7Ï07ҲT LUҰ19Ă1 Ӭng dөQJFiFNӻ thuұWYjF{QJQJKӋ sinh hӑFÿӇ Jk\QKLӉPQKkQWҥRFiFORjLnҩm ngoҥi cӝng sinh OrQFk\WUӗng nhҵm hӛ trӧ WURQJF{QJWiFWUӗng rӯng ÿmÿѭӧc nhiӅXQKjNKRDKӑFNKDLWKiFWUrQWKӃ giӟ
Trang 175ѬӠ1*ĈҤI HӐ&%È&+.+2$
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PHҤ0+2¬1*3+,3+ӨNG
6¬1*/Ӑ&9¬*Æ<1+,ӈM MӜT SӔ /2¬,1ҨM
NGOҤI CӜNG SINH THU THҰP TӮ RӮ1*3+Ñ1*+Ӝ
ĈҪU NGUӖ1Ĉ$1+,075Ç1&Æ<7+Ð1*%$/ÈPinus kesiya
&KX\rQQJjQK&{QJQJKӋ Sinh hӑc 0mVӕ: 60420201
LUҰ19Ă17+Ҥ&6Ƭ
TP HӖ &+Ë 0,1+WKiQJQăP
Trang 31+,ӊ09Ө/8Ұ19Ă17+Ҥ&6Ƭ
&KX\rQQJjQK &{QJQJKӋ6LQKKӑF 0mVӕ : 60420201
7Ç1Ĉӄ 7¬,6jQJOӑFYjJk\QKLӉm mӝt sӕ ORjLQҩm ngoҥi cӝng sinh thu thұp tӯ rӯng
SKzQJKӝ ÿҫu nguӗQĈD1KLPWUrQFk\WK{QJEDOiPinus kesiya
NHIӊM VӨ 9¬1ӜI DUNG:
1 Thu thұSYjSKkQOұp quҧ thӇ FiFORjLQҩm ngoҥi cӝng sinh trong khu rӯQJWK{QJED
OiPinus kesiya thuӝc rӯQJSKzQJKӝ ÿҫu nguӗQĈD1KLP
2 Ĉӏnh danh FiFFKӫng nҩPSKkQOұSWKjQKF{QJEҵQJSKѭѫQJSKiSsinh hӑFSKkQWӱ YjFKӑQUDORjLQҩm tiӅPQăQJFKRP{KuQKJk\QKLӉm
3 ;k\GӵQJFk\SKiWVLQKORjLP{Wҧ KuQKWKiLÿӇ ÿӏQKGDQKFKtQK[iFÿӃQORjLÿӕi vӟi
2 chӫng nҩPÿmFKӑn
4 KhҧRViWVLQKOêÿLӅu kiӋQQX{LFҩy) 2 chӫng nҩPÿmFKӑQYj[k\GӵQJP{KuQKJk\
nhiӉm nҩPOrQFk\WK{QJFRQPinus kesiya
Trang 4LӠI CҦ0Ѫ1
Em xin gӱi lӡi cҧPѫQFKkQWKjQKÿӃn thҫy Phҥm NguyӉQĈӭF+RjQJÿmQKLӋt
WuQKJL~Sÿӥ, chӍ dҥy cho em nhӳng kiӃn thӭc, kinh nghiӋm, kӻ thuұWWURQJQJKLrQ
cӭXYjÿӏQKKѭӟng cho em trong suӕt thӡi gian thӵc hiӋQÿӅ WjL
Em gӱi lӡi cҧPѫQVkXVҳFÿӃQF{/r7Kӏ Thӫ\7LrQYuF{ÿmOX{QTXDQWkP
WKHRG}LYjJySêFKRHPUҩt nhiӅu trong suӕt thӡi gian thӵc hiӋn luұQYăQ
Em xin cҧPѫQWҩt cҧ Thҫ\&{WURQJEӝ P{Q&{QJQJKӋ sinh hӑc ± Khoa Kӻ
thuұW+yDKӑFÿmWUX\ӅQÿҥt cho em nhӳng kiӃn thӭc khoa hӑc TXêEiX trong suӕt
nhӳng QăPHPQJӗLWUrQJKӃ giҧQJÿѭӡng
Xin gӱi lӡi cҧPѫQÿӃQFiFDQKFKӏFiFEҥQYjFiF em tҥi ViӋn NҩPYj&{QJ
nghӋ sinh hӑc ÿm WҥR ÿLӅu kiӋn vӅ Fѫ Vӣ vұt chҩt, thiӃt bӏ kӻ thuұW FNJQJ QKѭ OX{Q
nhiӋWWuQKJySêYjFKLDVҿ kinh nghiӋPWURQJTXiWUuQKPuQKWKӵc hiӋn luұQYăQӣ
ÿk\
CuӕLFQJFRQPXӕn gӱi lӡi cҧPѫQÿӃQJLDÿuQKYuÿmOX{QErQFҥQKOX{QWLQ
WѭӣQJYjOjQJXӗQÿӝng lӵc lӟQÿӇ FRQFyWKӇ KRjQWKjQKOXұQYăQQj\
Trang 57Ï07ҲT LUҰ19Ă1
Ӭng dөQJFiFNӻ thuұWYjF{QJQJKӋ sinh hӑFÿӇ Jk\QKLӉPQKkQWҥRFiFORjLnҩm ngoҥi cӝng sinh OrQFk\WUӗng nhҵm hӛ trӧ WURQJF{QJWiFWUӗng rӯng ÿmÿѭӧc nhiӅXQKjNKRDKӑFNKDLWKiFWUrQWKӃ giӟL7URQJQJKLrQFӭXQj\FK~QJW{LÿmWKXthұSYjSKkQOұp quҧ thӇ nhӳQJORjLQҩm ngoҥi cӝQJVLQKÿѭӧFWuPWKҩy trong khu
vӵc trӗQJWK{QJED Oi Pinus kesiya thuӝc rӯQJSKzQJ Kӝ ÿҫu nguӗQĈD1KLPĈj Lҥt), chӑQ UD KDL ORjL Qҩm tiӅP QăQJ Oj Suillus luteus Yj Scleroderma
sinnamariense, tiӃQ KjQK NKҧR ViW ÿLӅu kiӋQ QX{L Fҩ\ Yj [k\ Gӵng P{ KuQK Jk\
nhiӉPQKkQWҥROrQFk\WK{QJFRQPinus kesiya 7 tuҫn tuәi KӃt quҧ QJKLrQFӭu cho
thҩy nhӳng ҧQKKѭӣQJWtFKFӵc tӯ KDLORjLQҩPQj\OrQFiFGӳ liӋXWăQJWUѭӣng cӫa Fk\FKӫ Sau 12 tuҫQJk\QKLӉm nҩPOrQFk\WK{QJFRQ, tҩt cҧ nhӳQJFk\ÿѭӧFJk\nhiӉPÿӅXÿmxuҩt hiӋn cҩXWU~F[kPQKLӉm cӫa nҩPWUrQFiFÿҫXFKySUӉ KӃt quҧ Qj\OjWLӅQÿӅ cho nhӳQJQJKLrQFӭXVkXKѫQWLӃQÿӃn viӋc ӭng dөng kӻ thuұWJk\nhiӉm nҩm ngoҥi cӝQJVLQKOrQFk\WK{QJFRQSKөc vө FKRF{QJWiFWUӗng rӯng
SUMMARY
There were many experiments using biological technique for artificial innoculation of a host plant with specific ectomycorrhizal fungi (EMF) in plantation strategies over the world In this research, we collected and isolated the
ectomycorrhizal sporocaps found in plots of natural forest of Pinus kesiya in Da Nhim Forest Enterprise (Da Lat), then chose two species ± Suillus luteus and
Scleroderma sinnamariense for innoculation to 7-week-old P kesiya seedlings
Identification of these two species were based on morphology and phylogenetic
analysis In this research, we also report the optimum culture conditions of S luteus and S sinnamariense The results show that both of these EMF species had positive
effects on plants productivity comparing with non-innoculated ones After 12 weeks
of innoculation with S luteus and S sinnamariense, all the roots of inoculated
seedlings appeared structures of EMF colonisation This results will be the premise for EMF application in platation strategies in the future
Trang 6LӠ,&$0Ĉ2$1
7{LWrQ3KҥP+RjQJ3KL3KөQJKӑFYLrQFDR KӑFQJjQK&{QJQJKӋ6LQKKӑFNKyDÿӧWNKRD.ӻWKXұW+yDKӑFWUѭӡQJÿҥLKӑF%iFKNKRDWKjQKSKӕ+ӗ
&Kt0LQK[LQFDPÿRDQF{QJWUuQKQJKLrQFӭXQj\GRFKtQKW{LWKӵFKLӋQVӕOLӋXOjNӃW TXҧ QJKLrQ FӭX WKӵF Vӵ FӫD W{L GѭӟL Vӵ KѭӟQJ GүQ NKRD KӑF FӫD TS 3KҥP1JX\ӉQĈӭF+RjQJ
7{L[LQFKӏXKRjQWRjQWUiFKQKLӋPYӅNӃWTXҧQJKLrQFӭXWURQJOXұQYăQWӕWQJKLӋSFӫDPuQK
7S+͛&Kt0LQKQJj\18 WKiQJQăP
+ӑFYLrQWKӵFKLӋQ
3KҥP+RjQJ3KL3KөQJ
Trang 7MӨC LӨC
Trang phө EuD
NhiӋm vө luұQYăQWKҥc Vƭ
Lӡi cҧPѫQ i
7yPWҳt luұQYăQ ii
LӡLFDPÿRDQ iii
Mөc lөc iv
Danh mөc tӯ viӃt tҳt vii
Danh mөc bҧng ix
Danh mөFKuQKҧnh xi
Mӣ ÿҫu 1
&KѭѫQJ TәQJTXDQWjLOLӋu Nҩm rӉ ngoҥi cӝng sinh (Ectomycorrhizas) 3
1.1 6ѫOѭӧc vӅ nҩm rӉ cӝng sinh (Mycorrhizas) 3
1.1.1 6ѫOѭӧc vӅ nҩm rӉ ngoҥi cӝng sinh 3
1.1.2 +uQKWKiLKӑFYjVӵ SKiWWULӇn cӫa nҩm rӉ ngoҥi cӝng sinh 5
1.1.3 1.1.3.1 HӋ rӉ nҩm (Ectomycorrhizal roots) 5
1.1.3.2 ThӇ VLQKEjRWӱ (Reproductive structures) 9
9DLWUzFӫa nҩm rӉ ngoҥi cӝng sinh 13
1.1.4 Nҩm rӉ ngoҥi cӝQJVLQKWUrQFk\WK{QJEDOiPinus kesiya 14
1.2 6ѫOѭӧc vӅ Fk\WK{QJEDOiPinus kesiya 14
1.2.1 Mӝt sӕ chi nҩm ngoҥi cӝng sinh vӟLFk\WK{QJEDOiPinus kesiya 15
1.2.2 1.2.2.1 Pisolithus 15
1.2.2.2 Suillus 16
1.2.2.3 Scleroderma 17
1.2.2.4 Rhizopogon 18
1.2.2.5 Laccaria 19
1.2.2.6 Hebeloma 20
Trang 8Mӝt sӕ QJKLrQFӭXOLrQTXDQ 21
1.2.3 1JKLrQFӭXFiFSKѭѫQJSKiSJk\QKLӉm EMF 21
1JKLrQFӭu vӅ (0)WUrQQKӳQJFk\WKXӝc chi Pinus 23
&KѭѫQJ Vұt liӋXYjSKѭѫQJSKiS 6ѫÿӗ WKtQJKLӋm 25
2.1 ĈӕLWѭӧQJQJKLrQFӭu 26
2.2 2.3 Vұt liӋu 26
+yDFKҩt 26
2.3.2 ThiӃt bӏ 26
3KѭѫQJSKiSQJKLrQFӭu 28
2.4 Thu thұp quҧ thӇ nҩm 28
2.4.1 0{Wҧ cҩXWU~Fÿҥi thӇ cӫa quҧ thӇ 29
2.4.2 3KkQOұSYjOѭXWUӳ mүu quҧ thӇ nҩm 29
2.4.3 Ĉӏnh danh bҵng kӻ thuұt sinh hӑFSKkQWӱ 30
2.4.4 0{Wҧ KuQKWKiLFiFFҩXWU~FYLWKӇ cӫa quҧ thӇ nҩm 33
2.4.5 KhҧRViWVLQKOêQҩm 34
2.4.6 *k\QKLӉm nҩm ngoҥi cӝng sinh OrQ Fk\WK{QJFRQPinus kesiya 35
2.4.7 &KѭѫQJ KӃt quҧ YjELӋn luұn Thu thұSYjSKkQOұp mүu quҧ thӇ nҩm 36
3.1 Ĉӏnh danh bҵng sinh hӑFSKkQWӱ YjFKӑn nҩPFKRP{KuQKJk\QKLӉm 38
3.2 KӃt quҧ giҧLWUuQKWӵ ÿRҥn ITS 38
3.2.1 KӃt quҧ giҧLWUuQKWӵ ÿRҥn LSU-28S cӫa chӫng 41Yj 42
3.2.2 KӃt quҧ [k\GӵQJFk\SKiWVLQKORjLFӫa hai chӫng nҩPÿѭӧc chӑn 43
3.2.3 ĈӏQKGDQKKuQKWKiLPүu quҧ thӇ cӫa hai chӫng nҩPÿѭӧc chӑn 45
3.3 Chӫng 4131 (Suillus sp.) 45
3.3.1 Chӫng 5208 (Scleroderma sp.) 47
3.3.2 KhҧRViWVLQKOêKDLFKӫng nҩPÿѭӧc chӑn 48
3.4 *k\QKLӉm Suillus luteus YjScleroderma sinnamariense WUrQFk\WK{QJEDOi 3.5 Pinus kesiya 52
KӃt quҧ thu nhұQFk\WK{QJFRQPinus kesiya 52
3.5.1
Trang 9KӃt quҧ kiӇPWUDKuQKWKiLFKySUӉ nhiӉm nҩm 553.5.2.
&KѭѫQJ KӃt luұQYjNLӃn nghӏ
KӃt luұn 584.1
KiӃn nghӏ 584.2
&KѭѫQJ7jLOLӋu tham khҧo 59
&KѭѫQJ Phө lөc
Trang 10DANH MӨC TӮ VIӂT TҲT
PCR Polymerase Chain Reaction
Trang 11PDA Potato-Dextrose agar
Trang 12DANH MӨC BҦNG
BҧQJ7K{QJWLQPӗi sӱ dөng trong PCR 31
BҧQJ&{QJWKӭc phӕi trӝn cho mӝt phҧn ӭng PCR 32
BҧQJ&iFQJKLӋm thӭc cho khҧRViWÿLӅu kiӋQQX{LFҩy nҩm 34
Bҧng 3.1: Sӕ Oѭӧng mүu quҧ thӇ nҩm ngoҥi cӝng sinh thu thұSYjSKkQOұp 36
Bҧng 3.2: KӃt quҧ giҧLWUuQKWӵ ÿRҥQ,76FiFFKӫng nҩPSKkQOұSWKjQKF{QJ 39
Bҧng 3.3: Tәng hӧp sӕ mүXSKkQOұSYjTXҧ thӇ WѭѫQJӭQJWKHRFiFFKLÿmÿӏnh danh 41
Bҧng 3.4: KӃt quҧ giҧLWUuQKWӵ ÿRҥn LSU-28S cӫa hai chӫng nҩPÿѭӧc chӑn 42
BҧQJ/ѭӧng sinh khӕLNK{Fӫa hai chӫng Suillus luteus YjScleroderma sinnamariense khi khҧRViWӣ nhӳQJP{LWUѭӡQJGLQKGѭӥQJÿLӅu kiӋn nhiӋWÿӝ Yj S+NKiFQKDX 49
BҧQJ&iFWK{QJVӕ WăQJWUѭӣng cӫDFk\WK{QJFRQPinus kesiya NKLJk\QKLӉm vӟi hai chӫng nҩm 6OXWHXVYj6VLQQDPDULHQVH so vӟLFk\ÿӕi chӭng 53
BҧQJ&{QJWKӭFP{LWUѭӡng khҧRViWÿLӅu kiӋQQX{LFҩy nҩm
BҧQJ7rQYjPmJHQFӫDFiFFKӫng sӱ dөng trong bӝ dӳ liӋu dӵQJFk\SKiW VLQKORjLFKRFKӫng 4131
BҧQJ7rQYjPmJHQFӫDFiFFKӫng sӱ dөng trong bӝ dӳ liӋu dӵQJFk\SKiW VLQKORjLFKRFKӫng 5208
Bҧng 6.4: KhӕLOѭӧng sinh khӕi kh{Fӫa Suillus luteus khҧRViWWUrQQKӳQJP{L
WUѭӡQJGLQKGѭӥQJNKiFQKDX
Bҧng 6.5: KhӕLOѭӧng sinh khӕLNK{Fӫa Suillus luteus khҧRViWӣ nhӳQJÿLӅu kiӋn
nhiӋWÿӝ NKiFQKDX
Bҧng 6.6: KhӕLOѭӧng sinh khӕLNK{Fӫa Suillus luteus khҧRViWӣ nhӳQJÿLӅu kiӋn
S+NKiFQKDX
Bҧng 6.7: KhӕLOѭӧng sinh khӕLNK{Fӫa Scleroderma sinnamariense khҧRViWWUrQ
nhӳQJP{LWUѭӡQJGLQKGѭӥQJNKiFQKDX
Bҧng 6.8: KhӕLOѭӧng sinh khӕLNK{Fӫa Scleroderma sinnamariense khҧRViWӣ
nhӳQJÿLӅu kiӋn nhiӋWÿӝ NKiFQKDX
Trang 13Bҧng 6.9: KhӕLOѭӧng sinh khӕLNK{Fӫa Scleroderma sinnamariense khҧRViWӣ
Trang 14DANH MӨ&+Î1+ҦNH
+uQK&ҩXWU~FFѫEҧn cӫa mӝt hӋ nҩm rӉ ngoҥi cӝng sinh 5
+uQK0{SKӓQJTXiWUuQK[kPQKLӉm cӫa ECM vӟi rӉ Fk\FKӫ 8
+uQK9zQJÿӡi cӫa mӝWORjLQҩm tҥo thӇ VLQKEjRWӱ dҥQJPNJ 10
+uQK0{Wҧ mӝt quҧ thӇ dҥng nҩm cөc 12
+uQK0{Wҧ mӝt sӕ SKѭѫQJSKiSJk\QKLӉm nҩm rӉ 23
+uQK6ѫÿӗ WKtQJKLӋm 26
+uQK.KXYӵc thu nҩPÿӏnh vӏ bҵng GPS 28
+uQK0{SKӓng cҩXWU~FFҳt dӑc mӝWPNJQҩm 34
+uQK$+uQKFKөp thӵFÿӏDYjNKXҭn lҥc sau 3 tuҫQQX{LFҩy cӫa nhӳng chӫng nҩPSKkQOұSWKjQKF{QJ 36
+uQK.Ӄt quҧ ÿLӋn di sҧn phҭP3&5ÿRҥQWUunh tӵ ITS mӝt sӕ mүXP{Wӯ quҧ thӇ nҩm 38
+uQK.Ӄt quҧ ÿLӋn di sҧn phҭP3&5ÿRҥQWUuQKWӵ LSU tӯ quҧ thӇ nҩm cӫa hai chӫQJYj 42
+uQK&k\SKkQORjLFKRFKӫQJYjFKӫng 5208 44
+uQK+uQKWKiLFKӫng 4131 46
+uQK+uQKWKiLFKӫng 5208 47
+uQK&Kӫng Suillus luteus YjScleroderma sinnamariense khi khҧRViWӣ nhӳng P{LWUѭӡQJGLQKGѭӥQJÿLӅu kiӋn nhiӋWÿӝ YjS+NKiFQKDX 48
+uQK%LӇXÿӗ P{Wҧ Oѭӧng sinh khӕLNK{WUXQJEuQK cӫa hai chӫng nҩm NKLQX{L cҩy ӣ nhӳQJP{LWUѭӡQJGLQKGѭӥQJNKiFQKDX 50
+uQK%LӇXÿӗ P{Wҧ Oѭӧng sinh khӕLNK{WUXQJEuQK cӫa hai chӫng nҩPNKLQX{L cҩy ӣ nhӳQJÿLӅu kiӋn nhiӋWÿӝ NKiFQKDX 51
+uQK%LӇXÿӗ P{Wҧ Oѭӧng sinh khӕLNK{WUXQJEuQKFӫa hai chӫng nҩm khi QX{LFҩy ӣ nhӳQJÿLӅu kiӋQS+NKiFQKDX 51
+uQK&k\WK{QJFRQPinus kesiya sau khi Jk\QKLӉm vӟi Suillus luteus Yj Scleroderma sinnamariense so vӟi nhӳQJFk\NK{QJJk\QKLӉm 52
Trang 15+uQK%LӇXÿӗ P{Wҧ FiFGӳ liӋu cӫDFk\WK{QJFRQPinus kesiya NKLFKROk\
nhiӉm hai chӫng nҩm so vӟLFk\ÿӕi chӭng 54
+uQK+uQKWKiLUӉ nҩm cӝQJVLQKWUrQFk\WK{QJFRQPinus kesiya sau 12 tuҫn Jk\QKLӉm vӟi Suillus luteus YjScleroderma sinnamariense 56 +uQK+uQKWKiLJLҧi phүXFKySUӉ nҩm Suillus luteus YjScleroderma
sinnmariense WUrQFk\WK{QJFRQPinus kesiya 57
+uQK.Ӄt quҧ xӱ OêWUuQKWӵ ÿRҥn ITS cӫa chӫQJWK{QJTXDF{QJFө
Trang 16MӢ ĈҪU
RӉ nҩm cӝQJVLQKOjPӝWKuQKWKӭc cӝng sinh giӳa nҩPYjUӉ Fk\ӣ nhӳng thӵc vұt bұc cao7URQJÿyQҩPJL~SFk\FKӫ hҩSWKXFiFFKҩWGLQKGѭӥng thiӃt yӃu tӯ ÿҩWYjWӯ Fk\FKӫ, nҩm nhұn nhӳng hӧp chҩt carbon tӯ TXiWUuQKTXDQJKӧSÿӇ tӗn tҥi [1, 2@7KHR6PLWKYj5HDG(2008)FyGҥng cӝng sinh cӫa nҩm rӉ vӟi thӵc vұt, WURQJÿy nҩm rӉ ngoҥi cӝng sinh (Ectomycorrhizas, EC0 ÿѭӧF[HPOjKuQKWKӭc cӝng sinh chiӃPѭXWKӃ YjFyêQJKƭDÿһc biӋt quan trӑng vӟLFiFORjLWKӵc vұt thuӝc
hӑ 7K{QJKӑ Dҿ, hӑ BҥFK'ѭѫQJYjKӑ LiӉu tҥLFiFNKXUӯQJ{QÿӟLYjFұn nhiӋt ÿӟi [3] 7KHR QKѭ WKӕQJ Nr FӫD +DOO Yj FV FiF QKj QJKLrQ FӭX ÿm WuP UDkhoҧQJORjLQҩm cӝQJVLQKFyJLiWUӏ cao vӅ mһWGLQKGѭӥng hoһFGѭӧFWtQK[4] 7UrQWKӃ giӟLÿmFyUҩt nhiӅXQJKLrQFӭu Jk\QKLӉm nҩm ngoҥi cӝng sinh nhҵm
hӛ trӧ VLQKWUѭӣng FKRFk\FRQWURQJJLDLÿRҥQÿҫu7URQJÿLӅu kiӋn tӵ QKLrQQҩm
rӉ cho thҩy nhiӅu ҧQKKѭӣQJWtFKFӵc OrQFk\FKӫ QKѭNtFKWKtFKVLQKWUѭӣng cӫDFk\nhӡ JL~SFk\WăQJFѭӡng hҩp thu chҩWGLQKGѭӥQJYjQѭӟFWURQJÿҩWJL~SWăQJÿӅ NKiQJFӫDFk\YӟLVkXEӋQKYjWăQJVӭc chӕng chӏu vӟLÿLӅu kiӋn khҳc nghiӋt cӫa P{LWUѭӡng [5] .KLQKuQQKұn vӅ ÿLӅu kiӋn tӵ nhirQӣ ViӋt Nam, nguӗQWjLQJX\rQnҩm lӟn ӣ ViӋW1DPÿѭӧF[HPOjNKiÿDGҥQJÿһc biӋt mӝWYjLQJKLrQFӭXEDQÿҫu cho thҩy hӋ nҩm ӣ khu vӵc Nam Bӝ Yj7k\1JX\rQUҩWSKRQJSK~>@.LӇu rӯng
Fk\ Oi NLP ѭD ViQJ FKӏu hҥQ ÿLӇQ KuQK ӣ /kP ĈӗQJ Oj Fk\ WK{QJ ED Oi (Pinus
kesiya ÿѭӧF [HP Oj QpW ÿһF WUѭQJ FKR NLӇu thҧm thӵc vұt ӣ ÿk\ >@ Vӟi nhӳng
tiӅP QăQJ WtFK Fӵc cӫa nҩm ngoҥi cӝQJ VLQK ÿӕi vӟL Fk\ WK{QJ Yj F{QJ WiF WUӗng rӯQJ FK~QJ W{L WKӵc hiӋQ ÿӅ WjL ³6jQJ OӑF Yj Jk\ QKLӉm mӝt sӕ ORjL Qҩm ngoҥi
cӝng sinh thu thұp tӯ rӯQJSKzQJKӝ ÿҫu nguӗQĈD1KLPWUrQFk\WK{QJEDOiPinus
kesiya´ vӟLFiFPөFWLrX
- Ĉӏnh danh mӝt sӕ ORjLQҩm ngoҥi cӝng sinh trong rӯQJWK{QJEDOiPinus
kesiya thuӝc rӯQJSKzQJKӝ ÿҫu nguӗQĈD1KLP
- ;iFÿӏnh mӝt sӕ ORjLQҩPFyWLӅPQăQJtrong viӋc hӛ trӧ VLQKWUѭӣng cho Fk\FRQÿһc biӋWÿӕi vӟLFiFORjLQҩPFyJLiWUӏ vӅ mһWGLQKGѭӥng hoһc GѭӧFWtQKFyWKӇ tұn dөng
Trang 17- ;k\ GӵQJ P{ KuQK Jk\ QKLӉm nҩm ngoҥi cӝQJ VLQK WUrQ Fk\ WK{QJ FRQ
Pinus kesiya
ĈӅ WjL mӝt phҫQÿѭӧFWjLWUӧ tӯ dӵ iQQJKLrQFӭu ³7iFÿӝng cӫa quҫQ[mQҩm ngoҥi cӝng sinh tӟi sӵ WiLVLQKFӫa rӯQJWK{QJWҥi ViӋW1DP´do ViӋn NҩPYj&{QJnghӋ sinh hӑc thӵc hiӋQFQJYӟi Ban quҧQOêUӯQJSKzQJKӝ ĈD1KLP
Trang 18
&KѭѫQJ
TӘNG QUAN 7¬,/,ӊU
Trang 19Nҩm rӉ ngoҥi cӝng sinh (Ectomycorrhizas)
rӉ, mycorrhiza OjPӕi quan hӋ giӳa hӋ rӉ cӫa thӵc vұWYjQҩm [8]
RӉ nҩm (tӭc cҩXWU~FQҩP[kPQKLӉm WUrQUӉ Fk\ OjKLӋQWѭӧng cӝng sinh giӳa nҩPYj thӵc vұt phә biӃn trong tӵ QKLrQ1ҩm cӝng sinh nhұQÿѭӧc tӯ Fk\FKӫ thӵc vұW FiF hӧp chҩW FDUERQ FiF FKҩW GLQK Gѭӥng Fk\ FKӫ thӵc vұW FNJQJ QKұQ ÿѭӧc GLQKGѭӥQJYjQѭӟc cҫn thiӃt tӯ hӋ sӧi cӫa nҩm hҩp thu ÿѭӧc [9] RӉ YjQҩPFQJtӗn tҥL Oj NӃt quҧ cӫa sӵ tiӃQ KyD FKXQJ Ĉm Fy QKLӅX F{QJ WUuQK NKRD Kӑc chӭng PLQKYDLWUzFӫa nҩm cӝng sinh mang lҥi nhӳng lӧLtFKWROӟn, thiӃt thӵFÿӕi vӟLTXiWUuQKVLQKWUѭӣQJYjSKiWWULӇn cӫDFk\WURQJÿLӅu kiӋn bҩt lӧi cӫDP{LWUѭӡng, bӣi vұy chӍ WURQJÿLӅu kiӋQÿҩWÿDLNK{KҥQQJKqRGLQKGѭӥQJWKuQҩm rӉ mӟLSKiWKX\tӕWYDLWUzFӝng sinh HuQKWKӭc cӝQJVLQKQj\ÿmYjÿDQJÿѭӧFQJKLrQFӭu (vӅ SKkQloҥi, sinh hӑFSKkQWӱ, ҧQKKѭӣng cӫDFK~QJÿӕi vӟi thӵc vұW« Yjӭng dөQJYjRthӵc tiӉn sҧn xuҩWQ{QJ± OkPQJKLӋp ӣ nhiӅXQѭӟFWUrQWKӃ giӟi
CytWQKҩWKuQKWKӭc cӝng sinh cӫa nҩm rӉ ÿmÿѭӧc ghi nhұQWURQJÿyQKӳng KuQKWKӭc phә biӃn nhҩt bao gӗm [10]:
- Nӝi cӝng sinh (vesicular ± arbuscular mycorrhizas, VAM)
- Ngoҥi cӝng sinh (ectomycorrhizas, ECM)
- Cӝng sinh vӟi nhӳQJFk\WKXӝc hӑ Phong lan (orchid mycorrhizas)
- Cӝng sinh vӟi nhӳQJFk\WKXӝc bӝ Ĉӛ TX\rQHULFRLGP\FRUUKL]DV
- Nӝi ngoҥi cӝng sinh (ectendomycorrhizas)
Trang 20cӝng sinh chӫ yӃu thuӝFQJjQKQҩPĈҧm (Basidiomycota), mӝt sӕ ORjLWKXӝFQJjQKnҩm T~L$VFRP\FRWD [12]
HӋ nҩm rӉ ngoҥi cӝng sinh rҩWÿDGҥngKjQJQJjQORjL(0F (ectomycorrhizal fungi) ÿmÿѭӧFNKiPSKiQKѭQJvүQFKѭDFydӵ ÿRiQFKtQK[iFYӅ sӵ phong ph~cӫa QKyPQҩPQj\ 1JKLrQFӭu hiӋn tҥi cho thҩ\FyNKRҧQJORjLQҩm ngoҥi cӝQJVLQKÿmÿѭӧc ghi nhұQWUrQWKӃ giӟLYjGӵ ÿRiQPӭFÿӝ ÿDGҥng cӫDFK~QJFythӇ OrQ ÿӃQ ÿӃQ ORjL >@ ĈLӅX Qj\ Fy QJKƭD Oj FRQ QJѭӡi chӍ mӟi biӃWÿӃn khoҧng mӝt phҫn ba sӕ ORjLQҩm ngoҥi cӝng sinh tӗn tҥLWUrQWUiLÿҩWQj\7URQJÿyFyNKRҧng 80% nҩm ngoҥi cӝQJVLQKKuQKWKjQKFҩXWU~FVLQKVҧQFzQJӑi OjWKӇ VLQKEjRWӱ hay quҧ thӇ nҩPÿmÿѭӧFQJKLrQFӭu Quҧ thӇ nҩPÿѭӧFiSGөng WURQJQJKLrQFӭXÿӏnh danh hӋ nҩm rӉ NKLFK~QJFӝng sinh vӟi mӝWORjLWKӵc vұt QjRÿy7X\QKLrQQKӡ YjRVӵ tiӃn bӝ cӫa kӻ thuұt sinh hӑFSKkQWӱFiFQKjQJKLrQcӭXSKiWKLӋn sӵ ÿDdҥng cӫa nҩm trong rӉ FDRKѫQQKLӅu so cҩXWU~FTXҧ thӇ nҩm PjWDTXDQViWÿѭӧc [12] Do sӵ chӑn lӑFYj WtQKWKtFKQJKLNKiFQKDXSKҥm vi tӗn tҥLFk\FKӫ NKiFQKDXPӝt sӕ ORjLQҩm chӍ cӝng sinh vӟi mӝt chi thӵc vұt nhҩWÿӏnh, trong khi mӝt sӕ ORjLFyWKӇ cӝng sinh vӟi nhiӅu QKyP thӵc vұt NKiFQKDXNhӳng
ORjL Fy WKӇ cӝQJ VLQK WUrQ QKyP WKӵc vұt gӗm FiF ORjL WKXӝc chi Amanita,
Cantharenlus; nhӳQJORjLFӝQJVLQKWUrQ-3 QKyPWKӵc vұt QKѭBoletus, Ramaria;
YjFKӍ cӝQJVLQKWUrQPӝt QKyPWKӵc vұt QKѭSuilus, Russula [14] MӝWFk\FyWKӇ
FyQKLӅu loҥi nҩm cӝQJVLQKFQJ mӝWO~Fÿ{LNKLFy thӇ OrQÿӃn 15 lojL>@
7KHR QKѭ QJKLrQ Fӭu cӫD 7HGHUVRR Yj cs (2013) *HPO Yj cs (2017), nҩm ngoҥi cӝng sinh hiӋQFyKѫQFKL>@7URQJÿyPӝt sӕ FKLWKѭӡng gһSYjÿѭӧc
QJKLrQ FӭX VkX EDR Jӗm: Pisolithus, Suillus, Laccaria, Hebeloma, Rhizopogon,
Tuber, Scleroderma, Amanita, Paxillus, Cantharellus, Lactarius, Cenococcum, Hysterangium, Thelephora
HӋ rӉ nҩm ngoҥi cӝQJVLQKSKiWWULӇQѭXWKӃ ӣ nhӳng khu rӯQJOiNLPQKӳng khu vӵF Q~L FDR Fy QKLӋW ÿӝ thҩS (&0 FNJQJ ÿѭӧF WuP WKҩy nhiӅu ӣ nhӳng khu rӯQJFk\OiUӝQJYQJ {Qÿӟi hoһFĈӏa Trung HҧLYjFNJQJ[Xҩt hiӋn ӣ mӝWYjLKӋ VLQKWKiL[DYDQKRһc rӯQJPѭDQKLӋWÿӟi hoһc cұn nhiӋWÿӟi [10]
Trang 21+uQKWKiLKӑFYjVӵ SKiWWULӇn cӫa nҩm rӉ ngoҥi cӝng sinh
1.1.3.
1.1.3.1 HӋ rӉ nҩm (Ectomycorrhizal roots)
Mӝt hӋ nҩm rӉ ngoҥi cӝng sinh bao gӗm hӋ sӧi nҩPOLrQNӃWWURQJÿҩt, hӋ rӉ nҩPYjFҩXWU~FVLQKVҧQKD\FzQJӑLOjTXҧ thӇ nҩm [10] RӉ nҩm ngoҥi cӝng sinh WKѭӡQJKuQKWKjQKӣ ÿҫXFKySUӉ cӫDFk\FKӫSKkQEӕ NK{QJÿӗQJÿӅXWURQJÿҩt, chӫ yӃXWuPWKҩy nhiӅu trong lӟSÿҩt mһt chӭa nhiӅXPQFKӭ kh{QJFyWURQJFiFtҫQJÿҩWSKtDGѭӟi [17, 18]
CҩX WU~F UӉ nҩm ngoҥi cӝQJ VLQK SKiW WULӇQ WUrQ Fѫ Vӣ WѭѫQJ WiF YӅ mһt di truyӅn giӳa nҩPYjFk\FKӫ Hҫu hӃt nhӳQJFk\FyQҩm ngoҥi cӝQJVLQKÿӅXFyNLӇu SKkQQKiQKWҥo rӉ ErQWӭFOjFyQKӳng rӉ ngҳn mӑc ra tӯ mҥQJOѭӟi nhӳng rӉ GjLYjGj\KѫQTheo WilcR[ Yj.XELNRYD UӉ GjLYjUӉ ngҳn vӅ FѫEҧQFycҩX WU~F JLӕQJ QKDX QKѭQJ UӉ ngҳQ WKѭӡQJ Fy ÿѭӡQJ NtQK QKӓ KѫQ Yj SKiW WULӇn chұP KѫQ QKLӅu so vӟi rӉ GjL Sӵ SKiW WULӇn chұm cӫa rӉ ngҳQ Oj Fҫn thiӃW ÿӇ cho
&È&7+¬1+3+ҪN CӪA MӜT Hӊ NҨM Rӈ NGOҤI CӜNG SINH
+uQKWKiLEjRWӱ
+uQKWKiLTXҧ thӇ +uQKWKiLTXҧ thӇ
Quҧ thӇ
dҥng
nҩPPNJ
Quҧ thӇ dҥng nҩm cөc
Nҩm rӉ OLrQNӃt vӟi Fk\FKӫ
+uQK1.1: CҩXWU~FFѫEҧn cӫa mӝt hӋ nҩm rӉ ngoҥi cӝng sinh [10]
Trang 22SKpSQҩPFyÿӫ thӡLJLDQKuQKWKjQKOLrQNӃt cӝng sinh vӟi rӉ Fk\FKӫ &KLOYHUVYjGust, 1982) [10] Mӕi quan hӋ ngoҥi cӝng sinh ÿѭӧFKuQKWKjQKNKLORjLQҩm EMF WѭѫQJWKtFKYjUӉ Fk\FKӫ tiӃS[~FYӟLQKDXFQJYӟLÿLӅu kiӋQP{LWUѭӡQJWKtFKKӧp cho viӋc cӝQJVLQK4XiWUuQKFӝQJVLQKÿѭӧc P{ tҧ QKѭVDX [10, 19]:
1 9zQJÿӡi cӫa mӝt hӋ nҩm rӉ ngoҥi cӝng sinh bҳWÿҫu tӯ sӵ nҧy mҫm Sau khi
hӋ nҩm rӉ bӏ chӃWÿLFyWKӇ GRÿӝng vұWăQJһSÿLӅu kiӋn bҩt lӧLP{LWUѭӡng WKD\ ÿәL«