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Critical Review
MYSID CRUSTACEANS AS POTENTIAL TEST ORGANISMS FOR THE EVALUATION
OF ENVIRONMENTAL ENDOCRINE DISRUPTION: A REVIEW
T im A . V e r s l y c k e , * ! N a n c y F o c k e d e y , J C h a r l e s L . M c K e n n e y , J r .,§ S t e p h e n D . R o a s t ,[|
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a n d C o l in R . J a n s s e n i
■¡•Laboratory o f E n v iro n m e n tal T oxicology and A q u atic E cology, G h e n t U n iv ersity , J. P la te a u straa t 22, B -9000 G hent, B elgium
JD e p a rtm e n t o f B io lo g y , M arin e B iology S ection, G hent U n iv ersity , K rijg s k a n 2 8 1 -S 8, B -9000 G h en t, B elgium
§U .S. E n v iro n m en tal P ro tectio n A gency, N a tio n a l H ealth and E n v iro n m en tal E ffects R esearch L ab o rato ry , G u lf E co lo g y D ivision,
1 S ab in e Island D rive. G u lf B reeze, F lorida 32561 -5 2 9 9 , U S A
HSchool o f B io lo g ic al S cien ces, U n iv ersity o f P lym outh, D rake C ircus, D evon P L 4 BAA, U n ite d K ingdom
# F la n d e rs M arin e Institute, V ism ijn Pakhuizen 4 5 -5 2 , B -8400 O stend, B elgium
(R e c e iv e d 20 J u n e 2003; A c c e p te d 2 9 S e p te m b e r 2003)
A b stra ct— A n th ro p o g e n ic c h em ica ls that d isru p t the h o rm o n al system s (endocrine d isru p to rs) o f w ild life species re c en tly have
b eco m e a w id e ly in v e stig ate d and p o litic a lly ch arged issue. Inv erteb rates acco u n t fo r ro u g h ly 95 % o f all anim als, y e t surprisingly
little effo rt h as b een m ad e to u n d erstan d th eir v alu e in sig n alin g p o te n tia l en v iro n m en tal end o crin e d isru p tio n . T h is om issio n largely
can be attrib u ted to th e high d iv e rsity o f in v erteb rates a n d the shortage o f fu ndam ental kn o w led g e o f th e ir e n d o crin e system s.
In se cts and c ru stacean s a re e x ce p tio n s and, as su ch , ap p ea r to be e x ce lle n t c an d id ate s for ev alu atin g the e n v iro n m e n ta l consequences
o f ch em ically in d u ced e n d o crin e d isru p tio n . M y sid sh rim p (C rustacea: M y sid acea) m ay serv e as a v ia b le surrogate for m any
cru stacean s and h a v e b een p u t fo rw a rd as su itab le test organism s for the evalu atio n o f e n d o crin e d isruption by sev eral researchers
and re g u la to ry b o d ie s (e.g ., the U .S . E n v iro n m en tal P rotection A gency). D espite th e lo n g -sta n d in g u se o f m y sid s in to x icity testing,
little in fo rm atio n ex ists on th eir e n d o crin o lo g y , and fe w studies have fo cu sed on the p o te n tia l o f th ese an im als for ev alu atin g the
effe c ts o f h o rm o n e -d isru p tin g c o m p o u n d s. T h erefo re, the question rem ains as to w h e th e r the c u rre n t stan d a rd ize d m ysid endpoints
can be u se d o r ad ap ted to d e te ct en d o crin e d isru p tio n , or if new pro ced u res m u s t be d eveloped, sp ecifically d ire c te d a t evaluating
h o rm o n e-reg u lated en d p o in ts in th ese anim als. T h is re v ie w sum m arizes the e cological im portance o f m ysids in e stu a rin c a n d m arine
eco sy stem s, th e ir u se in to x ic ity testin g and env iro n m ental m o nitoring, and th e ir e n d o crin o lo g y and im p o rtan t horm o n e-reg u lated
p ro cesses to h ig h lig h t th eir p o te n tia l u se in a sse ssin g environm ental end o crin e disruption.
K e y w o rd s — M y sid s
E n d o c rin e d isru p tio n
B io m ark ers
R eg u latio n
E n v iro n m en tal m onito rin g
c i p ie n ts o f e n d o c r in e d i s r u p to r s in th e ir s e a w a r d tr a n s p o r t. O f
IN T R O D U C T IO N
th e e s tu a r i n e o r g a n is m s t h a t c o u ld b e a d v e r s e ly a f f e c te d b y
A n th r o p o g e n ic c h e m ic a ls t h a t d i s r u p t th e h o r m o n a l s y s te m s
th e s e c o m p o u n d s , c r u s ta c e a n s a r e g o o d c a n d i d a te s f o r th e
( e n d o c r in e d is r u p to r s ) o f w i ld lif e s p e c ie s r e c e n t ly h a v e b e ­
s tu d y o f p o te n tia l im p a c ts . C r u s ta c e a n s a r e c o m m o n in f r e s h ­
c o m e a w id e ly in v e s t ig a te d a n d p o litic a lly c h a r g e d is s u e [ 1 -
w a te r , e s tu a r i e s , a n d s h a ll o w c o a s ta l w a te r s a n d f o r m v ita l lin k s
3 ], I n v e r te b r a te s a c c o u n t f o r r o u g h ly 9 5 % o f a ll a n im a ls [4 ],
in a q u a t ic f o o d w e b s [ 1 6 - 2 1 ] . In a d d itio n , c r u s ta c e a n s a re
y e t s u r p r is in g ly littl e e f f o r t h a s b e e n in v e s t e d to u n d e r s ta n d
s u s c e p tib le to th e e f f e c ts o f e n d o c r in e d i s r u p to r s [1 3 ] . A n in ­
th e ir v a lu e in s i g n a l in g p o te n tia l e n v ir o n m e n ta l e n d o c r in e d is ­
te r n a tio n a l S o c ie ty o f E n v i r o n m e n ta l T o x ic o lo g y a n d C h e m ­
r u p ti o n [ 5 - 1 2 ] , A lth o u g h g r o w th , r e p r o d u c tio n , d e v e l o p m e n t,
i s tr y w o r k s h o p o n e n d o c r in e d is r u p tio n in in v e r te b r a te s h e ld
a n d o th e r a s p e c ts o f i n v e r te b r a te p h y s io lo g y a r e k n o w n to b e
in T h e N e th e r la n d s in 1 9 9 8 [7 ] id e n tif ie d in s e c ts a n d c r u s ta ­
u n d e r h o r m o n a l c o n tr o l, t h e e n d o c r in e s y s te m s a n d h o r m o n e s
c e a n s a s p o te n tia l o r g a n is m s f o r e v a l u a tin g c h e m ic a lly in d u c e d
p r o d u c e d a n d u s e d in in v e r te b r a te s a r e n o t d ir e c tly a n a lo g o u s
e n d o c r in e d is r u p tio n b y v ir t u e o f th e w e a lt h o f in f o r m a tio n
to th o s e o f v e r te b r a te s [1 3 ] . In in v e r te b r a te s , th e s e le c tio n o f
s u ita b le te s t m e th o d s a n d s p e c ie s f o r e v a l u a tin g e n d o c r in e d is ­
a v a i la b le o n th e ir e n d o c r in o lo g y c o m p a r e d w ith o th e r in v e r­
r u p ti o n is c o n f o u n d e d b y d iv e r s ity . T h e u s e o f a lim ite d n u m b e r
t e b r a te s [ 9 ,1 2 ,2 2 - 2 4 ] .
o f s p e c ie s a s r e p r e s e n ta tiv e o f th is d i v e r s i t y is a n a iv e a p p r o a c h
O f th e c r u s ta c e a n s , m y s id s h r im p h a v e b e e n p u t fo r w a r d
d e s ti n e d to f a ilu r e in th e a b s e n c e o f s u ita b le s a f e g u a r d s [2 ,9 ].
a s s u ita b le te s t o r g a n is m s f o r th e e v a l u a tio n o f e n d o c r in e d is ­
H e n c e , th e k e y c h a l le n g e f o r e n v ir o n m e n ta l a s s e s s m e n t is to
ru p tio n [ 7 ,9 ,2 5 ] . T h e U .S . E n v i r o n m e n ta l P r o te c tio n A g e n c y
fin d in v e r te b r a te s p e c ie s , s e le c te d f r o m m u ltip l e le v e ls o f e c o ­
e s ta b lis h e d th e E n d o c r i n e D i s r u p to r S c r e e n in g a n d T e s tin g
s y s te m f u n c tio n , to e f f ic ie n tly m o n ito r a n d e v a l u a te th e c o m ­
S ta n d a r d iz a tio n a n d V a lid a tio n T a s k F o r c e to c o o r d in a te a n d
p l e x i t y o f p o te n tia l e n v ir o n m e n ta l e f f e c ts o f e n d o c r in e - d is ­
c o n d u c t th e s c ie n tif ic a n d te c h n ic a l w o r k n e c e s s a r y to v a lid a te
th e s c r e e n s a n d te s ts r e c o m m e n d e d b y th e E n d o c r i n e D is r u p to r
r u p ti n g c h e m ic a ls a t a r e a s o n a b le f in a n c ia l c o s t [1 4 ].
M a n y a n th r o p o g e n ic p o llu ta n ts h a v e th e w o r l d ’s o c e a n s a n d
S c r e e n in g a n d T e s tin g C o m m itte e . T h e S ta n d a r d iz a tio n a n d
s e a s a s a fin a l s in k , a n d a r e c a r r ie d th e r e th r o u g h r iv e r i n e a n d
V a lid a tio n T a s k F o r c e r e c o m m e n d e d a tw o - ti e r e d a p p r o a c h fo r
e s tu a r i n e c o n d u its [ 1 2 ,1 5 ] , E s tu a r ie s a r e i n t r in s ic a lly a n d c o m ­
d e te r m in in g w h e t h e r a c h e m ic a l is a n e n d o c r in e d is ru p to r, a n d
m y s id s w e r e p r o p o s e d a s a s u ita b le i n v e r te b r a te a s s a y in th e
m e r c ia lly im p o r ta n t e c o s y s te m s a n d a r e a m o n g s t th e f ir s t r e ­
t i e r 2 te s t in g (in v iv o te s t in g ) ( h ttp :// w w w .e p a .g o v /s c ip o ly /
o s c p e n d o ) f o r a tw o - g e n e r a tio n r e p r o d u c tiv e a n d d e v e l o p ­
* To w h o m c o rre sp o n d en c e m ay be a d d ressed
(tirri.verslycke(g)ugent.be).
m e n ta l to x ic ity te s t. R e c e n tly , a d r a f t r e v ie w p a p e r w a s c o m ­
1219
1220
E n viro n . T oxicol. Cheni. 2 3, 2004
T.A . V erslycke e t al.
p i l e d o n m y s id lif e - c y c le t o x ic ity t e s t in g [1 3 ] , a n d th e tw o -
b e n th ic (c m ) p r e y ite m s , a n d m y s id s o f te n p r o g r e s s i v e ly r e ­
g e n e r a t io n m y s id lif e - c y c le a s s a y w a s p r o p o s e d to th e O r g a ­
p la c e c o p e p o d s in th e d ie t o f m a n y p o s t l a r v a l a n d j u v e n ile
n iz a tio n f o r E c o n o m ic C o o p e r a t io n a n d D e v e lo p m e n t a s a n e w
c o m m e r c ia l fish s p e c ie s [ 1 9 ,2 6 ,4 4 ,4 5 ] , In a d d itio n , m y s id s m a y
O r g a n iz a t io n f o r E c o n o m ic C o o p e r a t io n a n d D e v e lo p m e n t te s t
s e r v e a s p r e y f o r l a r g e r c r u s ta c e a n s , m a r in e m a m m a ls , o r w a d ­
g u id e lin e . D e s p ite th e l o n g - s ta n d in g u s e o f m y s id s in to x ic ity
in g b i r d s [ 2 6 , 2 7 ,3 2 , 4 6 - 4 8 ] .
te s t in g , littl e in f o r m a tio n o n th e ir e n d o c r in o lo g y h a s b e e n p u b ­
E s tu a r in e m y s id s h a v e a f le x ib le p h y s io lo g y th a t r e s p o n d s
lis h e d a n d f e w s tu d ie s h a v e f o c u s e d o n th e p o te n tia l o f th e s e
to a h o s t o f d y n a m ic a ll y c h a n g i n g e n v ir o n m e n ta l v a r ia b le s ,
a n im a ls f o r e v a l u a tin g th e e f f e c ts o f h o r m o n e - d i s r u p t in g c o m ­
c h a r a c te r is tic o f th e c o m p le x c h e m is tr y o f e s tu a r ie s . T e m p e r­
p o u n d s . T h e r e f o r e , th e q u e s ti o n r e m a in s a s to w h e t h e r th e
a tu r e a n d s a li n ity a r e th e d o m in a n t e c o lo g ic a l v a r ia b le s , a n d
m a y a c t e ith e r s in g ly o r in c o m b in a tio n to m o d if y th e p h y s ­
c u r r e n t s t a n d a r d i z e d m y s id e n d p o in ts c a n b e u s e d o r a d a p te d
to d e te c t e n d o c r in e d is r u p tio n , o r i f n e w p r o c e d u r e s m u s t b e
io lo g ic a l a n d e c o l o g ic a l p r o p e r ti e s o f e s tu a r i n e o r g a n is m s as
d e v e l o p e d , s p e c if ic a lly d ir e c te d a t e v a l u a tin g h o r m o n e - r e g u ­
w e ll a s r e s p o n s e s to x e n o b i o t i c e x p o s u r e . T h e r e f o r e , e m p iric a l
la te d e n d p o in ts in th e s e a n im a ls .
d e te r m in a tio n o f th e o p tim a l s a li n ity a n d te m p e r a tu r e c o n d i­
T h is p r e s e n t r e v ie w p r o v id e s a n o v e r v ie w o f th e a v a ila b le
tio n s o f e s tu a r i n e m y s id s is e s s e n tia l f o r th e d e v e l o p m e n t o f
in f o r m a tio n o n m y s id s r e le v a n t to th e is s u e o f e n d o c r in e d i s ­
o p tim u m la b o r a to r y c u ltu r e o f th e s e o r g a n is m s a n d th e ir u s e
r u p tio n , in c lu d in g th e ir e c o lo g ic a l r o l e in m a r in e a n d e s tu a r in e
in t o x ic ity a n d h a z a r d a s s e s s m e n t. F o r e x a m p le , th e o p tim a l
e c o s y s te m s , th e ir u s e in t o x ic ity te s t in g a n d e n v ir o n m e n ta l
s a lin ity a n d t e m p e r a tu r e c o n d itio n s f o r g r o w th o f A m e r i c a ­
m o n ito r in g , a n d th e ir e n d o c r in o lo g y . A c a s e is m a d e f o r th e ir
m y s i s b a h ia ( f o r m e r ly M y s i d o p s i s b a h i a ) t h r o u g h its e n tire
p o te n tia l u s e in a s s e s s in g th e e n v ir o n m e n ta l c o n s e q u e n c e s o f
lif e c y c le [4 9 ] a r e c o r r e la te d w ith r e s i s t a n c e p a tte r n s to th e s e
e n d o c r in e - d is r u p tin g c h e m ic a ls .
d o m i n a n t e n v ir o n m e n ta l v a r ia b le s [5 0 ] a n d d i s tr ib u tio n o f th is
s p e c ie s in e s tu a r ie s . M o r e o v e r , te m p e r a tu r e a n d s a li n ity in te r­
M Y S ID B IO L O G Y A N D E C O L O G Y
a c t to m o d if y th e r e p r o d u c t i v e c a p a c ity o f th i s s p e c ie s [5 1 ].
M y s id s ( M a la c o s tr a c a : P e r a c a r id a : M y s id a c e a ) a r e r e la ­
M Y S ID S A N D T O X IC O L O G Y
t iv e ly s m a ll ( w ith th e m a jo r ity o f th e s p e c ie s b e in g b e tw e e n
5 a n d 2 5 m m in le n g th ) , s h r im p lik e c r u s ta c e a n s , o fte n r e f e r r e d
to a s o p o s s u m s h r im p b e c a u s e th e o o s te g i te s f o r m a v e n tr a l
f e m a le m a r s u p iu m f o r c a r r y i n g th e d e v e l o p in g e m b r y o s . T h e
l a tte r f e a tu r e d is tin g u is h e s m y s id s f r o m o th e r s h r im p lik e c r u s ­
t a c e a n s . M y s id s a r e id e n tif ie d f r o m o th e r p e r a c a r i d s ( A m p h i­
p o d a , I s o p o d a , C u m a c e a , a n d T a n a id a c e a ) b y th e p r e s e n c e o f
a s ta t o c y s t ( c o n ta in in g la r g e e n d o g e n o u s s ta t o lith s , th e p r im a r y
e q u ilib r iu m o r g a n s fo r m y s id s ) o n th e p r o x im a l p a r t o f th e
u r o p o d a ! e n d o p o d . M y s id s a r e d i s tr ib u te d fr o m 8 0 ° N to 8 0 °S
a n d o c c u r in v a r io u s a q u a t ic e n v ir o n m e n ts , in c lu d in g f r e s h ­
w a te r , g r o u n d w a te r , b r a c k is h , e s tu a r i n e , c o a s ta l, a n d o c e a n ic
h a b ita ts [ 2 6 - 2 8 ] . M a u c h lin e a n d M u r a n o [2 8 ] p u b lis h e d a
w o r ld lis t o f m y s id s in 1 9 7 7 (7 6 5 s p e c ie s d is tr ib u te d b e tw e e n
—
1 2 0 g e n e r a ) ; h o w e v e r , th is n u m b e r is e v e r in c r e a s in g
M y s id s a r e s e n s itiv e to s o m e c h e m ic a l c o n ta m in a n t s a t e n ­
v ir o n m e n ta lly r e l e v a n t c o n c e n tr a tio n s a n d h a v e b e e n u s e d in
r e g u la to r y to x ic ity t e s t in g f o r m o r e th a n 2 0 y e a r s [ 1 5 , 4 3 ,5 2 6 6 ]. T h e U .S . E n v i r o n m e n ta l P r o te c tio n A g e n c y a n d th e A m e r ­
ic a n S o c ie ty f o r T e s tin g a n d M a te r i a ls b o th h a v e a d o p te d th e
s u b tr o p ic a l A . b a h ia a s a k e y t e s t in g s p e c ie s f o r c o a s ta l a n d
e s tu a r i n e m o n ito r in g , a n d s t a n d a r d g u id e s f o r c o n d u c t in g lif e ­
c y c le to x ic ity te s ts w ith th is s p e c ie s h a v e b e e n d e v e lo p e d
[ 1 3 ,6 7 - 7 2 ] , A lth o u g h a r e l a t i v e l y la r g e a m o u n t o f p u b lis h e d
to x ic ity d a ta is a v a i la b le f o r A m e r i c a m y s i s s p e c ie s , r e la tiv e ly
lim ite d d a ta a r e a v a i la b le o n th e s e n s itiv ity o f o th e r m y s id
s p e c ie s to to x ic a n ts [6 4 ] . H o w e v e r , th e a v a i la b le e v id e n c e s u g ­
g e s ts th a t m y s id s a r e g e n e r a l ly m o r e s e n s i t i v e to to x ic s u b ­
th r o u g h im p r o v e d s a m p l in g te c h n iq u e s a n d e x p lo r a tio n o f n e w
s ta n c e s th a n m a n y o th e r te s t s p e c ie s [ 4 3 ,7 3 - 7 5 ] , T o x ic ity te s t
h a b ita ts . T h e p r e s e n t c o u n t is m o r e th a n
p ro c e d u re s
1 ,0 0 0 s p e c ie s b e ­
have
been
p u b lis h e d
fo r
N e o m y s is
m e r c e d is
l o n g in g to a p p r o x im a te ly 1 6 0 g e n e r a (h t t p :// c r u s ta c e a .n e t/ ). A
[ 5 2 ,7 6 ] , M y s i d o p s i s in t i i [ 5 9 ] , H o l m e s i m y s i s c o s t a t a [6 2 ],
c o m p r e h e n s iv e d a ta b a s e o n th e w o r l d m y s id f a u n a ( N e m y s ,
h ttp :// in tr a m a r .u g e n t.b e / n e m y s ) , c o n ta in in g lin k s to r e le v a n t
A m e r i c a m y s i s b i g e l o w i [7 7 ] , N e o m y s i s i n t e g e r [ 6 4 ,7 5 ,7 8 ] ,
in f o r m a tio n ( i.e ., ta x o n o m ic a l , m o r p h o l o g ic a l, e c o l o g ic a l, b io ­
[ 8 0 ,8 1 ] , N e o m y s is a m e r i c a n a [ 8 2 ,8 3 ] , a n d N e o m y s i s a w a t­
g e o g r a p h ic , lite r a tu r e , p ic to r ia l, a n d m o le c u la r in f o r m a tio n ) ,
s c h e n s i s [8 4 ] (T a b le 1). I n a d d itio n , m e th o d s f o r m a in ta in in g
o n th e s p e c ie s le v e l p r e s e n tl y is b e in g c o n s tr u c t e d (T. D e p r e z ,
v ia b le p o p u la tio n s o f d i f f e r e n t m y s i d s p e c ie s u n d e r la b o r a to r y
G h e n t U n iv e r s ity , S e c tio n M a r i n e B io lo g y , G h e n t, B e lg iu m ,
c o n d itio n s
p e r s o n a l c o m m u n ic a t io n ) .
[ 4 6 , 5 9 ,7 5 ,7 9 ,8 5 - 8 9 ] . R e c e n tly , a s tr o n g c o r r e la tio n w a s r e ­
In g e n e r a l, m y s id s a r e r e g a r d e d a s o m n iv o r e s a n d f e e d o n
T e n a g o m y s is
n o v a e - z e a la n d ia e
have
been
d e s c rib e d
[7 9 ], P r a u n u s fle x u o s u s
by
se v e ra l re se a rc h e rs
p o r te d b e tw e e n th e to x ic r e s p o n s e o f d a p h n id s a n d m y s id s (R 2
p h y to p la n k to n , Z o o p la n k to n , a n d o r g a n ic d e tr itu s [ 2 6 , 2 7 ,2 9 -
= 0 .9 4 1 , n = 2 8 ; 9 6 - h m e d ia n le th a l c o n c e n tr a tio n s f o r A .
3 1 ]. P e la g ic f o r m s f il te r p a r tic le s d u r in g s w i m m in g , w h e r e a s
b a h i a a n d 4 8 - h m e d ia n le th a l c o n c e n tr a tio n s f o r D a p h n i a m a g ­
b e n th ic s p e c ie s h a v e b e e n o b s e r v e d a c t iv e ly h u n tin g a n d g r a b ­
n a ) f o r p e s ti c id e s a n d o r g a n ic s , e m p h a s iz in g th e u s e o f m y s id s
b in g s m a ll p a r tic le s [2 7 ] , M y s id s fo r m i m p o r ta n t lin k s in th e
in f u tu r e t o x ic ity te s t in g [9 0 ].
f o o d w e b s o f a q u a t ic e c o s y s te m s a n d o fte n f e e d s e le c tiv e ly
M y s id s h a v e b e e n u s e d s u c c e s s f u lly to m e a s u r e v a r io u s
f o r s iz e o r s p e c ie s ( o r b o th ) o f p r e y [ 2 6 ,3 2 ], C o n s e q u e n tly ,
s u b le t h a l to x ic a n t e f f e c ts , s u c h a s g r o w th , s w i m m in g c a p a ­
th e y h a v e th e p o te n tia l f o r s tr u c tu r in g Z o o p la n k to n c o m m u ­
b ility , f e e d in g b e h a v io r, m o ltin g , e n e r g y b u d g e t, r e p r o d u c tio n ,
n itie s [ 3 3 ,3 4 ] a n d in f l u e n c in g th e s t r u c tu r e o f p h y to p la n k to n ,
s e x u a l m a tu r ity , a n d v i t e llo g e n e s is ( d e s c r ib e d in d e ta il in th e
ty c h o p la n k to n , a n d m e io f a u n a l c o m m u n itie s [ 3 2 , 3 5 - 4 1 ] . M o s t
m y s id s u tiliz e o r g a n ic d e tr itu s to a c o n s id e r a b le e x te n t a n d a re
f o ll o w i n g p a r a g r a p h s a n d s u m m a r iz e d in T a b le 2 ). A ls o , fie ld
c a p a b le o f r e m in e r a liz in g a s u b s ta n tia l p o r ti o n o f th e n o n r e ­
l is h e d [ 1 7 ,9 1 - 9 4 ] .
s tu d ie s a n d c a g in g e x p e r im e n ts w ith m y s id s h a v e b e e n p u b ­
f r a c to r y d e tr itu s s u s p e n d e d in t h e w a t e r c o lu m n o r b u r ie d in
B e c a u s e o f th e ir e c o l o g ic a l im p o r ta n c e , w id e g e o g r a p h ic
th e s u r f a c e s e d im e n ts [ 2 9 ,3 7 ,4 2 ,4 3 ] . M y s id s iz e is in te r m e d ia te
d is tr ib u tio n , y e a r - r o u n d a v a i la b il ity in th e fie ld , e a s e o f tra n s ­
b e tw e e n m e s o z o o p la n k to n ic ( p m ) a n d e n d o b e n t h is o r e p i-
p o r ta tio n , a b ility to b e c u ltu r e d in th e la b o r a to r y , a n d s e n s i­
E nviron. T oxicol. Cheni. 23, 2004
M y sid s and e n d o crin e d isru p tio n : A re v ie w
1221
Table 1. C an d id ate m y sid te st sp ecies fo r to x ic ity te stin g w ith details on th e ir na tu ra l h a b ita t an d cu ltu rin g
S p ecies nam e
D istrib u tio n
A m e ric a m y sis bah ia
( = M y sid o p sis bahia)
A m e ric a m y s is b i g e lo w i
( = M y sid o p sis b ig elo w i)
A m e rica m y sis a lm yra
( = M y sid o p sis a lm yra )
H o lm e sim y sis c o sta ta
( = A c a n th o m y sis scu lp ta )
M y sid o p sis in tii
M ysis m ixta
N eo m y sis a w a tsc h e n sis
N e o m y sis m e rc e d is
N eo m y sis in teg er
P ra u n u s fle x u o su s
C o astal estu aries an d em baym e n ts ran g in g fro m th e G u lf
o f M ex ico to N arrag an sett
(R I, U S A ) [250]
E a ste rn c o a st o f th e U S A from
M A (G eo rg es B ank) to FL
often to g e th e r w ith A. b ahia
[250]
E a ste rn co ast o f the U SA , in ­
sh o re w a ters alo n g the entire
co ast o f G u lf o f M ex ico and
n o rth w ard a lo n g A tlantic
c o a st to P ata p sc o R iv e r
(M D ) [250]
P rin cip al sp ecies o f th e genus,
fro m so u th ern C alifo rn ia
(U SA ) to B ritish C olum bia
(C an ad a) [122, 253]
E astern P acific fro m South
A m e ric a to the so u th ern C al­
ifo rn ia c o a st o f th e U S A [59,
250]
E astern (fro m W h ite Sea to Ic e ­
land) and W estern (G reen­
land coastal w aters do w n to
C ape C o d ) A tla n tic regions
[26]
P acific co ast o f Jap an , K orea,
and U S A [115]
N o rth e a ste rn Pacific co ast o f
th e U S A (so u th ern A la sk a to
G o v io ta Bay, C A ) [52]
H a b ita t d e scription
C o m m ercial culture
C ulture
protocol
M arine (> 1 5 % 0), < 2 0 - 3 4 DC
[4 9 -5 1 ]
Yes
[46,89,251]
M arine (30-35% «), 2 -3 0 °C
No
[46]
M arin e (10-20% o), > 2 0 °C
Yes
[85,88,252]
M arine, planktonic, lives w ith in
su rfa c e can o p y o f kelp
N o, fie ld -co lle c te d anim als
a v ailab le [253]
[122,253]
M arine, e p ib e n th ic, optim al
tem perature 2 0 -2 2 °C , o p ti­
m al salin ity 28-35% «
No
[59]
B rackish, low salinity, c o ld w a ­ N o
ter
[114]
M arine, estuarine
No
[115]
No
[52]
No
[75]
No
[123]
F reshw ater, estuaries, and
coastal lakes, p la n k tonic/epibenthic, e u ry h a lin e (< 0 .5 to
>25% «), 6-22*C
N o rth e rn E u ro p ean estuaries
M arin e, e stu a rin e, freshw ater,
an d c o asta l w aters; o lig o h a ­
h y p erb en th ic, e u ry h a lin e
(< 0 .5 to >25% «), cold w a ter
lin a and fresh w ater lakes [20,
(< 2 0 °C ) [116]
43]
N o rth ern E uropean co astal w a ­ H y p e rb e n th ic /p lan k to n ic, e u ry ­
haline, eu ry th e rm [254, 255]
ters
tiv ity to c o n ta m in a n t s , m y s id s a r e a p p r o p r ia te t o x ic ity te s t o r­
e ra l s p e c ie s (A . b a h ia , A m e r i c a m y s i s a l m y r a , A . b ig e lo w i, IT.
g a n is m s .
c o s ta t a , M . i n tii, N . m e r c e d i s , a n d N . i n t e g e r ) a re c o n s id e r e d
f o r th e ir p o te n tia l u tilit y in e n d o c r in e - d is r u p tio n te s tin g [1 3 ].
C A N D ID A T E M Y S ID T E S T S P E C IE S F O R E N D O C R IN E -
F r o m th is re v ie w , i t m a y b e c o n c l u d e d th a t, a l th o u g h A . b a h ia
D IS R U P T IO N R E SE A R C H
h a s m a n y s tr e n g th s , l im it e d e c o l o g ic a l r e le v a n c e f o r h ig h - la t-
G e n e r a l s e le c tio n c r ite r ia f o r th e m o s t a p p r o p r ia te m y s id
itu d e a n d lo w - s a lin e s y s te m s p r e c lu d e its g e n e r a l u tilit y . H o w ­
s p e c ie s f o r to x ic o lo g ic a l te s t in g a r e g iv e n b y N im m o a n d H a ­
e v e r, g iv e n th e h ig h d e g r e e o f s t a n d a r d i z a tio n in A . b a h ia ,
m a k e r [1 5 ] a n d R o a s t e t a l. [4 3 ] . T h e s e c r ite r ia in c lu d e a v a i l­
p r o g r e s s in d e v e l o p m e n t o f s t a n d a r d i z e d t e s t p r o to c o ls f o r e n ­
a b le w h e n r e q u ir e d ; a lr e a d y a d a p t e d to l a b o r a to r y c o n d itio n s ,
d o c r in e - d is r u p tio n t e s t in g s h o u ld b e f a s te s t in th is s p e c ie s .
e lim in a tin g a n ( e x p e n s iv e ) c o n d itio n in g p h a s e ; c o lle c tio n f o r
T a b le 1 s u m m a r iz e s th e d is tr ib u tio n , h a b i t a t d e s c r ip t io n , a n d
l a b o r a t o r y te s t in g w ill n o t d e c i m a te f ie ld p o p u la tio n s ( o r d e ­
a v a i la b le c u ltu r e p r o to c o ls f o r o th e r c a n d i d a te m y s id te s t s p e ­
s tr o y h a b ita t d u r in g c o lle c ti o n ) ; e a s ily tr a n s p o r te d ; lif e h is to r y
c ie s.
is s h o r t, m a k in g i t p o s s ib le to s tu d y th e e f f e c ts o f a p o llu ta n t
o n v a r io u s a s p e c ts o f r e p r o d u c tio n ; d ie t is k n o w n a n d r e a d ily
c o n tr o lle d ; a n d e c o l o g ic a lly im p o r ta n t. In a d d itio n , th e im ­
M Y S ID E N D O C R IN O L O G Y A N D H O R M O N E -R E G U L A T E D
E N D P O IN T S
p o r ta n t c h a r a c te r is tic s fo r th e s e le c tio n o f a s u ita b le t e s t s p e c ie s
T h e u s e o f h o r m o n e s to r e g u la te b io lo g ic a l p r o c e s s e s is a
fo r id e n tif y in g th e e f f e c ts o f e n d o c r in e d is r u p tio n in th e e n ­
s tr a te g y c o m m o n to v e r te b r a te s a n d in v e r te b r a te s . A lth o u g h
v ir o n m e n t a r e g iv e n b y D e F u r e t a l. [7 ] a n d in c lu d e p r im a r y
t h e e n d o c r in e s y s te m s o f in v e r te b r a te s r e g u la te m a n y o f th e
m o d e o f r e p r o d u c tio n , c u ltu r e in th e la b o r a to r y , g e n e r a t io n
s a m e p r o c e s s e s in v e r te b r a te s ( d e v e lo p m e n t, g r o w th , a n d r e ­
tim e , s iz e , k n o w le d g e o f e n d o c r in o lo g y , a n d s ta n d a r d m e th o d s
p r o d u c tio n ) , s o m e e n d o c r in e - r e g u la te d p r o c e s s e s a r e u n iq u e
a v a ila b le . S o m e a ttr ib u te s d e s c r ib e d in th e la tte r p u b lic a tio n
to s p e c if ic g r o u p s o f in v e r te b r a te s . F o r e x a m p le , m o ltin g , d ia ­
(e .g ., m o d e o f r e p r o d u c tio n o r k n o w le d g e o f e n d o c r in o lo g y )
p a u s e , a n d l im b r e g e n e r a t io n
d o n o t a llo w f o r d is c r im in a tio n a m o n g c a n d i d a te m y s id s p e ­
c e s s e s a s s o c ia t e d w ith s o m e i n v e r te b r a te g r o u p s th a t a r e r a r e
c ie s . A v e r y u s e f u l d o c u m e n t in th is c o n te x t is a d r a f t r e v ie w
o r a b s e n t a m o n g v e r te b r a te s [7].
p a p e r o n lif e - c y c le to x i c i t y te s t in g w ith m y s id s in w h ic h s e v ­
a r e e n d o c r in e - r e g u la te d p r o ­
M o s t o f th e c u r r e n t k n o w le d g e o f c r u s ta c e a n e n d o c r in o lo g y
E n d p o in t
U se in m ysids
R eferen c e s/co m m e n ts
[69,253]
[69]
[69]
[62,67,253]
[52,67,76]
[55,59]
[53,64,75,78]
[7 9 ,8 0 -8 4 ]
L ife -c y cle testin g
A. b a h ia (S)
A. b ig e lo w i (S ), A . alm yra (S)
H. c o sta ta (S )
M. in tii
Ar. in te g e r
[68,70,72,253]
[68]
[56,68,71]
[55,59]
[160]
T w o -g en eratio n testin g
A. b a hia, A. bigelow i, A. alm yra (S in prep )
M. in tii
H. co stata, N. m ercedis, N. integer
[13,219,256]
M eth o d sh ould be developed
P ro b ab ly im p rac tic a l because o f lo n g g e n eratio n tim e
F e cu n d ity (b ro o d size)
A. b a hia, A . bigelow i, A. alm yra (S)
IT. c o sta ta (S )
N . m e rc e d is
M . in tii
N. in teger, P ra u n u s fle x u o su s
M ysis m ix ta
N e o m y s is a w a tsc h e n sis
[68,70]
[71,122]
[257]
[55,59]
[123,258]
[114]
[115]
E m b ry o n ic d ev elo p m en t
A. bahia
M e so p o d o p sis s la b b eri
[205]
[208]
Sexual m atu rity
A. b a h ia (S )
H. c o sta ta
[5 1 ,58,60,68,218]
[122]
E gg d ev elo p m en t lim e
N. integer, P. fle x u o su s
M. m ix ta
M. in tii
[123]
[114]
[55,59]
Sex ratio an d in tersex u ality
A. b a h ia (S)
N. in te g e r
G ro w th , b iom ass
A. bahia, A. bigelow i, A. a lm y ra (S)
H. c o sta ta (S)
N. m e rc e d is
M. in tii
M. m ix ta
N. in teg er
N. a w a tsc h e n sis
P. fle x u o s u s
T e n a g o m y sis no v a e-zealandiac
O th er species
A. b a h ia
N. in teg er
M. m ix ta
S ir ie lla a rm a ta
N. a w a tsc h e n sis
A. bahia
[68,219,253]
[2 2 3 -2 2 5 ]
[68,70]
[56,71,122]
[52]
[55,59]
[124]
[107,116]
[84,115,150,151]
[114]
[79]
[1 1 7 ,1 1 9 -1 2 1 ,1 3 2 ]
M o lt tim e a n d su ccess
E n ergy m etab o lism
0 :N ra tio , C :N ratio
R esp iratio n
N. m e rc e d is
N. in teg er
[259]
[111,140]
[140]
[138]
[115]
[63]
[260]
[1 5 3 ,1 5 5 ,1 6 4 -1 6 7 ,1 6 9 -1 7 5 ,1 7 8 -1 8 1 ]
T.A. Verslycke et al.
A m e ric a m y sis b a h ia (S )a
A m e r ic a m y s is b ig e lo w i (S)
A m e ric a m y sis a lm y ra (S)
H o lm e sim y sis c o sta ta (S)
N eo m y sis m e rc e d is (S)
M y sid o p sis in tii
N e o m y sis in te g e r
O th e r species
Environ. Toxicol. Chem. 23, 2004
S u rv iv al (acute)
T im e to first b ro o d rele a se
1222
Table 2. L ist o f p otential endpoints for e v a lu a tin g th e e ffe c ts o f e n docrine d isruptors and th e ir use in m ysids
E nd p o in t
U se in m ysids
R eferen c e s/co m m e n ts
B io ch em ical co m p o sitio n
M. m ix ta
N. a w a tsc h e n sis
P. fle x u o su s
L e p to m y sis lingvura
M y sis relicta
G a stro sa cc u s brevifissura
[114,182]
[1 5 0 -1 5 2 ]
[81]
[54]
[176,261]
[163]
E c d y stero id m etabolism
A. bahia
N. in teg er
S. a rm ata
S.R . T ubertyb a n d C .L. M cK enney,c unpu b lish ed data
A. G b e k iere e t al.,d unp u b lish ed data
[139]
Steroid m etabolism
N. in teg er
[162,185,198,199]
P 450 enzym es
N. in teg er
[162,185,198]
V itello g en esis
A. bahia
S. a rm a ta
N. in teg er
[215]
[139]
A. G h e k iere e t a l.,J unp u b lish ed data
O sm o reg u latio n
A. bahia
N. in teg er
P. fle x u o su s
O th e r species
[259,262,263]
[264]
[265]
M o rp hology, h istology
A. bahia, A. b igelow i
N. in teg er
1248]
[77]
[225]
N. in teg er
[4 3 ,2 2 6 -2 3 0 ,2 3 2 ]
A. b a h ia
M. m ixta
[231,241]
[239,242]
O th er beh av io ral en d p o in ts
M atin g , groom ing, sw arm ing, b u rro w in g ability, p re d a to r-p rc y dynam ics
[2 3 3 -2 3 8 ,2 4 3 -2 4 7 ]
a S = m eth o d is stan d ard ized .
b C en ter fo r E n v iro n m en tal D ia g n o stics and B io rem ediation, U n iv e rsity o f W est F lo rid a , P en saco la, F L , U SA .
CU .S . E n v iro n m en tal P ro te c tio n A gency, N a tio n a l H ealth an d E n v iro n m en tal E ffects R esearch L ab o rato ry , G u lf B reeze, F L , U SA .
d L a b o ra to ry o f E n v iro n m e n tal T o x ico lo g y an d A q u a tic E cology, G hent U n iv ersity , G hent, B elgium .
Environ. Toxicol. Cheni. 23, 2004
Sw im m in g behavior
F e e d in g b eh av io r
Mysids and endocrine disruption: A review
Table 2. C o n tin u e d
1223
1224
E n viro n . T oxicol. Cheni. 2 3, 200 4
T.A . V e rsly ck e e t al.
is b a s e d u p o n s tu d ie s w ith d e c a p o d s s u c h a s c ra b s , lo b s te r s ,
e f f e c ts o n m a n y a s p e c ts o f o r g a n is m a l f u n c tio n , in c lu d in g
c ra y fis h , a n d s h r im p , a n d h a s b e e n r e v ie w e d p r e v i o u s l y [ 7 , 9 5 -
p h y s io lo g y , b e h a v io r, a n d c h a n g e s in b io c h e m ic a l c o m p o s itio n
1 0 0 ]. T h e m a in b io lo g ic a l p r o c e s s e s , s u c h a s g r o w th , m o ltin g ,
[ 1 3 4 .1 3 5 ] , M o ltin g is r e g u l a t e d b y a m u l tih o r m o n a l s y s te m
a n d r e p r o d u c tio n , a r e c y c l ic a n d f a ir l y w e ll u n d e r s to o d in b e n -
b u t is u n d e r th e im m e d ia te c o n tr o l o f m o lt- p r o m o tin g s te r o id
th ic a n d te r r e s tr ia l m a la c o s tr a c a n s , s u c h a s d e c a p o d s , is o p o d s ,
h o r m o n e s ( e c d y s te r o i d s ) s e c r e te d b y a n e c d y s ia l g la n d , c a lle d
a n d a m p h ip o d s [ 2 2 ,1 0 1 ,1 0 2 ] , T h e s e b io lo g ic a l p r o c e s s e s a re
th e Y - o r g a n ( th e h o m o lo g u e o f th e p r o t h o r a c i c g la n d in in s e c ts
r e g u la te d b y a c o m p le x e n d o c r in e s y s te m [ 9 6 ,1 0 1 ], B a s ic a lly ,
in p u ts f r o m t h e e n v ir o n m e n t a r e in te g r a te d b y th e c e n tr a l n e r ­
[1 0 3 .1 3 6 ] , T h e Y - o r g a n s e c r e te s e c d y s o n e , w h ic h , o n r e le a s e
v o u s s y s te m ; n e u r o tr a n s m itte r s a n d n e u r o m o d u la to r s g o v e r n
th e r e le a s e o f n e u r o p e p tid e s , w h ic h c o n tr o l th e p r o d u c tio n o f
h o r m o n e s b y th e e n d o c r in e g la n d s [1 0 3 ] . T h e m a in c r u s ta c e a n
e n d o c r in e c e n t e r s in c lu d e th e Y - o r g a n , m a n d ib u la r o r g a n , a n ­
in th e h e m o ly m p h , is c o n v e r te d in to a c t iv e 2 0 - h y d r o x y e c d y s o n e ( s y n o n y m s : c r u s te c d y s o n e a n d e c d y s te r o n e ) . C ir c u la tin g
t i t e r s o f 2 0 - h y d r o x y e c d y s o n e v a r y im p r e s s i v e ly d u r in g th e
m o lt c y c le [ 1 0 3 ,1 3 4 ] . T h e Y - o rg a n p r o d u c e s tw o o th e r e c d y s ­
t e r o id s , 3 - d e h y d r o e c d y s o n e a n d 2 5 - d e o x y e c d y s o n e , w ith h e
d r o g e n ic g la n d , X - o r g a n , a n d s in u s g la n d [7 ,9 7 ], U n f o r t u ­
la tte r f o r m in g th e im m e d ia te p r e c u r s o r to th e a c t iv e p o n a s te r-
n a te ly , e n d o c r in e g la n d s o r s ite s o f h o r m o n e p r o d u c tio n in
o n e A [1 3 3 ]. M o r e s tu d ie s h a v e b e e n d o n e o n th e e f f e c ts o f
m y s id s a r e la r g e ly u n d e s c r ib e d .
T h e e f f e c ts o f o r g a n ic a n d in o r g a n ic c o n ta m in a n t s o n c r u s ­
c o n ta m in a n t s o n m o ltin g a n d lim b g e n e r a t io n th a n o n a n y o th e r
h o r m o n e - m e d ia te d p r o c e s s in c r u s ta c e a n s [8 ,1 3 5 ,1 3 7 ] ,
ta c e a n f u n c tio n s r e g u la te d b y h o r m o n e s a r e b e in g in v e s t ig a te d
M o lt s ta g in g , b a s e d o n c h a n g e s in th e in te g u m e n t, h a s b e e n
w ith in c r e a s in g f r e q u e n c y a n d s e v e r a l s h o w p r o m i s e a s b io -
d e v e l o p e d f o r v a r io u s c r u s ta c e a n s a n d is g e n e r a l ly d iv id e d in to
m a rlc e rs o f e n v ir o n m e n ta l c o n ta m in a t io n a n d e n d o c r in e d i s ­
f o u r m a j o r p e r io d s : p o s tm o lt, in te r m o lt , p r e m o lt, a n d m o lt ( e c -
r u p ti o n [ 7 ,8 ,1 0 4 ,1 0 5 ] . U n f o r t u n a te ly , r e la tiv e ly f e w d a ta a r e
d y s is ) . M y s id m o lt s ta g e s h a v e b e e n d e s c r ib e d f o r S i r ie lla
a v a i la b le o n th e h o r m o n a l c o n tr o l o f b io lo g ic a l p r o c e s s e s in
a r m a t a [ 1 3 8 ,1 3 9 ] , M y s is m i x t a [ 1 4 0 ] , a n d N . in te g e r [1 4 0 ].
m y s id s . H a v in g s a id th a t, c e r ta in e n d p o in ts r e le v a n t to th e
In m y s id s , e c d y s is is i n s ta n ta n e o u s , w ith th e e n tir e c a r a p a c e
t e s t in g o f s u s p e c te d e n d o c r in e d is r u p to r s , s u c h a s s u r v iv a l,
lif tin g u p a n d th e m y s i d s lid in g o u t o f th e o ld c u tic le w h ile
fe c u n d ity , s e x u a l m a tu r a tio n , a n d b io m a s s in c r e a s e , a r e a lr e a d y
s w im m in g . F o r f e m a le m y s id s , in te g u m e n ta l d e v e l o p m e n t d u r ­
s ta n d a r d i z e d p r o c e d u r e s f o r s o m e m y s id s , s u c h a s A . b a h i a ,
in g m o lt p r e p a r a tio n , m a r s u p ia l b r o o d d e v e l o p m e n t, a n d d e ­
A . b ig e lo w i ( p a r tl y ) , A . a l m y r a , I I c o s ta t a , a n d N . m e r c e d i s ,
v e lo p m e n t o f n e w e g g s in th e o v a r y a r e s y n c h r o n iz e d , f a c il­
a n d m a n y o th e r e n d p o in ts o r s p e c ie s a re p r o m i s in g [7 ], T h e
ita tin g m o lt s ta g i n g [1 0 1 ] . T o d a te , o n ly o n e s tu d y h a s q u a n ­
u s e o f p o te n tia l m y s id h o r m o n e - r e g u l a te d e n d p o in ts a s b i o ­
tif ie d e c d y s tc r o id tite r s d u r in g th e m y s id m o lt c y c l e a n d th is
m a r k e r s o f e x p o s u r e o r e f f e c ts o f e n d o c r in e d is r u p to r s a r e
s tu d y w a s w ith S. a r m a t a [1 0 1 ] , H o w e v e r , b o th e c d y s o n e a n d
d is c u s s e d in d e ta il in t h e f o ll o w i n g p a r a g r a p h s a n d a r e s u m ­
2 0 - O H e c d y s o n e h a v e b e e n id e n tif ie d in TV. in t e g e r (A . G h e k -
m a r iz e d in T a b le 2 . A lth o u g h m a n y , i f n o t a ll, o f th e s e e n d ­
ie r e e t a l., L a b o r a to r y o f E n v i r o n m e n ta l T o x ic o lo g y a n d A q u a t­
p o in ts m a y in d ic a te a r e s p o n s e to a n e n d o c r in e d is r u p to r , m o s t
ic E c o lo g y , G h e n t U n iv e r s ity , G h e n t, B e lg iu m , u n p u b lis h e d
a ls o v a r y in r e s p o n s e to e x p o s u r e to o th e r s tr e s s o r s a n d th is
d a ta ) a n d A . b a h ia ( S .R . T u b e rty , C e n t e r f o r E n v ir o n m e n ta l
is f u r th e r c o n f o u n d e d b y th e i n te r r e la te d n e s s (i.e ., n o n in d e ­
D ia g n o s ti c s a n d B i o r e m e d ia t io n , U n iv e r s ity o f W e s t F lo r id a ,
p e n d e n c e ) o f s o m e o f th e s e e n d p o in ts [1 3 ] . T h e k e y to th e
P e n s a c o la , F L , U S A , a n d C .L . M c K e n n e y , J r., U .S . E n v i r o n ­
in te r p r e ta tio n o f th e s e e n d p o in ts a s in d ic a to r s o f e n d o c r in e
m e n ta l P r o te c tio n A g e n c y , N a tio n a l H e a lt h a n d E n v i r o n m e n ta l
d is r u p tio n w ill b e to c r e a t e f o r e a c h s p e c ie s a la rg o d a ta b a s e
E f f e c t s R e s e a r c h L a b o r a to r y , G u l f B r e e z e , F L , u n p u b lis h e d
o f w h a t c o n s ti tu te s th e n o r m a l u n s tr e s s e d r e s p o n s e , a n d w h a t
d a ta ).
c o n s ti tu te s a n o r m a l r e f e r e n c e s ite o r p o p u la tio n w h e n w o r k in g
u n d e r f ie ld c o n d itio n s .
A s m e n tio n e d p r e v io u s ly , m o ltin g is c o n tr o lle d b y e c d y s ­
te r o id s [1 3 6 ] , E c d y s te r o id s c o n tr o l th e a c t iv ity o f s p e c ific
g e n e s a t th e tr a n s c r ip tio n a l le v e l b y i n te r a c tin g w ith th e in ­
G r o w th a n d m o lt i n g
t r a c e llu la r e c d y s te r o id r e c e p t o r [1 0 3 ,1 3 5 ,1 4 1 ] . In a r th r o p o d s ,
M o s t c o m m o n ly , g ro w th is m e a s u r e d e ith e r b y in c r e a s e s o f
th e e c d y s o n e r e c e p t o r is in th e s a m e g e n e f a m ily a s th e v e r­
d r y w e i g h t o r b o d y le n g th p e r tim e in te r v a l [ 5 9 ,1 0 6 - 1 0 8 ] a n d ,
te b r a te th y r o i d r e c e p t o r b u t, in te r e s tin g ly , s te r o id a l e s tr o g e n s
f o r c r u s ta c e a n s , is o f te n e x p r e s s e d in te r m s o f i n te r m o lt p e r io d
d o n o t a g o n iz e o r a n ta g o n iz e th e e c d y s te r o id r e c e p t o r [1 4 2 ].
a n d g r o w th f a c to r ( p e r c e n ta g e i n c r e a s e in b o d y s iz e a t th e m o lt)
E v i d e n c e e x is ts [1 4 2 ] th a t s o m e n o n s te r o id a l e n v ir o n m e n ta l
[2 6 ] . G r o w th c u r v e s ( s u c h a s th e v o n B e r ta la n f f y e q u a tio n )
e s tr o g e n s a r e e c d y s te r o id a n ta g o n is ts ( e .g .,l in d a n e , b is p h e n o l
c a n b e f itte d to tire g r o w th d a ta [ 2 0 ,1 0 9 ,1 1 0 ] , a llo w in g c o m ­
A , d ie th y lp h th a la te , a n d jö,/P - D D T ) . In a d d itio n , s e v e ra l c la s ­
p a r is o n s o f th e d i f f e r e n t g r o w th p a r a m e te r s b e tw e e n tr e a t­
s e s o f p h y to c h e m ic a ls a n ta g o n iz e e c d y s o n e a c tiv ity [1 3 7 ], T h e
m e n ts . A lth o u g h s e v e r a l s tu d ie s h a v e f o c u s e d o n g r o w th a n d
a p p a r e n t u b iq u ity o f t h e a n tie c d y s te r o id a l a c t iv ity o f e n v ir o n ­
m o ltin g in m y s id s u n d e r n a t u r a l c o n d itio n s [4 9 ,1 0 7 ,1 1 .1 —1 2 3 ],
m e n ta l c h e m ic a ls n e c e s s it a te s in v e s t ig a ti o n in to th e ir p o te n tia l
e x p o s u r e e x p e r im e n ts a ls o h a v e c o n f ir m e d th e s e n s itiv ity o f
e f f e c ts o n c r u s ta c e a n s [ 1 4 3 ,1 4 4 ] , M a n y p e s ti c id e s , g e n e r a l ly
th e s e e n d p o in ts in to x ic o lo g y [ 5 2 , 5 6 ,6 1 , 8 4 ,9 2 ,1 0 6 ,1 0 8 ,1 2 4 -
c la s s e d a s in s e c t g r o w th r e g u la to r s , f u n c tio n a s e c d y s o n e a g ­
1 3 1 ], F o r m y s id s , r e d u c e d g r o w th is th e m o s t c o m m o n s u b -
o n is ts [ 7 ,1 1 ], A s s u g g e s te d b y Z o u a n d F in g e r m a n [1 3 6 ] , fu tu r e
le th a l r e s p o n s e to t o x ic a n t e x p o s u r e a n d th is h a s im p o r ta n t
in v e s t ig a ti o n s o f m o ltin g in c r u s ta c e a n s s h o u ld h a v e tw o e m ­
im p lic a tio n s f o r r e p r o d u c tiv e s u c c e s s b e c a u s e f e c u n d ity is r e ­
p h a s e s , o n e e x a m in in g in te r a c tio n s b e tw e e n p o te n tia l e n d o ­
la te d d ir e c tly to f e m a le b o d y s iz e [ 2 0 ,8 4 ,1 1 8 ,1 2 3 ] . In c r u s ­
c r in e d is r u p to r s a n d th e e c d y s o n e r e c e p to r , a n d o n e fo c u s in g
ta c e a n s , s ig n if ic a n t g r o w th o c c u r s o n ly a s a r e s u lt o f m o ltin g ;
th e r e f o r e , d is r u p tio n o f m o ltin g m a y r e s u lt in a lte r a tio n s in
o n th e p o s s ib le im p a ir m e n t o f e c d y s te r o id o g e n e s is b y th e s e
a g e n t s . In v itr o a s s a y s c a n d e te r m in e q u ic k ly w h e t h e r a c h e m ­
g r o w th [ 1 3 ,1 3 2 ],
ic a l h a s ( a n ti) e c d y s te r o id a l a c t iv ity [ 1 4 5 ,1 4 6 ] . G iv e n th e c u r­
E c d y s te r o id s (th e m o ltin g h o r m o n e s in c r u s ta c e a n s ) a ls o
r e n t m e th o d s f o r q u a n tif y in g e c d y s te r o id s b y u s in g im m u n o ­
f u n c tio n in th e c o n tr o l o f r e p r o d u c tio n a n d e m b r y o g e n e s is
a s s a y [ 1 3 9 ,1 4 4 ,1 4 7 ] a n d th e a v a i la b le m e th o d s f o r m o lt s ta g in g
[9 7 ,1 3 3 ] ; th e r e f o r e , th e c r u s ta c e a n m o lt c y c l e h a s p r o f o u n d
[ 1 3 8 ,1 4 0 ] , i t s h o u ld b e p o s s ib le to e v a l u a te th e p o te n tia l in ­
M ysids and e n d o crin e d isru p tio n : A review
E nviro n . T oxicol. Cheni. 23, 2004
1225
te r a c tio n o f th e s e c h e m ic a ls w ith th e p r o c e s s o f m o ltin g in
p e s ti c id e c h lo r p y r i f o s (T .A . V e r s ly c k e e t a l., u n p u b lis h e d d a ta ).
m y s id s .
T h e c e l lu la r e n e r g y a llo c a ti o n a s s a y a ls o w a s r e c e n t ly v a lid a te d
B e c a u s e e c d y s te r o id s a r c u s e d a s m a j o r e n d o c r in e - s i g n a lin g
in th e f ie ld ( S c h e ld t e s tu a r y , T h e N e th e r la n d s ) [9 4 ] , T h e e c o ­
m o le c u le s in c r u s ta c e a n s [7 ] , a n d littl e is k n o w n o f t h e i r o th e r
lo g ic a l r e le v a n c e a n d u tilit y o f s h o r t- te r m b io in d ic a to r s o f m e t­
w ith
a b o lic p r o c e s s e s in A . b a h i a h a v e b e e n d e m o n s t r a te d a fte r
( a n ti) e c d y s te r o id a l a c t iv ity a ls o w i l l a f f e c t o th e r h o r m o n e - r e g ­
c h r o n ic e x p o s u r e to p e s ti c id e s [ 6 3 ,1 2 5 ,1 2 6 ,1 2 9 ,1 3 0 ] . In th e s e
u la te d p r o c e s s e s in c r u s ta c e a n s . S u p p o r t f o r th is h y p o th e s i s is
s tu d ie s , p e s ti c id e - e x p o s e d j u v e n i l e m y s id s h a d a g r e a te r r e l i ­
fu n c tio n s ,
it
m ay
be
e x p e c te d
th a t
a
c h e m ic a l
p r o v id e d b y M u a n d L e B la n c [ 1 4 4 ] , w h o d e m o n s t r a te d th a t
a n c e o n th e m o r e e n e r g y - r ic h l ip id s u b s tr a te s d u r in g m a tu r a tio n
th e f u n g ic id e f e n a r im o l a lte r e d e m b r y o d e v e l o p m e n t in d a p h n m a jo r a d v a n t a g e o f u s in g e c d y s te r o id m e ta b o lis m a s a n e n d ­
to s u p p o r t e le v a te d m e ta b o lic d e m a n d s , r e s u l t i n g in le s s lip id
m a te r ia l a v a i la b le f o r g a m e te p r o d u c tio n a n d r e d u c e d r e p r o ­
d u c tiv e s u c c e s s . U n e x p o s e d m y s id s s h if t to w a r d m o r e p ro -
p o in t is t h a t i t p r o v id e s a m e a n s o f e v a l u a tin g th e im p a c t o f
te in a c e o u s s u b s tr a te s d u r in g m a tu r a tio n , a s d e m o n s t r a te d fo r
e n v ir o n m e n ta l c h e m ic a ls o n c r u s ta c e a n s (a n d p o te n tia lly o th e r
A . b a h i a [6 3 ] a n d N . i n t e g e r [ 1 5 5 ,1 5 9 ] , T h e s e c h a n g e s in
a r th r o p o d s ) , w h ile n o t n e c e s s a r il y a f f e c tin g v e r te b r a te s . B e ­
m e ta b o lic s u b s tr a te u s a g e c a n b e m e a s u r e d b y m o n ito r in g th e
id s b y in te r f e r in g w ith e c d y s te r o id m e ta b o lis m . H o w e v e r , o n e
c a u s e ( a n ti) e c d y s te r o id a l a c t iv ity h a s b e e n p r o v e n in v itr o fo r
o x y g e n to n itr o g e n r a tio [ 1 2 5 ,1 6 0 ] , th e lip id a n d p r o te in c o n ­
c e r ta in c h e m ic a ls [1 4 5 ] , a n d d is r u p tio n o f m o ltin g h a s b e e n
t e n t [1 5 5 ] , o r th e c a r b o n to n itr o g e n r a tio o f th e t e s t o r g a n is m
o b s e r v e d a s a r e s u lt o f c h e m ic a l e x p o s u r e , th e s e c h e m ic a ls
[1 1 4 ], O n th e o th e r h a n d , h y p e r g ly c e m ia is a c o m m o n r e s p o n s e
s h o u ld b e te s t e d in e x p o s u r e s w ith m y s id s . In th e s e e x p o s u r e s ,
to e n v ir o n m e n ta l o r f u n c tio n a l h y p o x ia a n d c o n ta m in a n t e x ­
e n d p o in ts s u c h a s i n te r m o lt p e r io d , g r o w th , m o r p h o l o g ic a l a b ­
p o s u r e in n u m e r o u s d e c a p o d s , a n d it is t h o u g h t to b e tr ig g e r e d
e r r a tio n s , e c d y s o n e tite r s , p r o te in c o n c e n tr a tio n s , in te g u m e n t
b y th e a c tio n o f c r u s ta c e a n h y p e r g ly c e m ic h o r m o n e o n v a r io u s
d e v e l o p m e n t, a s w e l l a s r e la te d e n d p o in ts s u c h a s v ite llo g e n -
t a r g e t tis s u e s [ 8 ,1 4 9 ]. T h e a m in o a c i d s e q u e n c e o f c ru s ta c e a n
e s is , s h o u ld d e te c t in v iv o e f f e c ts o f c h e m ic a ls o n e c d y s te r o id
h y p e r g ly c e m ic h o r m o n e is h ig h ly h o m o lo g o u s w ith t h a t o f th e
m e ta b o lis m a n d m o ltin g in m y s id s . In v itr o a s s a y s s h o u ld a id
th e m e c h a n is tic u n d e r s ta n d i n g o f c h e m ic a l a c tio n to b e tte r
m o lt- in h ib itin g h o r m o n e , a n o th e r p r o d u c t o f th e s in u s g la n d s
in c r u s ta c e a n s , in d ic a tin g p o s s i b l e in v o lv e m e n t in th e c o n tro l
a llo w d is tin c tio n b e tw e e n e n d o c r in e - s p e c if ic a n d p h a r m a c o ­
o f m o ltin g a n d r e p r o d u c t i o n [1 4 9 ] . S e v e r a l in v e s t ig a to r s h a v e
lo g ic a l e ffe c ts .
e x a m in e d th e e f f e c ts o f m e ta ls a n d o r g a n ic c o n ta m in a n t s o n
b l o o d g lu c o s e c o n c e n tr a tio n s a n d c r u s ta c e a n h y p e r g ly c e m ic
E n e r g y m e ta b o lis m
h o r m o n e tite r s in c r u s ta c e a n s [8 ], C h a n g e s in b l o o d g lu c o s e
B io m a r k e r s lin k e d w ith p h y s io lo g ic a l e n e r g e t ic s p r o v id e
le v e ls in m y s id s e x p o s e d to p o te n tia l e n d o c r in e d is r u p to r s m a y
in f o r m a tio n o n k e y p r o c e s s e s in th e o r g a n i s m ’s e n e r g y a c ­
in d ic a te d is r u p tio n o f h o r m o n a l a c t iv ity o th e r th a n th a t a s s o ­
q u is itio n a n d e x p e n d i tu r e , a n d p o s s ib ly a ls o e lu c id a te th e m o d e
c ia te d w ith m o ltin g o r r e p r o d u c tio n [1 3 ],
o f a c tio n o f th e to x ic a n t. U n d e r n o r m a l c o n d itio n s , s p e c if ic
T h e m e th o d s d e s c r ib e d a b o v e a r e tr a n s f e r r e d e a s ily to o th e r
a m o u n ts o f e n e r g y a r e a llo c a te d to b a s a l m e ta b o lis m , g r o w th ,
a n d r e p r o d u c tio n a n d th e r e f o r e , th e o r e tic a l ly , c h a n g e s in m e t­
m y s id s p e c ie s . E n d p o i n ts r e l a t e d to e n e r g e t ic p r o c e s s e s a r e
r e la tiv e ly e a s y to m e a s u r e , b u t a b e t t e r a n d h o lis tic u n d e r ­
a b o lic t u r n o v e r a n d s p e c if ic a llo c a ti o n s s h o u ld b e lin k e d to
s ta n d i n g o f tire r o le o f th e d i f f e r e n t h o r m o n e s in v o lv e d in
e f f e c ts a t h i g h e r le v e ls o f e c o lo g ic a l o r g a n iz a tio n [1 4 8 ]. A
e n e r g y m e ta b o lis m , s u c h a s c r u s ta c e a n h y p e r g ly c e m ic h o r­
la r g e b o d y o f in f o r m a tio n is a v a i la b le o n th e n e u r o e n d o c r in e
m o n e , is n e e d e d to e v a l u a te th e p o te n tia l im p a c t o f h o r m o n e -
p a th w a y s o f p h y s io lo g ic a l r e g u la tio n in m a c r o c r u s t a c e a n s . A l­
m im ic k in g s u b s ta n c e s o n m y s id s . In th is c o n te x t, n e w im ­
th o u g h ty p ic a l a n d w e ll - s t u d i e d c h a l le n g e s to e n d o g e n o u s e n ­
m u n o a s s a y s f o r d e te r m in a tio n o f c ir c u la tin g h o r m o n e s in th e
e rg y m e ta b o lis m in c lu d e e n v ir o n m e n ta l h y p o x ia , fu n c tio n a l
h e m o ly m p h , s u c h a s c r u s ta c e a n h y p e r g ly c e m ic h o r m o n e , a re
( in t e r n a l) h y p o x ia , c h a n g i n g e n e r g e t ic r e q u ir e m e n ts , d is tu r ­
p r o m i s in g [1 6 1 ],
b a n c e to w a t e r b a la n c e a n d io n h o m e o s t a s is a n d c h a n g e s in
O th e r e n d p o in ts r e la te d to m e ta b o lis m in m y s id s h a v e b e e n
te m p e r a tu r e ( f o r r e v ie w , r e f e r to M o r r is a n d A ir r ie s s [1 4 9 ] ),
e x p o s u r e to to x ic a n ts a ls o w ill r e s u lt in a n e n e r g e t ic c h a lle n g e .
s tu d ie d . H ig h a c e ty lc h o lin e s te r a s e a c t iv ity in S i r i e l l a c la u s i ,
B e c a u s e e n e r g e t ic p r o c e s s e s a r e h o r m o n e - r e g u l a te d , th e y a re ,
tic u la r ly s u ite d f o r r e s e a r c h b a s e d o n b io m a r k e r s in tire m a r in e
in d ic a tin g a h ig h m e ta b o lic r a te , id e n tif ie d th is m y s id a s p a r­
b y d e f in itio n , s e n s itiv e to h o r m o n e d is r u p tio n a n d s e v e r a l m e a ­
e n v ir o n m e n t [1 7 ]. In a d d itio n , N . i n t e g e r h a s b e e n u s e d a s a
s u r e m e n ts o f e n e r g y r e s e r v e s a n d c o n s u m p t io n m a y s e r v e as
v ia b le a lte r n a tiv e m o d e l f o r th e p a r tia l r e p la c e m e n t o f v e r te ­
u s e f u l b io m a r k e r s o f e n d o c r in e - d is r u p tin g s u b s ta n c e s in c r u s ­
ta c e a n s [1 1 ]. H o w e v e r , o n c e a g a in , th e u tilit y o f b io e n e r g e tic
b r a te a n im a ls in m e ta b o lic s tu d ie s w ith ille g a l g ro w th p r o ­
e n d p o in ts w ill d e p e n d s tr o n g ly o n e s t a b lis h m e n t o f a c o n s e n ­
s p o n s e s h a v e b e e n s tu d ie d in m y s id s in r e la tio n to a v a r ia b le
s u s d a ta b a s e o f w h a t th e n o r m a l b io e n e r g e tic s ta t e is f o r m y s ­
e n v ir o n m e n t a n d to x ic e x p o s u r e [ 6 3 ,8 1 ,8 4 ,1 1 5 ,1 2 5 ,1 6 0 ,1 6 3 —
id s.
1 6 5 ].
m o te r s a n d v e t e r i n a r y d r u g s [1 6 2 ], F in a lly , r e s p ir a to r y r e ­
A lte r a tio n s to th e e n e r g y m e ta b o lis m o f m y s id s h a v e b e e n
A lth o u g h u n d o u b te d ly h a v in g e n v ir o n m e n ta l r e le v a n c e a n d
u s e d s u c c e s s f u lly a s a n in d ic a to r o f s tr e s s to to x ic a n t e x p o s u r e
b e in g f a ir l y e a s ily e x tr a p o la te d to h ig h e r le v e ls o f b io lo g ic a l
in A . b a h ia [ 6 3 ,1 2 5 ,1 2 6 ,1 2 9 ,1 3 0 ] , P. f l e x u o s u s [8 1 ] , N . a w a t ­
o r g a n iz a tio n , th e m a jo r d is a d v a n ta g e o f e n d p o in ts r e la te d to
s c h e n s i s [ 1 5 0 - 1 5 2 ] , a n d N . i n t e g e r [ 1 5 3 - 1 5 6 ] . I n A . b a h ia ,
e n e r g y m e ta b o lis m is t h e i r d if f ic u lty in m e c h a n is tic a lly e x ­
P . f l e x u o s u s , a n d N . a w a t s c h e n s i s , w e ig h t- s p e c if ic re s p ir a tio n ,
p la in in g h o r m o n e - r e g u l a te d r e s p o n s e s a s c a n b e e x p e c te d fro m
a m m o n ia e x c r e t io n r a te s , a n d o x y g e n to n itr o g e n r a tio s h a v e
b e e n m e a s u r e d a f te r t o x ic a n t e x p o s u r e . In N . in te g e r , R o a s t
e x p o s u r e to e n d o c r in e d is r u p to r s . M a n y a b io tic a n d to x ic
s tr e s s o r s a f f e c t th e e n e r g y m e ta b o lic p r o c e s s e s o f o r g a n is m s
e t a l. [1 5 3 ] u s e d s c o p e f o r g r o w th [1 5 7 ] , w h e r e a s V e r s ly c k e
[ 1 1 4 , 1 2 7 ,1 5 2 - 1 5 6 ,1 6 4 ] , w h ile n o t n e c e s s a r il y b e in g r e la te d to
a n d J a n s s e n [1 5 5 ] a n d V e r s ly c k e e t al. [1 5 6 ] u s e d th e c e l lu la r
d is r u p tio n in n o r m a l h o r m o n a l r e g u la tio n . T h e r e f o r e , th e s u c ­
e n e r g y a llo c a ti o n a s s a y [1 5 8 ] . B o th m e th o d s a r e p r o m is in g
c e s s f u l u s e o f b io m a r k e r s f o r th e e v a l u a tio n o f e n d o c r in e d is ­
a n d w e r e r e c e n t ly v a lid a te d in N. i n t e g e r a f te r e x p o s u r e to th e
r u p ti o n w ill b e lim it e d b y th e a m o u n t o f b a c k g r o u n d d a ta o n
1226
E n viro n . T o x ic o l Cheni. 2 3 , 2004
T .A . V e rsly ck e et al.
n a tu r a l v a r ia tio n a n d n o r m a l le v e ls o f t h e e n d p o in ts in q u e s ­
io u s r a te s o f te s t o s te r o n e c o n v e r s io n to a n d r o s te n e d io n e in
tio n , a n d a ls o b y th e f u n d a m e n ta l u n d e r s ta n d i n g o f th e to x ic a n t
d a p h n id s . F u t u r e s tu d ie s a r e n e e d e d to r e v e a l i f th e s e c o n v e r ­
a c tio n a t th e ( s u b ) c e llu la r le v e l. I n th is r e g a r d , m a n y s tu d ie s
s io n s a r e a f f e c te d b y a g e , g e n d e r , r e p r o d u c tiv e s ta t e , o r c h a n g e s
c o n ta in in g b a c k g r o u n d in f o r m a tio n o n th e b io c h e m ic a l c o m ­
in th e a b io tic e n v ir o n m e n t. N o t e th a t a n a n d r o g e n r e c e p t o r h a s
p o s itio n ( e .g ., p r o te in s , lip id s , a n d s te r o ls ) o f m y s id s h a v e b e e n
n o t b e e n f o u n d o r c lo n e d in c r u s ta c e a n s . T h e r e f o r e , id e n tif i­
p u b lis h e d [ 1 6 6 - 1 8 2 ] , b u t in f o r m a tio n o n n e u r o p e p tid e a n d
c a tio n a n d c h a r a c te r iz a tio n o f th e a n d r o g e n r e c e p t o r s h o u ld b e
h o r m o n a l le v e ls in m y s id s n e e d s d e v e l o p m e n t.
a p r io r ity f o r r e s e a r c h to e x p lo r e th e u s e f u ln e s s o f s e x s te r o id s
fo r e v a l u a tin g e n d o c r in e d is r u p tio n in c r u s ta c e a n s a n d o th e r
Steroid metabolism a n d cytochrome P450
P o ll u ta n ts m a y e x e r t r e p r o d u c tiv e e f f e c ts th r o u g h i n te r f e r ­
e n c e w ith n o r m a l s te r o id m e ta b o lis m [ 1 8 3 - 1 8 7 ] , F o r in v e r ­
te b r a te s , s e v e r a l s tu d ie s h a v e f o c u s e d o n p o l lu ta n t- i n d u c e d a l­
te r a tio n s in s te r o id m e ta b o lis m . T h e s e c h e m ic a ls o f te n in te r­
f e r e w ith th e m ic r o s o m a l P 4 5 0 m o n o o x y g e n a s e s y s te m , a ls o
c a lle d th e m ix e d - f u n c tio n o x y g e n a s e s y s te m . T h e m ix e d - f u n c tio n o x y g e n a s e s y s te m is in v o lv e d n o t o n ly in th e m e ta b o lis m
o f o r g a n ic to x ic a n ts b u t a ls o in s te r o id m e ta b o lis m ; c o n s e ­
q u e n tly , in d u c tio n o r in h ib iti o n o f th e m ix e d - f u n c tio n o x y ­
g e n a s e s y s te m a ls o m a y h a v e r e p e r c u s s io n s f o r th e h o r m o n a l
c o n tro l o f r e p r o d u c tio n . In s e a s ta r s , a lin k a g e w a s d e m o n ­
s tr a te d b e tw e e n im p a ir e d r e p r o d u c tiv e s u c c e s s , p o llu tio n - m o d ­
u la te d e n d o c r in e f u n c tio n , a n d in d u c tio n o f th e m ix e d - f u n c tio n
o x y g e n a s e s y s te m [1 8 7 ] , I n g a s tr o p o d s , m u c h w o r k o n s te r o id
in v e rte b r a te s .
S tu d ie s o v e r th e l a s t 3 0 y e a r s h a v e e s ta b lis h e d th e im p o r ta n t
r o le o f c y to c h r o m e P 4 5 0 in th e b io tr a n s f o r m a tio n o f x e n o b io tic s a n d e n d o g e n o u s c o m p o u n d s (s u c h a s e c d y s te r o id s ) in
c r u s ta c e a n s ( f o r a r e v i e w o n c a i s t a c c a n P 4 5 0 , r e f e r to J a m e s
a n d B o y le [1 8 4 ] ). A lth o u g h n o s tr u c tu r a l in f o r m a tio n o n c y ­
to c h r o m e P 4 5 0 in c r u s ta c e a n s is a v a i la b le , i t is c le a r t h a t th e y
a re in v o lv e d in s e v e r a l s te p s in th e b io s y n th e s is o f e c d y s te r o id s
a n d o th e r p h y s io lo g ic a ll y i m p o r ta n t s u b s tr a te s in c r u s ta c e a n s
[2 0 1 ] . M o r e s tu d ie s a r e n e e d e d to u n d e r s ta n d th e e f f e c ts , i f
a n y , o f v a r io u s c la s s e s o f e n v ir o n m e n ta l a n d o th e r c h e m ic a ls
th a t a r e k n o w n m o d u la to r s o f c y to c h r o m e P 4 5 0 e x p r e s s io n o r
a c tiv ity . N e w m o le c u la r to o l s , s u c h a s p r im e r - b a s e d re v e r s e
tr a n s c r i p t i o n - p o l y m e r a s e c h a i n r e a c t io n p r o c e d u r e s a n d e x ­
p r e s s io n o f P 4 5 0 s in h e t e r o lo g o u s s y s te m s , s h o u ld r e s u lt in
m e ta b o lis m h a s b e e n i n itia te d b y th e o b s e r v a ti o n o f tr i b u ty ltin -
b e tte r in s ig h ts in to th e f u n c tio n a n d e x p r e s s io n o f P 4 5 0 s in
in d u c e d im p o s e x , a s ta te o f p s e u d o h e r m a p h r o d i s m in w h ic h
th e c o n te x t o f e n d o c r in e d is r u p tio n . I n a d d itio n , in v iv o m e t­
f e m a le s e x h ib it f u n c tio n a l s e c o n d a r y m a le c h a r a c te r is tic s . A l ­
a b o lic s tu d ie s w ith d i f f e r e n t s u b s tr a te s ( te s to s t e r o n e a n d e c ­
th o u g h th e u n d e r ly in g m e c h a n is m b y w h i c h tr i b u ty ltin c a u s e s
d y s o n e ) w ill p r o v i d e v a lu a b le to o ls f o r e v a l u a tin g th e e ffe c ts
o f to x ic a n t e x p o s u r e , p a r tic u la r ly w h e n l i n k e d w ith e f f e c ts o n
im p o s e x in g a s tr o p o d s h a s n o t b e e n e lu c id a te d c o n c lu s iv e ly ,
th e w e ig h t o f e v id e n c e is in f a v o r o f th e c y to c h r o m e P 4 5 0 d e p e n d e n t a r o m a ta s e in h ib iti o n h y p o th e s i s [ 1 8 7 - 1 9 2 ] ,
A lte r a tio n s in s te r o id m e ta b o lis m h a v e b e e n s tu d ie d in D .
m a g n a [ 1 8 6 ,1 9 3 - 1 9 7 ] a n d in th e b l u e c r a b C a ll i n e c t e s s a p id u s
[1 9 1 ] . Tn d a p h n id s , c h a n g e s in s te r o id m e ta b o lis m c o u ld p r o ­
h ig h e r le v e ls o f b io lo g ic a l o r g a n iz a tio n . A lth o u g h in f o r m a tio n
o n th e id e n tity o f P 4 5 0 s a n d t h e i r f u n c tio n a l r o l e in m y s id s
is , to o u r k n o w le d g e , n o n e x is t e n t, m y s id s s h o u ld b e a g o o d
m o d e l to s tu d y th e s e m e c h a n is m s . F r o m th e p r e l i m i n a r y s tu d ­
ie s w ith N . i n t e g e r b y V e r s ly c k e e t al. [1 85,1 9 8 ,1 9 9 ] , s u ff ic ie n t
v id e a n e a r ly in d ic a tio n o f p o te n tia l r e p r o d u c tiv e to x ic ity a f te r
in f o r m a tio n is a v a i la b le to s u g g e s t th a t m y s id s h a v e an e n ­
s u b le th a l
d is ru p to rs
z y m a tic b io tr a n s f o r m a tio n s y s te m t h a t r iv a ls th a t o f o th e r in ­
[ 1 8 6 ,1 9 3 ,1 9 4 ,1 9 7 ] . V e r s ly c k e e t a l. [1 8 5 ] r e p o r te d te s t o s te r o n e
v e r te b r a te s a n d v e r te b r a te s . M e ta b o lic s tu d ie s w ith p h y s io lo g ­
m e ta b o lis m a n d th e p r e s e n c e o f v e r te b r a te - ty p e s te r o id s in N .
ic a lly r e le v a n t s u b s tr a te s t h a t a ls o m e a s u r e h o r m o n e - r e g u l a te d
ex p o su re
to
s u s p e c te d
e n d o c rin e
in te g e r a n d d e m o n s t r a te d th e p r e s e n c e o f a c o m p le x s te r o id
e f f e c ts a t a h ig h e r le v e l o f b io lo g ic a l o r g a n iz a tio n (i.e ., r e ­
h y d r o x y la s e s y s te m c o n s is tin g o f d if f e r e n t P 4 5 0 is o z y m e s . T h e
p r o d u c tiv e s u c c e s s ) w o u ld b e v a lu a b le in e v a l u a tio n o f e n v i­
r e m a r k a b le d i v e r s ity o f te s t o s te r o n e h y d r o x y la tio n e x h ib ite d
r o n m e n ta l e n d o c r in e d is r u p tio n .
s h o u ld s tim u la te f u r th e r s tu d ie s o n th e in d u c tio n , s te r e o s p e c ific ity , a n d r e g u la tio n o f th e e n z y m e s y s te m s o f N . i n t e g e r a n d
R e p r o d u c t io n a n d v i te llo g e n e s is
o th e r m y s id s . M o r e r e c e n tly , a lte r a tio n s in th e p h a s e I a n d II
A lth o u g h th e m a in n e u r o s e c r e t o r y c e n t e r s a n d th e s in u s
te s t o s te r o n e m e ta b o lis m in N . in te g e r a f te r a c u te e x p o s u r e to
g la n d in m y s id s r e s e m b l e th e s e f r o m d e c a p o d s , s e x u a l d if f e r ­
t r i b u ty ltin h a v e b e e n d e m o n s t r a te d [1 9 8 ] . I n a d d itio n , m e ta ­
e n tia tio n in ju v e n i l e s a n d m y s id r e p r o d u c tio n a r e m o r e lik e
b o lic s tu d ie s w ith N . i n t e g e r h a v e b e e n u s e d r e c e n t ly in e x ­
th o s e o f a m p h ip o d s a n d is o p o d s a n d a r e s t r i c t l y lin k e d to th e
m o lt c y c le [1 0 1 ]. In m y s id s , e m b r y o n ic a n d p o s te m b r y o n ic
p o s u r e e x p e r im e n ts w ith o th e r c h e m ic a ls , s u c h a s n o n y lp h e n o l
a n d m e th o p r e n e , a n d a ls o in th e f ie ld [1 9 9 ].
T h e p r e s e n c e o f s e x h o r m o n e s h a s b e e n s u g g e s te d in m a n y ,
d e v e l o p m e n t o c c u r s in th e f e m a le m a r s u p iu m a n d in c lu d e s fiv e
c o n s e c u tiv e s ta g e s f r o m
o v ip o s itio n
to th e j u v e n i l e
s ta g e
i f n o t a ll, a r th r o p o d s [1 8 3 ] . V e r te b r a te - ty p e s te r o id s (s u c h as
[ 2 6 ,1 1 8 ,2 0 2 - 2 0 5 ] , J u v e n il e s a r e lib e r a te d im m e d ia te ly b e f o r e
1 7 ß - e s tr a d io l, te s t o s te r o n e , a n d p r o g e s te r o n e ) h a v e b e e n m e a ­
s u r e d in s e v e r a l m a la c o s tr a c a n c r u s ta c e a n s [ 7 ,1 8 4 ] . A lth o u g h
e c d y s is o f th e m o th e r, s h o r tly a f t e r w h i c h s h e la y s a n e w b a tc h
th e la c k o f a r o le f o r v e r te b r a te s e x s te r o id h o r m o n e s in a r­
f o r a n e w b a tc h o f o o c y te s o n th e s e c o n d d a y o f th e m o lt c y c le .
t h r o p o d s h a s b e e n h i g h l i g h t e d [ 7 ,1 8 3 ] , f r a g m e n te d e v id e n c e
S e c o n d a r y v ite llo g e n e s is is n o t o n ly c y c l ic a l, a s in o th e r c r u s ­
s u g g e s ts th a t s o m e o f th e s e c o m p o u n d s m a y f u n c tio n a s h o r ­
ta c e a n s [ 2 0 6 ] , b u t a ls o s t r i c t l y lin k e d to th e m o lt c y c le , o f f e r in g
m o n e s in c r u s ta c e a n s [7 ,1 4 3 ,1 8 5 ]. E n d o g e n o u s a n d r o g e n s m a y
a n e x a m p le o f th e ty p e 2 p a tte r n ( e .g ., A m p h ip o d a , I s o p o d a ,
b e th e p r e c u r s o r s f o r o th e r h o r m o n e s ; th e r e f o r e , e x p o s u r e to
a n d D e c a p o d a ) f o r th e r e g u la tio n o f s im u lta n e o u s g o n a d a l a n d
e x o g e n o u s a n d r o g e n s ( o r a n d r o g e n m im ic s ) c o u ld e l i c i t a c t iv ­
ity th r o u g h r e c e p t o r s o th e r th a n th e a n d r o g e n re c e p to r. A l­
s o m a t ic g r o w th in c r u s ta c e a n s [ 2 0 6 ,2 0 7 ] , C u z in - R o u d y a n d
th o u g h th is h a s n o t b e e n d e te r m in e d in c r u s ta c e a n s , V e rs ly c k e
th e m y s id S. a r m a t a a s a b io lo g ic a l m o d e l f o r th e s tu d y o f
e t a l. [1 8 5 ] fo u n d e v id e n c e o f a s e x - s p e c if ic p r o d u c tio n o f
h o r m o n a l c o n tr o l o f m o lt a n d r e p r o d u c tio n , w h ic h s h o u ld b e
a n d r o g e n s , s u c h a s t e s t o s te r o n e a n d a n d r o s te n e d io n e , in N .
e x te n d e d f o r o th e r m y s id s p e c ie s . In a d d itio n , W o r th a m a n d
i n te g e r . S im ila r ly , L e B la n c a n d M c L a c h la n [2 0 0 ] r e p o r te d v a r ­
P r ic e [2 0 5 ] a n d G r e e n w o o d e t a l. [2 0 8 ] p u b lis h e d s tu d ie s on
o f e g g s in th e m a r s u p iu m . A s e c o n d a r y v ite llo g e n ic c y c le s ta r ts
S a le u d d in [1 0 1 ] p u b lis h e d a n e x c e l l e n t re v ie w ’ o n th e u s e o f
E nviro n . T oxicol. Cheni. 23, 2004
M y sid s and en d o crin e d isru p tio n : A rev iew
t h e in v itr o c u ltu r e o f m y s id m a r s u p ia l d e v e l o p m e n ta l s ta g e s
1227
la tio n s [ 1 1 , 5 8 ,6 0 , 6 3 ,1 0 8 ,1 2 5 ,1 2 6 ,1 2 8 ,1 3 0 ,2 1 8 ,2 1 9 ] (T a b le 2 ).
a t d if f e r e n t te m p e r a tu r e s , T h e s e a s s a y s s h o u ld b e e v a lu a te d
I n h ib ite d r e p r o d u c tio n is t h e m o s t s e n s itiv e , s u b le th a l p o p u ­
f u r th e r a s a m e a n s o f d e t e c t i n g e f f e c ts o f c o n ta m in a n t s o n
l a tio n r e s p o n s e o f A . b a h i a c h r o n ic a lly e x p o s e d to p e s ti c id e s
m a r s u p ia l d e v e l o p m e n t in m y s id s .
[6 3 ] . N u m e r o u s s tu d ie s h a v e d e s c r ib e d th e u s e o f r e p r o d u c tiv e
In g e n e r a l, f e w s tu d ie s h a v e b e e n c o n d u c t e d o n th e e f f e c ts
e n d p o in ts in m y s id s a f te r to x ic e x p o s u r e a n d c h a n g e s in th e
o f c o n ta m in a n t s o n g o n a d a l m a tu r a tio n o f c r u s ta c e a n s [8 ];
a b io tic
e n v ir o n m e n t
h o w e v e r , m u c h a tte n tio n h a s b e e n g iv e n r e c e n t ly to v i t e l l o ­
c h r o n ic
assay s,
g e n in , th e p r e c u r s o r t o th e y o lk p r o te in v ite llin in e g g - la y in g
s c r ib e d f o r A . b a h i a , th e s e s h o u ld b e a p p lic a b le to o th e r m y s ­
[ 1 5 ,1 2 3 ,1 3 1 ,2 2 0 ] .
in c lu d in g
A lth o u g h
s ta n d a r d
r e p r o d u c tiv e e n d p o in ts , a re d e ­
in v e r te b r a te s a n d v e r te b r a te s , a s a n i n d ic a to r o f e x p o s u r e to
id s , a lth o u g h th e lo n g e r lif e c y c l e in o th e r s p e c ie s m a y r e s tr ic t
e s tr o g e n i c x e n o b io tic s [ 5 , 2 0 9 - 2 1 6 ] , C o n tr o l o f v ite llo g e n e s is
is b e in g s tu d ie d in te n s iv e ly b e c a u s e y o l k is a n e x c e lle n t m o d e l
th e ir u s e in r o u ti n e te s tin g .
T h e lif e h i s t o r y o f A . b a h i a is v e r y a m e n a b le to d e m o ­
f o r s tu d y in g m e c h a n is m s o f h o r m o n a l c o n tr o l a t th e c e llu la r
g r a p h ic m o d e lin g b e c a u s e o f r a p i d g r o w th , e a r ly s e x u a l d if ­
a n d m o le c u la r le v e ls [ 5 ,2 1 5 ] , T o a s s e s s th e p o te n tia l a d v e r s e
f e r e n tia tio n ( a t 14 d ) a n d r e p r o d u c tio n ( c o m m e n c in g a ro u n d
e f f e c ts o f x e n o b io tic s o n c r u s ta c e a n r e p r o d u c tio n , it is im ­
17 d ) , a n d f r e q u e n c y o f b r o o d p r o d u c tio n ( a v e r a g e o f fiv e to
p o r ta n t to m e a s u r e a c c u r a te ly v ite llo g e n in a n d v ite llin in c r u s ­
s e v e n p e r f e m a le ) o v e r th e f u ll lif e s p a n o f 9 0 d [2 2 1 ,2 2 2 ],
ta c e a n m o d e ls (a n o v e r v ie w o f c r u s ta c e a n s p e c ie s f r o m w h ic h
v ite llin , v ite llo g e n in , o r lip o v ite lli n h a s b e e n is o la te d o r p a r­
T h e s e e n d p o in ts p r o v i d e u s e f u l in f o r m a tio n f o r p r e d ic tin g p o p ­
t i a lly c h a r a c te r iz e d is g iv e n b y T u b e r ty e t a l. [2 1 5 ] ). R e c e n tly ,
t h e r v a lid a tio n is n e e d e d in m u ltig e n e r a tio n a l l a b o r a to r y s tu d ­
a q u a n tita tiv e e n z y m e - lin k e d im m u n o s o r b e n t a s s a y w a s d e ­
v e lo p e d f o r th e m y s id A . b a h ia b y u s in g p o ly c lo n a l a n tis e r a
[2 1 5 ] . I n a d d itio n , s tu d ie s a r e u n d e r w a y to c h a r a c te r iz e a n d
p u r if y v ite llin o f th e m y s id N. i n t e g e r (A . G h e k ie r e e t a l.,
L a b o r a to r y o f E n v i r o n m e n ta l T o x ic o lo g y a n d A q u a tic E c o l ­
o g y , G h e n t U n iv e r s ity , G h e n t, B e l g iu m , u n p u b lis h e d d a ta ).
F u tu r e l a b o r a t o r y a n d fie ld s tu d ie s w ith m y s id s a r e n e e d e d to
e v a l u a te th e u s e o f th e s e im m u n o a s s a y s f o r in v e s t ig a ti n g e f­
f e c ts o f x e n o b io tic s o n c r u s ta c e a n v ite llo g e n e s is . A g o o d e x ­
a m p le o f th is is g iv e n b y O b c r d ö r s tc r e t a l. [ 2 1 3 ,2 1 4 ] , w h o
r e p o r te d th e e f f e c ts o f c h r o n ic p y r e n e e x p o s u r e o n m o ltin g
a n d r e p r o d u c tio n a s s e s s e d in t h e g r a s s s h r im p P a l a e m o n e te s
p u g i o b y u s in g a m o n o c lo n a l e n z y m e - lin k e d im m u n o s o r b e n t
a s s a y f o r v ite llin . O th e r s tu d ie s a ls o h a v e f o u n d a n im p a c t o f
x e n o e s tr o g e n s o n th e p r o d u c tio n o f c r u s ta c e a n p r o te in s (e .g .,
v ite llin a n d c y p r is m a j o r p r o te in ) , w h ic h a r e t h o u g h t to b e
u n d e r e s tr o g e n c o n tr o l [ 5 ,2 1 7 ] . F u tu r e w o r k o n s e q u e n c e d e ­
te r m in a tio n o f v ite llo g e n ic g e n e s a n d th e ir h o r m o n a l a c tiv ity
w i l l p r o v id e in te r e s tin g in s ig h t in to th e v ite llo g e n ic p r o c e s s
in m y s id s . S tu d y o f g e n o m ic a n d n o n g e n o m ic e f f e c ts o f e c ­
d y s te r o id s o n o v a r ia n m a tu r a tio n is a n o th e r p o te n tia l a r e a o f
w o r k . S y n e r g is tic a n d a n ta g o n is tic a c tio n s o f th e d if f e r e n t n e u ­
r o p e p tid e s , a n d th e m a n d ib u la r o r g a n c o n tr o l o v e r m o ltin g a n d
r e p r o d u c tio n , a r e o th e r a r e a s r e q u ir i n g f u r th e r s tu d y a s a b a s is
u la tio n - le v e l e f f e c ts o f r e p r o d u c tiv e to x ic a n ts . H o w e v e r, fu r­
ie s a s w e ll a s i n c o r p o r a tio n o f o th e r p o p u la tio n g r o w th p a ­
r a m e te r s s u c h a s d e n s it y d e p e n d e n c e , p r e d a tio n , m ig r a tio n , a n d
c o m p e titio n , b e f o r e c o n c l u s io n s c a n b e f o r m u la te d th a t a re
r e le v a n t f o r n a tu r a l e n v ir o n m e n ta l c o n d itio n s . P r e li m in a r y
tr a n s g e n e r a tio n a l r e s p o n s e s o f A . b a h i a to a p e s ti c id e a c tin g
a s a j u v e n i l e h o r m o n e a g o n i s t h a v e b e e n r e p o r te d [2 1 9 ] . S u r­
v iv a l, g r o w th , d e v e l o p m e n t, a n d r e p r o d u c tio n o f th is e s tu a r in e
m y s id w e r e m o n ito r e d t h r o u g h a n e n tir e lif e - c y c le e x p o s u r e
to f e n o x y c a r b a n d d u r in g t h e s e c o n d g e n e r a t io n w it h o u t a d ­
d itio n a l e x p o s u r e . J u v e n i l e m y s i d g r o w th , a n d c a r b o n a n d n i­
tr o g e n a c c u m u la tio n , a s w e l l a s m y s id s u r v iv a l th r o u g h th e
f ir s t b r o o d p r o d u c tio n , w e r e s ig n if ic a n tly a f f e c te d b y f e n o x ­
y c a r b . O n th e o th e r h a n d , m a tu r a tio n tim e , s e x d e te r m in a tio n ,
a n d y o u n g p r o d u c tio n w e r e n o t s ig n if ic a n tly a lte r e d d u r in g th e
lif e - c y c le e x p o s u r e . H o w e v e r , s e c o n d - g e n e r a tio n a d u lts , e x ­
p o s e d lo f e n o x y c a r b o n ly a s d e v e l o p in g e m b r y o s a n d j u v e ­
n ile s , p r o d u c e d f e w e r y o u n g a n d c o n ta in e d s ig n if ic a n tly f e w e r
m a le s . T h e s e r e s u lts d e m o n s t r a te c le a r ly th e n e e d f o r tr a n s g e n e r a tio n a l s tu d ie s w ith m y s id s to f u ll y u n d e r s ta n d th e p o ­
te n tia l c h r o n ic im p a c t o f e n d o c r in e d is r u p to r s .
D e ta i le d in f o r m a tio n a n d th e s h o r t lif e c y c le o f A . b a h ia
c le a r ly f a v o r th e u s e o f th i s s p e c ie s in th e in itia l d e v e l o p m e n t
a n d f u r th e r v a lid a tio n o f p o p u la tio n m o d e ls b a s e d o n r e p r o ­
f o r u s e o f c r u s ta c e a n s f o r e n d o c r in e - d is r u p tio n te s t in g in th e
d u c tiv e e n d p o in ts . A c o n c i s e d r a f t o f a d e ta ile d r e v ie w p a p e r
f u tu r e [1 3 ],
h a s b e e n p r o d u c e d b y U .S . E n v i r o n m e n ta l P r o te c tio n A g e n c y
[1 3 ] o n a r e c o m m e n d e d p r o to c o l a n d a d d itio n a l d a ta n e e d s fo r
L i f e - c y c l e t e s t i n g a n d p o p u l a t i o n a n d f i e l d s tu d ie s
D e s p i t e s u p e r f ic ia l r e s e m b l a n c e to d e c a p o d s h r im p s , m y s id s
a r e m o r e c lo s e ly r e la te d to a m p h ip o d s a n d is o p o d s , a n d a re
g r o u p e d to g e th e r in th e s u p e r o r d e r P e r a c a r id a . A ll th r e e o rd e rs
a r e g o o d c a n d i d a te s f o r to x ic o lo g ic a l te s tin g , a n d a m p h ip o d s
a n d m y s id s a r e u s e d r o u tin e ly . H o w e v e r , f o r e n d o c r in e - d is ­
a t w o - g e n e r a tio n lif e - c y c le t e s t w ith A . b a h i a in th e c o n te x t
o f e n d o c r in e d is r u p to r s . In th i s r e v ie w , th e f o llo w in g e n d p o in ts
a n d th e ir p r e f e r r e d m e th o d s f o r q u a n tif ic a tio n a re g iv e n : s u r ­
v iv a l, m o ltin g f r e q u e n c y , r e p r o d u c tio n ( s e x u a l m a tu r ity , tim e
to f ir s t b r o o d r e le a s e , b r o o d s iz e , a n d n u m b e r o f o f f s p r in g
p r o d u c e d ) , m e ta b o lic d is r u p tio n , d is r u p tio n in s te r o id m e ta b ­
r u p ti o n te s tin g , e s p e c ia lly f o r lif c - c y c lc te s ts , m y s id s o ff e r
o lis m , v ite llo g e n in in d u c tio n , c y to c h r o m e P 4 5 0 le v e ls , a n d
c le a r a d v a n t a g e s o v e r a m p h ip o d s . M o s t m a r in e a m p h ip o d s
b lo o d g lu c o s e le v e ls . T a b le 2 s u m m a r iz e s th e p o te n tia l e n d ­
u s e d in to x ic o lo g ic a l t e s t in g m u s t b e c o lle c te d fr o m th e ir n a t­
p o in ts f o r e v a l u a tin g e n v ir o n m e n ta l e n d o c r in e d is r u p tio n in
u r a l h a b ita ts b e f o r e u s e in te s ts . A lth o u g h t h e y c a n b e h e ld
m y s id s .
f o r a fe w w e e k s b e f o r e te s t in g , th e y g e n e r a l ly a r e n o t c u ltu r e d
T h e u s e o f m y s id s in fie ld s tu d ie s h a s b e e n e x tr e m e ly lim ­
f o r te s ts . C o n v e r s e ly , s e v e ra l m y s id s p e c ie s h a v e b e e n c u ltu r e d
ite d . M c K e n n e y e t a l. [9 2 ] a n d C la r k e t a l. [9 1 ] p e r f o r m e d
in th e la b o r a to r y a n d u s e d in lif e - c y c le te s ts [1 3 ] . S e v e ra l
e x p e r im e n ts w ith c a g e d m y s id s to e v a l u a te th e le th a l a n d s u b ­
m e a s u r e s o f r e p r o d u c tiv e p e r f o r m a n c e c a n b e u s e d to a s s e s s
le th a l r e s p o n s e s o f A . b a h ia d u r in g fie ld a p p lic a ti o n s o f fe n -
s u b le th a l r e s p o n s e in lif e - c y c le te s t in g , in c lu d in g s e x u a l m a ­
ih io n , a n o r g a n o p h o s p h a te in s e c tic id e . T o o u r k n o w le d g e ,
tu r ity . th e tim e to f ir s t b r o o d r e le a s e , th e tim e r e q u ir e d fo r e g g
th e s e a r e th e o n ly p u b l i s h e d s tu d ie s o n in s itu e x p o s u r e s w ith
d e v e l o p m e n t (a n d its s e p a r a te p h a s e s ) , fe c u n d ity , b r o o d s u c ­
c a g e d m y s id s . In a d d itio n , s tu d ie s t h a t h a v e in v e s t ig a te d b io ­
c e s s , a n d a lte r a tio n s in r e p r o d u c tiv e c h a r a c te r is tic s in p o p u ­
m a r k e r r e s p o n s e s in f ie ld - e x p o s e d m y s id s a ls o a re v e r y lim ite d
1228
E n viro n . T oxicol. Cheni. 2 3 , 200 4
T .A . V ersly ck e e t al.
[ 1 7 ,9 4 ], C le a r ly , f ie ld v a lid a tio n o f th e b io m a r k e r s d e s c r ib e d
in th is r e v ie w is a s tr o n g r e s e a r c h n e e d f o r th e fu tu r e .
C O N C L U S IO N
T h is r e v ie w c le a r ly d e m o n s t r a te s th e e c o l o g ic a l r e le v a n c e
a n d th e p o te n tia l u s e o f m y s id s h r im p a s te s t s p e c ie s f o r th e
M o r p h o lo g y a n d h i s t o l o g y
e v a l u a tio n o f e n v ir o n m e n ta l e n d o c r in e d is r u p tio n a n d a s a p o ­
M o r p h o lo g ic a l c h a n g e s r e s u l t i n g f r o m e x p o s u r e to c o n ta m ­
te n tia l s u r r o g a te f o r m a n y o th e r c r u s ta c e a n s . T h e h ig h ly s ta n ­
in a n ts h a v e b e e n d o c u m e n te d f o r m a n y ta x a , in c lu d in g a r­
d a r d iz e d u s e o f m y s id s in t o x ic ity t e s t in g is a n im p o r ta n t a d ­
th r o p o d s , b u t h a v e n o t b e e n c o n s id e r e d w id e ly in m y s id to x ­
v a n ta g e a n d r e s e a r c h s h o u ld b e d ir e c te d a t e v a l u a tin g th e c u r­
ic o lo g ic a l s tu d ie s a s a m e a s u r a b le e n d p o in t [1 3 ] . G e n tile e t al.
r e n t s t a n d a r d i z e d e n d p o in ts , s u c h a s s u r v iv a l, g r o w th , a n d r e ­
p r o d u c tio n , p r e f e r a b ly th r o u g h a n e n tir e lif e c y c l e , w ith a
[7 7 ] r e p o r te d m o r p h o l o g ic a l a b e r r a tio n s a t th e o n s e t o f s e x u a l
m a tu r ity in A . b a h ia a n d A. b i g e l o w i e x p o s e d to c a d m iu m in
n u m b e r o f e n d o c r in e d is r u p to r s . I n th i s c o n te x t, a n u m b e r o f
th e la b o r a to r y . In a d d itio n , fie ld o b s e r v a ti o n s o f in te r s e x u a lit y
r e f e r e n c e c h e m ic a ls , c h o s e n f o r t h e i r p o s s ib le m o d e o f a c tio n
a n d v a r ia b le te ls o n m o r p h o l o g y w e r e r e p o r te d in N . in te g e r
(i.e ., e c d y s o n e a g o n is t, e s tr o g e n a n ta g o n is t, j u v e n i l e h o r m o n e
f r o m d if f e r e n t E u r o p e a n e s tu a r i e s a n d th e B a l tic S e a [ 2 2 3 2 2 5 ], M o s t o f th e te ls o n d if f e r e n c e s m a y b e e x p la in e d b y r e ­
a g o n is ts , a n d o th e r s ) w a s p r o p o s e d b y D e F u r e t a l. [7 ] fo r
e v a l u a tin g r e la tiv e e n d p o in t s e n s i t i v i t y to p o te n tia l e n d o c r in e -
g e n e r a t io n o f p a r ts d a m a g e d b y p r e d a tio n a n d c a n n o t b e r e la te d
d is r u p tin g c o m p o u n d s . I n a d d itio n , e v id e n c e o f tr a n s g e n e r a -
d ir e c tly to p h y s io lo g ic a l p e r tu r b a tio n s d u r in g m o ltin g . S ti ll, a
tio n a l e f f e c ts h a s b e e n p u b lis h e d a n d p r e s e n tl y a tw o - g e n e r ­
g e n e t ic o r e p ig e n e tic b a s is c a n n o t b e r u le d o u t c o m p le te ly
a tio n lif e - c y c le p r o to c o l is b e in g in v e s t ig a te d w ith t h e s ta n d a r d
[2 2 5 ] . T h e d e g r e e o f f lu c tu a tin g a s y m m e tr y in m y s id s h a s b e e n
s p e c ie s A . b a h ia . H o w e v e r , a n e x te n s iv e l i s t o f n o n s ta n d a r ­
p r o p o s e d a s a q u a n tif ia b le m e a s u r e o f m o r p h o l o g ic a l a b e r r a ­
tio n s a n d is t h o u g h t to a r is e f r o m e n v ir o n m e n ta l o r g e n e tic
s tr e s s d u r in g d e v e l o p m e n t [1 3 ], B e c a u s e t h e r e s u lts fr o m e a r ­
li e r s tu d ie s o n m o r p h o l o g ic a l a b e r r a tio n s c o u ld n o t g iv e a c le a r
d iz e d e n d p o in ts h a s b e e n p u b lis h e d a n d s h o u ld b e in v e s t ig a te d
fu r th e r . S o m e o f th e s e e n d p o in ts , s u c h a s d is r u p tio n o f e c ­
d y s te r o id m e ta b o lis m a n d e m b r y o n ic d e v e l o p m e n t, m ig h t d if­
f e r e n tia te f o r in v e r te b r a te - s p e c if i c e f f e c ts o f c h e m ic a ls . T h e
m e c h a n is tic e x p la n a tio n f o r th e o b s e r v e d e f f e c ts , p r e lim in a r y
s e le c tio n o f w h ic h m y s id s p e c ie s to u s e w ill b e a b a la n c e o f
s tu d ie s e x a m in in g d if f e r e n t p o te n tia l c h a r a c te r is tic s w o u ld firs t
its e c o lo g ic a l r e le v a n c e a n d its e a s e o f u s e f o r m e a s u r i n g th e
h a v e to b e p e r f o r m e d in m y s id s , b e f o r e f u r th e r c o n s id e r i n g
s e le c te d e n d p o in ts . C le a r ly , th e a m o u n t o f a v a i la b le in fo r­
th is e n d p o in t.
m a tio n a n d th e r e l a t i v e l y s h o r t lif e c y c l e o f A . b a h i a f a v o r th e
u s e o f th is s p e c ie s , b u t its n a r r o w s a li n ity a n d te m p e r a tu r e
r a n g e l i m i t its u s e in c o ld e r w a t e r o r lo w - s a lin ity te s tin g . V a r­
B e h a v i o r a l a n d o t h e r e n d p o in ts
io u s o th e r m y s id s p e c ie s a r e p r o p o s e d in th is r e v ie w , to g e th e r
D is r u p tio n o f m y s id s w i m m in g a n d p o s itio n m a in te n a n c e
b e h a v i o r h a s b e e n in v e s t ig a te d in l a b o r a to r y s tu d ie s w ith N.
w ith a lis t o f p o te n tia l e n d p o in ts to e v a l u a te th e e f f e c ts o f
e n d o c r in e d is r u p to r s in th e s e a n im a ls . T h e s e s h o u ld s tim u la te
i n t e g e r e x p o s e d to s u b le t h a l c o n c e n tr a tio n s o f c h lo r p y r i f o s (an
th e s c ie n tif ic c o m m u n ity to e x p lo r e f u r th e r th e u s e o f m y s id
o r g a n o p h o s p o r o u s p e s ti c id e ) a n d
s h r im p a s a n in v e r te b r a te m o d e l f o r th e e v a l u a tio n o f e n v i­
c a d m iu m
[ 2 2 6 - 2 3 0 ] , A l­
th o u g h th e m o d e o f a c tio n o f th e to x ic a n t o n s w i m m in g r e ­
ro n m e n ta l e n d o c r in e d is r u p tio n .
m a in s u n k n o w n , t h e a u th o r s s p e c u la te d th a t th e d is r u p tio n in
c h lo r p y r i f o s - e x p o s e d m y s id s w a s p r o b a b ly d u e to th e in h ib i­
to r y a c tio n o n a c e ty lc h o lin e s te r a s e . I n a d d itio n , C r ip e e t al.
[ 2 3 1 ] r e p o r te d a r e d u c tio n in th e m a x im u m s u s ta in e d s w i m ­
A ck n o w led g e m e n t— T h is stu d y w as su p p o rte d b y a g ran t from the
F lem ish In stitu te for th e P rom otion o f Innovation b y S cience and
Technology to T.A. V erslycke.
m in g s p e e d o f A . b a h i a a f te r e x p o s u r e to s u b le t h a l le v e ls o f
tw o p e s ti c id e s . O th e r a u th o r s h a v e in v e s t ig a te d t h e s w a r m in g
b e h a v i o r o f m y s id s in l a b o r a to r y o r f ie ld s tu d ie s [ 2 3 2 - 2 3 8 ] ,
F o r m y s id s , d is r u p tio n o f s w i m m in g a n d s w a r m in g b e h a v io r
m a y le a d to in c r e a s e d p r e d a tio n o r d is p la c e m e n t f r o m o p tim u m
s ite s in th e e s tu a r y [4 3 ],
O th e r b e h a v io r a l r e s p o n s e s th a t h a v e b e e n m e a s u r e d in
m y s id s in c lu d e f e e d in g a c t iv ity [ 3 9 , 2 3 9 - 2 4 2 ] , g r o o m i n g b e ­
h a v io r [2 4 3 ] , b u r r o w i n g a b ility [2 4 4 ] a n d p r e d a t o r - p r e y d y ­
n a m ic s [2 4 5 ] . T h e u s e o f b e h a v i o r a l r e s p o n s e s a s a m o n ito r in g
to o l, h o w e v e r , h a s littl e u tilit y u n le s s b e h a v i o r a l c h a n g e s a re
u n d e r s to o d w ith in a n e c o lo g ic a l c o n te x t; th a t is , h o w w e ll th e
p a tte r n s a re u n d e r s to o d w ith in th e c o n te x t o f a n a n i m a l ’s n a t ­
u r a l lif e h a b its a n d e c o lo g ic a l r e q u ir e m e n ts [2 4 6 ] a n d i f th e
c h a n g e s c a n b e r e la te d c le a r ly to in te r n a l r e s id u e le v e ls or
e n v ir o n m e n ta l le v e ls o f s p e c if ic c o n ta m in a n t s [2 4 7 ],
S e v e r a l s tu d ie s h a v e b e e n p u b lis h e d o n o s m o tic re g u la tio n
in m y s id s (W e b b e t al. [2 4 8 ] a n d r e f e r e n c e s th e r e in ) a n d th e
i n te r a c tio n b e tw e e n o s m o r e g u la tio n a n d c h e m ic a l e x p o s u r e
[ 6 6 ,2 4 9 ] . O th e r h o r m o n a l r e s p o n s e s a n d d is tu r b a n c e s in c r u s ­
t a c e a n s , s u c h a s c o lo r c h a n g e s ( o n e o f th e e a r lie s t s tu d ie d
p h e n o m e n a th a t p r o v id e d d e f in ite p r o o f o f a h o r m o n e - m e d ia te d
p r o c e s s in a c r u s ta c e a n ) , r e tin a l p ig m e n ts , a n d lim b r e g e n e r ­
a tio n a re d is c u s s e d in a r e v ie w b y F in g e r m a n e t a l. [8 ], H o w ­
e v e r, th e u s e o f th e s e e n d p o in ts in m y s id s a w a it s f u r t h e r s tu d y .
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