Biology of T Cruzi Annurev.mi.27.100173.002023 73

7/29/2019 Biology of T Cruzi Annurev.mi.27.100173.002023 73

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BIGY F 7RPNSM RUZ!

Zn Brener

Darano d arasoloa CB. , Unvrsdad dra d nas Gras adso d das ras Bo Horo Bras

ONTNTSINTRODUCTION ............... TRYPANOSOMA CRUZI-IFE CYCLE IN THE VERTEBRATE ....

ine Strtur of ruzi Bltram Fo 48The Signicance of cruzi Polymorphism ...... 5

cruzi-Inracular 5Hoss of crui ............. 34ato Inuning the Cyl in the Vtbte .... . ....... .. 3

TRYPANOSOMA CRUZI-IFE CYCE IN THE INVERTEBRATE ... Ft Whh M nun Dlpmnt in th Vt . ........ . . Sm Chaacttis f zi nftion in the nrtbrate. 3Deelopme rui i Aboal erbrae Ho ine Srctur f rui Delopmental Stage in the Vor . . . . . . ... 360

EVEOPMEN IN TISSUE CUTURE ....... A ian Embryo nfcton ........ 60Tiue Culture nftion ............ 60

Iuee of Temperature i Mhgenei 6nuence ofnteen .............................. 36Pune Mtabosm in rui ntellular o .. . . . . . . 362Biochmisry of rui-Harbong Ces .. .... . . . . . ..... . . 362xmntal Chmothay ....... 362in Stutur f u f rui . 363CUTIVATION OF TRYPANOSOMA CRUZI .... Nea Ded Mia: Grwth at ....... 364Grwh and Dierentiation 36s f Dr

A o Cutur o .. . 66DNA-MediatHeter-transfoation of Culture orms? 367Cultiatin f Amatigte Stag i Ce Mium ... 367nfvit rui Cultur ....... 368

TOHEMITR AND METAOISM ....... 69Nuclic Aid Role of Kinetoplast-DNA and Dykintoplasty in rui . 36

Nucarand Kinolas DNA frm rui ucle Aid Metabom in

rui ........

Amin A nd otin . ..... 37Lipid ... . ... .. 7Carbohydra Meabolism and Respiraion .

OTHER AET .......... Chmohrapy . ... 3 rui Conenital Transmiion .......... Antigen Contitution ....... 77Vanatin Againt crui ............... 37

t of378CONCUING REMARKS .......

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8

RUI

ypnsm ui the se of Chgs disese is n nbeten dvesy Immnologil o hemothepeti poteton is not vilble nd it is beoming lie

ote highe ypnosomtide fvote fo stdies intended to elidte homitohondil DNA oins in eglting eltions beteen ife in the invetebteveto nd in the mmmlin ost hns lgely to its esy ltivtion ndppently indenite etention of infetivity fo veto nd mmmlin hosts: it ismh moe menbe to epeiment mniption thn the svin typnosomes, espeilly those of the T uigop It shes ith ishmni menbility to nlysis of dption to intelllity u nde onditions not elldeed n fom in lte infetive metyli typomstigotes mophogenetipoess f moe dilt to eet ith slivin typnosoes his oining ofgent ptil nd timely biologil motives mes T u despite the hdsttending its se n eve moe popl ojet even mong oes not speillyonened ith pophylis nd thepy

TRYPANOSOMA LIE LE I E VERERE

T cruzi tiies disoios i te etete ost te stigote ite stges tily y iy diisio, e ge ito odiidig tostigotesi emege fo te psitied tisses ito te loodstem, ee tey iltefo vible peiod befoe penetting ells nd esming te yle In the te

phse, the ost hbos bndnt tisse nd boodstem foms in the oni phsepsites e se nd ndetetble in blood e o psitemi of the oniphse eslts fom bildp of stong immnity nd stble blne beteen hostnd psite Spontneos e seems e if it hppens t ll, sine blood psiteshve been deteted by indiet lboto methods in te igh popotion ofepeimentl nd honi hmn infetions

ne ruure Bre r

e mophology of uitypomstigotes by ligt miosopy nd ts polymopism e been m stdied (21 2 12 191. oe eet ide 0)

nd i et l 121 desibed te ne stte of uibloodstem foms ndtied to oelte mopologi ndings ith ognee fntion

uibloodstem foms hve, pposed to the ote sfe of the tilminyell membne, dise lmentos ot of vible thiness 500 A) 0) hih ppently filittes pinoytosis nd s lso been elted to theoene of the sessive sfe ntigens epoted in the elpsing bloodstempopltions of T ui 6). Bt fo ui no onvining evidene of shntiges o vition in the ose of infetion hs ppeed Sbpelil miotbles lying A beneth the sfe membne e obseved, eept in some

es of the esevoi poet A gop of miotbles sees in lose eltionshipith the bsl body nd in ppent ontinity it th sbplli iotbe0).

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OLOG OF PAOOMA C 9

Two tpcl bsl bods r ound n cruzi bloodsrm orms 0) Snctrpomstgots do not multpl n th bloodstrm th occurrnc o thos twoorgnlls sms not to ndct lrpcton ntcptng cll dson rthr thr rhps stgl structurs A pculrt unrportd or othr trpnosom

scs s h rsnc o cnrl lmns h ntr nd xtrcolsmtcrgons o th bsl bod crossng h trnsonl zon bwn th wo rnsrsllts h gllum hs th 9 + 2 strotcl xonm structur. A lttclccssor lmnous coml whn h mmbrn shh o h gllum longsd th xl lmnts ls n th sgmnt o th gllum ttchd to th surcmmbrn. Th gllum s bound o th pllcl b comlx o wo trlmnrmmbrns nd b dsmosomlk brous condnsons. A h conc surcth sublculr mcrotubuls o th gllum r thcnd mrdl 1, 0).

Th rsror round th bs o th gllum hs on bn consdrd to bn trpnosomtds st o srton nd!or xcrton s wl s st or ngstono mcromolculs b pnoctoss 198) psumbl n uz tromstgots thrsror cts th sm s suggstd b h bsnc o subllculr mcrotubulsundrlng prs o h rsror th dns dbrs nd doublwlld scls ttchd to th rsror ll nd th multsculr bods n th surroundngctolsm 11 10) So r ctostom hs not bn dctd n cruz bloodstrm tromstgots. A nw tubulommbrnous structur hs bn dscrbdormd b oston o to cll mmbrns whch rs rom n ngnon oh so ll dsls som chrcscs n common h cosomsuch s ts mrgnc rom th rsror wll nd ts culdsc tubulr shp 1 1).

or h dn o th to srucurs cnno b dnl sblshd ongto sm mrkd drncs th ubulommbrnous structur shos mcrotubulsonl t ts px t s lnd b two mmbrn unts nstd o sngl unt cllmmbrn nd scls do not urround ths structur.

A tpcl Golg complx ls btwn ntoplst nd nuclus mctons oth grnulr ndolsm rtculum r on ound clos to th Golg suggstngdrct communcton btwn both structurs nd scrtor ctt o th Golgrus n h romstgos 10) A Golg comlx s mor dlopd ndrsns mor scls n stou orms hn n slndr ons 11). Vckrmn 1)suggstd tht th ncrs n Golg comlx n brce brod orms rsults rom

locl ntgnntbod rcton lctd b th roducton o xontgns n cz such n nrnc cnno b md ds sngl so r unconrmdror on rls o solubl ntgn b bloodstrm orms 16).

crz ntolst morholog hs bn dtld 18 15) ts sz nd shm r durng dlomntl sgs robbl cusd b drncs n mounsoA nd rrngmn o A loos. n bloodstm tpomsgos th loopsr on rrngd n to lrs mrtng bstl sct Clo contctbtwn nuclus nd ntoplst dsntgrton o th mmbrns nd possblxchng o gnc mtrl btwn both orgnlls h bn dscrbd n crzcultur pmstgots but nr n bloodstrm orms 15) Slndr orms h rthr dlod ostor mtochondrl tubul ntmtl rltd to th longtsh o th rst's ostror nd hrs sou oms h shor nd lss

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350 BRENR

developed posto mtochondon A sngle tubula anto mtochondon xtendng om the knetoplast to the paast's anto end has bn obsevd hoccunc o bucatons a moe snuous cous outgowths and etculatonssuggests a ag mtochonda aa n stout T oms

Dnse ganua bods sla to th bacclom o ova bods n T beand ongolense (21) w also dscbd n boh slnd and sou . hosbaclom bods have ben suggested to coespond to xtamtochonda peoxsomelke oganelles contanng th glycophosphate oxdas gung n thtemnal espaton o slnde beoms whch lack a unctonal cytochoesystm. Mutvscula bodes omd by a vaable numbe o vescles smla tothose ound n th olg complx and nveloped by a dns amophous substancehav also ben poted (21) Those bods ognatd om condnsatons ovescs ound n cos assocaton wth th tubues o th olg compl, maypogess o h svo and phaps gu n scon anspot o th xtoOth dscbd vscls (121 10) sembl those poted by Hbet (94) nypaooma thee and ae consdeed o b ysosoms. ytochmcal tchnqueshav no howv ben usd on T o dcng spcc lysosoal acvtyThe occunc at th agla pockt o vscs cos to dns amophous matalstongly suggests pnocytoss n ths egon (121, 10)

aa et al 21esce oun eonate ctolasc otusons ounded ba mebane sla to the pelcle ovelyng the agellate claly connectd to theagllum and aglla mmban. hy sm tansn suctus smla o thlopoda n be (227. Th uncton s unclea thy have ben suggstd to

patcpate n adheson o the paastes to the substatum hpng thm to passthough apllay walls and to pentat ost lls (121) Flpodalke stutusn clos contact wth pp clls hav been epoted n an n vvo study o zn the lve o expemntally nected me 187. ompason o slnde and stoutoms o z showed lttle detctable vaatons n the shape and sze o thenuclus and deveopment o th ough endoplasmc etculum and olg compexn spt o th physologcal dences btwn thes oms (I2)

Some poblems ased by the ew epots on T n stuctu may dsvuh nvestgaton 8 chaacteaton o the baclo bods whch bea someanalogy wth the glycophosphatcontanng oganlls o beslnde

oms a unctonal glyceophosphat system n bloodstam oms wouldbe ath puzlng snce t s genally acceptd that cytochomemeated espaton pvals thoughout the l cycle (b the uncton o th lopoda and ocondtons avong devlopment o ths tansnt stuctues n the bloodsteamos ( the natue o he tubuloembanous stucue and s laonshp o hesevo hstochmcal chaactaton o th nyme actvty othe lysosomk bods and oth cytoplasmc ncusons and (e) studes on the possble localzaton o xoantgns on th suae coat o th bloodstam oms by tnconugatd antbody (216).

The Snne of u Pomophsmhagas (41 st potd dmophsm n bloodstam typomastgotes theoccunc o boh slnde and boad oms Many auhos hav snc studed hs

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OOG OF PAOOMA C inteestng mophoogic viins 12, 162, 19 Bene & ha 2 tudedseven deent stins of cu nd descbed, besides sende nd bod typomstigotes, vey stout fom not yet thooughy investigted Bene 2, studyingseve stins in expementy infected mice obseved tht etive

pecentges o the dieent foms in the boodstem shoed t est thee distbu-ion pens in some sns ende os pedominated fom onse of infectionto the anal death n othe tan lende o edonated dung the tday o neton but then gadualy dnhed whle the ooton o tout oteady eaed nally n two othe tan boad o edonated dung theente neton A la vaaton had been eoted by lva 191 oe oela-ton wa obeved beteen the edonane o eah o n the deent tannd some chcteistics of the expeiment infection, such s dution of peptentpeiod, couse of psitemi, nd motity tes 1 Anims noulated wthstins shoing pedominnty stout foms deveoped gduy scending psitemi the nims died ith psites teeming in the bood Stins usuy shoingpedominnty lende o had an eay hgh aatea ollowed by a hadecese in nmbe of psites and n ieguy o psitemi theee hosedeent atten evaled ae eght yea of antenane 2

he gnane o thee deent o ontoveal Chaga 1 uggetedthat lende and boad o gny exual doh Fo But howevelende tyoatgote eeented young aate whh gadually would develonto boad atue o eye Olvea 12, wong wth tue ultueneted wth uz eoted that the ealy ntaellula tyoatgote ae

boad and hot thee o ay ethe be beated by e ute o nteadhange nto ende tyoatgote when allowed to develo longe n oe e-nt hos ces ende foms e he s o ppe in the couse of expeimentinfections but e not usuy found in the boodstem of nims dispying cuied immunity But hen immunity is depessed, s in cotisoneteted nims,n ey med incese of sim foms is obseved 12 hese fcts my bepetinent to the suggestion tht sende foms of Aficn typnosomes e vintsginst hich ntibodies hve not yet been oduced hees the stumpy fomsoiginte fom sende ones by the ction ntbdes 9 uiboad ohoweve ae vey uney to ognate o ende tyoatgote togh hotantbode' ohogenet aton ne both o ae obeved n neted tueutue 12 1 1 2 1 1 ntbode ght neethee at a eet agent enatng the e entve lende o dung hon neton 2

t not yet known whethe eah o an yeld deendent ohologallyla to t aent The nse cn come ony fom investigtions ith conpopions Inocution of ony bod foms, isoted by micomnpution, hsinduced infections shoing sende nd bod bood typomstigotes 191 Eectsofenvionment upon phenotypic typomstigote chcteistics hve not been thooughy investigted empetue doe nuen the etive nume of the dieent

foms n tisse te t 26 slende foms gety incese hees t 7 stoutfoms pedomine 21 1 The deent ohologe o the blood tyoatgote aaently denote hy-

ologal deene When cu bood tyomstigotes e intvenousy inocu

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2 RENER

latd in nma mi nd m diappa m t bldtam witin a wu ud m ul y u m ul t bldm m dy wut p t t lld pply ptt l bt 2) mmu am ta

uy jtd d m pd tmy i t t mmumanm y dappd m bd wt 2 a nutn,wa u m uad many day 2) nma a, tan wna numb tut m dipayd i an nduin paaitmia wi ainwin pdmnantly ld m pd ly a li, anint paaimia26) a ny ut at lnd and u bld typmatit maydpay di ba n t at ad aud mmunp.

umb pt w dlp pmt t ttmd t

ppy pptinl t numb bd m n ntd bd mdnu d dlpmnt t dnt m in t nbat t 26 9 ) In up typnm t ld typmait dn dlp i t y , wn ppay t la an bi APanytm wi m ni uial in w inmn Cly ttumpy m build up a ytm ttan ym d dlp ba 4) Hw lid anay mdalnym aiity and piy ai i t mplily dit T typmati man t b tudid.

Pld mannan up typanm by yin paa ianiml d it mnmpi ad unabl t dlp in , pbabybau i lnd m t midial appaau dat pd5) utd n a t matial bwn nulu nd ipa, mt ly uin pimit ta wn t pimity bt anll i uid t p intpla A abl t d mitnd m b wuld p y t t pim d u t. Cydpmt bt bt d tb t m l il i wi tain plympim ad apaity i duiid a t la i

ya paa in abay animal (3.

u-IneuJ e

ypmi a nt a wid an t ll but mul and liam m pd Hw p p tu bld apila adpt ll y udd. Kllt 06) lwd i atpat myli ultu ypm in HL ll d tu wtd l pti d. bldm typmtia dubwld ul, multiul bd, d bm 2) bu

tud a lain a dtmi ll ptatin. pib l tani lpdiali tutu i atamnt bld pait t l a bdiud 2 87

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BIOLOGY OF TRPANOSOMA CRUZ 353

Aer penetratng the ce, trypomastgotes soon change nto amastgotes whchudergo bna ssion about very 12 hr 2, 1). I tssue cuture cels theumber of arasits th d of a cyc, whch usuay lasts 4to5 days, may varyfrm a ddg c z 128) Trafrmat rmas

gs sm bg am sas js wh h ce s ld wh amasgote forms, th who cc bg awas md to c u 2) Not aamagot eem cme t dv rmasgte; lectro mcroscoyof cu th myocardum of nocuatd anmas showed that 5% of thinracluar amastgotes wer cleary degnerated ) he normal amasgotesside the cel-induced ctoasmc lss a clear halo oe surroudd th parasit2) he capsuek sructure that somtmes sems to circe the intraceluaramasg sags fLshmania donovani179) has ot b obsrvd i cuNevrhss, i ssu cutur, c m wthstad arasism wihou aar

damage, at ea h ear age, ce he rea m ad e hw rmamoss. Paraszd cs howvr vua d ar ruur and parase rase(28 Kbr (05 suggsd ha rred psdocss berat a urotoxiwhch ma ac arb gag ces

ntracluar fusiform and roud amastigots hav ben descibd n tissu cuurcls 19). I the host h amastigote show rrguar shaped miochodria wthfew csae, vacuoles, rbosomes, and dense ipid bodies 16) Phagocytosed parasites show irrguarits membranes and kntoast odcatos he e strucur f h ktat-chdrim f cui th ect f fcted tuecuture has be dscbd 1) cyostom was dtected i itraceluar stagesf cuicg HLa cs 131) a reg dg c th srcur has be fud bdsram frms 121 30). Fre amasgtes, robab fmruurd cs th boodsram of hav fectd aimas, hav rortd17).

racuar morphognss and transformaon of amastgo stages no tromastgotes hav be suded i ifeced tssue culture ad i rtoal iqudreasd from cuar eemts i nfcd anmas 12 91225) doube rasformatio of amastgots to trpomastgote s belved to occur: the rst thfuifrm amastgts td ega alg th aglar e of grwh, ther keto

las mgrats to the aglat'S ostror ed, ad a sledr tromastgot s thdrc frmed th cd rce he rd amasg devl a vace earh toas his gows ad veua ruurs, gvg orgin hrough an ufodig mchaism to broad rmastgotes. uch a vacoe has aso b dscribdas a rarfac f h coasm (3) or a Vshad sg 5) h ufodgprocss and h presenc of such vacuol hav o, howvr, be detectd bycro mcrosco of tracuar stag. he rc of tca emagt sags he cs o he verbrat hos s dmads ivsgai. ccordigto Ekeles 59) eimastgotes are vry scarc vrtbrat host tssues, whch

trg gge a drect rafrma f rd frm t trmastgts emage ag ccrrg as a remar th sdr rmagsmorhogeess has, howver, b dscribd tssue cultur 191)



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H

u wde o ge bede e ou wd mamm belgg eeal ode h ae feed wh le ae eeal dome amal

a b bd b aually mally fd w gam a f m a 26 abb am 2 ad gua g. Ayal aue e followed by cho ogee myod 2 dlao ofw muu g 50 d euoggl e 202 may dued feed mce h eec e feco eemble e huma deae Teuefule of T uz feo of youg ad dul dog fo udyg hologalad eleocdoga hge d fo hemeaeu ud h bee e-oed 8 78. bu mokey ue e exemel cue dee d ewad de y myad 0 wee Rh mey dee afaal fe w elaely mld ue l 2

Amhba 2 ad bd 7 ae oed efay A ubawh lye uulue fm bu bldeam fm bee debed omal hcke eum 220 222 bu h o bee o fa lealy eled o hebd efoe Sou Ame d ( oduu uld be feedwh uz (4 whee Noh Amec Ghonotus lzad wee ealyinfeed y feedin e infeed ny wid ni widpedo he Ame oe e feed w gelle moolgally decal o f uma g. feo w hee ocalled uk yaome bee deeed oe mmmala ee fom eeal ode olleed

edem ea d e ely fee fom uma hag' deae eg omee of e Ued Sae. May of e a e bee oled d elydl e de wh oe of u fom um mohlgy f bldam fm f allula ag delm ulu lfe yl amd ad e . y awee oe le ule h um a me f hem eem allfe labay amal M f u a a y lly be ame li e yemac oo owee l ude dcuo ce hefey fo m would he o be aumed (49.

Factors InRuencing the Cye in the Vrtebratef e eba lealy ued by e aula T ua.Tee eg edee ae ome exee oo of deeula. The ua efed he lau ae uually lae fw g bably ludg eegeeou ula led by he ggaual aage of he aae bewee dee ho Neehele e udehoe cod he oe dly dguhg a d bable elee fo e o fom whc he ae e oled my odefo ce degee of homogeey of he oulo Thu dee ae of

blod yomgoe moology moly ae d oue of aema hebee eoed amal exemelly culaed w dee uz 8 Pedomaly myoo o euloo 0 a he

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55bn b fu Bn 01 obv m n n g ofl gnglon lon n m noul wh n n obl ol-on of hogn wh numb of nou bloo omgo h bnvw b hl 158 who how h wo nnll n n of

nfon m nu ong o h n. In h nfon of lowm n mo on m ng h nouum n hon, h ho h m l bu no vn b ogv uonn h numb of noul Anl om nhn n h uh om fo u o ouon, w m oun fo h n hvo In of n n ong ommun h ul mong h vou n noul n loo nml, n whn n mmunoogl u 01

h ho x 9 g 8 n gn onuon 6 m nun hou of u nfon n h vb Som uho hv b hnun of nvonmnl mu 108 1 n onl uuon0 on h o of T fo h of loo u n n nm k v hgh mu C owmu 8C ul n hgh m n mol h nmol of m nf wh uin wh hloomn h lobn bu o h hohm on of h ug 1 Nuon nhmn on vmn A n ln m ou hgh m nh n mo v lon, wh ovn n nohn n hno nun 8.

Infon, nu b ouln olnonolJ , -nl xb u nfon n xmnl nf m 2. n hoh hn, n moun of nfon obbl l b h nngllul nvolvmn hv bn n m 18

An o homon, l oon, hv ong n known o nhnT uiu nfon 8 lhough n unbl o hon nfon 9 Som hml n hl gn hv nl bn u o u h of mmunouon n xmnl Chg Chohmgull n m mol n v of mo n m n hu h 0 No of hon n n h hon h w A u of h of gmm on lohohmhon n moun h ou of hon Chg w ou n m n wh ou n n mm on nlohohm nu, onl n nml noul wh h C n, n uh h b oub of nn m omn b hgh mo-l. h u on h n ugg ng u b h omnn n h n of ou omgo whh f mo n o h hommun mnm 5 A no molul on n unb h hml mmunouv gn h ul nbo onnon

n mmunouon woul obbl b un o mnn, ub-n lvl h m of h n omnn omgo n buno h ou h ou fom of C n Ahon n ou



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36 r dd o a h modrg hro da a ama Nooa hmom d o dad aara of anod hghr arma ad horrvva of m oad wh T u rog ggg ha hmdd mm a a moa ro h fo (189)

TYPNOOM Z/ CYC N H NB

Bas fars of uidvom h vrbra ho Hmpt, Rdv-da hav og b kow (3 Da (3 wa rs o or a omrhvao of h dvom of u hmaohago raomd a a rahrhma h gd oodram form asg aog h bg dgvra hrogh rvrsb morhooga rasfomaos d sq ahag dvog a aa oo of h g h h omah mo oodromago oo hag o mago ad a fw o od form h h k of h aga oao o of av dvdg mago whh sor h aar ra fo a h rma a ooo of h masgo dra o fv maromago whh ar mad h fs aog wh masgos

hs dso ad ar agd


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L S 7

change thus reopenng he probem of yngamy n u discussed n some oChaga' eary paper. A prevou aemp o demonrae syngamy n u warepored by Amren (6) who fed traomds on cuure o wo derent drugrean ran recovered he ageaes rom he fce and cuvaed hem Bu

furher expermn faed to demonsrae resance o he cuure orm o odrug and tereore genetc ecange was no demonsraed A cuure form maynot e utale stages for knd of expement yngamy coud not e dentelyexcuded So far drug reance or oher genec marers have no ye een reporedn oodstream orms

Te ndng reported y Brener (7) on te oer and omewat reeme tepenomenon repored y Ader (2) who found ha under e acton of speccanterum T epmastgote tage cange nto esmanod wc agglutnateand form arge multnucleaed syncya maes he fuson of oogcal memranes

as descred may have more genera mpcaons han o a u cyce. Themecansm of ce adeon ce uon and urer omaon o poyaryocytesan eteroaryon y erent agent uc a yoectn or nacvated Sendavrus has een revewed by Pooe et a (6) who empaszed te mporance of theudy o expermentay nduced aeraon n e negry o poproen memranewhch precede ce uon

e eature n he omach o ratom aong wh he areay nown exenvemupcaon o epmagoe wc occur muaneouy n e nene ed ote uggeton tat ot ee of evens are eer ndependent phenomena poslyoccurrng as concomtan cyces or phas of the ame cycle Thee recen osea

on undercore e need for ctca reexamnaon o T deeopmen n enveerae ost noay mophogene of te early stages n te ood-contanngomach he reaonshp beween hese form and he mulplyng epmagoesfound n he mdgut and e orgn and morpogenes o e nctve tage

Facto Which May Infuene Developmnt n th Vtor

e o-pare eatonp n z veco a not ye een sasacorycared; dcuon abou condton whch aect ay to develop n he nverterate hos are necary raer pecuatve Some factor nuencng te cyce ne vecto are aeady nown and w deaed ere

Fuctuaton n necton rate of derent tratomd speces known o ave ngested arge numers of u bloodstream forms ave been repored qute oeneg e ame z ran neced 0 o PagyJ mg and ony7 ofRhod x ed on expermenay nfeced anmas (56) A tudyo the sensvty of deren spece o vecor n he xenodagnos of humancronc Cagas dseae owed a nymp of fa came 8 psveas compared wt ony oR. ox Apar from ths varyng capactyof the deren vectors n aowng fu deveopment of he paraste ome bgs fromucepe aoaoybed coone may be ghy eacoy to necon

even after repeaed eedng on ood of anma wth hg paraema. Pps &Bertram (15) repored hat n a group o bugs wch dd not conract the neconae e nae o deam paa ony 1 o e pecmen were

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8 B

cd r scod cv m som o h rmg scs coudgv v r hr r v ms wh dr T cruzi srs. Svrumbs o nf/n a g suscb aa vco wch ad d oxma ocuad mc acu has ad gsd

mo oodsram omasgos aso ad vhss o com cd(6).

Th gograca og o vcor ad ras sras hs suggs asucg cc. Das () dmosad a a Vzua sra o rao-soms cd a sgca g cag o R ou om ua(6.8) a nftans a megsts o Baa g (8. ), wras Br sr ws sgc mor cv or h oc bugs. rdvos h adu a ava sags bu w a dcras suscb o Rpxus rva a ach succssv sag ds casg ak o bood a

ymsgs (9) As dgs h h rv sg

ms sra h s sa Das () suggs a ad ood dgso ma am haas a so accou o h gav sus. I scs sowg a ad moss h gs d a o dga a ss bo omasgwas cosodg g ()

Smoc mcroogasms occu gu o u vcos 8 ).r 33 su uc o o oos smos N o h dvom ad ao o u R oxu sowg ha haas mud mo s os. Howv r rvs ss cor or dra o masgs

msgos, sc h c rms w gr u sr ag scs Th ro ad Nocadia d ma oh Gramosv acra gu o romds (8) rms uucd.

T oss h gc acors o h vcr r vvd h cc o u vba os as b vokd Ps & ram (9) Tod ha xma co a o og o a gou o vcoswhch ad o g cd ar gsg cu was sgca owr ()a a obad o wo vo oao 85

Aoug som auos av dcd o sgca dc usas rsc caac o c vcors () r s s vdc a sd aboad ryomasgos bav d va o ad a somc a k o occu w oaos o booam omasgos,wh rdmac o hr orm r h rams Sva (9) rord h h cor's dgsv rac ms broad rms so dvo msgs orc ag o ud orms ws ms s ms usua k morhoog movms or r ds d dgra o rar. Asgc hghr rcg (8 .6) o us d o cur c wh do mc ocuad w sas g doma sd oms a w a g a ag umb o sou oms ( o gav scs) ().

Howvr ro a h dr oms as w as a durg boodgso cou o b ooug vsgad sc umd ouaos coud o ov o vcos.

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BOOG OF OSOM Z 5

e e u Inn n e In veebe

hprid iri bw ici a ad iiai h r ifciacycic rpaig ha uuay b cidrd a rgy ucd by

hiar udrg by h ic ad by praur. ia ) rprd ha i mgistus macycic rypmai ar dcd h ih h dayi aa a ad h h h day i adu pci. Siiar ruwr baid wih atoma protracta 6). Phiip 57) hw, drad h dpm T uzii h c b dircy rad irma praur whar h iar Iciiy drmid by icuai uariy rad fc ard fr a ag f R. pxus h h day a0C, h hid day a 5C ad h cd day a 5C daa ar rpri wih ic d haiy icd h ad may b ra rcr wih w parai ia

iiai f macycic rypaig i gray aud pri durigh cr ir i. Luz Brba () r ia dad ha infestans pt imiaig ifci aa 8 day ar ifci dpig aai pid O h h had, hwr, y .8% f R proxusicd aa raid hir ifci ar big rad h adu a rubid a w arai pd 5) Icd ymph f inftansaid h hirih ad iih day ar a a haiy icd ichwd a ard dcra piiiy .6% ad 50% fr h iar ad 6.0%ad 0.0% fr hird iar, rpciy ) wihadig h daa, h

iprac h cr pci i rpc pric f rui ifci ha b hruhy iiad Liwi h a acycic ypaigiiai ha y b iiad r ha hi umbr b imad,wig ac f a riab mhd fr cui h aga, dpi a fwap cr ici r dii i h c 5).

uidpma ag d rach h aiary gad r h hcbu ay b dcd i h apighia ub 5). Eidc ru raiiby h bi duiid, rprd by ary auhr, ha b riwd by Pipi 6)wh pracicay rud u hi pibiiy S ap ijc bd cuurm i h hc rud i h aa uria fr y a iid prid5, 06) I pi crria rcrd, iracuar ag ar w cidrd cur i ruicr 5). harfu c h ibah ha b rcrdd m auray r primay ifcd bu

aa ay ri i h cr i fr a a i day afr dah8) rh cy r ria fr ar brd h ria ruicy dcribd 0) ar ry iy a a Batothidia which iciamid (39).

Devepen u n n Inveebe

T ru' ma irbra h ar riamia Rduva) a haphau. Hwr, hr Arhrpda ha b priay ifcd wih h ai [riwd by Pipi 6)] icudi hr Hmipra (Cicida imx



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0

las and other bebugs; Lygaeiae: Caa apos Diptera (Hippobosciae: Mophags ov; Lpopa Pyraliae Acarina (Ixoiae, incluingticks belonging to hpphas! Ohoos, an Ambyomma Mos of thosearthroos are obviously abnormal hosts unable to support the full parasite cycle.

oeve fee of Cimex hemipterus and ltulaus fed on epeenaynfee nma had many meay ypomagoe hh poved nfeve ome (21 anmon of T cui o noma dog y naay nfeed kRhiichu nuinu) ha ao een epoed (11). A poe oe fo nonng e h a Muc domic a e a epemenay nfeed u inhas een ded by pn (6 n hs compehense evew on ansmsson of .

Golman (81 inoculate pupae of a plant-feeing Lpdopa Phosamayha with T culture forms an reporte the presence of free parasites inthe boy ui an migut as well as intracellular stages

Fne Se of Deveopmen Se n e Veo

anaa (15 ede dyng he ae of epmagoe and ypomagoe n R oxu ao poned o he andane and e of he mohondaonaed h he ay n he veeae e fom ak 8) deede ne strucue of sphaeomasigoes, eJimasigoes and yomasigoes in ou and peened a deaed dy of kneopa mophoogy dng hedeveopmena age ndng he aangemen of he DA oop n a ageype of keopa dvon and eaonhp eeen h DAonanng

ure an fJrmaon of miochonria

EVELE I IUE ULURE

The inracellular evelopmen of in avian embros an in tissue culture haspovded ngh no naea mophogene nene of deen fao onnaea go meaom of naea age ohemy of nfeede and dg avy agan ea and naea paae

Avn Emo Ineon

Chk emyo ave een ed y eea ao o popagae cuzi eveedby pn (60] espe a few unsuccesfu aemp avian eyo have povedsusceptible o infecton showing agellaes in he bloo (74 an viing forms invarious organs as well as in he choroallntoic membrane (6 7 Bloo nasonseems transitor an is characterize by very low parasiemia (60 Chicenshatche rom noculate eggs ha a ransient paasema (6.

Te e nfeon

naea czideveopmen a oaned eary n pama o e e

and hangng dop e (12 16 of emyon and ad e [eveed yPpkn (10)]. ae monoaye e e nfeon ee oaned n pmaye fom ypned e penon ( 1) o e ne (10 15 191)

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BL F TPANSMA

Deen ssues m vaus anmas and m man have pved susepbe naeua nen wh 2 9. Neva e a 4 ead nfeedsee emb ssues f hk d , hum emb sk muse, humkde, d hum m Amg pm uues hk emb es d

vus ssue e nes, human hea es wee nsdeed he bes (8.nua wee genea suspensns f mea pmasge fms havesed

fm bdag uue Sa deees T crui uues n espe nfe-v f sse uue es hve bee eped (2. uue fms mned nhe aba f ng peds end se he nfev f ssue uue es ban a gea ed f ehabng paase, een sae fm he veebaehs wee emmended as nua 9. Bdseam fms dd n appaenpdue susaed pgessve eua e, d ve md paassmesued m un f bd ng pmasges 2 Ohe epe

mens, hweve, shwed a e ne deved fm a mune hndsama behgh susepbe nen wh bdseam ms, snge paases eadg eu gwh suh es 2 Se ua nssg f pases ew eesed fm feed es d egu pdue hgh fe eshe nfev f hese fms s unea. Mea pmasges fm andHL meda ee sgnan me neve mnes hea es han msbaed m veda need ssue uue ud medum 9.

nun prur n rphn

Ms papes deang wh deean f amasges n pmasges andwh mphgeess f he deen pmasges n ssue uue hve beemeed pevus 28 , 7, 7, , 227. he uee f dee ubempeaue upn naeua gwh f he Baz sa was nvesgaedb Neva e a (2 A amasge ms sead muped and hanged npmasges, eua fe pgessed and he ppn need esehed 20 0% e 20 ds. As esu eeased ageesesg aumued n he ud fm eed uues; 0 pmsge pe meud be deeed ae 2 weeks. A 8, hweve, he peenage f nfeedes ws usua < 0 msges ud dvde bu emaned hs sage whehe de pmsge fs pgessve e vs Auee b empeue, hweve, ws bseved sm emes whhe Y san ( hs dsepan suggess a paae wh hemsensvmehansms p vuses and emphases he need fuhe expemens whsevea sans asean whehe hemsensv n he amasgepmasge ansfman nvves muana adapan he nuene f deenempeaues n he peeage f naeua nfen n mnke hea es hasbeen suded (9. he mphgee f empeue n ssue uuems was as desbed b es e a wh eped h a nfeed

es eeased sede pmasge fms dena hse n he bdseam fhe veebe hs whees 7 uded eshmds d u ms-ges pedmed mg he pses beed fm upued es.



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36 Inene Ineen

acua owh o u a mu chodacoma c wa uacd b h hh v of rfo ducd b a hc oucoddobesrnded poyinosinc-poycyidyc cid (124). The ck o proeciongins nrcer prooo incdng T orded by his nvr genmigh be expned by he c h nereron spposedy hindes synhess o vrpoeins by dsrbng he rnson o mRA on he hos ce boom I hecase o he prasies proein synhesis s coded by her ncec cds ad synhieby heir own rbosomes

Pe e n Ine F

A od h boh of ucod ad oucod b cu cuurfom fomd houh h ua of oou a h mdum 3

de novo phwys beng bes negigby ow. Yoned 229) nvesiged hebosynhess o prnes nd pyrmidnes by inrcer orms o T n inecedoney her es o deermne he meboic phwys whch ssy he hghdemnd or nceic cids by dviding prses By he se of abd c addenne s we s meboc nhbos e.g. zseine nd he inonceosde oomc wa dmorad auoradorahca h he preeren bosynhec phwy o prne componds n nrcer sges s ppreny de novo synhess wheres exrcer orms eep sng he svge phwyA radua racm of h d ovo bohc ahwa b ava m o

occur a o a amao raform o romao.Bemstr rurbng es

Prsied chick mcrophges hve more gycoyc ivy hn do noninecedc (220) Th vao frmao o caud b uco comumo ofho c aa b rher by he hgher gycoysis o he neced ces hemseves nd does no pper reed o he inensiy o inrcelr prsiism. cn accomh h who racuar cc v wh s ons oexogenos gcose vbe. he sme mno cds ccme n ineced cresad cubad coro (220)

Exeent emter

T ziineced monoyer sse cuur hav o b ud a a xrmamode or chemoherpy Despe obvos inconveniences sch a dcu d-ecing civiy o drg eboies he ipossibiy o u hh oubcomoud ad h abc of du xco ad of a mmu ro hmhod om advaa fo ud du acv. Thrauc civiymy be deeced gins boh exr nd inrcer orms Sned prprionsgve n ccre pcre o drg con gns he nrcer orms nd qni

v daa ma be obned by u rded dru cocrao. Sv Krchner(93 udd aco of mau cabdum ufa ad omc amoucod o cufcd mooa of mo ha c cv aa

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LG TS C 363

etra r raeuar rm wa determed by ut trymatte teuture ud ad ti me haraterti iraeluar et uh aditurbae the divdi rm Cardium ad tylmy amiuleidewere learly ave aai aellula m wherea maue eemed ave

y aai etraelular m Ntrura teted i eted rimary tue uture rm tryiized hie ell ad i eLa el rved hihy ative aatiue rm (22, 1 29 Th vitr ati itrura ha bee med i vv;eletr my he myardum me uaed wih T c ad reaedwh ruraze hwed he whle euee deeerave hae ad deru-ti the iraelular tae by hi aive mud (33

The ueule tiue ulture i hemtheray tude T ci wa alevdeed by a iveta the mde at tylmy amuleidewhh raaly ave aa eaellua arate bu emarably damatraelular rm (193 194 utterde e a (87 tudied thi deretial ueti

bity ad ueted that it miht be elaed ethe by udametal dereebetwee ure metbm bt develmeal tae r by metablii rar-ma te d a mre ave mud Whe huma hea iue ell wereubated wth trated dru tw mud wee baed me evdeeemered that e thee metabli rdut miht a aait iraelular tae.

Attemt ue ieted tiue a a i vitr ree te r ueul dru havebee rerted (14 6 tet ha bee eved whh ermit aemet druat cruzi ee t taellua wh, ell va ad aelluadevelmet, a well a the evaluati dru tiity u u ieted ell ( 14

he ue tue ulture reei wa , hwever edred by Mieth &eide at ( 19 w idered thi et imited value havi teted i eLa elme mud ative vv, wth eaive reult

able hw he rul baed w dere mud aive i vv hetudy ther atvy aat ra ad etraeluar rm tueuure

n Stutu of Tu Cultu Fom of uz

The truture the aelum ad ieatmthdr T c tueulture rm ha ee deried y Meyer 1 eide te uual u iet

at me etraellular trymatite hwed i their ietlat eletoluet rma arraed lie mthdrial rtae, imart t he truture abaee ae Thee rmat wer t berved he umeru amaerm taied i he aratized ell r i traelular trymatite It habee ueted that the etraelular arate ehaut a the etat DNAdur her le a utde the ell ad are uable at th tae, t ythezeew matera the aate mut heree d a ew h t reume their ie ye

A yttme trallular ci rm i HeLa ell ha bee deribed(131. The ruh edlami retiulum ad the me trymatite

rm tiue ulture are le develed tha i bldtream rm, reumably be-aue the eatie roduig truture are t uetly mulated by the viro dii l 3

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34 Actviy of differen ompounds agans nra- and exallular ssueulure foms

Compouns

Nofu

Pmaqune

(8nunolne)Crbdum(hennhrdnum sulfae)mnonuleosde

o Sylomyn"Cuzon(Bsqunaldne)

"rotyan

(Tralen nial)

Ts noheny rbonu)TyadnAnoerin

A

Ans Agansnalul eaellul

os oms+ +

+

O OF POO U

Referens

14, 22, 129

93

22, 93

87 93, 9

29

29

4887

Undnd mdi f cuiving T cu miny bd gs v bn viwd( 05) A fw us usd ucvd dipsic g-bd mdium (140) iquid mdium (13); T cuisms n qui bi mis. T civnss f diys mdi (07) is vidnc T ui my gow wiu mcomcus o nondiyzb ipids Tw comp undnd iquid mdi LITmdum (v nfusn-Typs) (3) nd bin- infusin + bod (1) n usd mss cuivon nd mpgnc sudis

Dn : Gwh

T cu ug ss s n mmmin ypnsms is si xcing nd s n bn cuivd in nd mdi Li & ubbw 1 1) ndmp & Li (84) cuivd i in mixu ppn Csmincids uu gw s nd bbi bd guum is mixu wsimpvd (1 1 7) by pcing ppn nd smin cids wi min cids vimins gucs nd miscnus cmpunds (dnin uci u puns ndpyimidins c) prsi ws subcuurd fr ng pids in suc mdi. Apty dnd mdum ws dsd cntnng mn pud fctn f sumbumin nd m juic (44) ws pcd by nwn gow fcs

incuding vimins RA cyidyic cid nd cinin is py dnd m

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Z 5dium in which casin hydolysa was h only undnd comonn suodgowh comaabl o ha in blood mdia o 5 yas (45 bu Bon & Pan(7 did no obain saisacoy gowh wih i.

Bon Pan (1 od ha yos-li inusion mdium dscibd

o dd no sui uzi unlss s ca sum was addd Egg yollcihin ad gow aciiy wn addd o ypos-li inusion om hcomosiion o gg yol lciin wc sggsd al o ay acids only saaand o som xn bna omod gow s suls w onmd nihy amb 2 wo amd o gow in Bon & Pans saa mdium no by nands & asllani (5 who od a saa couldno pac sum in L mdium. O incasd gow in a saa mdiumobably by nancing h biosynhsis oHay acids (. Wih ypos as h onlyunnd comonn li xac was lac by hiamin olic acid wiou

dcasing h gow a (7 again nands & asllani (5 could no con hs ndingsMons (132) sudd go o ou z srans in smsynic

mdia o Cii & Gossowicz (4, Bon Pan ( il & Subbaow ( and Lil & Olson 7. Bon & Pan's mdium supod gowh o all sainsmaing i h mos omising o dvlopmn o a dnd mdium Dincamong sains in ii & Gossowicz and Bon & Pans mdia oin o dinsin nuiional uimns

Gowth and Deentaton n Clte Fom

Soon a culiaion o T cruz i was cognizd a oigin in culus o inci macycc yomasigos is om imasigos (oms haing an anioundulaing mmban and ha macyclics a usually unabl o mulily Anincas in macyclics in old culus as bn commonly obsvd bu acoscolling moognsis o macyclic yomasigos could no b olysudid in h usual diasic culu mdia n a ss o as basd on us oh IT liuid mdium which supors as growh and allows s o lronicounrs ( 4, culus wih oiginal dnsiis anging om o 20 0/mlsow onnial gowh o ou days hn a saionay pas (. h numb

omacyclic ypomsigos dcasd duing h ponnial has as a sul onsiv muliplicaion o pimasigos h nd o ponnial gowh howv h numb o macyclic oms seadily incasd as a sul o h imasigo -- yomasigo diniaion which pocd as culus ag Inculus in an xonnial has by daily subculu h popoion o macycli oms was xmly low Subculus ino sh mdium o culusalady dislaying ongoing dirniaion ccd urr morphognsis o macycJics whas ans o a oo incoml mdium (lacalbumin-sum didno slow h diniaion (5. h obsaions suppo h ida ha hdiniaion avod by h saionay phas may dnd on xhausion o nui

ns hlss acumulaion o unnown acors in h mdium canno bignod sic ormaion o oganic acids may simula ansomaion o

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emge n rymage (7. cec an ruvc ac, ae gu-ceree meum hvng gh c H, ee u erenan, uggengrcan he rcrbc ac ce meabm n erenan (7) Recemen ver nun, uu eme n IT meum, b g her

nun, an cncman werng H rm 7. .7, rngl muaemeaccgene In h meum (HI, cuure rencube r 48 hrhen ncube 8, ha abu 5% meacclc rmage whn vea (7. The change n H eem een r he nene erenan ncereacemen he ver nun b g hear nun n ncreae mecgene

Fene en (, ung mum, hwe ha g hae rgnm were rger n hve hgher RA n en cnen where he nanr hae nhee ren n nucec ac much mre w whe

ung le nhe ren an nucec ac ha been ue b ncrrang abeeeucne n ren, Hurne n RA, an Hhmne n DA (. nH meum a H 7., whch ur grwh bu n g erenan, hehghe ncrrn rae were a he en enenal grwh he hen e-creae urng he nar hae When, hwever, wa grwn n HImeum a ., whch favod diniain in addin pak f in-rn e en he grwh hae here wa ecn e ren, RA,n DA nhe whch lw n rn rlel re erenn hbur nhe eem le r DA, be crre u remnn b he

mecclc rmge n he emge.In he bvemenne ue nl he Y rn eem hve been ue. hr

(4 cmre grh an erenn Y an MR rn mnne n vrr eren er aer lan rm eermenall nee anmal he gen-erl grwh characec were cnrme eren lae ha mlar grwhcurve neverhele, vaan n reec erenan were aaren. he Yrn mnane n r abu ear ha 00% meacclc rmage n he nr he, where her Y culure manne n r an 7 ear, reecve h m 2%ecclc. On he her hn, heF culure he Y ran ( e n blgr meum r 7 er, hen er n I cnenl h n eremel lw rrn mecclc %)MR uure mnne 5 er n I generll hwe hgher eren-n re hn he Y rn.

hee laer eermen arenl hw h bee uble rereng r mulng nuence revalng n he meum, mrke erence are nheren n heerenan enale everal culure, reveale b rlnge labrarmanenance n vr.

Eec fru n Grwh, erenan and Snhe f Pren andNue A f ulure Fr

Aem ure evemen cz culure rm wh nhbr ren an nuclec c nhe hve l uull been errme wh he am

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67f preventng nrmal mrphgeness, therey tanng pplatns f agellatenale t derentate t stll retanng haraterst antgenty (66, 69)

The nalty f tre frms t synthesze prne and pyrmdnenetdes, th nseent dependene n exgens ase (6, 168), ed me

athrs t try t nht nrpratn f ntrgens aes nt etde andnle ad th pne and pyrmde analg (6, 66, 69). he amnledef stylmyn ked nrpratn f adenne nt prne nletde and ead rra (F) and rra dexyrsde (FUDR) nhted nrpratn f ra r dne nt pyrmde nletde ad nle ad(66, 69, 1 68) The rapd ht y drg f DNA ythe drted rh: ee nreaed and ppatn grth ed (6, 66). ean antt mtmyn atnmyn D and prmyn ht grth y mnng th neads tmyn nhts synthess f DNA and prten and ater f NA et naanes grth as evdened y ytknes tht dpatn f DNAntanng rganelles (66, 69). rmyn al strngly nhted prten synthessy T pray y ts eknn atvty a an anag f amay tRNAAlthgh rather atve aganst tse ltre and verterate hst frm, theamnnede mety f prmyn dd nt nht ltre frms (66, 69).

Atnmyn, extenvey apped t T tres, trngy nhted RNAsynthes and at hgher nentratn, prten and DNA ynthess. tpatnas ked a demnstrated y nnnpratn f thymdne n DNA t,athgh t prevented nfetvty t te tre and anma t appaenty dd ntlesen antgenty hee eets ere rreversle nrma synthes a nt

resmed hen the drg as remved and the parastes pt n fresh eda (66 69)

DNA-M Hrs s?

In azng dehydrgenases n tre frms, Lehmann ( 1 1 5) fnd tat tt namnaed t aa tt ft frm htf nnntamnated res f the same srans. Befre expre t adIa thstran hed ny ne dehydrgenase eah ether str dehydrgenase r latdehydrgenae aer 11 day f ntat th adda hever, th tranntaned the t enyme As adda tsef pssesed the respeve prmtveenzymes akng n he sggested that genet transfer had taken paeeteen adda and .

lv Asg gs M

Trager (10 n dsng dte n tanng devepng stages fTrypansmatdae n elfree medm as are fnd n the verterate hst reveedsme f the envrnmental ndtns hh mght trgger derentatn tardthe pear frm f Lsmaa and ypasma say preent mammaanht. n tre, amastgte frms slated r aggtnated nt argemae have een repred spradaly (41, When as grn n a

medm tanng pe atem prepared rat, arge ne amatgte (eshmands pred y nay n apaed (2. Amastgtes asr n nfeted d kept at rm temperatre (9) and n the d f nfeted



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38 BE

( h l wy ad bld m h za vald ymag adly dvlg amag whh gaday hagd mag h wa m vd ha bad ad bdam ymag a m hag amag ha a

h ld (3Cva c ally ad dmay a hmam" l mda wa d ( y g h T mdm (Tya, a, hm, abb bd, 's issu uu ui ad isi u C a ( , ag wh a l ag 98 mag b-ad 9 amag a aag 4.C m l md gw T mdm a mag hwg ha ma wah m a mhg h dma amag wv, wha iml mdim (mdd mdm 99, hmyzd abb bd,h lama, ad h mby a, > 9% amag w bad,whav h ma. w md wh h mby a b lag h ama md a va hmam, v ma (3 ( d ayg a ma lg dvm amag l , h m babyhl ma h mag mag aay ba a a yamag a ( d a amag a aayb d vv a hgh m ha mag whh mbamag shmana ha a lw b qd vv a 37 haa h mag ha m ag hav a ga gy dmad b a

dad byh aably a hgh ma 98Inf of cz ulur

mg h cT agla m a l y h may yma-g a blvd b v h vba. Th vy ad maad g d wh ama aag. Th m dvdl wh m 13 ya bl ag (4, ad hal hav b bvd m lad wh v maylymag m maad ab 5 ya . m hwv adly dmh v (4 m gv v 8 agladd dv a a aama (4 Th aa vaiaby vy ma wd whh ymag may aly dvl lgm D m d a . 9 ya ad m 7 ya (4 ad a vaab b am awayga mb may ymag Hw a h ymag? y g h mayl ad lag m wh a mla mb v m Zd ( ad Cha (4 dvd a vl va maad d d h labay. Thdad vl m , h, md b ad by

vy h mayl ah ha by da h mb mayLgh m aal mda may g ag wh da vl 4.I ha b ggd ha gd maa aa mdm

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BOOG O TOSOM Z 369

ara eer e o eeo o ha he ererae ( eerhee,a nge aage hrough he eerae ho of aenuae cuue ha o g-cany creae ruece n exerme erforme aer reoaon a cuaon LI meum )

Acnomycneae T cz cuue form aaeny o fecy for mcewhe reanng agec comeence an ocuaon of reae ageae aorea ceran roecon agan chaenge wh uen oo form (68

Ceacu eece eman o e oane a o whehe T cz nee uene nure nce h ee faco fo mohogee ca e oay aaye h urn ma more ream o he ue of cuure fo rugeng cuue mgh hen more coey aroxmae o cono ncunguae gowh a oo a ue H a emeaue.

CYCHY ND MMNueic Aids: Role of Kinetoplast DNA and Dyskinetoplasty in crui

A emorae Schkrau (188, A from ac ryaoomacou e ocae y CC eygrae cefugao o a mao a aa aee a. he aee comoe whoe ae comoo e fomnuea A orreon o kneoa A a reeen a ea n T z- 20% of he oa exrace A (. h emakay age amou of exanucea A unuua among a or anma ce her mochora A oy0.5-10% of oa ce A Kneoa A ha een uggee o acae

cog of roe moee of oae enzyme a cyochrome nranoomaae reew: ewon (1); H Aneon (9 .

he keoa a eermna of enzyme aance n yanoomaae ha eecue n he aforemenone reew. here no reen reao o hk haT cz erge funamenay fom hee aern, herefore h uec w emeoe ony n oue.

ermaen o of eugenoe maera from he keoa (ykneoay common amog yaoome whch ae mehacay anme. Ac-ne ye a ehum n o kneoa A a may uce ykneoac

form n cyccay ranme ryanoome. h ecc ng ha ee uggee o e caue y he aence of hoe n he kneoa, y he ecuaronear rucure of he kneoa A, o y erece ewee ucear akeoa memane (1. A emonrae y Mhfor (16 ykneoac form oe he eugenoe e u ean he ouememaneeeoe. A now emonrae y hymne cororaon an eecron mcro-coc raoauograhy n T evas ykeoac ooream fom, A reen n he woaee keoa memrane ( 1 ; h conon hough o e characerc of he T vas ra. A for T cz coroaonue wh hmne howe er ga n he kneoa u no n he keo

a oue memane (15 I gou yaoome ooeam y-keoac form may muy eney he ererae ho u canoeeo eco o cuue whch acco wh he coce a gou

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30 BRENER

yomagoe mu elaoae a cyohomemeae eaoy yem o u-e an muly n he eco. he eoaa yanoome nung T uioay een on a unona yohome yem n al age n oh ho anae heefoe unae o muly n he ereae ho f ykneolac. Mhl-

fo (134) ueye aou genea of yanoomaae an aw ykneolacfom n a ee wh he exeon of cu ooeam yomagoe Inhee an of T cuia low ooon o ulue fom (00 o 0%) weeonaneouy ykneolac. Smla eu wee eoe y Ono e al (1)who oee < % of uch fom n cuue Cuue fom n meumeae wh aane ha a hgh ooon of ykneoa emagoe (080) hoe form whch coul no howee e uculue neehele couleenae an n ome cuue acoune fo he eene of 0% of ykneoa meaycl yomagoe (0). hee anorma meaycl wee nece

o HeLa ell an gae re whn 448 h o ykneolac amagoe u eryoon mo naellua age ha noma kneola an hee wa eence hamoe ha one geneaon of ykneoa aae ha een oue.

he eoe onaneou ykneola T cui looeam aaewee eoe y Deane Kloezel (0) who oee a few fom n a le fomhe loo of an aue h\man ae of Chaga eae A few ykneola yo-mastgot a amastgot oms av aso b obsv n a ots s o heeoneal exuae of mce nouae wh aaneeae culue (0) Al-hough ykneola fom hae no een egulaly oane n he mammalanho an n ue cuue, hee nng uggee ha an mae kneoa may

no aor eeloen of T cuin he eeae ho an hene ha eeneneo T cuon mohonral eaoy enyme mgh no e que a eenal auoe fo eeomen Moe ue on he ocuence of naua yneo-a aae an long em oeaon of he ehao of onaneou annue ykneoac fom n uue an ho may e neeay o eewhehe kneola DA al fo he aae heeoxenou eelomen ameae y mochonal enzyme

. Nue n Kneps DN /m u

oa DA omT cu

ulue om howe ee a nuea an whuoyan eny 1 7 10 wo aelle an wh uoyan ene of 1 699 an 1 686eeely (173) he gh aelle DNA wa no howee, eee n fueexemen aeaane ha een elae o ome unknown onon of hegowh meum (174) hee eul hae een onme y Ono e al (11) whofoun ene of 1 709 an 1 699 eeey, fo he nuclea an aelle anhe ae comoon of nuclea an heay aelle DA ha een eemne; heGC conen wa 10% an 398% (a eny) eecey (173 174) heeeo conc wh hoe of enane Caelan (6) who foun n DA ofwhole oganm a hgh ooon of aenne an hymne an GC of ony 7%.

cuikneoa A soat y oyaneny cenfugaon n ehum-CC eny gaen ome oen an oe ccle of 04 m monome zea wel a ong lnea lamen o 1 m ong (17) DA 699 wa fome of

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BOOG O TOSOM UZ

mncrces 0 /m long whereas the DNA 1688 was bult of lnear strands 1 1)Crcular DNA molecules have been detected n nuclear and knetoplast nuclecacds a study of the frequency dstrbuton of the crcular molecules showed a largerrange than reported n the prevous papers most of the molecules were 020.9

/m length and molecules between 0 1 1 /m and 12 m have been found Most ofhe neoplas DNA, however, consse of long lnear srands 1 1 Whe two trypanocds Berenl and hydroystlbamdne) nown to bnd selec-

tvely to netoplast DNA of trypanoome were used n uz cultures, DA1688 ncreased from 1 to 10 and DA 1699 decreased from 0 to %. Bothdrugs apparently enhance replcaton of one DNA and nhbt the other ). romthe DA content of each paraste, the number of crcles n the ru netoplasthas been estmated as ,1). The exstence of replcatn knetoplast DNAcrcles was descrbed for the rst tme n trypanosomes n Berenltreated T ruzculture forms 19). Berenl ncreased he proporon o replcatn molecules by 13 J

probably no by a general ncrease of DA synhess but rather by blockng tsreplcaton at Arch specc pons he lenghs of he replcang branches correspne o mulples of of he oal conour lengh of he crcular DNAmolecules, whh suggests a dscontnuous replcaton process

Some of the reported ndngs warrant addtonal nvestgaton ong to dscrep-ances: (a) The exstence, n derent expements of a sngle or two dstnct satelltebands detected by denstygradent centrfugaton of wolecell DN. Those twosatellte bands, dsplayng derent buoyant denstes and molecular comonents,hae been egularly reported by some autors 1 , ) but are not mentoned, for

nstance, by Ono et al 51) who detected only one band ( b A certanheterogenety of data for occurrence and sze of knetoplast crcular molecules.Although Rou DeJan 12) reported tht most of netoplast DA was formedby crcular molecules, Ono et al 1 5 1) clearly stated that netoplast DNA consstedmanly of long lnea DNA molecules and ha mncrcles were no he maorcomponent of the knetoplast. On the oter hand, most authors reported the presece of free or complex assocatons of mncrcles of0550 m contour lengthOno et all 11) hoever, although not separatng mncrcles fro nuclear andknetoplast sources descrbed crcular DNA molecues rangng from 0 0 to 1m. ) he codng propertes of the ncrcles ave not yet been dsclosed,

although they probably can code for a few polypeptdes 95). Those pecular DNAarrangements have been terpreted ether as evolutonary artfacts wthout genecfuncton or as an amplcaton of genes nvolved n mtochondral bogeness 95)Ther ndependent replcaton apparentl onfers upon them wth genetc sgn-cance 19 The presence of lnear DA laments mxed wh mncrc1es has beenexplaned eher as conamnaon of nucear orgn or by he supposon ha hsmaeup of DNA plays a role n replcaton of netoplast DA 2).

Nueic Ad Metabosm in cruzi

Purne nuceotdes may be syntheszed ether by converson of precursors e. gly-ne and CO de novo synthess) o by utlzaton of exogenous preformed purnessalvage pathway) Use of Cabeed glycne revealed that T ruz culture foms

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7

my synesize urines bu ery low re ue o rdiocie denine,oweer, s sown reormed urine uilizion is reerred or synesis ourne nuceodes (6). Using Cbeled urcl nd oroc cd s well s mebolcinibiors, Rey ernndes (68) suggesed culure orms cnno

synese yrimidnes de noo nd woud ereore deend on reormed yrm-dnes fo nuleode bosynes Ts deendence on reormed yrmidne seemsunusul since monoxenic Trynosomide cuied in dened medi reuireexogenous urine bu no yrmidine (98) eendence o on exogenousune nd ymdne bses s suggesed e ue o ese nucleodes ore biosynsis o nuceic cids my lso suly ddiionl energy or nboicrecions (96).

Te cemoeuc con o some selece nbos o rynosome nucleccid mebolism s been sudied in mice (99). iomycin C (n inibior o DNAsynesis), 6zurcil (inibior o de noo synesis o yrimidine nucleoides),

nd cnomycn D (nbor o N synss) dd no oong e le o eexerimeny ineced nims An n iro inbior o urine nucleoidesyness e minonuceosde o sylomycin eered some ciiy in exermen-ly ineed mce nging e cue disee no cnic one bu wouecing cure (70)

A A P

cuure orms conin 4% o roein by weig (9). Te roenconens o cuure orms in exonenil nd sionry ses o in LIT

medum sowed, reseciely, 67.0 nd 4.0 g X er gele (65) Culureorms exmined by er cromogry conined, mong oers, nine ndsric cid resumbly ormed by rnsminion () since rnsminse ciy d been obsered culue oms 3

e oysie nd rginne is meded by cie rnsor nd seems somewdieren rom eir ue by mmmln cels (90 9 ) Proeins nd some minocids my be uilzed by culure orms s energy sources in sugrree medi(08, 0) In medium deoid o glucose or in wic sugr deleion ws demonsred, e H rose, cused by roducion o mmoni rom nirogenous subsres(08) ccording o Zeedn (0) oidon o mino cids in orgnisms ing

crboxylic cid cyce is robby medid by rnsminse sysems. Honigberg(96) secules is oxidion is likey o be eroed only in ose blood-srem orms o rynosomes ing uncionl ricrboxylic cid cycle

Lp

ids n culure oms comrise u o 0. % o eir dry weig (9).Colesero s been idenied in cuure orms in medi conining is subsnce(9) Te resence o n ergoserolie serol (89 ws no conrmed by Wlm-so (). More recenly, oweer, six seros e been deeced in cuureorms grown on serumree mdium. ese, our were ergoserol comounds nde wo oers wer robby 4meyene7deydrocolesero nd 4eyi-dene7deydrocoeserol (09) colesero ws deeced ony in gelles grow

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Y TRYPAN CUZI suca u. ay ca{ay f xacs f uu fs sa sva ss sayaa

sayc a s .asqu craray f b a cuur frs rw au wr csr was ry avaab sw r y aray -ab ss A C a D vc A frac b csr 5 .Cuur frs a A a D srs bsra frs a A a C Pysrsar aary y ra srs ac ryass sc zs-rs cy csr ar f xus r a aary furr abz. Srs w srucura fuc ar rbaby ba xusy wras abcay acv ss a ky ars usy ss b

w vsa caracrscs f aas rw csf a

as s s cuva a w su sc s ay c aby a v f r ca srucurs.

Acvy f afua y Arc B aas su-s rsc f ss aass bra sc yas ar kw ac by aac srs scay sr srs cxs by cras raby f c -ra a r sca f wcuar cs 5 9.

Crbohyrte Metbsm Respirtio

s sr ysaccar us abs s aary sur by uza f fa a rar a by cabya 1 Saquans f a unc ysaccaca acs asxys a ucsa a b c cu fs Ar ysacca y aacs bu caa w ucc ac fas was r cuu fs 19. Sa aus f ucs .6 f ryaa a as av as b c 61}.

a f ucs by cuur a bsra frs f as br cuur fs fa uz as a acs sucrs ras arabs a aacs; ucs frucs a xys w u 14. Svra rucs f aarobc fraacc ac a succc ac cuur frs acc ac succ ac acc ac a sa aus f yruvcac bsra frsa b c rw a ]. Mrvr fw ycyc zys wr c T cuur frsxkas sxs sas sxkas aas sssas a ss yas [vw v Bra ]. ucs6sa a 6succ yrass ys f ssa away a as b r 6. A 6succsy as b scb 16 sus a yruvc ac a ycray

sa a fr by s ac. a f 6succ ac as wv b c by Ryy 11. s sa b u sa ba as sus by aa r by Maca aqura w ca aby f uu frs f w sras abab ucs y sw a uau sra csu ucs a

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74 Bcad labd gucC C a a hgh a ha h uva amv h hha ahway h Tulahu a wa bl guc caabm a cmad w 79 uva aA h Tuahu ai i caly h m ahgic h auh uggd ha,

alug a away i aiv i b ai, uaiav dc dcd h gluc caabc away may hav d whahgciy. A h cl, cabhyda mablim yam i caid u hug aabc glyclyi wh cv guc yuva hugh h hha ahway lwd by a abc dgada yuva va a uca cabxylc acid cyc, a acv cychm ym adduc C wv, dc bw h mal a yam m h b gu ad h h lw gu T lw, T hav b d 7 1 80 Thu, Aica yam hwd cyclcal

cag m a cychm a cychm mabic ahway i h didvlmal ag i vba h wha m hav a acivcycm ym dug whl cyc b . T dc avb dcd a hugh udy h vy h vau ci ad ag cyad (a hb a + a3 cycm, which may umaby dicawhh a ucal cychm ym i b by dc ccic caaciai di cycm ad by cycmical caaca iay ym uch a cchm xida NAD ad NADPdaha a w a gychat xda (9, 1 Such uggd mablc dc dmad a bi vw th ay cha chaacizg bh

gu ahg.Sld bldam m gu yam a cyaid iiv

i lc a ym i baby mdiad by cychm h mu iad a aicula cyaidiiiv glychha xida ymcaid i xamcda gau (8 vwd by gbg (. umy m igaig m h ld mag hwv, h xdai hi h cdm wh NAD diaha wa dmab byalium duci (1) uggig c a ucia icabxyicacid cycl. Thi iay wic aay ccu i h bldam, bably

ig a adaa h umy m h cd b acd byh aa h vba h m al hxu dvlm. bldam m gluc dgadd byd yuvc acd + gycl,ad ay qu


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BOL OF TROSOM CRUZ 75

t s vr unll tht th nwl nnsnst ultur orms wr usng nltronrnsr sstm mt tohr.

Th rsrton o loostrm n ulur orms o T zi s n snstv3 1 80 n th sto uotnt s hgh usull l .Gluos s gr b

bloostm n ultu oms bon uvt to mtu o lt t nsun s 180 tohom sstm hs bn tt n c utuorms ut ut unusull tohrom c sms snt 1 180. NDtohrom sstm ws tt n th mosoml hmorotn o T cruzi ulturorms 6 Th sm uthos howvr oul not tt mmmlnt a, b n roms th ult n sou tons. Nvrthlss vn tht rrbol l s tv n T cruzi ultur orms hs n rv ntton o vrous Krbs l nms .g. ml hrogns n umrs 1 0 sun hrogns ( 1 66 n sotr hrogns 1

nous n nms vov n cruzi boht mtosm shonn Fgu 1 . Ctohom os tvt ws v lo n homognts of cruuue nd w n need exenu toom c (180). toso nvstgtons of tohoms n cru boostm foms s s sltv su o th so nms n sn n o oms hv ntl no t n m. Howvr tohl mhos hv shown th rsnn h ntolst o T ulur orms s wll s n romstgots nmsgos o th vrbrt host o suh nms s tohrom os NDHn NDH hoss 10. lthough ths sus nt th sn o untonl tohrom sstm tohml nvstgtons must b onrm mor rt mnts n sum th stn o tohom sstm n lT crui vlomnt stgs sms untn.

h ossblt o n nog btwn th ng bohml bhos oslnr n bro tomstgots o th brucei grou n th nt cruziloosrm orms sms to gnor Thus nw stus on n snstvt nurthr hrtrton o rsrtor nms n shou m wtsom ttnton vot to omrng slnr n bro ors Suh stus woulrol b hmr b th bsn o ur monomoh T crui stns nb ults n srtng th sts om th bloo o nt nmls. Motl

romstgots w st om oo s n tts hv bn obtnb ssng boo of ns nt wth nt trnosoms ruce gmense rhodesense evns ongonse vvx lews though nnonhng oumn Ong to th sml n n u hg btwntnosoms n boo s crui s h on ss ng ston 3

rohrt mtolsm o th ntllul stgs o crui r of host otssu ultur lls hs rntl not t n nvstgt. Th suh su ss s suort th nvstgton o gluos ut n rsrtor quotntom wth Lesn onon om tssu (72 cru mstgot

oms m l obtn n vto 30 16 but sv roo o th nttwth ntu stgs hs not bn monstt so f sts fom tssuutus mo k to ov tbl mtl or ths stu

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36 BEE

I 8 U L O S E 5 _ P .6 _ PHOSPOGNC AD -GLUS

( Y R V )

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Terms parentheses are endproducts of z carbohydrate meabolsm.Enzymes involved in carbohydrate etabos derbed z hexokinase; phosphohose soas phosphohoas 4. adoas phosphotossoerase 6 phosphotose dehydrogenase 7. glucose-phosphate dehydrogena8 6-hohogconc ehydogene; 9. Wood chwed enzye c dehydogenase 1 fumarase succnc dehydrogena 1 soctrc dehydrogena1 mac enyme 1 one

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31/36

O SS

hemheap

O TOOM UZ

Drgs that wl re Chagas dsease are nt nwn wth ertanty. resent sreenng

meths aear eent fr etetng spressve atvty n the expemenalsease 20 2). wever, n ther parast sease has s aly msle hemthe-apsts stanar meths en fal t assess r atvty realstaly ease thespntanes lw parastema n the hrn phase arses hpes that are eventalyashe a f ene re means a rrespnng nerany as the reayf serlgal methds fr mntrn nfetns The dsmal state f aars an egaged frm the fa that endagnss s wely sed .e. allwg nnfetedratmds fee n he atent an s evel f sve agelate lansn the gt S negatve eslts ae nnlsve the patent ay have t e tten

repeately. Better arastgal and mre senstve mmnlgal ests areneeed Immnlga ests demand a far mre deale nwledge f the speantens f than s yet aaae.

A mre atanae gal s elmnatn f cruzi frm ane ld manynfetns have een trae srvval f cruzi n l ) Gentan vlet srtney ae t n enem areas. rsets f nn a ess etnaereferaly rless aent, apear g sne ne s nt n wh nearly nraelllar frms It seems stange tha the sreenn f rs f ths ppse hasn een nstte

cu ongental TansmssonT cruz as ther trypansmes had dne s learnng t dspense wth the nsetvetr as wtnesse y a strng nrease n the repted nmer f ngentaases 9) As a hghly sessfl araste, ruzi has mh se fr evn

Ann onuon

Sty f the antgen nstttn f several T cruzi strans slae frm man,tratmds, a wld anmals) has dente three mmngal types A, B andC) 8 6, ) hese types l nt e rrelate wth hst spees r wth

erahal strn f strans Crssretn sts n me l n sngsh the n vtr tye strans 8)An mmnlal relatnsh etween uzi an lewisihas een sgeste

6) m nlate wth /ii ltre frms were hghly restant t a crui halenge nfetn ledsn tests and mmneletphress shwedmmn antgens etween th sees Thse eslts wee nt nrme y Kletel Deane 0) wh reay nfete rats prely nate wth T swhT cruzi he athrs were als nae t resstane aganst T zi nme reeately nlate wth lisi lstream frms Sme eree f rssrtetn aganst rz halenge was serve aer natn f e -thdi fsult an ptomons ollosom ) as well as f a lvng ssensnf "ptomons psso a mnxen trypansmat slate frm a rev9)



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8 Vanaton agant ruz

accinaion ha been ied wih dieen pepaaion bloodeam om killed bchemical aen (93 o b eznhawn (1) ln cuue om aenuaed

b acinomin (8 amma adiaion 4 o epeaed in io paae (126)culue om killed b ehiolae (38 onic ocillaion (79) o peue (8iin aenuaed accine ae likel o cae cpic inecion which ma imulaean induce immune epone. Phical aen ued o killin and diupin aellae appaenl enhance he ecac o he anien neehele onl paialpoecion aain T ru challene ha been obained wih nonliin anien[eew Goble (80)].

Et o uz on Sontanu and Indu Tumo

Theonadico lieaue on he heapeuic eec o rui endooine and

ae on he deelopmen linan mo wa eiewed b Kaan e al (102).The auho epoed ha he popoion ponaneou mamma umo in miceinoculaed wih meican T ain wa inicanl le han in noninecedconol. hi deceae in he nume o ponaneou umo i pobabl caued bnonpcic imulaion o he eiculoenohelial em

NCDNG KS

The oeoin complexiie he ee moe cleal undeood nea inacabili ohe T rui heapeuic poblem hould no lead one o uccumb o Abandonhope all e who ene a Chaa' deae laoao." Ae all T ruzi canno haeexacl he ame phiolo a man; omewhee hee will be ound a chink in imeaboic amo

KNWN

D. S. . une aied in pepaaion o he Enlih manucip.

Liteatue ite

1 Abbo1, H e al 1 96 Bo Chil Paaitol 19 1 2-4

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29133108 Koodny M 1940 Am Hy

32C223109 Kon D on Band T Tobe J

99 amp Bioem hsio 300110

1 10 Kmar R Kne I K Aemann W

970 Am rap Me H 19932391 1 1 Labech F 19 Rev. Inst Me

p So Pulo 73452 1 abrh 1 66 Rv I M

p So ulo 2494 ham S 9 Nr 2

127374 Lehmann D L 963 r Me

Prsitol 572234 5 ehmann D 1970 p Me

rsito 4331341 1 le P A Sbbarow 1945 Bc

teal 05791 17 le P A Oleson 1 95 1 Bc

te/ 1709141 1 D T 97 Rev Bs

Mlo 1 77-71 19 E Borba A 19199 An

F M Uni. F Pn 1-114757

120 Naqia, 1964 Ptooo 1 1 50913

121 Mara T A Tafur W L Brener Z1972 nn rap. M Prsitol 42331

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124 MartneSa R ope V A ha 1970 S oM 134

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127 Meer 99 Rev Inst Med rap

So ulo 1 1 4418 Mr H O M O 1 948Psitolo 399194

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