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4 KKU Science Journal Volume 39 Number 1 <strong>Review</strong> √Ÿª∑’Ë 1 °≈‰°≈°“√∑”ß“π¢Õß‡Õπ‰´¡å∑’Ë‡°’Ë¬«¢âÕß°—∫‰Œ‚¥√‡®π‡¡µ“∫Õ≈‘ ¡„π‰´¬“‚π·∫§∑’‡√’¬ ª√–°Õ∫¥â«¬ (1) ‡Õπ‰´¡å‰π‚µ√®’‡π (nitrogenase) ´÷Ëßæ∫‰¥â„π‚§√ß √â“ß‡Œ‡∑Õ‚√´‘ µå (heterocyst) ∑”Àπâ“∑’Ëµ√÷ß ‚¡‡≈°ÿ≈‰π‚µ√‡®π·≈–‡ª≈’Ë¬π‰ª‡ªìπ·Õ¡‚¡‡π’¬·≈–‚¡‡≈°ÿ≈‰Œ‚¥√‡®π °≈‰°°“√∑”ß“π‡√‘Ë¡®“°Àπà«¬¬àÕ¬ ‰¥‚µ√®’‡π √’¥—§‡∑ (NifH) √—∫Õ‘‡≈Á°µ√Õπ¿“¬πÕ°®“°µ—«„ÀâÕ‘‡≈Á°µ√Õπ ferredoxin À√◊Õ flavodoxin ·≈â« àßÕ‘‡≈Á°µ√Õπ„ÀâÀπà«¬¬àÕ¬‰¥‰π‚µ√®’‡π (NifDK) ®“°π—Èπ‰¥‰π‚µ√®’‡π ®–√’¥‘«´å‚¡‡≈°ÿ≈ ‰π‚µ√‡®π‡ª≈’Ë¬π‰ª‡ªìπ·Õ¡‚¡‡π’¬ ·≈–‚¡‡≈°ÿ≈‰Œ‚¥√‡®π´÷Ëß„π°√–∫«π°“√µâÕß„âæ≈—ßß“π ATP (2) ‡Õπ‰´¡åÕ—æ‡∑§ ‰Œ‚¥√®’‡π (uptake hydrogenase, HupSL(C)) ´÷Ëßæ∫‰¥â∑’Ë‡¬◊ËÕ‰∑≈“§Õ¬¥å¢Õß ‡Œ‡∑Õ‚√´‘ µå ∑”Àπâ“∑’ËÕÕ°´‘‰¥´å‚¡‡≈°ÿ≈‰Œ‚¥√‡®π‰ª‡ªìπ‰Œ‚¥√‡®π‰ÕÕÕπ (H + ) ·≈–Õ‘‡≈Á°µ√Õπ (3) ‡Õπ‰´¡å‰∫‰¥‡√Á§—ππ—≈ ‰Œ‚¥√®’‡π (bidirectional hydrogenase) ´÷Ëßæ∫‰¥â∫√‘‡«≥‡¬◊ËÕ‰´‚µæ≈“ ´÷¡ ¢Õß‡´≈≈å ∑”Àπâ“∑’Ë√’¥‘«´å‰Œ‚¥√‡®π‰ÕÕÕπ°≈—∫‰ª‡ªìπ‚¡‡≈°ÿ≈‰Œ‚¥√‡®π °≈‰°°“√∑”ß“π¬—ß‰¡à∑√“∫ ·πà—¥ —ππ‘…∞“π«à“‡ªìπ°“√∑”ß“π§«∫§Ÿà°—π¢ÕßÀπà«¬¬àÕ¬‰Œ‚¥√®’‡π (HoxHY) ·≈–‰¥Õ–‚ø‡√ (HoxEFU) „π°“√√—∫Õ‘‡≈Á°µ√Õπ®“° NADH ·≈â« àßÕ‘‡≈Á°µ√Õπ„Àâ°—∫‰Œ‚¥√‡®π‰ÕÕÕπ‡°‘¥º≈‘µº≈ ‡ªìπ‚¡‡≈°ÿ≈‰Œ‚¥√‡®π·≈– NAD + (Tamagnini et al., 2007)
å ∫∑§«“¡ «“√ “√«‘∑¬“»“ µ√å ¡¢. ªï∑’Ë 39 ©∫—∫∑’Ë 1 5 ‰Œ‚¥√®’‡π ‡ªìπ‡Õπ‰´¡å„π°≈ÿà¡∑’Ë¡’‚≈À– π‘°‡°‘≈ (Ni) ‡ªìπÕß§åª√–°Õ∫„π‚§√ß √â“ß‡àπ‡¥’¬«°—∫ ‡Õπ‰´¡å¬Ÿ√’‡Õ (urease) ·≈–§“√å∫Õπ ¡ÕπÕ°‰´¥å ¥’‰Œ‚¥√®’‡π (carbon monoxide dehydrogenase) (Casalot and Rousset, 2001) °≈ÿà¡‡Õπ‰´¡å‡À≈à“π’È µâÕß°“√‚ª√µ’πµ—«à«¬ (auxiliary protein À√◊Õ accessory protein) „π°“√∑”ß“π ®“°°“√»÷°…“ §ÿ≥≈—°…≥– ¡∫—µ‘¢Õß‚ª√µ’πµ—«à«¬ æ∫«à“ ≈”¥—∫ π‘«§≈’‚Õ‰∑¥å·≈–≈”¥—∫°√¥Õ–¡‘‚π¢Õß‡Õπ‰´¡å¡’§«“¡ §≈â“¬°—π·≈–Õπÿ√—°…å (homology and conservation) „π°≈ÿà¡·∫§∑’‡√’¬ (Lutz et al., 1991) ‡Õπ‰´¡å‰Œ‚¥√ ®’‡π ¢Õß‰´¬“‚π·∫§∑’‡√’¬¡’‚ª√µ’πµ—«à«¬√à«¡∑”ß“π ‡àπ°—π §◊Õ °≈ÿà¡‚ª√µ’π Hyp (HypB, HypA, HypE, HypD, HypC ·≈– HypF) ´÷Ëß∂Ÿ°∂Õ¥√À— ·≈–·ª≈ √À— ®“°¬’π hypBAEDCF (Wünschiers et al., 2003) °≈ÿà¡‚ª√µ’π Hyp æ∫„π‰´¬“‚π·∫§∑’‡√’¬À≈“°À≈“¬ “¬æ—π∏ÿå Õ“∑‘ Trichodesmium erythraeum IMS101, Anabaena strain PCC 7120 ·≈– Synechocystis strain PCC 6803, Synechococcus strain 6301, Nostoc strain PCC 73102, Nostoc punctiforme ATCC 29133, Anabaena variabilis ATCC 29413 ·≈– Lyngbya majuscule CCAP 1446/4 (Happe et al., 2000; Linberg et al., 2000; Leitão et al., 2005; Tamagnini et al., 2007) ªí®®—¬∑’Ë àßº≈µàÕ°“√º≈‘µ‰Œ‚¥√‡®π’«¿“æ „π‰´¬“‚π·∫§∑’‡√’¬ ªí®®—¬¿“¬„π (Intrinsic factors) Õ“∑‘‡àπ °“√§«∫§ÿ¡°“√∑”ß“π¢Õß‡Õπ‰´¡å„π‡¡µ“∫Õ≈‘ ¡ ·≈–°“√ —ß‡§√“–Àå‡´≈≈åæ‘‡»… ‡ªìπµâπ ‡´≈≈å¡’°≈‰° „π°“√§«∫§ÿ¡°“√∑”ß“π¢Õß‡Õπ‰´¡åÕ—æ‡∑§ ‰Œ‚¥√‚√®’‡π ·≈–‡Õπ‰´¡å‰∫‰¥‡√Á§—ππ—≈ ‰Œ‚¥√®’‡π ‡æ◊ËÕà«¬√—°…“ ¡¥ÿ≈‰Œ‚¥√‡®π¿“¬„π‡´≈≈å Õ“∑‘ °“√ ≈¥·Õ§µ‘«‘µ’¢Õß‡Õπ‰´¡åÕ—æ‡∑§ ‰Œ‚¥√®’‡π §«∫§Ÿà °—∫°“√‡æ‘Ë¡·Õ§µ‘«‘µ’¢Õß‡Õπ‰´¡å‰∫‰¥‡√Á§—ππ—≈ ‰Œ‚¥√®’‡π (Tamagnini et al., 2002) °“√∑”„Àâ¬’π∑’Ë —ß‡§√“–Àå‡Õπ‰´¡åÕ—æ‡∑§ ‰Œ‚√®’‡π ‡ ’¬Àπâ“∑’Ë‰ª (hup gene knock-out) à«¬‡æ‘Ë¡ª√– ‘∑∏‘¿“æ°“√º≈‘µ ‰Œ‚¥√‡®π’«¿“æ (Tamagnini et al., 2002) ‰´¬“ ‚π·∫§∑’‡√’¬ Nostoc muscorum ∑’Ë‰¡à¡’¬’π hupL π—Èπ ‰¡à “¡“√∂ÕÕ°´‘‰¥´å‚¡‡≈°ÿ≈‰Œ‚¥√‡®π‰¥â (Axelsson et al., 1999) ‡¡◊ËÕ√–¥—∫°“√· ¥ßÕÕ°¢Õß‚ª√µ’π HupL ¢Õß Lyngbya majuscule ‡æ‘Ë¡¢÷Èπ æ∫«à“ ·Õ§µ‘«‘µ’ ¢Õß‡Õπ‰´¡åÕ—æ‡∑§ ‰Œ‚¥√®’‡π ‡æ‘Ë¡¢÷Èπ¥â«¬‡àπ°—π (Leitão et al., 2005) ·Õ§µ‘«‘µ’¢Õß‡Õπ‰´¡å‰π‚µ√®’‡π ·≈–‰Œ‚¥√®’‡π ∂Ÿ°¬—∫¬—Èß¥â«¬‚¡‡≈°ÿ≈ÕÕ°´‘‡®π∑’Ë‡°‘¥ ¢÷Èπ®“°°√–∫«π°“√ —ß‡§√“–Àå¥â«¬· ß ‰´¬“‚π ·∫§∑’‡√’¬®÷ß¡’°≈‰°„π°“√ª°ªÑÕß‰¡à„Àâ‡Õπ‰´¡å —¡º— °—∫‚¡‡≈°ÿ≈ÕÕ°´‘‡®π‰¥â‚¥¬µ√ß Õ“∑‘‡àπ (1) °“√ √â“ß‡´≈≈åæ‘‡»…¢÷Èπ„π‰´¬“‚π·∫§∑’‡√’¬π‘¥ µ√÷ß‰π‚µ√‡®π ‡√’¬°«à“ ‡Œ‡∑Õ‚√´‘ µå (heterocyst) (√Ÿª∑’Ë 2) à«¬„π°“√µ√÷ß‰π‚µ√‡®π„π ¿“«–∑’Ë¡’ ÕÕ°´‘‡®π (aerobic condition) ‡Œ‡∑Õ‚√´‘ µå‰¡à‡°’Ë¬«¢âÕß °—∫°√–∫«π°“√ —ß‡§√“–Àå¥â«¬· ß ¥—ßπ—Èπ ®÷ß‰¡à¡’ °“√ª≈¥ª≈àÕ¬ÕÕ°´‘‡®π ‡Õπ‰´¡å‰π‚µ√®’‡π ®÷ß∑”ß“π ‰¥âª°µ‘ (Lang et al., 1987; Fay, 1992; Adams, 2000) (2) ¡’°“√°√–µÿâπ°“√∑”ß“π¢Õß‡Õπ‰´¡å‰π‚µ√®’‡π ·≈–‰Œ‚¥√®’‡π „πà«ß ¿“«–∑’Ë‰¡à¡’· ß §«“¡‡¢â¡ · ßπâÕ¬À√◊Õ∑’Ë¡◊¥ ‡æ◊ËÕ≈¥‚Õ°“ —¡º— °—∫‚¡‡≈°ÿ≈ ÕÕ°´‘‡®π∑’Ë∂Ÿ°º≈‘µ¢÷Èπ¢≥– —ß‡§√“–Àå¥â«¬· ß ´÷Ëß °√–∫«π°“√π’È®–æ∫‰¥â„π‰´¬“‚π·∫§∑’‡√’¬ “¬æ—π∏ÿ ∑’Ë‰¡à “¡“√∂ √â“ß‡Œ‡∑Õ‚√´‘ µå (Bergman et al., 1997)
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