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       ºûÀ» ÀÌ¿ëÇÑ ¼¼Æ÷´Ü¹éÁú ±â´É Á¶Àý
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       ÀÌÁö¿¬ ±³¼ö        2014.07.29 17:57        12620
´Ù¿î·Îµå : ºûÀ» ÀÌ¿ëÇÑ ¼¼Æ÷´Ü¹éÁú ±â´É Á¶Àý_ÀÌÁö¿¬ ±³¼ö2.pdf(217 Kb)
 

ºûÀ» ÀÌ¿ëÇÑ ¼¼Æ÷´Ü¹éÁú ±â´É Á¶Àý

 

                                                                                     

                                                                                                                ¼­¿ï´ëÇб³ ¾àÇдëÇÐ ÀÌÁö¿¬ ±³¼ö

 

1. °³¿ä
¼¼Æ÷³»¿¡¼­ ¹ÙÀÌ¿ÀºÎÇ°µéÀÇ ±â´ÉÀ» ½Ã°£Àû °ø°£ÀûÀ¸·Î Á¤È®ÇÏ°Ô Á¶ÀýÇÒ ¼ö ÀÖ´Â ½Ã½ºÅÛÀ» È®¸³ÇÏ´Â °ÍÀº ÇÕ¼º»ý¹°ÇÐÀÌ Ãß±¸ÇÏ´Â Áß¿äÇÑ ¸ñÇ¥Áß Çϳª¶ó°í ÇÒ ¼ö ÀÖ´Ù (Medema et al., 2011). »ý¹°Çп¡ ÀÖ¾î ÃÖ±Ù ÁÖ¸ñ¹Þ°í ÀÖ´Â µÎ °¡Áö Å×Å©³î·ÎÁö´Â ºû¿¡ ¹ÝÀÀÇÏ´Â ´Ü¹éÁúÀ» ¿£Áö´Ï¾î¸µÇÏ¿© ±â´ÉÀ» Á¶ÀýÇÏ´Â optogenetics ºÐ¾ß¿Í ¿°±â¼­¿­ ƯÀÌÀûÀÎ DNA °áÇÕ ´Ü¹éÁúÀ» ÀÌ¿ëÇÑ genomic modificationÀ̶ó ÇÒ ¼ö Àִµ¥ ÇÕ¼º»ý¹°ÇÐÀû Á¢±ÙÀ» ÅëÇØ ÀÌ·¯ÇÑ ´Ü¹éÁúºÎÇ°µéÀ» È¿À²ÀûÀ¸·Î ºÐÇÒ Á¶¸³ÇÒ ¼ö ÀÖ´Ù¸é ¼¼Æ÷³»ÀÇ ´Ù¾çÇÑ ÇÁ·Î¼¼½º¸¦ È¿°úÀûÀ¸·Î Á¶Àý ÇÒ ¼ö ÀÖÀ» °ÍÀ¸·Î ¿¹»óµÈ´Ù (Moglich and Hegemann, 2013; Cho et al., 2013). ÀûÇÕÇÑ ÆÄÀåÀÇ ºûÀ» ¼±º°ÇÏ¿© ¼¼Æ÷³»ÀÇ ±¹ÇÑµÈ Áö¿ª¿¡ Á¶»ç(illumination)ÇÔÀ¸·Î½á ¿øÇÏ´Â ºÎÇ°ÀÇ ±â´ÉÀ» Á¶ÀýÇÏ¿© ¼¼Æ÷³» µ¶¼ºÀÌ ÀûÀ¸¸é¼­µµ ½Ç½Ã°£À¸·Î °¡¿ªÀûÀÎ º¯È­¸¦ ÀÏÀ¸Å³ ¼ö ÀÖ´Â ºñÆı«ÀûÀÎ (non-invasive) ¹æ¹ýÀº ºûÀ» ÀÌ¿ëÇÑ ¹æ¹ýÀÌ À¯ÀÏÇÏ´Ù°í ÇÒ ¼ö ÀÖ´Ù. Áö³­ ¼ö ³â°£ optogenetics ºÐ¾ß¿¡¼­´Â À¯ÀüÀÚ ÀçÁ¶ÇÕÀ» ÀÌ¿ëÇÑ ºû¿¡ ¹ÝÀÀÇÏ´Â ´Ü¹éÁúÀ» »ç¿ëÇÏ¿© ¼¼Æ÷ ±â´ÉÀ» Á¶ÀýÇϴ ÷´ÜÀÇ ¿¬±¸¸¦ º¸°íÇÏ°í ÀÖ´Ù (Moglich et al., 2009; Morgan et al., 2010; Ohlendorf et al., 2012; Zhou et al., 2012;Yin et al., 2013). º» º¸°í¼­¿¡¼­´Â optogenetics ºÐ¾ßÀÇ ÃÖ±Ù ¿¬±¸µ¿ÇâÀ» ºÐ¼®Çغ¸°íÀÚ ÇÑ´Ù.

 

2. ¼¼Æ÷ ´Ü¹éÁúÀÇ ±â´É Á¶Àý
Yang group¿¡¼­ ¹ßÇ¥ÇÑ LightOn À¯ÀüÀÚ¹ßÇö ½Ã½ºÅÛÀº Gal4-VVD µµ¸ÞÀÎ À¶ÇÕ¿¡ ÀÇÇØ ºû¿¡ ÀÇÇØ VVD µµ¸ÞÀÎÀÇ dimerizationÀ» Á¶ÀýÇÔÀ¸·Î½á UASG ¼­¿­¿¡ÀÇ °áÇÕ¿¡ ÀÇÇÑ transgeneÀÇ ¹ßÇöÀ» Á¶ÀýÇÏ¿´´Ù (Wang et al., 2012). LightOnÀº ³·Àº background¿Í ºü¸¥ kinetic, ±×¸®°í °¡¿ªÀû ¹ÝÀÀÀ» ÅëÇÑ À¯ÀüÀÚÀÇ inductionÀ» ºû¿¡ ÀÇÇØ Á¶ÀýÇÏ¿´°í ±¤¿øµµ ž籤¿¡ ºñÇØ ÈξÀ ³·Àº û»ö±¤À» ¿¬¼ÓÀûÀÎ illumination ÀÌ ¾Æ´Ñ  pulse¿¡ ÀÇÇØ Á¶ÀýÇÏ°í ºû¿¡ ±ØÈ÷ ¹Î°¨ÇÑ VVD ´Ü¹éÁúÀ» »ç¿ëÇÔÀ¸·Î½á µ¶¼ºÀ» ³·Ãâ ¼ö ÀÖ¾ú´Ù.

 

±×¸² 1. LightOn ½Ã½ºÅÛÀ» ÀÌ¿ëÇÏ¿© À¯ÀüÀÚ ¹ßÇöÀ» Á¶ÀýÇÑ ¿¬±¸¿¡¼­ÀÇ expression kinetic ¹× light pulse ȤÀº duration ½Ã°£ÀÇ Á¶Àý¿¡ µû¸¥ ¹ßÇö Á¤µµÀÇ Â÷ÀÌ.

 

Camilli group¿¡¼­´Â ¼¼Æ÷¸·ÀÇ Áö¹æ¼ººÐÀ¸·Î ¼¼Æ÷³» ´Ù¾çÇÑ ±â´ÉÀ» Á¶ÀýÇÏ´Â °ÍÀ¸·Î ¾Ë·ÁÁø Phosphoinositides (PIs)ÀÇ ¼¼Æ÷³» ·¹º§À» ºûÀ» ÀÌ¿ëÇÏ¿© ºü¸£°í ±¹¼ÒÀûÀ¸·Î ±×¸®°í °¡¿ªÀûÀ¸·Î Á¶ÀýÇÏ´Â ¿¬±¸¸¦ º¸°íÇÑ ¹Ù ÀÖ´Ù (Idevall-Hagren et al., 2012). ½Ä¹°¿¡¼­ À¯·¡ÇÑ ºû¿¡ ¹ÝÀÀÇÏ´Â ´Ü¹éÁúÀÎ cryptochrome 2 (CRY2)¿Í Àü»çÁ¶Àý ÀÎÀÚÀÎ CIBN ´Ü¹éÁú »çÀÌÀÇ dimerizationÀ» û»öÆÄÀåÀÇ ºûÀ» Á¶»çÇÔÀ¸·Î½á À¯µµÇÏ°í 5 ptaseOCRL (inositol 5 phosphatase domain of OCRL)¸¦ ¼¼Æ÷¸·À¸·Î À¯ÀÎÇÏ¿© PI(4,5)P2¿Í PI(3,4,5)P3 Å»ÀλêÈ­¸¦ À¯µµÇÏ¿´´Ù. 5 ptase ´Â ¼¼Æ÷¸·¿¡ Á¸ÀçÇÏ´Â PI(4,5)P2¿Í PI(3,4,5)P3 ·¹º§À» Á¶ÀýÇÏÁö¸¸ ¼¼Æ÷³»¿¡¼­ ±¹¼ÒÀû Á¶ÀýÀ» ÇÏÁö ¸øÇÏ´Â °ÍÀ¸·Î ¾Ë·ÁÁ® ÀÖ¾úÀ¸³ª ºû¿¡ ÀÇÇØ À¯µµµÇ´Â dimerization Çö»óÀ» ¿£Áö´Ï¾î¸µÇÏ¿© 5 ptase¸¦ ¼¼Æ÷¸·À¸·Î À̵¿½ÃÄÑ Phosphoinositides(PIs)ÀÇ ·¹º§À» °ø°£ÀûÀ¸·Î °¡¿ªÀûÀ¸·Î ±×¸®°í ºü¸£°Ô Á¶Àý °¡´ÉÇÏ¿´´Ù.

 

±×¸² 2. ºû¿¡ ÀÇÇØ 5-ptase ¸¦ plasma membraneÀ¸·ÎÀÇ À̵¿½ÃÄÑ PI(4,5)P2ÀÇ

Å»ÀλêÈ­¸¦ ºü¸£°Ô À¯µµÇÏ´Â °úÁ¤.

 

2013³â Zhang group¿¡¼­´Â Àü»çÁ¶ÀýÀ» À§ÇÑ ¸ðµâ·Î¼­ genomic anchor·Î Xanthomonas sp ¿¡¼­ À¯·¡ÇÑ TALEs (transcription activator-like effectors)°ú   Arabidopsis thaliana ¿¡¼­ À¯·¡ÇÑ cryptochrome 2 (CRY2)À» »ç¿ëÇÏ¿© ºûÀÌ ÀÖ´Â »óÅ¿¡¼­´Â TALE–CRY2 ´Ü¹éÁúÀÌ counterpartÀÎ CIB1 ´Ü¹éÁú¿¡ °áÇÕÇÏ¿© CIB1¿¡ ¿¬°áµÈ effector domain¿¡ ÀÇÇØ Àü»ç¸¦ È°¼ºÈ­½Ãų ¼ö ÀÖ´Â ¸ðµâÈ­ °¡´ÉÇÑ ½Ã½ºÅÛÀ» ¼Ò°³ÇÑ ¹Ù ÀÖ´Ù (Konermann et al., 2013).  ºûÀÇ ÆÄÀå, ºûÀÇ ¼¼±â, ºû¿¡ ÀÇÇÑ È°¼ºÈ­»óÅ À¯Áö±â°£ (duty cycle)µî¿¡ ´ëÇÑ Ã¼°èÀûÀÎ ¿¬±¸¿Í ´õºÒ¾î activation domain¿¡ ´ëÇÑ ¼±º°À» ÅëÇØ ÃÖÀûÈ­µÈ TALE–CRY2PHR/CIB1–VP64À» ã¾Æ³»¾ú´Ù. ÁÖ¸ñÇÒ Á¡Àº ¼ÒºÐÀÚ¿¡ ÀÇÇØ Á¶ÀýµÇ´Â TALE ½Ã½ºÅÛ¿¡ ºñÇØ on/off kineticÀÌ ºûÀ» ÀÌ¿ëÇÑ Á¶Àý¿¡¼­ ´õ È¿À²ÀûÀ¸·Î ÁøÇàµÉ ¼ö ÀÖ¾ú´Ù´Â Á¡ÀÌ´Ù.

 

±×¸² 3. LITE ½Ã½ºÅÛ µðÀÚÀΰú ºû¿¡ ÀÇÇÑ Àü»çº¯È­.

 

CRY2 ´Ü¹éÁúÀÇ °æ¿ì¿¡´Â ºû¿¡ ÀÇÇØ oligomerization ÇÁ·Î¼¼½º°¡ Á¶ÀýµÊÀ¸·Î½á ¼¼Æ÷³» ½Ã±×³ÎÀ» È°¼ºÈ­ ½ÃÅ°´Â °ÍÀ¸·Î ÃÖ±Ù¿¡ ¹àÇôÁ³´Ù (Bugaj et al., 2013). ¼¼Æ÷³» ½Ã±×³Î¸µÀÇ Á¶Àý¿¡ À־ oligomerizationÀº °¨µµ³ôÀº µðÁöÅÐÈ­µÈ ½ÅÈ£¹ÝÀÀÀ» Á¦°øÇÔ°ú ´õºÒ¾î ³ª³ë ½ºÄÉÀÏÀÇ ´Ü¹éÁú È°¼ºÀ» ¸¶ÀÌÅ©·Î ½ºÄÉÀÏÀÇ ¼¼Æ÷³» ±¸Á¶¿¡ ÀÇÇÑ Á¶Àý·Î ¿¬°áÇÔÀ¸·Î½á Áúº´¿¡ °ü¿©ÇÏ´Â È¿¼ÒÀÇ È°¼ºÁ¶Àý¿¡ ÀÖ¾î scaffold¸¦ Á¦°øÇÏ´Â °ÍÀ¸·Î ¾Ë·ÁÁ® ÀÖ´Ù. CRY2 ´Ü¹éÁúÀÇ PHR (Photolyase homology region) µµ¸ÞÀΰú ¿¬°áµÈ mCherry ´Ü¹éÁú(CRY2 mCherry)ÀÇ °æ¿ì HEK 293T ¼¼Æ÷³»¿¡¼­ û»ö±¤À» Á¶»çÇÑÁö 10ÃÊ ¾È¿¡ cluster¸¦ Çü¼ºÇÏ°í ºûÀ» Á¦°ÅÇÏ¸é ¼öºÐ¾È¿¡ Çظ®µÇ´Â °¡¿ªÀû °úÁ¤À» ÁøÇàÇÔÀÌ °üÂûµÇ¾ú´Ù. ÀÌ·¯ÇÑ °úÁ¤Àº ºûÀÇ À¯¹«¿¡µû¶ó Áö¼ÓÀûÀ¸·Î ¹Ýº¹µÉ ¼ö ÀÖ¾úÀ¸¸ç ºûÀ» Á¦°Å½Ã cluster ¼öÀÇ °¨¼Ò¿¡ ÀÖ¾î ¾à 5.5ºÐ¿¡ ÇØ´çÇÏ´Â ½Ã°£ »ó¼ö´Â ÀÌÀü¿¡ º¸°íµÈ ¼¼Æ÷³» È°¼ºÈ­µÈ Cry2 photoisomer ÀÇ lifetime (t1/2)°ú ÀÏÄ¡ÇÔÀ» ¾Ë ¼ö ÀÖ¾ú´Ù.

CRY2 mCherry Ŭ·¯½ºÅÍ Çü¼º ¸ðµâÀÌ oligomerization¿¡ ÀÇÁ¸ÇÏ´Â ½Ã±×³Î¸µ ½Ã½ºÅÛ¿¡ Àû¿ëµÉ ¼ö ÀÖ´ÂÁö È®ÀÎÇϱâ À§ÇØ LRP6 Cterminal domain (LRP6c)À» CRY2 mCherry¿¡ À¶ÇÕÇÔÀ¸·Î½á ºû¿¡ ÀÇÇØ À¯µµµÇ´Â Ŭ·¯½ºÅÍ Çü¼ºÀÌ Wnt/bcatenin ½ÅÈ£Àü´Þ°æ·Î¸¦ º¯È­½ÃÅ´À» È®ÀÎÇÏ¿´´Ù. ¶ÇÇÑ ¼¼Æ÷ÀÇ mobility¿¡ °ü¿©ÇÏ´Â °ÍÀ¸·Î ¾Ë·ÁÁø Rho GTPase Rac1ÀÇ oligomerizationÀ» ºû¿¡ ÀÇÇØ À¯µµÇÏ¿© HEK 293T ¼¼Æ÷³»¿¡¼­ cytoplasm¿¡¼­ ¼¼Æ÷¸·À¸·Î mCherryÀÇ Çü±¤ÀÌ À̵¿ÇÔÀ» º¸ÀÓÀ¸·Î½á ºû¿¡ ÀÇÇØ Rac1ÀÌ È°¼ºÈ­µÊÀ» ±Ô¸íÇÏ¿´´Ù.

            ±×¸² 4. ºû¿¡ ÀÇÇØ À¯µµµÇ´Â Cry2-LRP6c ÀÇ Å¬·¯½ºÅ͸µÀ» ÅëÇÑ

     Wnt/b-catenin ½Ã±×³Î¸µ °æ·Î Á¶ÀýÀÇ ¿¹.

 

3. °íÂû
Àü¼úÇÑ ¹Ù¿Í °°ÀÌ ÇÕ¼º»ý¹°ÇÐÀû ÀÀ¿ëÀ» À§ÇÑ ½ºÀ§Ä¡ÀÇ °³¹ß°ú È¿°úÀûÀÎ ºû Á¶Àý ½Ã½ºÅÛ È®¸³À» À§Çؼ­´Â ¹ÙÀÌ¿ÀºÎÇ°À¸·Î¼­ÀÇ ´Ü¹éÁúÀÇ Æ¯¼º ¹× ¼¼Æ÷³» ½ÅÈ£Àü´Þ¿¡ ÀÖ¾î ÀÛ¿ë±âÀü¿¡ ´ëÇÑ ¿¬±¸°¡ ¼±ÇàµÇ¾î¾ß ÇÒ °ÍÀÌ´Ù. ¼¼Æ÷ ³»¿¡¼­ ÀϾ´Â ´Ù¾çÇÑ ´Ü¹éÁúµéÀÇ ÇÁ·Î¼¼½º¿¡ ´ëÇÑ ÀÌÇØ¿Í ´õºÒ¾î ÇÕ¼º»ý¹°ÇÐÀû Á¢±ÙÀ¸·Î optogenetics¿¡¼­ ÇØ°á°¡´ÉÇÑ ¹®Á¦°¡ ¹«¾ùÀÎÁö ±×¸®°í ¼ÒºÐÀÚ È­ÇÕ¹°À» ÀÌ¿ëÇÑ Á¶Àýº¸´Ù ¿ì¼öÇÑ Á¡ÀÌ ¹«¾ùÀÎÁö¿¡ ´ëÇÑ ¹®Á¦ÀνÄÀÌ ¸Å¿ì Áß¿äÇÏ´Ù°í ÇÒ ¼ö ÀÖ´Ù.  ¶ÇÇÑ ºûÀ» ÅëÇÑ È¿À²ÀûÀÎ ¼¼Æ÷³» ´Ü¹éÁú ±â´É Á¶Àý¿¡ À־ÀÇ ºû ½ºÀ§Ä¡ÀÇ ±â´ÉÆò°¡´Â ºûÀÇ À¯¹«¿¡ µû¶ó ¸î ¹èÀÇ Â÷ÀÌ°¡ ³ª´Â°¡¿¡ ´ëÇÑ ±Ô¸í»Ó¸¸ ¾Æ´Ï¶ó ¾ó¸¶³ª »¡¸® È°¼º/ºñÈ°¼º »óÅ·ΠÀüȯµÉ ¼ö ÀÖ´ÂÁö¿¡ ´ëÇÑ ½Ã°£Àû Á¶Àý¿¡ ´ëÇÑ ¿¬±¸°¡ ÇʼöÀûÀ¸·Î ¼öÇàµÇ¾î¾ß ÇÒ °ÍÀ¸·Î »ç·áµÈ´Ù.  

 

4. Âü°í¹®Çå
1. Bugaj, L.J., Choksi, A.T., Mesuda, C.K., Kane, R.S., and Schaffer, D.V. (2013). Optogenetic protein clustering and signaling activation in mammalian cells. Nature methods 10, 249-252.
2. Cho, S.W., Kim, S., Kim, J.M., and Kim, J.S. (2013). Targeted genome engineering in human cells with the Cas9 RNA-guided endonuclease. Nature biotechnology 31, 230-232.
3. Idevall-Hagren, O., Dickson, E.J., Hille, B., Toomre, D.K., and De Camilli, P. (2012). Optogenetic control of phosphoinositide metabolism. Proceedings of the National Academy of Sciences of the United States of America 109, E2316-2323.
4. Konermann, S., Brigham, M.D., Trevino, A.E., Hsu, P.D., Heidenreich, M., Cong, L., Platt, R.J., Scott, D.A., Church, G.M., and Zhang, F. (2013). Optical control of mammalian endogenous transcription and epigenetic states. Nature 500, 472-476.
5. Medema, M.H., Breitling, R., Bovenberg, R., and Takano, E. (2011). Exploiting plug-and-play synthetic biology for drug discovery and production in microorganisms. Nature reviews Microbiology 9, 131-137.
6. Moglich, A., Ayers, R.A., and Moffat, K. (2009). Design and signaling mechanism of light-regulated histidine kinases. J Mol Biol 385, 1433-1444.
7. Moglich, A., and Hegemann, P. (2013). Biotechnology: Programming genomes with light. Nature 500, 406-408.
8. Morgan, S.A., Al-Abdul-Wahid, S., and Woolley, G.A. (2010). Structure-Based Design of a Photocontrolled DNA Binding Protein. Journal of Molecular Biology 399, 94-112.
9. Ohlendorf, R., Vidavski, R.R., Eldar, A., Moffat, K., and Moglich, A. (2012). From Dusk till Dawn: One-Plasmid Systems for Light-Regulated Gene Expression. Journal of Molecular Biology 416, 534-542.
10. Wang, X., Chen, X.J., and Yang, Y. (2012). Spatiotemporal control of gene expression by a light-switchable transgene system. Nature methods 9, 266-U264.
11. Yin, T., and Wu, Y.I. (2013). Guiding lights: recent developments in optogenetic control of biochemical signals. Pflugers Archiv : European journal of physiology 465, 397-408.
12. Zhou, X.X., Chung, H.K., Lam, A.J., and Lin, M.Z. (2012). Optical Control of Protein Activity by Fluorescent Protein Domains. Science 338, 810-814.




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