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CDDDDDBEDEEEF F6FNFRFGGLHTHVHnHpHrHHHHIXIZI`IIIIII0JHJVJpJvJJJJJJJK»ͭ饡 hbUhqL hbUhbUhbUhbU\hK hbUhL h!)h h!)hm h!)hSho h!)ho h!)hq h!)hc h!)he h!)hX h!)hQ'Y h!)h;Fh!)h;F64JJJJJJJJJJJJJJJJJJJJJJJJKKK$a$gdgegdgegdbU`gdK gdbUKKK"K8K:KHKtKKKKKKKKJLfLLLLLL&M8M>MHMbMMMMMMNVNPPPPPQQQQ8Q:QPQRQQQQۼh<h-h<5B*phh-hj5B*ph hj5h'l h'lh'lhjhj5 h'l5h8hmhm6h4 hV*&hV*&6]hmhV*& hqL5hqLh.#'hge5CJ aJ hbUhge1K8K:KrKtKPPQQ:QRQQ$$If`a$gd-`gd]X\gd53`gd.#'gd<`gd.#' QQQQQG777$$If`a$gd-kd$$IfTlF Fd NJ  tfff06    44 lapfffyt-TQQQQQQQQQQ$R&R,R.RXR\RdRRRRRRRRRRRRR2S4SSSSSS T TTTTT*T.TnTzT|TTTTŽ繵㵱}} hd80J&hKhK0J(hKhK0J'hKhK0J&hmhm6hK hmhmhmhV*&h.#'h-hh*6h-h'l6h-h6 hyuhyu h'lh'lh'lh<hjh-h<6h-hj61QQ&R.RZR\LLL$$If`a$gd-kd|$$IfTlF Fd NJ t06    44 lapyt-TZR\RRRR\LLL$$If`a$gd-kd8$$IfTlF Fd NJ t06    44 lapyt-TRRRRRS\WNIDgdKgdV*&`gd.#'gd<kd$$IfTlF Fd NJ t06    44 lapyt-TTTTTTUUUU"U$U(U8UDUFUHUJULUNUPUdUpUrUtUvUxUzU|UUUUUUUU@VVVVVVVVVV W"W0W2W>W@WBW庭h OJQJ^JhKhK0J'B*phhKhK0J&B*phhKhKOJQJ^JhKhK0J) hQ0J&hd80J&OJQJ^JhKhK0J&H* hd80J&hKhK0J&hKhK0J'hKhK0J(3SVBWDWFWHWlWnWWDXvXXX,YlYY% & F9D[$\$gdv & Fgdv & Fgdv`gd:gd:`gdl%9D[$\$`gd3 %9D[$\$gd[% & F 9D[$\$gd[% & F 9D[$\$gdKBWFWHWVWjWlWnWWWvXXXXXXXXYY Y"Y*Y,YZ`ZbZdZZZZZ[[[\\\Z\\\t\v\\\\$]&](]l]ķČķČČķxČČČķhhf30J'h-hf3B*phhf3 hp/hf3 hhf3CJ OJQJ^JaJ hhf30J) hw70J&hhf30J&hw7 hhf3 hlhl h:h:h:hlhc/"h.#'h[h[OJQJ^Jh[OJQJ^J/YZdZZ[^[[\Z\\\(]p]]] ^l^^^N_x_`D`t`` & F gdl & Fgdv% & F9D[$\$gdv & Fgdvl]n]p]]]]^^l^^^^^^^__L_N_x__D`p`r`t`````````aaaaa$bTbdbcHcJc\c^chcɼɼɜɼɘ}vvvv hhh*hvhh* h:h:hc/"h.#'hlhh hQ$hhf3CJ OJPJQJ^JaJ hhf30J'hhf30J&hf3 hhf3h-hf3B*ph hp/hf3hw7 hhf3CJ OJQJ^JaJ hhf30J).``aaaa$bdbb cjccdd edeee4fDfff&gxgg & F gdv & F gdv`gd:`gdl`gd,hhccccd dddddde efeefffffggggggghh hhhjjjjjkjkrktkkklPlNnbn,plpqgggggggghhjhlhjjvkxkNn20888e^egde^egdh h^h`gdhh^hgdIh^hgdc/"`gd.#'`gd.#'`gd3gdc/"alnoj duljini od 254 nm. Prikaz aparature za pripravu gela za DGGE je prikazan na Slici 4. Gel za DGGE se priprema tako da se pomijeaaju reagensi prema prora unu za razli ite gradijente kako je prikazano u Tablici 2. Ukupni volumen 100%-tne i 0%-tne otopine se izra una zbrajanjem potrebnog volumena tih otopina prema ~eljenom gradijentu (postotku) zaokru~enom na prvi cijeli broj, iji rezultati su prikazani u Tablici 3. Neposredno prije nanoaenja gela u suspenziju se doda 130L APS-a (stock 0,1g/mL) i 5,8 L TEMED-a. U suspenziju gela viaeg postotka dodaje se plava boja Dcode dye solution (100 L na 5mL - 300 L) kako bi se te dvije suspenzije razlikovale. Kao uzorci se koriste PCR produkti svakog soja, a kao standard se koriste pomijeaani sojevi jer DGGE nema komercijalne standarde. Nanoaenje uzoraka zapo inje kod postignutih 58 C, a sama eleketroforezu zapo inje kod postignutih 60 C. Nakon elektroforeze provodi se bojanje gela u etidijevom bromidu i skeniranje. DGGE elektroforeza se provodi pri 220 V kroz 3 sata, a zavraetkom elektroforeze gel se tretira na isti na in kao kod DNA elektroforeze. Tablica 2. Prikaz prora una reagenasa potrebnih za dobivanje 100%-tnih i 0%-tnih otopina volumena 34 mL za dobivanje gela za DGGE PRIPREMA GELA ZA DGGE (volumena 34 mL)REAGENSPo etna koncentracijaKona na koncentracija100 %0 %Urea7 M7,14 g/Formamid40 %6,8 mL/TAE50x1x340 L34 LAkriamid/bisakrilamid40 %8 %3,4 mL3,4 mLVodaNadopuniti do ukupnog volumenaUKUPNI VOLUMEN17 mL17 mL Tablica 3. Prikaz potrebnih volumena 100%-tne i 0%-tne otopine za dobivanje ~eljenog gradijenta gela 1 (30 % i 70 %) i gela 2 (40 % i 60 %) Volumen100 %Volumen 0 %%Ukupni volumenBroj gela3,413,620175,111,93017Gel 16,810,24017Gel 28,58,5501710,26,86017Gel 211,95,17017Gel 1  Slika 4. Prikaz aparature za pripravu gela za DGGE 3.2.3. Ekstrakcija proteina vezanih na S-sloj (eng. Surface-layer associated proteins, SLAP) pomou LiCl Ekstrakcija SLAP proteina modificirana je prema standardnoj LiCl S-sloj ekstrakcijskom protokolu za L. acidophilus (Goh i sur., 2009; Lortal i sur., 1992). Prekonone kulture stanica se centrifugiraju 10 minuta pri 2,236 x g (rcf) i 4 C. Talog stanica se ispire 2 puta s hladnim fosfatnim puferom pH=7,4. Dodaje se 10 mL 5 M litijevog klorida (LiCl) s ciljem odvajanja S-proteina i SLAP proteina od stanica, i 30 min se dr~e na 4 C uz povremeno protresanje. Centrifugira se 10 minuta pri 8,994 x g (rcf) i 4 C. Supernatanti svakog soja se prebace u Spectra/Por poroznu membranu (Slika 5) i dijaliziraju se u hladnoj destiliranoj vodi na magnetskoj mjeaalici 24 h, mijenjajui prvih 8 h vodu svaka 2 h. Sadr~aj svake membrane nakon dijalize se centrifugira na sobnoj temperaturi 30 min pri 20,000 x g (rcf). Talog se resuspendira u 1 mL 1 M LiCl pri 4 C i 15 minuta kako bi se razdvojili SLAP proteini od S-proteina koji nisu topljivi u 1 M LiCl. Suspenzija se centrifugira 10 minuta pri 20,000 x g. Supernatanti se prebace u Spectra/Por membranu i provodi se ponovna dijaliza. Sadr~aj membrane nakon druge dijalize se centrifugira 30 minuta pri 20,000 x g (rcf). Talog se resuspendira u 10 L reducirajueg reagensa i prokuha 10 min pri 80 C. Na gel za SDS-PAGE se pomou Hamiltonove igle stavlja 5 L svakog uzorka, 5 L standarda i 3 L reducirajueg reagensa. Na gel se nanose i uzorci nakon prve dijalize kako bi se provjerila u inkovitost provedene metode. Dio sadr~aja membrane nakon prve dijalize se centrifugira pri 20 000 o/min 30 min. Pomijeaa se 15 L supernatanta s 5 L reducirajueg reagensa, a talog se resuspendira s 10 L reducirajueg reagensa. Uzorci se prokuhaju 10 min na 80 C i nanose na gel, takoer Hamiltonovom iglom.  Slika 5. Prikaz pripremljenih ispitivanih uzoraka prije dijalize koji se nalaze u Spectra/Por poroznoj membrani pri vraeni na krajevima vezicom 3.2.4. Turbidimetrijsko odreivanje brzine rasta kultura odabranih izolata u razli itim uvjetima Prati se rast prekononih kultura BMK u mikrotitarskim plo icama pri razli itim temperaturama (30 C, 37 C i 45 C), sni~enom pH (pH=3) koji je podeaen 1M HCl-om, uz dodatak odreene koli ine soli (4,5 % i 6,5 % NaCl), te uz dodatak ~u nih soli (0,5 %, 1 % i 1,5 %). 15 L prekononih kultura odabranih izolata se nanose na 4 mikrotitarske plo ice pri emu se na dvije prati rast u svim pripremljenim medijima u 3 paralele te se one inkubiraju pri 37 C, na treoj se prati rast BMK u istoj MRS podlozi pri 30 C, a na etvrtoj se prati rast BMK u istoj MRS podlozi pri 45 C. Slijepu probu ini isti MRS medij. Nakon nanoaenja uzoraka, odreuje se po etna apsorbancija na ita u mikrotitarskih plo ica LKB 5060-006,  GDV pri 620 nm i plo ice se inkubiraju pri odreenim temperaturama. Mjerenje apsorbancije se provodi svaka 2 sata prvih 6 sati te na kraju procesa, odnosno 22. i 24. sat. Nakon 24 sata izra unavaju se srednje vrijednosti i standardna devijacija te se odreuje rast svakog ispitivanog soja ovisno o datim uvjetima. 3.2.5. Analiza proteoliti ke aktivnosti Prekonone kulture stanica se resuspendiraju u 5 mL otopine obranog mlijeka (2%). Kao kontrola se koristi 5 mL same otopine obranog mlijeka. Provodi se inkubacija preko noi pri 37 C. Prebaci se 1 mL bistre suspenzije i centrifugira se 5 min pri 10 000 o/min. Nakon toga supernatant se pomijeaa sa reducirajuim reagensom (omjer 15:5) koji sadr~i SDS za denaturaciju proteina. Uzorci se zagrijavaju 10 min pri 80 C. Pomou Hamiltonove igle se na gel za SDS-PAGE (eng. Sodium Dodecyl Sulfate Polyacrylamide Gel Electrophoresis) nanose uzorci. Kao standard se dodaju proteini niske molekulske mase zajedno sa reducirajuim reagensom. SDS-PAGE (4% gel za sabijanje i 12% gel za razdvajanje) se provodi pri konstantnom naponu od 200 V kroz 45 min u kadici za elektroforezu. Nakon zavraene elektroforeze, gel se boji u 0,1%-tnom metilenskom modrilu R-250 s 50% metanola i 7% octene kiseline kroz najmanje 3 sata. Nakon bojanja, gel se odbojava u 7%-tnoj octenoj kiselini nakon ega se gelovi skeniraju (Scanjet 3800; Hewlett Packard, CA, SAD). 3.2.6. Adhezija na mucin Adhezija probioti kih sojeva navedenih u Tablici 1. na mucin u in vitro uvjetima provedena je na mikrititarskoj plo ici sa 96 ja~ica (Slika 6). Mikrotitarsku plo ica je prethodno pripremljena na na in da se ja~ice tretiraju otopinom mucina koncentracije 10 mg/mL u karbonat/bikarbonatnom puferu (50 mmol/L, pH 9,6) te inkubiraju preko noi pri 4 C. Ja~ice se ispiru tri puta sa fosfatnim puferom i nakon toga se dodaje po 100 L fosfatnog pufera (PBS) sa 1% Tween 20 i inkubira se 1 sat pri sobnoj temperaturi kako bi se mucin imobilizirao. Iz ja~ica se pa~ljivo izvadi PBS sa Tweenom i nakon toga u svaku ja~icu se dodaje po 100 L prethodno pripremljene suspenzije sojeva u tri paralele. Izrasle kulture sojeva navedenih u Tablici 1. se nalaze u eksponencijalnoj fazi rasta, a apsorbancija im je podeaena tako da se dobiju uzorci koncentracije A620 =0,5 za svaki od ispitivanih sojeva. Stanice se resuspendiraju u 4 mL fizioloake otopine, odnosno u otopini 5 M gvanidin hidroklorida (M=95,53 g/mol) za skidanje S-proteina. Osim u ja~ice sa mucinom, suspenzije sojeva se takoer dodaju u ja~ice bez mucina ato slu~i kao kontrola adhezije. U slijepu probu umjesto stanica se dodaje fosfatni pufer. Tako pripremljene plo ice se inkubiraju preko noi pri 37 C. Neadhezirane stanice se uklanjaju uzastopnim ispiranjem sa 0,05 % Tween 20 u fosfatnom puferu. Adhezirane stanice se detektiraju dodatkom kristal violeta u svaku ja~icu, ispiranjem tri puta sa fosfatnim puferom i dodavanjem citratnog pufera (50 mmol/L, pH 4,0). Mjeri se apsorbancija na ita u mikrotitarskih plo ica LKB 5060-006,  GDV pri 620 nm, izra unava se srednja vrijednost i standardna devijacija.  Slika 6. Prikaz mikrotitarske plo ice sa 96 ja~ica u kojoj je provedeno ispitivanje adhezije probioti kih sojeva na mucin 4. REZULTATI I RASPRAVA 4.1. USPOREDBA ISPITIVANIH Lactobacillus SOJEVA PRIMJENOM GEL ELEKTROFOREZE U DENATURIRAJUEM GRADIJENTU (eng. Denaturing Gradient Gel Electrophoresis, DGGE) Usporeeno je 6 bakterijskih sojeva koriatenih u ovom eksperimentu, koja pripadaju rodu Lactobacillus. Primjenom gel elektroforeze u denaturirajuem gradijentu (eng. Denaturing Gradient Gel Electrophoresis, DGGE) mogue je razlikovati bakterijske vrste unutar istog bakterijskog roda. Stoga je najprije provedena izolacija DNA ispitivanih bakterijskih sojeva. Mjerenjem koncentracije DNA sa BioSpec Nano ureajem dobiveni su rezultati prikazani u Tablici 4. Zatim je proveden PCR za DGGE s HDA1 i HDA2 po etnicama u svrhu umna~anja fragmenata koje omeuju ove dvije po etnice i vizualizacija DNA vrpci na gelu. Kako bi se ispitala uspjeanost provedene PCR reakcije provedena je DNA elektroforeza sa dobivenim produktima PCR reakcije. Kao negativna kontrola, da fragment dobiven na gelu nije dimer po etnica, koriaten je uzorak koji ne sadr~i DNA. Gel je obojan otopinom etidijevog bromida i skeniran, a rezultati su prikazani na Slici 7. Nakon toga je provedena DGGE elektroforeza sa dva gela koja imaju razli ite gradijente (30 do 70 %; 40 do 60 %) pri emu je kao standard koriatena DNA svih ispitivanih sojeva BMK. Gel se takoer boja u etidijevom bromidu i skenira. Tablica 4. Prikaz rezultata mjerenja koncentracije DNA BioSpec Nano ureajem UzorakKoncentracija (ng/ L)Lactobacillus helveticus M9275,53Lactobacillus brevis ZG1215,29Lactobacillus plantarum D13556,53  Slika 7. Prikaz gela nakon provedene DNA elektroforeze sa produktima PCR reakcije primjenom HDA1 i HDA2 po etnica; S - standard; M92 - Lactobacillus helveticus; ZG1 - Lactobacillus brevis; D13 - Lactobacillus plantarum; K - negativna kontrola  Slika 8. Usporedba PCR produkata dobivenih primjenom HDA1 i HDA2 po etnica nakon provedene DGGE; S - standard; M92 - Lactobacillus helveticus; D6 - Lactobacillus brevis; ZG1 - Lactobacillus brevis; SF15B - Lactobacillus brevis; SF9B - Lactobacillus paraplantrarum; D13 - Lactobacillus plantarum Rezultati gel elektroforeze su potvrdili da je PCR reakcija uspjeano provedena jer su vidljive vrpce kod svih analiziranih sojeva. Negativnom kontrolom je utvreno da ne dolazi do dimerizacije po etnica HDA1 i HDA2 jer u zadnju ja~icu gdje nije stavljen uzorak DNA nema vidljive vrpce DNA (Slika 7). Provoenjem DGGE pri emu su umno~eni dijelovi DNA analiziranih sojeva denaturirani i razdvojeni prema molekulskoj masi, provedena je elektroforeza 2 gela koji sadr~e razli ite gradijente denaturirajueg agensa, a prikazani su rezultati na 30% do 70%-tnom gelu zbog bolje uspjeanosti provedene metode (Slika 8). DGGE metodom se mogu razlikovati vrste i iz toga razloga su radi mogunosti usporeivanja rezultata uzeti dodatni sojevi za analizu (D6 - L. brevis; SF15B - L. brevis; SF9B - L. paraplantrarum). Iz Slike 8 se vidi da na istom mjestu imaju vrpce sve L. brevis vrste (sojevi D6, SF15B, ZG1), dok na drugom mjestu imaju sve L. (para)plantarum vrste (D13 i SF9B) koji su jako srodne. Soj L. helveticus M92 ima tu regiju sli niju L. brevis vrstama. Standard ine DNA uzorci svih sojeva i zato su vidljive dvije DNA vrpce. 4.2. Ekstrakcija proteina vezanih na S-sloj (eng. Surface-Layer Associated Proteins, SLAP) S-proteini imaju molekulsku masu u rasponu od 40-200 kDa (Sra i Sleytr, 2000), no usporedbom sa S-proteinima ostalih Gram-pozitivnih bakterija, oni na povraini bakterija iz roda Lactobacillus su jedinstveni, zbog toga ato ne posjeduju SLH (eng. S-layer homologous) domenu (Boot i Pouwels, 1996) najmanji su od svih poznatih S-proteina (25-71 kDa) i imaju visoku pI vrijednost od 9.35 do 10.4 (vall-Jskellinen i Palva, 2005). Protokol za ekstrakciju SLAP-a je modificiran prema standardnom ekstrakcijskom protokolu za S-proteine za soj Lactobacillus acidophilus (Goh i sur., 2009; Lortal i sur., 1992). U cilju ekstrahiranja S-proteina i proteina koji su na njih vezani (SLAP) koriaten je 5 M LiCl koji je manje letalan za stanice od gvanidin hidroklorida (Ashida i sur., 2011; Beganovi i sur., 2011.; Goh i sur., 2009; Lortal i sur., 1992; Smit i sur., 2001; Taverniti i sur., 2013). Stanice su tretirane 5 M LiCl kako bi se ekstrahirali svi S-proteini i proteini koji su na njih vezani (SLAP), nakon ega se provodi dijaliza supernatanta u kojem se nalaze S-proteini i SLAP proteini. Nakon prve dijalize provedena je SDS-PAGE supernatanta (Slika 9) i taloga (Slika 10) kako bi se provjerila u inkovitost metode. Nakon toga preostali dio je tretiran s 1 M LiCl kako bi se odvojili S-proteini koji nisu topljivi u 1 M LiCl od SLAP proteina koju su na njih vezani, a topljivi su u 1 M LiCl. Nakon druge dijalize je ponovno provodena SDS-PAGE (Slika 11) u cilju provjere prisutnosti SLAP proteina.  Slika 9. Ispitivanje prisutnosti S-proteina i SLAP proteina nakon prve dijalize provoenjem SDS-PAGE elektroforeze supernatanta uzorka proteina nakon tretmana s 5 M LiCl; S -standard; M92 - L. helveticus; ZG1 - L. brevis; D13 - L. plantarum  Slika 10. Ispitivanje prisutnosti S-proteina i SLAP proteina nakon prve dijalize provoenjem SDS-PAGE elektroforeze taloga uzorka proteina nakon tretmana s 5 M LiCl; S - standard; M92 - L. helveticus; ZG1 - L. brevis; D13 - L. plantarum  Slika 11. Ispitivanje prisutnosti SLAP proteina nakon druge dijalize provoenjem SDS-PAGE elektroforeze supernatanta uzorka proteina nakon tretmana s 1 M LiCl; S - standard; M92 - L. helveticus; ZG1 - L. brevis; D13 - L. plantarum. Nakon provedene prve dijalize, talog i supernatant se nanose na gel za SDS-PAGE u cilju provjere uspjeanosti provoenja metode. Rezultati pokazuju da na gelu na koji je naneaen supernatant (Slika 9) nema proteinskih vrpci ato je dokaz da u supernatantu nema proteina. Na gelu na koji je naneaen reseuspendirani talog proteina (Slika 10) se vide proteinske vrpce kod soja L. helveticus M92 i L. brevis ZG1, pogotovo soja L. helveticus M92 koji je slu~io kao pozitivna kontrola jer sadr~i gene ijom ekspresijom nastaju S-proteini. Ovi rezultati upuuju na to da je metoda uspjeano provedena, odnosno da je uspjeano provedeno talo~enje S-proteina zajedno sa SLAP proteinima. Takoer na Slici 10, u usporedbi sa standardom, se vide proteinske vrpce sojeva M92 i ZG1 u rasponu od 45-97 kDa ato daje pozitivnu indikaciju da se ovdje radi o S-proteinima jer je njihova molekulska masa od 25-71 kDa. Na Slici 11 koja prikazuje gel nakon druge dijalize se vidi prisutnost jedne vrpce koja odgovara molekulskoj masi S-proteina ato upuuje na prisutnost SLAP proteina kod soja L. helveticus M92 i kod soja L. brevis ZG1. Za soj L. plantarum D13 nema vidljive proteinske vrpce ato takoer potvruje da taj soj nema S-proteine, a niti SLAP proteine. 4.3. Turbidimetrijsko odreivanje brzine rasta ispitivanih Lactobacillus sojeva u razli itim stresnim uvjetima Gastrointestinalni sustav je okolia pun stresnih uvjeta za bakterije, uklju ujui nizak pH, prisutnost proteoliti kih enzima, probavnih sokova itd., pri emu S-proteini imaju protektivnu ulogu kada su mikroorganizmi koji ih sadr~avaju izlo~eni ovakvim gastrointestinalnim uvjetima (Frece i sur., 2005). Ispitan je rast soja L. brevis ZG1 koji eksprimira S-proteine, L. helveticus M92 kao pozitivne kontrole za S-proteine i L. plantarum D13 kao negativne kontrole, pri razli itim temperaturama, sni~enom pH, uz dodatak odreene koli ine NaCl i uz dodatak odreene koli ine ~u nih soli. U obzir su uzeti upravo ovi uvjeti iz razloga ato glavnu prepreku prilikom oralne primjene probiotika ine niska pH vrijednost u ~elucu i velika koncentracija ~u nih soli u tankom crijevu koje inhibiraju rast mnogih Gram-pozitivnih bakterija. Takoer bakterije mlije ne kiseline se nalaze u mnogim fermentiranim proizvodima (sirevi, kiselo zelje) gdje je poviaena koncentracija soli, odnosno NaCl, a s obzirom da sojevi koriateni u ovom radu su autohtoni sojevi izolirani iz sira, ispitana je sposobnost njihovog rasta i u ovim uvjetima. Sojevi su nacijepljeni na MRS podlogu i uzgajani pri 30 C, 45 C i na sve pripremljene MRS podloge sa ~u nim solima, NaCl i sni~enom pH vrijednosti na pH=3 u tri paralele pri 37 C. Kao slijepa proba koriste se sojevi nacijepljeni na MRS medij bez dodataka. Nakon nanoaenja uzoraka mjeri se po etna apsorbancija, a mjerenja se provode svaka 2 sata prvih 6 sati jer su tada stanice u eksponencijalnoj fazi rasta. Zadnja mjerenja se provode 22. i 24. sata jer se tada stanice nalaze u stacionarnoj fazi rasta. Na kraju eksperimenta izra unaju se srednje vrijednosti i standardna devijacija i na osnovu dobivenih rezultata nacrtaju krivulje rasta svakog bakterijskog soja (Slike 12, 13, 14).    Slika 12. Rast probioti kih sojeva a) L. helveticus M92 b) L. brevis ZG1 i c) L. plantarum D13 u standardnoj MRS tekuoj podlozi pri 37 C, u MRS tekuoj podlozi pH=3 pri 37 C, te u standardnoj MRS tekuoj podlozi pri 30 C i 45 C    Slika 13. Rast probioti kih sojeva a) L. helveticus M92 b) L. brevis ZG1 i c) L. plantarum D13 u standardnoj MRS tekuoj podlozi, te u MRS tekuoj podlozi uz dodatak 4,5 % i 6,5 % NaCl    Slika 14. Rast probioti kih sojeva a) L. helveticus M92 b) L. brevis ZG1 i c) L. plantarum D13 u standardnoj MRS tekuoj podlozi, te u MRS tekuoj podlozi uz dodatak 0,5 %, 1 % i 1,5 % ~u nih soli Usporedbom ova 3 soja, najbolju sposobnost rasta u MRS tekuoj podlozi pri 45 C je pokazao soj L. helveticus M92 , u MRS tekuoj podlozi pri 30 C su najbolje rasli sojevi L. brevis ZG1 i L. plantarum D13 dok su sva tri soja pokazala podjednaku sposobnost rasta u tekuoj podlozi pri pH=3. Soj L. brevis ZG1 je imao jednak rast u MRS podlozi sa 4,5 % NaCl kao i soj L. plantarum D13, a najbolje je rastao u MRS podlozi sa 6,5 % NaCl. Soj L. helveticus M92 je najloaije rastao u obje podloge. Soj L. helveticus M92 je najbolje rastao u MRS hranjivoj podlozi sa 0,5 %, 1%, 1,5% ~u nih soli, zatim L. plantarum D13, a najloaije je rastao L. brevis ZG1. Ovi rezultati se djelomi no sla~u sa istra~ivanjem koje su proveli Grosu-Tudor i sur. (2016) jer je pokazano da S-proteini imaju zaatitnu ulogu kod sojeva M92 i ZG1 u odreenim nepovoljnim uvjetima gastrointestinalnog trakta. Grosu-Tudor i sur. (2016) su dokazali poveanje ekspresije S-proteina soja Lactobacillus acidophilus IBB 801 kada su stanica tretirane sa NaCl (0.6 % do 3 %) i ~u nim solima (0,05% do 0,2 %) pri 42 C usporedbom sa kontrolnom skupinom koja je uzgajana u optimalnim uvjetima pri 37 C. Dokazali su dobru izdr~ljivost stanica u stresnim uvjetima iako se poveanjem stresnih uvjeta ukupan broj stanica smanjivao. Najvee koncentracije S-proteina su naene kod soja koji je rastao pri 42 C, kada je u medij za rast dodano 0,05 % ~u nih soli ili 2 % NaCl ime su dokazali da u odreenim stresnim okolianim uvjetima mo~e doi do indukcije ekspresije S-proteina, koji vrlo vjerojatno odr~ava vijabilnost stanica u nepovoljnim uvjetima. 4.4. Analiza proteoliti ke aktivnosti ODABRANIH Lactobacillus SOJEVA Proteoliti ka aktivnost bakterija mlije ne kiseline je esencijalna za njihov rast u mlijeku i zna ajno dopridonosi okusu fermentiranih mlije nih proizvoda. Veina BMK izoliranih iz fermentiranih mlije nih proizvoda su auksotrofni mikroorganizmi za odreene aminokiseline koji se uzgajaju u podlogama kompleksnog sastava iji sastav ovisi o ekspresiji kompleksnog proteoliti kog sistema za razgradnju kazeina (Siezen, 1999; Detmers i sur., 1998). Cilj je bio ispitati proteoliti ku aktivnost soja L. brevis ZG1, pri emu je kao pozitivna kontrola koriaten soj L. helveticus M92, a kao negativna kontrola L. plantarum D13. Kulture stanica su inkubirane preko noi u otopini obranog mlijeka u svrhu ispitivanja proteoliti ke aktivnosti (Slika 15), odnosno provedena je SDS-PAGE elektroforeza supernatanata kultura nakon inkubacije u cilju utvrivanja prisutnosti glavnog proteina (kazeina) koji se nalazi u obranom mlijeku (Slika 16). b)  Slika 15. Odreivanje proteoliti ke aktivnosti u obranom mlijeku usporeivanjem uzoraka a) prije i b) nakon inkubacije preko noi pri 37 C M92 - Lactobacillus helveticus; ZG1 - Lactobacillus brevis; D13 - Lactobacillus plantarum; K - otopina obranog mlijeka  Slika 16. Utvrivanje prisutnosti kazeina u supernatantu nakon inkubacije pomou odabranih sojeva bakterije mlije ne kiseline SDS-PAGE elektroforezom; S- standard proteina niske molekulske mase; M92 - Lactobacillus helveticus; ZG1 - Lactobacillus brevis; D13 - Lactobacillus plantarum; K - otopina obranog mlijeka Iz prikazanih rezultata se mo~e pretpostaviti da soj L. helveticus M92 ima proteoliti ku aktivnost jer je doalo do bistrenja otopine nakon inkubacije preko noi pri 37 C (Slika 15). Provoenjem SDS-PAGE elektroforeze na gelu nije vidljiva proteinska vrpca (Slika 16) ato potvruje pretpostavku da je uslijed djelovanja proteaza ovog soja doalo do razgradnje kazeina zbog ega se on ne mo~e detektirati na gelu. Ovi rezultati su u skladu sa rezultatima Beganovi i sur. (2013). Nakon prekonone inkubacije doalo je do blagog bistrenja kod soja L. brevis ZG1 ato upuuje na potencijalnu proteoliti ku aktivnost ovog soja (Slika 15). SDS-PAGE elektroforezom su dobiveni isti rezultati kao kod soja L. helveticus M92 s obzirom da na gelu nije vidljiva proteinska vrpca (Slika 16). Kako bi se utvrdila potencijalna proteoliti ka aktivnost ovih sojeva potrebna su daljnja istra~ivanja jer BMK proizvode mlije nu kiselinu koja sni~ava pH vrijednost u mlijeku pri emu mo~e doi do talo~enja kazeina i iz tog razloga on nije detektiran u gornjem bistrom dijelu suspenzije. Soj L. plantarum D13 nema proteoliti ku aktivnost jer nije doalo do bistrenja, ato se vidi i usporedbom sa kontrolom (Slika 15). Na SDS-PAGE gelu kod soja L.plantarum D13 se vidi proteinska vrpca (Slika 16) ato je dokaz da ovaj soj ne posjeduje proteoliti ku aktivnost koja razgrauje kazein iz obranog mlijeka pri emu ga mo~emo detektirati na gelu. 4.5. Adhezija na mucin odabranih Lactobacillus sojeva Adhezija na sluznicu crijeva je jedan od glavnih svojstava odabira probiotika jer je potrebna za crijevnu kolonizaciju i takoer je va~na za modulaciju imunoloakog sustava i anatgonizam protiv patogena (Van Tassell i Miller, 2011; Bermudez-Brito i sur., 2012). Agregacija i adhezija su slo~eni procesi koji uklju uju nekoliko molekula prisutnih na povraini stani ne stijenke kao ato su: teihoninske, lipoteihoninske kiseline i (gliko)proteini (Li i sur., 2015). S-proteini sudjeluju u auto- ili koagregaciji bakterija, vezanju na razli ite povraine, uklju ujui gastrointestinani mucin, makromolekule izvanstani nog matriksa i epitelne stanice (Sakakibara i sur., 2007; Hynnen i Palva 2013; Shimotahira i sur., 2013; Zhang i sur., 2013), formiranju biofilma i interakciji s imunoloakim sustavom. Adhezija na epitelno i subepitelno tkivo je va~an po etni korak u uspjeanoj kolonizaciji crijeva sisavaca i drugih tkiva. Uloga S-proteina u u inkovitoj adheziji patogenih i nepatogenih bakterija ovisi i o supstratu i o soju (Kos i sur., 2003). Zbog va~ne uloge soja Lactobacillus u kompeticiji s patogenima, intenzivno se prou avala njihova sposobnost adhezije za proteine izvanstani nog matriksa kao sto su kolagen, laminin, fibronektin, ili fibrinogen (Muoz-Provencio i sur., 2009). S-proteini se nalaze u nekoliko vrsta bakterija roda Lactobacillus. S-proteini su naeni u: L. brevis, L. buchneri, L. helveticus i L. hilgardii i kod bivae Lactobacillus acidophilus grupe (Johnson i sur., 1980). S-proteini se mogu odvojiti s povraine Gram-pozitivnih bakterija tretmanom sa visokim koncentracijama gvanidin hidroklorida ili litijevog klorida, koji cijepaju vodikove veze izmeu S-proteina i sekundarne stani ne stijenke polisaharida (Sra i Sleytr, 2000). Iz tog razloga se krenulo u istra~ivanje prisutnosti S-proteina i sposobnost adhezije soja Lactobacillus brevis ZG1, pri emu se kao pozitivna kontrola koristio soj Lactobacillus helveticus M92 koji sadr~i S-proteine, a kao negativna kontrola soj Lactobacillus plantarum D13 koji ih ne sadr~i.  Slika 17. Ispitivanje in vitro adhezije probioti kih sojeva na mucin prije i nakon uklanjanja povrainskih proteina s 5 M gvanidin hidrokloridom; M92 - Lactobacillus helveticus; ZG1 - Lactobacillus brevis; D13 - Lactobacillus plantarum U mikrotitarskim plo icama u kojima je apsorbancija uzoraka za svaki od ispitivanih sojeva podeaena na A620=0,5 nakon provedenog ekperimenta, mjerenja apsorbancije, izra unavanja srednjih vrijednosti i standardne devijacije dobiveni su rezultati prikazani na Slici 17. Soj L. helveticus M92, koji je slu~io kao pozitivna kontrola, jer sadr~i gene ijom ekspresijom, nastaju S-proteini, tretiran sa fizioloakom otopinom je imao veu apsorbanciju u odnosu na stanice tretirane s gvanidin hidrokloridom (Slika 17) ato je u skladu s pretpostavkom da S-proteini uklonjeni GHCl-om slu~e za adheziju na mucin. Uklanjanjem S-proteina je izmjerena manja apsorbancija jer stanice u odsustvu tog proteina adheziraju manjim intezitetom za mucin. Soj L. plantarum D13 koji je slu~io kao negativna kontrola, jer ne sadr~i gene za ekspresiju S-proteina, tretiran sa fizioloakom otopinom je pokazao neznatno veu apsorbanciju (Slika 17) nego nakon tretmana sa gvanidin hidrokloridom, ato je u skladu s pretpostavkom da kod soja L. plantarum D13 budui da nema S-proteine, ne dolazi do zna ajnije promjene u vezanju za mucin. U njegovom vezanju na mucin najvjerojatnije posreduju teihoninske, lipoteihoninske kiseline ili neki drugi proteini odnosno glikoproteini na povraini stanica. Soj ija smo svojstva ispitivali je L.brevis ZG1 i on je pokazao sli ne rezultate kao i soj L. helveticus M92, odnosno doalo je do zna ajnog smanjenja apsorbancije nakon tretmana sa gvanidin hidrokloridom u usporedbi sa tretmanom sa fizioloakom otopinom (Slika 17). Ovi rezultati ukazuju na to da S-proteini soja L. brevis ZG1 sudjeluju u adheziji na mucina jer je njihovim uklanjanjem sposobnost adhezije za mucin znatno smanjena. Ovi rezultati su u skladu s pretpostavkom da vrste Lactobacillus helveticus i Lactobacillus brevis koje imaju S-proteine posjeduju bolja adhezijska svojstva od Lactobacillus vrsta kao ato je Lactobacillus plantarum koja nema S-proteine (Johnson i sur., 1980). 5. ZAKLJU CI DGGE analizom je mogue razlikovati vrste unutar roda Lactobacillus: Lactobacillus brevis ZG1, Lactobacillus plantarum D13, te Lactobacillus helveticus M92. Koriatenjem razli itih koncentracija litijevog klorida se mogu odvojiti S-proteini od proteina koji su na njih vezani (tzv. SLAP proteini), ato je dokazano za ispitivani soj L. brevis ZG1, kao i za pozitivnu kontrolu L. helveticus M92. Turbidimetrijskim odreivanjem brzine rasta kultura u razli itim stresnim uvjetima dokazano je da ispitivani soj L. brevis ZG1 ima dobru sposobnost rasta u kiselom pH (pH=3) i u MRS podlozi sa NaCl (4,5 % i 6,5%), dok je u MRS hranjivoj podlozi sa ~u nim solima (0,5 %, 1%, 1,5%) najloaije rastao od svih ispitivanih sojeva. Soj L. helveticus M92 je imao dobru sposobnost rasta pri poviaenoj temperaturi, u kiselim uvjetima i u hranjivoj podlozi sa ~u nim solima, ali ne i u podlozi sa NaCl-om. Soj L. plantarum D13 je imao dobru sposobnost rasta u kiselim uvjetima kao i u podlozi sa 4,5 % NaCl. Analizom proteoliti ke aktivnosti dokazano je da soj L. brevis ZG1 ima proteoliti ku aktivnost kao i kontrolni soj sa slojem S-proteina L. helveticus M92, dok kod soja L. plantarum D13 koji slu~i kao negativna kontrola za ekspresiju S-proteina, proteoliti ka aktivnost nije dokazana. 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