not
prove
any
connection
since
the
dermatologist
did
not
perform
a
mycological
examination.
Transmission
from
man
to
animals
remains
unproven,
though
possible,
according
to
Warner
[9].
A
woman
presented
with
tinea
corporis
localized
to
the
forearm
and
the
back
of
the
hand
before
the
arrival
of
a
cat
in
the
house.
After
a
short
time
the
cat
developed
patchy
loss
of
hair.
M.
canis
was
found
to
be
responsible
for
both
conditions.
On
the
other
hand,
in
four
cases
lesions
in
humans
were
not
accompanied
by
infection
in
the
family
pet
(three
cats
and
a
dog).
Our
data
showed
an
increase
in
infections
in
May
to
June
and
in
July
to
August,
and
a
further
peak
in
November
(Table
6).
There
appeared
to
Table
6.
Annual
incidence
of
lesions
in
cats
and
dogs
Samples
examined
Positives
n
%*
n
%**
December
22
10.8
9
11.0
January
22
10.8
4
4.9
February
18
8.9
6
7.3
March
23
11.3
8
9.8
April
17
8.4
3
3.7
May
19
9.4
9
11.0
June
21
10.3
10
12.2
July
20
9.9
14
17.1
August
8
3.9
7
8.5
September
11
5.4
3
3.7
October
10
4.9
3
3.7
November
12
5.9
6
7.3
*
Calculated
on
the
total
of
samples
examined.
t
Calculated
on
the
total
of
positives.
be
no
correlation
with
diet,
or
with
histories
of
vaccination
or
other
treatment.
None
of
the
isolates
showed
any
sign
of
reactivity
(proliferation
of
white
cottony
mycelia
or
forma-
tion
of
ascocarp
initials)
when
brought
into
contact
with
either
mating
type
of
A.
simii.
On
the
contrary,
the
two
species
seemed
to
be
antagonistic.
Discussion
Our
results
are
generally
in
line
with
those
pub-
lished
over
the
last
40
years.
They
confirm
that
M.
canis
is
the
principal
cause
of
dermatophytosis
in
cats
and
dogs
[9-23].
In
our
work
M.
canis
was
the
only
dermatophyte
species
responsible
for
skin
lesions
in
cats
and
dogs;
the
same
result
was
obtained
in
New
Zealand
[13],
and,
in
cats
only,
in
Denmark
[16],
Australia
[14]
and
the
midwest-
ern
United
States
[22].
Other
research
has
shown
that
different
dermatophyte
species
can
cause
this
type
of
lesion,
in
particular
Trichophyton
mentagro-
phytes
and
M.
gypseum
[9,
10,
12,
16,
17-23].
These,
however,
play
a
secondary
role,
being
implicated
in
only
small
numbers
of
cases.
Also
the
wider
occurrence
of
dermatophytic
lesions
in
cats
[13,
14,
16,
18],
in
the
young
of
both
cats
and
dogs
[13,
24]
and
the
higher
viru-
lence
in
dogs
[16,
18],
observed
during
our
work,
have
previously
been
noted
in
the
literature.
Higher
virulence
in
dogs
could
be
a
consequence
of
the
lower
level
of
specialization
of
the
parasite;
the
more
virulence
displayed
towards
a
given
host,
the
less
specific
is
the
parasite
[25-32].
We
again
confirmed,
however,
the
high
specificity
of
M.
canis
for
cats.
As
in
other
parts
of
the
world
[2,
33-37],
in
mycoses
38,
239-244
(1995)
DERMATOPHYTES
IN
CATS
AND
DOGS
IN
TURIN
243
Turin
[38]
cats
proved
quite
frequently
to
be
healthy
carriers
of
M.
canis.
Moreover,
from
work
done
in
1982
[39]
and
other
more
recent
research
not
yet
published,
we
can
state
that
M.
canis
is,
in
Turin,
the
main
cause
of
human
dermatophytoses.
Bearing
in
mind
the
finding
of
Rippon
[1]
that
M.
canis
becomes
less
virulent
after
about
four
human-to-human
passages,
and
the
consequent
need
for
renewed
contact
with
animal
hosts
for
the
spread
of
infection,
the
epidemiological
links
between
pets
and
the
human
population
in
Turin
seem
clear.
However,
to
demonstrate
such
links
on
a
wide
scale
would
require
a
programme
of
strict
collaboration
between
dermatologists
and
vets;
this
would
involve
checks
on
both
pets
and
humans
every
time
either
presented
cases
of
dermatophytosis.
At
present,
strain
identification
is
not
specific
enough
to
allow
reliable
identifi-
cation
of
the
source
[40].
Epidemiological
links
between
domestic,
stray
or
wild
cats
and
dogs
and
man
could
also
be
sought
by
using
a
regular
and
obligatory
checking
system.
One
could
also
in
this
way
obtain
better
data
on
the
seasonal
distribution
of
infections.
The
increase
in
late
spring,
summer
and
autumn
sug-
gested
by
our
data,
albeit
on
the
basis
of
very
few
cases,
could
be
related
to
the
seasonal
birth
of
kittens
[34].
According
to
McAleer
[14]
young
kittens
are
probably
the
main
source
of
infection.
However,
the
findings
of
other
studies
do
not
entirely
agree
with
this
theory.
According
to
Ainsworth
&
Austwick
[10],
Kaplan
&
Ivens
[41],
McAleer
[14],
Kristensen
&
Krogh
[16]
and
Stenwig
[18],
from
the
UK,
USA,
Australia,
Denmark
and
Norway
respectively,
the
peak
inci-
dence
is
in
the
autumn,
whereas
Baxter
[13]
did
not
find
any
significant
seasonal
variation
in
New
Zealand.
The
ratio
of
positive
cases
in
relation
to
the
total
number
of
suspected
cases
of
dermatophytosis
is
also
very
variable,
as
reported
in
the
literature.
Our
value
of
40%
is
similar
to
the
findings
of
Mantovani
&
Morganti
[12],
Baxter
[13],
Kristensen
and
Krogh
[16],
van
Cutsem
et
al.
[17],
Stenwig
[18]
and
Sparkes
et
al.
[23].
However,
methodologies
have
sometimes
differed
and
com-
parisons
are
therefore
of
little
value.
In
any
case,
the
almost
constant
low
incidence
emphasizes
the
difficulties
of
immediate
diagnosis
of
dermato-
phytic
lesions
and
the
need
to
perform
a
mycologi-
cal
examination
every
time,
in
order
to
determine
appropriate
treatment.
A
final
comment
concerns
the
mating
behaviour
of
our
isolates.
The
complete
absence
of
reaction
could
be
a
real
characteristic.
In
fact,
Weitzman
&
Padhye
[42],
Takatori
&
Hasegawa
[43]
and
Kubo
et
al.
[44]
all
found
that
a
significant
percent-
age
of
isolates
of
M.
canis
were
non-reactive.
Nevertheless,
the
problem
could
lie
with
the
strains
of
A.
simii
that
we
used,
because
Takashio
[45]
and
Weitzman
&
Padhye
[42]
reported
that
the
sexual
power
of
some
human
and
animal
dermato-
phyte
isolates
can
degenerate
over
a
period
of
time
if
they
are
subcultured.
Up
until
now
the
majority
of
M.
canis
isolates
tested
for
mating
behaviour
have
proved
to
be
negative
or
non-
reactive;
the
only
positive
isolates
have
been
detected
in
Japan
[44].
Acknowledgement
This
work
was
supported
by
MURST
60%.
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