ࡱ> lnghijk7 fnbjbjUU '7|7|f92lr r r r < ,Q :   OOOOOOO$R TdO!^!!O! P!!!!R  O!!O!!1K@f"O > ~(r !qM8OP0QMTU!TUO!Doc. dr. sc. Vlatka Rozman Poljoprivredni fakultet u Osijeku Rezistentnost ~ohara i mrava na insekticide i biotestovi Pojava rezistentnosti na insekticide  zaato? Rezistentnost je geneti ki uvjetovana pojava, a nastaje nakon ato je populacija insekata bila tijekom du~eg razdoblja tretirana istim sredstvom ili sredstvom istog na ina djelovanja. Pojednostavljeno bi se pojava rezistentnosti mogla opisati postupnim gubitkom u inkovitosti jednog isprva u inkovitog sredstva. Bitka s rezistentnoau na insekticide zapo ela je ve s prvim primjenama ovih sredstava (u zadnjih 50 godina). Du~om primjenom istog sredstva ili sredstva istog na ina djelovanja ubija se veina osjetljivih jedinki, dok one rezistentne pre~ive. Ove posljednje imaju geneti ku predispoziciju za rezistentnost na pesticide. Njihovi potomci e prenijeti geneti ku informaciju svojih roditelja. Takvom nenamjernom selekcijom iz generacije u generaciju popravlja se omjer u korist rezistentnih jedinki. Na kraju one prevladaju i sredstvo viae ne djeluje (Slika 1.)   Slika 1. Shematski prikaz postanka rezistentnosti ateto inja tijekom razvoja aest generacija (Maceljski i sur. 2002) Rezistentnost kao uroena osobina dopuata pojedinim insektima tolerantnost na iznadprosje ne doze sredstava. Rezistentni insekti su obi no izgledom jednaki i imaju iste navike kao osjetljivi insekti. Mo~e ih se jedino razlikovati po njihovoj sposobnosti tolerantnosti na ekstremne koncentracije sredstava. Treba napomenuti da kod nekih ateto inja postoje prirodne rezistentne jedinke (radi prisutnosti encima koji razgrauju neke tvari) na pojedina sredstva i mnogo prije njihove prve primjene. No, te su jedinke tako malobrojne da se uope ne zapa~aju. Postoji i unakrsna rezistentnost (eng. cross resistance) kada se rezistentnost izazvana jednim sredstvom odnosi i na druga sredstva istog mehanizma djelovanja, makar ona nikada nisu bila koriatena (Maceljski i sur. 2002). Osjetljivost populacija insekata na neki insekticid, esto mo~e varirati. Neki razlozi tome su poznati, dok ostali iziskuju daljnja pojaanjenja. Dva zna ajna imbenika su nesumnjivo sezonske promjene klime i utjecaj prehrane. Kao dio strategije pre~ivljavanja insekti su razvili brojne i u inkovite mehanizme rezistentnosti i obrane na veinu konvencionalnih kemijskih insekticida posjedovanjem gena za visok stupanj oksidaze, esterase, glutation-s-transferaze, insensitivne acetilholinesteraze (AChE) te neosjetljivost ~iv anog sustava na piretroide. Rezistentnosti kukaca na klororganske pesticide, organofosfate, karbamate, piretroide i regulatore rasta kukaca uklju uju sljedee: populacije kukaca mogu i hoe evolvirati rezistentnost u nove izazove koji su za njih po etno pogubni osobine ~ivota kroz povijest, dostupne zalihe genetske varijabilnosti i specifi nosti kontrolne situacije ine neke vrste visoko sklone evolvirajuoj rezistentnosti ne postoji jamstvo da siguran, u inkovit, jeftin insekticid nee biti zlorabljen kako bi se naglo inducirala rezistentnost na njega (Narayanan, K. 2007). Razli ite skupine insekticida imaju i razli ite na ine stvaranja rezistentnosti od strane insekata. Karbamati i organofosfati imaju isti mehanizam rezistentnosti, dok piretroidi imaju identificirana tri mehanizma stvaranja rezistentnosti, ato znatno ote~ava situaciju. Jedino na ato joa insekti nisu razvili mehanizam rezistentnosti su regulatori razvoja kukaca, ato bi trebalo biti jedno od moguih rjeaenja ovog problema (Lyon, 1991). Novija istra~ivanja ukazuju da do danas postoji preko 500 vrsta insekata rezistentnih na insekticide (Grafikon 1). Viaestruka rezistentnost (eng.  multiple resistance )  rezistentnost na viae od jedan insekticid i na sredstva iz viae od jedne kemijske skupine se rapidno poveava. Postoji viae od 1000 vrsta insekata rezistentnih po kombinacijama insekticida i vie od 17 vrsta insekata rezistentnih na sve glavne skupine insekticida (Bellinger, 1996). INCLUDEPICTURE "../../PROGRA~1/IncrediMail/Data/Runtime/Message/%7bF523B385-02E4-4631-8236-FA8ABD82302A%7d/Show/insect_resist_graph2.gif" \* MERGEFORMAT  Grafikon 1. Broj rezistentnih vrsta tetnika na insekticide (izvor: Douglas J. Futuyma; http://evonet.sdsc.edu/evoscisociety/insect_pests.htm) Kako su insekti postali rezistentniji, tako su se primjenjivale vie doze insekticida tako da se danas u SAD-u na njih troai viae od milijarde funti godianje. Rezistentnost je u inila nu~nim razvoj novih insekticida svaki po prosje noj cijeni tijekom 8  10 godina i to 20 do 40 milijuna $ za istra~ivanje i razvoj. Evolucija kukaca nametnula je veliki ekonomski teret (oko 118 milijuna $ godianje samo u SAD-u) i poveanu opasnost za okolia od kemikalija koje mogu ugroziti ljudsko zdravlje i prirodne eko-sustave. `toviae, neke vrste koje su bile rijetke, postale su ozbiljni atetnici jer su primjenom insekticida uniateni njihovi prirodni neprijatelji. Rezistentnost kukaca se naglo razvila (evolvirala) jer se poveala prirodna selekcija rijetkih mutacija koje ne napreduju u normalnim uvjetima, no dogaa se da omogue zaatitu od atetnih kemikalija. Entomolozi specijalizirani za evolucijsku genetiku razvili su strategije za odgaanje evolucije rezistentnosti. Naju inkovitija strategija zasnovana na evolucijskim modelima i dokazima ima svrhu osigurati atetne vrste tzv. "refuges" (pesticide-free) u kojima se suspektibilni genotipi mogu reproducirati, sprje avajui time preuzimanje rezistentnih genotipa. Iako se evolucija rezistentnosti mo~e odgoditi, vjerojatno je neizbje~na u mnogim slu ajevima. Zato moderne strategije integralne (cjelovite) zaatite kombiniraju pesticide s drugim opcijama. Kako odrediti rezistentnost na insekticide Za rutinski nadzor s ciljem detekcije po etne pojave rezistentnosti u prirodnim populacijama atetnika prikladno je primijeniti tzv. diskriminativnu dozu za koju se o ekuje da e uniatiti sve osjetljive primjerke. Odabrana doza odgovara dozi neato iznad LD(99.9) dobivenoj od regresivne linije za osjetljive vrste. Pri redovnom nadzoru rezistentnosti trebao bi biti detektiran ak i mali broj prisutnih rezistentnih jedinki. Po uzorku bi trebalo biti minimum 100 insekata u dvije serije od 50. Ograni en broj insekata mo~da nee biti dovoljan za detekciju niske razine rezistentnosti. Stoga, ako je mogue, treba dobiti dodatne uzorke. Ako, meutim, postoji sumnja ozbiljne rezistentnosti (npr. zbog propusta u tretmanu) jedan test s malim brojevima (10 do 20) mo~e osigurati neprocjenjivu ranu indikaciju. Ako su svi insekti uniateni na kraju razdoblja odr~avanja podtretmana, uzorak mo~e biti klasificiran kao  neutvrena rezistentnost . S druge strane, prisutnost pre~ivjelih insekata na kraju ovog testa treba smatrati kao dokaz rezistentnosti i dalje istra~ivati. Kod socijalnih insekata, kao ato su mravi, insekticidna rezistentnost je rjea pojava, stoga ato je reproduktivna samo kraljica. Kod nesocijalnih vrsta insekata, poput buha, svaka individua je reproduktivna i mogunost brze evolucije insekticidne rezistentnosti je znatno vea. Takoer, razvojni ciklus do reproduktivne forme je znatno krai, kod buhe za 21-28 dana, a kod mrava za 10-12 mjeseci te e krai razvojni ciklus pogodovati br~em razvoju insekticidne rezistentnosti (Lyon, 1991). Postupci sprje avanja pojave rezistencije kod ~ohara Porast insekticidne rezistentnosti njema kog ~ohara (Blatella germanica L.) predstavlja glavni problem za kemijsku industriju i proizvodnju insekticida (Lee et al. 2000). Ova vrsta je multi rezistentna na sve glavne skupine insekticida (Cornwell, 1976; Cochran, 1995). Velik broj znanstvenih lanaka o rezistentnosti B. germanica govori o biotestovima na primjenjene doze insekticida (Bennet & Spik, 1968; Batth, 1977; Rust & Reierson, 1991: Zhai & Robinskon, 1991). Mehanizam djelovanja rezistentnosti kod ove vrste ~ohara uklju uje dva poveana metabolizma (podizanje aktivnosti monooksigenaze, esteraze i glutation S-transferaze (GST)) i promjena mjesta intenziviranja acetilkkolinesteraze (AchE) te transport natrija (kdr-type). Ovakav mehanizam imaju vrste njema kog ~ohara rezistentne na DDT, organofosfate, karbamate i piretroide (Siegfried et al. 1990; Hemingway et al. 1993). Generalno, pojava rezistentnosti mo~e se sprje iti izbjegavanjem svake pretjerane primjene kemijskih sredstava, dakle zaatitom sukladnom na elima integrirane zaatite. Sredstva treba koristiti samo kada je to doista neophodno potrebno. Va~no je eae mijenjati skupinu upotrebljavanih sredstava. Primjena kombiniranih sredstava koja sadr~e dvije i viae aktivnih tvari, neprekora ivanje preporu enih doza sredstava te uporaba sredstava koja ne akode prirodnim neprijateljima, takoer usporavaju pojavu rezistentnosti (Maceljski i sur. 2002). Mjera koja nije preporu ljiva je poveanje doziranja pri rjeaavanju problema rezistentnosti. Takva praksa koja udvostru uje dozu u svrhu postizanja ekonomske koristi samo poveava problem ukoliko nije postignuto potpuno istrijebljenje. Svaki insekt koji je pre~ivio poveane doze mo~e razviti ak i viai stupanj rezistentnosti od one koja bi se pojavila uz uobi ajeno preporu en tretman. Opis biotestova, zaato su potrebni? Biotestovi su laboratorijski testovi koji slu~e u svrhu potvrivanja rezistentnosti insekata na neki od insekticida. Pojava insekata na koje nije bilo utjecaja u diskriminativnom testu mo~e biti posljedica prisutnosti neuobi ajeno tolerantnih jedinki normalne populacije. Ukoliko su uvjeti ekspozicije, fizioloaki status insekata i doziranje konzistentni, vjerojatnost jednog insekta u seriji od 100 na kojeg ne bi bilo utjecaja je manja od 0.1 (odnosno, manje od jednog u 10 testova). Va~no je utvrditi je li necjelovita reakcija posljedica ovakvih uzroka ili prave rezistentnosti. To se mo~e provjeriti na sljedei na in: Ovaj je test mogue ponoviti koriatenjem drugih uzoraka iste populacije insekata s tog mjesta. Mogunosti slu ajne pogreake kao reakcija odreene jedinke u svakom od sukcesivnih testova progresivno opada (manje od 0.01, 0.001, 0.0001 itd.). Pre~ivljavanje dvije ili viae jedinki ak je i manje vjerojatna. Stoga se kontinuirana pojava omjera jedinki na koje nije bilo utjecaja mo~e razmatrati kao dokaz rezistentnosti. Alternativno se insekte na koje nije bilo utjecaja u diskriminativnom testu mo~e zadr~ati i koristiti za uzgoj buduih generacija. Ako je njihova reakcija posljedica rezistentnosti pokazat e se da zna ajno vei omjer njihovog potomstva nee reagirati na diskriminativnu dozu. Kada ovi testovi poka~u da je neka populacija insekata rezistentna treba provesti opse~no testiranje radi utvrivanja stupnja postojee rezistentnosti, a test metoda sadr~i tri faze: Stvaranje baze podataka iz poznatih rezistentnih sojeva Uporaba diskriminativnih koncentracija za odreivanje rezistentnih sojeva Cjelovito definiranje rezistentnosti i usporedba rezultata mortaliteta na serije koncentracija insekticida rezistentnih i osjetljivih sojeva. Tip testa za utvrivanje rezistentnosti je kontinuirano izlaganje odraslih insekata, bez hrane, insekticidu odreene koncentracije koji se impregnira na filter papir. Dovoljan materijal za preliminarni test je minimalno 80 insekata podijeljenih u dvije grupe po 40. Insekti se broje i stavljaju u posude 1 sat bez hrane na 25 C i oko 70% r.v. Nakon toga se prebacuju na filter papir impregniran insekticidom odreene koncentracije te se bilje~i vrijeme po etka ekspozicije. Nakon to no odreenog perioda ekspozicije, utvruje se mortalitet. Ako nema pre~ivjelih, uzorak se ocjenjuje kao  nerezistentno . U protivnom, ako su uo ene pre~ivjele jedinke poslije perioda ekspozicije, tada se mora provesti test osjetljivosti na gradirane doze insekticida. Pre~ivjele jedinke se stavljaju na razvoj. Njihovi potomci se testiraju istim koncentracijama kao i u diskriminativnom testu. Ako je pre~ivljenje na viaem nivou od normalne osjetljivosti, o ekuje se da e cijelo potomstvo reagirati na diskriminativnu dozu. No, ako oni predstavljaju stvarni rezistentni fenotip, manji dio populacije e biti suzbijen u usporedbi s originalnim uzorkom kojeg smo ispitivali. Ovaj test pru~a korisnu potvrdu rezistentnosti u dvojbenim situacijama. Literatura Batth, S.S. (1977): A survey of Canadian populations of the German cockroach for resistance to insecticides. Canadian Entomologist, 109, 49-52. Bellinger, R.G. (1996): Pest Resistance to Pesticides, publication, Department for Entomology, Clemson University.  HYPERLINK "http://www.ipm.nesu.edu/safety/factsheets/resistance.pdf" www.ipm.nesu.edu/safety/factsheets/resistance.pdf Bennet, G.W. & Spink, W.T. (1968): Insecticide resistance of Geman cockroaches from various areas of Lousiana. Journal of Economic Entomology, 61, 426-431. Cochran, D.G. (1995): Insecticide resistance. Understanding and controlling the German cockroach. (ed. By M. K. Rust, J. M. Owens and D. A. Reierson), pp. 171-192. Oxford University Press, New York. Cornwell, P.B. (1976): The Cockroach. Insecticide and cockroach control, II. Associated Business Programmes, London. Hemingway, J., Small, G.J. & Monro, A.G (1993): Possible mechanisms of organophosphorus and carbamate insecticide resistance in German cockroaches (Dictyoptera: Blattellidae) from different geographicak areas. Journal of Economic Entomology, 86, 1631-1630. Lee, C.Y., Hemingway, J., Yap, H.H. & Chong, N.L. (2000): Biochemical characterization of insecticide resistance in the German cockroach, Blatella germanica, from Malaysia. Medical and Veterinary Entomology, 14, 11-18. Lyon, W.F. (1991): Pet Pest Management, Bulletin 586, Insect Resistance. http://ohioline.osu.edu/b586/b586_6.html Maceljski, M., Cvjetkovi, B., Igrc bar i, J., Ostoji, Z. (2002): Priru nik iz zaatite bilja; Zavod za zaatitu bilja u poljoprivredi i aumarstvu Republike Hrvatske; Hrvatsko Druatvo biljne zaatite Zagreb. 247 stranica. Narayanan, K. 2007: Insect resistance: Its impact on microbial control of insect pests.  HYPERLINK "http://www.fbae.org/Channels/Biotech_in_Plant_Disease_control/insect_resistance.htm" www.fbae.org/Channels/Biotech_in_Plant_Disease_control/insect_resistance.htm Rust, M.K. & Reierson, D.A. (1991): Clorpyrifos resistance in German cockroaches (Dictyoptera: Blattellidae) from restaurants. Journal of Economic Entomology, 84, 736-740. Siegfried, B.D. & Scott, j.g. (1990): Properties and inhibition of acetylcholinesterase in resistant and susceptible German cockroaches (Blatella germanica L.) Pesticide Biochemistry and Physiology, 38, 122-146. Zhai, J. & Robinson, W.H. (1991): Pyrethroid resistance in a field population of German cockroach, Blatella germanica (L.). 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