From www.bloodjournal.org by guest on June 18, 2017. For personal use only.
Basophil/Mast
Cell
By
Semisolid
the
(methylcellulose)
presence
derived
blood.
cells
in
strated
individual
cells
at various
(BMC)
microscopy.
to
have
pure
the
“eosinophil”
macrophage”
I).
constituted
granulocyte
colonies
systemic
colonies
cultures
of
from
and
(13/67)
may
help
the
fully
known,
there
is ultrastructural
from a common
for a substitution
across
species.’#{176} Mast
cell
tants
precursors
peripheral
blood,
of these
precursors
with systemic
mastocytosis
leukemia
(CML).
normals)
using
with
streptomycin
gradient
peripheral
SM,
in 0.9%
Iscove’s
ratio
figure
human
volume
4 with
venous
urticaria
methylcellulose
modified
(GIBCO,
Long
(GIBCO),
5 x
l05M
Blood.
Vol. 61,
No. 4 (April),
Island,
cultured
8 with
as previously
1%
v/v
N.Y.),
20%
v/v
(final
120
pg in
removed
These
glass
was
as
slides
like
previously
by an
or into
I 00 cl
(see below).
1:1-1:2
previously
colonies
in all cultures,
noted
picked
to a lower
was,
a
in analyses
100 jil).
Histamine
limit
of
100 iI
PBS
previously
of I 00 pg. Since
80-90
of
( I 00 pg in 80
in these
at 37#{176}C
with
the
zl, a cut-off
colony
assayed
I hr incubation
into
technique
practically,
a histamine-positive
cultures
diamine
was
oxidase
0.5 U/mI.
Stains
performed
using
Alcian
blue,
the Host
blue
purportedly
in part
the
after
following
for
Program,
University,
by a grant
procedures:
fixation
specific
Resistance
McMaster
in lead
mast
cells7;
Department
Hamilton.
from
May-
acetate
at pH
Luxol
ofMedicine
fast
and
Canada.
the National
Cancer
Institute
of Canada.
I8
Submitted
6
Address
calf
2 wk
inverted
(GM)
II)
radioisotope
is sensitive
GIBCO),
Astra
from
fetal
an
CFU-C.’4
colony
by
were
CML,
penicillin-
under
or 2% glutaraldehyde
to define
Grunwald-Giemsa:
sepa-
concentra-
per
Histochemical
described,’
medium,
2-mercaptoethanol
1983
was
pigmentosa,
cultures,
Dulbecco’s
serum
blood
I .077)
after
the methylcellulose
onto
has been
(kindly
Granulo-
identified
(type
from
placed
on individual
this assay
From
gravity
picked
by the enzymatic
assayed
(histaminase,
myeboid
),
II colonies
what
assays
I 20 pg was used
.zl =
lymphocytes.
were
like
(PBS)
blood
line’3
Assays
performed
>95%
METHODS
(specific
with
peripheral
Histamine
I
and
saline
cell
“neutrophil-macrophage”
(Eo)
human
superna-
phytohemagglutinin-
blood
cells
containing
included
described,’2
appearances
either
I to type
in keeping
marrow
(CM)
from
peripheral
pipette,
oftype
precursors
bone
of CSA
leukemic
CM
of 200-1000
(see Fig.
phosphate-buffered
The
methods
BMC
in
previously
and
“eosinophil”
Pasteur
of
Sources
morphological
.14.15
elongated
Supported
density
anticoagulated
(I
described23’
0.5’:
AND
or
with
but
these
medium
(CSA).
Golde),
as being
I)
described’6:
Cultures
subjects
(type
Histamine
increased
of patients
chronic
microscope
nature
conditioned
human
their
of
proliferation
by methods
colonies
by
vitro
of
granules
Use
T-lymphocyte
D.
normal
Pathology.
Ficoll-Hypaque
rated
and
Dr.
(CFU-C)
were
peripheral
precursors
demonstrating
in the blood
in
prepared
a human
stimulated
recently
in rat
of BMC
(SM)
MATERIALS
Cell
have
v/v
activity
by
and
(EM)
states.
5%-20%
from
pg in CML
in histamine-positive.
the
their
allergic
CM
supplied
cyte
clarify
content
cultures
microscopy
electron-dense
BMC
peripheral
in SM
± 0.08
colonies.
of
of
pigmen-
in SM
II colonies
Electron
of
further
(12/153)
histamine
to 0.29
UP.
features
regulation
placental
for their
compared
revealed
and
cells
account.
type
colonies
and
lion),
relais not
into
and
colony-stimulating
and evidence
in
type for the other
been described
by Zucken-Frankbin
blood”;
we report
on the existence
in human
numbers
the exact
mast cells
size
histamine-negative.
disorders
derived
evidence
precursor9
of one
pg.
8%
urticaria
controls
(p < 0.0001 ). Calculated
was approximately
1 per 2 x 106
normals
to
to
with
2 x 1 0 nucleated
colony
0.19
±
in
and
human
hemopoietic
colonies
consisting
of either
basophils or mast cells (BMC),
apart
from the rare occurrence
of small
clusters
of mast
cells
in cultures
of
human
bone marrow.67
BMC-like
cell lines have been
described
in long-term
munine
bone marrow
cultures
in
suspension,t
but their counterpart
in man has not yet
derivation
ontogeny
1.1
of
sinophibs,3
megakaryocytes,4
or mixtures
thereof5
have
been found to exist in peripheral
blood or bone marrow.
However,
there have been no descriptions
of growth
of
Furthermore,
although
human
basophils
and
was
in contrast
4 patients
in histamine-positive
not
single cell precursors
suspended
in semisolid
widely
used
to determine
human
hematolineages.’
Erythroid
or myeloid
precursors
and differentiate,
respectively,
into colonies
erythrocytes,’
neutrophils-monocytes,2
eo-
been identified.
tionship
between
cell
II)
myeloid
putatively
per
was
from
1 per
ultrastructural
type
chronic
and
individual
a patient
19%
with
colonies
I and
this
and 6 normal
of BMC
CFU-C
Taking
0.10
Histamine-positive
8 patients
GROWTH
from
media
is
poietic
cell
that grow
of mature
often
previously
“neutrophil-
of all (type
(SM).
from
more
(CML);
tosa
(UP)
frequency
Blood
J. Bienenstock
in cultures
blood.
By
Peripheral
and
in normal
baso-
the
than
type.
repeated
of
demon-
were
of
rather
(50/86)
in
in cultures
HE
II).
mastocytosis
periphstudies
of maturation.
colonies
colony
58%
colonies
colonies
appearance
(type
(type
colonies
BMC
overall
leukemia
colonies
of metachromatic
stages
S. Telizyn,
revealed
in human
histochemical
histamine-containing
in Human
Richardson,
cultures
present
and
populations
described
T
(CFU-C)
microscopy
M.
granulocyte
homogeneous
inverted
with
hemopoietic
precursors
Light
phil/mast
found
Denburg,
of histamine-containing
from
eral
J. A.
Precursors
Program.
versity,
© 1983
June
reprint
Department
Hamilton,
by Grune
10, 1982;
accepted
requests
to Dr.
November
8, 1982.
J. A. Denburg,
Host
of Medicine
Canada
LSN
& Stratton,
0006-497l/83/6104-0027$01
and
Pathology,
Resistance
McMaster
Uni-
3Z5.
Inc.
.00/0
775
From www.bloodjournal.org by guest on June 18, 2017. For personal use only.
DENBURG
776
Table
1 . Histoch
emistry
of Hista mine-Containing
Histamine
Colonies
dehydrated,
and
embedded
in
Spurr
resin,
all
within
ET AL.
the
Beem
capsule.
Content
(pg/Colony)
Type
Stain
Alcian
I
blue
RESULTS
Type))
120
.
±
120
-
120
<120
-
+
-
Astra
blue
±
-
+
-
Luxol
fast blue
-
-
+
+
Myelo
+
+
-
-
Eosinophil
-
+
+
Colony
Relation
Peroxadase
blue,
purportedly
peroxidase
cases,
both
electron
histamine
sodium
After
cacodylate,
tetroxide
then
were
cacodylate
Vt.)
hr.
sodium
and
performed
myelo-
or eosinophil
described.7’9’7
‘
in 0.lM
stained
chemical
staining
presence
In some
morphology/histochemistry
on an individual
I summarizes
to colony
type
of
or
colony.
of prominent
the
histochemical
and histamine
content.
individual
lA)
capsule.
aspiration
aspirated
in a Beem
Fixation
of
postlIxation
the
2% glutaraldehyde
into
(Ladd
was
Research
continued
glutaraldehyde
was
performed
sodium
cacodylate
for
en bloc
for
in saturated
I 5 mm
15 mm
Industries,
at
and
4#{176}C
for
wash
with
and
1% osmium
at 4#{176}C.
Samples
uranyl
30
in 0.2M
acetate,
histamine-positive
demonstrated
described
positive
histamine-positive,
metachnomatic
granules
dase-positive,
idase
staining
to changing
in SM or
maturation.9
were also
as demonstrated
(Fig.
11).
“
‘
partially
Type
blue,
the
homo-
colony
cells
from
red to
mucopolysacCML
type
colony
cells
by Alcian
(Fig.
I ,C-E),
resembling
B MC
colonies
findings
Histo-
by Alcian
blue, Astra
techniques
revealed
geneously
distributed
in histamine-positive
(Fig.
I,B-E).
A maturational
sequence
blue staining
corresponding
chanide
content
of granules
colonies
Burlington,
min-2
content
were
and
previously
and Histochemistry.’
Content
Table
according
type II colonies
(Fig.
or May-Grunwald-Giemsa
Microscopy
Individual
0.IM
for eosinophils’8:
by methods
microscopy
Electron
were
specific
stains,
-
Morphology
to Histamine
II
blue
was
previously
Cells
in histamineeosinophib-peroxi-
by bight granular
I, but not
penoxtype
II,
Fig. 1 .
Morphological
and cytochemical
characteristics
of histamine-positive
colonies:
(A) inverted
microscope
appearance
of type II
colony; (B) May-Grunwald-Giemsa
stain; (C-E) alcian blue/eosin
stain. showing
maturational
sequence.
(Continued
on page 777.)
(F-H)
electron
microscopic
appearance
of histamine-positive
colony cell; (I) eosinophil
peroxidase
stain. Electron
microscopy:
(F) TEM of a cell
representative
of those seen in histamine-positive
colonies.
Some granules
appear
partially
extracted
( x9000).
(G-H) Higher magnification
micrographs
of granules
in this cell show: (G) a granule
containing
particulate
material
(g) and two
darkly staining
myelin forms inside the
granule
(arrowheads).
The other
granule
contains
a uniformly
electron-dense
central
portion;
this granule
may be partially
extracted
( x 54.500).
(H) A multivesicular
body (mvb) and extracted
granules.
Some microtubules
(M) and portions
of the golgi apparatus
(g) are
present
in the cytoplasm
( x 60.000).
From www.bloodjournal.org by guest on June 18, 2017. For personal use only.
HUMAN
BASOPHIL/MAST
CELL
777
COLONIES
Fig.
colonies
in all cultures
contained
dase-positive
cells
(not
shown).
of Colonies.’
Histochemistry
Relation
and
negatively
to Histamine
In order
to examine
ultrastructure
of granules
in
SM
colonies
and
rebate
this to histochemistry
and
histamine
content,
electron
microscopy
ofcells
from ‘12
individual
colonies
(6 type I, 6 type II) also assayed
for
histamine
was undertaken.
In all, an average
of 200
cells was screened
by transmission
EM of fixed cobnies in each of the colonies
observed;
EM photos
were
obtained
of 30 separate
in appearance
according
histamine-negative
(type
cells, which
to colony
I) colony
were
type.
cells,
(Continued)
or mast cells (Fig. I , F-H).
Myelin
forms,
bar bodies,
and typical
granular
membranes
>90%
myeboperoxiHistamine-negative
type
II colonies
were
both
Luxol-blue-positive
eosinophil-peroxidase-positive,
but stained
with Alcian
blue or Astra
blue (Table
1).
Ultrastructure
Content
and
1.
quite uniform
In contrast
to
histamine-
positive
(type
II) colony
cells contained
more
numerous, electron-dense
granules,
with features
previously
described
as characteristic
of either
human
basophibs
mubtivesicuand pen-
odicity
were all observed
(Fig.
I, G-H).202’
Most cells
appeared
polymorphonuclear
(Fig.
I F), with mature
nuclear
chromatin
and a Golgi zone and microtububes
(Fig.
IG);
the polymorphonucleanity
suggests
that
these cells more likely fit the description
of basophils
rather
than mast
bined
basophil/mast
scnibed.9
Histamine
Content
Both type
were found:
4/24 CML
type I and
cultures
ble 2 and
number)
amount
in SM
cells,’#{176}’202’although
cell
features
I and
20/38
of Individual
type
SM
Fig.
into
II histamine-positive
type I and 30/48
I 20 pg histamine
2). Taking
account,
of histamine
type
I (0.29
with combeen
de-
Colonies
type I and 9/43 CML
9/108
type II colonies
contained
cells
have
SM
colonies
type II;
type II; and 3/45
from UP or normal
pen colony
(Ta-
colony
size (approximate
cell
the mean
(± SE) calculated
(pg/cell)
±
0.07;
was significantly
range
0-2)
or
higher
type
lb
From www.bloodjournal.org by guest on June 18, 2017. For personal use only.
778
DENBURG
Table
2.
Calculated
Histamine
Content
of Human
Peripheral
Colonies
Blood
According
Type
Granulocyte
Colony
to Colony
Cells
and Frequency
ET AL.
of Histamine-Positive
Type
I Colony
Type
I) Co)ony
Histamine
Clinical
Group
(pg/cell)
TA1,lONormal
0.10
TA 1 , 1OUrticaria
TA1,lOSystemic
myeloid
‘Number
leukemia
of histamine-positive
tP
<
0.05,
*p
§p
<
0.01.
range
<0.
mastocytosis
TA1,lOChronic
(1.06
0.08;
pigmentosa
compared
±
Histamine
3/28
10
0/ 1 7
0.17
±
O.O7
20/38
1.06
0.16
±
O.O7t
4/24
0.29
or UP, Student’s
Frequency
0.05
±
<0.
0.29
of colonies
colonies/number
to normals
Frequency
0.04
8/73
10
1/35
±
0.19
30/48
±
O.08
9/43
picked.
unpaired
t
test.
<0.0001.
±
0.19; range
0-7)
and CML
type II (0.29
range
0-1.5)
than in CML
type I (0.16
± 0.07;
0-1.6)
and
normalor
UP type
I on type
±
of most of the
type I and type
HISTAMINE
cor’imwr
individually
II colonies
picked
by his-
OF INDIVIDUAL
GRANULOCYTE
LONIES
1.0
0.7
Gm
tonis.
. _Eo_CoIonC
C
in human
quantitative
0
I
0.4
0
0.3
assay
-
.
NORMALS
0
lie,
blood
helps
for BMC
of individual
in
ofa
precur-
to delineate
precursors
.1 721
features
of
commensurate
mature
BMC
in most
normal
or UP
colonies.
to
be
colonies
present
SM,
BMC,
with
some
a
in semi-
in small
correlates
.25
and
well
ultra-
with a calculated
that expected
CML,
Although
and
amounts
in
of
rarely
histamine
methodology.’624
=
...
0
SSS.
isoi
URTICARIA
PIGMENTOSA
0
colonies.
BMC
CUNICAL
..
SYSTEMIC
MASTOCYTOSIS
Conversely,
(27)
CHRONIC
MYELOID
LEUKEMIA
GROUP
Fig. 2.
Histamine
content
of individual
granulocyte
colonies.
GM (type I) (0) and Eo (type II) colony
(#{149})
histamine
content
is
shown
in nanograms
per colony.
according
to patient
group.
Numbers
in parentheses
refer to total number
of colonies
in which
histamine
content
was undetectable
in a given condition.
Mean
histamine
content
(nanograms
per colony) for type II colonies
was
significantly
different
from all other
groups
(p < 0.01 ); for type I
SM colonies,
it was different
from
normals
or UP (p < 0.02).
Histamine
content
of both CML type I and II colonies
(nanograms
per colony)
was significantly
different
from normals
or UP (p <
0.02. Student’s
unpaired
t test).
colonies,
which
no
histochemical
criteria)
detectable
histamine
identified
a relatively
subset
of eosinophil-like
colonies
taminase-sensitive
histamine
not
#{149}S
0.2
601
peripheral
colonies
sinophil
colonies
(by
have picked
contain
and 2). We thus have
(21)
is shown
solid hemopoietic
cultures.
These
findings
supplement
the suspension
assay
for BMC
precursors
we have
described
in CML
peripheral
blood
cultures.2223
His-
current
0.6
8#{176}
groups
has
in
been
human
eo-
sinophils,24
such conclusions
have been based
on older
fluorometnic
assays
that underestimate
the histamine
content
of basophils
by tenfold
when
compared
to
0.8
0.06
presence
simple
said
0.9
0.1
The
son for histamine-positive
structural9’#{176}’20’2’
histamine
content
1.1
U
patient
DISCUSSION
tamine
content
with
accepted
1.2
C
0
0
in different
II (s0.1;
range
0-1.8)
colonies
(Table
2). Histamine-positive
colonies
in which
100%
cells
were
metachromatic
(“pure”
BMC)
invariably
contained
histamine
at a
level
equivalent
to >1 pg/cell
in a 200-1000
cell
colony.
Distribution
histamine-positive
tamine
content
Fig. 2.
in SM
that
found
almost
“pure”
eo-
that
we
(Tables
1
infrequent
contains
in most
invariably
a histype II
con-
tam homogeneous
cell populations,
appear
more
frequently
to be a subtype
of type
II (“eosinophil”)
colonies
overall;
in SM or CML,
higher
proportions
of
all colonies
(both type I and type II) contain
BMC
by
morphological,
histochemical,
and biochemical
cnitena compared
with normal
on UP colonies
(Table
2).
These
observations
are consistent
with
the accepted
notion of the common
origin of neutrophils,
eosinophils
as well as
progenitor;
commitment
basophils
from
a committed
granulocyte
they
also
suggest
that
specific
lineage
is probably
stages
of hematopoietic
with high
histamine
made
cell
content
at a number
differentiation.5
(1
pg/cell)
of different
Colonies
and tight
From www.bloodjournal.org by guest on June 18, 2017. For personal use only.
HUMAN
BASOPHIL/MAST
compact
cell
CELL
779
COLONIES
aggregations
(type
II)
in SM
were
pre-
dictabby
“pure”
BMC,
while
in other
instances
mal or UP or CML
cultures,
as well as some SM
(nortype I
colonies)
were “mixed,”
containing
BMC
as well as
other
cell types (Denburg
JA, in preparation).
This is
reflected
in the bower mean
histamine
content
per cell
in the latter
groups
in comparison
to SM
type
II
colonies
(Table
2). Although
we did find a relatively
high frequency
in CML
type
mine
of histamine-positivity
I or type II colonies,
content
and
mixed
cellular
the existence
of circulating
especially
in CML,
which
tion along neutrophil,
findings
supportive
by Aglietta
eosinophil,
of this concept
cell colonies.26
increases
patient
composition
hematopoietic
are capable
et al., using agar
et al. in a recent
Nakahata
mast
in SM type I or
their
lower
histaargue
for
Furthermore,
progenitors,
of differentia-
serial
in SM type I colony
histamine
content
as our
progressed
to end-stage
disease.
This
may
mean
that
there
is progressive
restriction
of lineage
commitment
to the BMC
pathway
as mast cell proliferation
accelerates,
a finding
not unlike
that which
we
have reported
in blastic
CML.23
The numbers
of circulating
BMC
precursors
in UP
are not different
from normal,
suggesting
that
prebeukemic
state
exists;
this is consistent
notion
that a spectrum
of mast cell proliferation
from
UP
frequency,
C/b6
cells,
to SM
based
and mast
cell leukemia.
on a plating
efficiency
Ficolb-Hypaque
a white
blood
in SM, a
with
the
exists
metachromatic
as well
eosinophib
peroxidase-positive
tive colonies
(Fig.
I); cells
as Luxob
cells
in these
characteristic
controversy
Ackerman
crystals
may be
fast
blue
in histamine-posicolonies
have
of BMC
exists
as
or
EM
(Fig.
I ). Although
to whether
or not
consensus
Recently,
has been
however,
et ab. have demonstrated
Charcot-Leyden
in human
basophibs,27
suggesting
that
there
more
commonality
between
eosinophibs
and
clarify
the ontogeny
well as in various
of these cells in the normal
state as
bone marrow
disorders.
It may also
shed
light
on the mechanisms
phibs
and
BMC
of mobilization
observed
in allergic
of eosino-
states.
ACKNOWLEDGMENT
David
Golde
Robertson
typed
Dolovich,
blood
and a
other
found
basophibs
than
heretofore
appreciated.
This
is in
accord
with numerous
observations
on the concurrence
of presence2t
or absence29
on abnormalities30
of both
these cell types in pathologic
states.
The application
of
both eosinophil
and BMC precursor
assays
may help to
Dr.
Precursor
of 20 CFU-
separated
peripheral
count
of S x l06/ml,
cell
uniformly
basophils
from
since we have
basophils
contain
peroxidases,
the
that mature
basophibs
do not.2125
in CML,6
and by
analysis
of mouse
we did observe
used
commonly
to distinguish
granubocytes
require
reevaluation,
appearances
considerable
and BMC
pathways;
have been reported
cultures
elegant
distribution
of colony
types
as described
in Table
2,
can be calculated
to be approximately
I per 2 x l0
nucleated
cells in SM and I per 2 x 106 in normal
peripheral
blood.
The specific
morphological
or histochemical
criteria
M.
Rosenthal,
kindly
provided
Mo conditioned
the manuscript.
M.
Fisher,
J. Senn,
We
wish
D. Hillyard,
and
W.
medium.
to thank
Drs.
H. Messner,
E. C. Wilson
W.
for their
Janice
R. Barr,
J.
Nicholson,
help
D.
in access
to
patients.
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1983 61: 775-780
Basophil/mast cell precursors in human peripheral blood
JA Denburg, M Richardson, S Telizyn and J Bienenstock
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