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OPSTA Mikrobiologijanovaa4pdf 1 UNIVERZITET CRNE GORE Biotehnički fakultet OPŠTA MIKROBIOLOGIJA Dr Mirjana Bojanić Rašović Podgorica, 2016. g. Sadržaj strana 1. Predmet, značaj i istorijski razvoj mikrobiologije...............5 2. Morfologija mikroorganizama………………….................21 3. Ekološki faktori koji djeluju na mikroorganizme...............75 4. Ishrana i metabolizam mikroorganizama.............................90 5. Rast i razmnožavanje mikroorganizama...........................114 6. Kretanje mikroorganizama.................................................125 7. Genetika mikroorganizama................................................126 8. Sistematika mikroorganizama............................................133 9. Staništa mikroorganizama u prirodi………………...........146 10. Mikroorganizmi sa posebnim svojstvima...........................148 11. Patogenost mikroorganizama.............................................153 12. Mikroorganizmi – trovači hrane.........................................163 13. Osnove zaraznih bolesti.....................................................171 14. Bakterije – uzročnici zaraznih bolesti životinja..................180 15. Gljive - uzročnici zaraznih bolesti životinja .......................201 16. Virusi - uzročnici zaraznih bolesti životinja........................206 17. Prion proteini – uzročnici zaraznih bolesti životinja...........214 18. Paraziti - uzročnici bolesti životinja....................................216 19. Mikrobiologija stočne hrane...............................................228 20. Uloga mikroorganizama u varenju hrane kod preživara.....235 21. Mikrobiologija mlijeka i proizvoda od mlijeka...................241 22. Mikrobiologija mesa i proizvoda od mesa ..........................258 23. Mikrobiologija ribe.............................................................248 24. Mikrobiologija jaja...............................................................264 Bibliografija..........................................................................266 2 Predgovor Ovaj udžbenik je rađen prema nastavnom planu i programu predmeta Mikrobiologija koga slušaju studenti Biotehničkog fakulteta Univerziteta Crne Gore u Podgorici, na studijskom programu Stočarstvo. Imajući u vidu da udžbenik obrađuje i teme iz opšte mikrobiologije, vjerujem da kao dodatna literatura može koristiti i studentima drugih srodnih fakulteta u upoznavanju sa mikroorganizmima. S obzirom na značaj koji imaju inženjeri stočarstva u preventivi, odnosno sprečavanju bolesti životinja i ljudi, proizvodnji stočne hrane, primarnoj proizvodnji i preradi namirnica animalnog porijekla, pripremanju obroka za ishranu životinja, obradi tečnog i čvrstog đubriva (stajnjaka), proizvodnji mikrobiološkog đubriva u cilju poboljšanja plodnosti zemljišta, kontroli proizvodnje i čuvanja stočne hrane, kao i hrane za ljude, poznavanje mikrobiologije im je neophodno u razumijevanju svih ovih oblasti. Mikroorganizmi se nalaze u svim životnim staništima, a zbog svoje izuzetno velike brzine razmnožavanja, značajno utiču na procese sinteze i razlaganja kako organskih, tako i neorganskih jedinjenja u prirodi. Zahvaljujući svojoj raznovrsnoj metaboličkoj aktivnosti, oni utiču na opstanak i odvijanje životnih funkcija svih drugih živih bića na zemlji - biljaka, životinja i ljudi. Neke vrste mikroorganizama mogu djelovati negativno, odnosno štetno, tako što izazivaju bolesti životinja, biljaka i ljudi ili mogu dovesti do kvarenja, odnosno truljenja stočne hrane i životnih namirnica. Druge vrste mikroorganizama su korisne i njihova aktivnost je neophodna u dobijanju konzervirane stočne hrane (silaža, sjenaža), konzervirane hrane za ljude (konzervacija voća, povrća, proizvoda od mlijeka, mesa i proizvoda od mesa), pri čemu utiču na dobijanje namirnica specifičnih svojstava - ukusa, konzistencije i mirisa. Svako klimatsko podneblje ima i svoju specifičnu mikrofloru koju karakterišu jedinstvene metaboličke osobine. Zahvaljujući tome, u različitim klimatskim područjima se proizvode i različiti prehrambeni proizvodi (jogurt, sir, kajmak, kobasice i dr.). Imajući u vidu gore navedeno, poznavanje mikroorganizama će omogućiti budućim inženjerima poljoprivrede da se uspješno bore protiv štetnih, truležnih i patogenih mikroorganizama, a da koriste potencijal korisnih mikroorganizama u svim oblastima poljoprivrede i prehrambene industrije. Kroz ukupno 24 tematske cjeline ovog udžbenika pokušala sam da studentima približim najvažnije činjenice iz istorijskog razvoja mikrobiologije, osnovna morfološka svojstva mikroorganizama, ekološke faktore koji djeluju na mikroorganizme, uticaj mikroorganizama na okolinu, način ishrane i metabolizam mikroorganizama, rast i razmnožavanje mikroorganizama, kao i način svrstavanja mikroorganizama u sistematske kategorije. Dalje, pokušala sam da približim sljedeće teme: pojam patogenosti mikroorganizama, imunitet životinja, osnovne osobine mikroorganizama i osnovne osobine parazita koji izazivaju bolesti životinja. Takođe je obrađen i značaj korisnih i štetnih mikroorganizama u procesu proizvodnje stočne hrane, kao i posebna uloga mikroorganizama u varenju hrane preživara. U poglavljima koja se odnose na mikrobiologiju mlijeka, mesa i jaja ukazano je na korisne i štetne aspekte djelovanja mikroorganizama koji se mogu naći u ovim namirnicama. Posebnu zahvalnost dugujem recenzentima, dr Veri Katić i dr Aleksandri Martinović, na temeljnoj i sveobuhvatnoj analizi teksta i veoma korisnim sugestijama i savjetima, koje sam sa velikim zadovoljstvom unijela u ovaj tekst. 3 Takođe, zahvaljujem Milošu Otaševiću i Marijani Maksimović na svesrdno pruženoj pomoći u obradi i prilagođavanju slika njhovoj namjeni. Nadam se da će mi uvaženi studenti i kolege, koji budu pročitali ovaj udžbenik dati svoje sugestije, što će doprinijeti da sljedeće izdanje bude još bolje i korisnije. S poštovanjem, Autor Podgorica, 2016. g. 4 PREDMET, ZNAČAJ I ISTORIJSKI RAZVOJ MIKROBIOLOGIJE Mikrobiologija je nauka o malim, golim okom nevidljivim živim organizmima, koji se nazivaju mikroorganizmi ili mikrobi. Dobila je ime od grčkih riječi micros - mali, bios - život i logos - nauka. Ona izučava: morfologiju, sistematiku, fiziologiju, ekologiju mikroorganizama, kao i njihovu ulogu u prirodi i životu čovjeka. Morfologija izučava izgled, građu i veličinu ćelije mikroorganizama. Sistematika se bavi klasifikacijom (svrstavanjem) mikroorganizama na osnovu njihovih srodničkih osobina. Fiziologija proučava metabolizam i životne funkcije mikroorganizama (disanje, rast, razmnožavanje, razmjenu materija sa okolinom itd.). Ekologija izučava uticaj faktora spoljašnje sredine na mikroorganizme, kao i načine njihovog prilagođavanja na te faktore u određenoj biocenozi. Biocenoza predstavlja skup svih živih organizama koji žive u jednom određenom prostoru - staništu, tj. biotopu. Mikrobiologija je nauka širokih mogućnosti. Sve dok mikroorganizmi nisu bili otkriveni, mnoge pojave koje su se dešavale u prirodi nisu bile objašnjene. Danas su mikroorganizmi našli široku primjenu u industriji, poljoprivredi, farmaciji i medicini. Mogu se otkriti samo uz pomoć optičkog ili elektronskog mikroskopa (slika 1). Maksimalno uveličanje optičkog mikroskopa iznosi 3000 puta. To omogućava razlikovanje čestica koje nisu manje od 0,1 μm. Slika 1. Optički - svjetlosni mikroskop http://bestcompoundmicroscopes.com/advantages-of- using-a-compound-light-microscope/ Savremeni elektronski mikroskopi imaju sposobnost razdvajanja do 0,15 nm, pa se pomoću njih mogu vidjeti ne samo sitni organizmi, već i finije strukture ćelija (slika 2). Uveličanje elektronskog mikroskopa iznosi 100.000 puta. 5 Slika 2. Transmisioni elektronski mikroskop (TEM) http://www.ru.nl/science/gi/facilities/electron-microscopy/transmission-ems/ Mikroorganizmi su svrstani u pet carstava: Eubakterije - prave bakterije (Eubacteria, Bacteria), Arheje (Archaea), Protozoe (Protozoa), Alge (Algae) i Gljive – plijesni i kvasci (fungi) (slika 3). Eubakterije i arheje pripadaju prokariotima (imaju prokariotsku građu ćelije), a protozoe, alge i gljive eukariotima (imaju eukariotsku građu ćelije). Virusi su acelularni mikroorganizmi i čine posebnu grupu mikroorganizama. Virusni sateliti, viroidi i prioni su subvirusne čestice. Slika 3. Šema podjele mikroorganizama prema građi ćelije (http://www.cen.ulaval.ca) Mikroorganizmi su široko rasprostranjeni u prirodi. Nalaze se u zemljištu, vodenim bazenima, na površini i unutar tijela čovjeka, životinja i biljaka, u hranljivim proizvodima, vazduhu itd. Široka rasprostranjenost mikroorganizama ukazuje na njihovu veoma značajnu ulogu u prirodi. Zahvaljujući njihovoj aktivnosti dolazi do: razlaganja različitih organskih materija u zemljištima i vodenim bazenima, kruženja materije i energije u prirodi, stvaranja plodnog zemljišta, kamenog uglja, nafte i mnogih drugih rudnih bogatstava. Zahvaljujući sposobnosti da proizvode mnogobrojne enzime, mikroorganizmi iskorišćavaju različite organske i neorganske materije. Usvajanje rastvorenih hranljivih materija vrše cijelom površinom svog organizma. Procese metabolizma mogu da vrše u aerobnim i 6 anaerobnim uslovima, u kisjelim, baznim i neutralnim sredinama, na temperaturama ispod nule i temperaturama do 1000C. Razmnožavaju se geometrijskom progresijom, što im omogućava brzo širenje i život u svim ekološkim sredinama. Zahvaljujući razvoju nauke i tehnike, znanja o mikroorganizmima su dostigla visok nivo, što je omogućilo njihovu primjenu u raznim oblastima. U industriji se mikroorganizmi koriste u proizvodnji antibiotika, vitamina, alkohola, vina, organskih kisjelina, proteina,
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