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18 KKU Science Journal Volume 39 Number 1 <strong>Review</strong> º≈‘µ‚ª√µ’π·≈â«‡¢â“‡°“–°—∫∫√‘‡«≥ enhancer ∑’ËÕ¬Ÿà „°≈â°—∫ RB ®“°π—Èπ virE ∑”Àπâ“∑’Ë √â“ß‚ª√µ’π (single strand DNA binding protein) ‡æ◊ËÕ‡¢â“‡°“–°—∫ T-DNA “¬‡¥’Ë¬«∑’Ë∂Ÿ°µ—¥ÕÕ°¡“ ∑”„Àâ T-DNA “¬‡¥’Ë¬«π—Èπ ¡’§«“¡¡—Ëπ§ß„π√–À«à“ß°“√ àß∂à“¬¬’π virB ∑”Àπâ“∑’Ë √â“ß‚ª√µ’π ·≈â«‡¢â“‰ª‡°“–°—∫‡¬◊ËÕÀÿâ¡‡¡¡‡∫√π¢Õßæ◊ ∑”„Àâ‡°‘¥√Ÿ ‡æ◊ËÕ àß∂à“¬ T-DNA ‡¢â“ Ÿà‡´≈≈åæ◊ (virE ∑”Àπâ“∑’§≈â“¬°—∫ tra gene) integration of T-strand in plant cell DNA ª≈“¬ 3' OH ¢Õß single strand T-DNA ®–‡¢â“‰ª synapsis °—∫ à«π¢Õß small homologies ¢Õß plant cell DNA ‡°‘¥®ÿ¥·µ°À—°∫π “¬Àπ÷Ëß¢Õß plant cell DNA ( “¬≈à“ß) ª≈“¬ 5' P ¢Õß single strand T-DNA ‡¢â“‰ª‡◊ËÕ¡µàÕ°—∫ plant cell DNA ∑’Ë‡°‘¥®ÿ¥·µ°À—° ´÷Ëß¡’√“¬ß“π«à“ virD Õ“®¡’ à«πà«¬„π°√–∫«π°“√ DNA repair nick and gap ¡’°“√‡◊ËÕ¡µàÕ∫√‘‡«≥ nick ·≈– gap ‚¥¬°“√∑”ß“π ¢Õß virD §≈â“¬°—∫°“√∑”ß“π¢Õß ligase ¿“¬„π à«π T-DNA ¡’¬’π°”Àπ¥°“√ √â“ß “√ opine ‡àπ nopaline synthase (nos), octopine synthase (ocs), agroclinoline synthase (acs) ·≈– agropine synthase (ags) à«π Õ◊ËπÊ ¿“¬„π Ti - plasmid ‰¥â·°à ®ÿ¥‡√‘Ë¡µâπ°“√ ®”≈Õß‚¡‡≈°ÿ≈ (ORI) à«π∑’Ë§«∫§ÿ¡°“√ àß∂à“¬ æ≈“¡‘¥ ‚¥¬«‘∏’§Õπ®Ÿ‡°—Ëπ ¬’π∑’Ë°”Àπ¥„Àâ‡´≈≈å ·∫§∑’‡√’¬ “¡“√∂„â “√æ«° opine ‡ªìπ·À≈àßæ≈—ßß“π ‡¡◊ËÕ·∫§∑’‡√’¬ àß T-DNA ‡¢â“‰ª„π‡´≈≈åæ◊·≈â« ‡´≈≈å¢Õßæ◊∫√‘‡«≥∑’Ë∂Ÿ°∫ÿ°√ÿ°®÷ß “¡“√∂ √â“ß “√ opine ¢÷Èπ‰¥âµ“¡π‘¥¢Õßæ≈“ ¡‘¥ Ti π—Èπ à«π Agrobacterium °Á “¡“√∂‡®√‘≠‡µ‘∫‚µ‰¥â‚¥¬„â “√ opine ∑’Ëæ◊ √â“ß¢÷Èπ‡ªìπ·À≈àßæ≈—ßß“π‰¥â ·≈–‡¡◊ËÕ°”®—¥ ·∫§∑’‡√’¬ÕÕ°‰ª æ◊¬—ß§ß —ß‡§√“–Àå “√ opine ‰¥â ‡π◊ËÕß®“° T-DNA ∂Ÿ° àß‡¢â“‰ªÕ¬Ÿà„π‡´≈≈åæ◊Õ¬à“ß ∂“«√·≈â« (Õ“√’¬å «√—≠êŸ«—≤π°å, 2541) „π à«π¢Õß T-DNA πÕ°®“°®–¡’¬’π∑’Ë °”Àπ¥°“√ —ß‡§√“–Àå “√ opine ·≈â« ¬—ß¡’¬’π∑’Ë °”Àπ¥°“√ √â“ßŒÕ√å‚¡πæ◊ (phytohormone) æ«° ÕÕ°´‘π·≈–‰´‚µ‰§π‘π¥â«¬ ‡ªìπ‡Àµÿ„Àâ‡´≈≈åæ◊∑’Ë‰¥â √—∫ T-DNA ¡’°“√‡®√‘≠‡µ‘∫‚µ·∫àß‡´≈≈åÕ¬à“ß√«¥‡√Á«·≈– ‰¡à®”°—¥ ∑”„Àâ‡°‘¥‡ªìπ°âÕπ‡π◊ÈÕ‡¬◊ËÕªÿÉ¡ª¡∑’Ë‰¡à “¡“√∂ æ—≤π“‡ªìπ¬Õ¥À√◊Õ√“°‰¥â ¬’π∑’Ë°”Àπ¥°“√ √â“ß ŒÕ√å‚¡πæ◊∑’Ëæ∫ §◊Õ trytophan monooxygenase (tms1), indoleacetamide hydrolase (tms2) ·≈– isopentenyl transferase (tmr) ‚¥¬æ∫«à“ ¬’π tms1 ·≈– tms2 °”Àπ¥°“√ √â“ß‡Õπ‰´¡å∑’Ë‡°’Ë¬«¢âÕß°—∫°“√‡ª≈’Ë¬π·ª≈ß “√ trytophan ‡ªìπ indoleacetamide ·≈–‡ª≈’Ë¬πµàÕ ‰ª‡ªìπ indoleacetic acid (IAA) ´÷Ëß‡ªìπ “√æ«° ÕÕ°´‘π à«π¬’π tmr ‡ªìπ‡Õπ‰´¡å∑’Ë∑”Àπâ“∑’Ë √â“ß isopentenyl adenosine (IPA) ´÷Ëß‡ªìπ “√æ«° ‰´‚µ‰§π‘π ¢Õ∫‡¢µ¢Õß T-DNA ∑’Ë®– àß∂à“¬‰ª ¬—ß‡´≈≈åæ◊®–§«∫§ÿ¡·≈–°”Àπ¥‚¥¬≈”¥—∫‡∫ ´È”Ê (terminal repeat) 25 §Ÿà‡∫ ®–¡’Õ¬Ÿà∑—Èß 2 ¢â“ß¢Õß T- DNA ‡√’¬° left border (LB) ·≈–right border (RB) (Õ“√’¬å «√—≠êŸ«—≤π°å, 2541) °“√ àß∂à“¬¬’π‚¥¬ Agrobacterium ´÷Ëß‡ªìπ«‘∏’°“√∑’ËÕ“»—¬∏√√¡“µ‘¢Õß ·∫§∑’‡√’¬‚√§æ◊∑’Ë¡’◊ËÕ«à“ A. tumefaciens à«¬„π°“√ àß∂à“¬¬’π‡¢â“ Ÿàæ◊ æ◊µâπ„À¡à∑’Ë‰¥â‡√’¬°«à“æ◊¥—¥·ª≈ß æ—π∏ÿå (transgenic plant) À≈—°°“√∂à“¬¬’π‚¥¬«‘∏’π’È§◊Õ π”‡π◊ÈÕ‡¬◊ËÕ¢Õßæ◊¡“∑”„Àâ‡°‘¥√Õ¬·º≈ ·≈â«‡≈’È¬ß∫π Õ“À“√‡æ“–‡≈’È¬ß√à«¡°—∫·∫§∑’‡√’¬ ®“°π—Èπ°”®—¥·∫§∑’‡√’¬ ¥â«¬¬“ªØ‘’«π– °àÕπ∑’Ë®–π”‘Èπ à«π¢Õßæ◊‰ª‡≈’È¬ß §—¥‡≈◊Õ°„πÕ“À“√ ‡´≈≈å∑’Ë‰¥â√—∫°“√∂à“¬¬’π®–‡®√‘≠ ‡µ‘∫‚µ‰¥â·≈â«π”‰ª—°π”„Àâ‡°‘¥‡ªìπµâπæ◊ ®“°π—Èπ µ√«® Õ∫°“√· ¥ßÕÕ°¢Õß¬’π‡æ◊ËÕ¬◊π¬—πº≈Õ’°§√—Èß Àπ÷Ëß ‚¥¬¬’π∑’Ë∂Ÿ° àß∂à“¬‚¥¬ à«π„À≠à¡’ 2 ·∫∫ §◊Õ sense ·≈– antisense ´÷Ëßæ◊π‘¥·√°∑’Ë‰¥â√—∫°“√∂à“¬ Ω“°¬’π‚¥¬ Agrobacterium §◊Õ Daucus carota (Õ“√’¬å «√—≠êŸ«—≤π°å, 2541) °“√ àß∂à“¬¬’π‚¥¬«‘∏’ Sense ·≈– Antisense Van der Kroll et al. (1990) æ∫«à“ °“√ ‡ª≈’Ë¬π·ª≈ß„¥Ê ∑’Ë‡°’Ë¬«¢âÕß°—∫¬’π dihydroflavonol - 4-reductase (DFR) ´÷Ëß‡ªìπ‡Õπ‰´¡å∑’Ë‡°’Ë¬«¢âÕß°—∫°“√ —ß‡§√“–Àå√ß§«—µ∂ÿ·Õπ‚∑‰´¬“π‘π ®– àßº≈µàÕ°“√
∫∑§«“¡ «“√ “√«‘∑¬“»“ µ√å ¡¢. ªï∑’Ë 39 ©∫—∫∑’Ë 1 19 · ¥ßÕÕ°¢Õß ’„π∑’Ë ÿ¥ ‰¡à«à“°“√‡ª≈’Ë¬π·ª≈ßπ—Èπ®– ‡°‘¥¢÷Èπ‡π◊ËÕß®“°°“√ àß∂à“¬¬’π‚¥¬«‘∏’ antisense ‡æ◊ËÕ ¬—∫¬—ÈßÀ√◊Õ≈¥°“√· ¥ßÕÕ°¢Õß¬’π (suppression) À√◊Õ àß∂à“¬¬’π‚¥¬«‘∏’sense ‡æ◊ËÕ‡æ‘Ë¡°“√ – ¡·≈–· ¥ßÕÕ° ¢Õß¬’π„Àâ¡“°¢÷Èπ (over expression) °“√ àß∂à“¬¬’π ‚¥¬„â«‘∏’ antisense π—Èπ‡ªìπ°√–∫«π°“√∂à“¬Ω“°¬’π ∑’ËµàÕ°≈—∫∑‘»∑“ß ‡¡◊ËÕ‡°‘¥°“√≈Õ°√À— ®“°¬’π∑’Ë∂à“¬ Ω“°‡¢â“‰ª ®–‰¥âÕ“√å‡ÕÁπ‡Õ “¬∑’Ë‡ªìπ§Ÿà ¡°—∫Õ“√å‡ÕÁπ‡Õ ¢Õß¬’π „πæ◊ ‡¡◊ËÕ‡°‘¥°“√®—∫‡¢â“§Ÿà°—π ∑”„Àâ‰¡à¡’°“√ ·ª≈√À— ‡ªìπ‚æ≈’‡ææ‰∑¥åÀ√◊Õ‚ª√µ’π ·≈–‰¡à‡°‘¥ °“√ √â“ß‡ÕÁπ‰´¡åÀ√◊Õ “√´÷Ëß‡ªìπº≈≈—æ∏å¢Õß¬’π¥—ß°≈à“« °“√ àß∂à“¬¬’π‚¥¬«‘∏’ antisense ‡æ◊ËÕ¬—∫¬—ÈßÀ√◊Õ≈¥°“√ · ¥ßÕÕ°¢Õß¬’π ‚¥¬∑’Ë antisense DFR gene ∑”„Àâ ª√‘¡“≥√ß§«—µ∂ÿ≈¥≈ß„πæ‘∑Ÿ‡π’¬ À√◊Õ°“√ àß∂à“¬ antisense DFR gene ‡¢â“ Ÿà‡¬Õ∫’√“ “¡“√∂‡ª≈’Ë¬π®“° ’·¥ß‡ªìπ ’¡æŸ ·≈– ”À√—∫æ◊ lisianthus æ∫«à“ ‡°‘¥°“√‡ª≈’Ë¬π ’®“°¡à«ß‰ª‡ªìπ¢“« πÕ°®“°π’È °“√ àß∂à“¬ antisense DFR gene ∑”„Àâ Torenia hybrida ‡ª≈’Ë¬π®“°¥Õ° ’πÈ”‡ß‘π‡ªìπ ’πÈ”‡ß‘πÕàÕπ‰¥â (Aida et al., 2000) ¥—ßπ—Èπ ®÷ßπ”¡“ª√–¬ÿ°µå„â„π°“√»÷°…“ °“√ àß∂à“¬¬’π DFR ‡¢â“æ◊ª∑ÿ¡¡“ ‚¥¬°≈‰°°“√‡°‘¥ ’„πæ◊‡°’Ë¬«¢âÕß°—∫‡Õπ‰´¡åÀ≈“¬π‘¥ ‡Õπ‰´¡å∑’Ë ”§—≠ §◊Õ ‡Õπ‰´¡å DFR ´÷Ëß∑”Àπâ“∑’Ë„π¢—ÈπµÕπ ªØ‘°‘√‘¬“√’¥—°—π ‡ª≈’Ë¬π “√µ—«°≈“ß (intermediate) dihydroflavonols ‡ªìπ leucoanthocyanidin ´÷Ëß “√ µ—«π’È‡ª≈’Ë¬π‰ªµ“¡≈”¥—∫ ®π°√–∑—Ëß‡ªìπ “√°≈ÿà¡ cyanidine, pelargonidine ·≈– delphinidin ´÷Ëß‡ªìπ √ß§«—µ∂ÿ·Õπ‚∑‰´¬“π‘π ‚¥¬¬’π DFR °”Àπ¥°“√ √â“ß ‡Õπ‰´¡å¥—ß°≈à“« ¥—ßπ—Èπ ∂â“¡’°“√ àß∂à“¬¬’π DFR ‚¥¬ «‘∏’ antisense ‡æ◊ËÕ¬—∫¬—ÈßÀ√◊Õ≈¥°“√· ¥ßÕÕ°¢Õß¬’π (suppression) Õ“®∑”„Àâ‰¡à¡’À√◊Õ≈¥°“√ √â“ß “√„π °≈ÿà¡ cyanidine, pelargonidine ·≈–delphinidin ·≈– àßº≈µàÕ°“√· ¥ßÕÕ°¢Õß ’„π∑’Ë ÿ¥ ”À√—∫°“√ àß∂à“¬¬’π ‚¥¬«‘∏’ sense ´÷Ëß‡ªìπ °√–∫«π°“√·∑√°¬’π∑’Ë¡’‘Èπ à«π§√∫∂â«π ¡∫Ÿ√≥å (full length gene) ‡¢â“ Ÿà ‘Ëß¡’’«‘µ∑’ËµâÕß°“√‡æ◊ËÕ„Àâ ‘Ëß¡’’«‘µπ‘¥π—Èπ “¡“√∂ √â“ß‚ª√µ’π´÷Ëß‡ªìπº≈≈—æ∏å ¢Õß¬’π∑’Ë·∑√°‡¢â“‰ªπ—ÈπÕÕ°¡“‰¥â ‚¥¬¬’π∑’Ë«à“π’ÈÕ“® ‰¥â¡“®“° ‘Ëß¡’’«‘µπ‘¥π—Èπ‡ÕßÀ√◊Õ ‘Ëß¡’’«‘µπ‘¥Õ◊Ëπ °Á‰¥â ªí®®ÿ∫—π‡∑§π‘§π’È∂Ÿ°π”¡“„â„π°“√ √â“ßæ◊ ¥—¥·ª≈ßæ—π∏ÿ°√√¡À≈“¬π‘¥ ‡àπ ¢â“« Golden Rice ·≈–ΩÑ“¬ BT ‡ªìπµâπ °“√ àß∂à“¬¬’π DFR ‚¥¬«‘∏’ Sense ‡æ◊ËÕ‡æ‘Ë¡°“√ – ¡·≈–· ¥ßÕÕ°¢Õß¬’π„Àâ¡“°¢÷Èπ (over expression) π—Èπ Õ“®∑”„Àâ¡’°“√ √â“ß “√„π°≈ÿà¡ cyanidine, pelargonidine ·≈– delphinidin ‡æ‘Ë¡¢÷Èπ àßº≈µàÕ°“√· ¥ßÕÕ°¢Õß ’∑’Ë¡“°¢÷Èπ¥â«¬‡àπ°—π √“¬ß“π¢Õß Holton (1996) »÷°…“°“√ àß∂à“¬ Sense DFR gene æ∫«à“ °“√ àß∂à“¬ sense DFR gene ®“° æ‘∑Ÿ‡π’¬ ∑”„Àâ¥Õ° ’¢“«¢Õß Dianthus caryophyllus ‡ª≈’Ë¬π‡ªìπ¥Õ° ’¡à«ß πÕ°®“°π’È¡’√“¬ß“π°“√ àß∂à“¬ ¬’π DFR ‚¥¬«‘∏’ RNA interference (RNAi) ´÷Ëß‡ªìπ °√–∫«π°“√¬—∫¬—Èß°“√· ¥ßÕÕ°¢Õß¬’π„π√–¥—∫ mRNA ‚¥¬À“°‡°‘¥Õ“√å‡ÕÁπ‡Õ “¬§Ÿà¢÷Èπ¿“¬„π‡´≈≈å Õ“√å‡ÕÁπ‡Õ “¬§Ÿà∑’Ë‡°‘¥¢÷Èππ’È®–∂Ÿ°∑”≈“¬‚¥¬°≈ÿà¡¢Õß ‡Õπ‰´¡åπ‘«§≈’‡Õ ∑’Ë‡√’¬°«à“ RNA-induce silencing complex (RISC) „Àâ‡°‘¥‡ªìπÕ“√å‡ÕÁπ‡Õ “¬§Ÿà‘Èπ‡≈Á°Ê ¢π“¥ª√–¡“≥ 22 π‘«§≈’‚Õ‰∑¥å ‡√’¬°«à“ small interfering RNA (siRNA) ´÷Ëß siRNA ∑’Ë‡°‘¥¢÷Èππ’È®– ‡ªìπµ—«°√–µÿâπ„Àâ‡°‘¥°“√∑”≈“¬ mRNA ∑’Ë¡’≈”¥—∫ ‡∫ ‡À¡◊Õπ°—πµàÕ‰ª ´÷Ëß‡◊ËÕ«à“°≈‰°∑’Ë‡°‘¥¢÷Èππ’È¡’ ª√–‚¬πå„π°“√à«¬„Àâ ‘Ëß¡’’«‘µ “¡“√∂µâ“π∑“πµàÕ ‰«√— ∫“ßπ‘¥‰¥â ·≈–¬—ß„â„π°“√§«∫§ÿ¡°“√ · ¥ßÕÕ°¢Õß¬’π„Àâ·µ°µà“ß°—π‰ªµ“¡à«ß°“√æ—≤π“ √–¬–µà“ßÊ ‰¥â ¡’√“¬ß“π°“√»÷°…“°“√ àß∂à“¬¬’π DFR ‚¥¬«‘∏’ RNA interference (RNAi) ‡æ◊ËÕ°“√ √â“ß °ÿÀ≈“∫¥Õ° ’πÈ”‡ß‘π‚¥¬ ∂“∫—π CSIRO ‚¥¬ Wang ·≈–§≥– (1997) ‰¥â„â RNAi (RNA interference) À¬ÿ¥°“√∑”ß“π¢Õß¬’π DFR „π°ÿÀ≈“∫ ’·¥ß ‡æ◊ËÕ ¬—∫¬—ÈßªØ‘°‘√‘¬“ cyanidin pathway §◊Õ °“√ √â“ß “√°≈ÿà¡ cyanidin ( ’·¥ß) „π¢≥–‡¥’¬«°—π°Á‡æ‘Ë¡°“√ —ß‡§√“–Àå “√„π°≈ÿà¡ delphinidin (´÷Ëß„Àâ ’πÈ”‡ß‘π) ‰¥âÕ¬à“ß ¡∫Ÿ√≥å ·≈–π—°«‘®—¬®“° ∂“∫—π«‘®—¬ Suntory ‰¥â„â
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