C o m m u n ic a tio n
T H E
N E M A T O C Y S T
p ré se n té e
O F
le 2 8
H Y D R A
m ai
ig ô o .
(P a rt II).
THE MATURATION OF NEMATOCYSTS ÍN HYDRA
by
A llis o n L . B urnett,
Laboratoire de Zoologie et de Biologie anim ale,
Université Libre de Bruxelles
ACKNO W LED G EM ENT.
T h i s w o r k w a s s p o n s o r e d b y a p o s t - d o c t o r a l g r a n t a w a r d e d to
t h e a u t h o r b y th e N a t io n a l S c ie n c e F o u n d a tio n
S ta te s o f A m e r ic a .
o f th e
U n ite d
T h i s g r a n t a llo w e d t h e a u t h o r to w o r k in
th e la b o r a to r y o f D r . P a u l B r ie n fo r a p e r io d o f o n e y e a r .
T h e a u t h o r w i s h e s to e x p r e s s h i s d e e p e s t t h a n k s to D r . P a u l
B r ie n
w h o se
m any
su g g e s tio n s a n d
k in d
e n co u ra g em e n t w ere
a g r e a t s o u r c e o f s t i m u l a t i o n to t h e a u t h o r d u r i n g t h e p r o g r e s s
o f th e s e s tu d ie s .
I n t r o d u c t io n .
M o s t p a p e rs in s c ie n tific lite r a tu r e w h ic h d e a l w ith th e s u b ­
je c t
of
th e
w ill c o n t a i n
d e v e lo p m e n t
so m e w h e re in
of
th e
th e ir
a n d « I m m a tu r e » n e m a to c y s ts .
n e m a to c y s ts
pages
th e
in
te rm s
c o e le n te ra te s
« M a tu re
»
H o w e v e r, n o s in g le a u th o r h a s
s h a r p l y d e f i n e d w h a t is m e a n t b y a m a t u r e n e m a t o c y s t o r h a s
g iv e n
a d e f in ite
c rite rio n
w h ic h
m a tu r e a n d m a tu r e n e m a to c y s ts .
le n t stu d ie s o n
a id
of an
th e
e le c tro n
w ill d if f e r e n tia te
b e tw e e n
n e m a to c y s t h a v e b e e n c o n d u c te d
m ic r o s c o p e .
im ­
I n r e c e n t y e a rs , s e v e ra l e x c e l­
O u ts ta n d in g
am ong
w ith
th e
th e s e s tu ­
d i e s a r e t h o s e o f S e m a l - V an G a n s e n (1 9 5 4 ), B o u il l o n e t a l. (1 9 5 8 ).
270
SO CIÉTÉ ROYALE ZOOLOGIQUE DE BELGIQUE
and
S l a u it e r b a c k
(1 9 5 9 a ,
1959 b ).
F a w c e t t (1 9 5 9 ),
and
A lth o u g h ,
a u th o rs
th e s e
and
Chapm an
have
T
il n e y
p re s e n te d
in
m u c h d e ta il a d e s c rip tio n o f th e fin e s tr u c tu r e a n d d e v e lo p m e n t
o f t h e n e m a t o c y s t c a p s u le , t h r e a d
and
o p e rc u lu m , th e
o f n e m a to c y s t m a tu r a tio n re m a in s in c o m p le te .
p ro c e ss
F o r it m u s t be
h e l d i n m i n d t h a t t h e m a t u r a t i o n p r o c e s s o f t h e n e m a t o c y s t is
n o t m e re ly o n e o f m o rp h o lo g ic a l d e v e lo p m e n t, b u t o n e o f c h e ­
m ic a l d e v e lo p m e n t a s
w e ll.
C le a rly , it
s tru c tu ra lly
p e rfe c t n e m a to c y s t w h ic h
e la b o ra tin g
th e
p o is o n
w ith in
th e
is
p o s s ib le
is s t i l l i n
c a p s u le .
ro u g h s tu d y o f n e m a to c y s t m a tu r a tio n
to
have
a
th e p ro c e ss o f
T h e re fo re ,
a
th o ­
m u s t in c lu d e jo in t c h e ­
m ic a l a n d a n a to m ic a l s tu d ie s .
A
s im p le
d e fin itio n
f o l lo w s , « A n y
m a tu re
».
of
a
m a tu re
n e m a to c y s t w h ic h
H o w ev e r,
W
e il l
n e m a to c y s t
m ig h t
be
as
is c a p a b l e o f d i s c h a r g i n g
(1 9 3 4 ) a n d
and
C hapm an
T
is
il n e y
(1 9 5 9 b ) h a v e s t a t e d t h a t i m m a t u r e n e m a t o c y s t s a r e c a p a b l e o f
e v a g in a tin g .
It
is
re g re tfu l
th a t
th e
la tte r
a u th o rs
w e re
not
m o r e s p e c ific i n d e s c r ib in g t h e s ta g e w h e n a n e m a t o c y s t is f ir s t
c a p a b l e o f d i s c h a r g i n g , b u t i t is o b v i o u s t h a t t h e s e i n v e s t i g a t o r s
h a v e in d e e d
o b se rv ed
d is c h a rg e d
n e m a to c y s ts w h ic h
w e re n o t
f u lly s c u lp tu re d .
W h e r e d o e s o n e e x p e c t t o f i n d a m a t u r e n e m a t o c y s t i n c o e le n te ra te s ?
w ill b e
It
fo u n d
a p p ea rs
in
th e
m ost
lik e ly
te n ta c le s ,
th e
e m p lo y e d to th e g r e a te s t a d v a n ta g e
re g io n o f th e
a n im a l.
A
a h y d r a w ill r e v e a l, a s K
s im p le
epner
th a t
m a tu re
p la c e
w h e re
ra th e r th a n
lo n g itu d in a l
n e m a to c y s ts
th e y
in
w ill b e
th e g a s tric
s e c tio n
th ro u g h
(1 9 4 3 ) h a s s t a t e d , t h a t n e m a t o ­
c y s ts a r e f o u n d in t h e p ro c e s s o f f o r m a t i o n in t h e d is ta l o r o ra l
■4 o f t h e b o d y . S e m a l -V an G a n s e n ( 1951 ) h a s p r e s e n t e d a m u c h
m o re e la b o ra te sc h e m e f o r g e n e ra l n e m a to c y s t d is tr ib u tio n .
She
c o n s id e rs e a c h r e g io n o f th e b o d y o f h y d r a s e p a ra te ly a n d o ffe rs
a
g e n era l
account
of
fo u n d
in
each
r e g io n .
g a s tric
c o lu m n
is r i c h
th e
b ase
and
o f t h e s e c e lls .
p e d u n c le
th e
num ber
She
s ta te s
and
k in d s
th a t
th e
of
n e m a to c y s ts
e p id e rm is
i n c n i d o h l a s t c e lls w h i l e t h e
a re
s p a rs e ly
p o p u la te d
by
of
th e
re g io n s o f
o r d e p riv e d
I n th e te n ta c le s , w h e re th e n e m a to c y s t b a tte rie s
a r e f o r m e d , t h i s a u t h o r c o n s i d e r s t h e n e m a t o c y s t s t o h e « r i p e ».
O n c e a n e m a t o c y s t is p a r t o f t h e n e m a t o c y s t b a t t e r y , t h e c e ll
c o n t a i n i n g t h e n e m a t o c y s t is n o l o n g e r c a l l e d a c n i d o h l a s t b y
S e m a l -V an G a n s e n b u t a n e m a t o c y t e .
T h e n e m a to c y te a lw a y s
h a s its p o s itio n in a n e m a t o c y s t b a t t e r y lo c a te d in t h e te n ta c le s
o f th e
h y d ra ;
th e
F re n c h
e q u iv a le n t fo r th e
te rm
n e m a to c y s t
( 1 9 5 9 -6 0 )
ANNALES 9 0
b a tte ry
is « b o u t o n
u rtic a n t
».
T h u s,
271
we
m ay
assu m e
fro m
S e m a l -V an G a n s e n ’s d e s c r i p t i o n t h a t a n e m a t o c y s t i n t h e g a s t r i c
re g io n
is i n
th e
s ta te
o f d iffe re n tia tio n
w h e re a s a
n e m a to c y s t
in a t e n t a c l e b a t t e r y is c o m p le te ly f o r m e d .
H o w e v e r, if o n e m a k e s a l o n g it u d i n a l s e c tio n t h r o u g h a h y d r a
a n d c o m p a r e s a s te n o te le in t h e te n ta c le w ith a s te n o te le in th e
g a s t r i c r e g i o n , i t w ill b e f o u n d , a t l e a s t a s f a r a s c a n b e d e t e r ­
m in e d w ith th e l ig h t m ic ro s c o p e , t h a t b o t h c o n ta in a f u lly e la ­
b o r a te d t h r e a d , c a p s u le , a n d o p e r c u lu m .
an
im p o rta n t
n e m a to c y s t
been
q u e s tio n
in
fo rm e d
th e
a ris e s ;
g a s tric
and
F ro m
is t h e r e
c o lu m n
a
w h e re
a n e m a to c y s t in
th e
th is o b s e rv a tio n
d iffe re n c e
th e
b e tw e e n
n e m a to c y s t
te n ta c le
b a tte ry
a
has
w h e re
it w ill b e e m p lo y e d ?
T h is p a p e r p u r p o r ts to a n s w e r th is q u e s tio n
h is to c h e m ic a l
te s ts
a p p lie d
to
w h o le
on
a n im a ls .
th e b a s is o f
By
e x a m in in g
w h o l e a n i m a l s i t is p o s s i b l e t o o b s e r v e t h e g e n e r a l d i s t r i b u t i o n
and
c h e m ic a l p a tte rn
fo u n d
in
h y d ra
M
The
a n im a ls
o f a ll f o u r o f th e
ty p e s o f n e m a to c y s ts
in ta c t.
a t e r ia l s a n d
e m p lo y e d
in
th e s e
M
etho ds.
s tu d ie s
w e re
h e a lth y , a d u lt
s p e c i m e n s o f t h e b r o w n h y d r a , P e l m a t o h y d r a o l ig a c t is . M e t h o d s
fo r
re a rin g
th e s e
a n im a ls
have
been
d e s c rib e d
in
a
p re v io u s
p a p e r (S e e B u r n e t t , 1 9 5 9 ).
T h e a n im a ls w h ic h w e re c h o s e n f o r h is to lo g ic a l e x a m in a tio n
w e re
1.
p re p a re d
R e la x
in
th e
th e fo llo w in g m a n n e r
h y d ra
fo r o n e
m in u te
:
in
a s o lu tio n
of
io
%
a lc o h o l.
2.
3.
F ix fo r
'4 h o u r i n
100 %
a lc o h o l.
B r in g th e h y d r a d o w n t h r o u g h a g r a d e d se rie s o f a lc o h o ls
to w a te r a n d im m e rs e th e m
f o r o n e o r tw o m in u te s in a
m e th y le n e b lu e s o lu tio n (c o n c e n tr a tio n
.01
% ). A c c o r d in g
t o t h e t y p e o f n e m a t o c y s t d e s i r e d t o s t a i n , i t is n e c e s s a r y
to
s ta in
m e th o d
th e
h y d ra
a d o p te d
S in g e r a n d
D
in
s o lu tio n s
fo r th e
em psey
o f d iffe re n t
p re se n t s tu d y
(1 9 4 7 — t a k e n
fro m
has
p . H ’s.
been
P ea rse,
The
th a t of
1954, p .
434 )M e th y le n e b lu e s o lu tio n s w e re b u ffe re d in th e p r e s e n t s tu d y
w ith
M ic rh o m e
B u ffe r T a b le ts
G u r r L td ., U p p e r R ic h m o n d
p u rc h ase d
fro m
Edm und
R o a d W . , L o n d o n , S .W .
14.
272
SOCIÉTÉ ROYALE ZOOLOGIQUE DE BELGIQUE
E n g la n d .
B u ffe r
e m p lo y e d
4.
in
s o lu tio n s
of
th e
fo llo w in g
p . H ’s
w e re
t h e f o l l o w i n g s t u d y : 3 .6 , 4 , 5 . 7 , 8 , 9 ,
io .
D if f e r e n tia te in 30 % a lc o h o l u n t il t h e b o d ie s o f t h e h y d r a
are
s ta in e d
a
lig h t b lu e .
At
th is
s h o u l d b e s t a in i n g d e n s e ly .
B r i n g r a p i d l y t o 100 % a l c o h o l t o
5.
tim e
th e
n e m a to c y s ts
p r e v e n t f u r t h e r lo s s o f
s ta in .
6.
C le a r in
7.
M o u n t w h o le a n im a ls in
T h is
m ay
x y le n e o r to lu e n e .
m e th o d
be
b a ls a m .
p ro v id e s fo r a r a p id
c o n s id e re d
s p e c if ic
fo r
s ta in in g
n e m a to c y s ts
p ro c e d u re
sin c e
w h ic h
th e s e
s tru c ­
tu r e s c o n c e n tr a te th e s ta in to a m u c h g r e a te r e x te n t t h a n
o th e r s tru c tu re in h y d ra .
any
S l i d e s p r e p a r e d b y t h i s m e t h o d w ill
o fte n b e g in to fa d e a f te r a p e rio d o f f r o m
o n e to tw o m o n th s ,
th e r e f o r e , i t is r e c o m m e n d e d t h a t t h e s lid e s b e e x a m i n e d w ith in
a w e e k o r tw o a f te r p r e p a r a tio n .
I t w a s a ls o n e c e s s a r y d u r i n g
su es o f o n e h y d r a
to
th e
th e s e s tu d ie s to g r a f t th e
tis s u e s
to
a n o th e r
o n e o r tw o
b e fo re th e a n im a ls w e re fix e d f o r h is to lo g ic a l s tu d y .
tis ­
days
T h e g ra ft
in a ll c ases c o n s is te d o f th e p e d u n c le o f o n e a n im a l w h ic h w as
p la c e d b e tw e e n th e g r o w th re g io n a n d b u d d in g r e g io n o f a n o r ­
m a l a n im a l.
F o r a d e s c r ip tio n o f th e s e g r a f t i n g te c h n iq u e s see
B u r n e t t ( 1 9 5 9 ).
O b s e r v a t io n s .
A
g ro u p
o f 250 h y d ra
w e re s ta in e d
t i o n s o f v a r i o u s p . H ’s a n d
scope.
A t
p .H
8,
th e
th e n
s te n o te le s
p e d u n c le o f th e a n im a l p lu s th e
th e
te n ta c le s w e re s ta in e d
bases
of
re g io n s
The
th e
of
s ta in in g
th is p .H
te n ta c le
th e
p lu s
a n im a l,
th o s e
in
o n ly
b lu e so lu ­
w e re
lo c a te d
in
th e
w e re
g a s tric
and
b u d d in g
at
a
p a tte rn .
th is
a s lig h t a ffin ity fo r th e
d id im p r e g n a te th e s ta in w e re in
of
a t th e
u n s ta in e d
s im ila r
th e
H
th e d is ta l
S te n o te le s lo c a te d
(*) f o l l o w e d
d esm o n em es h a d
th o s e w h ic h
w h ic h
m e th y le n e
u n d e r a lig h t m ic ro ­
s te n o te le s in
in te n s e ly .
h o w e v er,
o f is o rh iz a s
in
o b se rv ed
th e d is ta l
p .H .
At
s ta in :
H
of
th e te n ta c le s .
A t p .H
9
th e
s te n o te le s
m a n n e r a s th o s e a t p .H
and
is o rh iz a s
s ta in e d
in
th e
sam e
8 e x c e p t t h a t th e n e m a to c y s ts in
th e
(*) E xam ination of whole m oun ts of P. oligactis stained w ith m ethylene
blue fails to reveal any criterion whereby it is possible to distin guish a
holotrichous isorhiza from an atrichous form, thu s the general term
« isorhiza » w ill be em ployed in th is paper.
ANNALES 9 0 ( 1 9 5 9 - 6 0 )
p r o x i m a l 1/ 3 o f t h e
and
n e m a to c y s ts
p e d u n c le .
A t p .H
te n ta c le s w e re u n s ta in e d
s ta in e d
T h ere
273
o n ly
in
th e
on
lo w e r
t h is o c c a s io n ,
2/ 3 r e g i o n
o f th e
w a s n o s ta in in g o f d e s m o n e m e s a t th is p .H .
io o n ly th e
s te n o te le s h a d
an
a ffin ity fo r th e
s ta in ,
a n d th e s e s te n o te le s w e re lo c a te d in th e e x tr e m itie s o f th e t e n ­
t a c l e ( F i g . 3 a n d 4 ).
T h e r e w a s n o s t a in in g o f s te n o te le s w h ic h
w e re lo c a te d in th e p e d u n c le .
S ta in in g a t p .H
3 .6 . p r e s e n t e d
q u ite a d iffe re n t p ic tu re .
t h is p .H o n ly th e is o r h iz a s s ta in .
At
T h e y s ta in e d in te n s e ly in th e
b o d y r e g io n a s w e ll a s t h e te n ta c le s .
S te n o te le s w e re g e n e r a lly
u n a f f e c t e d a t t h i s p . H ( F i g . 1), h o w e v e r , t h e r e w a s , o n o c c a s i o n ,
a
lig h t
s ta in in g
te n ta c le s a n d
A t p .H
th e
s te n o te le s
g a s tric
p .H . T h e r e
in
th e
basal
re g io n s
of
th e
r e g io n .
4 is o rh iz a s c o n tin u e d
o f th e b o d y .
in
o f th e
in
to s ta in
in te n s e ly
in
a ll a re a s
D e s m o n e m e s i n t h e t e n t a c l e s a ls o s t a i n e d a t t h i s
is a n
in c re a s in g a ffin ity fo r th e s ta in
b y s te n o te le s
t h e b a s a l r e g io n s o f t h e te n ta c le s a n d in t h e g a s tr ic r e g io n .
A t p .H
5 th e r e w a s a f a ir ly in te n s e s t a in in g o f s te n o te le s in
t h e g a s t r i c r e g i o n o f t h e a n i m a l a n d a ls o i n t h e l o w e r % o f t h e
te n ta c le s .
D e s m o n e m e s a n d is o r h iz a s c o n tin u e to s ta in t h r o u g h ­
o u t th e te n ta c le s a n d t h e b o d y re g io n .
A ls o th e r e w a s a d e n s e
s ta in in g o f th e is o rh iz a s in th e p e d u n c le .
in g in
th is
D e sm o n e m e s a re la c k ­
r e g io n .
A t p . H 7 t h e r e is a g e n e r a l s t a i n i n g o f s t e n o t e l e s i n t h e e n t i r e
b o d y o f th e
a n im a l, h o w e v e r, th o s e s te n o te le s in
c o n c e n tr a te d
th e s ta in
re s e ls e w h e re in
in
th e
body
to a g r e a te r e x te n t th a n
t h e b o d y (S e e F i g . 4 ).
re g io n
e ith e r
do
not
te n ta c le s
H o lo tr ic h o u s is o rh iz a s
s ta in
T h o s e in th e te n ta c le s s ta in in te n s e ly .
th e
s im ila r s tr u c tu ­
or
s ta in
v e ry
fe e b ly .
A ls o it w as n o tic e d t h a t
is o r h iz a s in th e d e v e lo p in g te n ta c le s o f th e b u d s ta in e d w h e re a s
th o s e in
th e
g a s tric re g io n o f th e b u d
d id n o t.
A g r o u p o f a n i m a l s w h i c h h a d b e e n s t a r v e d f o r 14 d a y s w e r e
s ta in e d
a t p .H
p a tte rn
fro m
8, a n d
th e
th e re
n o rm al
w as n o
a n im a l.
d e v ia tio n
S te n o te le s
in
in
th e
th e
s ta in in g
lo w e r
%
r e g io n o f t h e s te n o te le s s till f a ile d to c o n c e n tr a t e t h e s ta in .
T h e p e d u n c le s f r o m 5 h y d r a w e re g r a f te d b e tw e e n th e g r o w th
re g io n s
of
th e
b u d d in g
re g io n s
of
5
n o rm a l
a n im a ls .
Two
a n im a ls w e re s ta in e d a t p .H
8 o n th e firs t d a y a fte r th e g ra ft
and
on
3 a n im a ls
w e re
s ta in e d
th e
seco n d
day.
th e o n ly s te n o te le s in th e g a s tr ic re g io n w h ic h h a d
fo r th e
s ta in
g ra fte d
p e d u n c le .
w e re th o s e in
th e
In
a ll
cases
a n a ffin ity
re g io n s j u s t b o r d e r i n g
on
th e
274
SOCIÉTÉ ROYALE ZOOLOGIQUE DE BELGIQUE
F ig .
1.
Photograph of whole ten tacle stained w ith m ethylene blue at pH 3.6.
Note th e Intense stain in g of th e oblong isorhizas and round desm onem es
and the lack of stain in th e large stenoteles.
F ig . 2.
Photograph of whole tentacle stained w ith m ethylene blue at pH 7.
Note th e general stain in g of all types of nem atocysts and th e densely
stain in g sten oteles in the center of each nem atocyst battery.
F ig . 3.
Photograph of th e extrem ity of a whole tentacle stained w ith m ethylene
blue at pH 10. Note th a t only th e sten oteles stain at th is pH.
F ig . 4.
Photograph of th e tentacles of a bud stained w ith m eth ylen e blue at
pH IO. Note th a t only th e sten oteles in the tip of th e tentacles concen­
trate th e stain
F ig . 5
Regenerated ten tacle of an anim al w hich had been severed thirty hours
previously through th e gastric region. Note th e stain in g of sten oteles after
treatm ent w ith m ethylene blue buffered to pH 8 .
F ig . 6
Proximal portion of a hydra w hich had been severed thirty hours pre­
viously through th e gastric region. Note th e stain in g of isorhizas (I) and
stenoteles (S) after treatm ent w ith m ethylene blue buffered at pH 8 .
ANNALES 9 0 ( 1 9 5 9 - 6 0 )
S e v e ral
h y d ra
w e re
e x c is e d
m id w a y
275
b e tw e e n
th e
b u d d in g
r e g io n a n d g r o w t h r e g io n , a n d t h e r e s u l ti n g s e c tio n s w e re a llo w ­
e d to r e g e n e r a te fo r a p p r o x im a te ly 30 h o u rs .
T h e s te n o te le s in
th e
p ro x im a l
b u d s o f th e
p o rtio n s
s ta in e d
d e v e lo p in g
te n ta c le s
at
The
p .H
8.
o f th e
s te n o te le s
in
th e
e x c is e d
d e v e lo p in g
p o r t io n s o f t h e e x c is e d d is ta l s e c tio n s s t a in e d in a s im ila r m a n ­
n e r a t t h e s a m e p . H (S e e F i g . 5 ).
Is o rh iz a s p o ssesse d a s im ila r
s t a in i n g c a p a c ity a s t h e s te n o te le s in b o t h
p ro x im a l a n d
d is ta l
s e c t i o n s (S e e F i g . 6 ).
T
ext
f ig u r e
I.
pH 4
pH 5
pH 7
pH 8
pH 9
pH 10
+
--+
—
+
+
—
+
+
—
+
T ----+ ---
—
—
Growth, Gastric S
Budding
D
Region
I
— 9
—
+
+
--+
+
+
+
+
+
?
____
____
____
—
—
-----
—
—
Proxim al 1/4
Region o í
T entacles
__ 9
+
+
+
___
+
+
+
+
____
—
+
+
+
____
D
I
—
—
-----
—
—
D istal 3/4
Region of
Tentacles
S
I)
I
___
____
____
+
+
+ ----+ —
— +
+
+
+
+
+
+
+
+
Proximal 3/4
Reg. of Tent.
S
—
--- +
+ —
+
+ —
+ ---
—
D istal 1/4
Reg. of Tent.
3
—
---
—
+
+
+
+
D istal 1/3
Reg. of Ped.
S
—
---
—
+
+
---
—
Proximal 2/3
Reg. of Fed.
S
—
—
—
+
+
+
—
pH 3.6
Peduncle
entire
S
D
I
S
—
—
S e v e ra l w h o le a n im a ls
s o lu tio n b u ffe re d a t p .H
th e te n ta c le s a n d
H o w ev e r,
th e
p in k ,
w h ile
s ta in e d
in
—
a to lu id in e
b lu e
T h e s te n o te le s in th e d is ta l Y\ o f
t h o s e i n t h e p e d u n c l e s t a i n e d a d e n s e v i o le t .
s te n o te le s
p in k , th o s e in
w e r e a ls o
7 .5 .
—
in
th e
g a s tric
re g io n
s ta in e d
a
lig h t
t h e m o s t b a s a l re g io n s o f t h e te n ta c le s a d a r k e r
th o se
re g io n a n d d is ta l Y
in
th e
m id -re g io n ,
i.e .,
b e tw e e n
th e
o f th e te n ta c le , s ta in e d d e e p re d .
b asal
H ence,
th e r e w as a g r a d u a l in c re a s e in m e ta c h r o m a s ia e x te n d in g fro m
2j 6
SOCIÉTÉ ROYALE ZOOLOGIQUE DE BELGIQUE
t h e b a s e o f t h e t e n t a c l e s t o w a r d s t h e d i s t a l -/\ r e g i o n o f t h e t e n ­
ta c le s.
S im ila rly , a g r a d ie n t o f in te n s ity o f m e ta c h r o m a s ia w as
w i t n e s s e d b e t w e e n t h e b u d d i n g r e g i o n a n d t h e u p p e r 1/ 3 r e g i o n
o f th e p e d u n c le w ith th e m e ta c h r o m a s ia in c re a s in g p ro x im a lly
fro m
t h e b u d d i n g r e g io n .
tra te
g ra p h ic a lly
th e
T e x t F i g u r e I is a n a t t e m p t t o i l l u s ­
re s u lts
e x p e rim e n ts c o n d u c te d
of
a ll
of
th e
fo re g o in g
s ta in in g
a t d i f f e r e n t p . H ’s.
I t m u s t a ls o b e m e n t i o n e d t h a t i n a ll o f t h e f o r e g o in g s t a in ­
in g re a c tio n s , th e s ta in w as lim ite d to th e c o n te n ts o f th e n e m a ­
to c y s t c a p s u le a n d h a d n o e ffe c t u p o n t h e c a p s u le p r o p e r.
D is ­
c h a r g e d n e m a to c y s ts e ith e r d id n o t s ta in , o r if th e p o is o n w as
n o t c o m p le te ly e x tr u d e d f r o m
t h e c a p s u le u p o n d is c h a r g e , s ta in ­
e d o n ly p a rtia lly .
D
I t is c l e a r f r o m
w h ic h
is c u s s io n .
th e p re c e e d in g o b s e rv a tio n s th a t n e m a to c y s ts
a re lo c a te d in
th e
body
o r g a s tr ic re g io n
o f th e
h y d ra
d o n o t p o ssess th e sa m e p ro p e rtie s as n e m a to c y s ts lo c a te d in th e
te n ta c le s .
A lth o u g h th e n e m a to c y s ts in th e g a s tr ic re g io n h a v e
e la b o r a te d
a
c a p s u le
and
s tin g in g
th re a d , it
a p p ea rs
th a t
th e
p o i s o n i n t h e c a p s u l e is n o t f u l l y e l a b o r a t e d u n t i l t h e n e m a t o ­
c y s ts h a v e r e a c h e d
s tra te d
by
th e
te n ta c le s .
E v i d e n c e f o r t h i s is d e m o n ­
t h e f a c t t h a t t h e r e is a g r a d u a l i n c r e a s e i n
a c id ity
o f s te n o te le s e x te n d i n g f r o m
th e b o d y re g io n to w a rd s th e e x tr e ­
m itie s
s im ila r
in
o f th e
th e s e
te n ta c le s .
s tru c tu re s
S e m a l-V a n
G a n sen
( 1 9 4 9 ), d e m o n s t r a t e d
te n ta c le s f r o m
A
le n d s
fu rth e r
( 1 9 5 i) ,
in c re a s e in
s u p p o r t to
e la b o ra tin g
a
m e ta c h ro m a s ia
th is
c o n v ic tio n .
th e o ry
t h a t th e n e m a to c y s ts o f h y d r a
of
B r ie n
re ac h
th e
th e b o d y re g io n w h e re th e y h a v e b e e n fo rm e d ,
n o t t h r o u g h a n a c tiv e m ig r a t io n d is ta lly v ia th e g a s tr o v a s c u la r
c a v ity as h a d b e e n s u p p o s e d b y e a rlie r a u th o r s , b u t th r o u g h th e
n o rm al
g ro w th
p ro cesses
of
th e
a n im a l.
A c c o rd in g
to
V an
G a n se n , a c tiv e ly m u l t i p l y i n g c e lls i n t h e g r o w t h r e g io n o f h y d r a
f o r c e c n id o h l a s t c e lls in th is r e g io n b a s a lly to w a r d s t h e p e d u n c le
a n d d i s t a l l y t o w a r d s t h e t e n t a c l e s . T h u s , t h e m i g r a t i o n is a n e p i ­
d e r m a l o n e a n d a ls o is a g r a d u a l p r o c e s s .
V a n G a n s e n 's t h e o r y
w ill b e d is c u s s e d in m o r e d e ta il in t h e f o llo w in g p a p e r in th is
se rie s , b u t f o r t h e
p re s e n t it m a y
b e m e n tio n e d
th a t th e
fa c t
t h a t n e m a to c y s ts s h o w a g r a d u a l in c r e a s e in m e t a c h r o m a s ia a n d
a c id ity as th e y e x te n d
fro m
th e g ro w th
re g io n
to th e
te n ta c le
re g io n m a y b e ta k e n a s e v id e n c e in s u p p o r t o f th is th e o r y .
n e m a to c y s ts
m ig ra te d
to
th e
te n ta c le s
v ia
th e
If
g a s tro v a s c u la r
277
ANNALES 9 0 ( 1 9 5 9 - 6 0 )
c a v ity th e p ro c e s s w o u ld b e a r a p id o n e , a n d w o u ld n o t e x p e c t
to f in d
a re g u la r c h e m ic a l c h a n g e
a lo n g
th e
e n tire
le n g th
of
th e te n ta c le .
I f i t is a s s u m e d t h a t g r o w t h p r o c e s s e s a r e r e s p o n s i b l e f o r t h e
m ig r a tio n o f n e m a to c y s ts in
se s.
h y d r a , th e fo llo w in g q u e s tio n a r i­
D o n e m a to c y s ts g r a d u a lly rip e n
p ro x im a lly a n d d is ta lly fro m
c h e m ic a lly as th e y
m ove
th e g r o w th re g io n b e c a u se a fix e d
t i m e is n e c e s s a r y f o r t h e r i p e n i n g p r o c e s s t o c o m p l e t e i t s e l f , o r
d o n e m a to c y s ts c h a n g e c h e m ic a lly b e c a u s e o f in flu e n c e s o f th e
p a r tic u la r b o d y r e g io n in w h ic h th e n e m a to c y s t h a p p e n s to b e
lo c a te d ?
m ic a l
If
a
n e m a to c y s t
d e v e lo p m e n t,
th e n
re q u ire s
it
a
s h o u ld
fix e d
be
tim e
p o s s ib le
fo r
by
its
che­
in h ib itin g
g r o w t h p r o c e s s e s t o b r i n g a b o u t t h e c o m p l e t e m a t u r a t i o n o f a ll
th e
n e m a to c y s ts
in
th e
body
of
h y d ra .
( 1959) h as
B u r n e tt
s h o w n t h a t s t a r v a t i o n w ill b r i n g a b o u t t h e i n h i b i t i o n o f g r o w th
p r o c e s s e s s i n c e e n e r g y f o r c e l l m u l t i p l i c a t i o n is l a c k i n g a f t e r a
p e r io d o f a p p r o x im a t e ly fiv e d a y s .
In th e p re s e n t stu d ie s it h a s
b e e n sh o w n t h a t a h y d r a w h ic h h a s b e e n s ta rv e d fo r fo u rte e n
d a y s s ta in s
a p p e a rs
s im ila rly
a t p .H
8 to
a n o r m a l a n im a l.
t h a t a n e m a t o c y s t w ill c o m p l e t e
m e n t o n ly
w hen
i t is u n d e r
th e
T h u s , it
its c h e m ic a l d e v e lo p ­
in flu e n c e
o f th e
te n ta c le
or
p e d u n c le r e g io n , o r, c o n v e rs e ly , n o t u n d e r th e in f lu e n c e o f th e
g a s t r ic t o r g r o w t h r e g io n .
S u p p o r t o f t h e f o r e g o i n g c o n v i c t i o n is g a i n e d f r o m
rim e n t
w h ic h
in v o lv e d
th e
tra n s fe rin g
th e e x p e ­
o f a p e d u n c le
th e g r o w th a n d b u d d in g re g io n s o f n o r m a l a n im a ls .
b e tw e e n
N e m a to ­
c y s ts b o r d e r in g o n th e g r a f te d p e d u n c le r a p id ly in c re a s e in a c i­
d ity .
F u r t h e r m o r e , i f a c u t is m a d e t h r o u g h
th e
m id d le o f a
h y d r a , n e m a to c y s ts o n e i t h e r s id e o f t h e in c is io n b e c o m e a c id
w ith in
th irty
p e d u n c le
h o u rs
in d ic a tin g
o r te n ta c le s
n e m a to c y s ts .
A
th a t
n e c e s s a rily
n e m a to c y s t
th e
fo rm a tio n
in v o lv e s
lo c a te d
in
th e
th e
a
new
« rip e n in g
of
» of
m id -re g io n
of
a
h y d r a w o u ld n o r m a lly ta k e a t le a s t a w eek to r e a c h th e re g io n
o f th e p e d u n c le
T h e re fo re ,
it
th ro u g h
is
sa fe
g ro w th
to
p ro c e s s e s a lo n e .
assu m e
th a t
a
n e m a to c y s t
in
th e
g r o w th re g io n o r g a s tr ic r e g io n o f a h y d r a , a lt h o u g h i t h a s e la ­
b o ra te d
its
c a p s u l e , is
h e ld
in
a
k in d
of
c h e m ic a l
abeyance.
O n l y w h e n a n e m a t o c y s t is r e m o v e d f r o m t h e s e a r e a s o r b r o u g h t
u n d e r th e
in flu e n c e o f th e
in h ib itio n
re le a s e d .
te n ta c le
o r p e d u n c le
re g io n
is t h i s
T h e n a t u r e o f t h i s i n h i b i t i o n is u n k n o w n a t t h e p r e s e n t t i m e .
I t is a
w e ll k n o w n
fa c t
th a t
th e
h y p o s to m e
re g io n
of h y d ra
SO CIÉTÉ ROYALE ZOOLOGIQUE DE BELGIQUE
su p p re sses
th e
fo rm a tio n
d ia te v ic in ity .
of
a n o th e r
h y p o s to m e
in
its
im m e ­
I t is a ls o q u i t e p o s s i b l e t h a t t h e r e is a s u b s t a n c e
in t h e g r o w th o r g a s tr ic re g io n s o f h y d r a w h ic h s u p p re s s e s th e
fu ll e la b o r a tio n o f a n e m a to c y s t p o iso n .
A s t h e n e m a t o c y s t is
p u s h e d b y g r o w th p ro c e ss e s to th e te n ta c le s a n d p e d u n c le , th is
i n h i b i t i o n b e c o m e s le s s a p p a r e n t .
ta c le s
and
p e d u n c le
of
th e
O n th e o th e r h a n d , th e te n ­
h y d ra
m ig h t
fu rn is h
a
s u b s ta n c e
n e e d e d f o r t h e c o m p le te d e v e lo p m e n t o f th e n e m a to c y s t p o is o n ,
a s u b s t a n c e w h i c h is l a c k i n g i n t h e g r o w t h a n d g a s t r i c r e g i o n s .
I t m u s t a ls o b e m e n t i o n e d
t h a t t h e n e m a t o c y s t is b e c o m i n g
m o re a n d m o re a c id a lo n g th e e n tir e le n g th o f th e te n ta c le .
w ill b e r e m e m b e r e d
te n ta c le
t h a t a t p .H
tip s h a v e a n
a ffin ity
It
io , o n l y t h e s t e n o t e l e s a t t h e
fo r m e th y le n e
b lu e .
I t is q u i t e
p o s s i b l e t h a t i n c r e a s e d a c i d i t y is a r e f l e c t i o n o f t h e p r o d u c t i o n
o f a m o r e p o t e n t p o is o n in t h e n e m a t o c y s t c a p s u le .
S in c e th e
n e m a to c y s ts m o s t o fte n e d e m p lo y e d in th e p ro c e s s o f s u b d u in g
p r e y a r e t h o s e i n t h e e x t r e m i t i e s o f t h e t e n t a c l e s , i t is o f g r e a t
a d v a n ta g e
to
th e
a n im a l
if
th e s e
n e m a to c y s ts
a re
th e
m o st
p o te n t o n e s in th e b o d y .
On
th e
o th e r h a n d , c h e m ic a l c h a n g e s a re
n e m a to c y s ts
such
p o is o n u p o n
d is c h a rg e .
h o llo w
s tru c tu re ;
as
d esm o n em es
a
u n k n o w n a t th e p r e s e n t tim e .
i.e .,
p ie rc e d
th e
and
lib e ra te
in
any
flu id , t h a t n a tu r e
o f w h ic h
is
I t is q u i t e p o s s i b l e t h a t t h e d e s ­
a n e m a to c y s t t h a t w as o n c e a p e n e tra n t,
p r e y 's
a lth o u g h
r e ta in s a p o is o n
w itn e s s e d
not
A l s o , t h e d e s m o n e m e is a h i g h l y
s p e c ia liz e d ty p e o f n e m a to c y s t.
c h a rg e ,
a ls o
do
N e v e r t h e l e s s , t h e d e s m o n e m e is n o t a
it c o n ta in s
m o n e m e e v o lv e d f r o m
w h ic h
tis s u e s
and
it
becom e
has
lib e ra te d
p o is o n
g re a tly
upon
d is ­
m o d ifie d ,
s t i ll
i n s i d e i t s c a p s u le .
I t w ill b e n o t i c e d t h a t i s o r h i z a s s t a i n a t p . H
3 .6 . a n d a ls o a t
p .H 8 in th e a r e a o f th e te n ta c le s a n d p e d u n c le .
T h e c o n flic tin g
n a tu r e o f th e s e re s u lts p r o b a b ly
is d u e t o t h e f a c t t h a t i t w a s
im p o s s ib le
d iffe re n tia te
in
th e s e
stu d ie s
to
c h o u s a n d a h o lo tr ic h o u s is o rh iz a .
c y s ts
m ay
have
q u ite
b e tw e e n
an
a tri-
T h e s e tw o ty p e s o f n e m a t o ­
d iffe re n t c o n te n ts
in sid e
th e ir
c a p s u le s .
H o w e v e r , S e m a l -V an G a n s e n (1 9 5 1 ) p o i n t s o u t t h a t t h e r e a r e n o
a t r i c h o u s is o r h iz a s in t h e p e d u n c le o f th e h y d r a .
I f t h i s is t r u e ,
t h e n i t is d i f f i c u l t t o e x p l a i n t h e s t a i n i n g c h a r a c t e r i s t i c s o f h o l o ­
tr ic h o u s is o r h iz a s in
th e
p e d u n c le a t h ig h ly a c id
and
a lk a lin e
p . H ’s.
In
c o n c lu s io n , it m a y
be
sa fe ly
s ta te d
th a t
th e
s te n o te le
of
h y d r a is n o t c a p a b l e o f c o m p l e t e l y e l a b o r a t i n g i t s p o i s o n u n t i l
ANNALES 9 0 ( 1 9 5 9 - 6 0 )
279
it re a c h e s th e re g io n o f th e te n ta c le s o r p e d u n c le .
T h e fa ct th a t
t h e r i p e n i n g p r o c e s s is a g r a d u a l o n e e x t e n d i n g f r o m
to
th e
re a c h
e x tre m itie s
th e
of
te n ta c le s
n o rm a l g ro w th
th e
te n ta c le s
fro m
th e
in d ic a te s
lo w e r b o d y
p ro cesses o f th e
c o n s id e re d
m a tu re
u n til
i t is
th e b ase
n e m a to c y s ts
re g io n s
a n im a l a n d
m ig r a t io n v ia th e g a s tr o v a s c u la r c a v ity .
not be
th a t
th ro u g h
not by
an
th e
a c tiv e
T h u s , a s te n o te le m a y
lo c a te d
in
th e
d is ta l
Ÿ
re g io n o f t h e te n ta c le s .
S ummary
A lth o u g h
b e fu lly
n e m a to c y s ts
d e v e lo p e d
im m a tu re
n o t fu lly
sin c e
in
th e
p o is o n
u n til
m a lly a n d d is ta lly fro m
an
o f c e lls i n
s u b s ta n c e
re g io n s o f t h e h y d r a
th a t th e re
be
w ith in
n e m a to c y s ts
of h y d ra
m ay
s till c o n s id e r e d
th e ir
re ac h
c a p s u le s
th e
is
te n ta c le
A s n e m a to c y s ts a re p u s h e d
p ro x i-
th is
r e g io n , th e y
becom e
m o re
T h e r e a p p e a rs to b e
p re se n t
g a s tric
in
th e
and
g ro w th
w h ic h in h ib it th e fu ll e la b o r a tio n
o f th e
O n t h e o t h e r h a n d , i t is
is a s u b s t a n c e , f o u n d
p e d u n c le b u t la c k in g in
a c id
m e ta c h ro m a s ia .
p o is o n w i t h in t h e n e m a t o c y s t c a p s u le .
p o s s ib le
re g io n
m ay
th e g r o w th r e g io n , b e c a u s e o f a r a p id
a n in c re a s e in
in h ib itiv e
g ro w th
c o n ta in e d
th e
o r p e d u n c le o f th e a n im a l.
m u ltip lic a tio n
th e
a n a to m ic a lly , th e y
e la b o ra te d
a n d sh o w
C o n c l u s io n .
a nd
in
th e
te n ta c le s a n d
th e g r o w t h a n d g a s tr ic re g io n s , w h ic h
is n e c e s s a r y t o c o m p l e t e t h e f o r m a t i o n o f t h e n e m a t o c y s t p o i s o n .
It
has
been
sh o w n
th a t
a
n e m a to c y s t,
lo c a te d
in
th e
g a s tric
re g io n o f a h y d r a , w h e n b r o u g h t u n d e r th e in f lu e n c e o f e ith e r
th e
p e d u n c le
o r te n ta c le re g io n s o f th e
a n im a l, b e c o m e s m o re
a c id w ith in a v e ry s h o r t tim e .
I t h a s b e e n c o n c l u d e d t h a t a m a t u r e n e m a t o c y s t is o n e t h a t
c o n ta in s
an
a c id
p o is o n
w ith in
its
c a p s u le .
to c y s t o c c u rs o n ly in th e te n ta c u la r a n d
Such
a
nem a­
p e d u n c u la r re g io n s o f
a h y d ra .
B IB L IO G R A P H Y .
BOUILLON, J., CASTIAUX, P. and VANDERMEERSSCHE, G. — Structure
subm icroscopique des cnidocils. Bull. micr. appl., 8. 61, 1958.
BRIEN, P. and RENIERS-DECOEN, M. — La croissance, la blastogénèse,
l ’ovogénèse chez Hydra fusca (Pallas). Bull. Biol, de la France e t de
la B elgique. 82, 293-386, 1949.
BURNETT, A. L. — H lstophysiology of growth In hydra. Journ. Exp. Zool.,
140, 281, 1959.
CHAPMAN, G. and TILNEY, L. — Cytological stu dies on the nem atocysts
of hydra. I. Desmonemes, Isorhizas, Cnidocils and Supporting Struc­
tures. Journ. Biophys. and. Biochm. C yt., 5, 69, 1959 a.
28o
SOCIÉTÉ ROYALE ZOOLOGIQUE DE BELGIQUE
CHAPMAN, G. and TILNEY, L. — Cytological stu dies on th e nem atocysts
of hydra. II. The Stenoteles. Journ. Biophys. and Biochem . C yt., 5. 79,
1959 b.
KEPNER, W. A., REYNOLDS, B. D., GOLDSTEIN, L. and TAYLOR, J. H. —
The structure, developm ent and discharge of th e penetrant of Pelm a­
toh ydra oligactis (Pallas). Journ. M orph., 72, 561-587, 1943.
SEMAL-VAN GANSEN, P. -— Le cnidosom e de l'Hydre e t le bouton urticant.
Bull. Acad. Roy. B elgique, 37, 650-664, 1951.
SEMAL-VAN GANSEN, P. — La structure des ném atocytes de l ’hydre d ’eau
douce. Bull. Acad. Roy. B elgique, 40, 269, 1954.
SLAUTTERBACK, D. and FAWCETT, D. — The developm ent of the cnldoblasts of Hydra. An electron m icroscopy stu dy of cell differentiation.
Journ. B iophys. an d Biochem . C yt., 5, 441-452, 1959.
WEILL, R. — C ontribution à l’étude des Cnldaires et de leurs ném atocystes. Lille : Trav. S ta t. Zool. W im ereux, 10-11, 1, 1934.

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