Download Blood Coagulation in Arthropods II. Phase Contrast

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Blood
II. Phase
Coagulation
Contrast
Microscopic
Coagulation
i-lAS
BEEN
inn tise
Sti(’k
simO)iVs
tO si
blood,
espe(’ially
of
SI-1O\VN
hemolymph
of
rikinsg
ausalogy
type
ta(’eans
plasma
o)f
pn’ooess
by
pinase
inemolymph
obseu’s-en’s.
ins
6
.
Tisey
Pt’0)0’(55
unsden’go
23
.
pn’eseunt-
papen’
is a sturdy
tlistn’i})utionl
of
tine
amonng
of
existeutce
of
inn
sped-ia!
s-aniouis
mN-iiannisms
tioni
possible
of
species
isen’e
A I )OU
:3t
I
vetor
Tine
Spe(i
nsiet lsod
Scope,
e.g.
nssenss
fronni
oons(! st udied
,
of drtowinng
connttoo’t
it
type
of
kinsti
coagulationi
c-oa-gulat.ioui
cell,
of
all hemo-
mio’roscope
goals
sven’e
lnotnsc
to
cricket
to
si’hethes
tine
of
tine
tine
isinctlsen
results
tine
observe
aunt!
ins(llsine
o-oagulatioun,
ssounld
zoological
stick
otlncr
sunggest
olassifica-
METIIODS
i nsseo-t s we re
of
after
ctopt
innflueunc’es
elensnensts
fou’mer
fact-on-s.
tos Possih)le
ioeusso)1ynnphn
In foreign
with
by
contrast
The
inn tine
AND
6 5 sPeoies
tos soons
pisase
aunol
or- ecologic
variatsee
altcn-ationss.
of inssects,
innsohsct!
i)etweeun
eiolutise
tine
innseots.
species
at
to) a special
ttesct’iI)ed
anal
svei’e
isla!sds
inemolympin
of the
commons
of
of
I)n’o(’ess
s’elationsshnips
anal
preseunce
a categon’y
inemolymph
insitiatioun
of tine
exceptions
meauns
I)1’
61
d)n(ten-s
o’oagulatiouo
inssects
tine
insects,
or coagurlatioun
inssect
u-ole ins tIne
MATERIAL
svhsole
of
n’ecognsized
a-s beiung n-estn’ioted
grousped
togetiner
as uniolen’g()inng
o’oagulatiouo
tIne
o’oagislationn
3#{176}ansy
spe(’ial
.
hsemolympln
iuusect-
of
\vithn tue
in
,
31)
anod
corpuscles
ins cnnspar’t, inn the inu(pinn tineiu- strusctune
which
alter’ationns
root
\it5
i)einsg
‘Vise
.
pt’0o’tss
f1iniiiiiea,
B(’IO.StOflla
t’vtes
o)f tine
type
inn cn’usta(’eanl
ssitii
of these
,
ooagulatious.
olescn’iptiono
tine
microscopy,
tlestrii)ed
associated
au’e n’apidly
follossctl
hr tine deselopmennt
of gn’annunlan’ cloutis
ins tine plasma
sisr’n’ouusdinsg
tinese isemocytes.
Jun tine mouse
cn-icket
annd inn tine stio’k
inssect,
the s’esults
tine
that
(lomesticus)
o’ountrast
of o’oagulations
o’lassificationi2#{176}
Ins tine
r)
(Gryilulus
o-orn’esponsdinng
to Han-dy’s
explosive
tt) play
aus impon’taust
if toot exc!ussive
foisnnd
si-as
Florkins,#{176}
-n’icket
as stuttied
Tait’s
con-punst’les.’7
hemocytes,
l)lOOtl,
tine
of
(1
aunt!
house
inorosiis),
siitin
ivitin
explosive
of isyaliuuc
tiono
(Cai’ausiris
of Insects
\L.I).
(Gn’goin’e
ins tine
on Hemolynnph
Species
(;RGIIE,
elsewheu’e
ot)agulatiOns
inssect
1larclv’s
CH.
Arthropods
Observations
Sixty-one
in
B!!
II
in
or
I he
(‘0)1 lect
O)f t lie
I inc oospect
toir,
to t
eol
t hi rougho
rtousclonn
t lie
nrc.
rul)l)ing,
process
nsmust
lie
unnc!er
t Inc msnicrotovoideCl
coorefully
or
Ze(l
nninsinssi
Fr
nnmo)st c)h)serVtot
or to leg
oos
sI ide
( ( )l(I
ohoeus
I lie
Cluicklv
setol
the
Ans tobst
tononi(’tol
gltoss
d n-oplet
Fronu
1)epoortnnemst
(onsgress,
Oxford,
te(1
inn(Iel)te(l
vtorious
innsects
uscrupt.
The
‘Iarchn
tonsd
expemnses
fixed
nsotrv
on
1951
1)r.
for’
J.
severtol
of
the
24
t
hoe st toge , ttnl(!
ry.
to
25,
observoot
severed
emno! of tons oontensnsa
on to st otnnc!oort!
ionns
nnnicroscopic
st tort e(!
inssmsne!itot
ely
st uolv
of Liege,
woos (lelivereo!
Belgiunmn.
tot t Inc Inst ernsat
lomoot I Anna-
195t).
for
Mr.
(!etermssinntot
reseoorcis
lyimsg
t lie
[Inoiversity
s i ns t Isis
; toccel)te(l
Leclero-q,
oowooy fronnm tine
of to I loins covergitoss
usg.
result
Jul-
27,
to
flowimsg
t lie e(Ige
of l3ioc’lsenniist
of Prelinsmi
Subnsnit
hsensolyunnpls
ott
st ii I spreoto!i
rood
tore
the
) , firnsslv
ovtos
\Ve
Hec’hoerclse
ionns,
Clropped
were
Pubhictot
(Is.
ionns;
to
poortly
Scienstifiolue.
1 173
ions
NLay
.Jeunsiaux
1)r.
21 , 191.
oonnd 1)r.
henry
defrayed
P.
Tagnnons
by
the
Houyez
for
Founds
for
retociinsg
supplying
thso mssttn-
Nootiounool
tie
Itt.
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1174
BLOOD
In
some
species,
leg or svings.
U5) and
In
forced
there
these
to the
thorax
slide.
room
the
for
and
used
long
periods
Suitable
bees,
individuals
and
samples
A Wild
by Mr.
cricket
of
the
Abel
(Gryllotalpa
some
the
insects,
phase
fixed
osmic
acid
was
August
PLATE
arid!
tions,
1.-Most
pre5)arootions
immediately
hemolymph
was
FIG.
1 -Mole
hemmer’s
Ringer.
blood
collection.
reto ring
severoo!
conc!it
ions
tirnnes.
various
objective
with
species
of butterflies,
40/0.66,
was
nnicrocanneroo
(35 mm.
picture
studiec!
beetle
water
Ph
the
used
Leitz
for
the
9 x 12
cm.
subsecuemotly.
motion
the
in
the
(Dysticus
filnms)*
present
performed!
paper:
the
mole
tine
nnneal
worm
niiai-yinalis),
categories
various
compared
stained
mm.
on
of inenmocytes
with
with
those
Gienisa
“Blood
International
from
or
Mooy
obtooinsecl
snmears
of
by nmeans
inenmolymph
Griinnso-alc!-Gieunmsa.
cootgulatiomn
Physiological
(3) and
plate
ins inssects”
Comogress,
(Gr#{233}goire
held
anol
inn Copenshagen,
3, fig.
nmediate
plasnmic
Fic.
the
of
spreading
Fn;.
diluted
location
a tremendous
nnicrogroophs
were
taken
injected
Hvaline
by
multiplication
prepooratiomns
proceeds
from
around
of tIne
with
the
with
squeezing
(no
in the
greater
hyaline
process
ac-
(see
three
other
Coalescenoce
dennsity
hemocyte
from
the
made
and
bright
seen
in
of
ins tine
inn-
center
area.
of clrawno
debris.
of tine
currenits
for
instracyto-
circular
of systems
by
See
with
denmse
refractive
the
covergiass.
straight
Cootgulatimsg
picture
aroumnd
developeol
compoorisonn
by
plotte
a
tIne
5, fig. 24,
hemocvtes.
supension
hemocyte
of
streamlines
hyaline
a
On
of cootgultttion.
expanded!
in
removool.
Area
process
an
of them
by
islanoo!s.
any
hemolymph
Meshsvork
under
pie-
ploosnmoo cotogulatiomn.
coagulation
after
away
produced
Meisen.
The
inn undiluted
of the
(one
of the
with
antenimsa.
thano
hemocytes
carried
fronn
circular
of the
various
of
diluted
removal
initiation
hemocytes.
shows
hemolymph
controlled
bet-weenn
prepara-
is smaller
minutes
islands
systems
rensoval.
out
Inn a few
(1) a small round! hemsmocyte
(lowerumnost-) ; (2)
vacuoles
and short
lappet-like
5)seudopodia;
hemocyte,
surroumocleci
by to highly
ref rooctive
hemolymph.
and
5.-Gryllotaipa
hemolymph
center
of extension
to
from
hemocyte)
pseudopodial
the
hyaline
1)icture
Diluted
anochored
of the
in
islands
Extension
Stages
originating
the
Henmolynnph
after
to the
coagulation
three
i)lack
hyaline
seen
No
cells.
gramoules.
to spreac!
or leg.
All
Orthop.)
15 minutes
hemocvtes
of these
12).
The
materials
loorge
are
3.-Gryllotalpa.
strings
hvalinse
Twenty-two
4.-Gryllotalpa.
gramouloor
#{231}jryllotalpa,
taken
around
vicinity
Fro;.
Ringer.
shown
in the picture:
with int-racytop!asmic
of nmulberry-like
granular
2.-Gryllotalpa.
allowed
severedanntemsnto
Meisemnheimer’s
(Gryllotalpa
(H)
isltonds
of loemolymphi
from
(Wild M/10)
with a microcamera
Leitz
(Mtokanm: 9 x 12 cm.)
The scale on the micrographs
represents
10 mnnicrouns.
the size
tvl)e
coagulation
with
of the
of hennocytes
hemocyt-e,
films
shedding
Microphotograph
another
aureole.
Fi;.
are thin
diluted
cricket
2, 3 and
categories
to grtonoular
gone
from
the
imnsects
on
microscopy
enlarged.
t-ure shows the relation
Two hvalinne
hemocvtes
count
of the dilution,
plasmic
the
coverg!ass
by pinasc contrast
amnd subsecuently
t rue
after
adequoote
agtoins
enlarged
by
were
ins 35
at
lifted
1950.
slide
figs.
of
and
film
cases
gigas.
vapours
projected
svith
takers
svere
the
niicroscope
cinematogrooph
Florkimn)
arnd
following
pictures
controost
with
A
the
10,
process
the
of anntemnna,
of time wing
hours
under
be obtained
svere
gryllotalpa),
and Blaps
-inolitor)
In
M
of 400
for
two
ivity
ol)served
not
microscope,
coagulation
severtonsce
joinnt
flies.
photomicrographs
Charles
(Tenebrio
*
and
at- a magnifications
Recording
of
contrast
The
after
the
during
-capt
process
could
ants
frons
at least
in semi
coagulation
svasps,
5)hase
(Makam)
kept
of hemolymph
bumblebees,
observationns.
observed
were
their
issuing
to the middle
of to coverglooss
which was genntly
generally
gave artifacts
(see l)elow).
from each specinnens,
placed
irs mnnoist clnannl)ers
at
made
process
II.
touched
method
also
ARTHROPODS.
of hennolynmpln
of blood
was
This
were
IN
rio shedding
a droplet
placed
down
onto the
Several
5)reparations
Previously
often
was
cases,
back
temperature
COAGULATION
an
of
engulfed
baker’s
sisrunmk
stnperpositiomn).
inemolymph,
Inn this
which
was
yeast
yeast
12
globule.
insect
iso
more
yeoost
Inours
before
Inst-racytohood
coagultoble.
under-
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CII.
GSnECOIIIE
:;
a:.ii
-
“I
l’i.crF:
1
1175
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1 176
BLOOD
COAGULATION
IN
ARTHROPODS.
II.
RESULTS
The result-s
recor(led
inn table
1 were collected
from more
preparations
of fresh
hemolymph.
The alterations
observed
in thin films of innsect hemolymph
described
unnder
specially
were
1
types
of
pictures
of
Plasma
Coagulation
Depending
on A iterations
in
Process
to the Whole
Plasma.
The
process
husustlred
si-ill be tenstatis’ely
inn svhich
these
alteuations
of the
i)e selected
with
that
Islands
Hernocytes.
hemolymph
to illustrate
described
or
Graiu u/ar
Extension
inn the
o,f
mole
the
Cloud.s’)
Coagulation
cri(-ket
(Gryliotaipa
the first type
in the house
of pictures.
It presetnts
a
cricket
(Gryiluius
(tomes-
from
anst-cnnnsa
t5),
the
betsveenn
drop
slide
of hemolymph
annd tsio kiunds
large
spherical
pale
four
granular
with
cytoplasm
active
are
rrh
first- stage
of the
these
hyaliune
hemocytes.
with
the spreadinng
film
to the glass surface
arid
of a rapid
expannsioin
annd
motion
: plate
small,
a fess
(plate
to
:olteratiorss
Three
in
I fig.
sharply
black
a categon-y
nnucleus
grainuslar
2, 3 : plate
witin
svith a relatisely
or proleukocytes,
I ), atsd
out!irned
,
scattered
I , figs.
spreading
along
3, fig.
of
arid
particles
12).
coagulatioun
process
takes
place
siithins
a fess se(’oustls ins
While
the other categories
of inemocytes
are still mosiung
of hemolymph,
the hyalinse
hemocytes
stop,
seem to fix
undergo
a susccessioun
of inntennse modifications,
(‘onnsistinng
boilinng motions
of the cytoplasmisubstaunce,
the forma-
6, 7, 8.-Gryllotalpa.
relatioum
is still
mosinng
categories
of cells
rounnnd hemocytes
hemocytes
a relatively
2
PLATE
Fro,s.
of hemocyt-es,
three
(small
containniung
i)roivniiatn
a severed
categories
There
cytoplasm
of large
hemocytes
hyalinne
exhibitinng
shed
annd (‘overglass,
the fluid,
cars be observed.
large
rnut’leus
asnd a dark
a
specimeuns
(Coagulation
Hyaline
of (-oagulation
gryllotaipa)
may
great
similarity
ticus,’tm’
Whets
in
fifteen
demounstrative.
Initiation
.
four
thans
invaline
stages
of
developnsennt
hemocytes,
of
recorded
by
tlme
motions
process
of
coagulootions
picture
by
Ch.
ins
Abel.
in-
frons
frannes
of 35 mm.
filni.
Figure
6: two expanmdinng
hytohinse hernoocytes
(11) oonsd
clark granular
hemocytes.
Plasma
riot yet altered.
Figure
7: later
stage.
Beanm-hike
)rojections
of cytoplasmic
blisters
by the two expandinog
hemocytes.
Developnnenst
of a
loorgenmeusts
three
large
grannuloor
coagulunm
cloud
tot the
secon(l
hvoolinne
Extenssions
(!O osot
FrG.
poirt
witholrairtol.
tions
At
categories,
inuclei
of the
(lividual
vooriation
. 10
-Mantis
of shrinkage
Fio
stages
Fn;.
hvahine
an(l
Tivo
lower
tue
running
the
side,
of one
The
the
svhole
hvaline
three
plasnsa
of them
form
preparations.
reliqiosa,
Orthopt.)
hemocytes
altered
tinn-ee
doork
by
a thins
other
cells.
Connpare
inn the thickness
of the coagulum.
religiosa
. Coagulat
ion islands
around
of their
sharply
out-hinsed
nucleus.
hyali
ne
hemocvtes
granular
in
anod
of those
Sixteens
nsinsutes
coaguluni
the
direction
of the
into
of
rows
plasma
after
of
hennolynnnph
prsrticles,
currents.
later
corogulunsi.
12 nminsutes
ptortobohic
ooftoor
cooogula-
helonoginig
differensce
in
witls
lnemocvtes
witlndroowal.
forming
the
stooge.
inennmocytes
inn the
vacuoles)
the
of o!eunser
around
groonnultor
ttokerm
(blisters,
them.
torea
a still
embedded
Microgra)h
hemocytes
around
the
8,
Fiyuie
I)assivelv
surrounded
Note
to be altered
fog.
The
tornd are
.
is stoortiing
of a thin
process,
reactions
(tot right)
plasnna
Note
relijiosa.
of
left-) under
to
without
ll.-t!anlis
rorrangemenst
threads,
the
vicinsitv
hemocyte.
the coagulations
marotis
(.lianlis
inn
Inemolvnnpin
other
(at-
cotogulations
9.-Pravinsg
islannd.
in the
of this
hemocvte
of the
take
startirsg
l)eripinery
bunsolles
size
figure
,
to
of
time
10 ins-
wit h vttri ous
Secondary
of
groonsular
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1 177
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1178
BLOOD
tionn annd retractionn
plasmic
of vacuoles
blisters
cessions
ansd
inn usearly
i)lebs.
all
modificatiorns
of the
granular
particles
in amounst
and
ins dennsity.
its close
relations
pictures,
are
clout!
to the
oftenn
develops
3;
plate
When
to
tine
directly
a thins
surrounding
Three
stages
fog
plasma
2, figures
the
process
thread-like
2, fig.
while
altered
hyalinne
begins,
whets
formationns,
resernblinng
pseudopodial
1 1 ). Subsequeunt-
organizations
of these
While
nearly
all
coagulation,
the
hyahinne
the
other
iii
figs.
unndergo
13 anid
cytoplasmic
extrusions
14).
They
area,
of short
processes
inn motion
of tine gransular
an-ca of iniginen- tlennsity
(plate
film
I figs.
2 aunt!
,
of inemolymph
structurres
(plate
formationns
in the
are
insto
the process.
or entrapped
by
1, plate
is still
assume
innto
1 fig.
4, plate
a mesinw-ork
takes
,
consistinng
the
,
of the
plasma
exclusive
part
coagulation
three
categories
of hemocytes
are shown
inn the field are surrounded
by granular
is no plasma
reactions
arounsd
the other
.
I 2.-Gryllotalpa
.
of extenssiono
Fro;.
side,
hornsec!
witlst!rawal.
at ranndom,
tiney
2, figs. 6-9 ; plate
3,
of va(’uohizations
ins the
,
ioemolynmplo
the
hyalinne
cricket.
developed
wit hdrtowosl
coooguhot
(Acrididae,
cloud
by the
mole
hemocytes
Inn this
picture,
3
of
grasshopper
granultir
played
: only the two hyalinne
hemot’ytes
present
clouds
or coagulationn
islansds
while there
categories
of hemot’ytes.
after
boundaries
Grannular
embeclo!ed
two
minsutes
(unnslsrorp
13-Short-
henmolvummph
Three
innitiatiorn
aunt! granninlar
(potocytosis)
aund of i)eam-hke
oft-cnn connnnected
with similar
ins the
1LATE
nosotiono
chiefly
of blisters
pseudopodia
inn the
rounid
hemocytes.
1 illustrates
innitiation
inns-oh-ed
(small,
Scattered
or agglirtirsated
in the coagulum
(plate
changes
other
actively
of hemocytes
occasionnal
formation
spiky
or !appet-like
developed
Figure
hemocytes
categories
cells) do snot take part
are passively
embedded
lower
coaguluirn
recorded
hemocytes
the
of
i unt-reases
of tine plasma
a circular-
off or
made
occasionally.
of the
FrG
shadiusg
aunt! progressiely
hemocytes,
in the
a c!oisd
6, 7 annd 8. Extennsions
place
hemolymph.
the coagunlatinng
materials,
carried
inn streamlinnes
by (‘uu’rennts,
shapes
: pearshaped
patches,
liming
up of graununlan’ substauscc
fortuitous
in
of
descrii)ed
(‘ricket.
insto
of developmennt
of cytoor inn sure-
a stsddenn
by
developiung
ins hyalinne
takes
arounnd
3, fig. 12),
the coagulation
preparatiorn
the surrounsdiung
fluid,
certainty
ins the mole
irs the
in plate
huslgiusg
simisltanseously
or followed
while
preparation
beam-like
place
inn a givenn
alteratiorns
illustrated
whole
spreading,
place
appears
take
into
with
accompannied
outlines,
II.
of a violernt
hemocytes
are
cell
ARTHROI’ODS.
changes
substansce
riot observed
-
These
IN
aund finnally
Similar
hyalinne
Discharge
of cytoplasmic
house
cricket,’4
‘ was
vanishing
COAGULATION
Orthopt.)
oorounsd
hennocytes
with
.
Cotogulost
ions
isloonsd
inog toreto).
Fifteenn
to Isytolinse
acciolenstoil
nninnutes
hennocyte.
relation
to
At
the
tofter
the
left
coooguiuros.
Fm. 14.-Earwig
(Forficula
auricularia,
I)er,napt.)
A hytohinne
lsersmocyte
surrounc!ed
ly to
thick
areot of coagultotinsg
nmaterials.
Generot!
extenmsions
of tine 5)OCCSS
fronnm
this
isltonsc!.
At
the extrenne
lower side, a ptossive
highly
vacuolated
hsennocvte
fronsn annother
cootegory.
Froos. 15, 16.-Wooter
scorpionm
(.Vepacinerea,
Hemipt.)
Fifteens
:onn(! 25 noninnutes
tofter
l)loodl
withnolrotwool.
Islansds
of coosgulationi
developeo!
tirounnol
hvoilinse
heumnocvtes.
Inn figure
15. extensions
Fn;
o)f the
.
henmolvnmpis
cyte.
No
process
17-Leaf
to
beetle
ivitiiclra’tvtol.
extensions
of
tine
left.
((‘hrqsoonela
Timinn
time process.
poole
Thick
coooguluns
staphylea
coagulation
ins figure
, (‘oleopt.
isloirmds
16.
C/i rysoinel.)
clevelopeol
Ninnety
tirouusol
nminsutes
to loytolinne
oofto-r
hsenno-
From www.bloodjournal.org by guest on June 18, 2017. For personal use only.
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From www.bloodjournal.org by guest on June 18, 2017. For personal use only.
1180
BLOOD
rfhe
type
of
velopmeint
described
of connspicuous
arid follosied
hemolymph
3,
reactions
COAGULATION
fig.
1 3)
inn the
islands
by a genseral
of all specimenss
ARTHROPODS.
II.
mole
cricket,
cinaracterizetl
of coagulations
coagulation
of Orthoptera
of the earsvig
(Forficula
scorpions
(Nepa
cinerea,
,
IN
arounid
in
all tine inyahinne
the preparations,
examirned
(plate
auricularia,
hemocytes,
(plate
15,
I 4);
tss-o
water
species
i)eetle
of Coleoptera
Tenebrionidae
(Tenebrio
molitor,
Blaps
gigasni),
of the
(Meloe
proscarabaeus,
Coleoptera)
arid
of t-wo species
of Hymenoptera
(Dolerus
2. Absence
the Hyaiine
pratensis,
of Islands
Hemocytes.
sp.)
Arge
(plate
3, figs.
3, fig.
16), of
of the
Tenthredinidae
plate
de-
si-as fouusd
inn tue
2, figs. 9-I I ; plate
Derinaptera-)
Hemiptera,
the
by
4, figs.
22,
oil
23).
of Coagulation.
Production
of Pseudopodiai
Occasional
Outgrowth
of Diffuse
Veils
Ta1ing
Meshworks
by
Origin
in
the
Plasma
Samples
of hemolymph
Cai-abidae)
may
from
be selected
the
Coagulation
islands
were
Hosvever,
hyalinne
hemocytes,
inn the mole cricket,
undergo
pseudopodia
exhibit
beetle
the
Cychrus
second
type
not
observed
in the films
similar
in their
cytologic
alteratiorns
of a different
of hemolymph
is still spreading
hemocytes
send
out long straight
them
black
cytoplasmic
produces
ins the growinng
grournd
to illustrate
out
granules.
pseudopodia
a high
between
threadlike
rostratus
of blood
from
this iunsect.
features
to those
found
nnatusre. Whets
the droplet
slide ansd
pseudopodia
coverglass,
carrying
Innt-enise brosvrsiann
motions
continuous
trepidatiouns.
degree
of thigmotropism
cars
the
of
size
reach
the
considerable
hemocyte
lengths,
from
towards
sometimes
which
they
solid
more
proceed.
strings,
cytes.
formed
By
exclusively
drawing
by the
out
the
pseudopodial
eytoplasmie
hvaline
arietis
FIGs.
19,
20,
Longic.).
Frinoges
of
21.-Larva
of
potato
Colorado
beetle
with
svit-h
of the
these
straigint
hyalinse
hemo-
pseusdopodia,
coagulating
plasnmma
(Leplinotarsa
Five to 7 minutes
after
blood
withdrawal.
severrol hyaline hemocytes.
Fig-are 15 shows
seen coating
found inn the larval
ma coagulation.
spinn(lle
shooped
agulunn
irsto
(Irawal.
stage.
Figure
(lark
bundles
Fio . 22.-A
rye
sp.
Coalescence
honnogersous
FIG.
times
together
the
around
fotnr
inennoc-tes.
Chrysoinel.).
lation
(Coleopt.
a husndred
joins
category.
meshworks,
innto
bodies,
4
PLATE
18.-Clylus
FIG.
than
expannsionns
substance
foreign
bits of chitineous
slip arournd
them
pseudopodial
for-
They
similar
structures
sent out by hemocytes
of the same
As shosvns ins plate
5, figure 24, there
arise polygounal
the hyaliune
along
with
of these
grannules
These
cytoplasmic
other hemocytes
and physical
interfaces
(dust particles,
crystals,
debris,
air bubbles).
They
adhere
to these
bodies
or bubbles,
and proceed
on their
exparssion
to farther
annchorages.
These
mationis
(Coleoptera
of pictures.
to
henmocyte
process
over
inn the
is passively
helc!.
of
carried
threads
(Hymenopt.
away
plasma
hemocytes.
of tloe
arournd
.
Twenty
aroun!
Dark
of
(‘oleop!.
corogulatinig
plasnmto
tore
the two kinds of hyotlinse Isemocytes
adult stage,
does riot insc!uce anny 1)1005-
Figure
from
Tenthredinidae)
coagulated
seven
imenmocytes
coagulooted
2! shows
the
materials
the
inytolinme
nninsutes
inn which
!roowinng
of
time
a
(‘0-
hemocytes.
oofter
inenmmolynmph
exparoc!e!
aund
scoottereci,
obviously
wit Is-
cvtol\-zing
(hazy
without
re-
process.
pratensis
surrounded
a loorge
absent-
hemocyte
hyaline
23.-Dolerus
cell,
extensions
coagulation
hyaline
large
a veil-like
of
niuclei)
tine
The
20 shows
decenilineata,
Frinnges
area.
(Hymenopt.
by
a dense
Tenthredinidac).
coatinsg
of
Inn tine
cotogulottec!
center
5)lasnsoo.
of
tue
Extennsiono
picture,
of
a
tine
From www.bloodjournal.org by guest on June 18, 2017. For personal use only.
CII.
I 151
GutEGOISIE
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From www.bloodjournal.org by guest on June 18, 2017. For personal use only.
I 182
BLOOD
cell
body
is greatly
OtiC
side
of
tlecreased
the
striungs.
particles,
normally
aund represenntinng
of cobwebs
Meanwhile,
these
24,
ansci ofteun
frequennt
probably
pseudopodial
systems.
26).
inn the
to take
The other
of
categories
pseu(lopodial
these
passive
anchorages
or they
cytes,
The type
was
rostratus
part-i(’les
(plate
become
visible
fluid
of the
of reactions
observed
beetles
traunsformed
(Dytiscus
veil-shaped
of this
to the
does
not
inn the buildiung
bodies,
they
sent oust- iy the
do unot (lisiust-egrate.
Odonata),
ins several
Coleoptera,
Cybister
structure,
coagulusm
strtrctunres
hyalinne
inn the hemolymph
of the grouund
specimenns
of nnymphs
o)f (lragoun
sp. arid
inst-o
meshsvorks.
cytoplasmic
filaments
ins the veils. They
and Libellukz
quadrimacukiki,
( caterpillars),
inn sevesal
species
of
by
particles
are scattered
(plate
5,
or tra(’tions
exerted
onn the coser-
of such
grandis
water
is progressively
pseudopodial
described
inn most
o)ni
grausinlar
(‘ircurmscril)ecl
of hemocytes
do riot actively
take pantsystems.
Like
the other
innert foreigun
for the
embedded
are
area
corntractile
(‘ounsisteuscy
ann appearansce
similar
Development
otnt
of
spe(’ially
inn Coleoplera,
strings
the appean’annce
5, fig. 25).
ins the flusid
plasma
btnlginsg
strinsgs
of mausy iunsects,
n-emnsarnts,
gi\e these
elasti(’
arid
up, assumes
outside
nnucleus
pseurtlopodial
i)lOOdl
The
Orthoptera.
place
IS.
to tine
these
glassy
veil,
on svhich
granular
I\Iechannical
agenncies,
as pressure
discloses
the jelly-like,
svhich,
by foldinng or rolling
observed
ARTHnR)poDs.
ret!uced
on
inn the
inistolytic
glass,
seem
IN
Adsorptions
spriunklet!
with dust
sloss alterations
a transparentfigs.
COAGUL.-TION
f-hem
arid
of tine (br
Roeselii)
hemo-
beetle
flies
species
among
tsp
au’e
Cychrus
(Aeshna
of Lepidoptera
most
specimens
beetle
of
(Geotrupes
stercorarius).
3.
1 and
Types
Frequennt
2
Combined
or inncidennta!
hemocytes,
combinned
of cells,
category
developmennt
with
arid
of islannds
product-ions
outgrosvth
of
of (-oagulat-ioul
pseudopodial
of plasma
veils,
around
mcshsso)rks
has
beens
by
ohservetl
hyalinse
the
of insects
listed
inn the precediisg
group,
for instance
in usymphns of tiragoin
various
Coleoptera
(Cybister:
plate
6, figs. 27 and 28; (‘iytus
arietis:
plate
18; Leaf beetles:
plate
3, fig. 17; plate
4, figs. 19-21)
ansd inn thnu’ce species
menoptera
Tenthredinidac
(plate
4, figs. 22, 23).
these
In
hut
various
generally
is
species,
munch
the
smaller
size
of the
than
islannds
of co)agtslatioun
Orthoptera.
As sisossns
inn the
Fnas.
24,
25,
after
26.-Grouund
beet
hennolvmph
le
(Cychrus
withclroowoil.
flies,15
4, fig.
of
Hg-
varies
greatly,
ins tine
pictunn’es,
5
PrATE
minutes
same
inn specimeuns
rostratiis.
Polvgonsool
( olo’:p(
meshwork
.
( ‘arab.
fornsned
).
Fiure
9()
24 :
exclusively
lnv
hoe
I
of severtol
hvooline
henmocytes
(H ) drtowns out inst (i st rttighit
t Isretoci-like
5)seudoexpansions
uunite(1 with sinmiloor fornnootionss
of o)ther lsytolinoe honssocytes.
Tue Pseu(lpodial
strings
are anscinore(l
t o foreign
bodies,
ot her (loork isensiocyt es tonsil bubbles
( lonng rtonsge
thigmotropisnn)
. Cell
body of t he hyahine
henmocytes
oftens
reduco’d
t o t hoe bulging
nsucletis.
cytoplasm
5)oolial
Other
6
exhibit
nminutes
sorption
the
of
highly
henmolvmph
poolial
veloping
protru(hmsg
tofter
blisters.
coarse
particles,
aolhesive
in tine
ars(l
lower
H
: rsuclei
Troinnsparenst
sprinnkleol
poort
groonsules
A
frequently
strirsg.
withclrtowtol.
meshwork
Black
withdroowool.
hennmolvnmpis
of tine
floootinng
of
gloossy
with
sctottereol
l)etooleol
veil
oo(lsorl)ed!
nmeshwork
free
hyoohinse
tolonig
tot
inn tine
iienssolynsnl)is
hennocytes.
(!evelope(l
bloock
inn figure
I he
strinng
Piqure
mo l)ltosnsnto
groinsules.
24.
st rinngs.
higher
b’iynre
25:
nonoognsifiototions.
of
26:
(‘oleoptera,
015
9() nninsutes
inssiole
A sinnniltor
Aol-
of I lie
veil
after
l)seu(1-
is seeno
(Ic-
From www.bloodjournal.org by guest on June 18, 2017. For personal use only.
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From www.bloodjournal.org by guest on June 18, 2017. For personal use only.
1184
BLOOD
COAGULATION
the plasma
reactions
often
by coagulating
materials,
4
No
.
Visible
Most
Plasma
samples
p1cc-us)
arid
appears
without
of the
No
hemolymph
II.
coatinng
of the hyaliune
extennsionn
to tine wisole
Reaction
of the
caterpillars
third
type of pictures
No visible
chasige
ARTHROPODS.
as a narrow
subsequent
Coagulation.
of several
IN
of the Hyaline
great
silver
of butterflies
svater
may
Hemocytes
i)eetle
(Hydrophilus
be selected
(plate
6, figs. 29, 30 annd plate
could
be observed
inn the plasma
to illustrate
7, fig. 32).
of these
inssects
a long stay
of the preparationns
in a moist
chamber.
rFine hyahinne
gennerahly
of large size, do not unndergo
amy alteratiorn
or they seund out
a few
short
actively
pseudopodial
movinng
several
hours.
Similar
absensce
ptera),
the
arid
With
the
cytoplasm,
these
hemocytes
of reactions
svater
Corixa
expansions.
irs their
was
boatman
geoffroyi,
found
arsd
Hemiptera),
exceptions
inn three
the
lesser
arid
of the
still
species
water
inn various
hemocytes
plasma.
es-ens after
inemocytes,
inscidenntally
black
appear
grannules
insert
of Pentatomidae
boatman
species
the
after
(Hemi-
glauca
(Notoneta
of Coieoptera.
DISCUSSION
mains
rFhe
results
of the
I Inn the hemolymph
cans easily
be distinguished
of all the
under
.
may
be sunmmarizetl
paler
tb
inucleus
snot exhibit
3. Inn species
appear
to
be
ansd
insects
hyaline
the
hemocytes
dark
ins which
cytoplasm
the
blood
highly
are of tsvo kiinds,
sequennt
plasma
The first kinid,
of the
does
labile
to
structure
plasma
contact
with
ivhich
coagulatiorn.
play
blood
each of them
being
reactiors.
described
previously
arouno!
into
threadl-!ike
other
Fio0s.
hyaline
29,
nmechanical
Inn
t hese
inn sshich
conntrast
categories
clot-,
30,
31.-Greathemolvniph
hemocvtes
product
of large
these
to
structures
silver
water
hemocytes
inyalinse
inemocytes
beetle
reaction.
(H)
round other
of
st raight
Figure
cells
t bread-like
appearance
-
or
ion
‘#{176}
is
rapid
of the
stnh-
characterized!
by
corogulateo!
foreign
bodies.
in
figure
24,
(Hydrophilus
inert
altered.
parallel
No
plasnma
5)seu(!opodlitol
pressure
arid squeezing
exerted
on the coverglass
in a otherwise
Inn spite
of a similoor
general alteration
in all hyaline
hemocytes
ansd development
of substantial
meshworks,
nso plasma
reactions
prepoorootionss
(‘oleopt.)
from
29:
Fiqure
10
without
withdrawal.
expannsions
out
strings.
henmocyte
conigulation.
-
is drawns
Pseud!oPo(!iool
hyalinse
isenmolymph
exception
nmateritil
Most-
piceus.
after
developec!
with
spherical
15 nmminnutes oofter
tond
islooniols
of the
30: 2 hours
variously
unntlergo
if unot exclusive
part
inn
material,
these alterationns
Tlmirtv-two
coagulat
A part
A large
of elements.
hyalinse
They
Orthoptera’tm
Coleopt.),
bubbles
scattered
svitis the
6
numerous
shown
withdrawal.
surfaces.
to a differennt
its tsvo
size.
the
plasma
ion
,
glass
related
Roeselii,
specimens
attached
resembled
after
(Cybister
henmocytes
structures,
surrounding
hvaline
beetle
.
sPecimens
nninut-es
any
28-Water
wit Isclrawal
ally
other
riot
an importannt
Inn the present
PLATE
27,
characterized
anny importannt
alterations
ins their
st-ructure.
inn which
hemolymph
coagulation
occurs,
these
alteratiorns
inn their
the insitiationi
of the
Fnns.
are
by a relatively
of blood
aund sharply
outliuned
nucleus,
a pale hyalinne
cytoplasm
grannules
exhibit
active
brownian
motions.
These
featusres
Ins those
as follosss:
innsects studied,
a categony
of hyahinne hemocytes
the phase
contrast
microscope
fn-om tine other
kinds
2.
These
iunvestigatioun
small
1)lack
larger
cells.
presennt-
Inert
Fiyiorc
by
31:
st ronng
insert hyaline
hemocyte.
preseust
ins tine preparat
ions,
occurreol.
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CS-I.
GuIEG0)IuIE
*&L:;,
i
-
PL;uTF:
6
I 185
From www.bloodjournal.org by guest on June 18, 2017. For personal use only.
1 186
BLOOD
a successions
hemocytes,
metnts,
COAGULATION
IN
ARTHROPODS.
II.
of various
local changes
inn the cytoplasm
of a snumber
of inyalinne
brought
about
by disturbansces
of the superficial
tennsionn ins these dc-
which
are
accompannied
or
rapidly
by tine developmennt
follosved
of a
thins fog, ins the plasma
surrourndinng
them
rapidly
thickenninng
innto an islannt! of
coaguslatirng
materials.
In various
species,
extennsiorn
of the process
briungs aboint
genneral
clottinsg.
The seeonnd kind of alteration
inn the hyalinne hemocytes
counsists
of tine extrusions
of thread-like
pseudopodial
expanssionss
simultaneously
by several
cells ant! consstitution
are
pseudopodial
of
followed
systems
by
of
Both
the
thin,
meshsvorks
formatiors
trarnsparennt,
kinids
of various
inn the
plasma
elastic,
of alterations
complexity.
slosvly
of the
These
circumscribetl
developinng
hyalinse
pseudopodial
veils.
hemocytes
assd
their
corresponndinng
plasma
-1 .
riot
they
reactions
earn occur
together.
Irs contrast
to the hyalinse
hemocytes,
take part inn the process
of coagulation.
alterations
by the
are passively
embedded
clusters,
strannds
or
inn
pseudopodial
34).
5. Irs the
ess,
innitiated
present
onnly
already
material,
urnder
the
place
are
J)ifferences
sponsii)le
in degree
the
the
other
in
with
transition
ol)served
part
of sensitiveness
played
by
categories
forms).
in a fesv
tinme had elapsed
of
specimens
since
to contact
the
However,
hyahinse
blood
cells
small
amounots
(small
Phraginatobia
Ftc;.
hvaline
from
hemocvtes
various
a thom
folded
otclsorbed.
veil
of similar
vitreous,
constrasts
proc-
of plasma
detectable
with
their
Agglomertotion
structure.
of
(No
“cell
material
tore attached
currents
in plasma.
in
Plasma
in lines.
On
intact
trannsformed
the
row
from
toppearannce
left
innto
side
reob-
together
exceptionally
elements,
oofter
a lonig
torso! Gunnnn,2#{176}it is possil)le
Forty-five
inensiolymph
unsooltered
agglutinooted
Fifty
figure
ins cooterpillars.
l)loosnsna
nninnutes
substance,
various
of
after
to
.
Other
tot randons.
adhesive
which
nninnutes
(conmmpare
scosttered
these
tofter
henmo-
circunnscribinsg
hsennocvtes
forrnationss
tore
ins Lepidoplera
inn Coleoptera.
(Lepidopt).
hemocytes
Etocln
were
occurred
(Lepidopt).
highly
on
fuliginosa
Arch-like
and
consistenscy
expansiorns
the
latter
Tooit
apparently
Schiff
and
aspect-
coagulation.”)
(pseudopodial
by
(Lepidopi).
intact
cornsisterncy,
refractive
nnumerous
of the
Prol)abb’
insoictivity
lnemocvtes,
nnateriotls
coagulation
in the
auricoina
granular
granular
of nouncoagulating
are
Phraqmatobia
and
are
tonic! the
7
shown
of glassy
elastic
highly
34.-Caterpillar.
F1G.
strinngs
round
no plasma
categories
Parallel
The
are
4cronycta
33-Caterpillar.
withdrawal.
embedoled
7,
isemocyt-es
degrees
reaction,
surftoces
process
of coagulatinsg
fuliginosa
In this specimen
is typical
for preparations
hemocytes
lvnmph
by
of hyalinne
foreign
in the
around
some
As suggested
withdrawal.
withdrawal.
34) . The picture
Nine
vesicular
cvtes
(plate
to be a conitiusuous
category
with
hemocytes
Orthoptera
of
the blood
32.-Caterpillar.
hensolymph
hesive
appears
or gatisered
adhesive
hemocytes
follosved
by various
clottinng
to a limited
PLATE
FnG.
32.
hyahinne
veils,
highly
microscope.
for
served
oftenn
inn the
the
of the Coagulation
Mechanism
doork
the
ins a sinngle
alterations
macroscopic
,
by
coagulationn
taking
These
a genneral
15)
.
from
is-ovens
blood
by alterations
(coagulocytes,r4
coagulation,
categories
of blood
cells tb
or agglutisnat-ed
at- random,
in the islansds of coagulationn,
tissue-like
structures
along
expannsionis
fig.
the other
Scattered
of the
a gloossy
Annotlser
inn 5)ooroollel
conosists
hyroline
of
torn axis
hemocytes).
brought
oobout
picture,
various
jelly-like
rows
specinmneno
fornsminng
nnoo(le
of
tos ins figure
to tissue-like
trtonnsptorenst
otci-
on winich other
memol)y benndinmg of tine structure
substoonsce.
inntact
inenmocytes
are
seen
From www.bloodjournal.org by guest on June 18, 2017. For personal use only.
(II.
CSIX)IItE
1187
I_.____J
---..
#{149}
4
A
_._)_-_‘
-,
.
‘,‘4’
-
,----
.:
#{176}-
Pr-s’n’F:
‘_li()
7
‘:
From www.bloodjournal.org by guest on June 18, 2017. For personal use only.
1188
BLOOD
the
thttt
oolinne
the
other
cootegories
henmocvtes,
nmigiot
inductions
loiter,
of
hoove
delay
trootionss
or
occur.
ARTHROPODS.
to nmore
cooogulation
Prehinninnary
grtoolual
cyto)lvsis
process
slmoore
observootionos,
tonic! cocainn
oolterationss
inn the
The
seenns
process
II.
to
which
hydrochloride
hyalinme
thnoons the
oo snmiooller
to
will
reported
ins todequoote
Isenmocytes.
be bounsd
le
concern-
Co:ogulootions
witlo
Up
cioooroges
Isv-
extenot
c)f
tookinsg
tine
5)100cc
cells.
inn these
The two
patterns
l)robotl)b
tore
of the
oxtiltite
the
to
IN
unsdergoinng
stages
potassium
prevenot
then
hemocytes,
coagulation.
tlmoot
fails
isenmolvnnph
of
ins later
5)loosnna
shown
COAGULATION
reaction
of ploosma
sul)sidittrv
observed
nnorphologic
ins insects,
variations
of
the
cotogultotiono
sanme
l’es
islooncls
Sloth
tonic! veils,
nnigiot
be
pro-
oluceol,
tos previousb
suggested
for the development
of islanmds of cotogulootion
oorounnc!
cxplosive
cells in crustoocea’727#{176}#{176} by the release
of sonme coagulations-insducinng
substoormce
innto
the surrounding
plasmos,
either
after rupture
of the hyaline
henmocytes
(Giylloolus,’#{176}.
‘5
or by
exuclootiorm
from the 1)erii)hery
of the expanding
hemocyte,
Gryllotalpa.
It is oorounnd
the
locool beam-like
ceilu!otr
expaission
thoot
tue process
of coagulootions
tttkes
1)100cc under
the
sinape
of a circular
ooreoo, tond
tIme thickest
coagu!um
occurs
irs the close nseighnborhoocl
of tine
invto!inse hemocyte,
where
the concentrations
of the osctive substoonoce
is expected
to be the
gretotest
(plate
1, figs. 2, 3).
mm the
second
cvtoplasmic
PhSica!
wouldl
oi)servationn
troonnsparennt
cal
torounnd
between
identity
the
exhibited
pseudopoclia!
variations
in
casionally
take
Tentative
Distribution
little
the
of
of veil-like
of the
inennocytes,
dirawing
of
the
ai)Out
tons innportoonst
increase
of
the
This
dispersion
tlnann when
the
and
loorge areas.
coagulunn
hyaline
plasma
hennocyte.
veils
seenms
to
be further
supported
up and crumpling
under
nmechannical
oogenncies, the thin
jelly-like,
elastic
and contractile
consistency
as the typiin species
with conspicuous
ploosma
coagulation,
in-
thickness
the aspect
hvalinse
over
veil-like
expanded
islands
by rolling
the same
Conversely,
assume
tine
brinngs
of the ooctive
sui)stance
tornd more
extended
coagulation
islands.
1w
strirogs
a relatively
that
veils
co)agulation
divic!ual
oolteratiorn
anno! to dispersion
for to thinner
limited
is
The
of
unto
interfaces
be responsible
reaction
i)y
kind
substance
the
coagulation
structures
(plate
Types
of
islands
2, fig.
Hemolymph
show
that
these
cttn
oc-
9).
Coagulation
among
the
Insects
Investigated
In
the
present
scribed.
Besides,
fication
of the
nnat-erial,
large
every
gradation
individual
material
was
variations
recorded
in
found
occurred
table
1 in
between
sharp-cut
types
the
in several
species.
of pictures
Therefore,
coagulatiorn
types
de-
a classi-
is not
presently
i)eeno
observed
from
different
possible.
Development
of
gradations
in
ith
but
orders,
hvaline
islarnds
to
the
process
by
the
of
veil-like
oi)served
is aimsent
total
investigated.
of
occasionally
by mechanical
study
of the
present
of
coagulationn
the
species
hennolymph
the
films
hyaline
combined
contribution
of
of the
all
of
the
time
speci-
coagula-
henmolymphs.
hemocytes
with
of
fornnsationn
followed
islands
of
of Odonata,
Lepidoptera,
Coleoplera.
spontaneously
irs Orthoptera.
In the
agencies
(looying of the coverglooss
seldom
nnostlv
the
species,
the
has
various
A general
in
Inn these
i)v
henmocytes
in
occurs
coagulation
species
place
Lepidoptera
meshworks
inn various
hyaline
substance
studied.
structures
or takes
around
coagulating
consistently
tonc! of Forficula
of pseudopodial
it is in(!uced
sure,
coagulation
of
in
the
is followed
format-iorn
s5)ecially
of
amount-
founnd
of Orthoptera
Constitution
tionn
not
hemocytes
mens
tiorm
was
islands
the
by
frecjuennt
coaguhotion,
This
insects
of altera-
of this
tue
onoto
was
type
order,
slide,
pres-
traction).
The
tribution
fooctors.
However,
lymph
coagulrttion.
Factors
material
l)ictures,
the
Responsible
order
for
does
not
zoological
Orthoptera
the Differences
show
tony striking
classification
appears
of
be highly
to
Exhibited
coinsciolensce
innsects,
evolutive
homogeneous
by Insects
hetweero
or
oos regoorc!s
as J?eyards
(lis-
ecological
inennno-
Hemoiymph
Coagulation
Differences
senssitivenness
inn the
to
foreignn
relottive
surfooces,
number
inn tine
of hvaline
aniounnt
hemocvtes
or
efficiermcy
in
of
time
blood,
inn degree
to cotogu!ootionn-inoc!ucirog
of
their
sub-
From www.bloodjournal.org by guest on June 18, 2017. For personal use only.
CII.
stance
expected
availoible
to
inn tue
serve(1
inn the
be
present-
ins these
Ioensolynspio,
elenmenits
nnight-
coagulootionn
be
process
of
of
As
shown
ins table
I , species
centooge
little
of elemenots
There
seennms
conotooct- with
cells appeared
for
necess:orv
to
I)ifferenmces
degree
of
as
inn tine
otnmounts
High
coagulunn
4, figs.
17,
art-ificiool
around
18).
Onm tIne
rupture
(Geotrupes
inn Orthoptera,
founsc!
expttnnsionos,
ster-
the
winile
winich
takes
1)100Cc sponot
Whets
the
tonic-
rostra(ins).
mtoteritols
tore
(hiluted
is
earn he olecreased
about
hemocytes
probtoi)le.
(Gryllotalpa)
in the
oobove
mo plasnmos
henmocytes
reIn-
coagulation.
with a relootively
low
pera conspicuous
coagulat
ions.
of the
hyaline
hemocytes
to
woos
(Cychrus
described
i)etween
of
in
(connpare
reactions
co-
exhibiting
1, fig.
1, to
in Hydrophilus,
occurs
to the
of Gryllotaipa
species
plate
Meisero-
l)roportionsally
hemolynsph
insect
henmolynmph
with
only
plates
even
a
3 and
follosving
inennocytes.
of Former
The
reports
ins the
the
conceptions
Several
degrees
type
isotnnd,
oh-
of Lepidoptera
and of Coleoptera.
ho
film of hemolynmph
(plate
6, fig. 31) is
tine
I)rinsg
loyalinse
isyoohinse
nmaterial
vooriootionss
Investigations
Hemoiymph
on
Coagulation
Insects
in
A rthro pods
flict-ing;
tion
of
beetle
those
other
Study
Other
of lability
species
squeezing
of dilutions
their
the
of
Comparative
and
have
senositivenoess
of cotogultoble
resembling
the
importansce
hyalirse
species
(Caiaiosins)
tos regards
grouro!
rates
in
of correlootion
anc!
of
of 1)seudopodliool
the
of coagulable
irnvolved
(!egree
Conversely,
for their conspicuous
cotogulontiorn
the size of tine cooogulations
isloonsds
pictures
narrow
of a highs
)ercenstage
category
extrusion
oomounts
fooctors
ioennolvnmpin
A high degree
stable
ins various
species,
dilution.
agulottionn
same
a strong
oohout
selected
1tinmger,*
to higin
coagulootions.
differences
instance.
bring
in other
of insects
heimer’s
the
exist
glass surfaces.
to he nmore
Hydrophil-u.s-,
ouslv
of
to
wit-in
mo
or
the
insects.
Ins tine present
nnaterial
there
is mo evidence
nsumber
of the invalinne hemocvtes
inn the
exhibit
oonnd inn tine
sonic
t-ive
corarius)
1189
GREGOIRE
and
on Ioenmolvnnpio
ors its mechanism
are
literature
in complexity
zoological
were
cooogultotions
controversial.
reported
with
irs innsects
little
or
no
are
scanty
relation
torso! cons-
i)etween
eooogula-
classification.
pointed
out ins specimens
of Coleoptera
(Hgof Dytiscus
narginalis,4
#{176}Orthoptera#{176} (Stenobothrus
parallelus)
, Heoniptera#{176} (iVotonecta
glauca)
, Nen’roptera-Odonata#{176}
(larvae
of Libellula
depressa,
Aeshna
grandis),
various
caterpillars
of Lepidoptera#{176} (Bombyx
rubi, quercus
and
trifolii,
Vanessa
anliopa,
Chelonia
caja,
Pieris
brassicae
and
rapae,
Saturnia
cynthia,
Deilephila
elpenor),
Diptera#{176}(Chironomus
plumosus)
and Hymenoptera
(larval
honey
bee).7
Clotting
of the
blood
was
stated
wit-isout
details
in Coleoptera
(larvae
of Oryctes
nasicornis,nn
Meloe
proscarabaeus#{176}.
ins Orthoptera#{176} (Gryllotalpa
vulgaris)
, in Heniptera#{176} (Ne pa
Absensce
of coagulation
drophilus
piceus,.
cinerea)
, irs various
pu(Iica
Yeager,
dispar
Shull
roach
Periplaneta
cells,
tine
fusifornn
and
Knightsn
reporteti
cording to the coagulation
coagulootiorn
a cell
irssects
tionn
and/or
took
a gltoss
*
0.1;
anc!
in which
These
consiste(l
sliole
Special
KCI:
or
CaCI
solution
Bombyx
that
blood
castrensis,
Chelonia
coagulatiorn
of a cellultor
coagulation.
Many
threou!like
upons the blood
ma-
to nnegligible
in the
coagulum
of the
by
blood
pseuolopodia,
cock-
the
cells
bloool
lose
thneir
rogglutintote
into
riot
the
without
for
plasmro
cotogulations
are
groups
insects25:
coagulation,
tonnd usuoolly
shoorl)ly
drop
tue
of 47 differenst
species
of
insects,
Yeooger
he classified
into
three
tentative
groups
toe: group one inocludes
insects
exhihitinsg
no l)lasma
cell coagulations.
The insects
of group
two had
a negligible
plasnmoo
of submerging
2: 0.2.
snmears
formations
forns
the otoninnals
process
oi)served
only
tioree
pyri,
clisinntegroote.
mo or only
tons(! observing
Ringer
0.2;
thoot
the
up,
appotrently
a conspicuous
ploo(’e.
observtotionss
on
mo
coagulationn
round
oI)servatiorns
and
of
oi)vious
shapes,
and
on stairsed
coissists
nmot undergoing
clumps,
spreood out
Fronm microscopic
(Saturnia
via u/a).
Farroor3#{176} showed
discoidool
was
larva
of Lepidoptera#{176}
Harpyqia
orientalis
5)Itosma
or
,
blood
mortisaga,#{176}
caterpillars
Liparis
,
of insect
Blaps
#{176}
use
of insect
d!elinnited.
blood
while
sonme
Tine
group
degree
nmethod
in to larger
three
of
includes
cell
cotogula-
used
drop
for tinese
of oil
(Nujol)
of to coverslip.
bidist-illed
water:
1000;
NaCI:
7.5;
NtoIICO
3:
From www.bloodjournal.org by guest on June 18, 2017. For personal use only.
1190
BLOOD
According
cellular
From
IN
ARTHROPODS.
II.
to Beard2
the hemolynsph
of Galleria
inellonella
coosgulates
by forming
with only inscidental
formation
of plasma
gelatin.
Inn Popiliajaponica
gel in which
the cells tore passively
ioeld with nno osgglutinsations.
of the literature,
it appears
that inn nmost species
of insects
studie(!,
blood
agglutinations
occurs
a plasma
there
COAGULATION
this
review
coagulation
has
been
described
as
I)ensg
eitlmer
a cellultor
togglutinsations
inn whiclo
special
no
part was recognized
to be played by a special kind of cells, or a ploosnmto coagultotion,
1)0th
considered
as two physiologically
distinct
processes,
which
cars occur
irso!epennclenntly
or
together.3
Cell agglutination
was described
in agreement
with Loeb’s
oi)servationss#{176}
of clotting
in various
arthropods
and especially
in the arachnid
Limulus.
Accorolinsg
to Loch),
the whole
process
consists
in first
instance
of agglutirsation
of cells
(cell
coagulatiorm
or Fredericq’s
10. 51 first
coagulation),
which
is or is not
followed!
by a plasma
coagulation.
Unncler
the innfluence of various
mechanical-generally
strong-agenscies,
oni clusters
of hemocytes,
such
as agitation
of the blood
with
a point
of a needle30tractiorn
ons the slides
between
which
the blood has been pressed”r, the cells themselves
can be transfornsed
into structureless
gelatinous
masses, resembling
a fibrin clot (Loeb’s
cell fibrin)
witis
secondoory
development
of fibrin-like
fibrils. No special part was recognized
to he played ins the process
by to category
of cells, all elements
being
involved
in the formation
of Loeb’s
cell fibrino.
The results of this study are at variance
with
those
reported!
ai)ove.
Inn the present
ma2
‘
terial,
plasma
coagulation
was
observed
under
the
phase
constroost-
microscope
inn species
in
which none was recorded
either macroscopically#{176} or by means of the light nnicroscope3’
tond
a separation
of the process into a cellular
agglutination
and to plasnma
cotogulations
could
riot
be observed.
Differences
in the procedures
used are probably
responsible
for sonic of the divergences.
In the present studies,
as far as histologic
observations
are concerne(l,
the delicate
morphologic relations
between
hemocytes
and plasma were often corsfusedl
by a nmere gennt-le traction
of
the
coverglass
much
as
In
such
ithout
along
itself.
and
therefore
mechanical
Pictures
reactions
When
hemocytes
described
applying
alterations
have
already
above.
mechanical
described
by
of oomorphous
the contribution
In species
were
avoided
as
masses
in which
the
burtonn’s56
anic!
modiunm
ooccompansiecl
by
tue
plasnma
similar to those use! by Loch,
Yeooger
ct aS. , the
namely
the destructions
of toll tine henmocytes,
formathreads,
are founsd. No distirsctions
is thens possiI)le
tos regards
categories
of cells to the process.
oouthors,
and
various
hemolymph
Nolf’s32
alterationss
procedures
these
of the
undergorse
coagulation
was
only of to cellular
coagulation,
it is not exeludeol
agulunm around
hyaline
isemocytes
cans sometinmes
the
disturbtonnces
of the hemocytes
was
observed
to occur
tot random,
arid without
amy direct- relatiorn
to the process
of cotogulation
to the cellular
or first- coagulation
of the
literooture,
occur
disintegration
corresponding
hvaline
tion
slide,
agglutination
conditions,
cellular
winen
the
possible.
etorly
assunmptionss
(Geddes’#{176}) of cells,
but
thoot
coint-ains
described
tistot
have
the
snmall
in the
literature
a microscopic
occurre(I.
clot
first
tnnmounsts
tos connsistinng
undetected
Tinis
developedl
is
of cotogultoted
l)!asmto
would
unot
co-
Hoolli-
support
to nmmere pins-
substoonsce
olistinict
frons
cells.
Orm the
other
agulation
sidlere(!
innsect
illustrated
the
the
the
presennt
to
the
Veooger
vhich
seenme!
a-gulationm
in this
already
insect#{176},
to
no there
sosnme
be
is
gransular
o’oiunciclennt
took
for
a striking
the
with
place
analogy
ansd
l)recil)itate
without
the
of
tous(I 1’.unigiit
inn various
a5)pearedl
of
fornmatiors
coagulationi
the
of to cell
inn tine
the
or i(Ienntical
ins tine
nninoutes
eunviroinmeint
appearoonsce
i)etweers
observed
to few
imme(lioote
the
hemolvmph
wit-hits
to be sinsiltor
tolso
Belostoma,
,
tine (lescril)tiono
Veonger
cocons-
‘
three.
ooppeoors
as
Knsight
in the
instensely
insect-
stootecl
group
Orthoptera,
order
with
whicis
fiuuuinea,
their
of
ano!
a yellowish
nmost-
to
nnaterial
to
of Belostousa,
Betas/ama
of coagulationn
I)elonginsg
dleveloped
lug
exception
process
tnre inn agreenment
results
presernt
water
1 . Accordirng
have
As
,
inn the
cinerea,
blood
of
ann inmterestinng
as
Inm this
table
hsansol,
picture
process
wtoter
to that
scorpion
otiser
species
tine
withdroiwal
after
ins the
pltisnssoo
of cert:oins
surrounnoliung
best
.Vepa
listed! irs
of the
onnsol seemed
cells,
to
cytolysis
grooniules.
of
The
co-
cotogulurns.
house
cricket
of olevelopnmeust
touno! in
the
stick
of coagulation
From www.bloodjournal.org by guest on June 18, 2017. For personal use only.
Cu.
islands
iii
torounsd
crustaceoo
nmodes
of
1)1000!
without
any
with
des
Invooline
jellying
5)lasma
cooogulation
sinnple
ploosnna
blood
special
jellying
process
this
oI)serve(!
tlsree
chist inset
pltosnma
inn two
inssignsificarnt.
winich
sue-
The
occur
“explosive
time
that
tine blooc!
corpuscles
of tue hloool
corpus-
of tIne
clots,
Hardy’s
wisich
of
relatively
globultor
corpuscles,
an(!
tot- letost
(3) jellying
;
beinng
occurs
rsmtoteriool,
togglutinootions
; (2) agglutinototiono
5)riniitively
blood
preseist
Tait#{176}#{176}
recognized
of tIne plasnma
of localized,
to
inn the
cell-agglutinsationn
conssists
to seconso!
: (1)
jellvinng
preliminary
relootion
stooge
also
annd Gunrm.2729
of the
general
the
inn imineditote
Tait
ins crustooceto
subseojuenit
sul)sequent
stages,
recorded
Tait,
coagulation
cessive
looter
hemocvtes,
liv Iloordy,’7
1191
GREGOIRE
first
arounc!
or
corpuscles.”
involves
At
a
tine
whole
of
the
re-
was
fouru!
irs several
nnmooinsinng pltosnnto.
Inn the
species
I)resennt
anol
On
the
nsooterial,
(‘onisist
the
of the
Nature
irocess
ly occurred
enst
inovolved
inn tine
i nn Orthoplera
ilyaline
C of Tooit
type
to nnd 1)ermaptera.
Hcmocytes
Involved
in
the Initiation
tine
phoose
of the Plasma
Alterations
Inn fresh
invoolinne
cvtes.
On fixed
(ploot-e
6)
coons be
tine
These
closssificationss
of innsect
loio!es
, ‘
In
‘2
These
cells
seenn
blood
smears
reltot-ivelv
highs
iii
casional
of
tine
identified
tine
of
overlooked
and
present
the
the
nuclei
founsd
of these
cellular
on glass
surrounsdec!
vicinity
colorless
hvaline
less
existing
innclusionss
literature
undergo
Rooseboomt#{176}
outlined
inn
several
ann(! stainninng.”
a
“leucocytes
inenmatologic
they
tlntot
Forficula,
granules
of a iovalinne
tine
in
alteratiorns
of sharply
to few
existenoce
with
heenn
rapid
Carausius
percentage
presensce
susl)ect
of
of fixation
speciesno
gennerally
2. 8.
cristal-
res , ‘ ‘ ‘ ‘leucocvtes
a cyt oplasme
poussi#{233}reux . “
with suspensions
of india ink, milk glohules
or
observed
to insgest particles
in their
cytoplasnn,
numnnulai
injected
to have
tore
the
of henno-
ion sonne
they
oonsy of the
within
the
niicroscope,
cootegories
ooppetor
hut
nsethods
with
other
still
altered,
on cloossical
conotroost
tine
differenmces
witln certooinoty
1, fig. 5), they were
(!escrihed
elsewhere.8-
orm toccounot
frons
distinoctly
similarities
insclusiorns
Gryllotalpa
baker’s
yeast- (lltote
like the ph:ogocytes
proi)ai)ly
based
otuest-ionable
a
of
tore
i)e ranged
hemocytes,
‘leucocvtes
‘
urnoler
difficulty
tine cytologic
isemocytes
cannnot
exisibit
specinmens
without
snnears,
hnytolinne
elennensts
22, 24 They
ni,
extonminle(l
recognized
rors(! stooirsed
thought
:op)oorenst.
no,
of loennolvnmpho
l)repoorationss
hennocvtes
i)OdlV.
ito
by
all her
bits
structures
Tinese
ono insect
surfttces.
blood,
However,
preparations
a
of cytoplasnn.
lead
remnsants
the
()c-
author
could
to
perhaps
l:e
isennocytes.
SUMMARY
1
.
Thins
films
61 differcnnt
process
of freshly
species
were
coagulation
of
picture
recordinsgs
were
2. Inn the isemolymph
hemocytes
the
(-ann easily
other
cytes
kinnds
witis
orn fixations
3,
Inn the
hemocytes
do
be
unot
cells.
were
ins which
exhibit
tastIer
tine
of about
the
Control
phase
of
unnder
Attempts
hi tine
the
phase
ann importaust
if nnot exclusive
innsects
bebonigiung
a category
constrast
to
annd tine
by motions
of inyahinse
microscope
from
of idennt-ificatioun
of these
hyalinne
(‘lassificationns
of insect
i)lood
cells
memobased
insconnclusise.
the
anny
1)100(1
important
clotting
alterations.
coagulat-ionn
occurs,
the cells of tinis category
appear
to glass surfaces.
Tisey unndergo
rapid
modificationns
the present
material
lated
to a differeunt
together.
360
cosntrast
microscope
the
oi)servat-ionns
ins a fesi species.
innsects
investigated,
recognnized
described
stainninng
innsects
hemolymph
performed
of all
of blood
elements
aunt!
drassnn
observed
iisvestigatetl.
part
inn tine
innitiationn
does
nnot occur,
Inn species
tinese
to i)e iniginly labile
inn tiieir struncture
of tine
plasma
hyalinse
inn which
i)lood
to coustact
svinich play
coagislations.
these
alteratiouss
are of two kinsds, eat-is of them
appearannce
of tine plasma
i-eactions.
Tiney
cann occtrr
i)einng
alone
Inn
reor
From www.bloodjournal.org by guest on June 18, 2017. For personal use only.
1192
BLOOD
COAGIJLATION
IN
.
ARTHROPODS.
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1194
BLOOD
COAGULATION
IN
ARTHROPODS.
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CH.
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1196
BLOOD
COAGULATION
IN
ARTHROPODS.
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From www.bloodjournal.org by guest on June 18, 2017. For personal use only.
CH.
4.
do
at
Inn (‘onntrast-
to
tine
hyalinnc
hemocyt-es,
snot take
part
ins the
process
of
raundom,
they
are passively
emi)etlded
Inn the
5.
process,
These
preseust
material,
innitiat-e(l
alteratiouns
i
ma-d’ros(’opid’
microscope.
6. The
resunlts
at
cOaguslatu)nn,
svith
Tait,
as
degrees
to a limiteti
svith
former-
Tait
coagulatious,
detectable
data
from
onnly
inn ivinich
as tsvt) physiologically
or together.
from
of hemocytes.
plasma
reactions
cells
agglutinnated
to be a conntinnusoiss
category
of
tlistinnct
(inn tine other
hannd,
tine
unnder
the
coagulationn
the
other
1)100(1
(‘ells,
processes,
tine presennt
of
svhio’h
results
inas
a specific
ausd
(-ann
shosv
inn crustaceans
i)lood
by
of cells, Hardy’s
explosive
the
(listinigusished
of blood
or
appears
inn a single
type
of coaguslatioun
tles(’ribed
a-nnd Gunsun, inn si-hit-h a spet’ial
(‘ategory
analogy
categories
as beinng eithen’ a celluslar
agglutinnatioun,
inn which
rio
to he played
by a special
category
of cells, or a plasma
(‘ounsidere(l
1)0th
inntlepennderntly
occur’
place
by various
variance
innset’t- 1)100(1 si-as (leso-rii)ed
special
part si-as recognnized
other
(‘oagunlatious
taking
(-lOttinmg
are
the
o’oagurlationn.
Scattered
inn the (‘oaguslum.
hemolympin
alterationss
are follosved
a genneral
1197
GREGOIRE
ann
Han-dy,
c-ells,
importannt
part
to
stonsg
(he
play.
R1;FF;Rl-:xc’Es
( A BE
1 , L. ,
1 AR5
ANI)
2
a
M.
deae.
4
ND
.J
Bull.
LHO5TE,
: Count
.
a I ‘et 01(10!
nh )ut ions
rsiorphsOlOgi(ue
Belgique
Biol.
Frtonnce
et
saunguirne
cinez l’Inmoogo
37,
82:
de Jorficula
clu
1948.
auricularia.
Soonng 17:
1946.
GABE,
_,
.*
(/eeeuolineala
LItOSTE,
596;
?iI .
,
Say.
,J. : Lto cytologic
(hrysoinela
11ev.
AN!)
J.
Lu-IOSTE,
caflto(l.
de
bioh.
: Conntributions
8:
184,
J. 0. WAKEI.1x
ANI)
ARNOLt),
inarjina1is
tonnc! Hydrophilus
BARRATT,
I)ytiseus
o’o l’#{233}tude nnorphologi(ue
(lu
sang
des
.l!anti-
1949.
G.
: A studs-
pieces.
of
the
of certain
blood
J. Microscop.
Quart.
Sc.
Coleopteroo:
56:
149,
1910-
1911.
H.
5 BEARD,
‘
-
inssects.
: Fx5)erinmental
Anstot.
Bee.
: 1xperimenntool
47,
ohservtotionss
736,
101:
ons
hsenmolynsmpim
cotogulationn
inn two
species
of
1948.
observations
ons cooogulootions
of
irssect
isensolvmph.
Phvsiol.
Zo#{246}I.23:
1950.
G. A., BRnu;s.
7 BnsHon,
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1951 6: 1173-1198
Blood Coagulation in Arthropods: II. Phase Contrast Microscopic
Observations on Hemolymph Coagulation in Sixty-one Species of Insects
CH. GRÉGOIRE
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