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2.1.8 Phn loi to lam (Vi khun lam) Ngnh c khong 150 chi vi 2000 loi (Fott, 1971).

Vit Nam pht hin c 344 loi v di loi (Dng c Tin, 2002). Tn ti mt s h thng phn loi to lam. Theo h thng ca Geitler (1932), to lam c mt lp (Cyanophyceae) v c chia lm 4 b: Chroococcales, Dermocarpales, Pleurocapsales v Hormogonales. H thng ca Fritsch (1945) chia to lam thnh 5 b ( tt c u thuc 1 lp Cyanophyceae): Chroococcales, Chamaesiphonales, Pleurocapsales, Nostocales v Stigonematales, cch phn chia ny cng c Desikachary (1967) tn thnh. Khc vi cc tc gi trn, Gollerbakh (1977) chia to lam thnh 3 lp: (1)Chroococcophyceae, (2)Chamaesiphonophyceae v (3) Hormogoniophyceae. Theo h thng ny, to lam gm c 10 b, trong ng ch l lp th (3) ng tch b Nostocales thnh 2 b: Nostocales v Oscillatoriales - iu m cc h thng trc cha lm. Van Den Hoek v cs (1995), khng ch da vo cc du hiu hnh thi, v cu to ca t bo v cch thc sinh sn ( cc tiu ch m cc h thng truc ng s dng ), m cn s dng cc c im cu trc siu hin vi ca t bo, c bit ch trng n ch tiu phn t ( AND) phn loi v xy dng h thng, theo to lam (VKL) c mt lp Cyanophyceae vi 5 b, l: Chroococcales, Pleurocapsales, Oscillatoriales, Nostocales v Stigonematales. Cch phn chia ny cng ph hp vi quan im ca mt s tc gi khc nh Anagnostidis K. & Komarek (1985, 1988, 1990). Da trn vic so snh trt t cc nucleotid trong 16s rARN ca 30 dng to lam thuc 5 b nu trn, kt qu cho thy cc b Pleurocapsales, Nostocales v Stigonematales dng nh l cc b "t nhin", mi mt b l mt nhm gm nhiu dng c mi quan h ln nhau, cn cc b Chroococcales v Oscillatoriales th khng ng nht (heterogeneous) v bao gm cc dng khng c quan h huyt thng. B Chroococcales B ny bao gm cc loi n bo hoc tp on (c cu trc palmelloid); cng c gp mt s loi m trong cc t bo hnh thnh m (cm) nh cht nhy (bn cht l ng cao phn t polysaccharid). Sinh sn bng cch chia i t bo (bao gm c vic hnh thnh cc nanocyt- cc ni bo t v ny chingoi bo t). Hnh 24. Gloeoeocapsa (du mi tn v mi tn ch bao nhy) (Theo Linda & Lee, 2000) Chi Gloeoeopsa (Hnh 24) L tp on c cu trc palmelloid, mi tp on cha mt s t bo c dng hnh cu n bn cu, c mu, phn tng v c bao bc trong bao nhy. Bao nhy bao bc khng ch cho tng t bo n l m n cn bao quanh c tp on.

S phn chia t bo c thc hin trn 3 mt ct. Chi c khong 40 loi, gp trn cc tng , trn tng nh, trong t v sui. Chi Microcytis Chi ny thng chim u th trong thc vt ni ca cc ao h nhit i. Loi M. aeruginosa l loi ton cu, s phong ph ca n trong s n hoa ca nc thng gy hu qu xu cho kinh t v sc kho (Collins, 1978; Reynolds et all, 1981). Kch thc, hnh dng v s sp xp ca cc t bo trong tp on bin thin rt ln. S lng t bo trong mt tp on c th ch vi t bo, c khi n hng ngn. Hnh 25. Microcystis (x 600, nh ca tc gi) T bo c dng hnh cu, hnh trng hoc qu nang, kch thc cu t bo thng t 3-10 m, Cc t bo c lin kt vi nhau bng cht nhy to nn tp on (25% th tch ca tp on l cht nhy) (Hnh 25). T bo ca Microcystis cha cc khng bo kh nn c mu en, i lc chng chm xung y ao h (vo ma thu do s tch lu hydratcarbon lm tng t trng ca to, vo ma xun khi nhit ca nc tng, to li ni ln). Khi cn non tp on c dng hnh cu, khi trng thnh v gi bin dng mnh. Theo Reynold et all (1981), hm lng O2 ti u cc qun th non Microcystis sinh trng khi nc n hoa l t 1- 4mg/lit, ngoi ngng trn th n c tc ng ngc li (vi qun th). S sinh trng ny cng c kch thch bi s tng ln ca nhit nc. Microcystis aeruginosa v M. flos-aqae thng gy hin tng n hoa nc cc h c hm lng dinh dng thch hp. Khi thiu photpho th cc khng bo kh Microcystis v Aphanizomenon u khng c hnh thnh. Cc khng bo kh ca Microcystis thng b xp mnh bi p sut trng. Chnh yu t ny gip ta gii thch s sinh trng dy c ca chng trn b mt nc. Chi Merismopedia (Hnh 26) Gp nc ngt, nc mn, k c cc bi ct ven bin. N l thnh phn c trng ca qun x m ly. T bo hnh trng hoc hnh cu, lin kt cht ch vi nhau thnh cc dy trong mt mt phng. Cc tp on lin kt vi nhau bng o nhy, cc o nhy ny khng cch xa nhau ( cc t bo ring l khng thy c o nhy r rng).

Hnh 26. Merismopedia; Hnh 27. Chamaesiphon (u mi tn ch ngoi bo t) ( Theo Linda & Lee, 2000) Chi Chamaesiphon (Hnh 27) Sng thnh m trn b mt thc vt thu sinh, trn to khc v trn cc tng , trn v gip xc trong cc h nc ngt v sui. T bo hnh tr, c o m rng pha nh, t cc bo t trn c to ra (exospore). B Oscillatoriales Tn dng si, sinh sn bng on to (hormogonium= hormogone). Si ch c mt kiu phn nhnh - l phn nhnh gi v khng c t bo d hnh cng nh khng c bo t. Chi Oscillatoria (Hnh 28) Trong chi ny, cc trichom c dng hnh tr trn v t do, chng khng bao gi to thnh tp on. Khi nhn di knh hin vi quang hc, hu ht cc loi ca n dng nh khng c o (bao), nhng trn thc t chng c bao rt mng, tn khng phn nhnh. Trichom gm cc t bo n gin, khng phn ho.T bo c chiu rng (theo mt ct ngang) ln hn chiu di, khng c eo tht gia cc t bo cnh nhau, trichom khng phn bit "chn" v "u", nhng t bo tn cng mt s loi c th l hnh cu, hnh nn hoc gin ra to thnh m. Hnh 28. Mt s loi thuc chi Oscillatoria 1. Oscillatoria sancta ; 2. O. limosa ; 3. O. princeps ; 4. O. brevis ; 5. O. formosa ( Theo Gollerbakh, 1977) Tn cng ca trichom i khi nghing v mt bn. Osccillatoria khng tm thy t bo d hnh hoc bo t. Sinh sn bng cch to ra cc hormogonium, l kt s hnh thnh cc a phn ct, lm 2 mt ti mt s im dc trichom ( Hnh 29). Chi c trn 100 loi, phn b rng trong nc ngt, trong bin, sui nc nng v nhng ni b nh hng ca nc
Hnh 29. Osccillatoria - a phn ct 2 mt (Theo Desikachang, 1959)

qu theo

thi.
lm

Chi Spirulina (Hnh 30) To c tm thy trong nc ngt v nc mn, c trong cc ao h c nng Na2CO3 cao, trong cc h thuc vng m ly, trong bn nho. Trichom a bo, hnh tr trn

hoc nhiu un cong dng l xo, vch ngn ngang m (b che khut), t bo tn cng ca trichom un trn, khng c chp. Loi Spirulina platensis l thc n truyn thng ca c dn quanh h Chad (Nam Phi). H ny c nng Na2CO3 cao (kim mnh). Hm lng protein ca to cao (4570% trng lng kh) nn c nui trng trn qui m ln nhiu nc trn th gii lm ngun thc n b sung cho ngi, gia sc v gia cm.

Hnh 30. Spirulina Hnh 31. Lyngbya (Theo Linda & Lee, 2000) (Theo Gollerbakh, 1997) Chi Lyngbya (Hnh 31) Trichom ca Lyngbya ging vi trichom ca Oscillatoria, nhng khc ch n c bc trong mt o nhy dy v chc. Chi c trn 100 loi, sng nc l, nhng cng gp c nc ngt v bin. Chi Microcoleus ( Hnh 32) Trichom ging vi Oscillatoria, nhng khc ch mt s hoc nhiu trichom to thnh tp on , chng c bc trong o nhy chung, cng v chc. Hnh 32. Microcoleus Theo Linda & Lee, 2000) B Nostocales To lam thuc b ny c dng si v sinh sn bng s hnh thnh hormogonium. Si c phn nhnh gi, nhng khng c phn nhnh tht. C hai loi t bo d hnh v bo t ngh (akinete) u c to ra trn si. Chi Nostoc (Hnh 33) Nostoc sng nc ngt v t, chng gp nhiu trong t canh tc . Mt s loi sng trong t m cng vi ru, l thnh phn ca a y hoc ni cng sinh vi thc vt bc cao. Si ca Nostoc khng phn nhnh, n un cong v tht ngng su

cc vch ngn ngang trng ging nh mt chui ht.Trn si c hnh thnh t bo d hnh v thnh thong c bo t ngh. Hnh 33. Nostoc sp. (Theo Kumar, 1999) Cc trichom cnh nhau bn vo nhau to thnh mt tp on c bao nhy chung, bao nhy c hnh dng xc nh. Tp on c th trn ging nh chm nho. Sinh sn bng hormogonium v bo t ngh. Chi Anabaena (Hnh 34) V hnh dng, ging vi Nostoc (khng phn nhnh) nhng khc ch cc si khng thng xuyn nhm li vi nhau to thnh tp on c bao nhy chung nh Nostoc. Cc trichom ca chi Anabaena hu ht u thng v chuyn ng, trong khi Nostoc th ch c hormogonium chuyn ng m thi. T bo c dng hnh cu, trng hoc tr trn v c cc ht cyanophycin. Cc loi sng tri ni c khng bo kh. Hnh 34. Anabaena sp (Theo Kumar, 1999) T bo d hnh c hnh thnh phn gia hoc cui si - l nhng ni to tin hnh c nh nit kh quyn. Sinh sn bng cch t on ngu nhin v bng hormogonium. Mt s si to ra mt chui cc bo t ngh (akinetes) k st t bo d hnh hoc gia hai t bo d hnh. Chi c trn 100 loi, phn ln sng ao h nc ngt v trong rung la. Chi Scytonema (Hnh 35) Khc vi cc chi Nostoc v Anabaena, Scytonema c phn nhnh gi, c th c mt hoc hai nhnh gi t mi si to. Hnh 35. Scytonema(Theo Linda & Lee, 2000) Trong vng gia ca mi nhnh gi c hnh thnh t bo d hnh. T bo ca trichom t nhiu c dng hnh vung. Si c bao nhy dy v chc. Sinh sn bng hormogonium, n c thot ra u mt ca si hoc ca nhnh gi. To c mt nc ngt v nc mn, trong t, trn v cy v cc c cht khc. Chi Rivularia (Hnh 36) Si c dng bp mng (to phn gc, thon dn v vut nhn phn cui), u cng gc ca mi si c mt t bo d hnh, k st l akinete. Mi si u c bao nhy ring, chng hp li to tp on c hnh bn cu, trong cc trichom sp xp theo li

phng x. Si thng phn nhnh gi gc. Sinh sn bng hormogonium v bo t ngh. To phn b rng, h, sui nc ngt v ven bin. Hnh 36. Rivularia (Theo Gollerbakh, 1977) B Stigonematales B bao gm nhng to lam c phn nhnh tht, c t bo d hnh v vch ngn ngang gia cc t bo to cc h lm. Hu ht cc loi trong b c sinh trng nh, sng trong t, st mt t hoc trong nc ngt. Chi Stigonema (Hnh 37) Hu ht cc loi ca chi ny sng trn mt t v trn cc tng c nc nh git. Mt s loi (v d, Stigonema hormoides) si gm mt dy t bo, cn hu ht c mt s dy (Hnh 37a,b).

(a). Stigonema hormoides (b). Stigonema sp. ( Theo Kumar, 1999 ) ( Theo Kumar, 1999 ) Si phn nhnh tht, c bao nhy dy, cc t bo d hnh thng xut hin c si b v si nhnh. Sinh sn bng hormogonium, mt s loi hnh thnh bo t ngh (akinete). Hormogonium ch c to ra cc si nhnh, cn cc bo t ngh c to ra cc si b. Chi Hapalosiphon (Hnh 38) Si b ca chi Hapalosiphon hu nh ch c mt dy t bo (cng c khi hai dy), cn chi Westiella ch c mt dy m thi. chi Fischerella si b li c nhiu dy t bo (Hnh 39).

Hnh 37.

Hnh 38.

Hapalosiphon

Hnh 39. Fischerella


(Theo Gollerbakh, 1953)

( Theo Kumar, 1999 )

2.2 Ngnh Prochlorophyta (= Prochlorobacteria)

Ngnh ny c Lewin (1976,1977) xng, cc i din ca ngnh sng cng sinh vi hi tiu bin. C th n bo, c cu to ging vi Cyanobacteria nhng khc ch khng c phycobiliproteins v phycobilisomes. T bo cha dip lc a v b v beta-carotene v mt ny ging vi to lc v lc lp ca thc vt bc cao. Chi Prochloron (Hnh 40) Prochloron l chi in hnh, n l mt trong nhng c th n bo c bit n nhiu nht v c th l t tin tin nhn ca lc lp to lc v thc vt bc cao. N c dng hnh cu, ng knh ti 30 micron. T bo khng c nhn, ty th, th golgi v li ni cht. s xp hai hoc vi ci to thnh thnh lamell (ging to lc). ADN khng tp trung vng trung tm nh Cyanophyta m nm gia cc s. Sn phm d tr - polysaccharide tinh bt ging vi cyanophycin. Ngnh ch c 1 lp - Protochlorophyceae vi 3 chi : 2 chi Prochloron v Prochlorococcus sng bin, cn chi Prochlorothrix sng nc ngt.

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Ghi ch: Mng sinh cht ADN Vch t bo Thylacoid Carboxysome Hnh 40. A. Prochloron ; B. Dng si sng t do ca Prochlorophyta ( Theo Kumar, 1999 )

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