1984 lectin-induced agglutination of trophozoites of different species and strains of entamoeba

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  • 8/4/2019 1984 Lectin-Induced Agglutination of Trophozoites of Different Species and Strains of Entamoeba

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    Z e i t s c h r i f t f~irZ Parsitenkd (1984)70:147-152 P a r a s i t e n k u n d e

    P a r a s i t o l o g y R e s e a r c h,9 Springer-Verlag 1984

    Lectin-induced agglutination of tropho zoitesof different species and strains of EntamoebaE . G h a d i r i a n * a n d E . M e e r o v i t c hInstitute of Parasitology, Macdonald Campus of McGill University,21, 111 Lakeshore Road, Ste-Anne-de-Bellevue Que., Canada H9X 1CO

    A b s t r a c t . I n v i t r o a g g l u t i n a b i l i t y o f t r o p h o z o i t e s o f t h r e e E n t a m o e b ah i s t o l y t i c a s t r a i n s , c u l t i v a t e d u n d e r a x e n i c c o n d i t i o n s i n t h e p r e s e n c eo f c o n c a n a v a l i n A ( C o n A ) , w a s s h o w n t o b e re l a t e d t o t h e d e g re e o ft h e ir p a t h o g e n i c i t y fo r e x p e r i m e n t a l a n im a l s a n d o f t h e c o n c e n t r a t i o no f C o n A . S e v e n s tr a in s o f E . i n v a d e n s t e s t e d a l s o a g g l u t i n a t e d i n t h ep r e se n c e o f C o n A , a n d t h e d e g re e o f a g g l u t i n a ti o n w a s p r o p o r t i o n a lt o t h e c o n c e n t r a t i o n o f t h e le c ti n . T h r e e s t r a in s o f E . h i s t o l y t i c a - l i k e" L a r e d o - t y p e " a m e b a e , a s t r a i n o f E . t e r r a p i n a e , a n d a s t r a i n o fE . m o s h k o v s k i i a g g l u t i n a t e d v e r y s l i g h t l y , o n l y i n t h e p r e s e n c e o f t h eh i g h e s t c o n c e n t r a t i o n o f C o n A t es te d .

    Introduct ionO f a ll t h e s p e ci es o f t h e g e n u s E n t a m o e b a , o n l y E . h i s t o l y t i c a a n d E . i n v a d e n sa r e k n o w n t o b e p o t e n t i a l l y p a t h o g e n i c f o r t h e ir h o s ts . W h i l e al l t h e s t r a in so f E . i n v a d e n s a r e e q u a l l y p a t h o g e n i c f o r c a r n i v o r o u s r e p t i l e s ( S t e c k 1 9 6 3 ;B ak er 1 9 6 5 ) , s t r a i n s o f E . h i s t o l y t i c a d i f f e r i n t h e i r p a t h o g e n i c i t y f o r m a n .A t t e n u a t i o n o f E . h i s t o l y t i c a s t r a i n s a f t e r p r o l o n g e d c u l t i v a t i o n i n v i t r o ,e s p e c i a l l y u n d e r a x e n i c c o n d i t i o n s , m a y b e d u e t o s e v e r a l f a c t o r s , a s y e tn o t f u ll y u n d e r s t o o d . A g o o d c o r r e l a ti o n h a s b e e n f o u n d b e t w e e n t h e p a t h o -g e n i c i ty o f s o m e s t ra i n s o f E . h i s t o l y t i c a f o r m a n a n d t h e ir a b il it y to p r o d u c ec e c a l u l c e r s i n r a t s ( M i z g i r e v a 1 9 6 6 ) . O n t h e o t h e r h a n d , B o s a n d H a g e(1 9 7 5 ) fo u n d t h a t a l l s t r a i n s o f E . h i s t o I y t i c a t e s te d b y t h e m , w h e t h e r f r o mp a t i e n t s o r c a r r ie r s , w e r e p a t h o g e n i c w h e n i n o c u l a t e d in t o h a m s t e r l iv e r.W h i l e t h e p r e c i s e r e a s o n f o r t h e e x p r e s s i o n o f t h e p a t h o g e n i c p o t e n t i a lb y E . h i s t o l y t i c a ( a n d b y E . i n v a d e n s ) is n o t k n o w n , m a n y f a c t or s a c c o m p a -n y i n g t h e p a t h o g e n i c ( i n v a s i v e ) p r o c e s s h a v e b e e n i d e n t i f i e d . A m o n g t h e s ea r e t h e o x i d a t i o n - r e d u c t i o n p o t e n t i a l a t t h e in t e s t in a l m u c o s a l s u r fa c e* P r e s e n t a d d r e s a n d o f f p r i n t r e q u e s t s t o." Montreal General Hospital Research Institute 1650Cedar Avenue, Montreal, Que., Canada H3G 1A4

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    148 E. Gh adir ian and E. M eerovi tchTable 1. Strains of Entamoeba histolytica a n d o f E. histolytiea-like strains used in the presents tudySpecies Strains ReferencesEntam oeba histolytica IP-1062 0 0 : N I HD K BEntam oeba histolytica- L A ( L a r ed o )li ke " L a r e d o - t y p e " H u f fJAEntamoeba invadens P ZIP-1IP-2B NSiSSivL165 ~Entam oeba terrapinae MEntamoeba mo shkovskii F I C

    Ghad i r ian an d M eerov i tch (1978)Ghad i r ian and M eerov i tch (1979)Deschiens (1937)Drey er (1961)Beaver et al. (1956)En tne r and M os t (1965)Meerovi tch (1958)Meerovi tch (1958)Meerovi tch (1958)M cConn ach ie (1958)G ray e t a l. (1966)G ray e t a l. (1966)Diamond (1960)Meerovi tch (1958)Lachance (1959)

    a Originally identified as E. terrapinae by Diamond (1960)

    ( M e e r o v i t c h e t a l. 1 97 6 ), t h e c o m p o s i t i o n o f t h e c o n c o m i t a n t b a c t e r i a l f l o r ai n t h e i n t e s t i n e ( P h i l li p s e t a l. 1 9 5 8 ) , d i e t o f t h e h o s t ( R o s s a n d K n i g h t1 9 73 ) , a n d t h e s u r f a c e m e m b r a n e p r o p e r t i e s o f t h e a m e b a e t h e m s e l v e s . S e v -e r a l s t u d i e s h a v e s h o w n t h a t t r o p h o z o i t e s o f p a t h o g e n i c s t r a i n s o f E . h i s t o l y -t i ca a g g l u t i n a t e d i n t h e p r e s e n c e o f r e l a t i v e ly l o w c o n c e n t r a t i o n s o f t h el ec ti n , c o n c a n a v a l i n A ( C o n A ) , w h i le a t t e n u a t e d o r n o n - p a t h o g e n i c s t ra i n so f E . h i s t o l y t i c a a n d o f o th e r E n t a m o e b a s p ec ie s d i d n o t ( M a r t i n e z - P a l o m oe t a l. 1 9 7 3 ; T r i s s l et a l. 1 9 7 7 ; B o s a n d v a n d e G r i e n d 1 9 7 7 ; D a s 1 9 7 7 ).T h e p r e s e n t r e p o r t d es c r ib e s a d d i t i o n a l o b s e r v a t i o n s o n th e a g g l u t i n a b i l i t yo f t r o p h o z o i t e s o f se v e ra l s t ra i n s o f E . h is t o l y t ic a , E . h i s t o l y t i c a - l i k e " L a r -e d o - t y p e " a m e b a e , E . i n v a d e n s , E . t e r r a p i n a e , a n d E . m o s h k o v s k i i i n t h ep r e s e n c e o f C o n A .

    M a t e r i a l s a n d m e t h o d sTro phoz oites f rom 3-day-old axenic cul tures of E. h istolytica growing in the TPS-1 med ium(D iam ond 1968) a t 37 ~ C an d fr om 8-day-old axenic cu ltures of the o the r species of Entamoebagrow ing in the TY I-S-33 (D iam ond e t a l . 19 78) a t 25 ~ C were used in the exper iments. Th estra ins of these ame bae tha t w ere employed are shown in Table 1 .The am ebae were washed three t imes with phosp hate-buffered sa line, pH 7 .2 (PBS) bycentr i fugat ion a t 200 g fo r 7 ra in . Their nu m ber in the f ina l sediment w as determined withthe a id of a hem ocytom eter , then they were resuspended in PBS to a f ina l con centra t io nof 5 x 105 ce lls /ml. T he " L ar ed o- typ e" s t ra in L A amebae, because o f the ir small s ize , wereresuspended to a co nce ntra tion o f 2 x 106 cells/ml. By the tryp an blu e exclusion test, the viabi-l ity of the organisms was es timated a t ~ 97%.Solu t ion o f the lect in , C on A (Sigma, St . Louis , M issouri) was prepared in PBS. Dilu t ionsof 5 , 20, and 40 gg/m l were prepa red in th is so lvent for use in the exper iments.

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    Con A effect on Entamoeba spp. 149Table 2. Agglutinability of trophozoites of different strains of E. histolytica by concanavalin AStrain Source and year Agglutination with Co n A

    of isolation (gg/ml) ~2.5 10 20

    IP-106 Am ebic dysentery, 1965 2 3 4200 : N IH Am ebic dysentery, 1949 1 2 3D K B Am ebic dysentery, 1924 0 0 1See Materials and Methods for the scoring system

    The agglutination reactions were carried out in Tissue Culture Chamber slides (Lab-TekProducts, Miles Labo ratories , Illinois) according to a modification of the me thod o f Martinez-Palo m o et al. (1973). Volum es o f 0.1 ml of suspensions of all species contained 5 x 104 cells,except for the strain LA am eba e which were used at a concentration of 2 x 105 cetls/0A ml.The suspensions of am ebae were mixed with equal volumes o f Con A in PB S. Thus, theconcentrations of Co n A used in the reactions were 2.5, ~0, and 20 !zg/ml. Th e cham ber slideswere agitated for 5 min, then incubated for 30 rain with occasional shaking - E. histolyticaat 37~ and the oth er ameb ae at 25 ~ C. At the end o f the incubation period, the cham berslides were examined u nde r low po we r in an inverted microscope. T he degree of agglutinationwas recorded on a scale from 0 to 4, according to the following criteria: 0, no clumps ofagglutinated am eba e observed; l, few clumps o f 4~10 am eba e; 2, larger clumps o f 10-20am ebae; 3, ~ 70 % of am ebae in large clumps of 20-60; 4, almost 90% of amebae in verylarge clumps of > 60.

    All the tests were repeated five times, the results representing the averages of five observa-tions.R e s u l t s a n d d i sc u s s i o nT a b l e 2 sh o w s t h e r es u l t s o f e x p o s i n g t h e t r o p h o z o i t e s o f th e E . h i s t o l y t i c as t r a i n s to t h e t h r e e c o n c e n t r a t i o n s o f C o n A . I t is e v i d e n t t h a t t h e a g g l u t i n a -t i o n l ev e l o f t h e h i g h l y p a t h o g e n i c s t r a in , I P - 1 0 6 , i n c r e a s e d p r o g r e s s i v e l yi n t h e p r e s e n c e o f in c r e a s in g c o n c e n t r a t i o n s o f C o n A . T h e 2 0 0 : N 1 H s t ra i nc el ls a g g l u t i n a t e d s o m e w h a t l es s, r e a c h i n g t h e s co r e o f 3 a t C o n A c o n c e n t r a -t i o n o f 2 0 b tg /m l . S t r a i n D K B , o r i g i n a ll y , b u t n o l o n g e r p a t h o g e n i c , a g g l u t i -n a t e d o n l y t o th e s c o re o f i w h e n m i x e d w i t h t h e h i g h e st c o n c e n t r a t i o no f C o n A .

    T a b l e 3 s u m m a r i z e s t h e r es u l ts o f te s ts w i t h t h e r e m a i n i n g s p ec ie s o ft h e E . h i s t o l y t i c a t y p e . T h e s t r a i n s o f E . i n v a d e n s , e x c e p t f o r 1 65 a n d S i v L ,a g g l u t i n a t e d i n th e p r e s e n c e o f 2 .5 l -tg /m l o f C o n A ; t h e d e g r e e o f a g g l u t i n a -t io n w a s p r o p o r t i o n a l t o th e c o n c e n t r a t i o n o f t h e le c ti n. O f th e t h r e e s t ra i n so f E . h i s t o l y t i c a - I i k e , " L a r e d o - t y p e " a m e b a e , o n l y t w o ( H u f f a n d J A ) a g g lu -t i n a t e d s l i g h lt y w i t h 2 0 g g / m l o f C o n A . E n t a m o e b a t e r r a p i n a e s t r a i n Ma n d E . m o s h k o v s k i i s t r a in F I C r e a c te d v e r y w e a k l y ( < 1 ) o n l y w h e n t h eh i g h e s t c o n c e n t r a t i o n , 2 0 g g / m l , o f C o n A w a s e m p l o y e d .O u r r e s u lt s c o n f i r m t h o s e o f s e v e ra l p r e v i o u s w o r k e r s . T h u s , M a r t i n e z -P a l a m o e t al . ( 1 9 73 ) f o u n d t h a t in v a s i v e s t r a i n s o f E . h i s t o l y t i c a w e r e a g g l u t i -n a t e d s t r o n g l y b y C o n A , w h i l e n o n i n v a s i v e s t ra i n s a g g l u t i n a t e d s l i g h tl yo r n o t a t al l. T h e y r e l a t e d a g g l u t i n a t i o n t o a c l u s t e r i n g o f s p e c if i c r e c e p t o r s

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    1 50 E . G h a d i r i a n a n d E . M e e r o v i t c hT a b l e 3 . A g g l u t i n a b i l i t y o f a m e b i c t r o p h o z o i t e s o f sp e c ie s o t h e r t h a n E. h i s to ly t i ea b y c o n c a n a v -a l i n A aS p e c ie s a n d s t r a i n S o u r c e a n d y e a r A g g l u t i n a t i o n w i t h C o n A

    o f i s o l a t i o n ( l~ g / m l) a2 .5 1 0 2 0

    E. invadensP Z S n a k e , 1 9 4 7 1 3 4I P - 1 S n a k e , 1 9 5 2 1 2 3I P - 2 T u r t l e , 1 9 5 5 1 2 31 6 5 T u r t l e , 1 9 4 7 0 1 2B M S n a k e , 1 9 53 1 2 2S i s L i z a r d , 1 9 6 4 1 2 3S i v L L i z a r d , 1 9 6 4 0 1 2" L a r e d o - t y p e "L A H u m a n , 1 96 1 0 0 0H u f f H u m a n , 1 95 6 0 0 1J A H u m a n , 1 96 5 0 0 1

    E. terrapinaeM T u r t l e , 1 9 5 7 0 0 < 1

    E . m o s h k o v s k i iF I C S e w a g e , 1 9 5 9 0 0 < 1

    a S e e M a t e r i a l s a n d M e t h o d s f o r t h e s c o r i n g s y s t e m

    f o r C o n A o n t h e c ell m e m b r a n e o f t h e a m e b a e a n d s u g ge s te d t h a t v ir u le n c eis a f u n c t i o n o f s u r f a c e p r o p e r t i e s . T h e r e s u l t s o f D a s ( 1 97 7 ) w e r e s i m i l a rt o o u r s , i n t h a t t r o p h o z o i t e s o f i n v a s i v e s t r a i n s o f E. h i s to ly t ica a n d E. inva-dens w e r e a g g l u t i n a t e d i n t h e p r e se n c e o f C o n A , a n d t h o s e o f n o n i n v a s i v es t ra i n s o f th e f o r m e r d i d n o t .

    T r i ss l e t a l. ( 1 97 7 ) d e t e r m i n e d t h a t p a t h o g e n i c s t r a i n s o f E. h i s to ly t icap o s se s s s e v e r a l c e ll su r f a c e p r o p e r t i e s t h a t d i f f e r f r o m t h o s e o f a m e b a e b e -l o n g i n g t o s p e c ie s w h i c h a r e e i t h e r n o n p a t h o g e n i c o r n o t i n fe c t iv e f o r h u -m a n s . T h e s e w o r k e r s ' i n t e r p r e t a t i o n o f s u sc e p t ib i li ty o f a m e b a e t o a g g l u t in -a t e w i t h C o n A r e l a t e d th e p r e s e n c e o n th e i r s u rf a c e s o f s tr o n g a n d u n i f o r mC o n A - b i n d i n g s i te s a n d a l a c k o f d e t e c t a b l e r e p u ls i v e c h a r g e s . O u r r e s ul tsd i f f e r f r o m t h o s e o f T r is s l e t al . w i t h r e g a r d t o E. invadens s t r a in P Z . F a i l u r eo f th i s s tr a i n t o a g g l u t i n a t e s t r o n g l y w i t h C o n A r e p o r t e d b y t h e s e w o r k e r s ,a n d s t r o n g a g g l u t i n a t i o n o b s e r v e d b y u s , m a y b e d u e t o t h e f a c t t h a t t h e yu s e d a c l o n e - d e r i v e d c u l t u r e a n d w e e m p l o y e d t h e p a r e n t s tr a in . I t is c o n c e iv -a b l e t h a t t h e i r c lo n e c u l t u r e w a s d e r i v e d f r o m a n a m e b a w i t h c h a r a c te r i s ti c su n l i k e t h o s e o f t y p ic a l E. invadens, i .e ., t h e P Z s t r a i n is a m i x t u r e o f o r g a n -i s m s w i t h d i f f e r e n t b i o l o g i c a l p r o p e r t i e s . H o w e v e r , t h e p o s s i b i li t y o f v a r i a -t i o n d e p e n d i n g u p o n c o n d i t i o n s p r e v a i li n g in c u l t u re , w i t h o r w i t h o u t s e le c -t i o n , s h o u l d a l s o b e c o n s i d e r e d . O u r r e s u l t s w i t h r e s p e c t t o t h e " L a r e d o -t y p e " a m e b a e a n d E . m o s h k o v s k i i a r e in a g r e e m e n t w i t h t h o s e o f T ri ss l

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    C o n A e f f e c t o n E n t a m o e b a spp. 151e t a l . ; i n ad d i t i o n , E . t e r r a p i n a e , a c o m m e n s a l o f tu r t le s , w h i c h i s a l s o in f e c -t iv e b u t n o t p a t h o g e n i c f o r s n a k e s, r e a c ts w i t h C o n A i n th e s a m e w a ya s th e " L a r e d o - t y p e " a m e b a e a n d E . m o s h k o v s k i i .A c k n o w l e d g e m e n t s . T h i s w o r k w a s s u p p o r t e d b y a N a t u r a l S c ie n c es a n d E n g i n e e r in g R e s e a r c hC o u n c i l g r a n t to E . M e e r o v i t c h a n d a M c C o n n e l l s c h ol a r sh i p t o E . G h a d i r i a n . R e s e a r c ha t th e I n s t i t u t e o f P a r a s i t o l o g y is s u p p o r t e d b y t h e N a t i o n a l S c i e n c e a n d E n g i n e e r i n g R e s e a r c hC o u n c i l o f C a n a d a a n d t h e F o n d s F C A C p o u r l ' a i d e e t l e s o u t i e n 5L l a r e c h e r c h e .

    Re f e r enc e sB a r k e r D C ( 19 65 ) T h e i n c i d e n c e o f Entam oeba invadens i n a n a t u r a l p o p u l a t i o n o f r e p ti l e s

    i n s o u t h w e s t M a d a g s c a r . P r o g r e s s i n P r o t o z o o l o g y , E x c e r p t a M e d i c a I n t e r n a t i o n a l C o n -g res s Se r i e s N o . 9 1 , p . 1 9 5B e a v e r P C , J u n g R C , S h e r m a n H J , R e a d T R , R o b i n s o n T A ( 19 56 ) E x p e r i m e n t a l E n t a m o e b ahis to ly t ica i n fe c t io n s in m a n . A m J T r o p M e d H y g 5 : 1 0 0 0 - 1 0 0 9B o s H J , H a g e A J ( 1 97 5 ) V i r u l e n c e o f b a c t e r i a - a s s o c i a t e d , Crith id ia-associa ted , a n d a x e n i cEntam oeba h i s to ly t i ca : e x p e r i m e n t a l h a m s t e r l i v e r i n f e c t i o n s w i t h s t r a in s f r o m p a t i e n t sa n d c a r r i e r s . Z P a r a s i t e n k d 4 7 : 7 9 - 8 9B o s H J , v a n d e G r i e n d R J ( 1 97 7) V i r u l e n c e a n d t o x i c i t y o f a x e n i c Entam oeba h i s to ly t i ca .N a t u r e 2 6 5 : 3 4 1 - 3 4 3D a s S R (1 97 7) C o n c a n a v a l i n A - i n d u c e d a g g l u t in a t i o n i n p a t h o g e n i c a n d n o n - p a t h o g e n i c a n a e r -o b i c a n d a e r o b i c a m o e b a e . I n t J P a r a s i t o l 1 : 5 7D e s c h i e n s R ( 1 9 3 7 ) C o n s i d e r a t i o n s s u r l ' a c t i o n p a t h o g ~ n e d ' u n e s o u c h e d ' a m i b e s d y s e n t e r i q u e s( S o u c h e : C D o b e l l D . K . B . ) B u l l S o c P a t h o l E x o t F i li a l e s 3 0 : 5 6 2 - 5 6 4D i a m o n d L S ( 1 96 0) T h e a x e n i c c u l t i v a t i o n o f t w o r e p t i l i a n p a r a s it e s , Entamoeba terrapinae

    San d e r s & C l ev e l an d , 1 9 3 0 , an d E. invadens R o d h a i n 1 9 3 4 . J P a r a s i t o l 4 6 : 4 8 4D i a m o n d L S ( 19 68 ) T e c h n i q u e s o f a x e n ic c u l ti v a t io n o f Entam oeba h i s to ly t i ca S h a u d i n n , 1 9 0 3a n d E. h is to ly t ica- l ike a m o e b a e . J P a r a s i t o l 5 4 : 1 0 4 7 - 1 0 5 6D i a m o n d L S , H a r l o w D R , C u n n i c k C C ( 19 78 ) A n e w m e d i u m f o r t h e a x e n i c c u l t iv a t i o no f Entam oeba h i s to ly t i ca a n d o t h e r E n t a m o e b a . T r a n s R S o c T r o p M e d H y g 7 2 : 4 3 1 - 4 3 2D r e y e r D A (1 96 1) G r o w t h o f a s tr a i n o f Entam oeba h i s to ly t i ca a t r o o m t e m p e r a t u r e . T e x a sR e p B i o l M e d 1 9 : 3 9 3 3 96E n t n e r N , M o s t H ( 19 65 ) G e n e t i c s o f E n t a m o e b a : c h a r a c t e r i z a t i o n o f t w o n e w p a r a s i t ic s t r a i n sw h i c h g r o w a t r o o m t e m p e r a t u r e ( a n d a t 3 7 ~ C ) . J P r o t o z o o l 1 2 : 1 0 - 1 3G h a d i r i a n E , M e e r o v i t c h E ( 19 78 ) B e h a v i o r o f a x e n i c IP - 1 0 6 s t r a in o f Entam oeba h i s to ly t i eai n t h e g o ld e n h a m s t er . A m J T r o p M e d H y g 2 7 : 2 4 1 - 2 4 7G h a r d i r i a n E , M e e r o v i t c h E (1 97 9) P a t h o g e n i c i t y o f a x e n ic a l l y c u l t i v a t e d Entam oeba h i s to ly -tica, s t r a i n 2 0 0 : N I H , i n t h e h a m s t e r . J P a r a s i t o l 6 5 : 7 6 8 - 7 7 1G r a y C W , M a r c u s L C , M c C a r t e n W C , S a p p i n g t o n T (1 96 6) A m e b i a s is i n t h e K o m o d o D r a g o n ,Var anus kom odoens i s . I n t Z o o Y e a r B o o k 6 : 2 7 9 - 2 8 3L a c h a n c e P J ( 1 9 5 9 ) A C a n a d i a n s t r a i n o f E n t a m o e b a m o s h k o v s k i i C h a l a i a , 1 9 4 1 . C a n J Z o o l3 7 : 4 1 5 - 4 1 7M a r t i n e z - P a l o m o A , G o n z a l e z - R o b l e s A , d e l a T o r r e M (1 97 3) S e l e ct iv e a g g l u t i n a ti o n o fp a t h o g e n i c s t r a i n s o f Entam oeba h i s to ly t i ca i n d u c e d b y C o n A . N a t u r e ( N e w B i o lo g y )2 4 5 : 1 8 6 1 8 7M c C o n n a c h i e E (1 95 8) F a c t o r s a f f e c t i n g t h e g r o w t h a n d b e h a v i o u r o f a x e n ic s t r a in s o f E n t a -moe ba invadens R o d h a i n , 1 93 4. P a r a s i t o l o g y 4 8 : 4 2 3 - 4 3 2M e e r o v i t c h E (1 95 8) S o m e b i o lo g i c a l r e q u i r e m e n t s a n d h o s t - p a r a s i t e r e l a t i o n s o f E n t a m o e b ainvadens. C a n J Z o o l 3 6 : 5 1 3 5 23M e e r o v i t c h E , E a t o n R D P , M c L a u g h l i n J (1 97 6) F u n c t i o n a l a s p ec t s o f h o s t - p a ra s i te r e l a ti o n si n a m o e b i a si s . P r o c e e d i n g s o f t h e I n t e r n a t i o n a l C o n f e r e n c e o n A m e b i a s i s , M e x i c o C i ty ,O c t o b e r 1 97 5, I n s t i t u t o M e x i c a n o d e l S e g n r o S o c i a l , p p 6 2 8 - 6 3 5

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    152 E. Ghadir ian and E. MeerovitchMizgireva MF (1966) Comparative vurulence of Entam oeba h i s to ly t i ca strains isolated frompatients and different groups of carriers. Med Parazitol Parazitar Bolez (Moscow)35 : 201-204Phillips BP, Wolfe PA, Bartgis IL (1958) Studies on the ameba-bacteria relationship in amebia-sis. II Some concepts in the etiology of the disease. Am J Trop Med Hyg 7 : 392-399Ross GW, Knight R (1973) Dietary factors affecting the pathogenicity of Entam oeba h i s to ly t icain rats. Trans R Soc Trop Med Hyg 67:560-567Steck F (1963) Die Ameobendysenterie der Reptilien. Acta Trop (Berl) 20:115-142Trissl D, Martinez-Palomo, Arguello C, de la Torre M, de la Hoz R (1977) Surface propertiesrelated to Concanavalin A-induced agglutination. A comparative study of several Enta -m o e b a strains. J Exp Med 145 : 65~665

    Accepted July 26, 1983