From www.bloodjournal.org by guest on June 16, 2017. For personal use only.
The
By
Antigenic
Structure
II. Histocompatibility
Rabbit
Blood
SHthLEY
the
N homografts
PREVIOUS
in STUDIES
rabbits
homologous
blood
platelets
to
and
the
skin
is known
and
humoral
be
jection
of a tissue
available
clearly
reactions
The
object
of the
number
blood
present
In
group
this
donor
which
the
per
ml.
cent
time
platelets
injection
of the
the
plasma
was
poured
which
humoral
factors
accelerated
re-
present
no
with
method
certainty,
other
histocompatibility
The
in outbred
results
anti-
obtained
blood
of a skin
skin
Suspensions
blood
drawn
one
was
grafts
Free
were
rabbits,
on
of
indicated
platelets
homograft
in
the
in a
The
blood
plate’let
rpm
for
10
sediment,
was
off,
and
then
and
the
centrifuged
rich
nlasma
The
again
at
was
at
platelet
button
0.7
thus
were
recipient
collected
animals
in
at
rpm
(170
1000
another
x
was
for
g)
for
20
test
transferred
rpm
(1650
obtained
x g)
again
of
1111.
EDTA
chloride)
SOdiunl
into
was
2
into
cent
rpm
pipetted
supernatant
3000
animal
per
1000
centrifuged
centrifuged
platelets
in
determined.
donor
in
was
minutes.
then
the
frolil
blood
occasion,
Leukocytes
ethylenediamine-tetraacetate
tubes.
1000
discarding
platelet-rich
It
during
METHODS
intradermaliy,
of homologous
supernatant
at
contain
of homologous
rejection
time
AND
conducted
injected
of cardiac
test
4 C. f The
plasma
and
disodium
to
centrifuged
tube,
experiments
of Platelet
transferred
at
of
rejection
Eighteen
(1.5
is at
be tested
directly
antigens.
of animals.
rabbits
Preparatioll
There
investigation.
MATERIALS
from
lead
phenomenon
can
blood
rejection.
that
the previous
intradermal
rabbit
could
induce
shortening
significant
however,
and
that
the
which
produce
( s)
factor
immunity
as to whether
the
not,
of skin
towards
that
histocompatibility
transplanted.3
transplantation
of homograft
question
was
could
produced
tile
subsequently
by which
transplantation
of immunity
demonstration
is a complex
are
with
DAMESHEK
Donocan
that
degree
contain
immunity
identified
by acceleration
gens
platelets
Janet
This
in common
blood
Platelets.
in
WILLIAM
of
it was a demonstrated
elicited
significant
later
infused.1’2
transplantation
cannot
AND
assistance
antigens
that
that
cellular
than
share
conclusion
BALDINI
technical
I
platelets
MARIo
EBBE,#{176}
With
of Jilood
Antigens
Platelets
tul)e
to
10
minutes.
30
minutes.
for
resuspended
afl(l
minutes
in
and
another
This
The
1-2
ml.
From
the
Blood
Research
Laboratory,
Pratt
Clinic-New
England
Center
Hospital,
and
Department
of Medicine,
Tufts
University
School
of Medicine,
Boston,
Mass.
The results
contained
in this paper
were presented
at tile 1961 Meeting
of the American
Society
of Hematology
in Los Angeles;
and partially
reported
in abstract
form
in J. Gun.
Invest.
40:1035,
1961.
Aided
by a grant from the Atomic
Energy
Com.nsi.wion,
#AT
(30-1)
1276.
Submitted
Dec. 14, 1961; accepted
for publication
Jan. 17, 1962.
tenure
of The Medical
Foundation
Research
Feiiowship.
Refrigerated
Centrifuge,
Model
PR-2,
International
Equipment
Co., Boston,
Mass.
f/ic
548
BLOOD,
VoL.
19,
No.
5 (MAY),
1962
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ANTIGENIC
0.85
STRUCTURE
per
directly
tinder
suspension
phase
contrast
mixed
with
was
and
examined
nwthod,
tile
injected
into
the
Without
multiple
sites
of
magnification
leukocytes.
If
tile
Platelet
were
leukocytes
experiments
preparatlons
experience,
seen
were
percentage
by
either
SCCI1
it
because
the
micro-
by two
exaniinecl
a drop
of the
with Wright’s
stained
leukocytes
if
beginning
increased
any
dried,
no
of these
Only
contrast
( 2)
and
a slide,
Ofl
discarded.
At the
with
x
phase
of leukocytes
EDTA,
were
was
Was
it
necessary
of
contamination
of
contaminated
considerably.
adjuvant:
on
(2)
With
of complete
posterior
the
adjuvant:
Freund’s
thorax
of
glassware
The
shaved
saline
skin
The
saline
adjuvant,
the recipient
used
for
after
the
the
suspension
of
the
of
lower
suspension
then
of
injected1
was
platelets
back
of
the
injected
recipient
platelets
was
intradermally
mixed
with
into
multiple
subcutaneously
into
rabbits.
an equal
sites
volume
over
the
animals.
collection,
separation
and
injection
the
of
platelets
was
siliconized.
Homografting
weeks
Two
skin
rabbit
platelet
injection,
the recipient
animals
were
challenged
with
a
platelet
donor
(“specific”
skin)
on one ear and one from another
skin)
on the other
ear.
Each
graft,
of full
skin
thickness,
was
measured
2-3
x 1 cm. in size. In these experiments
tile onset of graft
as basic
parameter
for measuring
tissue
sensitization.
The
onset
of
by daily gross inspection
and was judged
by the onset
of cyano-
from
hoinograft
(“nonspecific”
rejection
shape
and
was taken
rejection
was
ovoid
in
sis,
later,
430
by
of Platelets
(1)
Skin
549
IL
globulin
for
one-half
but
decreased
All
at
rabbit
was
animal.
approximately
In/ection
of
suspension
recipient
leukocytes,
samples
microscope
a drop
microscopically
platelet
discard
with
PLATELETS
NaCI.
Platelet
counts
were done on this suspension
suspension
was carefully
examined
for the presence
( 1 ) 1.8 Ill1fl.,
either
IIfldillIte(l
or (lillited
1 : 10 with
methods:
to
BLOOD
cent
The
scopy.
stain,
OF
which
and
determined
brown
edema,
discoloration,
to complete
progressed
dryness,
necrosis
scaling,
the
of
or
partial
graft.
Each
(usually
of
peripheral)
these
necrosis
changes
was
any one of
two or more
of the above criteria
were considered
significant
of initial
graft
breakdown,
and
1 PltiS
changes
in three
or more
of the parameters
were
considered
significant.
Signs
of graft
breakdown
were not considered
significant
if they were later reversed.
The onset of homegraft
rejection
was,
thus,
determined
retrospectively,
and
as
“rejection
time”
the time
was taken at which
there appeared
definite,
significant
signs of graft breakdown
which
then
progressed
without
reversal
to complete
necrosis
of the
graft.
quantitated
these
on
criteria
0-4
a
were
not
plus
basis.
considered
Minor
changes
significant.
Two
(1
plus
plus
2 pius)
involving
recorded
or
cllanges
in
RESULTS
By
ized
these
criteria,
recipient
the
rabbits
a standard
deviation
defined
above,
was
technic,
skin
The
iously
of
by
of 2.6 days
(fig.
of skin
3 to 8 days
(fig. 1).
followed
regularly
times
immunized
times
from
of 1.1
usually
autografts
rejection
average
rejection
ranged
survived
homograft,
an
ranged
period
i.e.,
from
and
onset
of rejection
in one day. By
indefinite
homografts,
non-immun-
of 5.8 days
in
1 to
of
time.
animals
6 days
as
this
prev-
with
an
1).
times
of “specific”
and
injected
with
platelets
Nineteen
the time
animals
limits
imposed
mean
nonimmunized
for
for
skin
The
rejection
mals
previously
were
in 34
an average
In these
animals,
the
by complete
necrosis
12 second-set
a skin
homografts
with
so
by
studied.
two
rabbits.
Of
standard
Of
the
“nonspecific”
presented
are
these,
skin
homografts
in table
I and
11 rejected
deviations
remaining
eight
011
both
either
rabbits:
grafts
side
(1)
in anifigure
1.
within
of the
five
re-
From www.bloodjournal.org by guest on June 16, 2017. For personal use only.
550
EBBE,
REJECT/ON
TIME
OP
SKIN
HOMOGRAPTS
SENSITIZED
Non-
Immunized
I
BALDINI
IN
AND
DAMESHEK
PLATELET
RABBITS
Immunized
To Plotelets
Immunized
11
To Skin
ii
Specific
Skin
Non-
‘:‘
8-
Specific
Skin
i_.;:_i5:I5555555555
S
7.
(1)
6
Ui
5.
I-
z
4.
IC-)
Ui
3.
0
-)
Ui
2
Fig.
grafts
1.-In
42 per cent of the platelet
sensitized
(2) were
rejected
significantly
earlier
than
as rapidly
as in animals
specific”
the
skin
experiments
jected
graft
graft
immunized
homografts
were
by
rejected
rabbits,
the
in nonsensitized
“specific”
skin homoanimals
(1) and
skin homograft
(4).
The “nonnormal
in only 10 per cent of
a previous
sooner
than
(3).
tile “specific’
homograft
rapidly
(1-3
days),
while
the “nonspecific”
was rejected
normally
(5.-fl (lays);
(2) one rabbit
rejected
the “specific”
rapidly,
(1 day)
while
the “nonspecific”
graft
was
maintained
for
longer
than
normal
rapidly
(1-2
days).
The
group
of
grafts
in
there
was
exhibited
sued.
time
animals
a different
not
days);
immunized
fashion
infrequently
significant
The
(13
mechanism
to
than
of such
(3)
two
animals
platelets
also
nonimmunized
a period
symptoms
and
of
appeared
controls.
of 2, 3 or 4 days
breakdown
changes
will
rejected
perhaps
to
In
during
before
which
the
animals
the
graft
necrosis
revealed
by
a
grafts
reject
these
complete
be
both
en-
studies
in
progress.
In summary,
42 per cent of the platelet-sensitized
rejection
of the “specific”
skin graft
earlier
than
time
itized
grafts
values
below
two
standard
deviations
of the
animals
non-sensitized
showed
onset
of
animals,
with
mean
for
animals.
On the contrary,
only 10 per cent of the
were
rejected
sooner
than normal
(fig. 1).
value
“nonspecific”
the
nonsens-
skin
homo-
From www.bloodjournal.org by guest on June 16, 2017. For personal use only.
ANTIGENIC
too
STRUCFURE
OF
The
addition
of
small
a number
BLOOD
PLATELETS
Freund’s
adjuvant
of experiments
551
U
to the platelet
injection
evaluation
at this time.
for
was
applied
in
DiscussioN
From
the
Intradennal
above
results,
in/ection
of
in a significant
grafted
from
A clear,
after
number
the same
positive
platelet
be
in small
doses.
In
the
result
proposed
size
that
the
animals
brief
was
significant
period
probably
Other
injection
of rejection
have
technical
given,
the
the
skin
be
by
fo this.
the
number
The
occurrence
in two
of the
a reflection
dividual
of accelerated
recipients
in
of
antigenic
specificity
in our
previous
of
had
been
overlap.
A
antigens
the
investigations
in which
(13
thought
titer
of
jected
days).
Although
very
rapidly
Further
studies
tized
animals
are
endothelium
past4c4d
and
in this
A relationship
immunity
elicited
question
relationship
is still
in
was
clearly
for
(table
may
be
demonstrated
the
and
one
red
that
probably
incomplete
skin
was
studies
in-
observed
were
skin.1’2
In
an unusually
only
such
homograft
was
of
survival
done
in
one animal,
long period
the
of
may
be
experiment,
it
suggesting
however,
a high
was
re-
1).
of skin homograft
possibility
that
between
isosensitization
by tissue
transplantation
debated
platelets
in common
was
vascular
damage
rabbits
skin
platelets
produced
in this animal,
The
“specific”
homograft,
animal
have
antigens
it is possible
that
to
was
is drawn
phenomenon
homologous
viable
for
occurred
was
on the histology
warranted.
The
to the platelet-sensitized
phenomenon.
this
this
that
“enhancement”
humoral
antibodies.42’43
with
single
and
of positive
However,
obtained
conclusion
“nonspecific”
platelet
animals
which
had been
grafted
with
“nonspecific”
skin homograft
appeared
time
of the
a
platelet-immunized
results
injected
similar
common
The
of platelets
injected,
the
the time
schedule
adopted,
and the
antigens.
rejection
which
non-
4b
It is realized
of positive
that
in
others.4’
as of now,
believe
only
multiple.
homografts
influenced
the percentage
to elucidate
these
points.
we
grafts
animals.
antigens
may
the animal
species
used)
may have
also
results.
Further
studies
will be necessary
that
they
cannot
be attributed
to chance,
blood
platelets
contain
histocompatibility
later
skin grafts
in nonsensitized
for the demonstration
of
time
(the
number
of injection,
of
result
of
homologous
condition
skin
skin
cent
responsible
skin-rejection
reasons
the site
per
experiments
been
induced,
towards
homologous
42
partial
in
to another
transplantation
this
than
of take
of the
an unfavorable
of
of
only
carry
for
onset
may
rabbit
immunity
rejection
platelets
earlier
used
of
in
reasons
the
grafts
shortening
animals.
platelet
blood
occurred
of skin
a degree
obtained
the
experiments,
is apparent:
front
one
accelerated
was
that
animals
larger
of
However,
above
following
platelets
of animals,
donor.
sensitization
It could
sensitized
the
blood
the
rejection
in platelet-sensiblood
platelets
and vascular
repeatedly
in the skin
prominent
induced
has
for
ce1ls.#{176}’ Blood
feature
by blood
long
been
the
suggested
homografts
white
platelets
in the
applied
of the
rejection
transfusions
suspected.5
blood
have
cells,6
never
and
While
the
been
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552
EBBE,
Table
1.-Rejection
Time
of Skin
BALDINI
in Platelet
Homografts
Number
Injected
of
Platelets
(x 10’)
Freund’s
Adjuvant
383
2.352
-
387
2.4675
-
Z-
400
402
2.008
1.7425
436
1.124
.$&
436’
437
0.7025
1.815
438
301
1.141
1.96875
411
2.5
412
2.46
-
L.
1.752
381
2.556
385
386
409
410
2535
2.52
1.44
2.53
395
403
5.908
4.284
Rejection
time
in 34
-
6
5
5
5
5
5
5
5
8
8
-
7
4
-
4
4
6
4
-
6
6
+
6
6
+
+
1
1
2
2
3
1
5
6
6
5
5
131
2
1
2
1
-
--
-
non-sensitized
Rabbits
Time
(Days)
“Nonspecific”
graft
6
6
-
S
378
DAMESHEK
Sensitized
Rejection
“Specific’S
graft
-
Animal
Number
AND
animals.
Average
value
5.8
=
days
±
1.10
S.D.
01
Not
included
previously
in figure
1.
considered
in this
of histocompatibility
The
experiments
of inducing
herein
a state
patibility
respect
and
reported
also
of immunity
antigens-are
not
of
histocompatibility
However,
antigeniticity,’4
genicity25’26
Histocompatihility
of
with,
proteins
substances
are,
Although
jection
was
presence
investigation
carried
contained
nuclei
fragments
nuclear,
of course,
almost
in
previous
experiments
obtained
by
constituents.
and
cell
themselves.
in
(nuclear
to he
devoid
mitochondrial
meml)ranes,
both
It is conceivable
cellular)
and
microsomal
as
at
present
time,
demonstrated
components.21’25’26
in
that
the platelets
these
antigens
point
the
in
disrupted
has
cell
Such
homograft
re-
cytoplasm,
the
demonstrated
may also
been
in this
present
be
recently
given
H-2
antigens
were
found
and
cellular,
rather
than
earlier
cells
fractions,2224b
experiments,
may
all
have
of
consist
materials.
accelerated
of
to be
anti-
to
in biological
fractions
not
believed
to lack such
lipid
others,26
of
the
was previously
been
shown
originally
and presence
The precise
been
with
nuclear
that
of
nuclei
and
to this
most
is,
capable
have
by
Evidence
antigens
is, histocom-
cell
present
sensitization
Herzenberg;2’
and
which
has since
universally
antigens
to the view
that
by
antigens
DNA,
carbohydrate
of histocompatihiiitv
lends
support
Herzenherg
thought
do not contain
a nucleus,
be clearly
demonstrated.152#{176}
antigens
possibly,
not
in cytoplasmic
even
demonstrate
exclusively
defined.24b
for tissue
nature
completely
responsible
been
skin-homografts-that
towards
believed.6’12-14
In fact, blood
platelets
of DNA
in blood
platelets
could
never
chemical
have
antigens.”
to
in
by
be
the
membrane
contaminated
which
were
be-
From www.bloodjournal.org by guest on June 16, 2017. For personal use only.
ANTIGENIC
STRUCTURE
lieved
hrane
OF
to be themselves
in platelets
has
mitochondria
are
observations
excludes
finding
be
and
the
possibility
histocompatibility
antigens
regularly
is also
with
acid
phosphatase
Blood
platelets
to red
cell
onstrated
ence
possess,
by
and
have
less
meaning
in
various
evidence
contain
contents
such
high
of acid
are
conantigens
containing
platelets
a complex
antigenic
platelet
platelets,38’39
antigens.
on
high
as blood
and
in human
“types”
These
current
could
tissues
memand
anti-
doses
of
phosphatase
known
to
contain
37b
of histocompatibility
“groups”
that
platelets
that
possess
antigens”
others
platelets.2M’2
of histocompatibility
blood
concept,
therefore,
blood-group
blood
location
Stetson’3
abundantly.34
antigens.2”24
A cell
microscopic
observation,27
emphasize
that
and
this
in
to
cellular
l)y Basch
consistent
553
II
present
here
structure
gens.
The
to
summarized
chemical
in no way
PLATELETS
carriers
of transplantation
been
seen
by electron
known
are
cerning
BLOOD
tile
the
our
of
practice
of
tests
platelet
already
indicate
of human
serologic
addition
In
antigens
experiments
Classification
basis
structure.
specific
in
blood
vitro
pres-
platelets
may,
in
therefore,
than
transfusions
demthe
previously
prospected.40’4’
SUMMARY
The
rejection
time
sensitized
had been
by one
carefully
determined
by
graft
homografts
gross
the
donors
platelet
inspection
and
sensitized
(“specific”
tized
animals
based
than
normal
rabbits,
grafts)
were
in 42 per
cent
were
the
rejected
of the
the
platelet
donors
in only 10 per cent
The
results
histocompatibility
in
the
Ofl
rabbits
previously
blood
platelets
which
graft
rejection
time
was
onset
interpreted
antigens.
Le
tempore
de rejection
previemente
de sanguine,
sensibilisate
le quales
of
definite
(“nonspecific”
of the experiments.
grafts)
as
demonstration
signs
of
plus
normal
Le
graffo
esseva
le declaration
sanguine
contine
esseva
were
blood
que
10 pro
antigeno
in nonsensi-
cento
animals
rejected
sooner
platelets
contain
esseva
mesurate
determinate
de definite
in conilios
plachettas
leucocytos.
per inspection
a oculo
signos
de deterioration
de plachettas,
le homograffos
“specific”)
esseva
rejicite
a un
in
le animales
nonsensibilisate
del
como
cutanee
tempore
in
Le homograffos
ab animales
altere
que
“nonspecific
) esseva
rejicite
post tui teml)ore
interpretate
platelet
from
injection
intradermal
de homologe
essite
liberate
meticulosemente
de
per medio
(graffos
precoce
in solmente
resultatos
cutanee
del
super
sensibilisate
de plachettas
than
homografts
that
the
INTERLINGUA
un
in le graffo.
In le conilios
ab le donatores
cento
del experimentos.
(Ic plachettas
(graffos
IN
from
earlier
The
de homograffos
rejection
basate
hoinografts
significantly
per
habeva
Le tempore
de
inerme
e esseva
significativemente
skin
experiments.
SuIAnIo
que
measured
was
injection
of homologous
free of leukocytes.
Tile
breakdown.
In
other
than
of skin
intradermal
prepared
42
pro
Ic donatores
plus i)reve
experimentos.
tin
a histocompatihilitate.
demonstration
que
Ic 1)lachettas
(IC
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554
EBBE,
BALDINI
AND
DAMESHEK
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556
EBBE,
Shirley
Ebbe,
M.D.,
Senior
Research
Blood
Research
Laboratory,
In.s’tructor
in Medicine,
Tuft.c
Boston,
Mario
ratory,
Baldini,
New
Tufts
University
Dameshek,
\Villiani
tori,,
M.D.,
England
New
England
Tufts
University
Fellow
School
of Medicine,
School
AND
Hematology,
Hospital:
Medieine,
Mass.
Director,
Hospital;
Center
in
New
England
Center
University
School
of
Associate
Center
M.D.,
BALDINI
Director,
Blood
Research
LaboAssociate
Professor,
Boston,
Blood
Hospital;
of Medicine,
Mass.
Labora-
Research
Professor
Boston,
of
Medicine,
Mass.
DAMESHEK
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1962 19: 548-556
The Antigenic Structure of Blood Platelets. II. Histocompatibility Antigens
in Rabbit Blood Platelets
SHIRLEY EBBE, MARIO BALDINI, WILLIAM DAMESHEK and Janet Donovan
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