Dai Cuong Ve Nam. Basidiomycotina

Gio trnh mn Nm hc
By: Nguyn Vn Thnh Cao Ngc ip Nguyn Vn B
Gio trnh mn Nm hc
By: Nguyn Vn Thnh Cao Ngc ip Nguyn Vn B
Online: < http://cnx.org/content/col10923/1.1/ >
CONNEXIONS
Rice University, Houston, Texas
This selection and arrangement of content as a collection is copyrighted by Nguyn Vn Thnh, Cao Ngc ip, Nguyn Vn B. It is licensed under the Creative Commons Attribution 3.0 license (http://creativecommons.org/licenses/by/3.0/). Collection structure revised: August 14, 2009 PDF generated: February 5, 2011 For copyright and attribution information for the modules contained in this collection, see p. 94.
Table of Contents
1 i cng v nm mc . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 2 Nm roi - nm trng (ngnh ph Chytridiomycotina) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 3 Ngnh ph Nm tip hp (Zygomycotina = lp Zygomycetes) . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . 17 4 Ngnh ph Nm Bt Ton (Deuteromycotina = lp Deuteromycetes) . . . . . . . . . . . . . . . . . . . . . 31 5 Ngnh ph nm m (Basidiomycotina = Lp Basidiomycetes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45 6 Ngnh ph Nm Bt Ton (Deuteromycotina = lp Deuteromycetes) . . . . . . . . . . . . . . . . . . . . . 69 7 Vai tr hu dng ca nm trong ch bin thc phm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83 Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93 Attributions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
iv
Chng 1
i cng v nm mc
1.1 Hnh dng, kch thc, cu to ca nm mc

1.1.1 Hnh dng v kch thc
Mt s t nm th n bo c hnh trng (yeast=nm men), a s c hnh si (filamentous fungi=nm si), si c ngn vch (a bo) hay khng c ngn vch (n bo). Si nm thng l mt ng hnh tr di c kch thc ln nh khc nhau ty loi. ng knh ca si nm thng t 3-5m, c khi n 10m, thm ch n 1mm. Chiu di ca si nm c th ti vi chc centimet. Cc si nm pht trin chiu di theo kiu tng trng ngn (Hnh 1.1). Cc si nm c th phn nhnh v cc nhnh c th li phn nhnh lin tip to thnh h si nm (mycelium) kh sinh x x nh bng. Trn mi trng c v trn mt s c cht trong t nhin, bo t nm, t bo nm hoc mt on si nm c th pht trin thnh mt h si nm c hnh dng nht nh gi l khun lc nm (Hnh 1.2)
1 This
content is available online at <http://cnx.org/content/m30247/1.1/>.
CHNG 1. I CNG V NM MC
Figure 1.1
Hnh 1.1 Si nm v cu to vch t bo si nm (theo Samson v ctv., 1995)
Figure 1.2
1.2
Hnh 1.2. Mt s dng khun lc nm (theo Samson v ctv., 1995)
1.2.1 Cu to
T bo nm c cu trc tng t nh nhng t bo vi sinh vt chn hch khc c m t v trnh by nh Hnh 1.3
Figure 1.3
Hnh 1.3 Cu to t bo nh si nm Fusarium (theo Howard R J & Heist J R., 1979) (Ch thch: MT: vi ng, M: ty th, SC: b Golgi, V: bng(ti) nh, P: mng sinh cht 4 lp) Vch t bo nm cu to bi vi si chitin v c hoc khng c celluloz. Chitin l thnh phn chnh ca vch t bo hu ht cc loi nm tr nhm Oomycetina. Nhng vi si chitin c hnh thnh nh vo enzim chitin syntaz (Hnh 1.4). ***SORRY, THIS MEDIA TYPE IS NOT SUPPORTED.*** Hnh 1.4. Con ng tng hp chitin T bo cht ca t bo nm cha mng ni mc (endoplasmic reticulum), khng bo (vacuoles), ty th (mitochondria) v ht d tr (glycogen v lipid), c bit cu trc ty th t bo nm tng t nh cu trc ty th t bo thc vt. Ngoi ra, t bo nm cn c rib th (ribosomes) v nhng th khc cha r chc nng. T bo nm khng c dip lc t, mt vi loi nm c ri rc trong t bo mt loi sc t c trng m Matsueda v ctv. (1978) u tin ly trch c v gi l neocercosporin (C29H26O10) c mu tm nm Cercosporina kikuchi. T bo nm khng nht thit c mt nhn m thng c nhiu nhn. Nhn ca t bo nm c hnh cu hay bu dc vi mng i phospholipid v protein dy 0,02 [U+F06D]m, bn trong mng nhn cha ARN v ADN.
1.3 Dinh dng v tng trng ca nm mc

Hu ht cc loi nm mc khng cn nh sng trong qu trnh sinh trng. Tuy nhin, c mt s loi li cn nh sng trong qu trnh to bo t (Buller, 1950). Nhit ti thiu cn cho s pht trin l t 2oC n 5oC, ti ho t 22oC n 27oC v nhit ti a m chng c th chu ng c l 35oC n 40oC, c bit c mt s t loi c th sng st OoC v 60oC. Ni chung, nm mc c th pht trin tt mi trng acit (pH=6) nhng pH ti ho l 5 - 6,5, mt s loi pht trin tt pH < 3 v mt s t pht trin pH > 9 (Ingold, 1967). Oxi cng cn cho s pht trin ca nm mc v chng l nhm hiu kh bt buc v s pht trin s ngng khi khng c oxi v d nhin nc l yu t cn thit cho s pht trin.
5 Theo Alexopoulos v Minns (1979) cho bit nm mc c th pht trin lin tc trong 400 nm hay hn nu cc iu kin mi trng u thch hp cho s pht trin ca chng. Nm mc khng c dip lc t nn chng cn c cung cp dinh dng t bn ngoi (nhm d dng), mt s sng st v pht trin nh kh nng k sinh (sng k sinh trong c th ng vt hay thc vt) hay hoi sinh (saprophytes) trn xc b hu c, cng c nhm nm r hay a y sng cng sinh vi nhm thc vt nht nh. Theo Alexopoulos v Mims (1979) cho bit ngun dng cht cn thit cho nm c xp theo th t sau: C, O, H, N P, K, Mg, S, B, Mn, Cu, Zn, Fe, Mo v Ca. Cc nguyn t ny hin din trong cc ngun thc n v c n gin nh glucoz, mui ammonium... s c nm hp thu d dng, nu t ngun thc n hu c phc tp nm s sn sinh v tit ra bn ngoi cc loi enzim thch hp ct cc i phn t ny thnh nhng phn t nh d hp thu vo trong t bo.
1.4 Sinh sn ca nm mc
Ni chung, nm mc sinh sn di 2 hnh thc: v tnh v hu tnh. Trong sinh sn v tnh, nm hnh thnh bo t m khng qua vic gim phn, tri li trong sinh sn hu tnh nm hnh thnh 2 loi giao t c v ci.
1.4.1 Sinh sn v tnh

The Alexopoulos v Mims (1979), nm mc sinh sn v tnh th hin qua 2 dng: sinh sn dinh dng bng on si nm pht trin di ra hoc phn nhnh v sinh sn bng cc loi bo t. Mt s loi nm c nhng bo t c trng nh sau: a. Bo t ti (bo t bc)(sporangiospores): cc bo t ng (zoospores) (Hnh 1.5 a, b, c) c nm Saprolegnia v bo t ti (sporangiopores) nm Mucor, Rhizopus (Hnh 1.6) cha trong ti bo t ng (zoosporangium) v ti bo t (sporangium) c mang bi cung ti bo t (sporangiophores).
Figure 1.4
Hnh 1.5 Bo t ng (theo Samson v ctv., 1995) ***SORRY, THIS MEDIA TYPE IS NOT SUPPORTED.*** Hnh 1.6. Bo t ti (b) Mucor circinelloides, a. cung bo t ti (theo Samson v ctv., 1995) b. Bo t nh (conidium): cc bo t nh khng c ti bao bc ging nm Aspergillus, Penicillium, ... Hnh dng, kch thc, mu sc, trang tr v cch sp xp ca bo t nh thay i t ging ny sang ging khc v c dng lm tiu chun phn loi nm. Cung bo t nh dng bnh c th khng phn nhnh nh Aspergillus (Hnh 1.7) hay dng th phn nhnh nh Penicillium (Hnh 1.8). Bo t nh hnh thnh t nhng cm (cluster) trn nhng cung bo t nh Trichoderma (Hnh 1.9).
Figure 1.5
Hnh 3.3. Cc kiu cung bo t nh ca Aspergillus. a. 1 lp, b. 2 lp, c. phin, d. tia, e. t (theo Samson v ctv., 1995)
Figure 1.6
Hnh 1.8. Bo t nh v cung bo t nh Penicillium chrysogenum (theo Samson v ctv. 1995) ***SORRY, THIS MEDIA TYPE IS NOT SUPPORTED.*** Hnh 1.9. Cung bo t phn nhnh Trichoderma. a. T. viride, b. T. koningii, c. T. polysporum, d. T. citrinoviride (theo Samson v ctv. 1995) ging Microsporum v Fusarium, c hai loi bo t nh: loi nh, ng nht gi l tiu bo t nh (microconidia) (Hnh 1.10 a) , loi ln, a dng gi l i bo t nh (macroconidia) (Hnh 1.11 b)
Figure 1.7
c. Bo t tn (Thallospores): trong nhiu loi nm men v nm mc c hnh thc sinh sn c bit gi l bo t tn. Bo t tn c th c nhng loi sau: 1. Chi hnh thnh t t bo nm men: Cryptococcus v Candida l nhng loi bo t tn n gin nht, gi l bo t chi (blastospores) 2. Ging Ustilago c nhng si nm c xut hin t bo c vch dy gi l bo t vch dy cn gi l bo t o (chlamydospores) (Hnh 1. 11 c). V tr ca bo t vch dy si nm c th khc nhau ty loi. 3. Ging Geotrichum v Oospora c si nm ko thng, vung hay ch nht v t bo vch dy gi l bo t t (arthrospores) (Hnh 1.12)
Figure 1.8
Hnh 1.12 Bo t t (theo Samson v ctv. 1995)
1.4.2 Sinh sn hu tnh

Sinh sn hu tnh xy ra khi c s kt hp gia hai giao t c v ci (gametes) c tri qua giai on gim phn. Qu trnh sinh sn hu tnh tri qua 3 giai on: Tip hp t bo cht (plasmogamy) vi s ha hp 2 t bo trn (protoplast) ca 2 giao t Tip hp nhn (karyogamy) vi s ha hp 2 nhn ca 2 t bo giao t to mt nhn nh bi (diploid) Gim phn (meiosis) giai on ny hnh thnh 4 bo t n bi (haploid) qua s gim phn t 2n NST (nh bi) thnh n NST (n bi). Theo Machlis (1966) tt c cc giai on trn k c giai on to c quan sinh dc c iu khin bi mt s kch thch t sinh dc (sexual hormones). C quan sinh dc ca nm mc c tn l ti giao t (gametangia) c 2 loi: c quan sinh dc c gi l ti c (antheridium) cha cc giao t c (antherozoids), cn c quan sinh dc ci gi ti non (oogonium) cha giao t ci hay non, khi c s kt hp gia giao t c v non s to thnh bo t, bo t di ng c gi l bo t ng (zoospores). Kiu hai si nm c gii tnh c v ci tip hp nhau sinh ra bo t c tn l tip hp t (myxospores), tip hp t l c trng ca nhm nm Myxomycetes (Hnh 1.13).
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Bo t sinh dc khi hnh thnh c dng ti gi l nang (ascus) v ti ny cha nhng bo t gi l bo t nang (ascospores). Nang v bo t nang l c trng ca nhm Ascomycetes (Hnh 1.14) . Trong nhm Basidiomycetes, 4 bo t pht trin phn tn cng ca cu trc th qu gi l m (basidium) v bo t c gi l bo t m (basidiospores) (Hnh 1.15) Nhm Nm bt ton (Deuteromycetes=Deuteromycotina)) gm nhng nm cho n nay cha bit r kiu sinh sn hu tnh ca chng. ***SORRY, THIS MEDIA TYPE IS NOT SUPPORTED.*** Hnh 3.2.1 Cc kiu hnh thnh tip hp t Mucoraceae. a-f. Rhizopus. g-h. Zygorhynchus, i. Absidia, j. Phycomyces (theo Talbot, 1995)
Figure 1.9
Hnh 1.13. Bo t nang Saccharomyces cerevisiae (theo Samson v ctv. 1995)
11
Figure 1.10
Hnh 1.14. Cc kiu bo t m. a. Astrea, b. Bovista, c. Agaricales, d. Clavulina, e. Dacrymyces, f. Sistotrema, g. Repetobasidium, h. Xenasma, i-n. bo t m c vch, n. Puccinia. (theo Kreisel, 1995)
1.5 V tr v vai tr ca nm mc
Nm mc c nh hng xu n cuc sng con ngi mt cch trc tip bng cch lm h hng, gim phm cht lng thc, thc phm trc v sau thu hoch, trongch bin, bo qun. Nm mc cn gy h hi vt dng, qun o... hay gy bnh cho ngi, ng vt khc v cy trng. Tuy nhin, cc qui trnh ch bin thc phm c lin quan n ln men u cn n s c mt ca vi sinh vt trong c nm mc. Nm mc cng gip tng hp nhng loi khng sinh (penicillin, griseofulvin), acit hu c (acit oxalic, citric, gluconic...), vitamin (nhm B, riboflavin), kch thch t (gibberellin, auxin, cytokinin), mt s enzim v cc hot cht khc dng trong cng nghip thc phm v y, dc ... c s dng rng ri trn th gii. Ngoi ra, nm cn gi vai tr quan trng trong vic phn gii cht hu c tr li mu m cho t trng. Mt s loi thuc ging Rhizopus, Mucor, Candida gy bnh trn ngi, Microsporum gy bnh trn ch, Aspergillus fumigatus gy bnh trn chim; Saprolegnia v Achlya gy bnh nm k sinh trn c. Nhng loi nm gy bnh trn cy trng nh Phytophthora, Fusarium, Cercospora.... c bit nm Aspergilus flavus v Aspergillus fumigatus pht trin trn ng cc trong iu kin thun li sinh ra c t aflatoxin. Bn cnh tc ng gy hi, mt s loi nm mc rt hu ch trong sn xut v i sng nh nm n, nm dc phm (nm linh chi, Penicillium notatum tng hp nn penicillin, Penicillium griseofulvum tng
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hp nn griseofulvin...), nm Aspergillus niger tng hp cc acit hu c nh acit citric, acit gluconic, nm Gibberella fujikuroi tng hp kch thch t gibberellin v mt s loi nm thuc nhm Phycomycetina hay Deuteromycetina c th k sinh trn cn trng gy hi qua c th dng lm thin ch dit cn trng. Ngoi ra, nhng loi nm sng cng sinh vi thc vt nh Nm r (Mycorrhizae), gip cho r cy ht c nhiu hn lng phn v c kh tan v cung cp cho nhu cu pht trin ca cy trng. Nm cn l i tng nghin cu v di truyn hc nh nm Neurospora crassa, nm Physarum polycephalum dng tng hp ADN v nhng nghin cu khc.
1.6 Phn loi nm mc

u tin, nm c sp xp theo tin ha nh m hnh di y: (Hnh 1.15) Dayal (1975) lit k 7 c tnh phn loi nm mc nh sau: 1. 2. 3. 4. 5. 6. 7. c im hnh thi k ch c th c im sinh l c im t bo hc v di truyn hc c im khng huyt thanh c tnh sinh ha chung phn loi s hc
Figure 1.11
13 Hnh 1.15 Cy di truyn pht sinh ngnh cho thy nm mc c mi lin h gn vi thc vt (PLANTAE) v ng vt (ANIMALIA) (theo Hawkswort v ctv., 1995) Theo Gwynne-Vaughan v Barnes (1937) chia nm thnh 3 lp chnh: Phycomycetes, Ascomycetes v Basidiomycetes da trn khun ty c vch ngn ngang hay khng v c im ca bo t. Theo Stevenson (1970) phn loi nm trong ngnh Mycota gm 6 lp: Chytridiomycetes, Oomycetes, Zygomycetes, Ascomycetes, Basidiomycetes, v Deuteromycetes. Gn y, Kurashi (1985) nhn mnh n tm quan trng ca h thng ubiquinon trong phn loi nm mc cng nh ng dng k thut sinh hc phn t kho st a dng di truyn v qua mi lin h di truyn phn loi li cho chnh xc hn.
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Chng 2
Nm roi - nm trng (ngnh ph Chytridiomycotina)

1
1 This
15
16
CHNG 2. NM ROI - NM TRNG (NGNH PH CHYTRIDIOMYCOTINA)
Chng 3
Ngnh ph Nm tip hp (Zygomycotina = lp Zygomycetes)

3.1 c tnh chung ca ngnh ph Nm tip hp

y l nhm nm k sinh trn ng vt, thc vt v c trn nm khc Hu ht nm cho khun ty pht trin v phn nhnh; c mu nu, xm, trng T bo nm cha y cc thnh phn nh ti th, nhn, rib th, ht lipid, mng ni mc Mng t bo ch yu l chitosan chitin. Chitosan c nhiu b Mucorales v Entomophthorales nhng khng c b Zoophagales Nm khng c trung th (centrioles) Sinh sn v tnh vi bo t trong ti hay bc (sporangiospore) cn gi l bo t bt ng (aplanospores), cha rt nhiu bc hay ti bo t (sporangia). S t loi nm sinh sn vi bo t vch dy (chlamydospore), bo t nh (conidia) Sinh sn hu tnh vi s phn chia giao t (2 giao t pht trin t khun ty khc nhau). Hai giao t hp nhau thnh bo t c vch dy gi l bo t tip hp (zygospore) nn gi l lp nm tip hp (lp Zygomycetes). Bo t tip hp chng chu s kh hn v nhng yu t bt li ca mi trng; v bo t c mu c trng nhiu loi nm nht nh.
3.2 Phn loi

Webster (1980) phn loi ngnh ph hay lp nm tip hp ch c 2 b Mucorales v Entomophthorales B Mucorales B Mucorales bao gm nhng loi ph bin trong t nhin nh t, khng kh, xc b thc vt... trong c nhiu loi cng c ch cho con ngui. Khun ty phn nhnh v c vch ngn ngang, trong t bo cht vi thnh phn nu c tnh chung ca lp ny, t bo cht c thm ti cha dch (cisternae) c nhim v ging nh b Golgi; Sinh sn hu tnh vi tip hp t (zygotes)(giao t a nhn hay nhiu nhn nh bi [diploid]). Theo Martin (1961) phn chia b ny gm c 9 h nhng Hesseltine v Ellis (1973) chia b ny thnh 14 h khc nhau trong h Mucoraceae quan trng nht. H Mucoraceae Nhng loi nm thuc h ny c nhng c tnh chung l v t bo cha chitin, chitosan; nm c ti bo t ln (sporangia) cha cung hay li (columella) v bo t tip hp hin din hu ht cc loi trong h; Hesseltine v Ellis (1973) chia h Mucoraceae thnh 20 ging trong chi Rhizopus v chi Mucor l quan trng nht.
1 This
17
18
CHNG 3. NGNH PH NM TIP HP (ZYGOMYCOTINA = LP ZYGOMYCETES)
3.2.1 Ging [Chi] Rhizopus

Ging ny c t nht 120 loi v th c m t trong Rhizopus stolonifer (R. nigricans) l loi ph bin trong thin nhin v c m t tng i k; Rhizopus stolonifer thng hin din bnh m c nn thng c gi l mc bnh m, n cn hin din trong t, trong tri cy h, c.... n cn k sinh trong r khoai ty, to, du, c chua nhiu khi chng cn gy ra bnh trn ng vt nui. Hu ht nhng loi Rhizopus l nhng loi thc vt hoi sinh (saprophytes), chng pht trin khun ty bao ph phn bn ngoi ca c cht (v d nh bnh m), khun ty ca Rhizopus stolonifer c mu trng, phn nhnh, a nhn v khng c vch ngn ngang . Hu ht cc si khun ty c dng nh si bng vi khi cn non (hnh 3.1), sau pht trin su vo c cht th phn chia thnh 3 dng khun ty Da petriBnh mKhng bonhnv khun ty
Figure 3.1
19 Hnh 3.1. Nm Rhizopus pht trin bnh m c (a), si khun ty nm vi nhiu nhn cng nh tng trng (b) (Sharma, 1998) : khun cn (rhizoids), khun ngang (stolon) v cng mang ti (bc) bo t (sporangiophores)(hnh 3.2). khun cn l khun ty n su vo c cht tng t nh r cy n su vo t nhng chng pht trin cn hn. khun ngang l khun ty nhng pht trin chiu ngang, bn trn mt c cht, chng ni tng nhm nm vi nhau. Cng mang ti bo t l khun ty mc thng ln khng, chng pht trin t trung tm im xut pht ca khun ngang v khun cn, mi cng mang ti bo t pht trin tn cng l ti bo t (sporangium), y l giai on sinh sn v tnh.
3.2.1.1 Cu trc bn trong ca khun ty Khun ty c cu trc hnh ng (hnh 3.1b) vi vch khun ty cu to bng chitin, siu cu trc ca vch khun ty cho thy chng cu to bng vi si (microfibrillar), chy song song bn b mt ni vi nhau bng mng plasma mng; Ht nguyn sinh (protoplast) l nhng ht bao gm nhn, ht d tr, ti th, rib th, mng ni mc v nhng khng bo (vacuole) v nhng ht ny tp trung nhiu nh tng trng hay u khun ty.
20
Figure 3.2
Ti bo t cng bo tcung,li khun ngang
21 khun cn Hnh 3.2. Ba loi khun ty ca nm Rhizopus l khun cn (rhizoid), khun ngang (stolon) v cng bo t (sporangium)(Sharma, 1998) 3.2.1.2 Dinh dng Khun cn tng hp v phng thch nhiu enzym trong c nhng enzym phn hy tinh bt thnh ng n; mi trng vi nhiu nit hu c v v c s gip Rhizopus tng hp nhiu protein hn. 3.2.1.3 Sinh sn v tnh (Asexual reproduction) c tnh ca ging ny l hnh thnh nhng cng mang bc bo t (sporangiophores) v ti (bc) bo t (sporangium). Bo t khng c roi, gn nh trn, ng nht, a nhn nm trong ti mu en gi l ti bo t, mt ti bo t pht trin n c v tn cng ca cng mang bc bo t (hnh 3.2) v bc bo t c mu en nn cn gi l mc en. 3.2.1.4 Sinh sn hu tnh (Sexual reproduction) Bt u giai on sinh sn hu tnh bng s tip hp (conjugation) v kt qu to nn bo t tip hp (zygospore), qu trnh sinh sn hu tnh chia ra 2 trng hp nh sau: D tn (heterothallic) trong 2 ni khc nhau t 2 si nm khc (tm gi l + v - ) kt hp vi nhau ng tn (Homothallic) trong 2 ni kt hp t mt si nm nh trng hp Rhizopus sexualis. Trong nhng loi d tn, hai khun ty khc nhau cho ra 2 bo t khc nhau + v - s kt hp li vi nhau thnh th nh bi (diploid) v pht trin thnh ti giao t non (progametangia) gi l th tip hp (zygophores)(hnh 3.3).
22
Figure 3.3
gametangia = ti giao t Hnh 3.3. Sinh sn hu tnh vi trng hp d tn trong 2 bo t + v - kt hp vi nhau t 2 khun ty nm khc nhau to nn bo t tip hp (Sharma, 1998)
23
Figure 3.4
Columella = li, promycelium = tin khun ty Hnh 3.4. Bo t ny mm cho ra cc tin khun ty v to ra cc bo t c nhn n bi (Sharma, 1998) Bo t tip hp (zygospore) mc mm bng cch ph v v bo t (hnh 3.4) pht trin thnh mt khun ty hnh ng mc thng ln khng gi l tin khun ty (promycelium); Tin khun ty bt u gim phn cho cc nhn n bi (n nhim sc th [NST]) v hnh thnh ti bo t tn ngn v tu bo t ny cha bo t c hai loi + v - . Trong trng hp ng tn (nh Rhizopus sexualis) th th tinh xut pht t mt khun ty (hnh 4.5) v to nn bo t tip hp ring bit kt hp vi nhau. S pht trin tin khun ty nm R. sexualis tng t nh nm R. stolonifer.
24
Figure 3.5
Hnh 3.5. Sinh sn hu tnh vi trng hp ng tn nm Rhizopus sexualis (Sharma, 1998)
3.2.2 Chi Mucor

Mucor l nhm nm hoi sinh trn xc b hu c c bit trong d dy ca nga v tru b (Mucor mucedo), nhiu loi pht tn trong t nh Mucor racemosus v Mucor spinosus, nm ny cng c mt trn bnh m c, tht, ph mt, nc tri cy... nhiu loi gy ra bnh mycormycosis trn ngi v gia sc; Tuy nhin nhiu loi nm cng c ch nh Mucor rouxii phn hy tinh bt thnh ng; c tnh pht trin ca Mucor ging nh Rhizopus, v d nh chng pht trin khun ty trn bnh m c trong 24 gi. 3.2.2.1 Sinh sn v tnh (Asexual reproduction) Nm Mucor sinh sn v tnh nh nm Rhizopus bng cch thnh lp cng mang bc bo t v bo t vch dy (chlamydospore). - Cng mang bc bo t vi nhng bo t bt ng hnh thnh trong ci bao hay bc bo t (sporangia); mi bc bo t pht trin tn ngn, khng phn nhnh v cng mang bc bo t pht trin ring bit, khng cng nhm (hnh 3.6) nhiu khi c nhiu loi c bit c th mang bc bo t phn nhnh nh Mucor racemosus (hnh 3.7) v Mucor plumbeus.
25
Figure 3.6
bc bo t Hnh 3.6. Cng mang bc bo t vi 1 bc bo t (Sharma, 1998) Trong t bo cht cha nhiu nhn nhng bo t ch c 1 nhn, tu bo t i sang mu nu khi bo t trng thnh v d dng v ra phng thch bo t theo gi, nhiu khi bo t dnh vo chn cn trng pht tn ti nhng ngun thc n khc v khi c iu kin thun tin, bo t ny mm cho ra mt khun ty mi. Khng ging nh nhng loi khc trong ging Mucor, Mucor rouxii c bo t ny mm nh nm men trong iu kin k kh, c bit khi c s hin din ca kh CO2; tuy nhin , khi c oxi th bo t ny mm cho ra mt khun ty bnh thung.
26
Figure 3.7
Hnh 3.7. Th mang bc bo t vi nhiu bc bo t sporangial wall = v ti bo t sporangiospores = bo t branched sporangiophore = cng mang bc bo t phn nhnh chlamydospore = bo t vch dy sporangium = bc bo t - Bo t nang ch thnh lp khi khun ty to ra nhng t bo c thnh dy nh trng hp Mucor racemosus (hnh 3.7).
27
Figure 3.8
Hnh 3.8. S sinh sn hu tnh (ng tn) Mucor Progametangium = tin giao t Gametangium = giao t Zygospore = bo t tip hp 3.2.2.2 Sinh sn hu tnh (Sexual reproduction) Trong sinh sn hu tnh, Mucor c nhng c im chung vi Rhizopus, M. genevensis v nhiu loi khc l nhng loi ng tn (tt c sinh ra t mt khun ty v thnh lp bo t tip hp)(Hnh 3.8), tuy nhin, M. mucedo v nhng loi khc li l d tn (hnh 3.9)
28
Figure 3.9
Hnh 3.9. S sinhsn hu tnh (d tn) nm Mucor (Sharma, 1998) Hai ging Rhizopus v Mucor trong h Mucoraceae c nhng im khc bit c bn sau:
29
ging Rhizopus C khun cn C khun ngang Thc n c hp thu t khun cn Cng bo t pht trin ring bit vi khun cn Bo t dnh trn cung bo t v kh phn tn
ging Mucor Khng c khun cn Khng c khun ngang Thc n c hp thu t b mt khun ty Cng bo t pht trin ring bit v khng cng tp hp thnh nhm Bo t d pht tn theo gi
Table 3.1
3.2.3 Tm quan trng ca b Mucorales

1. 2. 3. 4. 5. 6. 7. 8. 9. 10. Cc ging trong b ny gy ra mt s bnh trn khoai ty, du, to, v nhiu loi tri cy khc Ht ging lun nhim cc bo t ca cc ging trong b ny Rhizopus l tc nhn nhim mc trn bnh m Cc ging nm cn gy ra mt s bnh trn ngui v gia sc Nhiu loi trong ging Rhizopus tng hp acit lactic v acit fumaric nh Rhizopus oryzae v R. stolonifer Nhiu loi trong ging Rhizopus v Mucor dng sn xut ru Nhiu loi trong ging Actinomucor v Mucor dng sn xut Tempeh v Sufu Nhiu loi ca ging Blakeslea tng hp nhiu -carotene Nhiu loi trong b ny c kh nng k sinh trn nhiu loi nm khc Rhizopus stolonifer c dng sn xut corticoid
30
Chng 4
Ngnh ph Nm Bt Ton (Deuteromycotina = lp Deuteromycetes)

1
4.1 Gi thiu chung

Ngnh ph Deuteromycotina gm mt h thng cc nhm nm b thiu hoc khng pht hin c nhng c im ca nm hon chnh (tip hp, nang hoc m); nhng nm ny khng mang bo t tip hp (zygospore), bo t nang (ascospore), hoc bo t m (bo t nh th sinh - basidiospore). Nm ny thiu giai on sinh sn hu tnh trong vng i nn ngi ta gi chung l nm khng hon chnh hay Nm bt ton (Imperfect fungi). Cc c th ch sinh sn bng hnh thc v tnh, ch yu l bng bo t nh (conidia) pht trin trn cung bo t nh (conidiophores). Sutton (1973) ngh Deuteromycotina nh "l mt tp hp cc kiu nm sinh sn bng bo t vi dng khng hp nhn bi s gim nhim"
4.1.1 c im chung
- Deuteromycotina c m t bi trn 15.000 loi (Ainsworth, 1973) phn ln sng trn cn; Mt s ln nm bt ton thu sinh (Alatospora, Tricladium, Pyricularia) tm thy trong c mi trng bin v nc ngt, a s cc c th hoi sinh hoc k sinh, l nguyn nhn gy mt s bnh trn thc vt v ng vt. - Ngoi tr dng n bo ging nh nm men ca Blastomycetes, hu ht tt c Deuteromycotina cn li u c h khun ty (mycelium) tht, gm c s pht trin si, phn nhnh v vch ngn si nm (hypha) - H si nm thng c gian bo hoc ni bo v mi t bo cha nhiu nhn. - Vch ngn trn tt c cc loi c kho st hu nh ging vi Ascomycotina,c mt l thng gia mi vch. - Hon ton khng c sinh sn hu tnh, sinh sn ch yu bng dng bo t c bit l bo t nh (conidia); Bo t l bo t nh bt ng, pht trin bn ngoi cung bo t nh, v phn ny th Deuteromycotina ging nh Ascomycotina. Bo t nh c hnh dng, kch thc, mu sc thay i. . . n c th trong sut hoc c mu sc thay i, n nhn hoc a nhn, c vch ngn ngang, dc hoc khng; N c th c hnh trng (oval), thun di, hnh cu, dng sao, dng hi cong, dng si, hnh a, dng cun xon hay nhng dng khc. - Bo t nh c sinh trc tip t cung bo t hoc t mt vi kiu th qu nh; b si bo t (synnema) (hnh 6.1), cm cung bo t (arcevulus) (hnh 6.2), gc cm bo t nh (sporodochium) hoc ti bo t phn (pycnidium). Nhng th qu ny l cc m mm gi trong phm vi ni bo t c sinh
1 This
31
32
CHNG 4. NGNH PH NM BT TON (DEUTEROMYCOTINA = LP DEUTEROMYCETES)
ra. Sutton (1973) pht hin ch c 3 kiu th qu l ti bo t phn, cm cung bo t v lp cht m (stroma) - Gii tnh i ng (Parasexuality) (d tnh) c m t trn mt s Deuteromycotina; di hin tng ny, c s hnh thnh cc u tch hp cht nguyn sinh, tip hp nhn v n bi ho ti mt thi im c bit hoc mt v tr c bit trong vng i ca nm. Hin tng ny c cp mt cch gin tip bi Pontecorvo v Roper (1952) v nhng nghin cu b sung ca Pontecorvo (1956,1958), Davis (1966). Mt vi c s bn ngoi ca hin tng ny bao gm s thnh lp ca hai kiu nhn (heterokaryotic) (d hch), si nm; c s tip hp nhn v nhn ln ca nhn lng bi, c s xy ra phn bo gin phn (mitosis), s "tuyn chn" ca nhn lng bi v cui cng l s n bi ho mt vi nhn lng bi trong khun ty.
Figure 4.1
Hnh 6.1. Mt b si bo t (synnema) ca Arthrobotryum (Sharma, 1998)
33
Figure 4.2
Hnh 6.2. Nm Collectrichum [A], Fusarium [B], Endothia [C](Sharma, 1998)
4.1.2 Tm quan trng

Phn ln cc nm trong ngnh Deuteromycotina u c tc ng trc tip hoc gin tip n cuc sng hng ngy ca con ngi. Mt s nm thuc lp Hypomycetes nh mt s loi thuc ging Alternaria gy bnh g st khoai ty, c chua v mt s cy trng h Solanaceae; Pyricularia gy bnh o n la; Cercospora gy bnh m l bng vi, thuc l, C. apii gy l lot ngoi da ngi. Ging Fusarium gy bnh thi ma, thi qu c chua (Fusarium solani), thi kh khoai ty, h hnh ti; Cc c th thuc chi Colletotrichum gy bnh lot cy, C. lagenarium gy bnh thi hng bu b (Sharma, 1989; Lng c Phm, 2002). c t nm Fusarium, Alternaria gy xut huyt ni quan (d dy, rut, gan,. . .), ri lon thn kinh.
34
4.2 Phn loi

Hnh dng, kch thc, vch ngn, mu sc v s trang tr ca bo t l tiu chun chnh phn loi Deuteromycotina; Song song vi vic da vo hnh thi ca bo t th s pht trin ca chng (tn v kiu pht trin phi nguyn bo, Kendrick, 1971), hnh dng v s pht sinh ca cung bo t nh cng nh s t hp ca chng trong dng th qu xc nh (b si bo t (synnema), cm cung bo t (arcevulus), gc cm bo t nh (sporodochium) hoc ti bo t phn (pycnidium) cng l nhng c im phn loi quan trng. Ainswoth (1973) chia ngnh ph Deuteromycotina theo 3 lp: Khun ty khng pht trin hoc pht trin yu; dng c th ging nh nm men v c s ny chi: Blastomycetes Khun ty tht; khng ny chi; si nm bt dc hoc sinh bo t trn cung, khng c s tp trung thnh ti bo t hay cm cung bo t: Hypomycetes Khun ty tht; bo t tp trung trong ti bo t hoc trn cm cung bo t: Coelomycetes Alexopoulos v Mims (1979) a ra 3 lp ph hnh thc l Blastomycetidae, Hypomycetidae v Coelomycetidae.
4.2.1 Lp Hypomycetes
4.2.1.1 c tnh chung Phn ln cc c th sng hoi sinh trn thc vt hoc k sinh. Si nm pht trin mnh, c vch ngn v phn nhnh. Ch yu sinh sn bng bo t (Moniliales) nhng mt s ch sinh sn bng phn on (fragmentation) nh Rhizoctonia v Sclerotium. Bo t ca chng kh hoc nhy nht. C ti bo t ln cm cung bo t u khng c trong s sinh sn ca bt k c th no.
4.2.1.2 Phn loi Alexopoulos v Mims (1979) cng nhn 2 b hnh thc (Moniliales v Agromycetales) di phn lp hnh thc Hyphomycetidae, cc c th ca Hyphomycetes sinh ra bo t c t trong b hnh thc Moniliales nhng nhng dng thiu bo t v sinh sn bng phn on si nm th c t vo b hnh thc Agromycetales. 4.2.1.3 c im ca b Moniliales Phn ln c th hoi sinh hoc k sinh v bo t ca chng pht trin trn nhng si nhnh chuyn bit l cung bo t (sporophore) hoc cung bo t nh (connidiospore), chng c ngh thnh 4 h hnh thc (form-class) sau: 1. Moniliaceae: cung bo t tch ra t mt si no hoc khng c; bo t v h si nm trong sut hoc c mu sng, i din Monilia. 2. Dematiaceae: bo t v h si nm mu sm. i din Altenaria, Curvularia, Cercospora, Helminthosporium, Drechslera. 3. Tuberculariaceae: bo t v cung bo t nh c sinh ra t cm cung bo t. i din Fusarium. 4. Stilbellaceae: bo t v cung bo t nh pht trin trong b cung bo t nh. i din Graphium.
35 4.2.1.4 a. Ging ALTERNARIA * c im Nhm nm ny kh ph bin, c nhiu loi hoi sinh v gy cht tng phn cy trng. Alternaria l tc nhn gy nhim chnh trong nui cy trong phng th nghim. Bo t ca chng rt ph bin trong bi bm trong nh, trong khng kh v l tc nhn chnh gy d ng (Hyde v Williams, 1946), mt s bnh v da v vi ri lon nghim trng c th ngi.Nhiu loi Alternaria k sinh trn thc vt. Trn cc c th thuc h Solanaceae (khoai ty), Alternaria cho triu chng bnh r st sm hn l Phytophthora infestans (thuc lp Oomycetes, tc nhn gy bnh r st mun (late-blight) khoai ty), ch ring Alternaria c gi l bnh r st sm. Triu chng sm ca bnh l nhng m nh mu vng nu trn l, sau lan rng to nhng vt hnh nhn ng tm; Ton b phin l, cung l, gn l v thm ch c h thng mch dn cng tn thng t gy do b nhim. Phn cn li ca ng mch c mu nu.
Figure 4.3
Hnh 6.3. Alternaria solani trn khoai ty (Solanium tuberosum)(Sharma, 1998) Alternaria alternata (=A.tenuis) l nguyn nhn gy bnh m en trn la m trong khi A.triticina gy bnh r st (thi kh l) (Bhownik, 1969). A. brassicae v A.brassicicola tn cng trn ht Brascica (h ci bp) cn A.solani (hnh 6.3) gy bnh r st sm trn khoai ty v cc loi khc thuc h Solanaceae; Mt vi loi Alternaria khc (vi k ch ca chng trong ngoc n) l A. citri (trn l h cam qut Citrus sp.), A..helianthi (trn hng dng Helianthus annuus) v A. palandui v A. porri gy chy l trn hnh ty, ti. H si nm Mu nu sng, mnh, phn nhnh mnh, si nm c vch ngn trc ht l gian bo, sau c th tr thnh ni bo.;Mi t bo thng c nhiu nhn. Theo Knox-Davis (1979) th nhng t bo sinh dng ca A.brasicicola cha t mt ti nhiu nhn, u mt t bo si nm c 27 nhn v nhng t bo gi c n 33 nhn. Sinh sn Ging Alternaria ch yu sinh sn bng cch to bo t nh; Giai on hon chnh ca Alternaria l Pleospora infectoria (hnh 6.4) mt loi nm Loculoascomycetous (Webster, 1980).
36
Figure 4.4
Hnh 6.4. Mt nha bo t phng (ascus) ca Pleospora infectoria mt loi nm Loculoascomycetous giai on hon chnh ca Alternaria (Sharma, 1998) Bo t nh pht trin trn cung bo t nh ngn, sm mu v thng v nh hnh; Mt bo t pht trin nh l chi ngn ca t bo nh trn cung bo t nh (hnh 6.5. A-D); N khng pht trin bng cch tht eo v m rng phn cht t bo ca cung bo t. Nhng bo t non c phn cch bng vch ngn ngang (hnh 6.5. E) vi s pht trin ca vnh hnh khuyn vo bn trong (Campbell,1970); Trung tm mi vch ngn c mt l thng ni cht gia cc t bo ca bo t, sau mt s t bo phn cch bi vch ngn dc (hnh 6.5. F). Nhm bo t vi vch ngn v dc nh th c gi l dng qu du (muriform) hoc bo t li (dictyospore), thng th phn chp bo t ny chi v cui cng to thnh si ca bo t (hnh 6.5. G). i khi chi c th pht trin t t bo thp hn hay gn vo bo t to nn nhnh ca si bo t (hnh 6.5. H-I). S m rng thm ca si bo t s ngng khi c s bt kn l nn bo t; Bo t chn l mt qu th nhiu nhn c vch ngang v dc; N c bao quanh bi 2 lp vch, tng ngoi c sc t ca t bo, tng trong trong sut (Campbel, 1969, 1970). Theo Knok-Davies (1979) cung bo t chn cha vi nhn (0-3) trong khi bo t cha 1-2 nhn, Purkayastha v cng s (1980) nghin cu siu cu trc b mt ca 5 loi Alternaria gy bnh (A.longissima, A.cassiae, A.tenuissima, A.raphani v A.sonchi); Cc ht bo t c pht tn nh gi, gp iu kin nhit v m thch hp bo t ny mm to t 5 n 10 ng (hnh thnh si nm)(hnh 6.5. J).
37
Figure 4.5
Conidiophore = cng mang t1ui bo t, bud = chi, conidia = bo t nh, germ tube = ng mm, lateral bud = chi hng, young conidium = bo t nh non Hnh 6.5. A-F s pht trin bo t nh ca Alternaria solani; G, 2 bo t nh thnh chui ca A.brascicae; H, bo t ny chi ca A.brasicicola; I, chui bo t phn nhnh ca A.brascicae; J, bo t ny chi ca A.brasicicola (Sharma, 1998) Kim sot bnh thi li S lun phin ma v l c li v bnh ch yu t t trng; Thuc phun tr nm tt nht l loi c cha ng hoc km, cch khong 15 ngy trong phm vi kim sot d phng. Azariah v cng s (1962) ch trng s dng hn hp Bordeux trong khi Mathur v cng s (1971) th gii thiu phun Zineb v Dithane M-45. 4.2.1.5 b. Ging CERCOSPORA c im Ging nh Alternaria, Cercospora cng l mt chi ln trong h Dematiaceae, c i din bi trn 2000 loi (Ellis, 1971) nhng s ln xut hin rt nhiu v hu nh ng dng (Webster,1980). Cercospora l nguyn nhn gy bnh m l trn c chua, rau dip, khoai ty, bng vi, la, u phng, t, u trng ct
38
(piegon pea - arhar), c ci ng, thuc l. . . v nhiu cy trng kinh t quan trng khc; C. personata l tc nhn gy bnh m gch nu u phng (Arachis hypogea), C.gossypina gy bnh m l trn bng vi (Gossypium herbaceum) v C. oryzae gy bnh gch nu trn la, C. apii gy bnh trn ngi v c th l nguyn nhn gy nhng vt l lot trm trng trn mt trng rt kinh khng. (Emmons v ctv, 1975). H si nm pht trin mnh, phn nhnh v c vch ngn mng, si nm ni bo, gic mt phn nhnh tm thy C. personata; H si nm c bn trong v bn ngoi tm thy C.arachidicola. Vo thi im hnh thnh bo t nh, si nm tp trung thnh khi dy c dng qu cu gi l cht nn (stroma), cht nn pht trin bn di lp biu b trong nhng l hng di kh khu ca l; Bo t nh pht trin trn vch ngn nhng cung bo t mu sm, c nhng bin i rt ln v kch thc ca bo t v cung bo t; Bo t di, mnh, hp, thon nhn v cha rt nhiu vch ngn ngang (hnh 6.6). S pht trin ca nhng cung bo t ghp thnh cm sm mu, cong gp nh u gi, thng chng th ra ngoi cht nn ca t bo l cy ch, s phng thch bo t khi cung bo t nh to vt so nh ni n gn vo, bo t pht tn hiu qu nh cc git ma, gp iu kin nhit v m thch hp, mi bo t ny mm v to nn h si nm mi.
Figure 4.6
Conidiophore = cng mang ti bo t Hnh 6.6. Cung bo t v bo t ca Cercospora beticola (Sharma, 1998)
4.3 Ging CURCULARIA

c im N cng l mt thnh vin ca h Dematiaceae ca b Moniliales, n c gii thiu trn 30 loi;
39 Curvularia tm thy trn la (Benoit v Mathur, 1970) v nhiu cy trng khc. L tc nhn gy bnh m l, bnh r st (thi kh), bin dng ht, bin mu (bc mu) ht v thm ch thi r; Giai on hon chnh c bit l dng loi ca Cochliobolus, mt thnh vin ca Loculoascomycetes. Cung bo t ng thng, si ln (marconematous) v si n (mononematous). Bo t xon thnh vng trn cung bo t. Bo t thng cong. C 3 bo t trn mt l nhiu nht (hnh 6.7); S li ln ca rn ht bo t trn gp mt vi loi nh C.combopogonis, i khi cung bo t pht trin trn cht nn.
Figure 4.7
Hnh 6.7. Cung bo t v bo t ca Curvularia lunata (Sharma, 1998)
4.4 Ging PYRICULARIA

c im Pyricularia l mt thnh vin ca h Dematiaceae, b Moniliales; P. oryzae l nguyn nhn chnh gy bnh o n (n lp) la; Nm bnh thng git cht hon ton cy con, i khi trn la trng thnh, n cng nhim trn nhiu thc vt nh c mn tru voi (Euleusine coracana) v k (Setaria italica). H si nm pht trin v phn nhnh, c vch ngn, si nm ni bo hoc gian bo, t bo thng nhiu nhn, cung bo t thng n, di, mnh, c hoc khng c vch ngn, v thng khng phn nhnh. Mt nhm cung bo t mc trn cht nn, bo t mu nu nht, dng qu l ngc v c 2 vch ngn (to 3 ngn) (hnh 6.8); Mi bo t gn vi cung bo t bi rn ht (hilum) nh nh li. Bo t c phng thch khi m rt cao c bit vo ban m, c th s v ra ca rn ht gy phng thch bo t.
40
Massarina, mt nm Loculascomycetes c gii thiu nh giai on hon chnh ca Pyricularia aquatica, trong khi giai on hon chnh ca P.grisea l Magnaporthe grisea li l nm Pyrenomyceteous.
Figure 4.8
Conidia = bo t nh, conidiophore = cng mang ti bo t Hnh 6.8. Cung bo t v bo t ca Pyricularia oryzae (Sharma, 1998)
4.5 Ging FUSARIUM

c im Fusarium l chi ln nht trong Tuberculariaceae, chng hoi sinh hoc k sinh trn nhiu cy trng, cy n tri v rau. N l nguyn nhn chnh lm ho r cy ch. H si nm lan to khp m mch v lp kn mch g. S lp mch g s cn tr qu trnh chuyn vn nc lm ho cy (hnh 6.9 A), Fusarium cng sn xut mt s cht c tit vo mch dn cy ch cng c th gy ho r, nhiu loi thc vt b Fusarium tn cng (hnh 6.9. A). Sau y l vi loi Fusarium gy bnh ho l v cy ch (trong ngoc n): F. udum (trn u sn Cajanus cajan), F.oxysporum bv. licopersici (trn c chua Lycospersicon esculentum), F. lini (trn cy lanh Linum usitatissimum) F. solani (trn khoai ty Solanum tuberosum) v F. orthaceras (trn u m-u Th Nh K Cicer arietium). H si nm phn nhnh, c vch ngn, si nm thng khng mu, chuyn mu nu khi gi. H si nm sn sinh c t tit vo h mch gy ho cy ch.
41
4.5.1 Sinh sn
Fusarium sinh sn v tnh trung bnh gia 3 kiu bo t v tnh l bo t nh ln (Macroconidia), bo t nh nh (Microconidia) v bo t vch dy (hu bo t - Chlamydospores).Macroconidia di, nhiu nhn, hnh lim hoc thn cong sinh ra t cung bo t. u v cui bo t ln thun nhn (hnh 6.9 C); Mt vi loi bo t ln tch ri v khng gn trn cung bo t, nhng t bo sinh bo t ln gi l th bnh (phialide) (hnh 6.9.B). Tiu bo t nh thng n nhn i khi 2 ngn, hnh cu hoc hnh trng c sinh ra t mt th bnh hay nhng cung bo t phn nhnh hoc khng phn nhnh (hnh 6.9 D); Tiu bo t nh thng c gi trong mt nhm nh v tiu bo t nh ca Fusarium rt ging bo t ca Cephalosporium v th giai on ny thng c qui vo nm Cephalosporium. Bo t vch dy (hnh 6.9 E) hnh trn hoc hnh trng, vch dy, nm tn cng hoc chen gia cc si nm gi. Chng c th pht trin n hoc thnh chui, chng tch ra v mc cc ng mm nu bo t gp iu kin thun li, Hu bo t hay bo t vch dy rt bn v tn ti c lp trong thi gian di.
Figure 4.9
Hnh 6.9. A, Fusarium udum gy bnh ho l (trn u sn Cajanus cajan); B, cung sinh bo t v bo
42
t nh ln; C, i bo t nh (macroconidia); D, tiu bo t nh (microconidia); E, bo t vch dy (hu bo t)(Chlamydospore)(Sharma, 1998) Lp Coelomycetes c tnh chung 1. 2. 3. 4. 5. Nhm ny k sinh v hoi sinh trn thc vt c mch trn cn. Mt s k sinh bc hai trn nm khc. Tn l th qu tht, h si nm c vch ngn. Khng c t bo chi. Bo t v cung bo t sp xp trn ti bo t hoc cm cung bo t. Ti bo t b mt hay nm su bn trong, hnh cu, tri phng hoc hnh da, mt vch to nhng t bo cng ng knh. 6. Cm cung bo t nm bn trong cht nn thiu phn bn v trn vch (Sutton,1973) 7. Bo t n bo, rng sm, trong sut hoc c sc t t bo. Phn loi Sutton (1973) ngh Coelomycetes vo 2 b: 1. Melanconiales: th qu kiu cm cung bo t. 2. Sphaeropsidales: th qu kiu ti bo t phn. c im b Melanconiales 1. 2. 3. 4. Cc c th k sinh hoc hoi sinh trn thc vt Th qu kiu cm cung bo t. M cht nn gii hn c s ca th qu. Th qu di lp cutin, di biu b, hoc di chu b v v ra bi s rn nt ca m vt ch.
Melanconiales c mt h Melaconiaceae (Sutton, 1973) vi ging Colletotrichum c m t y. c im b Sphaeropsidales 1. 2. 3. 4. Tnh cht ch yu l th qu dng ti bo t phn. C th c kch thc hin vi k sinh hoc hoi sinh. Mng bao quanh th qu l nhu m gi. Bo t nh dng t cc vch ngn.
4.6 Ging COLLETOTRICHUM

c im N c m t c 11 loi (von Arx,1957; Sutton, 1973). Nhng Alexopoulos v Mims (1979) th xut trn 1000 loi hnh thc trong ging ny c m t trc y, phn ln chng trng tn. Theo kin gn nht ca Baxter v cng s (1985), Colletotrichum c gii thiu c 21 loi: C. coccodes, C. dematium, C. gloeosporioides, C. graminicola, C. falcatum v C. capsici. . . l nhng loi thng gy bnh thn th (anthracnose). Bnh thi ma y l bnh thng gp ma do C. falcatum, nm tn cng chnh vo thn v l (hnh 6.10), phin l tr nn nht hoc sm v r xung, thn c nhiu vt nt v lp mng si pht trin dc thn; S ho ch yu b mch, i khi vo tn trong rut v thn b thi, rt ngn ti cc t. Si nm
43 Ni sinh, si nm mnh, phn nhnh, khng mu, c vch ngn, si nm c ni bo v gian bo; Nhiu ht du c sn xut trong mi t bo ca h si nm; Khi chn si nm tr nn sm mu v bn xon li thnh dng cht nn nh di lp ngoi cng. Sinh sn Colletotrichum ch sinh sn v tnh bng bo t nh, bo t nh pht trin trn cung bo t trong dng th qu l cm cung bo t (hnh 6.11 A-C); Cm cung bo t c dng a phng, mt sau c cu trc phn mn, mi cm cung bo t gm lp cht nn, b mt sn sinh cung bo t trong sut (hnh 6.11.C). Cung bo t khng c vch ngn ko di n bo, dng lim, cong, bo t trong sut. Cng vi bo t v cung bo t l cc lng cng trn mi cm cung bo t, lng di cng, thun nhn, khng phn nhnh v a bo cu trc nh t cng (hnh 6.11.A-B) v Frost (1964) m t mt vi loi ca Colletotrichum c hoc khng c lng cng c th c kim sot bi s thay i m.
Figure 4.10
Hnh 6.10. Colletotrichum falcatum. A, cm bo t trong thn; B, cm bo t trn l (Sharma, 1998) S hnh thnh mt s ln ca bo t gy nt gy trn biu b vt ch, gp iu kin thun li, mi bo t mc t mt n nhiu ng mm hnh thnh h si nm (Hnh 6.11.D); a bm l dng ca Colletotrichum trong nui cy (Sutton, 1962, 1968) Si nm gi i khi hnh thnh vch dy, mu nu sm, hnh cu hoc khng u gi l hu bo t (Chlamydospores), N c th tn cng hoc chen gia si nm v tn ti trong thi gian di v khi tch ra chng cng mc mm hnh thnh si nm mi. Theo von Arx (1957) Sclerotia cng l mt dng c bit ca Colletotrichum,
44
Glomerella tucumanensis l mt nm ti (Ascomyceteous) c coi nh giai on hon chnh ca Colletotrichum falcatum.
Figure 4.11
Hnh 6.11. Cm bo t ca Colletotrichum lindemuthianum; B, cm bo t ca C. faltacum; C, cung bo t v bo t nh ca C. graminicola; D, bo t ny mm (Sharma, 1998)
Chng 5
Ngnh ph nm m (Basidiomycotina = Lp Basidiomycetes)

1
5.1 c tnh tng qut

1. Cc loi nm thuc ngnh ph ny sng trong t, hoi sinh hay k sinh. Nhm hoi sinh gy ra triu chng lm mc cy..., nhm k sinh gy bnh r, chy l, mc nh ca.... 2. Nhm ny ch sng trn k ch thc vt trong t nhin 3. Khun ty phn nhnh, pht trin v c vch ngn ngang, cm su vo trong k ch ht cht dinh dng, chng c mu cam, vng.... khun ty c s cp, th cp.... 4. Vch t bo cu to bi cc si chitin v glucans vi mi lin kt 1,3 v 1,6 [U+F062]-D-glucosyl 5. Cc si khun ty qun cht vo nhau to nh mt hnh dng ca r cy (rhizomorph) 6. Sinh sn v tnh vi nh bo t, bo t chia t (arthrospore), bo t vch mng (oidia), on khun ty v mc mm 7. Khng c c quan sinh dc c bit, hp nhn ch l s tip hp dinh dng (somatogamy) hay s tip tinh (spermatization) 8. c tnh bo t l nhng m bo t, chng pht trin mt M, m c th khng c vch ngn ngang (holobasidia) hay c vch ngn ngang (phragmobasidia), lun lun c 4 bo t m trong mt m, mi m bo t c mt nhn v ny mm ngay trong khun ty u tin. 9. V mt kinh t, ngnh ph NM M va gy hi va hu ch vi hng triu tn hoa mu b hi b bnh r v m l, chng tn cng c cy lng thc ln cy rng nhng c nhm c ch nh cc loi nm n nh nm trng Agaricus bisporus, Volvariella volvaria vi trn 300.000 tn cung cp cho con ngui nhng cng c loi nm c c t.
5.2 Khun ty v hp nhn (nhn kp)

C 3 loi khun ty bc 1, bc 2 v bc 3
5.2.1 Khun ty bc 1
Chng pht trin t s ny mm ca mt m bo t, gm nhng t bo n nhn cn gi l ng nhn (homokaryon), tuy nhin trong giai on khun ty a nhn sau phn chia cc vch ngn ngang thnh
1 This
45
46
CHNG 5. NGNH PH NM M (BASIDIOMYCOTINA = LP BASIDIOMYCETES)
t bo n nhn; m khng bao gi pht trin trn khun ty bc 1.
5.2.2 Khun ty bc 2 v nhn kp

Khun ty bc 2 gm nhng t bo nhn kp v pht trin bi s hp nhn ca 2 t bo n nhn. Trong nhng loi d tn, t bo hp nhn khi nhng khun ty bc 1 ca nhng loi khc nhau nhng trng hp ng tn (homothallic) th s hp nhn xy ra gia hai khun ty ca hai khun ty bc 1. Qu trnh phi hp ca khun ty bc 1 thnh khun ty bc 2 hay nhn kp gi l nhn kp ho (dikaryotization) hay nh bi ho (diploidization).
47
Figure 5.1
48
Hnh 5.1. Qu trnh nhn kp nm m (A - F), khun ty th cp to ra M v BO T M (Sharma, 1998) Nhn kp ho trong ngnh ph Basidiomycotina c th xy ra t s hp nhn ca: 1. 2. 3. 4. 5. 6. t bo dinh dng ca hai khun ty xut pht t khun ty bc 1 ca hai dng khc nhau (hnh 5.1) Hai m bo t ca hai dng khc nhau Mt bo t vch mng ca dng A v mt t bo ca khun ty bc 1 ca dng B Mt bo t m ny mm v mt t bo n bi ca mt m Hai t bo n bi ca mt m Hai m hnh thnh t s ny mm ca bo t than (smut spore) ca dng A v dng B
T bo nhn kp ca khun ty bc hai phn chia to ra nhng t bo nhn kp t s phn ct ng thi ca hai nhn; m pht trin t nhng t bo nhn kp ca khun ty nhn kp.
5.2.3 Khun ty bc 3
Khun ty bc 2 ca mt s nm m tin ho s to ra m nang (basidiocarps) gi l khun ty bc 3.
5.3 To mu (Clamp connection)

Mu c hnh thnh trong hu ht cc loi ca ngnh ny, n hnh thnh trong sut s phn chia t bo khun ca khun ty bc hai, thng thng mt t bo phn chia trong khun ty b gii hn thnh t bo hon chnh. S hnh thnh mu tri qua cc bc sau (hnh 5.2): 1. Cng lc vi phn chia t bo nhn kp s xut hin mt on di gia hai nhn X v Y, on hnh thnh nh mt ci MU 2. Nhn Y di chuyn ra ngoi v to thnh mt MU 3. Nhn X v Y ng thi phn chia 4. Nhn Y va phn chia trong mu v nhn X va c phn chia tin v phi nhn Y, nhn Y trong mu tip hp vi nhn X. 5. S hnh thnh vch t bo ngn chia gia hai nhn X v Y vi nhn X v Y, phn chia t bo m v t bo con. 6. T bo con vi hai nhn X v Y tin ra phi trc.
5.4 M (Basidia)
5.4.1 Cu trc
m l mt b phn, c quan hay mt t bo nm; mang mt s bo t m trn b mt ca n. S bo t m ny c hnh thnh cc buc sau: hp t bo cht v hp nhn (karyogamy) ri gim phn v s bo t m l 4. Tuy nhin, chi Dacrymyces v Calocera c mi m ch cha 2 bo t m. Theo Talbort (1954), mi m c th chia lm 3 phn: TIN M (Probasidium), ni nhn s phn chia M M (Metabasidium), ni nhn s gim phn TA CUNG (Sterigma), phn trung gian gia hai trn Thng thng M c dng bu dng hay hnh thn (hnh 5.3)
49
Figure 5.2
Hnh 5.2. Qu trnh thnh lp mt MU trong mt khun ty (Sharma, 1998)
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Figure 5.3
Hnh 5.3. Cc loi M (Sharma, 1998)
5.4.2 Cc loi m
C hai loi m c cc nh khoa hc cng nhn, l: TON M (Holobasidium) : m khng c vch, ch l mt t bo n c VCH M (Phrabmabasidium) l mt m c nhiu t bo ko di, khng c vch ngn Mi vch m cha mt 1 v tr u tin phn chia gi l SINH M (Hypobasidium) v sau ny l NGOI M (Epibasidium)
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5.4.3 Pht trin ca mt TON M
Figure 5.4
m hnh thnh v pht trin trong th c mt lp bao bn ngoi gi l BO TNG (Hymenium) (hnh 5.4) Hnh 5.4. Cc giai on pht trin ca mt TON M (Sharma, 1998) Mt s t bo ca bo tng pht trin thnh mt M, thng thng t bo s to nn mt mu ri ko di ra sau nhn tip hp s tin hnh gim phn cho ra 4 nhn n bi v pht trin thnh 4 m bo t.
5.4.4 S pht trin ca VCH M

R v mui than (smut) cha nhng vch m, mt vch m trong than pht trin vi s ny mm ca mt bo t nh bi c vch dy, chung quanh c mt lp t bo nh bi ca mt khun ty (nh bi); Hai nhn trong mt bo t phi hp thnh mt nhn hp t nh bi. Bo t ny mm vi mt ng mm hay mt ngoi m (epibasidium). Trong giai on ny, v tr hnh thnh u tin ca bo t c gi l NI M (Hypobasidium); nhn nh bi tin hnh gim phn thnh 4 nhn n bi ri di chuyn vo trong mt ngoi m, sau phn on thnh 4 t bo n bi, t mi t bo ca ngai m pht trin mt cung (sterigma) v ti u mi cung s pht trin mt m bo t.
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5.5 BO T M (Basiospore)
5.5.1 Hnh thi
Bo t m c cu trc n bi nhng c mt s ging li cha n 2 nhn v trng h Dacrymytaceae, bo t m khng c vch ngn (Reil, 1974). Bo t m c hnh cu, bu dc, chai.... vi nhiu mu sc khc nhau nh vng, xanh, tm, nu, hay khng mu v vch trn lng. S lng bo t m c to ra t mt qu th rt ln v d nh nm Agaricus campertris c n 1,8 t bo t m trong 2 ngy hay trung bnh 40 triu bo t/gi.
53
Figure 5.5
Hnh 5.5. M hnh tiu biu ca mt BO T M ct ngang (Sharma, 1998) Phn gc y ca mt bo t m gi l T (hilum)(hnh 5.5), k bn trn l PH T (hilar appendix). Theo Pegler v Young (1975) vch bo t m gm c 5 lp: NGOI B BO T (ectosporium), NGOI K B BO T (perisporium), TRUNG B BO T (exosporium), NI K B BO T (episporium), v NI B BO T (endosporium). Lp ngoi th sn si, lp gia khng mu v bo t m non ch c 2 lp: NGOI B v NI B n khi trng thnh th pht trin 5 lp.
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5.5.2 C ch phng thch ca bo t m

Nhiu c ch c ngh nhng nhiu nh khoa hc ng c ch BONG BNG hay BT bt ngun t ph rn. Theo Buller (1909, 1922), bong bng ban u l dung dch (hnh 5.5) v tng kch thuc ln ln cho n khi m bo t t ngt ri khi cng nn cn gi l BONG BNG BULLER. Nhng nghin cu ca Wells (1965) di knh hin vi in t cho thy bao bn ngoi git dung dch l mt lp mng ca cung v chnh nhng p lc ca cung bao ny s lm bo t m phng thch th nhng theo Olive (1964) v Ingold v Dunn (1968) cho rng nhng git ny thay v l dung dch li l kh CO2 v nh bung ra d dng mang theo cc bo t m, c ch ny c tn PHNG THCH N (explisive discharge). Theo van Niel v ctv (1972), git bong bng ny c th l kh c th l dung dch v c hai u rt d dng gip cho bo t phng thch ra ngoi.
5.6 Phn loi

Ainsworth (1973) chia ngnh ph ny thnh 3 lp sau: 1. Lp Teliomecetes : khng c bo m v thay th bng bo t vch dy (chlamydospore) tiu bu l ging Puccinia v Ustilago 2. Lp Hymenomycetes : c bo m v tiu biu l ging Agaricus v Volvariella 3. Lp Gasteromycetes : c bo m
5.7 Lp Teliomycetes
Tiu biu cho lp ny l nm R st (rust) v mui than (smuts) k sinh trn thc vt
5.8 B Uredinales
H Puccinaceae (bo t ng [teliospore])
5.8.1 Ging [Chi] Puccinia

* Puccinia graminis Nm ny thuc nm k sinh bt buc trn cy lng thc vi hn 700 loi. Nm Puccinia graminis c chu k sinh trng trn 2 k ch khc nhau (la m v du [Berberis vulgaris]), chng s to ra vt g st trn l (hnh 5.6) nhng giai on nh bi ca vng i nm ny trn la m (hnh 5.7). Bo t r (urediniospore) l l mt cu trc cung pht trin thnh th hnh bu dc, cu trn, mi bo t cha 2 nhn v bao bng mt lo v dy, t 50.000 n 60.000 bo t trong m bo t r (uredinium). Cc m bo t r c tm thy trn thn, l la m c mu r , en. T mt bo t r ny mm v cho ta mt bo t m, mi bo t m c cu trc nh, n nhn n bi v chng d dng bay vo trong khng kh.
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Figure 5.6
Hnh 5.6. Cc m r st trn l v thn la m (A v B) trn l du (C v D)(Sharma, 1998) ** Chu k sinh trng ca nm Puccinia gramini Puccinia graminis gy ra r st trn l v thn la m ch l k ch 1 v cy du tm (Berberis vulgaris) l k ch 2. Chu k sinh trng c 5 giai on trong 3 giai on trn la m v 2 giai on sau cy du tm. trn la m ch gm khun ty nh bi, khun ty pht trin trn l v thn la m s to ra cc bo t ng (teliospore), c vch dy v lng thung c dng gn trn v pht tn trong khng kh cng nh sng st kh lu. Khi bo t ng ny mm cho ra cc m bo t xut pht t on sinh m (hypobasidia) ri to ra hnh ng di gi l NGOI M (epibasidia), nhn nh bi di chuyn vo ngoi m ri phn chia
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thnh 4 nhn n bi trong m bo t v pht tn trong khng kh nhng n khng th ny mm trn la m v ch ny mm trn cy du tm v trn l ca cy du c cht dinh dng cn thit cho bo t m ny mm v pht trin. trn cy du tm, bo t m to thnh ci ti bo t phn (spermaforium).
5.9 B Ustilaginales 5.10 H Ustilaginaceae

5.10.1 Ging [Chi] Ustilago
Ging ny c hn 400 loi k sinh trn thc vt v hu ht thuc h Graminae v Cyperaceae trong c nhiu cy lng thc quan trng; Triu chng th hin rt r l chng gy ra bnh MUI THAN trn ht vi nhng bo t than trong mt ci bc c v mng v khi gi thi mnh th bc v ra phng thch bo t vo trong khng kh (hnh 5.8).
57
Figure 5.7
Hnh 5.7. L v thn la m nhim nm Puccinia graminis vi cc m bo t ng (A-C), mt vi m bo t ng (urediniospore) ny mm vi mt ng mm xuyn vo nhu m l lu m (D - F)(Sharma, 1998)
58
Figure 5.8
Hnh 5.8. Triu chng nhim bnh mui than do Ustilago gy ra trn la m (A - B), trn bp (C), trn la kiu mch [oat](D), trn la mch (E), trn c ch [Cynodon dactylon](F), trn ma ng (F)(Sharma, 1998) Khun ty phn nhnh, c vch ngn ngang, c 2 loi khun ty: khun ty s cp l nhng khun ty hnh thnh t s ny mm ca m bo t vi nhng t bo ch cha mt nhn n bi v vy khun ty ny cn gi l khun ty n bi (monokaryotic mycelium), chng chuyn sng khun ty th cp hay l cht, khun ty th cp cha nhn nh bi v thng gp k ch, khun ty ny cn gi l khun ty nh bi (dikaryotic mycelium). Qu trnh chuyn t khun ty n bi sang khun ty nh bi cn gi l hin tng nh bi ho (diploidization = dikaryotization) trong 2 nhn ca 2 dng khc nhau trong t bo n bi bt cp thnh t bo nh bi, qu trnh ny xy ra di nhiu hnh thc sau:
59 1. 2. 3. 4. 5. 6. Phi Phi Phi Phi Phi Phi hp hp hp hp hp hp gia 2 khun ty s cp ca 2 dng khc nhau (hnh 5.9) nh trng hp Ustilago maydis gia 2 ng t 2 m bo t ny mm nh trng hp U. anthearum gia 2 t bo n bi ca mt m bo t ca 1 dng v 1 ng mm t 1 dng khc nh trng hp U. hordei 2 m bo t t ti m bo t phn nh trng hp U. nuda gia 1 m bo t v 1 t bo m bo t t dng khc nh trng hp U. violacea
Figure 5.9
Hnh 5.9. Nhng trng hp nh bi ho ca Ustilago nh U. maydis [A], U. anthearum [B], U. hordei [C-D], U. nuda [E], U. violacea [F](Sharma, 1998) ** Sinh sn Ustilago maydis (gy bnh than bp) Khun ty nh nh bi trn bp tip tc ny chi trn tri bp to thnh cc khi u (hnh 5.10), khi cc khun ty th cp pht trin cc m bo t ng hnh thnh vi cc dng cu, trn vch dy, hai nhn n bi hp thnh mt nhn nh bi sau cc bo t ny ny mm cho ra mt ng di gi l TIN KHUN TY (promycelium), nhn nh bi di chuyn vo trong tin khun ty v phn chia thnh 4 nhn n bi, nhn ca mi t bo tin khun ty phn chia thnh 2 nhn con, mt i vo chi bn cnh v mt vn cn li t bo ch, chi s pht trin thnh m bo t, cn nhn trong t bo ch tip tc phn chia cho chi th hai, th ba..... m bo t trn, bu dc, v mng cha mt nhn n bi v khi ny mm cho mt khun ty n bi.
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Cc than bo t d b nc nng lm h, ch cn nc m 26oC n 30oC trong 4 - 5 gi hay 54oC trong 10 pht s lm cc bo t mt ny mm (cht) v vy cn ngm ht ging trong nc m, sch phng nga cc loi nm ny. Ngoi ra cn th dng bin php k kh cc bo t khng th h hp v mt kh nng ny mm.
5.11 Lp Hyphomycetes
y l lp ln nht trong ngnh ny, bo m pht trin tt nht. a s cc loi trong lp ny l hoi sinh, mt s rt t l k sinh. Bo t m chnh l bo t banh (ballstopore) Lp ny chia lm 2 lp ph sau: 1. Lp ph Holobasidiomycetidae 2. Lp ph Phegmabasidiomycetidae
61
Figure 5.10
62
CHNG 5. NGNH PH NM M (BASIDIOMYCOTINA = LP BASIDIOMYCETES) Hnh 5.10. Chu k sinh trng ca nm Ustilago maydis (Sharma, 1998) Lp ph Holobasidiomycetidae c 6 b trong b Agaricales l quan trng nht.
5.12 B Agaricales
B ny c nhng c im nh c vi di (pileus) khc nhau (hnh 5.11); trong sinh sn v tnh vi nhng m v bo t m hin din trong mt qu th gi l bo m (basidiocarp), tuy nhin s phn nhnh vi nhng rnh (gill) v cng c nhng vng (ring) v ni vi phn cui ca cng c mt bao (volva).
Figure 5.11
Hnh 5.11. Cu trc ca bo m ca mt s loi thuc b ny nh Coprinus atramentarius (A), Agaricus compestris (B), m hnh tiu biu ca mt bo m vi bao c m (Sharma, 1998) Nm c 2 loi n c v khng n c; loi nm n c c nhiu cht dinh dng vi nhiu protein v vitamin cng thm nhng hng v c trng.
5.13 H Agaricaceae
5.13.1 Ging [Chi] Agaricus campestus
Khun ty s cp l khun ty ngn, khng vch ngn, bo t m n bi ny mm cho ra nhng khun ty nhiu nhn. S phi hp 2 khun ty s cp ca 2 dng khc nhau to ra khun ty th cp (hnh 5.12), chng pht trin thnh N (knots) vi nhng khun ty dng r (rhizomorph) bn di v t y chng pht trin thnh qu th . Khun ty th cp bc 1 c th thnh khun ty th cp bc 2 v tim sinh trong t rt lu. * Sinh sn v tnh: rt him
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5.14 ** Sinh sn hu tnh

Trong nhm ny khng c c quan sinh dc, im chnh ca ging ny l D TN, s tip hp ca 2 khun ty ca 2 dng khc nhau hnh thnh khun ty th cp; t y pht trin thnh n vi nhng khun ty r bn di v pht trin thnh nhng qu th vi nhng rnh bn di (gill) v khi qu th bung d khi cc rnh dn ra v c mt vng bn di cng. Khi qu th bung d lc bo m chn, nu ct qu th ra, chng ta s thy s tp hp v b cht ca nhng khun ty v cc rnh c 3 vng khc nhau.
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Figure 5.12
65 Hnh 5. 12. Vng i ca nm Agaricus campestus (Sharma, 1998) Vng tr (trama) gm nhng khun ty b cht theo mt chiu nht nh v tn cng l phn d (pileus) Vng gia (subhymesium) ca rnh, y khun ty c nhiu nhn nh bi Vng ngoi (hymenium) ca rnh, cha mt hay nhiu lp t bo c nhn nh bi v tn cng gm nhng t bo sinh sn gi l M (hnh 5.13)
66
Figure 5.13
67 Hnh 5.13. Cu to bn trong ca bo m ca Agaricus campestus (Sharma, 1998) M l nhng t bo n nhn nh bi do s kt hp t nhn n bi sau gim phn thnh 4 nhn n bi v s phn chia 2 gii khc nhau giai on ny vi 2 bo t m l dng + v 2 l dng - , tn cng ca m l s pht trin thnh 4 cng (sterigmata) v nhn n bi di chuyn vo 4 cng ny v cui cng cng s pht trin thnh 4 bo t m theo nguyn tc PHNG THCH N v bo t m ny mm cho ra 1 khun ty s cp ca dng + hay dng - .
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Chng 6
Ngnh ph Nm Bt Ton (Deuteromycotina = lp Deuteromycetes)

1
6.1 Gi thiu chung

Ngnh ph Deuteromycotina gm mt h thng cc nhm nm b thiu hoc khng pht hin c nhng c im ca nm hon chnh (tip hp, nang hoc m); nhng nm ny khng mang bo t tip hp (zygospore), bo t nang (ascospore), hoc bo t m (bo t nh th sinh - basidiospore). Nm ny thiu giai on sinh sn hu tnh trong vng i nn ngi ta gi chung l nm khng hon chnh hay Nm bt ton (Imperfect fungi). Cc c th ch sinh sn bng hnh thc v tnh, ch yu l bng bo t nh (conidia) pht trin trn cung bo t nh (conidiophores). Sutton (1973) ngh Deuteromycotina nh "l mt tp hp cc kiu nm sinh sn bng bo t vi dng khng hp nhn bi s gim nhim"
6.1.1 c im chung
- Deuteromycotina c m t bi trn 15.000 loi (Ainsworth, 1973) phn ln sng trn cn; Mt s ln nm bt ton thu sinh (Alatospora, Tricladium, Pyricularia) tm thy trong c mi trng bin v nc ngt, a s cc c th hoi sinh hoc k sinh, l nguyn nhn gy mt s bnh trn thc vt v ng vt. - Ngoi tr dng n bo ging nh nm men ca Blastomycetes, hu ht tt c Deuteromycotina cn li u c h khun ty (mycelium) tht, gm c s pht trin si, phn nhnh v vch ngn si nm (hypha) - H si nm thng c gian bo hoc ni bo v mi t bo cha nhiu nhn. - Vch ngn trn tt c cc loi c kho st hu nh ging vi Ascomycotina,c mt l thng gia mi vch. - Hon ton khng c sinh sn hu tnh, sinh sn ch yu bng dng bo t c bit l bo t nh (conidia); Bo t l bo t nh bt ng, pht trin bn ngoi cung bo t nh, v phn ny th Deuteromycotina ging nh Ascomycotina. Bo t nh c hnh dng, kch thc, mu sc thay i. . . n c th trong sut hoc c mu sc thay i, n nhn hoc a nhn, c vch ngn ngang, dc hoc khng; N c th c hnh trng (oval), thun di, hnh cu, dng sao, dng hi cong, dng si, hnh a, dng cun xon hay nhng dng khc. - Bo t nh c sinh trc tip t cung bo t hoc t mt vi kiu th qu nh; b si bo t (synnema) (hnh 6.1), cm cung bo t (arcevulus) (hnh 6.2), gc cm bo t nh (sporodochium) hoc ti bo t phn (pycnidium). Nhng th qu ny l cc m mm gi trong phm vi ni bo t c sinh
1 This
69
70
ra. Sutton (1973) pht hin ch c 3 kiu th qu l ti bo t phn, cm cung bo t v lp cht m (stroma) - Gii tnh i ng (Parasexuality) (d tnh) c m t trn mt s Deuteromycotina; di hin tng ny, c s hnh thnh cc u tch hp cht nguyn sinh, tip hp nhn v n bi ho ti mt thi im c bit hoc mt v tr c bit trong vng i ca nm. Hin tng ny c cp mt cch gin tip bi Pontecorvo v Roper (1952) v nhng nghin cu b sung ca Pontecorvo (1956,1958), Davis (1966). Mt vi c s bn ngoi ca hin tng ny bao gm s thnh lp ca hai kiu nhn (heterokaryotic) (d hch), si nm; c s tip hp nhn v nhn ln ca nhn lng bi, c s xy ra phn bo gin phn (mitosis), s "tuyn chn" ca nhn lng bi v cui cng l s n bi ho mt vi nhn lng bi trong khun ty.
Figure 6.1
Hnh 6.1. Mt b si bo t (synnema) ca Arthrobotryum (Sharma, 1998)
71
Figure 6.2
Hnh 6.2. Nm Collectrichum [A], Fusarium [B], Endothia [C](Sharma, 1998)
6.1.2 Tm quan trng

Phn ln cc nm trong ngnh Deuteromycotina u c tc ng trc tip hoc gin tip n cuc sng hng ngy ca con ngi. Mt s nm thuc lp Hypomycetes nh mt s loi thuc ging Alternaria gy bnh g st khoai ty, c chua v mt s cy trng h Solanaceae; Pyricularia gy bnh o n la; Cercospora gy bnh m l bng vi, thuc l, C. apii gy l lot ngoi da ngi. Ging Fusarium gy bnh thi ma, thi qu c chua (Fusarium solani), thi kh khoai ty, h hnh ti; Cc c th thuc chi Colletotrichum gy bnh lot cy, C. lagenarium gy bnh thi hng bu b (Sharma, 1989; Lng c Phm, 2002). c t nm Fusarium, Alternaria gy xut huyt ni quan (d dy, rut, gan,. . .), ri lon thn kinh.
72
6.2 Phn loi

Hnh dng, kch thc, vch ngn, mu sc v s trang tr ca bo t l tiu chun chnh phn loi Deuteromycotina; Song song vi vic da vo hnh thi ca bo t th s pht trin ca chng (tn v kiu pht trin phi nguyn bo, Kendrick, 1971), hnh dng v s pht sinh ca cung bo t nh cng nh s t hp ca chng trong dng th qu xc nh (b si bo t (synnema), cm cung bo t (arcevulus), gc cm bo t nh (sporodochium) hoc ti bo t phn (pycnidium) cng l nhng c im phn loi quan trng. Ainswoth (1973) chia ngnh ph Deuteromycotina theo 3 lp: Khun ty khng pht trin hoc pht trin yu; dng c th ging nh nm men v c s ny chi: Blastomycetes Khun ty tht; khng ny chi; si nm bt dc hoc sinh bo t trn cung, khng c s tp trung thnh ti bo t hay cm cung bo t: Hypomycetes Khun ty tht; bo t tp trung trong ti bo t hoc trn cm cung bo t: Coelomycetes Alexopoulos v Mims (1979) a ra 3 lp ph hnh thc l Blastomycetidae, Hypomycetidae v Coelomycetidae.
6.2.1 Lp Hypomycetes
6.2.1.1 c tnh chung Phn ln cc c th sng hoi sinh trn thc vt hoc k sinh. Si nm pht trin mnh, c vch ngn v phn nhnh. Ch yu sinh sn bng bo t (Moniliales) nhng mt s ch sinh sn bng phn on (fragmentation) nh Rhizoctonia v Sclerotium. Bo t ca chng kh hoc nhy nht. C ti bo t ln cm cung bo t u khng c trong s sinh sn ca bt k c th no.
6.2.1.2 Phn loi Alexopoulos v Mims (1979) cng nhn 2 b hnh thc (Moniliales v Agromycetales) di phn lp hnh thc Hyphomycetidae, cc c th ca Hyphomycetes sinh ra bo t c t trong b hnh thc Moniliales nhng nhng dng thiu bo t v sinh sn bng phn on si nm th c t vo b hnh thc Agromycetales. 6.2.1.3 c im ca b Moniliales Phn ln c th hoi sinh hoc k sinh v bo t ca chng pht trin trn nhng si nhnh chuyn bit l cung bo t (sporophore) hoc cung bo t nh (connidiospore), chng c ngh thnh 4 h hnh thc (form-class) sau: 1. Moniliaceae: cung bo t tch ra t mt si no hoc khng c; bo t v h si nm trong sut hoc c mu sng, i din Monilia. 2. Dematiaceae: bo t v h si nm mu sm. i din Altenaria, Curvularia, Cercospora, Helminthosporium, Drechslera. 3. Tuberculariaceae: bo t v cung bo t nh c sinh ra t cm cung bo t. i din Fusarium. 4. Stilbellaceae: bo t v cung bo t nh pht trin trong b cung bo t nh. i din Graphium.
73 6.2.1.4 a. Ging ALTERNARIA * c im Nhm nm ny kh ph bin, c nhiu loi hoi sinh v gy cht tng phn cy trng. Alternaria l tc nhn gy nhim chnh trong nui cy trong phng th nghim. Bo t ca chng rt ph bin trong bi bm trong nh, trong khng kh v l tc nhn chnh gy d ng (Hyde v Williams, 1946), mt s bnh v da v vi ri lon nghim trng c th ngi.Nhiu loi Alternaria k sinh trn thc vt. Trn cc c th thuc h Solanaceae (khoai ty), Alternaria cho triu chng bnh r st sm hn l Phytophthora infestans (thuc lp Oomycetes, tc nhn gy bnh r st mun (late-blight) khoai ty), ch ring Alternaria c gi l bnh r st sm. Triu chng sm ca bnh l nhng m nh mu vng nu trn l, sau lan rng to nhng vt hnh nhn ng tm; Ton b phin l, cung l, gn l v thm ch c h thng mch dn cng tn thng t gy do b nhim. Phn cn li ca ng mch c mu nu.
Figure 6.3
Hnh 6.3. Alternaria solani trn khoai ty (Solanium tuberosum)(Sharma, 1998) Alternaria alternata (=A.tenuis) l nguyn nhn gy bnh m en trn la m trong khi A.triticina gy bnh r st (thi kh l) (Bhownik, 1969). A. brassicae v A.brassicicola tn cng trn ht Brascica (h ci bp) cn A.solani (hnh 6.3) gy bnh r st sm trn khoai ty v cc loi khc thuc h Solanaceae; Mt vi loi Alternaria khc (vi k ch ca chng trong ngoc n) l A. citri (trn l h cam qut Citrus sp.), A..helianthi (trn hng dng Helianthus annuus) v A. palandui v A. porri gy chy l trn hnh ty, ti. H si nm Mu nu sng, mnh, phn nhnh mnh, si nm c vch ngn trc ht l gian bo, sau c th tr thnh ni bo.;Mi t bo thng c nhiu nhn. Theo Knox-Davis (1979) th nhng t bo sinh dng ca A.brasicicola cha t mt ti nhiu nhn, u mt t bo si nm c 27 nhn v nhng t bo gi c n 33 nhn. Sinh sn Ging Alternaria ch yu sinh sn bng cch to bo t nh; Giai on hon chnh ca Alternaria l Pleospora infectoria (hnh 6.4) mt loi nm Loculoascomycetous (Webster, 1980).
74
Figure 6.4
Hnh 6.4. Mt nha bo t phng (ascus) ca Pleospora infectoria mt loi nm Loculoascomycetous giai on hon chnh ca Alternaria (Sharma, 1998) Bo t nh pht trin trn cung bo t nh ngn, sm mu v thng v nh hnh; Mt bo t pht trin nh l chi ngn ca t bo nh trn cung bo t nh (hnh 6.5. A-D); N khng pht trin bng cch tht eo v m rng phn cht t bo ca cung bo t. Nhng bo t non c phn cch bng vch ngn ngang (hnh 6.5. E) vi s pht trin ca vnh hnh khuyn vo bn trong (Campbell,1970); Trung tm mi vch ngn c mt l thng ni cht gia cc t bo ca bo t, sau mt s t bo phn cch bi vch ngn dc (hnh 6.5. F). Nhm bo t vi vch ngn v dc nh th c gi l dng qu du (muriform) hoc bo t li (dictyospore), thng th phn chp bo t ny chi v cui cng to thnh si ca bo t (hnh 6.5. G). i khi chi c th pht trin t t bo thp hn hay gn vo bo t to nn nhnh ca si bo t (hnh 6.5. H-I). S m rng thm ca si bo t s ngng khi c s bt kn l nn bo t; Bo t chn l mt qu th nhiu nhn c vch ngang v dc; N c bao quanh bi 2 lp vch, tng ngoi c sc t ca t bo, tng trong trong sut (Campbel, 1969, 1970). Theo Knok-Davies (1979) cung bo t chn cha vi nhn (0-3) trong khi bo t cha 1-2 nhn, Purkayastha v cng s (1980) nghin cu siu cu trc b mt ca 5 loi Alternaria gy bnh (A.longissima, A.cassiae, A.tenuissima, A.raphani v A.sonchi); Cc ht bo t c pht tn nh gi, gp iu kin nhit v m thch hp bo t ny mm to t 5 n 10 ng (hnh thnh si nm)(hnh 6.5. J).
75
Figure 6.5
Conidiophore = cng mang t1ui bo t, bud = chi, conidia = bo t nh, germ tube = ng mm, lateral bud = chi hng, young conidium = bo t nh non Hnh 6.5. A-F s pht trin bo t nh ca Alternaria solani; G, 2 bo t nh thnh chui ca A.brascicae; H, bo t ny chi ca A.brasicicola; I, chui bo t phn nhnh ca A.brascicae; J, bo t ny chi ca A.brasicicola (Sharma, 1998) Kim sot bnh thi li S lun phin ma v l c li v bnh ch yu t t trng; Thuc phun tr nm tt nht l loi c cha ng hoc km, cch khong 15 ngy trong phm vi kim sot d phng. Azariah v cng s (1962) ch trng s dng hn hp Bordeux trong khi Mathur v cng s (1971) th gii thiu phun Zineb v Dithane M-45. 6.2.1.5 b. Ging CERCOSPORA c im Ging nh Alternaria, Cercospora cng l mt chi ln trong h Dematiaceae, c i din bi trn 2000 loi (Ellis, 1971) nhng s ln xut hin rt nhiu v hu nh ng dng (Webster,1980). Cercospora l nguyn nhn gy bnh m l trn c chua, rau dip, khoai ty, bng vi, la, u phng, t, u trng ct
76
(piegon pea - arhar), c ci ng, thuc l. . . v nhiu cy trng kinh t quan trng khc; C. personata l tc nhn gy bnh m gch nu u phng (Arachis hypogea), C.gossypina gy bnh m l trn bng vi (Gossypium herbaceum) v C. oryzae gy bnh gch nu trn la, C. apii gy bnh trn ngi v c th l nguyn nhn gy nhng vt l lot trm trng trn mt trng rt kinh khng. (Emmons v ctv, 1975). H si nm pht trin mnh, phn nhnh v c vch ngn mng, si nm ni bo, gic mt phn nhnh tm thy C. personata; H si nm c bn trong v bn ngoi tm thy C.arachidicola. Vo thi im hnh thnh bo t nh, si nm tp trung thnh khi dy c dng qu cu gi l cht nn (stroma), cht nn pht trin bn di lp biu b trong nhng l hng di kh khu ca l; Bo t nh pht trin trn vch ngn nhng cung bo t mu sm, c nhng bin i rt ln v kch thc ca bo t v cung bo t; Bo t di, mnh, hp, thon nhn v cha rt nhiu vch ngn ngang (hnh 6.6). S pht trin ca nhng cung bo t ghp thnh cm sm mu, cong gp nh u gi, thng chng th ra ngoi cht nn ca t bo l cy ch, s phng thch bo t khi cung bo t nh to vt so nh ni n gn vo, bo t pht tn hiu qu nh cc git ma, gp iu kin nhit v m thch hp, mi bo t ny mm v to nn h si nm mi.
Figure 6.6
Conidiophore = cng mang ti bo t Hnh 6.6. Cung bo t v bo t ca Cercospora beticola (Sharma, 1998)
6.3 Ging CURCULARIA

c im N cng l mt thnh vin ca h Dematiaceae ca b Moniliales, n c gii thiu trn 30 loi;
77 Curvularia tm thy trn la (Benoit v Mathur, 1970) v nhiu cy trng khc. L tc nhn gy bnh m l, bnh r st (thi kh), bin dng ht, bin mu (bc mu) ht v thm ch thi r; Giai on hon chnh c bit l dng loi ca Cochliobolus, mt thnh vin ca Loculoascomycetes. Cung bo t ng thng, si ln (marconematous) v si n (mononematous). Bo t xon thnh vng trn cung bo t. Bo t thng cong. C 3 bo t trn mt l nhiu nht (hnh 6.7); S li ln ca rn ht bo t trn gp mt vi loi nh C.combopogonis, i khi cung bo t pht trin trn cht nn.
Figure 6.7
Hnh 6.7. Cung bo t v bo t ca Curvularia lunata (Sharma, 1998)
6.4 Ging PYRICULARIA

c im Pyricularia l mt thnh vin ca h Dematiaceae, b Moniliales; P. oryzae l nguyn nhn chnh gy bnh o n (n lp) la; Nm bnh thng git cht hon ton cy con, i khi trn la trng thnh, n cng nhim trn nhiu thc vt nh c mn tru voi (Euleusine coracana) v k (Setaria italica). H si nm pht trin v phn nhnh, c vch ngn, si nm ni bo hoc gian bo, t bo thng nhiu nhn, cung bo t thng n, di, mnh, c hoc khng c vch ngn, v thng khng phn nhnh. Mt nhm cung bo t mc trn cht nn, bo t mu nu nht, dng qu l ngc v c 2 vch ngn (to 3 ngn) (hnh 6.8); Mi bo t gn vi cung bo t bi rn ht (hilum) nh nh li. Bo t c phng thch khi m rt cao c bit vo ban m, c th s v ra ca rn ht gy phng thch bo t.
78
Massarina, mt nm Loculascomycetes c gii thiu nh giai on hon chnh ca Pyricularia aquatica, trong khi giai on hon chnh ca P.grisea l Magnaporthe grisea li l nm Pyrenomyceteous.
Figure 6.8
Conidia = bo t nh, conidiophore = cng mang ti bo t Hnh 6.8. Cung bo t v bo t ca Pyricularia oryzae (Sharma, 1998)
6.5 Ging FUSARIUM

c im Fusarium l chi ln nht trong Tuberculariaceae, chng hoi sinh hoc k sinh trn nhiu cy trng, cy n tri v rau. N l nguyn nhn chnh lm ho r cy ch. H si nm lan to khp m mch v lp kn mch g. S lp mch g s cn tr qu trnh chuyn vn nc lm ho cy (hnh 6.9 A), Fusarium cng sn xut mt s cht c tit vo mch dn cy ch cng c th gy ho r, nhiu loi thc vt b Fusarium tn cng (hnh 6.9. A). Sau y l vi loi Fusarium gy bnh ho l v cy ch (trong ngoc n): F. udum (trn u sn Cajanus cajan), F.oxysporum bv. licopersici (trn c chua Lycospersicon esculentum), F. lini (trn cy lanh Linum usitatissimum) F. solani (trn khoai ty Solanum tuberosum) v F. orthaceras (trn u m-u Th Nh K Cicer arietium). H si nm phn nhnh, c vch ngn, si nm thng khng mu, chuyn mu nu khi gi. H si nm sn sinh c t tit vo h mch gy ho cy ch.
79
6.5.1 Sinh sn
Fusarium sinh sn v tnh trung bnh gia 3 kiu bo t v tnh l bo t nh ln (Macroconidia), bo t nh nh (Microconidia) v bo t vch dy (hu bo t - Chlamydospores).Macroconidia di, nhiu nhn, hnh lim hoc thn cong sinh ra t cung bo t. u v cui bo t ln thun nhn (hnh 6.9 C); Mt vi loi bo t ln tch ri v khng gn trn cung bo t, nhng t bo sinh bo t ln gi l th bnh (phialide) (hnh 6.9.B). Tiu bo t nh thng n nhn i khi 2 ngn, hnh cu hoc hnh trng c sinh ra t mt th bnh hay nhng cung bo t phn nhnh hoc khng phn nhnh (hnh 6.9 D); Tiu bo t nh thng c gi trong mt nhm nh v tiu bo t nh ca Fusarium rt ging bo t ca Cephalosporium v th giai on ny thng c qui vo nm Cephalosporium. Bo t vch dy (hnh 6.9 E) hnh trn hoc hnh trng, vch dy, nm tn cng hoc chen gia cc si nm gi. Chng c th pht trin n hoc thnh chui, chng tch ra v mc cc ng mm nu bo t gp iu kin thun li, Hu bo t hay bo t vch dy rt bn v tn ti c lp trong thi gian di.
Figure 6.9
Hnh 6.9. A, Fusarium udum gy bnh ho l (trn u sn Cajanus cajan); B, cung sinh bo t v bo
80
t nh ln; C, i bo t nh (macroconidia); D, tiu bo t nh (microconidia); E, bo t vch dy (hu bo t)(Chlamydospore)(Sharma, 1998) Lp Coelomycetes c tnh chung 1. 2. 3. 4. 5. Nhm ny k sinh v hoi sinh trn thc vt c mch trn cn. Mt s k sinh bc hai trn nm khc. Tn l th qu tht, h si nm c vch ngn. Khng c t bo chi. Bo t v cung bo t sp xp trn ti bo t hoc cm cung bo t. Ti bo t b mt hay nm su bn trong, hnh cu, tri phng hoc hnh da, mt vch to nhng t bo cng ng knh. 6. Cm cung bo t nm bn trong cht nn thiu phn bn v trn vch (Sutton,1973) 7. Bo t n bo, rng sm, trong sut hoc c sc t t bo. Phn loi Sutton (1973) ngh Coelomycetes vo 2 b: 1. Melanconiales: th qu kiu cm cung bo t. 2. Sphaeropsidales: th qu kiu ti bo t phn. c im b Melanconiales 1. 2. 3. 4. Cc c th k sinh hoc hoi sinh trn thc vt Th qu kiu cm cung bo t. M cht nn gii hn c s ca th qu. Th qu di lp cutin, di biu b, hoc di chu b v v ra bi s rn nt ca m vt ch.
Melanconiales c mt h Melaconiaceae (Sutton, 1973) vi ging Colletotrichum c m t y. c im b Sphaeropsidales 1. 2. 3. 4. Tnh cht ch yu l th qu dng ti bo t phn. C th c kch thc hin vi k sinh hoc hoi sinh. Mng bao quanh th qu l nhu m gi. Bo t nh dng t cc vch ngn.
6.6 Ging COLLETOTRICHUM

c im N c m t c 11 loi (von Arx,1957; Sutton, 1973). Nhng Alexopoulos v Mims (1979) th xut trn 1000 loi hnh thc trong ging ny c m t trc y, phn ln chng trng tn. Theo kin gn nht ca Baxter v cng s (1985), Colletotrichum c gii thiu c 21 loi: C. coccodes, C. dematium, C. gloeosporioides, C. graminicola, C. falcatum v C. capsici. . . l nhng loi thng gy bnh thn th (anthracnose). Bnh thi ma y l bnh thng gp ma do C. falcatum, nm tn cng chnh vo thn v l (hnh 6.10), phin l tr nn nht hoc sm v r xung, thn c nhiu vt nt v lp mng si pht trin dc thn; S ho ch yu b mch, i khi vo tn trong rut v thn b thi, rt ngn ti cc t. Si nm
81 Ni sinh, si nm mnh, phn nhnh, khng mu, c vch ngn, si nm c ni bo v gian bo; Nhiu ht du c sn xut trong mi t bo ca h si nm; Khi chn si nm tr nn sm mu v bn xon li thnh dng cht nn nh di lp ngoi cng. Sinh sn Colletotrichum ch sinh sn v tnh bng bo t nh, bo t nh pht trin trn cung bo t trong dng th qu l cm cung bo t (hnh 6.11 A-C); Cm cung bo t c dng a phng, mt sau c cu trc phn mn, mi cm cung bo t gm lp cht nn, b mt sn sinh cung bo t trong sut (hnh 6.11.C). Cung bo t khng c vch ngn ko di n bo, dng lim, cong, bo t trong sut. Cng vi bo t v cung bo t l cc lng cng trn mi cm cung bo t, lng di cng, thun nhn, khng phn nhnh v a bo cu trc nh t cng (hnh 6.11.A-B) v Frost (1964) m t mt vi loi ca Colletotrichum c hoc khng c lng cng c th c kim sot bi s thay i m.
Figure 6.10
Hnh 6.10. Colletotrichum falcatum. A, cm bo t trong thn; B, cm bo t trn l (Sharma, 1998) S hnh thnh mt s ln ca bo t gy nt gy trn biu b vt ch, gp iu kin thun li, mi bo t mc t mt n nhiu ng mm hnh thnh h si nm (Hnh 6.11.D); a bm l dng ca Colletotrichum trong nui cy (Sutton, 1962, 1968) Si nm gi i khi hnh thnh vch dy, mu nu sm, hnh cu hoc khng u gi l hu bo t (Chlamydospores), N c th tn cng hoc chen gia si nm v tn ti trong thi gian di v khi tch ra chng cng mc mm hnh thnh si nm mi. Theo von Arx (1957) Sclerotia cng l mt dng c bit ca Colletotrichum,
82
Glomerella tucumanensis l mt nm ti (Ascomyceteous) c coi nh giai on hon chnh ca Colletotrichum falcatum.
Figure 6.11
Hnh 6.11. Cm bo t ca Colletotrichum lindemuthianum; B, cm bo t ca C. faltacum; C, cung bo t v bo t nh ca C. graminicola; D, bo t ny mm (Sharma, 1998)
Chng 7
Vai tr hu dng ca nm trong ch bin thc phm

1
7.1 GII THIU

S pht trin v hot ng trao i cht ca nm (nm men v nm mc) trong thc phm c th c nhng nh hng khc nhau. Mt khc hot ng ca nm c khai thc bi con ngi cho muc ch sn xut v ch bin thc phm. Thc tin thu c qu th nm rm cng nh s ng dng nm mc chun b thc phm ln men tri qua nhiu th k. Gn y l nm si cng nh nm men c nui cy thu c thc phm dinh dng giu m cho con ngi v vt nui. Nm ng vai tr quan trng trong s ln men cng nghip to ra nhiu loi phn ho t (enzim) v cc cht hu c khc. Nhiu trong s ny c ng dng nh thnh phn ca thc n. Gn y nht, k thut ti t hp DNA tr nn ph bin b sung nhng c tnh ca nm. Mt vi ng dng cho cng ngh thc phm s c cp.
7.2 Thc phm ln men bi nm

Ln men l mt trong nhng k thut lu i nht ca s ch bin thc phm v mang tm quan trng kinh t to ln. S xut hin, qui trnh ch bin, v s dng ca thc phm ln men c vit nhiu (Campbell-platt 1987, Steinkraus 1997). Mt vi sn phm ln men (ph-mat, bia, ru, nc tng) c kinh nghim sn xut mt lng ln, vi vic s dng ging chng u vit, mt khc nhiu thc phm ln men hy cn c sn xut s dng k thut truyn thng lu i di nhng iu kin n gin hoc ngay c nguyn s. Do nhiu nguyn nhn, ca nn kinh t v tnh cht yu cu ca sn phm, hu ht vic ln men thc phm khng th tin hnh mt cch li nhun di iu kin v trng. Thc phm ln men v th c th cha ng nhiu vi khun, nm men v nm mc, xut x t vt liu th, ging chng, s nhim trong tin trnh. Bng 7.1. Thc phm (chn lc) ln men bi ging hn hp (nm mc, nm men, vi khun)
Nm mc
Nm men
Vi khun
C cht
sn phm
s dng
ngun gc
continued on next page
1 This
83
84 Actinomucor elegans Amylomyces rouxii
CHNG 7. VAI TR HU DNG CA NM TRONG CH BIN THC PHM u h Endomyces spp. Hyphopichia spp. Zyg. rouxiiTorulopsis spp. Zyg. rouxii, Zyg. soyzae, Hansennula spp., Torulopsis spp., Candida spp. Pediococcus go (khng spp.Enterococcus nu) spp. Tetragenococcus u nnh halophila; + go/la Ent. faecalis mch Lactobacillus u nnh delbrueckii, + la m + Tet. mui halophilaPed. damnosus go nu chao Ragi thc m phm Trung Quc, VN Phng ng
ging chng lm ru
Aspergillus oryzaeA. soyae A. oryzaeA. soyae group
Miso
gia v
Phng ng Phng ng
Nc tng
gia v
A. oryzae
Hans. Leuc. anomala,Sacch.mesenteroiCerevisae des var sak, (sak) Lb. sak
Sak
ru
Nht
Monascus purpureusM. rubber,M. pilosius Penicillum roqueforti P. camemberti Yarrowia lipolytica Leuconostoc spp.
go nu
Angkak(go ln men vi mc ) Ph-mt kiu roquefort Phmt kiu Camembert Salani
cht to mu, gia v,thnh phn b dng thc phm m, gia v thc phm m, gia v thc m phm
Trung Quc, Nht
Bnh sa (hoa sa p)
Php
Candida BrevibacteriumBnh sa spp.Kluyveromyces linensLc. (hoa sa p) spp.Torulopsis lactis ssp spp cremoris Mirococcus tht spp.Staphylococcus xch) spp.Pediococcus spp.Lactobacillus spp. (xc
Php
P. nalgiovenseP. chrysogenum
u Chu A
continued on next page
85 Rh. oligoporusRh. chinensis,Rh. oryzae,Mucor indicus Trichosporon Klebsiella u nnh beigelii,Clavispora pneumolusitaniniaeEnase,Yar. terobaclipolytica ter cloacae,Lactobacillus spp.
Table 7.1
Tempeh
thc phm m,thc n nhanh
Indonesisa
thc phm ln men nm a ging chng, s kch thch t cht chuyn ho bi s phn gii tng h ca c cht hoc bi s phng thch ca cc sn phm phn gii; hoc s c ch bi s cnh tranh s to thnh ca cht khng sinh hoc cht trao i khng vi sinh l cc yu t quan trng ca s cn bng ca cc qun th vi sinh vt (Nout, 1995). Theo tnh t nhin vt l ca c cht, s ln men c th c phn bit thnh ln men b su hay ln men chm (liquid fermentation) v ln men b mt (solid-state fermentation). Trong ln men b su, cht lng phc v nh mt mi trng cho s phn b ng nht ca vi sinh vt v cho s truyn nhit v truyn khi. Ln men b su c s dng cho vic ch bin nc gii kht v nc st (sauces). Bng 1 lit k vi thc phm ln men trong nm ng mt vai tr thit yu. Thm vo , vt liu th ca chng, vi sinh vt ln men i din, kiu ca h thng ln men (b su hoc b mt) v lin h n nhm h vi sinh vt cho s ln men thnh cng cng c lit k. vng n i, tht lm chn bi nm mc v ph-mt l chim u th bi Aspergillus v Penicillium spp. Nm men ng vai tr trong cc sn phm nng v nc gii kht c cn. nhng vng nhit i v cn nhit i, thc phm ln men nm chim u th ng v ng nam chu . Rhizopus, Amylomyces, Mucor, Neurospora, and Monascus spp. thng c tm thy nh h vi sinh vt chc nng. Cc sn phm ln men bi nm men t vng nhit i bao gm thc n nhanh v nc gii kht c cn. Sau y l mt vi thc phm ln men chn lc.
7.2.1 Ru
S a dng ca ru khng ch v nhiu hn 5.000 loi nho (Vitis vinifera) m cn c bit n iu kin trng trt (a im, t, kh hu) v iu kin ln men. Nguyn tc c bn ca lm ru c tm tt hnh 7.1 Nho phi l khng b ln mc, tr phi s ch bin ca ru ngt Sauterne loi ny i hi mc Botrytis cinerea. Ru th thng thng ln men trn lp v. Thng th SO2 c thm vo 100-150mg/lt c ch s pht trin ca nm men biu sinh (Candida, Hanseniaspora, Kloeckera, Pichia, Rhodotorula, Saccharomyces, Torulopsis spp.) iu ny s thc y s chim u th tt ca nm men ru chn lc (Saccharomyces cerevisae thng ng hnh vi Torulopsis stellata) (cho vo sp x 106/ml dch nho. Lu rng nm men to mi (hng v) l cn bng vi nm men chc nng cho s ln men ru. Khi tt c ng ln men cn kit th s ln men ru dng li v nm men c ly ra ngn cn s mt mi t s t phn gii ca nm men. ru axit cao, vi khun axit lactic (Oenococcus oeni) c chng bin i axit malic thnh axit lactic, v th cho ra ru v ngt hn.
7.2.2 Ph-mt Camembert

y l mt ca nhiu ph-mt b mt chn. C ngun gc t Normandy, Php ln u tin n c lm bi Marie Harel vo nm 1791. Nm 1890, M. Ridel pht trin hp g ni ting to iu kin xut khu vi lng ln. Nguyn tc c bn ca sn xut ph-mt Camembert c vit ra nh hnh 7.2. Sau khi sa ng cn rt non (p thnh bnh sa), n c chng ging bi bo t ca Penicillium camembert bng cch phun nh sng m. Sau khi ngm trong nc mui v a vo iu kin thch hp, s pht trin ca mc bt
CHNG 7. VAI TR HU DNG CA NM TRONG CH BIN THC PHM u ti b mt ph-mt trong sut giai on . V cng ca ph-mt Camembert th mng v trng. Cc dng mc chng khc nhau c mu sc trong khong t xm lc nht ti trng tinh. Bn trong ca ph-mt phi l vng nht vi trung tm mu trng rn chc. Trong qu trnh chn, cc phn ho t (enzim) thu phn m v cht bo ca P. camemberti khuych tn vo trong ph-mt. 86
Figure 7.1
Hnh 7.1. Qui trnh lm ru nho Nhng tnh xp mm l v pH tng ln gy ra bi s phng thch ca NH3, nm men, c bit l Debaryomyces hansenii, thng hin din trong ph-mt v cng gp phn lm tng pH bi v s tiu th ca axt lactic. Cc phn ng thu phn m v hot ng ca enzim aminotransferase ng gp phn ln vo s pht trin hng v. Thng thng sn phm c tiu th sau 3-5 tun. Kim nh ging thun chng, tt c cc dng P. camemberti c bit l c kh nng sn xut ra c t nm (mycotoxin) axit
87 cyclopiazonic (CPA). Cht ny xem ra mang c tnh bn bi v cc dng thu thp nui cy lu khng mt i kh nng to ra CPA. Nhiu n lc tin hnh t c cc dng t bin CPA m tnh v cc ging chng c tuyn chn theo tiu chun ny. C hi ri ro ca vic ng c l rt nh. Ch mc rt thp ca CPA c th c pht hin ra ph-mt Camembert. iu ny c gii thch bi s khng n nh v ho hc ca n trong s hin din ca cht amines v s khuych tn ngho nn t v cng vo bn trong. Hn na, CPA th kh c to ra ti nhit bo qun <15 C.
Hnh 7.2. Quy trnh ch bin ph-mt Camembert
7.2.3 Chao (sufu)

Chao (sufu) cng c vit nh fu-ru, l do hoa sa u nnh ln men mc c mi kh nng (Su, 1986). Chao c sn xut ch yu Trung Quc cho tiu th ni a v xut khu, sn lng hng nm c tnh t nht 300 triu tn. Chao c tiu th nh mt cht gia v, v d vi cm im tm sng hoc bnh m hp. Chao c lm t khi u h (thu c t kt ta sa u nnh). u h c dng lm chao cng hn bi p hoc c s l hi nng nhanh, v tun t c chng ging trn b mt v m cao ti nhit 20-35 C, tu thuc vo a im v ma (trong nm). Cc loi nm mc tham gia ch yu l Actinomucor v Mucor spp. Nhng nh my sn xut ln c s dng ging thun chng, chng hn nh Actinomucor elegans, ngc li ti cc c s sn xut nh, s dng rm ln mc chng ln khi u h vi mt h vi sinh vt hn hp ca nm mc v vi khun. Actinomucor elegans khng pht trin tt nhit
CHNG 7. VAI TR HU DNG CA NM TRONG CH BIN THC PHM vt qu 25 C, cc mc khc v d nh Mucor hiemalis hoc Rhizopus chinensis c s dng nhit cao hn. Sau vi ngy khi u h c bao ph vi mt lp dy ca si nm nh bng gn. Sn phm trung gian c gi l pehtze-u h ln mc. Pehtze tr thnh mt ngun enzim thu phn m, v m u nnh phn ln c phn ct. Bc k tip mui v ngm nc mui vi mc ch bo qun sn phm trong khi cho php s trng thnh v mt enzim m cui cng s cho ra cu trc mm, mi v v nng theo yu cu. S l u tin vi mui (ht, bt) vi mc ch l lm tng ln nhanh chng hm lng mui khong 15%. Bc k tip cc pehtze mui c t vo cc keo/l v bm y vi nc mui hoc hn hp nc chan chao chn . Thnh phn ca nc mui chao chn nh hng mnh m n tnh cht ca sn phm cui cng. N lun cha 10-12% NaCl, v ru go 10% hm lng ethanol. Sn phm rt ni ting l chao , trong nc ngm chao ny c cha ang-kak (xem Bng 8.1). Ang-kak khng ch cho sc t cam v m cn cha ng nhiu phn ho t (enzim) tch cc gp phn vo s phn ct v to hng v ca chao. Chao c th c s hin din ca vi khun Tetragenococcus halophila. Sau nhiu thng chn, cc keo c lm sch bn ngoi, dn nhn v phn phi cho vic tiu th. 88
7.2.4 Tempeh
Tempeh c ngun gc t Java, Indonesia nhng cng c ph bin H Lan v n t c th trng u v c. Tempeh l mt bnh c th thi (ct) c, thu c bi phng tiu th ng k M, chu A php ln men b mt vi nm ca nhng ht u, ng cc hoc vt liu ph hp khc sau khi ngm ri nu. C cht c dng nhiu v ph bin nht l u nnh (Nout and Rombouts, 1990; Wood, 1998). Tempeh cung cp mt ngun m thc vt r tin giu dinh dng d tiu ho v an ton. N thng c n ch khi sau khi nu hoc rn/chin trong du. Qui trnh ch bin truyn thng c trnh by Hnh 7.3. u nnh c ngm v tch v, hoc tch v kh trc ri ngm (c gii ho nh H lan). Trong qu trnh ngm s ln men lactic t nhin din ra lm h thp pH ca u, lm cho chng ph hp hn cho s pht trin ca nm mc v chng li vi khun gy h hng v cc mm bnh. Sau khi nu si v lm ngui, u c chng ging truyn thng usar (Rhizopus spp. hin din trn l cy Hisbiscus, hoc ging thun chng trong bt go hoc bt khoai m. Mc chc nng chnh yu l Rhizopus oligosporus v R.oryzae, chng ny mm nhanh 37 C v s pht trin si nm nhanh ca chng m bo s chim u th v ln t cc dng tm nhim khc nh Aspergillus spp. trong vng 30 gi cc u ri rc c kt cht li nhau to thnh khi rn. R.oligosporus c kho st l thm nhp khong 2mm vo su bn trong u nu. Hot ng enzim ca Rhizopus spp. bao gm enzim thu phn protein, enzim thu phn cht bo, enzim thu phn cc hp cht cc-bon v phosphatase. Nh hot ng ca cc enzim ny, mt phn cc c cht mch di b thu phn thc y cho s tiu ho d dng. Qui trnh mi ci tin c bo co bao gm tin trnh bn lin tc cho ln men tempeh vi nhit c iu khin trong thng ln men xoay vng.
89
Figure 7.2
Hnh 7.3. Quy trnh ch bin tempeh
7.2.5 Nc tng u nnh

Nc tng u nnh ln men c ngun gc t Trung Quc. Nht c khong 3600 cng ty, trong c 5 cng ty ln nht sn xut 1,2 triu tn chim khong 1/2 sn lng hng nm . Nc tng c nhiu dng khc nhau, nhng Koikuchi-Shoyu l i din tt nht ca nc tng ln men (Yokotsuka and Sasaki 1998). N l mt cht lng nu xm trong vi thnh phn xp x: 22 B, 17% NaCl, 1,6% m tng s, 1% m formol, 3% ng kh, 2,3% ru alcohol v pH 4.7. V c bn ca qu trnh ch bin (Hnh 7.4) bao gm 3 giai on: lm koji (u nnh ln mc), ln men trong nc mui v lc trong. Lm koji l qu trnh ln men b mt ca hn hp u nnh nu/hp v bt m rang. N c chng ging vi bo t Aspergillus oryzae hoc A.sojae v c 25 C trong 2-3 ngy thu c s pht trin dy c v s to bo t xanh vng ch ra rng mc cao ca enzim c to ra. Nhng enzim ny bao gm peptidases, proteinases, glutaminase, amylase, pectinases v cellulases cn thit cho s thu phn tng phn ca m v cc hp cht carbon ca vt liu th.
CHNG 7. VAI TR HU DNG CA NM TRONG CH BIN THC PHM Giai on 2 ln men trong nc mui din ra trong m nc mui 22-25% NaCl vi sinh vt gy h hng khng th pht trin nhng s phn ct enzim vn din ra. Tuy nhin, trong 2 thng u 1520 C, vi khun axt lactic (Tetragenococcus halophila) v nhng thng tip theo 30 C, nm men a mui (Zygosaccharomyces rouxii) s pht trin, vo khong thi gian chn, nhng sn phm trao i cht ca chng thm vo thnh phn hng v thc cht thm ngon ca nc tng. 90
Figure 7.3
Hnh 7.4. Quy trnh lm Nc tng u nnh ln men
7.3 Sinh khi nm v s chuyn ho sinh hc

7.3.1 Nm rm
Nm rm (n c) rt phong ph v a dng, nhng ch mt vi loi c pht trin v thng mi ho. Vic sn xut nm rm thng mi c c tnh 1-2 triu tn. Agaricus spp. chim phn ln trong tng s nm sn xut. A. bisporus trng l c bit nhiu nht, nhng A. bitorquis chng chu vi-rt v A. bisporus m nu ang c tng ln s ch . Sau khi si nm chim ng c cht, s to tri c khi
91 ng bi s thay i iu kin mi trng (thong kh, m , nhit ). Qu th c to thnh trong s ca s ny mm vt qu khong 1-2 tun. Sau 4-5 t thu hoch, sn lng gim xung v vng i bt u li vi c cht ti mi. C cht c dc s dng nh thnh phn thc n gia sc giu protein. Qu th ln k l ca Basidiomycete, Ustilago maydis c thu thp v tiu th nh nm rm, v ph bin chu M Latinh.
7.3.2 m n bo (nm men, protein nm)

Nhiu dng nm men (Candida utilis, C. tropicalis, Yarrowia lipolytica, Kluyveromyces lactis) c th pht trin vi sn lng t bo cao trn c cht l sn phm ph cng nghip, v d nh nc nh thanh (whey) t lm ph-mat, nc thi t cng nghip tinh bt khoai ty, ru sulfite g, v cn b ca hp cht cac-bon. Mc d nhiu qui trnh cng nghip c cp bng, th nhng m n bo thu c hin ti l khng th cnh tranh vi cc m khc (v d nh u nnh). Si nm ca Penicillium chrysogenum v Aspergillus niger c to ra mt lng ln nh mt sn phm ph ca s ln men sn xut khng sinh, phn ho t (enzim) axt hu c... khun ty ca nm khng c l mt thnh phn thc n l tng bi v n c hm lng m th cao (xp x 12% trn trng lng ti). Khun ty c s dng nh thnh phn thc n gia sc. Ging nh vy, si nm ca dng Fusarium venenatum c s dng cho sn xut cng nghip protein cho ngnh dt may (Wiebe et al., 1996). Sn phm tm thy cho s ng dng nh tht thay th trong bnh nhn to thm, sp...S dng m vi sinh cho s tiu th ca con ngi v vt nui l b gii hn bi hm lng axt nucleic ca n. T chc WHO khuyn co mc ti a ca axt nucleic l 2% trong thc phm. Trong khi m vi khun cha mc kh cao axt nucleic, m nm (F. venenatum) cha khong 6-13%.
7.3.3 Lm giu thm m cho thc phm tinh bt v thc n gia sc

Do c kh nng phn ct vt cht cc-bon, nm l hu dng trong vic nng cao gi tr dinh dng ca cc sn phm ph nng-cng nghip sau ch bin, chng hn nh tinh bt cha trong cht cn b khoai lang (Yang et al., 1993) hoc b ma ng cellulose (Moo-Young et al., 1992). Cho mc ch thm vo cc ngun m r tin nh (NH4)2SO4 v urea c th bin i thnh m v th lm giu dinh dng thm cho thc n. Sn phm cui cng (sau vi ngy ln men) c th cha dng xp x 30% m th trn trng lng kh, khi s dng nm Aspergillus niger, Rhizopus spp. v Neurospora sitophila.
7.3.4 Thnh phn thc n v gia v c ngun gc t nm mc

a. Axt hu c Axt hu c c to ra bi s ln men c th phn thnh hai nhm: (i) c to ra thng qua l trnh axt carboxylic, (ii) c to ra trc tip t ng glucose. nhm (i), axt citric l axt hu c quan trng nht c to ra bi s ln men. Sn lng hng nm c tnh khong 400.000 tn c lm ch yu vi Aspergillus niger nhng Yarrowia lipolytica cng c s dng. H thng nui cy b mt cng nh b su (chm) c s dng cho s bin i nhng ngun carbon r tin (mt ng) v n-alkanes. Axt citric c s dng rng ri trong cng nghip thc phm, cht hng v. . . Trong nhm (ii) axt gluconic (50.000 tn/nm) c lm ra ch yu vi vi khun, nhng A. niger v A. foetidus c s dng trong cc qui trnh nui cy b mt v nui cy chm. Axt lactic (30.000 tn/nm) c lm ra ch yu vi vi khun lactic, nhng cng vi Rhizopus oryzae v c nhng ng dng rng ri trong cng nghip thc phm nh cht axt, nhn t bo qun, bt nng . . . (Mattey, 1992). b. Cht bo (lipids) Hm lng cht bo ca nm i khi c th cao n 60-80% sinh khi kh. Tuy nhin, cht du v cht bo ng v thc vt l r tin hn sn xut, v th ch nhng sn phm c sn th vic ln men l s quan tm v mt kinh t. Ni ring, kh nng tch lu axt bo khng no t nm l l th v quan im dinh dng. Nhiu nm mc c s dng thng mi to ra axt linolenic, trn ngun m v cac-bon r
CHNG 7. VAI TR HU DNG CA NM TRONG CH BIN THC PHM tin (b nho, tinh bt, mt r ng), Mucor javanicus v M. rouxii c th pht trin trong mi trng nui cy chm sn lng axt linolenic 0,33g/l. Nhng trng hp nh th hm lng lipid l 7-11% ca sinh khi kh v axt linolenic chim 17-37% trng lng lipid (Lindberg and Hansson, 1991). c. Phn ho t (enzim) Trong s nhng phn ho t enzim c to ra bi s ln men, chim a s l enzim thu phn m (proteases) v enzim thu phn hp cht cac-bon (carbonhydrases). Enzime thu phn m thu c t Aspergillus oryzae, Penicillium roquefortii v Mucor spp. c ng dng trong cht ty ra, v trong ch bin thc phm nh lm nhanh chn ph-mat, lm bnh m, s lm mm tht. Enzim thu phn hp cht carbon bao gm enzim thu phn tinh bt (-amylase, glucoamylase) c to ra bi Aspergillus oryzae v A. niger v c ng dng trong lm bnh m, bia ru, v bnh ko. Nhng carbonhydrases khc l cellulase c tit ra bi A. niger, Penicillium spp. v Trichoderma reesei; enzim thu phn pectin (pectinases) c to ra bi Aspergillus spp.; v glucanase c to ra bi A. niger, Penicillium spp. nhng enzim ny c ng dng thc y s tiu ho ca thc phm c si, s tinh lc nc tri cy v bia. . . Nhng enzim quan trng khc bao gm enzim thu phn cht bo (lipases) c to ra bi Mucor spp. v A. niger ng dng cho s pht trin hng v cc sn phm t sa. Glucose oxidase c to ra bi A. niger c nhiu ng dng trong thc phm v y hc. . . Mt khc, c s phong ph ca nhng c cht c th c to ra bi nm, bao gm axt amin, cc hp cht ng, cc sinh t vitamins, phm mu v cc thnh phn hng v. . . 92
INDEX
93
Index of Keywords and Terms

Keywords are listed by the section with that keyword (page numbers are in parentheses). Keywords do not necessarily appear in the text of the page. They are merely associated with that section. Ex. apples, 1.1 (1) Terms are referenced by the page they appear on. Ex. apples, 1
N Ngnh ph Nm Bt Ton, 4(31), 6(69)

Ngnh ph nm m, 5(45) Nm roi, nm trng, 2(15) Nm tip hp, 3(17)
V Vai tr, hu dng, nm, ch bin thc phm,

7(83)
i cng, nm mc, 1(1)
94
ATTRIBUTIONS
Attributions
Collection: Gio trnh mn Nm hc Edited by: Nguyn Vn Thnh, Cao Ngc ip, Nguyn Vn B URL: http://cnx.org/content/col10923/1.1/ License: http://creativecommons.org/licenses/by/3.0/ Module: "i cng v nm mc" By: Nguyn Vn B URL: http://cnx.org/content/m30247/1.1/ Pages: 1-13 Copyright: Nguyn Vn B License: http://creativecommons.org/licenses/by/3.0/ Module: "Nm roi - nm trng (ngnh ph Chytridiomycotina)" By: Cao Ngc ip, Nguyn Vn B URL: http://cnx.org/content/m30105/1.1/ Page: 15 Copyright: Cao Ngc ip, Nguyn Vn B License: http://creativecommons.org/licenses/by/3.0/ Module: "Ngnh ph Nm tip hp (Zygomycotina = lp Zygomycetes)" By: Cao Ngc ip, Nguyn Vn B URL: http://cnx.org/content/m30255/1.1/ Pages: 17-29 Copyright: Cao Ngc ip, Nguyn Vn B License: http://creativecommons.org/licenses/by/3.0/ Module: "Ngnh ph Nm Bt Ton (Deuteromycotina = lp Deuteromycetes)" By: Cao Ngc ip URL: http://cnx.org/content/m30258/1.1/ Pages: 31-44 Copyright: Cao Ngc ip License: http://creativecommons.org/licenses/by/3.0/ Module: "Ngnh ph nm m (Basidiomycotina = Lp Basidiomycetes)" By: Cao Ngc ip URL: http://cnx.org/content/m30252/1.1/ Pages: 45-67 Copyright: Cao Ngc ip License: http://creativecommons.org/licenses/by/3.0/ Module: "Ngnh ph Nm Bt Ton (Deuteromycotina = lp Deuteromycetes)" By: Cao Ngc ip URL: http://cnx.org/content/m30898/1.1/ Pages: 69-82 Copyright: Cao Ngc ip License: http://creativecommons.org/licenses/by/3.0/
ATTRIBUTIONS Module: "Vai tr hu dng ca nm trong ch bin thc phm" By: Nguyn Vn Thnh URL: http://cnx.org/content/m30069/1.1/ Pages: 83-92 Copyright: Nguyn Vn Thnh License: http://creativecommons.org/licenses/by/3.0/
95
Gio trnh mn Nm hc Nhm mc ch cung cp thm nhng kin thc c bn sinh vin hc tp tt mn L thuyt NM HC, Gio trnh c son theo th t phn loi ca ngnh NM v c nhng v d c th nhng loi nm tiu biu ca tng ngnh ph (hay lp) trong m t tng i y nhng c im sinh hc ca mi nhm nm thng qua nhng dng khun ty, cng mang ti (bc) bo t, cc loi bo t, tm tt nhng vng i vi nhng c tnh sinh sn hu tnh.... tiu biu v nu ln nhng khc bit r rt gia cc ngnh ph (lp) sinh vin c th so snh v nhn bit s khc nhau gia cc ging trong mt h hay gia cc lp trong ngnh. Gio trnh NM HC c son tng i chi tit sinh vin i hc v c hc vin Cao hc cc ngnh hc lin quan tham kho nhng thng tin cn thit n ngnh hc. Chng ti mong rng gio trnh s ng gp c nhng thng tin c th v mn hc ny v chc chn gio trnh s cn nhng thiu st, chng ti hy vng cc ng nghip gp cho gio trnh ngy cng hon thin hn. Ngoi ra, c nhng t c dch t cc t in Sinh hc Anh - Vit s gy s ng nhn, chng ti ch thch phn ting Anh. TM. Nhm bin son PGs. Ts. Cao Ngc ip
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Gio trnh mn Nm hc

By: Nguyn Vn Thnh Cao Ngc ip Nguyn Vn B

By: Nguyn Vn Thnh Cao Ngc ip Nguyn Vn B

Online: < http://cnx.org/content/col10923/1.1/ >

1.1 Hnh dng, kch thc, cu to ca nm mc

content is available online at <http://cnx.org/content/m30247/1.1/>.

Hnh 1.1 Si nm v cu to vch t bo si nm (theo Samson v ctv., 1995)

1.3 Dinh dng v tng trng ca nm mc

1.4.1 Sinh sn v tnh

Hnh 1.12 Bo t t (theo Samson v ctv. 1995)

1.4.2 Sinh sn hu tnh

Hnh 1.13. Bo t nang Saccharomyces cerevisiae (theo Samson v ctv. 1995)

1.6 Phn loi nm mc

Nm roi - nm trng (ngnh ph Chytridiomycotina)

content is available online at <http://cnx.org/content/m30105/1.1/>.

CHNG 2. NM ROI - NM TRNG (NGNH PH CHYTRIDIOMYCOTINA)

Ngnh ph Nm tip hp (Zygomycotina = lp Zygomycetes)

3.2 Phn loi

content is available online at <http://cnx.org/content/m30255/1.1/>.

CHNG 3. NGNH PH NM TIP HP (ZYGOMYCOTINA = LP ZYGOMYCETES)

3.2.1 Ging [Chi] Rhizopus

CHNG 3. NGNH PH NM TIP HP (ZYGOMYCOTINA = LP ZYGOMYCETES)

Ti bo t cng bo tcung,li khun ngang

CHNG 3. NGNH PH NM TIP HP (ZYGOMYCOTINA = LP ZYGOMYCETES)

CHNG 3. NGNH PH NM TIP HP (ZYGOMYCOTINA = LP ZYGOMYCETES)

Hnh 3.5. Sinh sn hu tnh vi trng hp ng tn nm Rhizopus sexualis (Sharma, 1998)

3.2.2 Chi Mucor

CHNG 3. NGNH PH NM TIP HP (ZYGOMYCOTINA = LP ZYGOMYCETES)

CHNG 3. NGNH PH NM TIP HP (ZYGOMYCOTINA = LP ZYGOMYCETES)

3.2.3 Tm quan trng ca b Mucorales

CHNG 3. NGNH PH NM TIP HP (ZYGOMYCOTINA = LP ZYGOMYCETES)

Ngnh ph Nm Bt Ton (Deuteromycotina = lp Deuteromycetes)

4.1 Gi thiu chung

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CHNG 4. NGNH PH NM BT TON (DEUTEROMYCOTINA = LP DEUTEROMYCETES)

Hnh 6.1. Mt b si bo t (synnema) ca Arthrobotryum (Sharma, 1998)

Hnh 6.2. Nm Collectrichum [A], Fusarium [B], Endothia [C](Sharma, 1998)

4.1.2 Tm quan trng

CHNG 4. NGNH PH NM BT TON (DEUTEROMYCOTINA = LP DEUTEROMYCETES)

4.2 Phn loi

CHNG 4. NGNH PH NM BT TON (DEUTEROMYCOTINA = LP DEUTEROMYCETES)

CHNG 4. NGNH PH NM BT TON (DEUTEROMYCOTINA = LP DEUTEROMYCETES)

4.3 Ging CURCULARIA

Hnh 6.7. Cung bo t v bo t ca Curvularia lunata (Sharma, 1998)

4.4 Ging PYRICULARIA

CHNG 4. NGNH PH NM BT TON (DEUTEROMYCOTINA = LP DEUTEROMYCETES)

4.5 Ging FUSARIUM

CHNG 4. NGNH PH NM BT TON (DEUTEROMYCOTINA = LP DEUTEROMYCETES)

4.6 Ging COLLETOTRICHUM

CHNG 4. NGNH PH NM BT TON (DEUTEROMYCOTINA = LP DEUTEROMYCETES)

Glomerella tucumanensis l mt nm ti (Ascomyceteous) c coi nh giai on hon chnh ca Colletotrichum falcatum.

Hnh 6.11. Cm bo t ca Colletotrichum lindemuthianum; B, cm bo t ca C. faltacum; C, cung bo t v bo t nh ca C. graminicola; D, bo t ny mm (Sharma, 1998)

Ngnh ph nm m (Basidiomycotina = Lp Basidiomycetes)

5.1 c tnh tng qut

5.2 Khun ty v hp nhn (nhn kp)

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CHNG 5. NGNH PH NM M (BASIDIOMYCOTINA = LP BASIDIOMYCETES)

t bo n nhn; m khng bao gi pht trin trn khun ty bc 1.

5.2.2 Khun ty bc 2 v nhn kp

CHNG 5. NGNH PH NM M (BASIDIOMYCOTINA = LP BASIDIOMYCETES)

5.3 To mu (Clamp connection)

Hnh 5.2. Qu trnh thnh lp mt MU trong mt khun ty (Sharma, 1998)

CHNG 5. NGNH PH NM M (BASIDIOMYCOTINA = LP BASIDIOMYCETES)

Hnh 5.3. Cc loi M (Sharma, 1998)