ࡱ> oy ~bjbj {{*22%%%%%&&&0()$&|+2"....22 2yyyyyyy$gfzI%2&2^222z%%..pQ|6ooo2)%.%.yo2yoowhjx.XO\.x,y|<|Zx2nN jx%jxdo222zzo222|22222222222222 R$: Sveu iliate u Zagrebu Prehrambeno-biotehnoloaki fakultet Preddiplomski studij Biotehnologija Ana Filipovi 6418/BT In vitro evaluacija otpornosti probioti kih sojeva na uvjete gastrointestinalnog trakta ZAVR`NI RAD Modul: Biotehnologija IV Mentor: Doc. dr. sc. Jasna Beganovi Pomo pri izradi: dr. sc. Andreja Leboa Pavunc Zagreb, 16. lipnja 2015. DOKUMENTACIJSKA KARTICA Sveu iliate u Zagrebu Zavrani rad Prehrambeno-biotehnoloaki fakultet Preddiplomski studij Biotehnologija Zavod za biokemijsko in~enjerstvo Labaratorij za tehnologiju antibiotika, enzima, probiotika i starter kultura Ana Filipovi 6418/BT In vitro evaluacija otpornosti probioti kih sojeva na uvjete gastrointestinalnog trakta Sa~etak: Cilj ovog rada je ispitati otpornost probioti kih sojeva iz roda Lactobacillus i Bifidobacterium, izoliranih iz liofiliziranih probioti kih pripravaka, na in vitro uvjete gastrointestinalnog trakta (GIT). Ispitano je pre~ivljavanje bakterijskih sojeva u simuliranom ~elu anom soku i simuliranom soku tankog crijeva, a zatim tolerancija na 0.3% i 0.6% ~u ni soli, te in vitro adhezija na glikoprotein mucin, ato mo~e doprinjeti iskazivanju funkcionalnih u inaka u domainu. Smrtnost probioti kih sojeva tijekom 6 sati izlaganja bakterijskih stanica simuliranim uvjetima GIT je bila u rasponu vrijednosti od 0.94-3.16 logCFU/ml, ato ukazuje na uspjeano pre~ivljavanje ispitivanih sojeva u simuliranim uvjetima GIT. Prema rezultatima, ispitane probioti ke bakterije, s obzirom na soj, su razli ito otporne na testirane koncentracije ~u nih soli. Nakon uzgoja u MRS tekuoj podlozi, pokazalo se da pojedini sojevi, primjerice Lactobacillus reuteri DSM 17938, iskazuju mogunost vezanja na mucin. Na temelju provedenih eksperimenata mo~e se zaklju iti da ispitane probioti ke bakterije uspjeano pre~ivljavaju nepovoljne uvjete GIT-a domaina, ato je jedan od glavnih kriterija za izbor probiotika u svrhu prirodnog uspostavljanja i odr~avanja ravnote~e crijevne mikrobiote. Klju ne rije i: Bifidobacterium, Lactobacillus, probiotici, uvjeti gastrointestinalnog trakta Rad sadr~i: 38 stranica, 6 slika, 3 tablice, 32 literaturna navoda Jezik izvornika: hrvatski Rad je u tiskanom i elektroni kom (pdf format) obliku pohranjen u: Knji~nici Prehrambeno-biotehnoloakog fakulteta, Ka ieva 23, Zagreb Mentor: Doc. dr. sc. Jasna Beganovi Pomo pri izradi: dr. sc. Andreja Leboa Pavunc Rad predan: 08.06.2015. BASIC DOCUMENTATION CARD University of Zagreb Final work Faculty of Food Technology and Biotechnology Undergraduate study Biotechnology Department of Biochemical Engineering Laboratory of Antibiotic, Enzyme, Probiotic and Starter Culture Technologies Ana Filipovi 6418/BT In vitro evaluation of the tolerance of the probiotic strains on the conditions of the gastrointestinal tract Abstract: The purpose of this research is to examine the tolerance of probiotics strains from Lactobacillus and Bifidobacterium genus, isolated from commercial freeze dried probiotic preparations, to in vitro conditions of the gastrointestinal tract (GIT). We observed the survival of bacterial strains in simulated gastric and simulated small intestine juice. In addition, we examined their tolerance to 0.3% and 0.6% bile salts and in vitro adhesion to glycoprotein mucin, the characteristics which can contribute to expression of functional effects in a host. The mortality of probiotic strains during six hours of exposure of the bacterial cells to simulated conditions of the GIT was in the range 0.94-3.16 logCFU/ml, which indicates successful survival of tested strains in simulated conditions of the GIT. According to results, the examined probiotic bacteria tolerated tested bile salt concentrations. After cultivation on MRS broth medium, it was found that certain strains, e.g. Lactobacillus reuteri DSM 17938 show a possibility of adhesion to mucin. Based on conducted research, we can conclude that tested probiotic bacteria successfully survive in inimical conditions in the GIT of a host, which is one of the main criteria for selection of probiotics with a purpose of building and maintaining natural balance of intestinal microbiota. Keywords: Bifidobacterium, Lactobacillus, probiotics, gastrointestinal tract conditions Thesisi contains: 38 pages, 6 figures, 3 tables, 32 references Original in: Croatian Final work in printed and electronic (pdf format) version is deposited in: Library of the Faculty of Food Technology and Biotechnology, Ka ieva 23, Zagreb Mentor: Assist. prof. Jasna Beganovi Technical support and assistance: PhD. Andreja Leboa Pavunc Thesis delivered: 08.06.2015 SADR}AJ  TOC \o "1-4" \h \z \u  HYPERLINK \l "_Toc421381584" 1.UVOD 1  HYPERLINK \l "_Toc421381585" 2. TEORIJSKI DIO 2  HYPERLINK \l "_Toc421381586" 2.1.Osjetljivost bakterija mlije ne kiseline na uvjete niskih pH vrijednosti 2  HYPERLINK \l "_Toc421381587" 2.2.Regulacija unutarstani nog metabolizma aminokiselina 4 2.3. HYPERLINK \l "_Toc421381588" Uvoenje egzogenog biosinteti kog puta 5  HYPERLINK \l "_Toc421381589" 2.4. Prekomjerna proizvodnja proteina geneti kim modifikacijama s ciljem zaatite stanica pri nepovoljnim uvjetima 5  HYPERLINK \l "_Toc421381590" 2.5.Funkcionalnosti stani ne membrane bakterija mlije ne kiseline 7  HYPERLINK \l "_Toc421381591" 2.6. Adaptacija bakterija mlije ne kiseline na stresne uvjete 7  HYPERLINK \l "_Toc421381592" 2.7.Dodatak sastojaka sa zaatitinim u inkom za poboljaanu adaptaciju na kisele uvjete 8  HYPERLINK \l "_Toc421381593" 3. EKPERIMENTALNI DIO 9  HYPERLINK \l "_Toc421381594" 3.1. MATERIJALI 9  HYPERLINK \l "_Toc421381595" 3.1.1.Radni mikroorganizmi 9  HYPERLINK \l "_Toc421381596" 3.1.2.Hranjive podloge 9  HYPERLINK \l "_Toc421381597" 3.1.3.Kemikalije 10  HYPERLINK \l "_Toc421381598" 3.1.4. Aparature i pribor 11  HYPERLINK \l "_Toc421381599" 3.2. METODE RADA 12  HYPERLINK \l "_Toc421381600" 3.2.1.Odr~avanje i uvanje mikroorganizama 12  HYPERLINK \l "_Toc421381601" 3.2.2.Odreivanje tolerancije probioti kih sojeva na ~u ne soli 12  HYPERLINK \l "_Toc421381602" 3.2.3.Pre~ivljavanje probioti kih bakterija u simuliranim uvjetima  HYPERLINK \l "_Toc421381603" gastrointestinalnog trakta 12  HYPERLINK \l "_Toc421381604" 3.2.3.1. Priprava simuliranoga ~elu anog soka i simuliranoga soka tankog crijeva 12  HYPERLINK \l "_Toc421381605" 3.2.3.2. U inak simuliranog ~elu anog soka i soka tankoga crijeva na pre~ivljavanje 12  HYPERLINK \l "_Toc421381606" 3.2.4 In vitro adhezija na protein mucin 13  HYPERLINK \l "_Toc421381607" 4. REZULTATI 14  HYPERLINK \l "_Toc421381608" 5. RASPRAVA 20  HYPERLINK \l "_Toc421381609" 6. ZAKLJU CI 22  HYPERLINK \l "_Toc421381610" 7.LITERATURA 23  TOC \o "1-3" \h \z \u  1.UVOD Bakterije mlije ne kiseline (BMK) se tradicionalno koriste u proizvodnji razli itih fermentiranih proizvoda. Osim toga mnogi sojevi iz razli itih sojeva BMK se zbog dokazanih pozitivnih u inka na zdravlje domaina smatraju probioticima (`uakovi i sur., 2009). BMK su prilikom proizvodnje i primjene u fermentacijama izlo~ene razli itim nepo~eljnim utjecajima okoline, a vrlo esto se radi o visokim koncetracijama vodikovih iona, odnosno kiselim uvjetima mikrookoliaa. Poboljaanje otpornosti BMK na niske koncentracije kiselina mo~e doprinijeti i funkcionalnim u incima probioti kog soja na domaina (Broadebent i sur., 2012; Wu i sur., 2014). Primjena suvremenih metodologija odnosno genomike i funkcionalne genomike omoguuje razumijevanje mehanizama tolerancije ovih bakterijskih sojeva na kisele uvjete prisutne u specifi nom mikrookoliau, a razvijaju se i razli iti pristupi za smanjenje njihove osjetljivosti na kiselost. Poseban naglasak je na modulaciju uvjeta specifi ne stani ne mikrookoline (primjerice razvoj amino-kiselinskog metabolizma, uvoenje egzogenog biosinteti kog kapaciteta i poveana proizvodnja proteina koji ubla~avaju osjetljivost bakterija) i zadr~avanje funkcionalnosti stani ne membrane (Wu i sur., 2014). Kako bi mogle provesti svoj utjecaj, probioti ke bakterije bi trebale biti sposobne pre~ivjeti u gastrointestinalnom traktu (GIT) stoga je nu~no da imaju obrambeni stani ni mehanizam kojim e se oduprijeti niskom pH u ~elucu, probavnim enzimima i ~u nim solima. Otprilike 2,5 L ~elu nog soka i 1 L ~u ne soli se dnevno izlu uju u ljudskom probavnom traktu zbog ega je tolerancija sojeva na te uvjete u GIT va~an selektivni kriterij prilikom odabira probioti kih sojeva. Da bi probioti ke bakterije mogle pre~ivjeti ekstremne uvjete GIT-a, po~eljna je njihova sposobnost adhezije na stanice crijevnog epitela i/ili mukusa poato se time omoguuje produ~eno vrijeme zadr~avanja u crijevima, zatim mogunost kompetitivne ekskluzije patogenih mikroorganizama te interakcije epitelnih i imuno stanica domaina (Jensen i sur., 2012) Cilj ovog zavranog rada je ispitati otpornosti probioti kih sojeva na uvjete GIT-a in vitro, primjenom razli itih eksperimentalnih modela. Bakterijski sojevi odabrani za ova istra~ivanja pripadaju bakterijama iz roda Lactobacillus i Bifidobacterium te su prisutni u razli itim komercijalno dostupnim probioti kih pripravcima namijenjeni za primjenu kod razli itih intestinalnih poremeaja, kod odraslih ljudi, djece ili dojen adi. Osim toga ispitana je i tolerancija odabranih probioti kih sojeva na razli ite koncentracije ~u ne soli i in vitro adhezija na glikoprotein mucin, ato mo~e utjecati na iskazivanje funkcionalnih u inaka u GIT domaina. 2. TEORIJSKI DIO 2.1. Osjetljivost bakterija mlije ne kiseline na uvjete niskih pH vrijednosti Bakterije mlije ne kiseline (BMK) su heterogena grupa Gram-pozitivnih bakterija koje ne proizvode spore i imaju niski udio GC parova baza (pb) u molekuli DNA, a fermentiraju izvore ugljika prvenstveno do primarnog metabolita, mlije ne kiseline (Gaspar i sur. 2013). U prehrambenoj industriji BMK se uglavnom koriste u fermentaciji hrane i pia, ali i proizvodnji bakteriocina i egzopolisaharida (`uakovi i sur., 2010; Zhu i sur., 2009). Uz to, BMK se takoer koriste za proizvodnju niskih koncentracija visoko-kvalitetnih kemikalija, a takoer i razli itih kemikalija standardiziranim procesima poput organskih kiselina, poliola i vitamina u velikim koli inama (Tablica 1.). Tablica 1. Primjena bakterija mlije ne kiseline u industrijskoj proizvodnji PRIMJENAPROIZVODISOJEVI BMKMlije na industrijaSir JogurtLactococcus lactis; Leuconostoc spp. Streptococcus thermophilus; Lb. delbrueckiiFermentacija hraneVino Kiselo povre Soja sos OcatOenococcus oeni PSU-1; Lb. acetolerans L. lactis; Leuconostoc mesenteroides; Lb. plantarum Lb. salivarius; Tetragenococcus halophilus; Lb. fermentum; S. thermophilus Lb. panis; Lb. pontisTekstilne kemikalijeMlije na kiselina 2,3 - butandiol 1,3  propandiol Etanol Jantarna kiselina ButanolLb. rhamnosus; Lb. brevis Lc. lactis Lb. reuteri Lb. plantarum Lb. plantarum Lb. brevisSastojci hraneAlanin -aminobutiri na kiselina Diacetil Acetaldehid BakteriociniLc. lactis Lb. brevis Lc. lactis Lc. lactis Lc. lactis L. lactisNutraceuticiKsilitol Manitol Sorbitol Riboflavin Folna kiselina Vitamin B12 Izoprenoidi FenilpropanoidiLc. lactis Lc. lactis; L. mesenteroides Lb. casei; Lb. plantarum Lc. lactis Lc. lactis Lb reuteri Lc. lactis Lc. lactisPolisaharidiEgzopolisaharid Hijaluronska kiselinaLc. lactis  BMK su esto izlo~ene nepovoljnim uvjetima tijekom industrijske proizvodnje, a isto tako prilikom primjene kod domaina u probavnom traktu (Kos i sur., 2008.). Meu brojnim nepovoljnim uvjetima okoline, pre~ivljavanje BMK pri niskim koncentracijama kiseline u specifi nom mikrookoliau je jedan od glavnih izazova, a upravo je tolerancija bakterijskih stanica na visoke koncetracije kiseline jedan od glavnih kriterija pri izboru probiotika (Prvez i sur., 2006). Tijekom fermentacije, BMK proizvode mlije nu kiselinu koja uzrokuje acidifikaciju, odnosno sni~avanje pH vrijednosti okoline, koje mo~e uzrokovati inhibiciju rasta bakterijskih stanica i mogue odumiranje stanica uslijed ulaska nedisocirane mlije ne kiseline u citoplazmu difuzijom (Serrazanetti i sur., 2009). Mlije na kiselina se unutar stanice razgrauje, utje e na promjenu stani nog pH i koncetraciju aniona citoplazme ato utje e na strukturu baze koja se sastoji od purina i uzrokuje promjenu u strukturi i aktivnosti esencijalnih enzima unutar stanica (Warnecke i Gill, 2005). Stoga, poboljaanje otpornosti BMK na uvjete kiselog okoliaa je od velike va~nosti prilikom industrijske primjene BMK. Kao odgovor na nepovoljne kisele uvjete, BMK su razvile stani ne mehanizme za prepoznavanje kiselih uvjeta koji im omoguuju prilagodbu na ekstremne uvjete i nagle promjene u okolini, odr~avanjem unutarstani ne pH homeostaze, funkcionalnosti stani ne membrane i poveane osjetljivosti proteina stresa ija se aktivnost inducira u nepovoljnim uvjetima (Lebeer i sur., 2008; O Sullivan i Condon 1997) (Slika 1.).  Slika 1. Mehanizmi adaptacije bakterija mlije ne kiseline na kisele uvjete mikrookoliaa Osim toga, pretpostavka je da adaptacija bakterijskih stanica na kiselost (engl. acid tolerance response, ATR) omoguuje zaatitu pri nepovoljnim uvjetima okoline prethodnim izlaganjem stanica umjereno kiselim uvjetima (De Angelis i sur., 2001). Primjena brojnih metoda molekularne biologije doprinose razumijevanju molekularnih mehanizama uklju enih u toleranciju bakterijskih stanica na kiselost, a vjerojatno pri tome va~nu ulogu ima ekspresija specifi nih proteina stresa kao generalni odgovor bakterijske stanice na openito nepovoljne uvjete, zatim odr~avanje funkcionalnosti stani ne membrane i regulacija metabolizma aminokiselina, ato se utje e na poboljaanje otpornost BMK na kisele uvjete (Wu i sur., 2014; Trip i sur. 2012). 2.2 Regulacija unutarstani nog metabolizma aminokiselina Regulacija unutarstani nog metabolizma aminokiselina uobi ajen je mehanizam koji BMK koriste prilikom prisutnosti nepovoljnih uvjeta okoline. Jedan od naj eae opisanih mehanizama regulacije je sustav argininske deiminaze (engl. arginine deiminase ADI) koje BMK koriste pri kiselim uvjetama. Ovim stani nim putem se arginin razgrauje, ato uzrokuje nakupljanje molekula NH3, CO2 i ATP. Nastajanje energije u obliku ATP-a omoguuje istiskivanje protona citoplazme H+-ATP-zom (Beganovi i sur., 2010). Prema pojedinim istra~ivanjima (Wu i sur., 2014) dodatak arginina utjecao je na uspjeanije pre~ivljavanje Lb. casei u kiselim uvjetima, jer je uo ena poveana aktivnost H+-ATPaze i unutarstani ne koncetracije ATP-a. Unutarstani no nakupljanje aspartata je joa jedna reakcija BMK u kiselim uvjetima, jer je uo en prirast bomase i poboljaanje pre~ivljavanja bakterijskih stanica pri niskom pH u prisustvu aspartata (Wu i sur., 2014). Regulacija aminokiselina razgranatog lanca (engl. branched-chain amino acids, BCAA) leucina, izoleucina i valina takoer je jedan od odgovora prilagodbe BMK u uvjetima niskih pH vrijednosti okoline. Uslijed kiselih uvjeta, enzimi IlvA, IlvC2, IlvD i IlvE, koji sudjeluju u metabolizmu aminokiselina razgranatog lanca, prekomjerno se sintetiziraju, a otklanjanje amino grupe iz molekula aminokiselina razgranatog lanca je pretpostavljeni mehanizam za odr~avanje unutarstani nog pH (Ganesan i Weimer, 2004). Takoer, ustanovljeno je da dekarboksilacija atiti stanice od atete uzrokovane kiselinom, stvaranjem ATP-a i utroakom jednog protona (Higuchi i sur., 1997). 2.3. Uvoenje egzogenog biosinteti kog puta Trenutne spoznaje o genomima BMK omoguuju razumijevanje regulacije gena BMK, ato omoguuje manipulaciju tolerancije kiselosti primjenom metaboli kog in~enjeringa. Brojna istra~ivanja upuuju na ulogu glutationa u zaatititi BMK od niza nepovoljnih djelovanja iz okoline poput visokih koncetracija kiseline, prisutnosti kisika, niskih temperatura ili visokih koncetracija soli (Wu i sur., 2014). Kako bi se dodatno istra~ila zaatitna uloga glutationa u nepovoljnim uvjetima, dva gena gshA i gshB, koji kodiraju za -glutamilcistein sintetazu i glutation sintetazu, iz Escherichia coli su eksprimirani u bakteriji L.lactis NZ9000. Prema rezultatima, rekombinantni soj je pokazao veu otpornost u kiselim uvjetima u usporedbi s sojem divljeg tipa (Fu i sur.,2006). Cardoso i sur. (2004.) su pokazali da osjetljivost na kiselost utje e na znatno nakupljanje trehaloze (mikoze) kod bakterije Propionibacterium freudenreichii u kiselim uvjetima. Stoga su Carvalho i sur (2011.) uveli trehaloza de novo biosinteti ki put P. freudenreichii u L. lactis kako bi se ispitao utjecaj proizvodnje trehaloze na toleranciju domainskog soja na kisele uvjete. Mutantni soj je pokazao veu toleranciju na kiselinu, pri pH 3.0 i hladnou, pri 4 C kao i uvjetima poviaene temperature pri 45C. 2.4. Prekomjerna proizvodnja proteina geneti kim modifikacijama s ciljem zaatite stanica pri nepovoljnim uvjetima Sekvencioniranjem genoma i stalnim razvojem naprednih tehnologija doprinosi razvoju otpornih industrijskih mikroorganizama na globalnoj ili sistemskoj bioloakoj razini. Kontinuirano se primjenjuju omi ki pristupi, genomika, transkriptomika, proteomika, metabolomika u kombinaciji s molekularnim tehnikama kako bi se ato bolje razumjeli fizioloaki mehanizmi BMK te se predla~u strategije za poboljaanje fizioloakih funkcija i stvaranje tolerancije na nepovoljne uvjete (Slika 2). Broadbent i sur. (2010) ispitali su adaptaciju L.casei ATCC334 na kisele uvjete primjenom metoda transkriptomike tako ato su bakterijske stanice izlo~li uvjetima kiselog mikrookoliaa te su prema rezultatima pokazali da dva gena koja sudjeluju u malolakti noj fermentaciji jabu ne u mlije nu kiselinu (mleS, malolakti ni enzim; mleP, malat/laktat antiporter tj. antiporter soli jabu ne i mlije ne kiseline) i genska nakupina osam gena za biosintezu histidina (LSEI_1426-1434) su zna ajno eksprimirani. Dodatno su potvrdili dobivene rezultate, dodavanjem 30mM malata ili 30mM histidina u zakiseljenu hranjivu podlogu pH vrijednosti 2.5 u kojem je zabilje~eno stostruko poveanje pre~ivljavanja stanica ovog soja nakon inkubacije tijekom 60 min, i veim od 107 poboljaanjem nakon dva sata (Broadbent i sur., 2010). Slika 2. Pojednostavljeni prikaz primjene omi kih pristupa za bolje razumjevanje fizioloakih mehanizama BMK (preuzeto od Wu i sur., 2014). Bakterije odr~avaju strukturu i aktivnost proteina pri uobi ajenim ili nepovoljnim uvjetima okoline uz pomo kompleksnih regulacijskih mehanizama koji obuhvaaju enzimsku aktivnost skupine regulacijskih proteina (Wu i sur., 2014), meu kojima su molekularni pratioci (engl. chaperons) i proteini popravaka DNA oate enja najdetaljnije istra~eni. Ekspresija dnaK gena iz bakterije E. coli u L. lactis NZ9000 utjecala je na poveanu toleranciju ovog soja na mlije nu kiselinu, NaCl i etanol (Abdullah-Al-Mahin i sur. 2010). Tian i sur. (2012) eksprimirali su gen koji kodira za mali toplinski aok proteina kod bakterije Streptococcus thermophilus u Lc. lactis, nakon ega je rekombinantni soj pokazao zna ajno bolje pre~ivljavanje u razli itim ispitivanim nepovoljnim uvjetima, visokih koncentracija kiseline, poveane temperature, visokih koncentracija etanola, soli i H2O2. Usporedba eksprimiranih proteina primjenom proteomi kih pristupa, pokazala je da je mutantni soj otporniji na visoke koncentracije kiseline u usporedbi s roditeljskim sojem Lb. casei i poja ano eksprimira proteine odgovorne za popravak molekule DNK: MutL, MutS2, UvrC, RecO (Wu i sur., 2014). Mogunost sekvenciranja itavog genoma rezultirala je poveanjem broja kompletno sekvencioniranih genoma BMK ato je temelj za detaljniju karakterizaciju i identificiranje gena koji se induciraju pri nepovoljnim uvjetima okoliaa. Takve spoznaje omoguuju geneti ke modifikacije BMK koje se odlikuju poboljaanom otpornoau pri nepovoljnim uvjetima. Va~no je naglasiti da je primjena bakterijskih sojeva koji su geneti ki modificirani pod posebnom legislativom te su GMO sojevi izlo~eni kriti kom mialjenju javnosti prilikom njihove primjene u industriji. 2.5. Funkcionalnosti stani ne membrane bakterija mlije ne kiseline Barijera svake stanice je stani na membrana koja dijeli unutraanjost stanice od njene okoline i time je prva na udaru prilikom djelovanja nepovoljnih uvjeta iz okoline (Beganovi i sur., 2010). Promjene u stani noj membrani mogu zaatititi bakterijsku stanicu modifikacijom fizikalno-kemijskih karakteristika membrane. U uvjetima visoke koncentracije kiseline, uo ena je poveana fluidnost stani ne membrane i udio jednostavnih nezasienih masnih kiselina, kao i poveana prosje na du~ina lanca kod bakterije Lb. casei (Wu i sur., 2014). Stoga, proizvodnja nezasienih masnih kiselina mo~e biti potencijalna metoda za poboljaanje tolerancije kiselosti u BMK. FabM, enzim odgovoran za proizvodnju jednostavnih nezasienih masnih kiselina, identificiran je kod S. mutans, a delecijski mutant u tom genu, je iznimno osjetljiv na visoke koncentracije kiseline u usporedbi s divljim tipom (Fozo i Quivey, 2004). Tipi an fenotip osjetljivosti ovog soja na kiselinu se smanjivao prilikom rasta u prisustvu jednostavnih nezasienih masnih kiselina (Fozo i Yuivey, 2004). Takoer, proizvodnja ciklopropanskih masnih kiselina bila je openiti odgovor na kisele uvjete (Broadbent i sur. 2010; Wu i sur., 2014). Ipak prema istra~ivanjima To i sur. (2011) sa sojevima L. lactis subsp. cremoris, soj divljeg tipa i cfa mutantu, ciklopropanacija nezasienih masnih kiselina nije neophodna za pre~ivljavanje u kiselim uvjetima. Stoga je potrebno daljenje ispitivanje mehanizama zaatitnog u inka ciklopropanskih masnih kiselina u kiselim uvjetima. 2.6. Adaptacija bakterija mlije ne kiseline na stresne uvjete Kao odgovor na nepovoljne uvjete okoline, BMK koriste sofisticirane mehanizme kako bi se adaptirale na stres. Kod veine bakterijskih sojeva uo eni su sli ni mehanizmi adaptacije iako su bakterijske stanice bile izlo~ene razli itim nepovoljnim uvjetima te je ustanovljeno da su ti stani ni mehanizmi meusobno povezani. Primjena predhodne prilagodbe, odnosno prethodno tretiranje soja s ubla~enim nepovoljim uvjetima, mo~e poboljaati otpornost bakterijskih sojeva prilikom ekstermnih nepovoljnih uvjeta, a i tkz. kombinirana zaatita, odnosno mogunost da jedna vrsta tolerancije na nepovoljne uvjete mo~e omoguiti zaatitu i protiv drugih nepovoljnih uvjeta. Broadbent i sur. (2010) su prethodnim izlaganjem bakterijskih stanicama pH vrijednostima u iznosu od 4.5 tijekom 20 minuta postigli poboljaanje pre~ivljavanja L. casei ATCC 334 pri izrazito kiselim uvjetima, odnosno pri pH 2.0. Zna ajno poveanje pre~ivljavanja bakterijskih stanica u izrazito kiselim uvjetima (pH 3.9) postignuto je i prethodnim izlaganjem stanica L. lactis subsp. lactis tijekom 30 minuta blago kiseloj okolini pH vrijednosti 5.5 (Hartke i sur. 1996). Kombinirana zaatita takoer se smatra u inkovitim pristupom poveanja stupnja tolerancije na kiselinu u BMK. Primjerice soj L. plantarum prethodno izlo~en ni~im visokim temperaturama pokazao je povean rast pri pH vrijednosti od 5.0 (De Angelis i sur., 2004). Prema iznesenom, predhodno izlaganje stanica blago kiseloj sredini potaknulo je toleranciju na visoke koncentracije kiseline ato je doprinjelo i zaatiti stanica i pri drugim nepovoljnim uvjetima. Iako je o ito da prethodna prilagodba i kombinirana zaatita doprinose zna ajnom poboljaanju pre~ivljavanja BMK u kiselim uvjetima, to ni molekularni mehanizmi u bakterijskoj stanici koji se pri tome odvijaju nisu u potpunosti jasni i potrebna je njihova daljna karakterizacija. 2.7. Dodatak sastojaka sa zaatitinim u inkom za poboljaanu adaptaciju na kisele uvjete Dodatak egzogenih protektivnih tvari, odnosno sastojaka sa zaatitinim u inkom, jedan je od pristupa za zaatitu stanica BMK u kiselim uvjetima koji poboljaava njihovu toleranciju na kiselinu. Istra~eni su brojni protektivni sastojci za zaatitu BMK u kiselim uvjetima uklju ujui aminokiseline, masne kiseline i aeere. Tako dodatak aminokiseline arginina poveava pre~ivaljavanje L. casei pri niskim pH vrijednostima (Wu i sur., 2014). Fizioloaka analiza pokazala je da arginin doprinosi izdr~ljivosti stanica u kiseloj okolini jer utje e na poveanu aktivnost H+-ATP-aza i unutarstani nu razinu ATP-a. Openito, regulacija arginin deiminaznim sustavom je u estali sustav adaptacije bakterija uklju ujui BMK na kiselinu, (Ryan i sur., 2009; Beganovi i sur., 2010;). Istra~ivanja adaptacije L. casei na kisele uvjete, uz dodatak aspartata u MRS hranjivu podlogu, utjecao je na prirast biomase i pre~ivljavanje stanica u kiselim uvjetima (Wu i sur., 2014). Analizom unutarstani nog sadr~aja, ustanovljena je poveana koncentracija intermedijera koji sudjeluju u glikolizi i ciklusu limunske kiseline, a kod bakterije L. casei je promijenjen metaboli ki put jer je povean pritok iz aspartata prema argininu (ADI sustav), a smanjen pritok iz aspartata prema asparaginu (Wu i sur., 2014). Kod bakterije L. rhamnosus dodatak Tween-80 u podlogu za rast rezultirao je 1000-strukim poveanjem pre~ivljavanja prilikom izlaganja bakterijskih stanica simuliranim sokovima GIT (Corcoran i sur., 2007). Analizom sastava masnih kiselina bakterije L. rhamnosus uo en je 55-erostruko vei sadr~aj oleinske kiseline i znatno vei omjer nezasienih/zasienih masnih kiselina u membrani stanica u prisustnosti Tween-80 (Corcoran i sur., 2007). Navedeni rezultati upuuju da dodatak sastojaka sa zaatitnim u inkom mo~e biti u inkovita strategija za poboljaanje rezistencije bakterija mlije ne kiseline u uvjetima niskih pH vrijednosti. 3. EKSPERIMENTALNI DIO 3.1. MATERIJALI 3.1.1. Radni mikroorganizmi Svi mikroorganizmi koriateni u ovom radu u in vitro eksperimentima, navedeni su u Tablici 2. Radi se o probioti kim bakterijama iz rodova Lactobacillus ili Bifidobacterium koje su prisutne u obliku liofiliziranih probioti kih pripravaka na tr~iatu. Navedeni sojevi pohranjeni su u Zbirci mikroorganizama Laboratorija za tehnologiju antibiotika, enzima, probiotika i starter kultura Prehrambeno-biotehnoloakog fakulteta, Sveu iliata u Zagrebu. Sojevi se uvaju pri -80 C u MRS tekuoj hranjivoj podlozi uz dodatak 15% (v/v) glicerola. Tablica 2. Bakterijski sojevi izolirani iz liofiliziranih probioti kih pripravaka koriateni u ovom zavranom radu Bakterijski sojeviOznake sojevaUvjeti rastaLactobacillus rhamnosus LGG -11MRS, 370C, aerobnoLactobacillus sp. 2A2AMRS, aerobnoBifidobacterium sp. 2B2BTOS, 37 0C, anaerobnoBifidobacterium longum 33TOS, 37 0C, anaerobnoLactobacillus reuteri DSM 17938 -44MRS, 37 0C, anaerobnoLactobacillus sp. 5/15/1MRS, 37 0C, anaerobnoBifidobacterium sp. 6B6BTOS, MRS, 37 0C, anaerobnoLactobacillus sp. 8/18/1TOS, MRS, 37 0C, anaerobnoBifidobacterium sp. 8/28/2TOS, MRS, 37 0C, anaerobnoL. reuteri Protectis (DSM 17938) -99MRS, 37 0C, anaerobnoLactobacillus rhamnosus GR1GR1MRS, 37 0C, anerobnoLactobacillus reuteri RC14RC14MRS, 37 0C, anerobnoBifidobacterium bifidum BB12BB12TOS, 37 0C, anerobno 3.1.2. Hranjive podloge U radu su koriatene optimalne hranjive podloge za odr~avanje, uvanje i uzgoj bakterija mlije ne kiseline iz roda Lactobacillus i bakterija iz roda Bifidobacterium: - MRS kruta hranjiva podloga - Man-Rogosa-Sharpe sastava: pepton 15,0 g/l; mesni ekstrakt 3,0 g/l; kvaa ev ekstrakt 5,0 g/l; glukoza 20,0 g/l; Tween 80 1,0 g/l; MgSO4 x 7 H2O 0,05 g/l; MnSO4 x 7 H2O 0,05 g/l; natrijev-acetat 5,0 g/l; agar 20,0 g/l, u destiliranoj vodi, pH vrijednost podloge je 6,5; sterilizacija se provodi pri 121C/15 min. - MRS - Man-Rogosa-Sharpe tekua hranjiva podloga istog je sastava kao MRS-agar, samo bez dodanog agara. - TOS  propionat agar: pepton iz kazeina 10 g/l; kvaa ev ekstrakt 1,0 g/l; kalijev dihidrogen fosfat 3,0 g/l; kalijev hidrogen fosfat 4,8 g/l; amonijev sulfat 3,0 g/l; MgSO4 x 7H2O 0,2 g/l; L-cistein monoklorid 0,5 g/l; natrijev propionat 15,0 g/l; galaktooligosaharidi (TOS)10,0 g/l; agar 15,0 g/l; u destiliranoj vodi. pH vrijednost podloge iznosi 6,7; sterilizacija se provodi pri 115C/15 min. 3.1.3. 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METODE RADA 3.2.1. Odr~avanje i uvanje mikroorganizama Probioti ki sojevi su uvani pri -80C u MRS tekuoj hranjivoj podlozi uz dodatak 15 % (v/v) glicerola. Dan prije eksperimenta probioti ki sojevi su inokulirani u svje~u optimalnu hranjivu podlogu te inkubirani pri optimalnoj temperaturi rasta, pri uvjetima navedenim u Tablici 2. 3.2.2 Odreivanje tolerancije probioti kih sojeva na ~u ne soli Tolerancija probioti kih sojeva na ~u ne soli je ispitana na dva na ina. Prvom metodom se prekonono uzgojeni bakterijski sojevi inokuliraju u mikrotitarske plo ice s 96 ja~ica koje sadr~e 200 (L MRS+0.3% ili MRS+0.6% konjugiranih ~u nih soli natrija u tri paralele. Rast inokuliranih sojeva se prati kroz 7 sati mjerenjem opti ke gustoe pri 620 nm (OD620). Rast probioti kih sojeva u MRS hranjivoj podlozi je kontrola rasta. Drugom metodom se prati rast probioti kih sojeva u 5 mL tri prethodno navedena tekua medija. Svakih sat vremena se mjeri OD620 te odreuje broj poraslih bakterijskih kolonija (CFU/ml) na MRS agaru u anaerobnim uvjetima pri 37C nakon 48 sati inkubacije. 3.2.3 Pre~ivljavanje probioti kih bakterija u simuliranim uvjetima gastrointestinalnog trakta 3.2.3.1 Priprava simuliranoga ~elu anog soka i simuliranoga soka tankog crijeva Simulirani ~elu ani sok je pripravljen suspendiranjem pepsina (3 g/L) u 0,5% otopini natrijevog klorida, kojoj je pH podeaen na 2,0 s koncentriranom klorovodi nom kiselinom. Simulirani sok tankoga crijeva je pripravljen suspendiranjem pankreatina (1 g/L), samog ili/i ~u nih soli (3,0 mg/mL govee ~u i) u 0,5% otopini natrijeva klorida, kojoj je pH podeaen na 8,0 s natrijevom lu~inom (Kos, 2001). 3.2.3.2 U inak simuliranog ~elu anog soka i soka tankoga crijeva na pre~ivljavanje probioti kih bakterija Suspenzije stanica odabranih bakterijskih kultura su izlo~ene djelovanju ~elu anog soka (pH = 2) 2 sata, a zatim su centrifugirane pri 3500 o/min i resuspendirane u simuliranom soku tankoga crijeva (3 mg/mL govee ~u i) tijekom 3 sata (Kos, 2001). Broj poraslih ~ivih stanica odreivan je indirektnom metodom. 3.2.4 In vitro adhezija na protein mucin Vezanje probioti kih sojeva na imobilizirani mucin je provedeno na mikrotitarskim plo icama s 96 ja~ica, koje su inokulirane bakterijskim stanicama poraslim do eksponencijalne faze rasta. U ja~ice mikrotitarskih plo ica se prethodno, dakle prije dodatka suspenzija bakterijskih stanica, dodaje glikoprotein mucin u koncentraciji od 50 g/mL u karbonantnom/bikarbonantnom puferu (50 mmol/L, pH 9.6) te se inkubira preko noi pri 40C. Potom se ja~ice isperu tri puta s fosfatnim puferom, te se mucin imobilizira tijekom sat vremena dodatkom 1% Tween 20 u fosfatnom puferu. 100 l suspenzije svakog soja, kojoj je podeaena OD620 na 1 u fosfatnom puferu, se doda u svaku ja~icu mikrotitarske plo ice koje se zatim inkubiraju preko noi pri 370C. Nakon uklanjanja neadosrbiranih odnosno nevezanih stanica (tri uzastopna ispiranja s 200 l 0.05% Tween 20 u fosfatnom puferu) mikrotitarske plo ice se suae pri sobnoj temperaturi i vezane (adhezirane) stanice se detektiraju bojanjem kristal violetom. Potom se mikrotitarske plo ice isperu s 100 ml citratnog pufera (50 mmol/L, pH 4.0) i odreuje se adsorbancija na ita u mikrotitarskih plo ica LKB 5060-006,  GDV pri 620 nm. Takoer je ispitana i in vitro adhezija probioti kih sojeva na mikrotitarskim plo icama s 96 ja~ica koje ne sadr~e vezani mucin kao kontrola u svim eksperimentima. Eksperimenti su ponovljeni tri puta kako bi se izra unala srednja vrijednost i standardna devijacija. 4. REZULTATI Mikroorganizmi prisutni u GIT ovjeka, ine intestinalnu mikrobiotu, koja zna ajno utje e na metabolizam i fiziologiju domaina, njegov imunoloaki sustav i otpornost na infekcije. Meutim, u GIT su prisutni nepovoljni uvjeti, u usporedbi s uvjetima optimalnim za rast pojedinih bakterijskih sojeva, poput niskog pH, brzog protoka i toksi nih ~u nih soli. Pre~ivljavanje nepovoljnih uvjeta u GIT neophodna je karakteristika probioti kih sojeva. Glavnu prepreku njihovom pre~ivljavanju osim kisele pH vrijednosti u ~elucu, ine i probavni enzimi poput pepsina ili pankreatina (Kos i sur., 2003). Stoga je kao bitno funkcionalno svojstvo, ispitana sposobnost pre~ivljavanja bakterijskih sojeva iz rodova Lactobacillus i Bifidobacterium u simuliranim uvjetima GIT. Odnosno praeno je njihovo pre~ivljavanje u simuliranom ~elu anom soku, pri pH vrijednosti 2,0 te u simuliranom soku tankog crijeva gdje se izlu uje sok guatera e i ~u . Ovisno o bakterijskom soju, smrtnost bakterijskih stanica izra~ena kao CFU/ml, izra unata kao razlika po etnog broja bakterijskih stanica i broja bakterijskih stanica odreenog nakon njihovog izlaganja nepovoljnin uvjetima GIT, se razlikovala (Tablica 3.). Smrtnost probioti kih sojeva tijekom 6 sati izlaganja bakterijskih stanica simuliranim uvjetima GIT je u rasponu vrijednosti od 0.94 - 3.16 log CFU/ml. Pri tome je za soj Bifidobacterium longum 3, u usporedbi s preostalim sojevima bila najvea, a za soj Lactobacillus 2A najmanja. No va~no je naglasiti, da eksperimentalno odreene vrijednosti smrtnosti bakterijskih sojeva, u ovom radu, se mogu okarakterizirati kao uspjeano pre~ivljavanje sojeva, s obzirom na broj CFU/ml koji je deklariran na probioti kim pripravcima. Tolerancija na visoke koncentracije ~u i jedan je od najva~nijih svojstava probioti kih bakterija, budui da ima u inak na pre~ivljavanje u tankom crijevu, a time i na njihove funkcionalne uloge u GIT (Ruiz i sur., 2013). Tako da je ispitana otpornost bakterijskih sojeva iz rodova Lactobacillus i Bifidobacterium na ~u ne soli, nacjepljivanjem sojeva na optimalne vrste i tekue MRS podloge s 0.3% ili 0.6% ~u nih soli te je usporeen njihov rasta s brojem poraslih bakterijskih kolonija na kontrolnim MRS podlogama pri optimalnim uvjetima (Slike 3-5). Prema rezultatima, ato je bilo i za o ekivati, dodatak odreene koncentracije ~u nih soli u MRS hranjivu podlogu za uzgoj utjecao je na smanjenje prirasta biomase u usporedbi s kontrolom, odnosno rastom odreenog soja u MRS hranjivoj podlozi (Slika 3). Izmeu ispitivanih sojeva odabrana su tri probioti ka soja koja najuspjeanije podnose dodane koncentracije ~u nih soli u MRS tekuoj hranjivoj podlozi, tijekom 7 sati inkubacije pri 37C, i radi se o Lactobacillus rhamnosus LGG, Bifidobacterium longum 3 i Lactobacillus rhamnosus GR-1 (Slika 4). Tim sojevima je odreeno i pre~ivljavanje, osim praenjem apsorbancije uslijed zamuenja podloge za rast, nacjepljivanjem alikvota stanica tijekom 7 sati (svaki sat vremena) na MRS kruti hranjivi agar (Slika 5). Dobiveni rezultati bili su u skladu s rezultatima eksperimenata dobivenim praenjem apsorbancije. Pri izboru sojeva za probioti ku upotrebu, jedan od va~nih kriterija je mogunost adhezije za crijevni epitel (Kos i sur., 2003; `uakovi i sur., 2009). Adhezija probioti kih sojeva na intestinalne epitelne stanice pokazala se kao po~eljno svojstvo, stoga je ispitana in vitro adhezija na glikoprotein mucin probioti kih sojeva iz rodova Lactobacillus i Bifidobacterium (Slika 6). Isti eksperiment je proveden na mikrotitarskim polo icama s 96 ja~ica koje ne sadr~e imobiliziran glikoprotein mucin, kao kontrolni test. Mucini su glikoproteini sluznice i dio su dinami nog, interaktivnog obrambenog sustava sluznice, te su prisutni na povraini sluznice GIT gdje predstavljaju mukozalnu barijeru izmeu lumena i intestinalnih epitelnih stanica. U usporedbi s kontrolom, ispitani sojevi veinom nemaju zna ajnu sposobnost vezanja in vitro na glikoprotein mucin. Ipak, prema rezultatima, isti u se probioti ki sojevi Bifidobacterium sp. 8/2, Bifidobacterium sp. 6B, Lactobacillus reuteri DSM 17938-4 i Bifidobacterium sp. 2B koji iskazuju mogunost interakcije s ovim glikoproteinom. Tablica 3. Pre~ivljavanje bakterijskih kolonija poraslih na MRS krutoj hranjivoj podlozi nakon izlaganja stanica simuliranim uvjetima GIT (direktan prijelaz iz simuliranog ~elu anog soka u simulirani sok tankog crijeva) Bakterijski sojeviPrijelaz iz ~elu anog soka u sok tankog crijeva (CFU/mL)}elu ani sok (t=0 h)}elu ani sok (t=2 h)Sok tankog crijeva (t=4 h)Smrtnost stanica log CFU/mL (0-6h)Lactobacillus sp. 13,41x1081,64x1082,63x1071,11Lactobacillus sp. 2A3,53x1088,14x1072,96x1071,08Lactobacillus sp. 2B2,12x1085,21x1069,78x1052,34Bifidobacterium sp. 32,52x1087,496x1071,75x1053,16Lactobacillus sp. 41,43x1083,83x1071,45x1062,00Lactobacillus sp. 5/13,89x1083,41x1071,04x1062,57Lactobacillus sp. 6B1,94x1085,84x1064,25x1052,6Lactobacillus sp. 7A3,65x1081,03x1084,13x1070,94Lactobacillus sp. 8/14,38x1088,48x1077,48x1061,77Lactobacillus sp. 8/22,71x1081,04x1083,41x1061,9Lactobacillus sp. 91,84x1086,13x1072,15x1061,93  Slika 3. Usporedba rasta bakterijskih sojeva u optimalnoj MRS tekuoj podlozi i u istoj tekuoj podlozi uz dodatak 0.3% ili 0.6% ~u nih soli. Sivo su ozna eni bakterijski sojevi koji bolje pre~ivljavaju dodatak ~u nih soli u usporedbi s preostalim ispitivanim probioti kim bakterijama.    Slika 4. Rast probioti kih sojeva a) Lb. rhamnosus LGG i b) B. longum 3 c) Lb. rhamnosus GR-1 u MRS tekuoj podlozi, te u MRS tekuoj podlozi uz dodatak 0.3% ili 0.6% ~u nih soli, praen tijekom 7 sati inkubacije bakterijskih sojeva pri 37 C.  Slika 5. Pre~ivljavanje bakterijskih kolonijaa) Lb. rhamnosusLGG b) Lb. rhamnosus GR-1 i c) B. longum 3, poraslih na MRS krutoj hranjivoj podlozi nakon inokulacije u MRS tekuu podlogu, te u MRS tekuu podlogu s 0.3% ili 0.6% ~u nih soli, odreeno tijekom 7 sati inkubacije bakterijskih sojeva pri 37 C. a)  EMBED Microsoft Excel Chart  b)  EMBED Microsoft Excel Chart  Slika 6. In vitro adhezija probioti kih sojeva ispitana na mikrotitarskim plo icama s 96 ja~ica a) kontrola; prazne ja~ice b) mucin; ja~ice sadr~e imobilizirani mucin. 5. RASPRAVA Mikrobiota gastrointestinalnog trakta (GIT) ovjeka nu~na je za pravilno funkcioniranje organizma zbog niza va~nih u inaka ove kompleksne mikrobne zajednice na zdravlje ovjeka (`uakovi i sur., 2009). Neki od tih pozitivnih utjecaja su modulacija imunoloakog sustava, zaatita od karcinoma debelog crijeva i crijevnih infekcija izazvanih prisutnoau bakterija E. coli, Clostridium, Salmonella, Listeria, Shigella, sinteza bakteriocina, sinteza vitamina B kompleksa i proteoliti kih enzima te poticanje bolje resorpcije minerala meu kojima su najva~niji kalcij, magnezij i ~eljezo. Meutim, mikrobiota GIT mo~e imati i ne~eljen u inak na ljudski organizam kada nastupi naruaavanje njene ravnote~e zbog prisutnosti patogenih mikroorganizama i dugotrajne terapije antibioticima. Upravo zbog toga se primjenjuju probioti ki sojevi koji su veinom okarakterizirani iz rodova BMK, prisutni u funkcionalnoj hrani ili kao probioti ki pripravci kako bi se uspostavila ponovna ravnote~a naruaene mikrobiote GIT (Parvez i sur., 2006; `uakovi i sur., 2009; Jensen i sur., 2012). Uvjeti niskih pH vrijednosti, odnosno pre~ivljavanje u uvjetima kiselog mikrookoliaa je vrlo esto izazov za BMK (Broadbent i sur., 2012; Wu i sur., 2014). Osim toga poboljaanje otpornosti probioti kih sojeva na visoke koncentracije kiseline klju no je u primjeni BMK kao bioterapeutika (De Angelis i sur., 2001; Kos i sur., 2003; `uakovi i sur., 2009). Sekvenciranje genoma bakterijskih sojeva koji pripadaju rodovima BMK doprinosi razumijevanju molekularne biologije BMK, a razvoj i primjena post-genomi kih pristupa ubrzali su identifikaciju gena i proteina koji su klju ni u odgovoru i adaptaciji bakterijskih stanica na nepovoljne uvjete. Ove spoznaje doprinose razvoju stategija koje e omoguiti poveanu otpornost BMK u nepo~eljnim uvjetima, a ato je osobito zna ajno prilikom iskazivanja funkcionalnosti probioti kih sojeva u GIT ovjeka. `toviae, specifi ni uvjeti fermentacije mogu se primijeniti na temelju razumjevanja fizioloakih odlika BMK kako bi im se poveala tolerancija na nepovoljne uvjete. Navedeni sistemski bioloaki pristup uz primjenu mikrobioloakih i molekularnih tehnika omoguuju razvoj BMK s poveanom otpornoau na ne~eljne uvjete ato rezultira i poveanjem funkcionalnosti probioti kih, ali i industrijskih va~nih starter kultura (Gaspar i sur., 2013). Pre~ivljavanje u nepovoljnim uvjetima prisutnim u GIT-u je jedan od najva~nijih zahtjeva koji sojevi BMK moraju zadovoljiti kako bi mogli biti kategorizirani kao probioti ki sojevi koji iskazuju pozitivne u inke na zdravlje domaina bez nepo~eljnih nuspojava i autolize (Serrazanetti i sur., 2009). Zato je ispitano ponaaanje i pre~ivljavanje izoliranih probioti kih sojeva roda Lactobacillus i Bifidobacterium in vitro u simuliranom ~elu anom soku i simuliranom soku tankog crijeva te tolerancija na ~u ne soli. Adhezija na povrainu epitelnih stanica bitan je preduvjet za kolonizaciju probiotioti kih sojeva u GIT i omoguava djelovanje probiotika na crijevnu mikrofloru i imunoloaki sustav domaina (Kos i sur., 2003), stoga je ispitana in vitro adhezija na glikoprotein mucin. Mucin je prisutan kao jedna od strukturnih komponenta na povraini sluznice GIT, gdje ini dio intestinalne barijere, te postoji mogunost da ova molekula ima ulogu receptora za adhezine probioti ke bakterije. Primjenom eksperimentalnih metodama u ovom radu se ustanovilo da su ispitane probioti ke bakterije otporne na ~u ne soli i sposobne za rast u njihovoj prisutnosti, ato se vidi iz prilo~enih grafova (Slike 3 do 5). Izmeu ispitanih sojeva, Lactobacillus rhamnosus LGG, Bifidobacterium longum i Lactobacillus rhamnosus GR-1 pokazuju najbolju toleranciju na ~u ne soli. }u  igra va~nu ulogu u fiziologiji crijevnih bakterija i utje u na njihovu funkcionalnost. To je posebno va~no za probioti ke bakterije jer njihovi pozitivni u inci ne smiju biti ometani u prisustvu ove fizioloake tekuine. Prisutnost ~u nih soli utje e na aktivnosti intestnalnih laktobacila i bifidobakterija te su ak neke od tih aktivnosti, kao ato je asimilacija kolesterola, izravno povezani s metabolizmom ~u ne soli kod tih bakterija. Razumijevanje mehanizama zbog kojih probioti ke bakterije mogu pre~ivjeti stres uzrokovan ~u nim solima joa uvijek se istra~uju (Ruiz i sur., 2013). Svi ispitani sojevi pre~ivljavaju u ~elu anom soku i soku tankog crijeva, ato zna i da ujedno dobro podnose niske pH vrijednosti prisutne u tim uvjetima. Treba naglasiti da sojevi Lactobacillus rhamnosus LGG -1, Lactobacillus sp. 2A i Lactobacillus sp. 8 iskazuju najmanju smrtnost (log CFU/ml) odnosno najotporniji su na simulirane uvjete GIT dok su bakterije Bifidobacterium longum, Lactobacillus sp. 5/1 i Bifidobacterium sp. 6B neato osjetljivijije na prisutne uvjete. Ispitani sojevi veinom nemaju zna ajnu sposobnost vezanja in vitro na glikoprotein mucin. Ipak probioti ki sojevi Bifidobacterium sp. 8/2, Bifidobacterium sp. 6B, Lactobacillus reuteri DSM 17938-4 i Bifidobacterium sp. 2B iskazuju mogunost interakcije s ovim glikoproteinom. Adhezija na povrainu intestinalne sluznice interakcijom bakterijskih stanica s mucinom ili proteinima ekstracelularnog matriksa mo~e biti va~an preduvjet za kolonizaciju probioti kih sojeva u intestinalnom lumenu, koji mo~e omoguiti ovim bakterijama i kompetitivnu prednost pred drugim mikroorganizmima u takvom ekosustavu (`uakovi i sur., 2010) Dakle, dobiveni rezultati provedenog ispitivanja su pokazali da ispitane bakterije iz roda Lactobacillus i Bifidobacterium, zadovoljavaju va~an uvjet prilikom selekcije probioti kih sojeva jer pre~ivljavaju u nepovoljnim uvjetima GIT-a domaina (niski pH, visoke koncetracije ~u ne soli, prisutnost probavnih enzima) i pri tome pojedini sojevi iskazuju mogunost adhezije na mucin in vitro; ato je jedan od glavnih kriterija za izbor probiotika u svrhu prirodnog uspostavljanja i odr~avanja ravnote~e crijevne mikrobiote. U kona nici, ispitivanje otpornosti na uvjete GIT sojeva iz roda Lactobacillus i Bifidobacterium pokazuje da su svi sojevi tolerantni na ekstremne uvjete GIT odnosno pre~ivljavaju u kiseloj sredini, pri visokim koncetracijama ~u ne soli i u prisutnost probavnih enzima. Navedeno dokazuje da ovi sojevi zadovoljavaju va~an selektivni kriterij za odabir probioti kih sojeva te se mogu koristiti kao funkcionalni dodaci prehrani ili terapijski pripravci s ciljem poboljaanja svojstava crijevne mikrobiote i odr~avanje njihove ravnote~e. Bakterije iz roda Lactobacillus i Bifidobacterium, osim ato su va~ni industrijski mikroorganizmi koji imaju mogunost biotehnoloake proizvodnje specifi nih metabolita, uz to su prepoznate kao va~ne probioti ke kulture, koje iskazuju po~eljne funkcionalne u inke na organizam domaina, ato otvara mogunost njihove primjene kao bioterapeutika (`uakovi i sur., 2010; Gaspar i sur., 2013). Imaju brojne mogunosti u industrijskoj proizvodnji i airok spektar primjene, od prehrambene industrije kao starter kulture ili funkcionalni dodaci prehrani te do medicinske primjene kao bioterapeutici, ato je sve atraktivnija tema brojnih znanstvenih istra~ivanja. 6. ZAKLJU CI 1. Unato  tome ato nizak pH ~eluca nepovoljno utje e na pre~ivljavanje bakterijskih sojeva, rezultati pokazuju da je broj ~ivih stanica nakon izlaganja simuliranom ~elu anom soku i soku tankog crijeva ostao visok kod svih ispitanih sojeva. Iako su neki sojevi imali vei udio smrtnosti od drugih, mo~e se zaklju iti da ispitivani sojevi uspjeano pre~ivljavaju simulirane uvjete GIT. 2. Ispitani probioti ki sojevi su otporni na 0.3% i 0.6% koncentracije ~u ne soli, meu kojima se isti u Lactobacillus rhamnosus LGG, Lactobacillus rhamnosus GR-1 i Bifidobacterium longum. 3. Sojevi Bifidobacterium sp. 8/2, Bifidobacterium sp. 6B, Lactobacillus reuteri DSM 17938-4 i Bifidobacterium sp. 2B pokazuju visok stupanj adhezije in vitro na glikoprotein mucin, ato mo~e upuivati na postojanje adhezina na povraini bakterijskih stanica koji specifi no prepoznaju ovaj glikoprotein. 7. LITERATURA Al-Mahin A,Sugimoto S, Higashi C, Matsumoto S, Sonomoto K (2010) Improvement of multiple-stress tolerance and lactic acid production in Lactococcus lactis NZ9000 under conditions of thermal stress by heterologous expression of Escherichia coli dnaK. Appl Environ Microb 76(13), str 4277 4285 Beganovi J., Guillot A., van de Guchte M., A. Jouan A., C. Gitton, V. Loux,K. Roy, S. Huet, H. Monod, V. Monnet (2010) Characterization of the Insoluble Proteome of Lactococcus lactis by SDS-PAGE LC-MS/MS Leads to the Identification of New Markers of Adaptation of the Bacteria to the Mouse Digestive Tract. 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