From www.bloodjournal.org by guest on June 17, 2017. For personal use only.
The
Effects
of
the Dog
of
Splenectomy
on the Red
with
Particular
Emphasis
Reticulocyte
Response
By MORTIMER
LTHOUCH
response
NUMEROUS
to sl)leIuectoIny’,
I)eell
neglected.
This
dogs
following
removal
quent
to Otiler
inferetuces
types
have
as
of
regarding
the
and
of
the
heniatologic
count
the
Based
certain
splenectonlized
on
has
reticuiocyte
with
compared
or phlebotomy.
drawn
in
beeiu written
the reticulocyte
a study
spleen
of simrgery
been
erythropoiesis
in
is primarily
of the
Cells
on
LORBER
PAPERS
have
the effect
on
report
Blood
that
occurring
on observed
the
often
level
of
subse-
differences,
mechanisms
concerned
in nonspienectomized
animals.
METHODS
Fifteen
formmied
adult,
male,
on
mongrel
individma!
animals
at
(logs
were
2 to
6 momitil
enlployed.
The
following
procedures
were
per-
imltervals:
l)og
A
Splemuectonly
B
Splenectomny,
C
Splenectomy,
Unilateral
Nephrectomiiy,
1)
Splemuectomy,
Unilateral
Nephrectomuy
Phlebotomy
( splenectomized
E
Phlebotomy
F,I
Splenic
C
Splemuic
Venoums
Routing
into
the
H
Splenic
Venoums
Routing
into
the
J
Phlebotonuy,
Vemloms
K,NI
Plllehotomy
L
Piulebotomily,
N,O
Unilateral
The
results
tomized
several
ant!
Unilateral
Nephrectomy
2
dime to tiue remiuoval
tiue
performed
from
initially
From
the
Center,
in
The
opinions
A
of
tile
Systemnic
Ciremmiation,
Circ.,
Pllltbotomy
Pillehotomy,
Splemmectomy
I)lmms Splemiectomny
of
or
(N,O),
of
6
dogs;
F-I,
se
per
the
at
of
omue
or
operation.
nuonths
to
tile
were
one
(J)
it
Naval
was
of
done
Research
of
tile
There
into
splemuectomy
splenic
as
Unilateral
blood
I)og
was
nephrectomy
subsequent
initial
splenec-
rerouted
resmmlts
removal
tile
Medical
was
plilebotomnizet!,
(3)
inclumding
were
procedumres.
blood
the
whether
previously.
in
(A-i))
Otiler
vemuous
to
muerely
(J-NI)
which
from
splenic
as
clogs
and
4
recovery
procedures,
Hemuuatoiogy,
surgery
in
Institute,
E
was
to
recovery
all
but
National
one
Naval
Maryland.
assertions
as
11
in
information
Four
fifteen
2 dogs
Bethesda,
1)ogs
spleen
(2)
total
Division
muot to be construed
service
at large.
in
splenectomizec!
phlebotomuy.
Nledical
previously
Ciremmiation
Systemic
following
affor!ing
circulation.
been
months
Systemmuic
observet!
( H,L)
(1)
thus
fromuu
had
were
others
groups:
were
which
the
Nephrectomy
circulation,’
portal
into
Unilateral
splenectomy
control
systemuic
fifteemu
Routing
Nephrectomy
initially
were
the
of
Philehotomny
official
contaned
or
reflecting
herein
are
the
views
the
private
of
the
ones
Navy
of
time
writer
I)epartment
Nov.
18, 1957;
accepted
for publication
Apr. 12, 1958.
The author
wishes
to express
his appreciation
to the mnembers
of the Division
Physiology
for performing
the surgery,
to the staff of the Division
of Biophysics
for the statistical
analysis
of the data,
and
to Dr.
N. Raphael
Shu!muami
of the
Henuatology
for his aid amid encouragement.
amid
the
or
are
naval
Submitted
972
of
Surgical
Research
Division
of
From www.bloodjournal.org by guest on June 17, 2017. For personal use only.
EFFECTS
OF
( E ) , were
formed
on
4
SPLENECTOMY
followed
10
Tue
animllals
sets
of
on
control
were
the
pipette
erythrocyte
counts
tocrit
divisiOml
were
of
All
sumrgery
nephrectomy
splenic
blood
of
the
splenic
formued
using
(7%
of
the
Iliac
procedures
done
were
per-
and
counts4
were
noting
values
were
entire
Ier
red
hemuuatocnit
also
division
fragility
numiuber
deter-
1000-3000
per
by
multiplying
erythrocytes
were
assmmnlec!
the
per
in
counts
was
erythro-
perfornued.
observation
cell
Double
and
cyanmethemoglobin
tile
of
blood
All
tumbes,
needle.
osmotic
10,000
period
dogs,
least
oxalate’
the
obtained
150,000±
At
procedures.
normoblasts
Drabkin’s
Blue,5
coumnt.
from
indices,2
Niethy!ene
water.
major
balanced
directly
of
cell
and
the
circulating
platelet
the
biscuits,
to
reticulocyte
using
the
genera!
and
left
renal
circulation
veins
from
marrow
was
ilemlla-
10
dogs
which
done
but
rarely,
calculating
in
fenuoral
domue
the
humntlrec!
were
cell
The
of
smaller
and
Routing
end-to-end
an
nephrectomuy.
A
I)og
technics.
by
22%-31#{231}’
vein.
in
specimnemis
Five
left
Splenectomy
surgical
accomplished
withdrawing
was
anestiuesia.
routine
following
the
volume
Nembutal
employing
anesthesia,
weight)
bone
intravenous
performed
into
tllereafter.
marrow
15
Wintrobe
for
blood
erythrocyte
performed
blood
crest
intervals
were
microscope
cells
in
modification
red
throughout
red
dog
prior
smears
pillebotomllized
were
body
and
valumes.
Pentothal
estimated
the
the
150,()00
was
lateral
and
New
present
reticulocyte
which
stained
Ahsolumte
In
)
nueat,
collected
( Crosby’s
by
surgery.
a valime
absolumte
smears
using
done.
iuorse
performed
blood
Wright
Phase
percentage
ummiderwent
amid
and
counts
of
were
hematocrit2
were
reticmmlocyte
a diet
on
performed.
Reticumlocyte
( A .E,H,L
dogs
fed
hemoglobin’
were
erythrocytes
and
counts,’
\Vintrobe
973
CELLS
( F-O).
performed
mllOrpholOgy,
mmuinations
BLOOD
determinations
platelet
),
RED
spienectomized
caged
were
except
muetluod
7
aninuals
henuatologic
determinations
cyte
ON
were
estimated
blood
phlebotomy
the
anastomilosis
ph!ebotomies
the
uiiii-
of
of
per-
volume
about
10%
of
E.
aspirated
the
at
differential
time
counts
of
were
sumrgery
domue
amld
omi
at
various
stained
Wrigiut
smears.
The
data
variates
were
are
nihcamlt
analyzed
ummuknowmu.
valmme being
statistically
The
dime
limits
solely
i)y
of
to
tile
chance
were
was
5 in
Cox
amid
Cociuramu
significance
sumch
t
that
the
lilethod’
when
likelihood
of
the
a
sig-
100.
RESULTS
Platelets.
to
22
in
weeks
which
was
four
of
the
prior
dogs
were
1),
circulating
other
for
splenectomy.
contrast,
in one
cally
durmg
after
unilaterally
remaining
controls
Hentatocrit,
operative
and
following
two
to three
or in Dog
dogs
whose
and
splenic
only
After
removal
of
six
dogs,
for
almost
latter
venous
In
five
6
tiss tie
occasional
not thereafter.
(table
The
required
circulation
former
to
were
one
of
two
(table
and
in the
longer
in
four
made.
with
was
By
a surgi-
seen
only
noted
in the
were
splenectomy.
occurred
three
in one
no
noted
Two
remaining
normoblast
None
Count.
2)
the
and
following
Erythrocyie
was
spleen,
regeneration.
were
animal
normoblasts
one
the
months
observations
nephrectomy.
the
circulating
during
E 15 months
and
depressions
unilateral
weeks
animal,
nephrectomized
Hemoglobin
hematocrit
former
phase.
an
and
of
splenic
or laparotomy.
the
which
altered
splenic
circulation,
the phase
of regeneration
all
of
maintenance
persisted
months,
of
one
counts.
in the
thrombocytosis
removal
normoblastemia
normoblasts
six
tile
autopsy
but
In
intermittent
followed
over
at
In all
differential
had
not
indicating
confirnled
to
a significailt
dogs,
occurred,
preoperative
these
The
splenectomulized
Norinoblasts.
absent
in
tiue
evelltnmal!y
Circulating
were
all
duration
Similar
maximal
following
dogs
regenerated
four
weeks
rerouted
post-
splenectomy
for
within
recovery.
experienced
a
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974
MORTIMER
greater
postoperative
weeks,
except
The regeneration
the
hematocrit
for
depression
one animal
of erythrocyte
which
and
and
developed
hemoglobin
regenerated
in
LORBER
two
mange
and required
values,
in general,
to
five
12 weeks.
paralleled
hematocrit.
Erythrocyte
Morphology.
( target
cytes
first
noted
and
most
)
cells
on
the
Splenectomy
which
usually
second
day
of them
decreased
seventh
postoperative
day.
months.
Thereafter,
although
level
for
the
entire
splenectomy,
greater
rare
decrease
and
Jolly
by
of
found
control
animals.
No
Reticulocytes.
active
The
erythropoiesis,
into
two
phases:
termed
the
turn
of the
termed
greatly
two
regeneration
phase
or
(table
2)
observed
are
regeneration,
to that range
phase.
As
animals,
maintenance
were
usually
phases
better
to
were
the
mean
value
of
expressing
a
were
no
Howell-
in
in
the
all
the
of any animal.
be evidence
of
from
was
divided
blood
loss
subsequent
to
to the procedure
absolute
reticulocyte
ratios
of the mean
means
and
fragility
response
and that
occurring
which
existed
prior
the
the
(table
unchanged
recovery
after
different
value
the erythrocytes
considered
to
during
in
however,
There
reticulocyte
occurring
months
resistance
osmotic
the
to four
a constant
cent
fragility
essentially
within
usually
E 22
bodies,
in
by
about
osmotic
maximal
dogs.
were
postprocedure
phase
at
Dog
changes
indices
each
animal
splenectomized
numerous
in two
1 to 30 per
the
when
more
reached
Howell-Jolly
appreciable
were
reticulocytes
the maintenance
between
individual
each
increased
of
of lepto-
polychromatophilia
was
about
an
value.
of the
or
their
became
including
from
minimal
that
phase
of
hematocrit
in
decrease
erythrocyte
Bartonellae
As
lost
appearance
polychromatophilic
They
number
developed
a slight
leptocytes
dogs.
and
in number
the
in only
bodies,
size
maximum
persisting
dogs
characterized
and
surgery.
of observation,
varied
animals.
All splenectomized
1)
in
in the
large
following
The
period
they
resulted
were
and
the
reand
values
differed
values
during
the
in
the
preprocedure
the
effect
of
the
TABLE
1 -The
Erythrocyte
Response
to Splenectomy
Each underlined
figure represents
the postoperative
day on which
the circulating
normoblasts tended
to stabilize
about the accompanying
mean value. The latter existed until the
next listed day. in Dog C the mean value for time last recorded
day persisted
thereafter
for
the entire period
of observation.
Dog:
Normobiasts:
Indices
A
B
Pre
0
0
Post
0
0
D
0
1
0
2
130#{176}
94#{176}
54
1
1,2020
64
308
0
(MCV,MCH,MCHC):
Pre
Post
Frag:
Osmotic
Pre
Post
% NaCI
No.
C
Days
Observed
#{176}Statistica!!y
significant.
84, 22, 34
64, 22, 34
.52--.36
.44-28
226
66, 22, 33
67, 22, 33
.52-36
.44-.24
197
64, 22, 33
63, 21, 33
66, 24, 36
68, 24, 35
.48-.36
.44-24
.48-.36
190
182
.44-.24
From www.bloodjournal.org by guest on June 17, 2017. For personal use only.
EFFECTS
OF SPLENECTOMY
ON RED BLOOD
975
CELLS
TABLE
2.--Reticulocyte
and Hematocrit
Data
The figures
for maximum
postprocedure
hematocrit
depression
(column
basis of 100 as the preprocedi#e
mean value.
if Dog F, which
developed
mange,
be omitted,
the mean time required
regeneration
of the heinatocrit
to occur (column
7) in the dogs with the
shunt into time systemic
circulation
is reduced
from 40 to 25 days.
1
2
3
4
5
6
4)
7
8
Maintenance
5)05i
m’
o.a
#{176}
n-
.o*
m
.2cs
0,
.‘#{176}
,
i.
1.,
.2
0,
E
Z
88
79
88
79.891*
101.615*
108,128*
3.2*
19.3*
79*
15
63.527*
B
C
25,315
5,265
13,665
22
18
34.191*
76.944*
D
20,906
87
51,725
2.5
17
79,330*
5.6*
3.8*
H
20.400
77
90,190*
L
2,650
71
96.706*
4,4*
36.5*
12.3
18
63,498
4.6
15
47.893*
89,060
68,955
1.6*
Splenectomy
A
Splenectomy
Splenectomy
Splenectomy
Splen.
Splen.
of Shunt
+ Nephr.
0
‘‘-
.
a0
a
I.
5)20
0i.
0-a
-.E
6
9
am.
I
,.i
the
0
,
5i
0
on
for complete
splenic
venous
Regeneration
‘
are
Mean
2.5*
6.5*
14.700
82
88,043
B
30,185
Spienectomized
C
93,308*
122,450*
3.1*
1.8*
Dogs
E
66.487
53,625
78
77
93
74,070*
1.4*
16
10
50,099
81
96,609
2.1
14
68,686
1.4
82,100
42,050
87
74
86.703
112,737*
1.1
2.7*
17
19
78,996
108,037*
2.6*
Phlebotomy
of
Mean
Unilateral
Nephrectomy
Splenect.
Mean
Splenic
Shunt
C
D
of
Venous
into the
1.0
62,075
81
99,720
1.9
18
93.517
1.8
F
G
20,152
6,528
71
71
17.068
26,572
0.9
4.1*
86
23,895
H
I
24,637
61.965
71
77
45,960
146,982*
1.9
2.4*
35
23
14
17,824*
27,355
38,153
1.2
2.7
1.1
28,321
73
59,146
2.3
40
26,807
1.4
21,120
38.960*
39,486*
1.8*
12
19,096
0.9
16.762
87
74
2.4*
20
21.600
1.3
18.941
81
39.223
2.1
16
20,348
1.1
16.950
7,987
23.456
40,575
86
88
82
81
18,387
14,055
37,831
22.365*
1.1
1.8
1.6
13
10
11,677
11,391
0.7
1.4
0.6
36
10
14,303
18.865*
0.5*
22,242
83
23,160
1.3
17
14,059
0.8
J
N
7,590
23,445
0
9,391
89
85
79
21,987*
38,411
26,366*
2.9
1.6
2.8*
19
28
21
8,565
36,566
21,118*
1.1
1.6
2.3
13,475
84
28,921
2.4
23
22,083
1.8
reticulocyte
count
Mean
Phlebotomy
Shunted
of
G
H
Dogs
Mean
Phlebotomy
J
K
L
M
of
Normal
Dogs
Mean
Unilateral
Nephrectomy
Normal
Dogs
of
Mean
Statistlcally
various
that
1.3*
Dogs
Systemic
Circulation
lute
1.8
0.6*
significant.
procedures
on
the
numbers
of reticulocytes.
It has been
reported
in
a
0.6
correlation
exists
iron
between
deficiency
the
than
anemia7
erythrocyte
were
and
level
changes
in the
pernicious
and
the
abso-
anemia8
subsequent
From www.bloodjournal.org by guest on June 17, 2017. For personal use only.
976
MORTIMER
reticulocytosis
despite
in
response
differences
Ilematocnt
in
values
of all
cent
of the preprocedure
whose
sp!enic
venous
hemorrhage
during
hematocrit
depression
main
factors
l)ut
OIlC
by
those
reticulocvte
but
the
of the
per
to
govern
whose
cent
the
had
cyte
elevation
which
( 1.3
were
Of the
F-O
(fig.
13 procedures
1, table
2),
lowed
by
significant
(M)
even
only
two
degree,
only
the
they
were
In marked
durmg
the
0.5
to 2.7 times
the
for
all animals
reticulocyte
times
the
levels
preoperative
a reticulocytosis
being
were
during
was
evidenced
rose
from
53.5
count,
change
per
cent
Dog
which
was
to 60.0
per
cent
his
41.0
per
his
reticulocytes
cent
occurred
occurred
Less
in
of a reticulo-
the
four
normal
control
(logs
were
folOne
at that
although
the
ranged
During
a signi-
control
from
the
the
in
a lesser
(L,M)
all
dog
time.
to
others
two
level,
during
the
2.5 to 6.5
splenectomized
was
not
pro-
0.6
to
4.1
maintenance
mean
being
1.2.
due
maintenance
times,
usually
of
the
day
to dehydration,
from
hematocrit
on
increased
7,000
the
first
more
than
phase
the mean
being
animals
exhibited
greater
a function
13 fold,
also
as
also
of regeneration.
in
hematocrit
increased
magnitude
and
as
animals
developed
a reticumlocytosis
to 36.5 times
the preprocedure
mean,
which
L whose
greater
Considering
In addition,
reticulocytosis
by
nevertheless,
in
phase
preprocedure
phases
mean
times
persist,
elevated
from
value.
Thus,
the
both
in the controls.
That
the observed
sion
12.3.
their
2.3
of reticulocytes
phase.
contrast,
the splenectomized
phase
of regeneration
of 2.5
mean
despite
the
\Vhereas,
that
but
the
in all
experienced
performed
on
dogs
(G,H,I,J,O)
reticulocytosis
during
was
levels
during
regeneration
level,
with
a mean
of 2.0.
the
4.6
no
the
of
procedure.
decrease
phase.
noted
tile
locyte
during
did
loss
rerouted,
regeneration
initial
mean
One
constant
blood
nephrectomy.
during
per
maximal
kept
was
of
than
splenectomy
procedures
on five
maintenance
was
cedures,
the reticulocyte
times
the preprocedure
phase
other
six
(G,O)
value.
thus
81-84
group
of dogs
due to greater
the
same
order
phlebotomized,
that
mean
being
the
group,
preoperative
phase
as their
a significant
animals
during
occurred
reticulocytosis
experienced
decrease
ficant
)
times
this
greater
unilateral
phlebotomized
was
flow
regenerative
in three
dogs
the
venous
undergone
similar,
was
despite
This
was of the
two of them
were
which
of the
fortunate
postoperative
anastomoses,
reticulocytosis
than
preoperatively.
subsequently
when
dogs
were
venous
sp!enic
during
therefore,
lowest
sole exception
rerouted.
In
of the
73
was,
the
groups
The
been
However,
dogs
value
one
was
It
animals,
creation
groups.
four
treatment.
mean.
blood
had
believed
of the
to
individual
LORBER
than
that
hematocrit
after
seen
depressplenectomy
and
whose
to 90,000.
Dog
reticuC had
postsplenectomy
12 times
day,
from
13,000
to 172,000.
It was
cant
noted
elevation
that
of
in
the
three
out
reticu!ocyte
unilateral
nephrectomy
performed
The mean
the phase
reticulocyte
of regeneration
elevation
and
of
four
splenectomized
level
two
amounted
somewhat
could
or
more
less
be
dogs,
induced
months
a further
by
following
to about
a twofold
than
that during
signifi-
phlebotomy
splenectomy.
increase
during
the maintenance
or
From www.bloodjournal.org by guest on June 17, 2017. For personal use only.
EFFECTS
OF
phase.
SPLENECTOMY
This
a higher
was
following
noted
ON
four
of which
had
tion
phase.
which
persisted
locyte
values
were
all
tile
all
coiutrol
20,301
in the
values,
further
Bone
latter.
any
slight
significant
also
marked
the
increased
been
was
noted
following
numbers
phase
overlap
of the
that
the
laparotom
in the
evident
between
(lid
There
not
was
months
unilateral
was
in
and
splenectomy
marrow.
and
ratio
‘
absolute
response
of several
mveloid-erythroid
of reticimlocytes
tile
animals.
suggested
of
reticugroup
former
a
duration,
nephrectomy.
ever
observed
peripheral
despite
blood.
DIScuSsIoN
Platelets.
The
dogs
persisted
and
has
been
spleens
been
experimental
are
longer
and
group,
in
after
in
accord
than
in
its
that
animals
trauma
or
duration
former
has
are
cases
they
are
the
removal
of
with
that
observation
been
splenectomized
finding#{176}’14”
reported
truly
othenvise
to be
free
of
healthy
that
splenectomv
the
It
is,
reticulocvtosis
platelet
levels
spleens.
reticumlocvtosis
to
disease.
observing
unusually
high
note
a matter
comparable
although
traumatized
following
of the
and
in people
whose
a variety
of hematologic
for
more
both
Ek and Ravner,”
were
unable
to
thrombocvtosis
in all
is a vell-known
experimental
following
the
of interest
that
of 18 people,
\rears
data
latter
animals
therefore,
in 11 out
several
both
However,
which
postopen1ti\ely
months
removed
In this
years.’5
thromhocvtosis
to five
observed
have
disorders.
of
significant
up
The
tends
in otherwise
present
to
normal
persist
animals
man.
Circulating
is rare
indicate
Although
Normoblasts.
in dogs
except
that
it
is more
The
tomized
loss has
animals
also been
months
reported
following
removal
of the
ago.
it was
magnitude,
claimed
which
disturbance
of unspecified
picture.20’22
No
after
noted
normoblastemia
the
immediately
after
complete
recovery
in man,17
bumt has not
in
present
findings
splenectomv
several
of
and
the
splenec-
from
postoperative
been
observed
several
blood
years
spleen.13
and
that
persiSte(I
both
that
stated9
splenectomy,
normoblastemia
Hemoglobin,
splenectomy
values
blood
intermittent
been
following
prevalent
thereafter.
Heniatocrit,
it has
immediately
in
Erythrocyte
contrast
resulted
for
to
in
several
a
was
Three
surgical
greater
decrease
months192#{176} to almost
mechanisms
mention
Count.
other
concerned
made
of the
or
of
one
of
red
presence
similar
blood
cell
due
to a
year,21
in maintaining
possible
decades
four
procedures
\
‘
mean
Whereas,
reticulocyte
cellularity
one.
the
splenectomized
any
in
megakaryocytosis
in the
in but
s
#{231}
regenera
maintenance.
during
scarcely
nonsplenectomized
study
of smears
in
the
during
difference
alteration
change
in
wa(
con
the
together,
38,033
there
postoperative
has
No
were
the
and
Careful
nonspecific
Wilich
tlle\’
during
phase
procedures
71,878
than
nonsplenectomized
level
performed
In addition,
reticulocytosis
maintenance
and
emphasizing
Marrow.
C1tIS
the
procedures
i)rocecltmres
with
in eight
of reticulocyte
secondary
regeneration
splenectomized
the
the
during
elevation
and
animals
procedures
during
initial
of
92,502
an
977
CELLS
of
secondary
trols,
the
BLOOD
incidence
similar
Considering
RED
the
of
normal
Barto-
From www.bloodjournal.org by guest on June 17, 2017. For personal use only.
978
MORTIMER
LORBER
Retics
x103
per Cu. mm
200
00
200
HCT4:
71
77
79
87
88
88
HCTm:
53.8
51.5
44.1
46.6
43.9
41.3
DOG:
L
H
B
D
A
C
PROCEDURE:
SPLENECTOMY
_____
10:
HCT4:
73
77
93
74
87
HCTm:
44.7
43.3
46.1
47.0
43.1
DOG:
B
C
E
D
C
PROCEDURE:
PHLEBOTOMY1
NEPHRECTOMY1
400
‘OF
SPLENECTOMIZED
DOGS
300
200
10:
HCT4:
71
71
71
77
74
87
HCTm:
49.0
50.4
48.2
39.0
49.8
47.6
DOG:
F
H
G
PROCEDURE:
100
G
I
H
SPLENIC
SHUNT
-
BYPASSING
PHLEBOTOMY
OF
SHUNTED
DOGS
LIVER
HCT4:
81
82
83
86
79
85
89
HCTm:
DOG:
49.5
M
56.9
L
53.3
K
50.8
J
47.0
0
48.9
N
49.3
J
PROCEDURE:
PHLEBOTOMY
OF
NORMAL
NEPHRECTOMY
OF
NORMAL
DOGS
Fic.
1.-Reticulocyte
data,
postprocedure
minimilum
as the preprocedure
value,
and
preprocedure
columns
encompass
the range
of the reticulocyte
mean
value.
The reticuiocyte
values
are illustrated
procedure,
regeneration
and maintenance
levels.
corner
of the figure
were
not followed
during
only
the
first
two
co!ununs
of
the
usual
DOGS
thr’.
hematocrit
miuean
values
by
three
However,
the
(HCT
hematocrit
and the
latter
enclosed
columns
Dogs
phase
+),
(HCTm)
L
dot
based
representing
and
and,
H
in
therefore,
on
values.
indicates
the
the
100
The
the
pre-
upper
left
each
has
From www.bloodjournal.org by guest on June 17, 2017. For personal use only.
EFFECTS
OF
nellae
SPLENECTOMY
in those
have
failed
loss
during
peared
ON
reports.
RED
In contrast,
to reveal
any
more
splenectomy.
in the
claimed.21
values,
In
addition,
thereby
also
Erythrocyte
the
ly agreed
of
such
The
nothing
cells ) siderocytes
further
shall
be
though
in man,1
they
were
described
rather
scarce
paper.
Reticulocytes.
following
to
be
either
vations,
the
of the
tion
was
bile
pigments,
dog
response
splenectomized
animals
rate,
during
the
the
rise
of
the
succeeding
there
was
suggesting
of
values
cell
new
further
rise
the
reticulocyte
the
experience
per
resultant
of the
bone
cobalt
compounds,28
se does
by
not
normal
elevation
marrow,
erythrocyte
Castle27
always
physiologic
of hemoglobin
as
have
others
or
following
that
mean
They
values
administration
use
by
contrast,
in
elevation,
hematocrit
values,
longer
primarily
interest
numbers
formation
or
noted
reticulocytosis
yeast
delivery
involvement
of caffeine,
of
a great
evidenced
in myelophthisic
the
reticuloto
Of pertinent
of increased
increased
processes.
the
By
no
in
occurred
reticulocyte
manufacture.
the presence
following
the
recovery
was
was
a
alterations
expected
represent
or
time
in
not
of increased
minimum.
at that
and
process
greater
been
circulation
life
excre-
hemin
That
did
hemoglobin
increase
or
the
evidently
yet
a definite
erythrocyte,
of increased
by Minot
and
cells
despite
obser-
formation
hemolytic
of others.23’26
postoperative
definite
urobilinogen
spleens,
to the
their
a
reticulocytosis
was
however,
species
latter
a manifestation
intact
erythrocytes
in
fecal
dogs
reported
or
the
from
blood
loss and
marrow
might
have
with
phase,
been
glycine-labeled
definite
entirely
in those
from
various
the
splenectomy
therefore,
regeneration,
maintenance
of reticulocytes
without
been
red
that
blood
regeneration
ratio
of the
phase
of
a
If it were
than
addition
no
demonstrate
the
C’t
of
in
of hemoglobin
and
studies
all
has
observed
following
reticulocytosis,
as has
the
a manifestation
is the statement
of red
rapid
rate
that
in
with
Al-
splenectomy,
increased,9
the
which
bodies.
found
response
whether
normal
leptocytes
about
of
level
The
unchanged21
to
The
not
in agreement
shown
in his
hemolysis.
erythropoiesis,
more
in the myeloid-erythroid
extent
was
unable
to increased
at a normal
in the
been
Drabkin,
dogs.’3’
to
of removal
and
result
in
general-
of
iron
slightly
as
of
changes
results
formation
reticulocyte
findings,
question
erythrocyte
was
the
no
it is now
striking
the
were
published.
present
previously
decreased.13
splenectomized
on
elevation
of Howell-Jolly
leptocytes,
been
alterations
It has
most
a constant
unchanged,”24
The
implied
destruction.
cytosis
reports
be
other
lag of hemoas had
also
stated#{176}that
occurrence
to
ap-
and,
significant
nonhemoglobin
said
dogs
splenectomy,
are
the
unlike
have
suggested
span
are
diminished,2
necessarily
and
to blood
anemia
reports.9’1#{176}
been
the
contain
said,
Diverse
reticulocytosis.’tm”6’17
among
others13’17’24
of the
control
subsequent
it has
which
in the
previous
following
and,
splenectomy
no
from
and
attributable
dog,
there
was no
cells
and
hematocrit
developments
latter
in this
control
blood
paper
was
duration
than
was
are
,
the
nor
occur
main
than
degree
Although
changes
spleen.10”7
( target
there
in this
anemia
one
red
differing
of erythrocytes
that
the
findings
the
and
the
Morphology.
appearance
the
splenectomized
than
in one splenectomized
globin
regeneration
behind
those
979
CELLS
marked
Neither
to be greater
been
BLOOD
arsenic
in
pernicious
or
From www.bloodjournal.org by guest on June 17, 2017. For personal use only.
980
MORTIMER
anemia.2m’
Unfortunately,
were
not
Another
been
reported
possible
decreased
IlOrteci
by
Jacobsen
and
have
could
by
cytosis
(lid
not
dvspoiesis
not
or
of great
importance
site
Having
increases
i.e.,
as
the
increase
stated
that
reticulocytes.4m
reticulocytosis
was
so that
appears
to,
most
throcytes.
This
therapy
Crtmz
reticulocyte
count,
of
that
first
to removal
spleen.’7
determine
but
coimnts46
reticulum
fotmr of
was inversely
the
splenectomized
of
forms
but
were
not
present
during
long
appreciable
As
unusually
al.42
that
short
more
cells
he
life
than
it
mature
cry-
in discussing
the
been
implied
by
anemia,
before
hematopoiesis
the
the
erythrocvte
through
suhsequentlv
been
long
usual
this
since
release
cells
even
been
repaired,
mechanism
level
to
the
by
emission
thoumgh
has heell
subsequment
observed
following
belief.
resulted
of more
hormonal
adhered
spleen
also controls
of normoblastemia,
reticulocyte
with
not
only
immature
on
basis
splenectomy.
as well,
that
number
differential
the
amount
of the cells were
done
the
normal
phlebotomized
nephrectomized
four
splenectomized
maintenance,
in an increased
ones
the
to the maturity
with
two
of
as before
an
et
marrow
(logs.
must
Minot
man.4
hone
an
represents
normal
has
in the
the
and
the
regeneration
have
that the
presence
proportional
dogs,
two of the unilaterally
suggested
that
in all
regeneration
youngest
classifying
from
of furtiler
he men-
did.
elevated
release
released
by
has
splenectomy
in the
reticulocyte
animals
and
The results
The
an
of a reticumlo-
nonsplenectomized
regulates
compatible
whether
of reticulocytes,
peril1ps
in which
or any
Jacohsen.
It has also
from
acetvlphenylhydrazine
Isaac,43
splenectomy
for a change
is likewise
much
regeneration
to
of whom’7’44’41
believe
the bone
marrow.
The
of the
splenectom
the
values
spleen
by
loss following
as evidence
The
possiprobably
animalstm
in reality
returned
IlematOcrit
the
are
expressed
h
of
reticulo-
normally,
animals
without
during
been
(logs
the
although
are,
reticulocytes
released
propounded
some
from
the blood
presented
To
has
possibilities,
(luring
in
count
anemia,
and
that
in recovers’
and
concept
others,
of cells
then
on
removed.
loss
true,
pam
agreed
or undergo
nonsplenectomized
were
reticulocyte
counts
The
in
cells
belief
hemoglobin
means,
blood
ye-
normoblasts.
of reticulocytosis
without
evidence
let another
apparent
paradox
now
blood
cells
of pernicious
who
noted
been
have
been
unaccom-
that
of
or splenectomized
has
all red
red
here
states,8
life
numbers
erythrocytes7
two
intact
noted
that
a small
a(lult
latter
should
have
occurred
not the case,
unless
tile
usually
span.
If
in
maturation
autoinimune
from
been
of
has
the
usually
.:vi
increase
could
as
increased
dm
It is assumed
into
destruction
regeneration
two
rate
The
some
occurrence
levels,
as has
been
values.1’
in either
of blood
considered
of erthrocyte
tioned;
It has
in
have
to
processes
increase
erythrocytes,
apparent
develop
destruction.
the
would
increased
either
of
one
their
cell
an
operative
important
cvte
of red
could
theoretically
This
hemolytic
reticulocyte
)
the
for
rel)resent
reticulocytoses
ble
Plum.’
accounted
concomitant
apparent
beyond
elevations
out
( maturation
and
reticulocytes
to rule
papers.
of the
“ripening”
circulating
I)aniel
studies
in those
cause
LOBBER
dogs
about
as controls
dogs
during
twice
Of
the
as
of
on
(table
3).
the phase
many
controls,
of
the
only
one
From www.bloodjournal.org by guest on June 17, 2017. For personal use only.
EFFECTS
OF
dog
( K)
the
left
largest
is not
locytes
SPLENECTOMY
developed
by
splenic
blood
was
reticulocyte
counts
It
differential
blood
cell
of
should
the
erythrocytes
animals.
maturation
reticulocytes
disappear
be true,
of the
from
tion
or
into
to
adult
expect
the
the
It
numbers,
3.-Differential
noted
to
peruse
the
in
latter
of
as
many
the
belief
that
Regeneration
Maintenance
DogC
Presplenectomy
Regeneration
Maintenance
DogD
Presplenectomy
Regeneration
Maintenance
MEAN
(splenectomy)
Presplenectomy
Regeneration
Maintenance
.01
as
25,000
did
sug-
were
removal
of the
release
of erythroid
cells
the delayed
reticulocyte
since
the
the
time
younger
before
splenectomy
the
absence
they
in
from
the
the
the
bone
may
the
they
of
normal
case,
marrow
then
internormo-
splenic
complete
peripheral
be
and
one
would
normally
blood,
albeit
to the
spleen
in Four
Splenectomized
and Four
of Counts
of 25 Reticulocytes)
increase
in time younger
reticulocytes
dogs during
the phase of regeneration
inhibi-
maturation
their
Counts
Ofl a Series
perhaps
to
Control
comDogs
(with
more
as compared
animals.
RETICULUM
Most
Maintenance
DogB
Presplenectomy
white
counted
splenectomy
than
the
nonsplenectomized
RETICULUM
Most
Least
2.8
6.7
4.3
repeated
data
of reticulocytes
which
were
Note
time two to almost
threefold
reticulum)
in all time splenectonmized
to a sinmilar increase
in but one of time control
DogA
Presplenectomy
Regeneration
1, whose
when
his
per cent),
of the differential
per slide were
following
loss
in
release
of
organ
way
Reticulocyte
(Based
Dog
times
( 9.1
reproducibility
following
increased
normoblasts
their
in
Nevertheless,
to support
because
an
If the
on
having
increases,
maturation.
to the
of
circulating
the
as good as that
25 reticulocytes
of
were
marrow
removal
counts
following
splenectomy,30’36
the circulation,
the longer
due
slight
That
a shift to the
numbers
o1 rctieu-
.
of large
resulting
in earlier
thus,
partly
explain
their
being
bone
that
tend
marrow,
might,
that
but
(0 )
releage
reticulocytes
from
blood
noted
entering
erythrocytes.
to find
in small
TABLE
bone
had
reticulocyte
reticulocytes.
it would
findings
as either
times
25
completing
by
Many
of the
emphasized
at
three
the systemic
circulation,
at
( 3.1 per cent ) and 409,000
immature
recovery
that
was
are on
other
preoperatively
differential
finding
spleen
affects
the
into
the circulation.
blasts
be
of more
during
If this
the
as
counts
was not
because
only
necessity
before
gest a release
usually
seen
the
981
CELLS
shift;
diverted
into
were
281,000
reticulocyte
count,
perhaps
because
BLOOD
again
as many
manifestation
indicated
respectively.
RED
a similar
being
half
an inherent
was
preted
ON
17.6
15.8
16.8
4.6
2.5
3.9
DogJ
Prephlebotomy
Regeneration
Maintenance
DogK
Prephlebotomy
3.3
14.4
7.3
9.3
4.6
14.8
17.2
0.9
3.2
3.6
7.1
4.9
18.4
15.7
18.7
3.0
2.0
1.4
2.3
5.3
17.7
17.2
5.0
2.5
2.6
18.4
4.0
3.0
17.0
5.0
Preprocedure
7.1
4.1
15.9
17.8
2.0
3.1
Regeneration
Maintenance
Regeneration
Maintenance
DogO
Prenephrectomy
Regeneration
Maintenance
DogN
Prenephrectomy
Regeneration
Maintenance
MEAN
(control)
Least
2.7
3.0
5.8
19.0
19.7
14.5
3.3
2.3
4.7
3.4
7.8
7.2
12.3
15.6
15.5
9.3
1.6
2.3
4.0
6.0
3.0
17.0
15.3
18.0
4.0
3.7
4.0
2.3
2.4
1.5
17.7
19.4
19.0
5.0
3.2
4.5
3.1
4.8
4.4
16.5
17.4
16.8
5.4
2.7
3.9
From www.bloodjournal.org by guest on June 17, 2017. For personal use only.
982
MORTIMER
plete
their
ing
development.
normoblasts
addition,
two
(if
splenectomy
The
are
absent
the
dogs
day,
fact
that
or very
had
despite
in nonspleneetomized
rare
no
suggests
depression
which,
within
18
animals
that
of
that
of
on
the
or
reticulocytes,
continued
rather
suggests
had
whose
to circulate
that
occurred,
into
Bone
the
Marrow.
increase
a change
permitting
precursors
and
are
thus
release
span
one
mechanism
two
had
days,
was
from
only
In
post-
they
or
of maturation
not
case.
first
a
It is somewhat
difrepresented
a passive
is but
site
of
in
but
2 of
Pearce
after
have
appeared.
of
more
in
release
in dogs
‘
evidence
hyperplasia
in the
in 1914,
marrow
‘
particular
usual
the
marrow
no
life
their
the
more
the
but
that
removed.
bone
It
marrow
younger
red
cell
circulation.
Although
of red
normal
because
circulatthe
surgery,
12 and 13 fold increase
in their
circulating
reticulocytes.
ficult for the author
to conceive
that this increase
merely
backlog
is not
hematocrit
hours
LORBER
Pepper47
agreement
present
changes
with
patients,
noted
no
subsequent
The
marrow
bone
12
and
splenectomy,
data,
the
recent
of
of bone
however,
subsequent
a number
macroscopic
reports
to
observation
after
afford
splenectomy,
of erythroid
removal
of the
years
spleen.16
SUMMARY
In splenectomized
were
the
1.
There
was
no
2. A
cytosis.
tion
persisted
There
is a suggestion
more
of
target
in
all
in
some
of
significant
the
bone
in response
alterations
probably
in
largely
the
from
The
hemo-
same
rate
as in
animals,
circulation.
most
are
from
and
the
due
marrow,
permitting
rather
changes
than
merely
of
their
are
to
of
due
the
than
on
a fall
bone
into
specifically
in
bone
hematocrit
except,
marrow
of
the
to
of
the
reticulocytes.
incurred
at surgery.
are unaccompanied
inhibition
entry
removal
from
rather
erythropoiesis,
the
normal
earlier
persisted
rare.
released
cells
dependent
the
re-
developed
which
of increased
ratio
a diminution
osmotic
also
were
the blood
loss
normoblastemia
myeloid-erythroid
to
regenera-
noted.
red
is not
regeneration
reticulocytosis
increased
Most
erythrocytes
mature
or even
during
regeneration,
were
of hemolysis
aforementioned
itself,
loss
by
bodies
changes
thrombo-
animals.
dogs.
active
was
associated
circulating
control
Howell-Jolly
marrow
reticulocytes
the
accompanied
reticulocytosis
a reflection
perhaps,
in part during
3. Postsplenectomy
portal
elements
erythrocyte,
circulating
splenectomized
during
to blood
Postsplenectomy
is not
4.
erythroid
at the
than
reticulocytes
of the
cells
the
of
duration
the
animals.
It is suggested
that:
1. In nonsplenectomized
2.
of
occurred
number
those
particularly
thereafter
No
that
than
present
normoblastemia,
the
a longer
numbers
were
marrow
depression
regeneration
in
for
immature
Varying
4.
and
increase
which
were
3.
spleen
on the
dogs.
sistance
lease
observations
postoperative
values,
significant
noted,
by
following
unusual
or hematocrit
control
and
CONCLUSIONS
made:
globin
the
dogs
AND
splenic
the
and
erythroid
are
re-
circulation.
removal
blood
of
from
the
the
From www.bloodjournal.org by guest on June 17, 2017. For personal use only.
EFFECTS
OF
SPLENECTOMY
ON
RED
BLOOD
SUMMARIO
In
canes
respecto
splenectomisate
al elementos
1. Occurreva
con
del
le mesme
2.
notate
circulante.
Iste
bocytosis.
Certe
durante
le
sequente
inusual
observationes
depression
hemoglobina,
o del
intensitate
Esseva
INTERLINGUA
esseva
facite
con
erythroide:
nulle
erythrocytos,
IN
le
983
CELLS
como
un
in le canes
augmento
augmento
observationes
esseva
le
valores
regeneration
del
se
faceva
de controlo.
in
plus
pare
in
e le
significative
persisteva
regeneration
post-operatori
hematocrite,
le
numero
longemente
indicar
que
plus
immatur
a oculo
de
del
que
le
reticulocytos
le associate
throm-
reticulocytos
que
illos
circulante
in
le
animales
de
controlo.
3.
Vane
nimmeros
resistentia
In
le
4.
del
presente
casos
specialmente
in certes
Esseva
notate
sequente
1. In
cellulas
esseva
majoritate
blastemia,
sequentemente
Le
de
osmotic,
il habeva
nulle
non-splenectomisate,
ossee
in responsa
reticulocytos.
post
3.
de
de
resulta
per
in
normo-
persisteva
esseva
medulla
de
del
de
non
subrar.
ossee.
erythrocytos
ab
ab
simplemente
per
matur
del
o mesmo
le
de
un
regeneration
chirurgic.
non es accompaniate
le medulla
ossee
in
le
le
liberate
cellulas
abassamento
durante
le intervention
mesura
un
hemolyse
e normoblastemia
myeloido-erythroide
grande
es
de
de
parte
per
erythrocytos
sanguine
depende
reflexion
diminution
medulla
de illos a in le circulation.
alterationes
es effectuate
se e non
le circulation
perdita
tin
causate
probabilemente
splen
del
le majoritate
a tin
post-splenectomic
del proportion
normal
del liberation
entrata
plus precoce
4. Le mentionate
del
de
active.
Isto
de Howell-Jolly
excepte-forsan-in
sanguine
Reticulocytose
alterationes
augmentate
a splenectoinia.
disveloppamento
significative
post-splenectomic
e non es
erythropoiese,
perditas
de
subjicite
es proponite:
animales
accelerate
tin
regeneration
Corpores
alterationes
per le medulla
plus tosto
que
2. Reticulocytosis
valores
del hematocrite
accompaniate
le canes
etiam
durante
le
del animales.
conclusiones
aye,
in omne
del
con
le
specificamente
le elimination
effecto
per
de
e
inhibition
de
le
sanguine
un
ablation
splenic
ab
portal.
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1958 13: 972-985
The Effects of Splenectomy on the Red Blood Cells of the Dog with
Particular Emphasis on the Reticulocyte Response
MORTIMER LORBER
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