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       ±è¼±¿ø ±³¼ö        2015.03.19 11:17        12273
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°æ»ó´ëÇб³ ±è¼±¿ø ±³¼ö

 

1. °³¿ä
ÇÕ¼º»ý¹°ÇÐ ±â¼úÀÇ ¹ß´Þ·Î ±âÁ¸ ´ë»ç°øÇÐ ±â¼ú·Î ±¸ÇöÀÌ Èûµé¾ú´ø ´ë»ç°æ·Î È帧ÀÇ ¿ªµ¿Àû Á¶Àý(dynamic regulation)ÀÌ °¡´ÉÇØÁ³´Ù. ´ë»çÈ帧ÀÇ ´Ù¾çÇÑ ¿ªµ¿Àû Á¶ÀýÀº ÀÏ·ÃÀÇ ´ë»çÁß°£Ã¼ °¨Áö±â¿Í À¯µµÀÎÀÚ(metabolite sensors and inducers)µé¿¡ ÀÇÇؼ­ Àü»ç, ¹ø¿ª, ¹ø¿ªÈÄ ¼ö½Ä´Ü°è(transcriptional, translational and posttranslational levels)¿¡¼­ ÁøÇàµÈ´Ù. ÀÌµé °¢ ´Ü°è¿¡¼­ÀÇ ¿ªµ¿Àû ´ë»çÁ¶ÀýÀº »óÀÌÇÑ ¹ÝÀÀ½Ã°£Ã´µµ(different dynamic time-scales)¸¦ °®´Â ´Ù¾çÇÑ ´ë»çÁ¶Àý ÀåÄ¡µé¿¡ ÀÇÇؼ­ ÀÌ·ç¾îÁö°í, ´Ù¾çÇÑ µ¹¹ßº¯¼öµéÀÌ °øÁ¸ÇÏ´Â »ê¾÷ÇöÀå¿¡¼­ÀÇ Àû¿ëÀ» À§Çؼ­´Â ÀÌµé ´ë»çÁ¶Àýȸ·ÎÀÇ °ß°í¼º(robustness)ÀÌ Áß¿äÇÑ Æ¯¼ºÀÌ µÈ´Ù. 


ÇÕ¼º»ý¹°ÇÐÀº ½Å¼ÓÇÏ°í Àú·ÅÇÑ À¯ÀüÀÚ ÇÕ¼º ¹× ºÐ¼®±â¼úÀÇ ¹ß´Þ·Î ¹ÙÀÌ¿ÀºÎÇ°ÀÇ ¼³°è-Á¦ÀÛ-½ÃÇèÀ» ÅëÇÑ À¯ÀüÀÚȸ·Î¿¡ ´ëÇÑ ÀÌÇØ¿Í »ê¾÷È­ °¡´É¼ºÀ» Á¦°íÇÏ°í ÀÖ´Ù. »ê¾÷±ÕÁÖ °³¹ß¿¡ ¼º°øÇϱâ À§Çؼ­´Â ¼öÀ²(yield), ³óµµ(titer), »ý»ê¼º(productivity)ÀÇ 3°¡Áö Áß¿äÇÑ ÀÎÀÚ¸¦ ¹Ýµå½Ã °í·ÁÇØ¾ß ÇÑ´Ù. Van DienÀº $1.00/lb °¡°Ý¼öÁØÀÇ ´ë·®»ý»ê »ý¹°Á¦Ç°ÀÇ »ý»ê¼º ±âÁØÀ¸·Î ¼öÀ²Àº ÃÖ´ë À̷мöÀ²ÀÇ 80%, »ý»ê³óµµ´Â 50 g/L, »ý»ê¼ºÀº 3 g/(L h)¸¦ Á¦½ÃÇÏ°í ÀÖ´Ù. »ó±â 3°¡Áö ÀÎÀÚ´Â ¼­·Î »óÃæµÇ´Â °³³äÀ¸·Î ƯÈ÷ ¼öÀ²°ú »ý»ê¼ºÀº »óÈ£ÀýÃæ(trade-off)ÀÇ °ü°è¿¡ ÀÖ´Ù(±×¸² 1). ZhuangµéÀº »ý»ê±ÕÁÖ ¼³°è¿¡¼­ ¼öÀ²À» ÃÖÀûÈ­ÇÑ °æ¿ì¿¡ »ý»ê¼ºÀº °¨¼ÒÇÏ´Â »óÈ£°£ÀÇ ³»ÀçÀû ÀýÃæ°ü°è¸¦ º¸°íÇÏ¿´´Ù[2]. ´ë»ç°øÇÐÀº ´ëºÎºÐÀÇ °æ¿ì¿¡ ¸ñÀû»ê¹° »ýÇÕ¼º°æ·Î¸¸À» ¿°µÎ¿¡ µÎ°í ±Ø´ëÈ­Çϱ⠶§¹®¿¡ ±Õü»ýÀ°¿¡ ÇÊ¿äÇÑ ´ë»çÀÚ¿øµéÀÌ ºÎÁ·À¸·Î ±ÕüÁõ½Ä°ú »ê¹°»ý»êÀÌ »óÈ£ÀýÃæ(trade-off)µÇ´Â °á°ú¸¦ °®°í ¿Â´Ù(±×¸² 2). µû¶ó¼­ »ý»ê¼öÀ²À» Çâ»ó¸¸À» ¿°µÎ¿¡ µÐ ´ë»çÁ¶Àý Àü·«µéÀº »ýÀ°¼Óµµ °¨¼Ò·Î ÀÎÇÑ Ã¼Àû »ý»ê¼º(volumetric productivity)ÀÇ °¨¼Ò·Ñ ±Í°áµÈ´Ù. ³·Àº üÀû »ý»ê¼ºÀº ¼³ºñÅõÀÚºñ ¹× ¿î¿µºñ ºÎ´ãÀÇ Áõ°¡·Î »ê¾÷È­ÀÇ Á¦¾à¿äÀÎÀÌ µÈ´Ù. 

 

±×¸² 1. ±ÕÁÖ ¼³°è ¾Ë°í¸®µë: ±ÕÁÖ¼³°è´Â ´ëºÎºÐ »ý»ê¼º º¸´Ù´Â ¼öÀ²Çâ»ó¿¡ ÁßÁ¡À» µÎ°í ÀÖ´Ù. »ýÀ°¼Óµµ¸¦ Èñ»ýÇϸ鼭 »ý»ê¼öÀ²À» Áõ°¡½ÃÅ°´Â À¯ÀüÀÚ Á¶ÀÛµéÀÌ ÁøÇàµÈ´Ù(a, solid arrow). ÀÌ·¯ÇÑ ±ÕÁÖÁ¶ÀÛµéÀº static techniques (static gene overexpression or deletion)À» ÅëÇؼ­ ÁøÇàÀÌ µÇ±â ¶§¹®¿¡ ¸ñÇ¥ ´ë»ç°æ·ÎÀÇ dynamic controlÀº ¾ø´Ù. ±×·¯³ª ÃÖ±Ù¿¡ dynamic flux balance analysis (dFBA)¿¡ ±â¹ÝÀ» µÐ »ý»ê¼ºÀ» °í·ÁÇÑ ±ÕÁÖ ¼³°è ¾Ë°í¸®µëÀÌ °³¹ßµÇ°í ÀÖ´Ù. ÀÌ ¾Ë°í¸®µëÀº product flux¿Í growth rate¸¦ ¼³Á¤ÇÏ°í ÀÌ·ÐÀûÀ¸·Î ±ÕÁÖ¸¦ ¼³°èÇÑ´Ù(a, dots). ±×¸®°í productivity, yield, titer¸¦ ¸ñÀûº¯¼ö·Î Çؼ­ ±ÕÁÖ¸¦ ÃÖÀûÈ­ ÇÑ´Ù(b).

 

±×¸² 2. ¾ß»ýÇü±ÕÁÖ¿Í ÀçÁ¶ÇÕ±ÕÁÖÀÇ ±âÁú ºÐ¹è: °¢ »ê¹°µé·ÎÀÇ »ó´ëÀûÀÎ ±âÁúÈ帧ÀÇ ¾çÀº È­»ìÇ¥ÀÇ ±½±â·Î Ç¥½ÃÇÔ. (a) ¾ß»ýÇü±ÕÁÖ¿¡¼­ ´ë»çÈ帧Àº ¹ÙÀÌ¿À¸Å½º·Î ´ëºÎºÐ È帣°í ¸ñÀû»ê¹°°ú ºÎ»ê¹°·Î´Â Á¦ÇÑµÈ ÀûÀº ¾ç¸¸ÀÌ È帧. (b) ÀçÁ¶ÇÕ±ÕÁÖ¿¡¼­ ´ë»çÈ帧Àº ¸ñÀû»ê¹°·Î ÁÖ·Î °¡µµ·Ï ´ë»ç Àç¼³°èµÊ. ´ë»çÈ帧ÀÇ ÃÖ´ëÇѵµ°¡ ±âÁú À¯ÀÔ¼Óµµ¿¡ ÀÇÇؼ­ Á¦ÇѵǾîÁö±â ¶§¹®¿¡ ¹ÙÀÌ¿À¸Å½º¿Í ¸ñÀû»ê¹°·ÎÀÇ ´ë»çÈ帧Àº ³»ÀçÀûÀÎ trade-offÀÇ °ü°è¿¡ ÀÖÀ½.

 

2. ÀÌ´Ü°è ´Éµ¿Àû ´ë»çÁ¶Àý(two-stage dynamic control strategy)
»ê¹° »ýÇÕ¼º°æ·Î °­È­·Î ÀÎÇÑ »ýÀ°¼ÓµµÀÇ ½É°¢ÇÑ °¨¼Ò¹®Á¦¸¦ ÇØ°áÇϱâ À§Çؼ­ »ýÀ°´Ü°è(growth stage)¿Í »ý»ê´Ü°è(production stage)·Î ±¸ºÐÇؼ­ ´ë»ç°æ·Î¸¦ Á¶ÀýÇÏ´Â À̴ܵ¿ÀûÁ¶Àý(two-stage dynamic control) ¹æ½ÄÀ» Àû¿ëÇÒ ¼ö ÀÖ´Ù. »ýÀ°´Ü°è¿¡¼­´Â »ê¹° »ýÇÕ¼º°æ·Î °­È­ À¯ÀüÀÚµéÀÇ ¹ßÇöÀ» ¾ïÁ¦Çؼ­ Àüü ´ë»çÈ帧ÀÌ ¾ß»ýÇü ±ÕÁÖ¿Í À¯»çÇϵµ·Ï Çؼ­ »ýÀ°¼Óµµ¸¦ Çâ»ó½ÃÅ°°í, ÀÏÁ¤ÇÑ ±Õü³óµµ¿¡ µµ´ÞÇÏ¸é »ý»ê´Ü°è·Î ÀüȯÇؼ­ »ê¹° »ýÇÕ¼º°æ·Î °­È­ À¯ÀüÀÚµéÀÇ ¹ßÇöÀ» ÅëÇؼ­ »ý»ê¼ºÀ» ±Ø´ëÈ­ÇÏ´Â °ÍÀÌ´Ù[3, 4]. µ¿Àû ´ë»çÁ¶Àý(dynamic control)ÀÇ ÀÌÀÍÀ» ºÐ¼®Çϱâ À§Çؼ­ Á¤Àû ´ë»çÁ¶Àý(static control strategy)¿Í µ¿Àû ´ë»çÈ帧 ±ÕÇü ºÐ¼®(dynamic flux balance analysis)¿¡ ÀÇÇÑ À̴ܵ¿ÀûÁ¶Àý(two-stage dynamic control strategy)ÀÇ »ý»ê¼ºÀ» ºñ±³ÇÑ °á°ú µ¿ÀûÁ¶ÀýÀü·«(dynamic control strategy)ÀÌ »ý»ê¼ºÀ» ´ëÆø Çâ»ó½ÃÅ°´Â °ÍÀ» ¾Ë ¼ö ÀÖ´Ù(±×¸² 3). ÀÌ·¯ÇÑ µ¿Àû´ë»çÁ¶ÀýÀº ÃÖ±Ù ÇÕ¼º»ý¹°ÇÐÀÇ ¹ßÀüÀ¸·Î À¯ÀüÀÚȸ·Î¸¦ ÀÌ¿ëÇÑ ´ë»ç°æ·ÎÀÇ ÇÙ½É È¿¼ÒµéÀÇ ¹ßÇö ¹× ´ë»çÈ帧À» ¿¬¼ÓÀûÀ̸鼭 ¿ªµ¿ÀûÀ¸·Î Á¶ÀýÀ» ÇÒ ¼ö ÀÖ´Â ¼öÁØÀ¸·Î±îÁö ¹ßÀüÇÏ¿´´Ù.

 

±×¸² 3. »ý»ê¼º Çâ»óÀ» À§ÇÑ Á¤Àû(static) ¹× µ¿Àû(dynamic) ´ë»ç°øÇÐ(metabolic engineering) Àü·«ÀÇ ºñ±³. (a) ¼÷½Å»ê »ý»êÀ» À§ÇÑ »ý»ê¼º ÇÑ°è. (b) static strategy (solid line)¿Í dynamic strategy (dashed line)¿¡ µû¶ó ¼³°èµÈ ±ÕÁÖµéÀÇ ÇÑ°è »ý»ê¼º ºñ±³. Specific product flux¿Í growth rate·Î »ý»ê±ÕÁÖÀÇ hypothetical operating pointsÀ» production envelope °î¼±(a, solid line)¿¡¼­ ¼±ÅÃÇÔ. Static strategyÀÇ »ý»ê¼ºÀº dynamic gene regulationÀÌ ¾ø´Â single-stage fermentation Á¶°ÇÀ¸·Î ¸ð»ç. Dynamic strategy´Â wild-type growth rates¸¦ °®´Â growth stage¿Í hypothetical strainÀÇ product fluxes¸¦ °®´Â production stage·Î ±¸¼ºµÇ´Â two-stage fermentationÀ» ÀǹÌÇÔ.

 

3. ´ë»ç¹°Áú °¨Áö(metabolite sensor) ±â¹Ý ¿¬¼ÓÁ¶Àý À¯ÀüÀÚȸ·Î
¿¬¼Ó´ë»çÁ¶Àý(continuous metabolic control)Àº ¼¼Æ÷ ³»ºÎ ¹× ¿ÜºÎÀÇ È¯°æº¯È­¸¦ µ¿ÀûÀ¸·Î °¨ÁöÇÏ°í º¯È­µÈ ȯ°æ¿¡ ¸ÂÃç¼­ ´ë»ç¸¦ Á¶ÀýÇÏ´Â °ÍÀ¸·Î »ê¾÷ÇöÀåÀÇ ºÒ±ÕÀÏÇÑ Á¶¾÷ȯ°æ°ú ¸Å¹ø µ¿ÀÏÇÏÁö ¾ÊÀº ¹ßÈ¿Á¶°Çµé¿¡ ¸ÂÃç¼­ ¼¼Æ÷°øÀåÀÌ ´Éµ¿ÀûÀ¸·Î ´ëÀÀÇÏ°Ô ÇÔÀ¸·Î½á ¸ñÀû»ê¹°ÀÇ »ý»ê¼ºÀÌ Ç×»ó ±Ø´ëÈ­ µÇµµ·Ï ÇÏ´Â °ÍÀÌ´Ù. Farmer & Liao°¡ ¶óÀÌÄÚÆæ »ý»ê¼º Çâ»óÀ» ¸ñÀûÀ¸·Î ÀÌ·¯ÇÑ ´Éµ¿Àû ¿¬¼ÓÁ¶Àý À¯ÀüÀÚȸ·Î¸¦ ÃÖÃÊ·Î »ç¿ëÇÏ¿´´Ù. À̵éÀº acetyl phosphate-activated sensor¸¦ °³¹ßÇؼ­ ÇØ´ç°æ·ÎÀÇ ´ë»çÈ帧À» °¨ÁöÇÏ°í ÇØ´ç°æ·Î ´ë»çÈ帧¿¡ ¸Â°Ô ¶óÀÌÄÚÆæ »ýÇÕ¼º°æ·Î ¹ßÇöÀ» Á¶ÀýÇÏ´Â À¯ÀüÀÚȸ·Î¸¦ ±¸ÃàÇؼ­ »ý»ê¼ºÀ» ȹ±âÀûÀ¸·Î Çâ»ó½ÃÄ×´Ù(±×¸² 4a). ¿¬¼ÓÁ¶Àý À¯ÀüÀÚȸ·ÎÀÇ ÇÙ½ÉÀÎ ´ÙÀ̳»¹Í ¼¾¼­(dynamic sensors)´Â ÇÙ½É ´ë»çÁß°£Ã¼¿¡ ¹Î°¨ÇÏ°Ô ¹ÝÀÀÇÏ´Â ¹ßÇö ÇÁ·Î¸ðÅ͵éÀ» Àü»çü ºÐ¼®À» ÅëÇؼ­ ¾ò°í À̵éÀ» »ýÇÕ¼º°æ·Î ¹ßÇöÁ¶Àý¿¡ Á¢¸ñÇÔÀ¸·Î½á ±¸ÃàµÉ ¼ö ÀÖ´Ù(±×¸² 4b). ±×·¯³ª ¿¬¼ÓÁ¶Àý À¯ÀüÀÚȸ·ÎÀÇ ´ÙÀ̳»¹Í ¼¾¼­¿Í ±¸µ¿ÀÚ(actuator)¸¦ ¸ñÀû¹°ÁúÀÌ º¯°æµÉ ¶§¸¶´Ù ´Ù½Ã °³¹ßÇØ¾ß µÇ±â ¶§¹®¿¡ ¸¹Àº ½Ã°£°ú ÀÚ¿øÀÇ ÅõÀÔÀÌ ¿ä±¸µÇ´Â ´ÜÁ¡ÀÌ ÀÖ´Ù. 

 

±×¸² 4. Dynamic control ±¸ÇöÀ» À§ÇÑ À¯ÀüÀÚȸ·Î. (a) acetyl phosphate-activated sensor ±â¹ÝÀÇ ¶óÀÌÄÚÆæ »ýÇÕ¼º À¯ÀüÀÚȸ·Î. (b) FPP-activated sensor ±â¹ÝÀÇ ¾Æ¸ðÆĵ𿣠»ýÇÕ¼º À¯ÀüÀÚȸ·Î.

 

4. °íÂû
ÃÖ±Ù ÇÕ¼º»ý¹°ÇÐÀÇ Áøº¸·Î ´ë»çÈ帧À» ´ÙÀ̳»¹ÍÇÏ°Ô Á¶ÀýÇÏ´Â ´Éµ¿Àû ´ë»ç°øÇÐ(dynamic metabolic engineering)ÀÌ ÁÖ¸ñÀ» ¹Þ°í ÀÖ´Ù. À̸¦ ÅëÇؼ­ ¹ßÈ¿°øÁ¤ÀÇ 3´ë ÇÙ½ÉÀÎÀÚÀÎ »ý»ê¼öÀ²(yield), »ê¹°³óµµ(titer), »ý»ê¼º(productivity)À» ȹ±âÀûÀ¸·Î Çâ»ó½Ãų ¼ö ÀÖ´Ù. ´ë»çÈ帧À» Á¶ÀýÇϱâ À§Çؼ­ µ¿ÀûµÇ¸ÔÀÓÁ¶Àý(dynamic feedback control)¿¡ ±â¹ÝÀ» µÐ ´ë»ç¹°Áú °¨Áö(metabolite sensor) ¿¬¼ÓÁ¶Àý°ú ÀÌ´Ü°è»ýÀ°Á¶°Ç(two-stage growth conditions)¿¡ ±â¹ÝÀ» µÐ ÀÌ´Ü°è ´Éµ¿Àû ´ë»çÁ¶Àý(two-stage dynamic control strategy)Àº ´Éµ¿Àû ´ë»ç°øÇÐÀÇ Áß¿äÇÑ Çٽɱâ¼úÀÌ´Ù. »ó±â µÎ ÇÙ½É ±â¼úÀº ¸ñÀû»ê¹°ÀÇ »ý»ê¼º(productivity) »Ó¸¸ ¾Æ´Ï¶ó, ¼öÀ²(yield)°ú ³óµµ(titers)µµ ´õºÒ¾î Çâ»ó ½Ãų ¼ö ÀÖ´Ù. µ¿Àû´ë»çÁ¶ÀýÀü·«(dynamic control strategies)ÀÌ ´õ È¿À²ÀûÀÌ°í ±¤¹üÀ§ÇÏ°Ô ÀÌ¿ëµÇ±â À§Çؼ­´Â µÎ°¡Áö Áß¿äÇÑ ¹ßÀüÀÌ ¿ä±¸µÈ´Ù. ù°´Â À¯ÀüÀÚ È¸·Î°¡ °ß°íÇؼ­ ±ÕÁÖ¿Í ¹ßÈ¿Á¶°Ç¿¡ »ó°ü¾øÀÌ ÀÏÁ¤ÇÑ ¼º´ÉÀ» ³ªÅ¸³»¾ß ÇÑ´Ù. µÎ ¹ø°´Â ¿¬¼ÓÁ¶Àý¿¡¼­ °¨Áö(sensing)¿Í ±¸µ¿(actuation)ÀÇ ¿ªµ¿¼º(dynamics)ÀÌ ´ë»çÈ帧À» È¿À²ÀûÀ¸·Î Á¶ÀýÇÒ ¼ö ÀÖÀ» Á¤µµ·Î ÃæºÐÈ÷ »¡¶ó¾ß ÇÑ´Ù. ÇâÈÄ¿¡ ÇÕ¼º»ý¹°ÇÐÀÇ ¹ßÀüÀº ÀÌ·± ¹®Á¦µéÀ» ÇØ°áÇÏ°í ´ë»ç°øÇаú »ý¹°°øÁ¤¿¡¼­ µ¿ÀûÁ¶ÀýÀÇ È°¿ëÀÇ ÆøÀ» ³ÐÈú °ÍÀÌ´Ù. º» µ¿Çâ¿ø°í´Â Venayak et al. (2015)ÀÇ ¡°Engineering metabolism through dynamic control¡±¿¡ ±â¹ÝÇÏ°í ÀÖÀ½.

 

5. Âü°í¹®Çå
1. Van Dien S: From the first drop to the first truckload: commercialization of microbial processes for renewable chemicals. Curr Opin Biotechnol 2013
http://dx.doi.org/10.1016/j.copbio.2013.03.002.
2. Zhuang K, Yang L, Cluett WR, Mahadevan R: Dynamic strain scanning optimization: an efficient strain design strategy for balanced yield, titer, and productivity. DySScO strategy for strain design. BMC Biotechnol 2013, 13:8..
3. Gadkar KG, Doyle Iii FJ, Edwards JS, Mahadevan R: Estimating optimal profiles of genetic alterations using constraint-based models. Biotechnol Bioeng 2005, 89:243-251.
4. Anesiadis N, Cluett WR, Mahadevan R: Dynamic metabolic engineering for increasing bioprocess productivity. Metab Eng 2008, 10:255-266.
5. Farmer WR, Liao JC: Improving lycopene production in Escherichia coli by engineering metabolic control. Nat Biotechnol 2000, 18:533-537.
6. Dahl RH, Zhang F, Alonso-Gutierrez J, Baidoo E, Batth TS, Redding-Johanson AM, Petzold CJ, Mukhopadhyay A, Lee TS, Adams PD et al.: Engineering dynamic pathway regulation using stress-response promoters. Nat Biotechnol 2013, 31:1039-1046.
7. Venayak N, Anesiadis N, Cluett WR, Mahadevan R.: Engineering metabolism through dynamic control. Curr Opin Biotechnol. 2015, doi: 10.1016/j.copbio.2014.12.022.




Total:118 page:(8/5)
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