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Phagocytosis of granules from disrupted mast cells.

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PHAGOCYTOSIS O F GRANULES
FROM DISRUPTED
MAST CELLS1
DOUGLAS E. SMITH AND YEVETTE S. LEWIS
Division of Biological and Medical Research, Argonne Natioiial Laboratory,
Lemont, Illinois
TW ENTY- FOU R FIGURES
I n the connective tissue throughout the mammalian organisni the tissue mast cell is characteristically round or
spindle-shaped and is filled with large cytoplasmic granules
that stain metachromatically with toluidine blue. Occasionally, however, there occur apparently abnornial mast cells
which show singly, or in combination, conglomeration of metachromatic granules, colorless or metachromatic vacuoles, or
alterations in shape. Such cells make up 1-2% of the total
mast cell population of untreated animals (Smith and Lewis,
'541, 54b, '55) and become significantly greater in number
in 1,cptone shock (Wilander, '3S), stimulation of urticaria pigmentosa lesions (Drennan, '31) , and treatment with x-rays
(Smith and Lewis, '53, '54a), nitrogen mustard (Schoch and
Glick, '%), bacterial pyrog-ciis (Stuart, '51), cort,isone (AsboeIIaiisen, '52 ; Cavallcro and Braccini, '51 ; Smith and Iiewis,
'<j4h, 'Xj),aiid ACT11 (Smith and Lewis, '54b, '5.5).
Recently Higginbotham et al. ('56) reported that the fibroblasts of the connective tissue selectively take up and dispose
of the cytoplasmic granules shed from mast cells disrupted
by various treatments. Tb seemed to us that such fibroblasts
closely resemble certain of the above-mentioned abnormal
mast ce11s. It appeared possible t o us, moreover, that some o r
all of the mast cells designated hitherto as abnormal o r degen'This work was performed under the auspices of the U. S. Atomic Energy
Coniniission.
93
94
DOUGLAS E. S M I T H AND YEVETTE S. LEWIS
erated might be in reality fibroblasts or other cells which
have ingested cytoplasmic granules from disrupted mast cells.
The present paper is concerned with a microscopic examination of (a) the fate of the cytoplasmic granules shed from
mesenteric mast cells broken up as a result of exposure to distilled water and of (b) abnormal mast cells in several tissues
from normal and treated animals.
METHODS
The mesentery of the Sprague-Dawley rat (male and female, weighing approximately 200 gm) was employed as the
site for studying the fate of the cytoplasmic granules released
from disrupted mast cells. Break-up of the mast cells was
brought about by the method of Fawcett ('54) whereby
distilled water is injected intraperitoneally into the unanesthetized animal. Rats were sacrificed by ether anesthesia at various times (table 1)after the administration of distilled water.
Whole mounts of mesentery were prepared as previously described (Smith and Lewis, '53). The tissues were fixed in
80% ethyl alcohol and stained with toluidine blue (.025% in
50% ethyl alcohol) or May-Grunwald stain. The preparations
were examined under the light microscope at magnifications
up to 1000 diameters. A similar study was carried out on fixed
whole mounts of mesentery from 150-gm male Sprague-Dawley
rats treated with 48/80.2 Twenty milliliters of Tyrode's solution containing 1mg of 48/80 was injected intraperitoneally,
and portions of mesentery were taken from animals sacrificed
at 10 minutes, 2, 6 and 24 hours after injection.
In addition to observations on fixed tissue, a microscopic
study was carried out in. vivo on the mesenteries of rats injected intraperitoneally with distilled water. See table 1 for
the schedule of the observations. F o r this study the animals
were anesthetized with Nembutal R (sodium pentobarbital)
and a loop of intestine was withdrawn through a mid-ventral
incision. The rat was then placed on its side on a holder con' A polymer of N-methylhomoanisylamine and formaldehyde, supplied through
the courtesy of Dr. Edwin J. de Beer of Wellcome Research Laboratories.
20
10
I n vivo preparations
In vivo preparations
x
X
x
15
x
X
x
30
X
x
x
1
x
x
2
x
x
4
x
x
6-7
Hours after injection
x
x
1
TIMES OF SACRIFICE AND 0BSER.VATION
Observations were made on 3 animals at each time interval checked.
20
10
injection
ML
INJECTED
Fixed and stained whole mounts
TISSUE
Minutes a f t e r
WATER
DISTILLED
Schedztle of observations of rat mesentery after injection o f distilled water
TABLE 1
x
x
x
x
3
x
x
4
x
5
Days after injection
2
x
X
7
E!
Q
96
DOUGLAS E. SMITH A N D YEVETTE S. LEWIS
taining a small lucite (methyl methacrylate) block over which
the mesentery w7as laid in such a manner that it was transilluminated on the microscope stage. Oxygenated Tyrode’s
solution a t 37°C was allowed t o drip continually on the mesentery. Vital staining of these preparations was readily accomplished by adding toluidine blue to the Tyrode’s solution.
Observations were made at magnifications up to 900 dianieters with water immersion objectives, and the phenomena
were recorded by time-lapse cinematography and still photography.
hlicroscopic examination was made of the abnormal mast
cells in whole mounts of fixed and stained mesentery and skin
of the Sprague-Dawley rat (200 gm) and mesentery, skin and
cheek pouch of the Syrian hamster (100 gm) at various times
after s-irradiation o r administration of ACTH o r cortisone,
Similar observations were made in untreated rats and liamsters. The s-irradiations consisted of single total-body exposures to 600 r or 1200 r to rats, and 600 r, 715 r or 1200 r to
hamsters. The radiation factors were 250 Bv, 15 ma, 0.; nim
Cu, and 3.0 mm Bakelite filters, 26.7 ern target distance, 1.5
mm Cn half-value layer and 215-225 r per minute. Cortisone
was administered subcutaneously or intramuscularly daily
for 1-4 days in dosages of 200 nig/kg. ACTH ~ - a 7injected
subcutaneously for 1-4 days in dosages of 100 or 400 I.U./kg.
T n a few experiments in vico examination., wcre made of
mesentcry of rats csposed to s-irradiation o r treated with
cortisone or ACTH.
RES1T~T-S
Fired and s t c( in e (7 7 i cs P+Z t P r 71 o f rci t .s i i z jcc t c d in f ra p e r it o n e a11i j 1 o i t h cl isti11 (1 u‘c, t-. Tithin 15 minutes after the
(2
administration of distilled water, the limitjng boundaries of
many mast cells are no longer intact and metachromatic
granules are seen free in the surrounding tissue. Some of
these mast cells are completely disrupted and their granules
occupy a space 11h to 3 times that of the volume of an intact
mast cell; the remainder show the loss of a fern granules into
the surrounding tissue. With the passage of time after in-
PHAGOCYTOSIS O F MAST CELLS
97
jection more and more mast cells are found in the disrupted
state until at 1-2 hours most of them are broken up and the
mesentery is filled with mast cell granules. No changes are
found in cells other than mast cells. No changes are apparent
in the staining properties of the disrupted mast cells and
their scattered granules except for the presence of an occasional enlarged, pink granule.
The fate of the freed mast cell granule is apparent as
early as 15-30 minutes after distilled water injection. At
this time metachromatic granules of the same size and shape
as the cytoplasmic granules of the mast cell are found in
macrophages, fibroblasts and leucocytes in the vicinity of the
broken mast cells. As time passes and the number of disrupted
mast cells increases in the preparations so does the number
of macrophages, fibroblasts and leucocytes that contain metachromatic material. At 6-7 hours leucocytes are present
throughout the tissue in impressive numbers (up to 200 and
more per 0.0625 mm2 as compared with 0-5 per 0.0625 mm2 at
15-30 minutes). Many of the leucocytes contain no metachromatic granules. The distribution of macrophages and
fibroblasts is not uniform throughout the tissue. Macrophages are found in large numbers and fibroblasts are few
near blood vessels, whereas fibroblasts are more numerous
away from the blood vessels. Many of these cells contain
metachromatic material. By 7 hours very few metachromatic
granules are found free in the tissues, and at 24 hours no
metachromatic material is found outside of fibroblasts, macrophages and leucocytes. Many of these cells contain such large
numbers of metachromatic granules at tv7o hours after treatment with distilled water that their nuclei are obscured, and
they thus resemble intact mast cells. Others show few metachromatic granules in their cytoplasm even at 24 hours.
Illustrations of metachromatic material in fibroblasts, macrophages and leucocytes are found in figures 1-16.
Alterations in the ingested metachromatic material are obvious as early as two hours after treatment. I n some macrophages, fibroblasts and leucocytes one now finds no discrete
98
DOUGLAS E. SMITH AND YEVETTE S. LEWIS
granules but rather metachromatic vacuoles, some of which
stain light red, others, purple. An individual cell may contain
from one to dozens of such vacuoles. By 7 hours more than
half of the macrophages, fibroblasts and leucocytes have at
least some of their metachromatic material in this state, with
the remainder in the form of discrete granules. At the end
of 24 hours most of the cells have practically all of their
metachromatic material in the form of vacuoles. Thereafter
progressively fewer cells are found to contain metachromatic
material in any form. By 4-5 days no such cells are found
and the mesentery has a normal appearance except for a complete absence of mast cells and an increased number of eosinophils. Eosinophils are found in increasing numbers up to
24 hours after treatment and are still present in elevated
numbers even at the end of one week. The eosinophils do not
contain metachromatic inclusions.
After the intraperitoneal injection of 48/80, large numbers
of cytoplasmic granules were released from the mast cells.
These granules were ingested and disposed of by macrophages, fibroblasts and leucocytes in the same fashion as
those released by treatment with distilled water.
Mesentery--in. vivo study. Within 10 minutes after the
intraperitoneal injection of 20 ml of distilled water, many of
the mast cells are disrupted and their granules are scattered
in the surrounding tissue. Here the granules show Brownian
movement and freely bounce about. At this time macrophages, fibroblasts and white blood cells containing some
metachromatic granules can be found. As more mast cells
break up, macrophages which were previously sessile and
unnoticed become motile and evident. They move toward
broken mast cells and engulf the granules with long protoplasmic extensions. Leucocytes also make their way toward
the loose granules by amoeboid movement. They do not surround the free mast cell granules with long pseudopodia but
rather show a marked bubbling of the cell surface when in
contact with the free granules; shortly thereafter the mast
cell granule is seen within the leucocytc. Lcucocytes are
PHAGOCYTOSIS O F MAST CELLS
99
frequently seen to traverse the walls of small blood vessels,
and they are accumulated in large numbers in the tissue alongside the vessel walls at 2 4 hours after injection of distilled
water. By 6-7 hours large numbers of leucocytes are visible
at considerable distances from blood vessels throughout the
mesentery. Fibroblasts show neither ambulatory nor surface
activity. Free mast cell granules appear to make their way
to fibroblasts by Brownian movement. After bouncing against
the limiting surface of the fibroblast for a few minutes, the
granules penetrate it and are seen within the cytoplasm of
the fibroblast. The morphology and staining properties o€
the mast cell granule do not appear to be changed upon their
entrance into the cytoplasm of the macrophage, fibroblast or
leucocyte. Once inside these cells, the granules continue to
show Brownian movement within the bounds of vacuoles. The
vacuoles vary in size and in the number of granules which
they contain. Later the granule (s) may disappear leaving
the vacuole filled with material which appears uniformly
purple. The phenomena observed after injections of 10 ml of
distilled water were essentially the same as those above, except that some cells lose only a few granules and appear intact
thereafter while other mast cells seem completely unaff ectcd
by the treatment. By the end of 24 hours when all of the
free mast cell granules have been ingested, no further activity
is evident and the living preparations thereafter closely resemble the fixed mesenteries of the same time period. 11lustrations of typical findings in the living preparations are
found in figures 17-24.
Fixed and staified tissues from untreated rats and hamsters. Normal skin, mesentery and cheek pouch clearly show
macrophages and fibroblasts which contain metachromatic
granules and vacuoles. These cells are similar in appearance
to those seen in the fixed and stained preparations of mesentery exposed to distilled water. One o r two such cells occur
for every 100 mast cells. Cells indistinguishable from mast
cells but containing conglomerations of granules and metachromatic vacuoles were found but rarely. Leucocytes con-
100
DOUGLAS E. SMITH A N D YEVETTE S. LEWIS
taining metachromatic material are not found in these
preparations.
Fixed and staimed tissues from rats and hanzsters treated
with z-radiation, cortisone and ACTB. The skin, mesentery
and cheek pouch of animals subjected to x-irradiation or injection with cortisone or ACTH showed large numbers of
macrophages and fibroblasts containing metachromatic granules and vacuoles. These cells were similar to those found
in the fixed and stained mesenteries exposed to distilled water.
The number of such cells varied with duration and severity
of treatment. I n the most striking instances the number of
macrophages and fibroblasts containing metachromatic material equalled or even exceeded the number of intact mast
cells found. As with the same tissues from untreated rats
and hamsters, cells indistinguishable from normal mast cells
except for conglomerations of granules and metachromatic
vacuoles were found occasionally. Leucocytes with or without
metachromatic material were seldom found. The i m vivo observations on the mesentery of x-irradiated and cortisone and
ACTH rats were in essential agreement with those on fixed
preparations.
DISCUSSION
The finding in both living and fixed mesentery that the
intraperitoneal injection of distilled water completely disrupts the tissue mast cells but causes no conspicuous effects
in other cells agrees with the report of Fawcett ('54, '55) on
fixed tissue. The observation that fibroblasts, macrophages
and leucocytes ingest mast cell granules immediately after
treatment with distilled water further indicates that general
cellular damage does not follow this procedure.
The present finding of increased tissue eosinophils after
distilled water treatment is at variance with the report of a
decrease by Fawcett ( '55). It is possible that this discrepancy
is due to the different strains of rats employed in the t ~ r o
studies.
PHAGOCYTOSIS O F MAST C E L L S
101
Both the living and fixed mesenteries of the rat clearly
show that fibroblasts, macrophages and leucocytes ingest and
dispose of the cytoplasmic granules released from mast cells
disrupted by distilled water. Prior reports have suggested
that macrophages alone (Pawcett, '54) or fibroblasts alone
(Higginbotham, '56) perform this phagocytic action, and,
to the best of our knowledge, leucocytes containing metachromatic material have not been reported.
The differential distribution of fibroblasts and macrophages with respect to their location near or away from
blood vessels agrees with previous findings (Maximow and
Bloom, '52). This differential distribution may account for
the failure of prior workers to note that both macrophages
and fibroblasts take up loose mast cell granules.
The findings after the injection of 48/80 indicate that the
debris of mast cells disrupted by different procedures is phagocytized in the same fashion.
The macrophages and fibroblasts containing cytoplasmic
granules from mast cells disrupted by distilled water or by
48/80 are grossly quite similar in appearance to the "abnormal" mast cells previously described as occurring in small
numbers in normal tissue (Smith and Lewis, '54b, '55) and i n
quantity after treatment with ACTH (Smith and Lewis, '54b,
'55) cortisone ( Asboe-Hansen, '52 ; Cavallero and Braccini,
'51; Smith and Lewis, '54b, '55) o r x-radiation (Smith and
Lewis, '53, '54a). I n the present study, examination of normal
and treated (ACTH, cortisone and x-irradiated tissue
revealed that macrophages and fibroblasts that contain metachromatic granules and vacuoles occur with frequencies similar to those described for "abnormal" mast cells in prior
studies. Only rarely does there appear a cell which contains
conglomerated metachromatic granules or metachromatic
vacuoles but which in other respects is a typical mast cell.
Thus practically all of the cells previously designated as
"abnormal" or degenerated mast cells (Wilander, '38 ; Cavallero and Braccini, '51 ; Drennan, '51 ; Stuart, '51 ; Schoch and
Glick, '53 ; Smith and Lewis, '53, '54a, '54b, '55) may well be
102
DOUGLAS E. SMITH A N D YEVETTE S. LEWIS
macrophages, fibroblasts and leucocytes that contain metachromatic material. Previous failures to identify these cells
correctly probably resulted from the limitations of the metachromatic stains employed. The cellular detail brought out
by the May-Grunwald stain used in the present study makes
possible more accurate identification of the various cell types.
It was previously considered (Wilander, ’38 ; Cavallero and
Braccini, ’51 ; Drennan, ’51; Stuart, ’51; Schoch and Glick,
’53; Smith and Lewis, ’53, ’54a, ’54b, ’55) that the “abnormal”
mast cells reflected a series of degenerative changes that occur spontaneously only rarely but with great frequency after
certain treatments, the final step in degeneration being break
up of the cell with release of the cell contents into the surrounding tissue. The present findings indicate that while such
a degenerative change occurs, the usual process, which is seen
only occasionally in untreated animals but frequently after
the aforementioned treatments, is one of disruption of the
mast cells with the freeing of their granules into the surrounding tissue where macrophages, fibroblasts and leucocytes
quickly ingest them. Since one is concerned with disruption
of mast cells in both the past and the present interpretations,
the proper identification of the cells does not require basic
revision of conclusions drawn from experiments (Wilander,
’38 ; Cavallero and Braccini, ’51 ; Drcnnan, ’51; Stuart, ’51 ;
Schoch and Glick, ’ 5 3 ; Smith and Lewis, ’53, ’54a, 54b, ’55)
in which these cells were not properly identified. Since the
remnants of a single disrupted mast cell may be taken up by
more than one macrophage or fibroblast, however, the prior
accountings (Smith and Lewis, ’54a, ’54b, ’55) in terms of
“abnormal” mast cells would tend to give a somewhat exaggerated idea of the number of mast cells being destroyed hy
treatment. Although it would appear that absolute quantitation of mast cell destruction as a result of treatment is not
possible from a measure of the increase in macrophages and
fibroblasts and leucocytes in a preparation, it is certain that
one can employ such counts to give qualitative and perhaps
rough quantitative information on the effectiveness of cxperi-
PHAGOCYTOSIS O F MAST CELLS
103
mental procedures. Failing to account f o r phagocytes containing metachromatic material, i.e., including them as mast
cells, obviously leads to errors in interpreting data involving
changes in mast cell number. It is possible that the increase in
mast cell reported by Bensley ('52) and by Schiirer ('46)
after histamine and heparin injection respectively are examples of such a misinterpretation. Higginbotham et al. ( '56)
have called attention to the possibility that such errors arise
from failure to account for fibroblasts which have ingested
inetachromatic material. Earlier claims (Devitt et al., '54)
that similar treatments produce no change in mast cell number may be explained in somewhat the same way. I n these
studies phagocytes containing metachromatic material were
not identified and cells of the "abnormal" type were considered to be artifacts of handling and counted as normal mast
cells.
The functional significance of the disruption of mast cells
by various treatments is not clear. I n general, such disruption
with release of granules into the surrounding material has
been considered t o constitute secretion (Wilander, '38 ; Cavallero and Braccini, '51 ; Drennan, '51 ; Stuart, '51 ; Schoch
and Glick, '53; Smith and Lewis, '53, '54a, '54b, '55). Indeed,
large amounts of histamine are released into the peritoneal
fluid after injection of the rat with histamine liberators and
distilled water (Fawcett, '54). On the other hand, solutions
of toluidine blue and protamine sulphate cause similar release
of histamine without morphological changes in the mast cells
of the mesentery (Smith, '58). It is possible that the latter
is the usual method of release of material (secretion) from the
mast cell and that rupture of the cell with release of granules
occurs only when marked changes occur in its environment.
SUMMARY
Microscopic examination was made of the fate of the cytoplasmic granules shed from mesenteric mast cells disrupted by
intraperitoneal injection of distilled water into the rat. Both
fixed and living preparations of mesentery were employed.
104
DOUGLAS E. SMITH AND YEVETTE S. LEWlS
I n addition microscopic examinations were carried out on the
“abnorma1” or degenerated mast cells of several tissues from
control and treated (x-radiation, ACTH, o r cortisone) rats
and hamsters.
Treatment with distilled water completely disrupted the
mast cells of the mesentery, while other cells appeared to be
undamaged. Fibroblasts, macrophages and leucocytes were
observed to take up and digest the shed mast cell granules.
These phagocytes were quite similar in appearance to the
“abnormal’ o r degenerated mast cells previously described as
occurring in normal and treated animals.
Mast cell granules released as a result of 48/80 administration met the same fate as those freed by distilled water treatmcnt.
The examination of “abnormal” mast cells in control and
treated animals indicated that most of these were macrophages and fibroblasts that contained nietachromatic material.
The results are discussed in relation to the question of the
phagocytosis of disrupted mast cells and of the significance
of the “abnormal” or degenerated mast cell.
LITERAT URF CITED
ASBOE-HANSEN,
G. 1952 The mast cell a n object of cortisone action o n connective tissues. Proc. SOC.Exp. Biol. Med., 80: 677-679.
BENSLEY,
S. H. 1952 On the origin of mast cells. Anat. Rec., 122: 310.
C., AND C. BRACCINI 1951 Effect of cortisone on the mast cells
CAVALLERO,
of the rat. Proc. SOC.Exp. Biol. Med., 7 8 : 141-143.
DEVITT, J. E., P. B. SAMUELS,w. J. PIROZYNSKI AND D. R. WEBSTER 1954
Morphology of tissue mast cells; the frequency of artifacts and the
influence of certain biologic agents. Am. J. Pathol., 3 0 : 391401.
DRENNAN,J. M. 1951 The mast cells in urticaria pigmentosa. J. Pathol. Bacteriol., 63: 513-520.
FAWCETT,
D. W. 1954 Cytological and pl~~lrniacological
observations on the release of histamine by mast cells. J. Exp. Med., 100: 217-224.
1955 A n experimental study of mast cell degranulation and regeneration. Anat. Rec., 121: 29.
AND W. S. S. SEE 1956 F a t e of shed
HIGGINBOTHAN,
R. D., T. F. DOUGIIERTY
mast cell granules. Proc. Soe. Exp. Biol. Med., 92: 256-261.
MAXIMOW,A. A., AND W. BLOOM1952 Textbook of Histology. Sixth Ed.
p. 57. W. B. Saunders, Philadelphia.
PIIAGOCYTOSIS O F MAST CELLS
105
SCHOCII,
E. P., AXD D. GLICK 1953 The effect of cold stress, ACTH, cortisollc,
pyroyeri and nitrogen mustard 011 tissue mast cells in the skin and
subcutaneous tissues of the rat. J. Invest. Dermatol., 20: 119-131.
S C H ~ E W.
R , 1946 Speicheruiig voii Heparin in den Zelleri des Iieticuloeiidothcls. Helv. Mecl. Aeta, 13: 161-171.
S ~ I I T I II).
, E. 1958 The nature of tlic secretory activity of the mast cell,
Amer. J. Physiol., 293: 573-575.
~ M I ' I T I , 1). E., AND Y.
LEWIS 1953 Effects of total-body x-irradiation o n
the tissue mast cell. Proe. Soc. Exp. Biol. Med., 6'2: 208-212.
-~
1954:~ Iiiflueiiec of total-body x-irradiation upon tissue iuast cell
iiuiribcr. Ibid., X5 I 306-307.
1954b Iiiflueiice of hypophysis mid adrrrinl cortex upoil tissue mast
crll of the rat. Ibitl., 8 7 : 515-518.
1955 Tiiflueiice of hypophysis and of adreii:rl cortex upon tissuc
mast cell of thc hamster. Ibid., 88: 631-634.
S T l i A R T , E. G.
1951 C'oiiiieetive tissuc in:~.st-cell response to b:rctcrinl pyrogrii,
ovalbumin aiid cortisone. Anat. Rec., 1 0 9 : 351.
~VILASDER, 0. 1938 Stnclicii iiber Heparin. Skand. Arch. Pllysiol., Xl : 1-89,
Suppl. 15.
s.
PLATE 1
EXPLANATION OF FIGURES
Pibro1)lasts f r o m fixed :ind staiiied whole niounts of iiieseiitery of the rat after
iiitr;ipeiitoiieal iiijectioii of distilled wxter. M:iy-Gruiiwilcl stain. X 1900.
1 A few mast cell granules i n n cell. Xotc the vacuole co~itniiiiiiga gr:iiiulc.
2
3-4
Maiiy mast cell granules iii a cell. Note t h a t
granules.
seine
vacuoles coiitaiii several
I ~ r g c xaccuuiulatioiis of mast cell gr;riiulcs iii cells. K o t e sinnll eongloiiic~rntioris of granules.
5
Kote large conylomeratioiis of granules.
G
A cell aliriost coinpletely filled with nietncliromatic material.
i-8
Exaiiiplcs of dissolution of iiictaeliroiiintic material. Vacuoles show varyiiig
degrees of mctachromasi:~ (pink t o red).
PLATE 2
EXPLASAI’ION
O F FIGURES
Mncroplinges :iiid leucocgtes f r o m fixed and stained mliole mounts of mesentery
of t h e rat aftcr intraperitouenl iiljeCtioii of tlistilled water. May-Griinmnld stain.
X 1900.
9
10
A niacropli:igc coiitaiiiiiig iiinst cell grauules. Note t h e vacuole coiit:iiiiing
a granule.
h m:icropliage coiitaiiiiiig
iii:iiig
inast cell granules.
11-12
J1:icropliages coutniiiiiig co~iglonic~r:itio~~s
of metaclironintic granules and
nietaclironiatic vacuoles.
13-14
Kcutropliils coiitainiiig iii:ist ccll granules.
15 A nionoc>yte coiit:iiiiing ni:ist ccll granules.
16 1,yinplloc.ytcs eoiitaiuiiig mast cell granules.
108
109
PLATE 3
EXPLANATION O F FIGURES
Fibroblasts, uiacropliagcs atid leucocytcs i n living m e s c n t e r ~of r a t a f t e r intriiperitoncwl iiijrctiou of d i s t i l l d water. Toluidine blur stain. X 1000.
17-18
Fibroblasts mhicli 1i:rve taken u p f r e e niast cell grnuules.
vacuoles about t h e granules.
19-20
Fi1)rol)l:ists sliowiug ~iict:~rliroiiiatic
inaterial in the process of dissolution.
The vacuoles exliibit varying degrws of inetacliroinasia.
21-22
bfacroplinges eontainiug met:ichromntic granules aud vacuoles.
23
A lcucocpte wliicli has just engulfed a niast cell
top1:rsiiiie vxtensions of the cell.
24
A lcucocpte contaiiiiiig niast cell granules.
110
K o t c tllc
~ K L I I U ~ C .S o t ( ’
tllr pro-
I’IIA(:OCYTOSIS O F MAST CELLS
u.
F,. shl1’PII
A S D Y. S. LEWIS
111
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