¿ÞÂÊ ¸Þ´º ŸÀÌƲ À̹ÌÁö
¿¬±¸µ¿Çâ

Home < ¿­¸°¸¶´ç < ¿¬±¸µ¿Çâ

       ¹ÚÅ׸®¾Æ ncRNA¸¦ ÅëÇÑ ½ºÆ®·¹½ºÀÇ Á¶Àý
       kimsg7596@kaist.ac.kr        
       ÀÌ¿µÈÆ ±³¼ö        2014.08.12 10:17        15199
´Ù¿î·Îµå : ¹ÚÅ׸®¾Æ ncRNA¸¦ ÅëÇÑ ½ºÆ®·¹½ºÀÇ Á¶Àý_ÀÌ¿µÈÆ ±³¼ö.pdf(297 Kb)
 

¹ÚÅ׸®¾Æ ncRNA¸¦ ÅëÇÑ ½ºÆ®·¹½ºÀÇ Á¶Àý

 

Çѱ¹°úÇбâ¼ú¿ø ÀÌ¿µÈÆ ±³¼ö

1. °³¿ä
¹ÚÅ׸®¾Æ´Â »ýÁ¸À» À§ÇùÇÏ´Â ¿©·¯ °¡Áö ȯ°æ½ºÆ®·¹½º¿¡ ³ëÃ⠵ǰí ÀÖ°í, À̸¦ ±Øº¹Çϱâ À§ÇÑ ´Ù¾çÇÑ ¹æ¹ýÀ» »ç¿ëÇÏ°í ÀÖ´Ù. ȯ°æº¯È­¸¦ °¨ÁöÇÏ¿© ´Ù¾çÇÑ À¯ÀüÀÚÀÇ ¹ßÇö°ú  ´Ü¹éÁúµéÀÇ È°¼ºÀ» Á¶ÀýÇÏ´Â ½ÅÈ£Àü´Þ ü°è¸¦ Á¶Á¤ÇÑ´Ù. ¼¼Æ÷³»ÀÇ RNA À¯Àüü (RNomics) ºÐ¼®À» ÅëÇؼ­, ´Ü¹éÁú ÇÕ¼ºÁ¤º¸¸¦ ¾ÏȣȭÇÏ°í ÀÖ´Â messenger RNAs (mRNAs)¿Ü¿¡ ¼¼Æ÷ ´ë»çÁ¶Àý¿¡ °ü¿©ÇÏ´Â ´Ù¾çÇÑ Å¸ÀÔÀÇ RNA°¡ Á¸ÀçÇÏ°í ÀÖÀ½ÀÌ ¾Ë·ÁÁ³´Ù. ´Ù¾çÇÑ ¹ÚÅ׸®¾Æ¿¡¼­ Áö¼ÓÀûÀ¸·Î ¹àÇôÁö°í ÀÖ´Â ÀÌ·¯ÇÑ ÀÛÀº ´Ü¹éÁúÀ» ÄÚµåÇÏÁö ¾Ê´Â ÀÛÀº RNA (small non-coding RNA: ncRNA)µéÀº ±âÁ¸ÀÇ 2°³-ÀÎÀÚ ½ÅÈ£Àü´Þ°è (two-component signal transduction system) ¹× ´Ù¾çÇÑ Á¶Àý´Ü¹éÁú°ú ´õºÒ¾î ½ºÆ®·¹½º ´ëÀÀÀ» À§ÇÑ Çٽɿä¼Ò·Î½áÀÇ ±â´ÉÀ» ÇÏ°í ÀÖ´Â °æ¿ì°¡ ¸¹¾Ò´Ù. (Abu-Qatouseh et al., 2010; Altuvia, 2007; Pichon and Felden, 2008; Raabe et al., 2011). °¡Àå Àß Á¤¸³µÇ¾î ÀÖ´Â ncRNAÀÇ ÀÛ¿ë±âÀÛÀº Ÿ°Ù mRNA¿Í ¿°±â½ÖÀ» ÀÌ·ç°Å³ª Ÿ°Ù ´Ü¹éÁú°ú »óÈ£ÀÛ¿ëÇÏ¿© RNA-´Ü¹éÁú º¹ÇÕü (Ribonucleoprotein; RNP)¸¦ ¸¸µå´Â °ÍÀÌ´Ù. ncRNA´Â À¯Àüü »ó¿¡ Á¸ÀçÇÏ´Â À§Ä¡¿¡ µû¶ó¼­ ½Ã½º·Î ÄÚµåµÇ´Â (cis-encoded) ncRNA ¶Ç´Â Æ®·£½º·Î ÄÚµåµÇ´Â (trans-encoded ncRNA)·Î ºÐ·ùÇϱ⵵ ÇÑ´Ù. ½Ã½º·Î ÄÚµåµÇ´Â ncRNA´Â Ÿ°Ù mRNAÀÇ ¹Ù·Î ¹Ý´ë°¡´Ú¿¡ Á¸ÀçÇÏ´Â ncRNA·Î Ÿ°Ù mRNA¿Í ¿ÏÀüÇÑ ¿°±â½ÖÀ» ÀÌ·ê ¼ö ÀÖ´Â ¹Ý¸é, Ÿ°Ù mRNA À¯ÀüÀÚ¿Í ¸Ö¸® ¶³¾îÁø À¯ÀüÀÚ¿¡¼­ Æ®·£½º·Î ÄÚµåµÇ´Â ncRNA´Â Ÿ°Ù mRNA¿Í ºÎºÐÀûÀÎ ¿°±â½ÖÀ» ÀÌ·é´Ù (Richards and Vanderpool, 2011). Æ®·£½º ÄÚµåµÇ´Â ncRNAÀÇ °æ¿ì, ±â´ÉÀ» À§Çؼ­ »þÆä·Ð ´Ü¹éÁúÀÎ Hfq°¡ ÇÊ¿äÇÑ °æ¿ì°¡ ¸¹´Ù. 102°³ ¾Æ¹Ì³ë»êÀ¸·Î ÀÌ·ç¾îÁø Hfq ´Ü¹éÁúÀº AU°¡ ¸¹Àº ´ÜÀÏ°¡´Ú¼º RNA¿Í »óÈ£ÀÛ¿ëÇÑ´Ù°í ¾Ë·ÁÁ® Àִµ¥ (Geissmann et al., 2006; Moller et al., 2002), ¹ÚÅ׸®¾Æ Á¾°£¿¡ º¸Á¸¼ºÀÌ ³ôÀ¸¸ç ½ºÆ®·¹½º ´ëÀÀÀ» Æ÷ÇÔÇÑ ´Ù¾çÇÑ ¼¼Æ÷±â´É¿¡ ¿µÇâÀ» ¹ÌÄ£´Ù. hfq µ¹¿¬º¯ÀÌ´Â ´Ù¾çÇÑ º´¿ø¼º ¹ÚÅ׸®¾ÆÀÇ º´¿ø¼ºÀ» ¸Å¿ì °¨¼Ò½ÃŲ´Ù. mRNAÀÇ 5¡Ç ¸»´Ü ºñ¹ø¿ª ¿µ¿ª (5¡Ç untranslated regions; 5¡Ç UTR)Àº ±¸Á¶º¯È­ µîÀ» ÅëÇؼ­ mRNAÀÇ ¹ßÇöÀ» Á¶ÀýÇÏ´Â °æ¿ì°¡ ¸¹Àºµ¥, ncRNA°¡ ÀνÄÇϴ Ÿ°Ù¿µ¿ª ¶ÇÇÑ 5¡Ç UTRÀÌ µÇ´Â °æ¿ì°¡ ¸¹¾Ò´Ù. º» ¿¬±¸µ¿Çâ¿¡¼­´Â ¹ÚÅ׸®¾Æ ncRNA°¡ ½ºÆ®·¹½º¿¡ ´ëÀÀÇϱâ À§ÇÏ À§ÇÏ¿© ¼¼Æ÷´ë»ç¿¡ ¾î¶»°Ô °ü¿©ÇÏ°í ÀÖ´ÂÁö¸¦ ¸î°¡Áö ½ÇÇè °á±«µéÀÇ ¿¹¸¦ ÅëÇØ »ìÆ캸°íÀÚ ÇÑ´Ù.

 

2. ¿Âµµ ½ºÆ®·¹½º¿Í ncRNA
¹ÚÅ׸®¾Æ´Â ¿Âµµº¯È­¿¡ ¹Î°¨ÇÏ°Ô ´ëÀÀÇϱâ À§ÇÑ º¹ÀâÇÑ Çǵå¹é Á¶Àý±âÀÛµéÀ» °¡Áö°í ÀÖ´Ù. ¡®RNA ¿Âµµ°è (RNA thermometers)¡¯¶ó°í ºÎ¸£´Â RNA ¿µ¿ªÀÇ ¿Âµµ¿¡ µû¸¥ ±¸Á¶º¯È­¸¦ ÅëÇÑ Çǵå¹é ½Ã½ºÅÛÀº, ¿Âµµº¯È­¿¡ ºü¸£°Ô ´ëÀÀÇÒ ¼ö ÀÖ´Ù. RNA ¿Âµµ°è´Â ÁÖ·Î 5¡Ç UTR ¿µ¿ª¿¡¼­ ¸®º¸Á» °áÇÕÀÚ¸®¸¦ Æó¼âÇÏ´Â ÇüÅ·Π2Â÷±¸Á¶¸¦ Çü¼ºÇÏ°í ÀÖ´Ù°¡, ³ôÀº ¿Âµµ¿¡¼­ ±¸Á¶°¡ º¯È­ÇÏ¿© ´Ü¹éÁú ¹ßÇöÀÌ °¡´ÉÇϵµ·Ï ÇÑ´Ù (Kortmann and Narberhaus, 2012). ROSE ¿ä¼Ò (Repression of the heat shock gene expression element: ROSE)´Â ¿­Ãæ°Ý ´Ü¹éÁú¿¡¼­ ÈçÇÏ°Ô ³ªÅ¸³ª´Â RNA ¿Âµµ°èÁß Çϳª·Î, SD ¼­¿­ (Shine-Dalgarno sequence) ±Ùó¿¡ U(U/C)GCU °øÅë¼­¿­À» °®°í 60~100 ntÀÇ ¿µ¿ªÀ¸·Î Á¸ÀçÇÑ´Ù. 2°³ ÀÌ»óÀÇ ¸Ó¸®ÇÉ ±¸Á¶°¡ Æ÷ÇԵǴµ¥, SD ¼­¿­À» Æ÷ÇÔÇÏ´Â 3¡Ç ¸Ó¸®Çɱ¸Á¶°¡ °í¿Â¿¡¼­ Ç®¾îÁö¸é¼­ mRNA ¹ø¿ªÀ» ÃËÁø½ÃŲ´Ù (Waldminghaus et al., 2009). 4U ¿ä¼Ò (fourU element)´Â Salmonella agsA (aggregation suppressing A) mRNAÀÇ 5¡Ç UTR¿¡ ³ªÅ¸³ª´Âµ¥, ³·Àº ¿Âµµ¿¡¼­ ³× °³ÀÇ ¿ì¸®µò ¿°±â¼­¿­ÀÌ SD ¼­¿­ÀÇ AGGA ¿°±â¼­¿­°ú ÇÔ²² ¸Ó¸®ÇÉ ±¸Á¶¸¦ Çü¼ºÇÑ´Ù. ¸¶Âù°¡Áö·Î ¿Âµµ°¡ ³ô¾ÆÁö¸é ±¸Á¶°¡ ºÒ¾ÈÁ¤ÇØÁö°í mRNAÀÇ ¹ßÇöÀÌ ÃËÁøµÈ´Ù.
RNA ¿Âµµ°è´Â Àú¿Â ½ºÆ®·¹½º¿¡¼­µµ Áß¿äÇÑ ±â´ÉÀ» ÇÑ´Ù. Àú¿Â¿¡¼­ ¹ßÇöµÇ¾î RNA »þÆä·ÐÀÇ ±â´ÉÀ» ÇÏ´Â CspAÀÇ °æ¿ì, Àú¿Â¿¡¼­ ¿ÏÀüÈ÷ ´Ù¸¥ ÇüÅÂÀÇ 5¡Ç UTR ±¸Á¶¸¦ °¡ÁüÀ¸·Î½á ¼¼Æ÷³» mRNAÀÇ ¾ÈÁ¤¼ºÀÌ Áõ°¡µÇ¾î ´Ü¹éÁú ¹ßÇöÀÌ Áõ°¡µÈ´Ù (Giuliodori et al., 2010).
Æ®·£½º·Î ÄÚµåµÇ´Â ncRNAÀÎ DsrA´Â µÎ°¡Áö Àü»çüÀÇ ÇüÅ·ΠÁ¸ÀçÇϴµ¥, ¿Âµµº¯È­¿¡ µû¶ó µÎ°¡Áö Àü»çüÀÇ ºñÀ²ÀÌ ´Þ¶óÁö°í, À̸¦ ÅëÇØ ½ºÆ®·¹½º ´ëÀÀÀ» À§ÇÑ RpoSÀÇ ¹ßÇöÀ» ÃËÁø½ÃŲ´Ù (Repoila and Gottesman, 2001).
Åä¾ç¹Ì»ý¹°ÀÎ L. monocytogenesÀÇ prfA mRNAÀÇ °æ¿ì, »ç¶÷ÀÇ Ã¼¿ÂÀÎ 37¡É¿¡¼­ 5¡ÇUTRÀÇ ±¸Á¶°¡ º¯È­µÇ¸é ¹ßÇöÀÌ ÃËÁøµÇ°í º´¿ø¼ºÀ» ÀÏÀ¸Å°´Â ¿©·¯ °¡Áö À¯ÀüÀÚ¸¦ ¹ßÇö½ÃÅ°°Ô µÈ´Ù (Johansson et al., 2002).

±×¸² 1. ¿Âµµ ½ºÆ®·¹½º¸¦ ±Øº¹Çϱâ À§ÇÑ ncRNAÀÇ ÀÛ¿ë±âÀÛ. agsA mRNA´Â RNA ¿Âµµ°èÀÎ 4U ¿ä¼Ò¸¦ ÅëÇؼ­ ³·Àº ¿Âµµ¿¡¼­´Â ¸®º¸Á»ÀÇ °áÇÕÀÌ ÀúÇصǰí, ³ôÀº ¿Âµµ¿¡¼­´Â ÁöÆÛ¿Í À¯»çÇÑ ¸ÞÄ¿´ÏÁòÀ¸·Î ºÒ¾ÈÁ¤È­µÇ¾î ¸®º¸Á»ÀÇ °áÇÕÀÌ °¡´ÉÇØÁö°í À¯ÀüÀÚ°¡ ¹ßÇöµÈ´Ù.

 

3. ´ë»ç»ê¹°/¾çºÐ ½ºÆ®·¹½º¿Í ncRNA
¹Ì»ý¹°µéÀº Á¾Á¾ ¿µ¾ç°áÇÌÀ¸·Î ÀÎÇÑ ¼ºÀåÁ¤Ã¼»óÅ·Π¹ß°ßµÈ´Ù. ¼¼Æ÷³»¿¡ ±Û·çÄÚ¿À½º-6-Àλê (glucose-6-phosphate)ÀÌ ÃàÀûµÇ¸é ¼¼Æ÷¼ºÀåÀÌ ¸ØÃß°í Á×À½¿¡ À̸£±âµµ ÇÑ´Ù (Englesberg et al., 1962). ÀÌ·¯ÇÑ ½ºÆ®·¹½º »óÅ¿¡¼­ Hfq-°áÇÕ ncRNA Áß ÇϳªÀÎ SgrS ncRNA°¡ ¹ßÇöµÇ°í, ±Û·çÄÚ¿À½º ¿î¹Ýü ÁßÀÇ Çϳª¸¦ ÄÚµåÇÏ´Â ptsG mRNA¿Í »óÈ£ÀÛ¿ëÇÏ¿© ¸®º¸Çٻ갡¼öºÐÇØÈ¿¼Ò E (RNase E) ÀÇÁ¸ÀûÀÎ ¹æ½ÄÀ¸·Î ptsG mRNA¸¦ ºÐÇØÇÑ´Ù (Maki et al., 2010; Morita et al., 2005). SgrS ncRNA´Â SgrT¶ó´Â ªÀº ÆéŸÀ̵带 ¹ßÇöÇϱ⵵ Çϴµ¥, SgrT´Â PtsG ´Ü¹éÁúÀÇ ±â´ÉÀ» ÀúÇØÇÑ´Ù (Wadler and Vanderpool, 2011).
ź¼ÒÀúÀå Á¶Àý½Ã½ºÅÛÀÇ ÇÙ½É ¿ªÇÒÀ» ÇÏ´Â CsrA ´Ü¹éÁúÀÇ È°¼ºÀº BarA/UvrY 2°³-ÀÎÀÚ ½ÅÈ£Àü´Þ°è¿¡ ÀÇÇؼ­ °£Á¢ÀûÀ¸·Î Á¶ÀýµÈ´Ù. ź¼Ò°áÇÌ»óȲ¿¡¼­ UvrY¿¡ ÀÇÇØ µÎ°¡Áö ncRNA, CsrB¿Í CsrC°¡ ºü¸£°Ô ¹ßÇöµÇ´Âµ¥, ÀÌ µÎ°¡Áö ncRNA´Â ´Ù¼öÀÇ CsrA ´Ü¹éÁú °áÇÕÀÚ¸®¸¦ °®°í À־ CsrA ´Ü¹éÁú°ú °áÇÕÇÔÀ¸·Î½á ÇØ´çÀÛ¿ë¿¡ °ü¿©ÇÏ´Â À¯ÀüÀÚÀÇ ¹ßÇöÀ» ÀúÇØÇÏ°í, ±Û·çÄÚ¿À½º ½Å»ýÇÕ¼º¿¡ °ü¿©ÇÏ´Â À¯ÀüÀÚµéÀ» ¹ßÇöÇÑ´Ù (Pernestig et al., 2003).
Caulobacter crescentusÀÇ CrfA ncRNA´Â ź¼Ò°áÇÌ »óȲ¿¡¼­ ¹ßÇöµÇ¾î CC3461 ¼ö¿ë±â mRNAÀÇ 5¡Ç UTR°ú ¿°±â½ÖÀ» ÀÌ·ç¾î, mRNAÀÇ ¼¼Æ÷³» ¾ÈÁ¤¼ºÀ» Áõ°¡½ÃÄѼ­ ´õ ´Ù¾çÇÑ Á¾·ùÀÇ Åº¼Ò¿øÀÌ ¼¼Æ÷³»·Î À¯À﵃ ¼ö ÀÖµµ·Ï ÇÑ´Ù (Landt et al., 2010).
Staphylococcus, Macrococcus, Bacillusµî¿¡¼­ Àß º¸Á¸µÇ¾î ÀÖ´Â RsaE ncRNA´Â ¾çºÐÀÌ ºÎÁ·ÇØÁö´Â Áö¼ö¼ºÀå±â ÈĹݺο¡¼­ ¹ßÇöÀÌ Áõ°¡µÈ´Ù. 17 ntÀÇ ´ÜÀÏ°¡´ÚºÎºÐ ¾çÂÊÀ¸·Î ¸Ó¸®Çɱ¸Á¶¸¦ Çü¼ºÇÏ°í ÀÖ´Â ÀÌ ncRNA´Â, opp3 ¿ÀÆä·ÐÀÇ Ã³À½°ú ¸¶Áö¸· mRNAÀÎ opp3B, opp3A¿Í ¿°±â½ÖÀ» ÀÌ·ç¾î ¹ßÇöÀ» ÀúÇؽÃÄѼ­ ¿µ¾ç°áÇÌ»óÅ¿¡ Àß ÀûÀÀÇÒ ¼ö ÀÖµµ·Ï ÇÑ´Ù (Bohn et al., 2010).
¾Æ¹Ì³ë»êÀÇ ´ë»ç´Â ÇÙ»êÀ̳ª È¿¼Ò º¸Á¶ÀÎÀÚµîÀÇ »ý¼º¿¡ ÇʼöÀûÀ̱⠶§¹®¿¡ Á¤È®ÇÏ°Ô Á¦¾îµÉ ÇÊ¿ä°¡ ÀÖ´Ù. Gcv ¿ÀÆä·Ð (glycine cleavage operon)ÀÎ gcvTHP ¿ÀÆä·ÐÀº ±Û¶óÀ̽ÅÀ» ´ë»ç½ÃÄÑ ¼¼Æ÷³» ź¼Ò¿ø»ýÇÕ¼º¿¡ ÀÌ¿ëµÇ´Â 5,10-¸ÞÆ¿·»Å×Æ®¶óÇÏÀ̵å·Î¿±»êÀ» »ý¼ºÇϴµ¥ ÇʼöÀûÀÌ´Ù. ¼¼Æ÷³»¿¡ ±Û¶óÀ̽Š³óµµ°¡ ³ô¾ÆÁö¸é Hfq¿Í »óÈ£ÀÛ¿ëÇÏ´Â GcvB ncRNAÀÇ ¹ßÇöÀÌ È°¼ºÈ­µÇ°í ªÀº ÆéŸÀ̵峪, ´Ù¸¥ ¾Æ¹Ì³ë»ê, µ¶¼º¹°Áú°ú Ç×»ýÁ¦µîÀÇ ¼¼Æ÷À¯ÀÔ¿¡ °ü¿©µÇ´Â À¯ÀüÀÚµéÀÇ ¹ßÇöÀ» ¾ïÁ¦ÇÑ´Ù (Urbanowski et al., 2000). GcvB ncRNA´Â mRNA Ÿ°ÙµéÀ» Á¶ÀýÇÏ´Â µÎ°¡Áö ¾ÈƼ¼¾½º¿µ¿ªÀ» °¡Áö°í Àִµ¥, ~1%ÀÇ Salmonella mRNA°¡ G/U°¡ ¸¹Àº ¿µ¿ªÀ» ÅëÇØ Á¶ÀýµÊÀÌ º¸°íµÇ¾ú°í, cycA mRNA´Â µÎ°¡Áö ¾ÈƼ¼¾½º¿µ¿ªÀ» ¸ðµÎ »ç¿ëÇÏ¿© Á¶ÀýµÈ´Ù. À̴ Ÿ°Ù mRNA¿Í ¿°±â½ÖÀ» ÀÌ·ç´Â À¯¿¬¼ºÀ» ³ªÅ¸³»ÁÖ´Â ¿¹·Î ¾ÕÀ¸·Î ´õ ´Ù¾çÇÑ Å¸°Ù mRNA°¡ ã¾ÆÁú ¼ö ÀÖÀ½À» ÀǹÌÇÑ´Ù.

±×¸² 2. ÀϹÝÀûÀÎ ¼ºÀåÁ¶°Ç¿¡¼­ CsrA ´Ü¹éÁúÀº ÇØ´çÀÛ¿ë¿¡ ÇÊ¿äÇÑ À¯ÀüÀÚ¸¦ ¹ßÇö½ÃÅ°°í, ±Û·çÄÚ¿À½º ½Å»ýÇÕ¼º¿¡ °ü·ÃµÈ À¯ÀüÀÚµéÀÇ ¹ßÇöÀ» ÀúÇØÇÑ´Ù. ź¼Ò¿øÀÌ ºÎÁ·ÇÑ ½ºÆ®·¹½º »óȲ¿¡¼­ CsrB/CsrC ncRNA´Â CsrA¿¡ °áÇÕÇÏ¿© CsrAÀÇ È°µ¿À» ¹æÇØÇÑ´Ù. ½ºÆ®·¹½º»óȲ¿¡¼­ ¹þ¾î³ª¸é CsrB/CsrC´Â CsrD¿Í RNase E¸¦ ¸Å°³·ÎÇÏ¿© ºÐÇصȴÙ.

 

4. ¼¼Æ÷¸· ½ºÆ®·¹½º¿Í ncRNA
±×¶÷À½¼º ¹ÚÅ׸®¾ÆÀÇ ¼¼Æ÷¿Ü¸· (outer membrane; OM)Àº µ¶¼º¹°ÁúÀÇ À¯ÀÔÀ» ¸·´Â Áß¿äÇÑ ¿ªÇÒÀ» ÇÑ´Ù. Æ÷¸° ´Ü¹éÁúµéÀº OM¿¡ Åë·Î¸¦ ¸¸µé¾î ¾çºÐÀ» Èí¼öÇÏ°í Æó±â¹°À» ¹èÃâÇÑ´Ù. E. coli¿Í Salmonella¿¡¼­´Â ¥òE ÀÎÀÚ ½ÅÈ£Àü´Þü°è¸¦ ÅëÇؼ­ OM ´Ü¹éÁúµéÀÇ »óÅ°¡ °¨½ÃµÈ´Ù. À߸ø Çü¼ºµÈ OM ´Ü¹éÁúµé (¼¼Æ÷¸· ½ºÆ®·¹½º¸¦ ¾ß±â)ÀÌ ¸¹¾ÆÁö¸é ¥òE ÀÎÀÚ°¡ È°¼ºÈ­µÈ´Ù. Àå³»¹ÚÅ׸®¾Æ¿¡¼­ º¸Á¸¼ºÀÌ ¸Å¿ì ³ôÀº OmpA ´Ü¹éÁúÀº ¼ºÀåÁ¤Áö±â¿¡¼­ Æ®·£½º·Î ÄÚµåµÇ´Â ncRNAÀÎ MicA¿¡ ÀÇÇؼ­ RNase E ÀÇÁ¸ÀûÀ¸·Î ±Þ°ÝÈ÷ ¹ßÇöÀÌ ÀúÇصȴ٠(Rasmussen et al., 2005). ¥òE ÀÎÀÚ¿¡ ÀÇÇØ ¹ßÇöµÇ´Â ¶Ç ´Ù¸¥ ncRNAÀÎ RybB ncRNA ¶ÇÇÑ ´Ù¼öÀÇ OM ´Ü¹éÁúµéÀÇ ¹ßÇöÀ» Á¶ÀýÇÏ¿© ¼¼Æ÷¸· ½ºÆ®·¹½º »óȲ¿¡¼­ OM ´Ü¹éÁúµéÀÇ ¾çÀ» º¯È­½ÃŲ´Ù (Papenfort et al., 2010). CyaR ncRNA´Â ompX mRNAÀÇ ¹ø¿ªÀ» ¾ïÁ¦Çϸç, V. choleraeÀÇ VrrA ncRNAµµ OmpAÀÇ ¹ßÇöÀ» ÀúÇؽÃÅ°°í, OMV (outer membrane vesicle)ÀÇ »ý¼ºµµ ÃËÁø½ÃÄÑ ¼¼Æ÷¸· ½ºÆ®·¹½º¸¦ ´õ¿í ¿ÏÈ­ÇÑ´Ù.

±×¸² 3. Æ®·£½º·Î ÄÚµåµÇ´Â ncRNAÀÎ VrrA´Â ¼¼Æ÷¸· ½ºÆ®·¹½º¿¡ ´ëÀÀÇÏ¿© ¹ßÇöÀÌ À¯µµµÈ´Ù. VrrA ncRNA´Â ompT, tcpA, ompA mRNAÀÇ ¸®º¸Á» °áÇÕÀÚ¸®¿µ¿ª¿¡ °áÇÕÇÏ¿© À¯ÀüÀÚ ¹ßÇöÀ» ÀúÇØÇÑ´Ù.

 

5. »êÈ­½ºÆ®·¹½º¿Í ncRNA
OxyR Àü»çÀÎÀÚ´Â H2O2¿¡ ÀÇÇؼ­ Ƽ¿Ã-ÀÌȲȭ »êȭȯ¿ø (thiol-disulfide redox) ½ºÀ§Ä¡¸¦ ÅëÇØ È°¼ºÈ­µÇ¾î, Ä«Å»¶ó¾ÆÁ¦ (catalase), ¼ö»êÈ­°ú»êÈ­¹° ȯ¿øÈ¿¼Ò, ÃÊ»êÈ­¹° ºÒ±ÕµîÈ­È¿¼Ò (superoxide dismutase) µîÀÇ ¹ßÇöÀ» Áõ°¡½ÃÄѼ­ »êÈ­½ºÆ®·¹½º¿¡¼­ ¼¼Æ÷¸¦ º¸È£ÇÑ´Ù (Pomposiello and Demple, 2001). OxyS ncRNA´Â »êÈ­½ºÆ®·¹½º·Î ÀÎÇؼ­ ¹ßÇöÀÌ À¯µµµÇ¾î ¸¹Àº À¯ÀüÀÚµéÀÇ ¹ßÇöÀ» Á¶ÀýÇÏ°í µ¹¿¬º¯ÀÌ°¡ À¯¹ßµÇ´Â °ÍÀ» ¾ïÁ¦ÇÑ´Ù (Altuvia et al., 1997).
¸¹Àº º´¿ø¼º ¹ÚÅ׸®¾ÆµéÀº ¼÷ÁÖÀÇ ´ë½Ä¼¼Æ÷¾È¿¡¼­ »êÈ­½ºÆ®·¹½º¿¡ ³ëÃâµÇ°Ô µÇ´Âµ¥, B11, B55, F6, ASpks¿Í °°Àº ncRNAµéÀÌ °¨¿° Ãʱ⿡ ¹ßÇöµÇ¾î¼­ ´Ù¾çÇÑ À¯ÀüÀÚÀÇ ¹ßÇöÀ» Á¶ÀýÇÑ´Ù (Arnvig and Young, 2009).

 

6. öºÐºÎÁ· ½ºÆ®·¹½º¿Í ncRNA
öºÐÀº ¼¼Æ÷¼ºÁ¤°ú ºÐÈ­, ÀüÀÚÀü´Þ°è, »ê¼Ò¿î¹Ý, ¼¼Æ÷´ë»ç, Çص¶µîÀÇ ¼¼Æ÷È°µ¿¿¡ ¹Ýµå½Ã ÇÊ¿äÇÑ ¿ä¼ÒÀÌ´Ù. ÇÏÁö¸¸ ³ôÀº ³óµµÀÇ Ã¶ºÐÀº È°¼º»ê¼ÒÀÇ Çü¼ºÀ» À¯¹ßÇÏ¿© ¼¼Æ÷¿¡ Çظ¦ ³¢Ä¡±âµµ ÇÑ´Ù. ´ëºÎºÐÀÇ ¹ÚÅ׸®¾Æ¿¡¼­ Fur (ferric uptake regulator) ´Ü¹éÁúÀº öºÐÀÇ Ç×»ó¼º À¯Áö¿¡ Á߽ɱâ´ÉÀ» ¸Ã°í ÀÖ´Ù. öºÐÀÌ ÃæºÐÇÒ ¶§´Â öºÐ Èí¼ö À¯ÀüÀÚµéÀ» ¾ïÁ¦ÇÏ°í, öºÐ ÀúÀå ´Ü¹éÁúµéÀÇ ¹ßÇöÀ» Áõ°¡½ÃŲ´Ù. RyhB ncRNA´Â öºÐºÎÁ· ½ºÆ®·¹½º¿¡¼­ ¹ßÇöµÇ´Â ncRNA·Î Fur ´Ü¹éÁú¿¡ ÀÇÇØ ¹ßÇöÀÌ Á¶ÀýµÈ´Ù. öºÐÀÌ Ç³ºÎÇÒ ¶§ RyhB ncRNAÀÇ ¹ßÇöÀÌ À¯µµµÇ°í sdhCDAB, acnA, fumA, bfr, ftnA, sodB mRNA µî öºÐ´ë»ç¿Í °ü¿©µÈ 18°³ ¿ÀÆä·Ð, 56°³ ´Ü¹éÁúÀÇ ¹ßÇöÀ» Á¶ÀýÇÑ´Ù (Masse et al., 2005). RyhB ncRNA´Â fur mRNAÀÇ ¹ßÇöµµ ¾ïÁ¦ÇÏ´Â À½¼ºÇǵå¹é½Ã½ºÅÛµµ º¸¿©Áشٴ Á¡ÀÌ ÁÖ¸ñÇÒ ¸¸ÇÏ´Ù (Vecerek et al., 2007).
V. cholerae, S. flexneri, S. dysenteriae µî¿¡¼­ RyhB ncRNAÀÇ »óµ¿Ã¼°¡ ¹ß°ßµÇ°í ÀÖ´Ù. ±× ±â´Éµµ ´õ ´Ù¾çÇÏ°Ô ¹àÇôÁ®¼­ ¹ÙÀÌ¿ÀÇʸ§ Çü¼ºÀ̳ª ÁÖÈ­¼º ¿òÁ÷ÀÓ¿¡µµ °ü¿©ÇÏ°í ÀÖ°í (Davis et al., 2005; Mey et al., 2005), ³»»ê¼ºÀÇ °¨¼Ò¿Í º´¿ø¼º ¾ïÁ¦¿¡µµ °ü¿©ÇÑ´Ù (Murphy and Payne, 2007).

 

7. pH ½ºÆ®·¹½º¿Í ncRNA
À§»êÀÇ ±ØÇѻ꼺Á¶°ÇÀ» °ßµ®³»´Â °ÍÀº ¹ÚÅ׸®¾ÆµéÀÌ Àå³»¿¡ Á¤ÂøÇϱâ À§ÇØ Áß¿äÇÏ´Ù. ¸î¸î ±×¶÷¾ç¼º±Õ ¹× ±×¶÷À½¼º±ÕÀº ±Û·çŽ»êÀ» ÀÌ¿ëÇÑ ³»»ê¼º ½Ã½ºÅÛ (gad system)À» °¡Áö°í À־ ³ôÀº ³óµµÀÇ H+ ½ºÆ®·¹½º·ÎºÎÅÍ ¼¼Æ÷¸¦ º¸È£ÇÒ ¼ö ÀÖ´Ù. GadA, GadB µÎ °³ÀÇ ±Û·çŽ»ê Żī¸£º¹½ÇÈ­È¿¼Ò´Â ±Û·çŽ»êÀ» ÀÌ¿ëÇØ ¥ã-¾Æ¹Ì³ëºÎÆ¿»êÀ» ¸¸µé¸é¼­ H+¸¦ Á¦°ÅÇÑ´Ù. GadE, GadX, GadW, CRP, H-NS, ¥òS µîÀÌ º¹ÀâÇÏ°Ô °ü¿©ÇÏ¿© ³»»ê¼º½Ã½ºÅÛÀ» Á¶ÀýÇÑ´Ù. E. coli¿¡¼­ Hfq¿Í »óÈ£ÀÛ¿ëÇÏ´Â GadY ncRNA´Â gadX mRNAÀÇ 3¡Ç ¸»´ÜºÎÀ§¿¡ ½Ã½º·Î ÄÚµåµÇ´Â ncRNA·Î gadX mRNAÀÇ 3¡Ç ºÎÀ§¿¡ »óº¸ÀûÀ¸·Î °áÇÕÇÏ¿© mRNA¸¦ RNase EÀÇ ºÐÇطκÎÅÍ º¸È£ÇÑ´Ù (Opdyke et al., 2004). GadXÀÇ ÃàÀûÀº ÇÏÀ§ÀÇ ³»»ê¼º °ü·ÃÀ¯ÀüÀÚ¸¦ ¹ßÇö½ÃÄÑ ³·Àº pH¿¡¼­ »ýÁ¸¼ºÀ» Áõ°¡½ÃŲ´Ù (Opdyke et al., 2011). GcvB ncRNA ¶ÇÇÑ ¾Ë ¼ö ¾ø´Â ±âÀÛÀ¸·Î ¥òS ÀÇ ¹ßÇöÀ» Áõ°¡½ÃÄÑ ³·Àº pH¿¡¼­ÀÇ »ýÁ¸¼ºÀ» Áõ°¡½ÃÅ°¸ç (Jin et al., 2009), ArcZ, DsrA, RprA ncRNA¸¦ ÅëÇÑ ¥òS ÀÇ ¹ßÇöÁõ°¡ ¿ª½Ã pH 2.0¿¡¼­ÀÇ ¼¼Æ÷ÀÇ »ýÁ¸¼º¿¡ Å« ¿µÇâÀ» ¹ÌÄ£´Ù (Bak et al., 2014). ³ôÀº pH¿¡¼­´Â ¾Æ¹Ì³ë»ê Å»¾Æ¹ÎÈ­¿Í ´ç¹ßÈ¿¸¦ ÅëÇÑ »ê »ý¼ºÁõ°¡, ¼¼Æ÷¸·ÀÇ Æ¯¼ºº¯È­ µîÀ» ÅëÇØ ½ºÆ®·¹½º¸¦ ±Øº¹ÇÑ´Ù. alx mRNAÀÇ 5¡Ç UTR¿¡´Â PRE ¸®º¸ ½ºÀ§Ä¡ (pH responsive RNA element)°¡ À־ ³ôÀº pH¿¡¼­¸¸ Alx ´Ü¹éÁúÀ» ¹ßÇö½ÃŲ´Ù (Nechooshtan et al., 2009).


 8. CRISPR: ½ºÆ®·¹½º °ü¸®¸¦ À§ÇÑ »õ·Î¿î ±âÀÛ
¹ÚÅ׸®¾Æ´Â ¹ÙÀÌ·¯½º, ÆÄÁö, ÇÃ¶ó½º¹Ìµå µîÀÇ Ä§ÀÔÀ» °ßµ®³»±â À§Çؼ­ ´Ù¾çÇÑ Àü·«µéÀ» »ç¿ëÇØ¿Ô´Ù. ±×Áß CRISPR/Cas ½Ã½ºÅÛ (clustered regularly interspaced short palindromic repeats/CRISPR-associated proteins)Àº ncRNA¸¦ ±â¹ÝÀ¸·Î ÇÑ ¹æ¾î½Ã½ºÅÛÀÌ´Ù (Barrangou and Horvath, 2012; Makarova et al., 2011; Westra et al., 2012). CRISPR À¯ÀüÀÚ ÀÚ¸®´Â ¸¹°Ô´Â ¼ö¹é°³ÀÇ °£°Ý ¶ç¿ì°Ô ¿°±â¼­¿­µé (spacer sequences)ÀÌ ¹Ýº¹¼­¿­ (repeat sequence) »çÀÌ·Î ±ÔÄ¢ÀûÀ¸·Î ¹è¿­µÇ¾î Àִ Ư¼ºÀ» º¸ÀδÙ. CRISPR ¿µ¿ª¿¡¼­ Àü»çµÈ ¼±±¸ crRNA´Â °¡°øµÇ¾î¼­ ¿Ï¼ºµÈ crRNA°¡ µÇ°í, ÁøÇà»ý¹°ÀÇ RNAi ½Ã½ºÅÛ°ú À¯»çÇÑ ¹æ½ÄÀ¸·Î Cas ´Ü¹éÁúµé°ú ÇÔ²² ¿ÜºÎ¿¡¼­ À¯·¡ÇÑ RNA ¶Ç´Â DNA¸¦ Àý´ÜÇÑ´Ù. CRISPR/Cas ½Ã½ºÅÛÀº °í¼¼±Õ·ù¿¡¼­´Â Ç×»ó Áö¼ÓÀûÀ¸·Î ¹ßÇöµÇ°í ÀÖÀ¸³ª (Hale et al., 2008; Tang et al., 2002), ƯÁ¤ ½ÅÈ£¿¡ ÀÇÁ¸ÀûÀÎ °æ¿ìµµ ¸¹´Ù. Thermus thermophilusÀÇ CRISPR/Cas Àü»çü´Â ÆÄÁö¿¡ °¨¿°µÈ ÈÄ¿¡ ¹ßÇöÀÌ Áõ°¡µÇ°í, ´ëÀå±Õ¿¡¼­´Â H-NS¿¡ ÀÇÇØ casAÀÇ ¹ßÇöÀÌ Á¶ÀýµÇ°í Àֱ⵵ ÇÏ´Ù (Pul et al., 2010). ¼¼Æ÷¸· ½ºÆ®·¹½º¿¡¼­ À¯µµµÇ´Â BaeSR 2°³-ÀÎÀÚ Á¶Àý½Ã½ºÅÛ¿¡ ÀÇÇؼ­ CRISPR/Cas ¹ßÇöÀ» ÃËÁø½ÃÅ°´Â °æ¿ìµµ º¸°íµÇ¾ú´Ù (Perez-Rodriguez et al., 2011). ¾î¶² ´Ù¸¥ ½ºÆ®·¹½ºµéÀÌ CRISPR/Cas ½Ã½ºÅÛÀÇ ¹ßÇöÁ¶Àý°ú ¾î¶² ±âÀÛÀ¸·Î ¿¬°áµÇ¾îÀÖ´ÂÁö ¾ÆÁ÷ ¹àÇôÁöÁö ¾Ê¾ÒÁö¸¸, ¾ÕÀ¸·Î ½ºÆ®·¹½º¿Í CRISPR/Cas ¹ßÇö»çÀÌ¿¡ ¸¹Àº ¿¬°üÀÌ ¹àÇôÁú °ÍÀ¸·Î ±â´ëµÈ´Ù.

 

 9. °íÂû
ÃÖ±Ù ncRNA ¿¬±¸ºÐ¾ßÀÇ ¼º°úµéÀº ½ºÆ®·¹½º Á¶Àý°ú ¹ÚÅ׸®¾Æ º´¿ø¼º µî¿¡ ncRNA°¡ ¾î¶»°Ô °ü¿©ÇÏ°í ÀÖ´ÂÁö¸¦ ÁüÀÛÄÉ ÇÑ´Ù. RNA ¿Âµµ°è, ½Ã½º ¶Ç´Â Æ®·£½º·Î ÄÚµåµÇ´Â ncRNA ¹× CRISPR µîÀº ´Ù¾çÇÑ ½ºÆ®·¹½º¿¡ ´ëÀÀÇÏ¿© ±¸Á¶ÀûÀ¸·Î º¯È­µÇ°Å³ª, Ÿ°Ù mRNA·Î ºÎÅÍ ¹ßÇö ¶Ç´Â ´Ü¹éÁúÀÇ È°¼ºÀ» Á¶ÀýÇÑ´Ù. ncRNA¸¦ È°¿ëÇÑ ½ºÆ®·¹½ºÀÇ ´ëÀÀÀº ´Ü¹éÁú Àü»çÀÎÀÚ¸¦ ÀÌ¿ëÇÑ ½ºÆ®·¹½º ´ëÀÀ°ú ºñ±³µÇ´Â Á¡µéÀÌ ÀÖ´Ù. ncRNA´Â ª°í ´Ü¹éÁú·Î ¹ø¿ªµÇÁö ¾Ê±â ¶§¹®¿¡ ¼¼Æ÷³» ¿¡³ÊÁö ¼Ò¸ð°¡ Àû°í, ÇÕ¼º°ú ºÐÇØ°¡ ºü¸£´Ù. ¶ÇÇÑ ncRNAµéÀº Àü»çÈÄ ´Ü°è¿¡¼­ Ÿ°ÙµéÀ» Á¶ÀýÇÏ´Â °æ¿ì°¡ ¸¹¾Æ¼­ ½ºÆ®·¹½º ´ëÀÀ¿¡ ´õ ½Å¼ÓÇÒ ¼ö ÀÖ´Ù. ¶ÇÇÑ ncRNAµéÀº Ãß°¡ÀûÀÎ Á¶Àý´Ü°è¸¦ °®°Ô ÇÏ´Â ±â´Éµµ ÇÑ´Ù. ncRNA¿Í Àü»çÀÎÀڴܹéÁúÀÌ ¸ðµÎ °ü¿©ÇÏ´Â Á¶Àý½Ã½ºÅÛÀÇ °æ¿ì ¹ßÇöÀÇ ´©¼ö°¡ ÃÖ¼ÒÈ­µÇ°í µ¿¿ªÇÐÀûÀ¸·Îµµ Àü»çÀÎÀÚ Çϳª¸¸ »ç¿ëÇÏ´Â °Íº¸´Ù ÀåÁ¡ÀÌ ¸¹´Ù (Levine and Hwa 2008).
´ë¿ë·® ½ÇÇè±â¼ú°ú, »ý¹°Á¤º¸ÇÐÀû ±â¹ýµéÀÇ ¹ß´Þ°ú ÇÔ²² ¹ÚÅ׸®¾Æ ½ºÆ®·¹½ºÁ¶Àý¿¡ °ü¿©ÇÏ´Â ´Ù¾çÇÑ ncRNAµéÀÌ ´õ ¸¹ÀÌ ¹ß°ßµÉ °ÍÀ¸·Î º¸ÀδÙ. ¾à¹° À¯»çü¿¡ ´ëÇÑ ¸®º¸½ºÀ§Ä¡¸¦ È°¿ëÇÏ¿© Áã¿¡¼­ S. aureusÀÇ º´¿ø¼ºÀ» °¨¼Ò½ÃÅ°°Å³ª (Mulhbacher et al., 2010), Cas9°ú ÇÕ¼º °¡À̵å ncRNA¸¦ ÀÌ¿ëÇÑ CRISPR ½Ã½ºÅÛÀ» ¿°»öü Á¶ÀÛ¿¡ È°¿ëÇϰųª (Cong et al., 2013; Jinek et al., 2012; Mali et al., 2013)ÇÏ´Â »õ·Î¿î ±â¼úµé·Îµµ ¸¹ÀÌ Àû¿ëµÇ°í ÀÖ´Ù. ´õ ³ª¾Æ°¡ ncRNA¸¦ Áø´Ü ¸¶Ä¿³ª ¾à¹° Ÿ°ÙÀ¸·Î ÇÏ¿© ¹ÚÅ׸®¾ÆÀÇ º´¿ø¼ºÀ» Á¶ÀýÇÏ´Â ¿¬±¸°¡ »õ·Î¿î µµÀüÀÌ µÉ °ÍÀ¸·Î »ý°¢µÈ´Ù.

 

Âü°í¹®Çå

1. Abu-Qatouseh, L.F., Chinni, S.V., Seggewiss, J., Proctor, R.A., Brosius, J., Rozhdestvensky,

   T.S., Peters, G., Von Eiff, C., Becker, K., 2010. Identification of differentially expressed small 

   non-protein-coding RNAs in Staphylococcus aureus displaying both the normal and the

   small-colony variant phenotype. J. Mol. Med. 88, 565–575.

2. Altuvia, S., 2007. Identification of bacterial small non-coding RNAs: experimental

   approaches. Curr. Opin. Microbiol. 10, 257–261.

3. Altuvia, S., Weinstein-Fischer, D., Zhang, A., Postow, L., Storz, G., 1997. A small, stable

   RNA induced by oxidative stress: role as a pleiotropic regulator and antimutator.

   Cell 90, 43–53.

4. Arnvig, K.B., Young, D.B., 2009. Identification of small RNAs in Mycobacterium tuberculosis.

    Mol. Microbiol. 73, 397–408.

5. Bak, G., Han, K., Kim, D., Lee, Y., 2014. Roles of rpoS-activating small RNAs in pathways

   leading to acid resistance of Escherichia coli. MicrobiologyOpen. 3, 15-28.

6. Barrangou, R., Horvath, P., 2012. CRISPR: new horizons in phage resistance and strain

    identification. Annu. Rev. Food Sci. Technol. 3, 143–162.

7. Bohn, C., Rigoulay, C., Chabelskaya, S., Sharma, C.M., Marchais, A., Skorski, P., Borezée-

    Durant, E., Barbet, R., Jacquet, E., Jacq, A., Gautheret, D., Felden, B., Vogel, J., Bouloc, P.,

    2010. Experimental discovery of small RNAs in Staphylococcus aureus reveals a

    riboregulator of central metabolism. Nucleic Acids Res. 38, 6620–6636.

8. Cong, L., Ran, F.A., Cox, D., Lin, S., Barretto, R., Hsu, N.H.P.D., Wu, X., Jiang, W.,

   Marraffini, L.A., Zhang, F., 2013. Multiplex genome engineering using CRISPR/Cas systems.

   Science.

9. Davis, B.M., Quinones, M., Pratt, J., Ding, Y., Waldor, M.K., 2005. Characterization of the

    small untranslated RNA RyhB and its regulon in Vibrio cholerae. J. Bacteriol. 187, 4005–4014.

10. Englesberg, E., Anderson, R.L., Weinberg, R., Lee, N., Hoffee, P., Huttenhauer, G., Boyer,

     H., 1962. L-arabinose-sensitive, l-ribulose 5-phosphate 4-epimerase-deficient mutants of

     Escherichia coli. J. Bacteriol. 84, 137.

11. Geissmann, T., Possedko, M., Huntzinger, E., Fechter, P., Ehresmann, C., Romby, P.,

     2006. Regulatory RNAs as mediators of virulence gene expression in bacteria. Handb. Exp.

     Pharmacol., 9–43.

12. Giuliodori, A.M., Di Pietro, F., Marzi, S., Masquida, B., Wagner, R., Romby, P., Gualerzi,

     C.O., Pon, C.L., 2010. The cspA mRNA is a thermosensor that modulates translation of the

     cold-shock protein CspA. Mol. Cell 37, 21–33.

13. Hale, C., Kleppe, K., Terns, R.M., Terns, M.P., 2008. Prokaryotic silencing (psi)RNAs in

     Pyrococcus furiosus. RNA 14, 2572–2579.

14. Hoe, C.-H., Raabe, C.A., Rozhdestvensky, T. S., Tang, T.-H., 2013. Bacterial sRNAs:

     Regulation in stress. Int J Med Microbiol. 303, 217-229.

15. Jinek, M., Chylinski, K., Fonfara, I., Hauer, M., Doudna, J.A., Charpentier, E., 2012. A

     programmable dual-RNA-guided DNA endonuclease in adaptive bacterial immunity. Science

     337, 816–821.

16. Johansson, J., Mandin, P., Renzoni, A., Chiaruttini, C., Springer, M., Cossart, P., 2002. An

      RNA thermosensor controls expression of virulence genes in Listeria monocytogenes. Cell

      110, 551–561.

17. Jin, Y., Watt, R.M., Danchin, A., Huang, J., 2009. Small noncoding RNA GcvB is a novel

      regulator of acid resistance in Escherichia coli. BMC Genomics 10, 165.

18. Kortmann, J., Narberhaus, F., 2012. Bacterial RNA thermometers: molecular zippers and

     switches. Nat. Rev. Microbiol. 10, 255–265.

19. Landt, S.G., Lesley, J.A., Britos, L., Shapiro, L., 2010. CrfA, a small noncoding RNA

     regulator of adaptation to carbon starvation in Caulobacter crescentus. J. Bacteriol. 192,

     4763–4775.

20. Levine, E., Hwa, T., 2008. Small RNAs establish gene expression thresholds. Curr. Opin.

     Microbiol. 11, 574–579.

21. Makarova, K.S., Haft, D.H., Barrangou, R., Brouns, S.J.J., Charpentier, E., Horvath, P.,

     Moineau, S., Mojica, F.J.M., Wolf, Y.I., Yakunin, A.F., Van der Oost, J., Koonin, E.V., 2011.

     Evolution and classification of the CRISPR-Cas systems. Nat. Rev. Microbiol. 9, 467–477.

22. Maki, K., Morita, T., Otaka, H., Aiba, H., 2010. A minimal base-pairing region of a bacterial

     small RNA SgrS required for translational repression of ptsG mRNA. Mol. Microbiol. 76, 782–

     792.

23. Mali, P., Yang, L., Esvelt, K.M., Aach, J., Guell, M., Dicario, J.E., Norville, J.E., Church, G.M.,

     2013. RNA-guided human genome engineering via Cas9. Science.

24. Massé, E., Vanderpool, C.K., Gottesman, S., 2005. Effect of RyhB small RNA on global iron

      use in Escherichia coli. J. Bacteriol. 187, 6962–6971.

25. Mey, A.R., Craig, S.A., Payne, S.M., 2005. Characterization of Vibrio cholerae RyhB: the

     RyhB regulon and role of ryhB in biofilm formation. Infect. Immun. 73, 5706–5719.

26. Morita, T., Maki, K., Aiba, H., 2005. RNase E-based ribonucleoprotein complexes:

     mechanical basis of mRNA destabilization mediated by bacterial noncoding RNAs. Genes

     Dev. 19, 2176–2186.

27. Mulhbacher, J., Brouillette, E., Allard, M., Fortier, L.-C., Malouin, F., Lafontaine, D.A., 2010.

     Novel riboswitch ligand analogs as selective inhibitors of guanine-related metabolic

     pathways. PLoS Pathog. 6, e1000865.

28. Murphy, E.R., Payne, S.M., 2007. RyhB, an iron-responsive small RNA molecule, regulates

     Shigella dysenteriae virulence. Infect. Immun. 75, 3470–3477.

29. M©ªller, T., Franch, T., H©ªjrup, P., Keene, D.R., Bächinger, H.P., Brennan, R.G., Valentin-

     Hansen, P., 2002. Hfq: a bacterial Sm-like protein that mediates RNA–RNA interaction. Mol.

     Cell 9, 23–30.

30. Nechooshtan, G., Elgrably-weiss, M., Sheaffer, A., Westhof, E., Altuvia, S., 2009. A pH-

     responsive riboregulator. Gene Dev. 23, 2650–2662.

31. Opdyke, J.A., Fozo, E.M., Hemm, M.R., Storz, G., 2011. RNase III participates in GadY

     dependent cleavage of the gadX–gadW mRNA. J. Mol. Biol. 406, 29–43.

32. Opdyke, J.A., Kang, J., Storz, G., 2004. GadY, a small-RNA regulator of acid response

     genes in Escherichia coli. J. Bacteriol. 186, 6698–6705.

33. Papenfort, K., Bouvier, M., Mika, F., Sharma, C.M., Vogel, J., 2010. Evidence for an

     autonomous 5ꠗ target recognition domain in an Hfq-associated small RNA. Proc. Natl.

     Acad. Sci. U. S. A. 107, 20435–20440.

34. Perez-Rodriguez, R., Haitjema, C., Huang, Q., Nam, K.H., Bernardis, S., Ke, A., DeLisa,

     M.P., 2011. Envelope stress is a trigger of CRISPR RNA-mediated DNA silencing in

     Escherichia coli. Mol. Microbiol. 79, 584–599.

35. Pernestig, A., Georgellis, D., Romeo, T., Suzuki, K., Tomenius, H., Normark, S., Melefors,

     Ö., 2003. The Escherichia coli BarA-UvrY two-component system is needed for efficient

     switching between glycolytic and gluconeogenic carbon sources. J. Bacteriol. 185, 843–853.

36. Pichon, C., Felden, B., 2008. Small RNA gene identification and mRNA target predictions in

     bacteria. Bioinformatics (Oxford, England) 24, 2807–2813.

37. Pomposiello, P.J., Demple, B., 2001. Redox-operated genetic switches: the SoxR and OxyR

     transcription factors. Trends Biotechnol. 19, 109–114.

38. Pul, U., Wurm, R., Arslan, Z., Geissen, R., Hofmann, N., Wagner, R., 2010. Identification and

     characterization of E. coli CRISPR-cas promoters and their silencing by H-NS. Mol.

     Microbiol. 75, 1495–1512.

39. Raabe, C.A., Hoe, C.-H., Randau, G., Brosius, J., Tang, T.-H., Rozhdestvensky, T.S., 2011.

     The rocks and shallows of deep RNA sequencing: Examples in the Vibrio cholerae RNome.

     RNA 17, 1357–1366.

40. Rasmussen, A.A., Eriksen, M., Gilany, K., Udesen, C., Franch, T., Petersen, C., Valentin-

     Hansen, P., 2005. Regulation of ompA mRNA stability: the role of a small regulatory RNA in

     growth phase-dependent control. Mol. Microbiol. 58, 1421–1429.

41. Repoila, F., Gottesman, S., 2001. Signal transduction cascade for regulation of RpoS:

     temperature regulation of DsrA. J. Bacteriol. 183, 4012–4023.

42. Richards, G.R., Vanderpool, C.K., 2011. Molecular call and response: The physiology of

     bacterial small RNAs. Biochim. Biophys. Acta 1809, 525–531.

43. Tang, T.-H., Bachellerie, J.-P., Rozhdestvensky, T., Bortolin, M.-L., Huber, H., Drungowski,

     M., Elge, T., Brosius, J., Hüttenhofer, A., 2002. Identification of 86 candidates for small non-

     messenger RNAs from the archaeon Archaeoglobus fulgidus. Proc. Natl. Acad. Sci. U. S.

     A. 99, 7536–7541.

44. Urbanowski, M.L., Stauffer, L.T., Stauffer, G.V., 2000. The gcvB gene encodes a small

      untranslated RNA involved in expression of the dipeptide and oligopeptide transport

      systems in Escherichia coli. Mol. Microbiol. 37, 856–868.

45. Ve¢§cerek, B., Moll, I., Bläsi, U., 2007. Control of Fur synthesis by the non-coding RNA RyhB

      and iron-responsive decoding. EMBO J. 26, 965–975.

46. Wadler, C.S., Vanderpool, C.K., 2007. A dual function for a bacterial small RNA: SgrS

      performs base pairing-dependent regulation and encodes a functional polypeptide. Proc.

      Natl. Acad. Sci. U. S. A. 104, 20454–20459.

47. Waldminghaus, T., Gaubig, L.C., Klinkert, B., Narberhaus, F., 2009. The Escherichia coli

      ibpA thermometer is comprised of stable and unstable structural elements. RNA Biol. 6,

      455–463.

48. Westra, E.R., Swarts, D.C., Staals, R.H.J., Jore, M.M., Brouns, S.J.J., Van der Oost, J.,

     2012. The CRISPRs, they are a-changin¡¯: how prokaryotes generate adaptive immunity.

     Annu. Rev. Genet. 46, 311–339.




Total:118 page:(8/5)
54 Á¤º¸ ÀÌ¿µÈÆ RNA ±¸Á¶ ±â¹Ý ¼¼Æ÷³» ¿Âµµ°¨ÀÀ¼¾¼­ 15.08.21 12081
53 Á¤º¸ ÃÖÁ¾Çö ¹Ì»ý¹°À» ÀÌ¿ëÇÑ ÀÌŸÄÜ»ê »ý»ê ¹× µ¿Çâ 15.06.30 13833
52 Á¤º¸ ÀÌÆòõ Whole-Genome and Whole-Population ºÐ¼®À» ÀÌ¿ë.. 15.05.27 11954
51 Á¤º¸ ³ëÁ¤Çý ¼¼±ÕÀÇ ¹ø¿ª ½ºÆ®·¹½º ¹ÝÀÀ 15.05.27 11035
50 Á¤º¸ À±¿©ÁØ »ýÇÕ¼º°ú À¯±âÇÕ¼º À¶ÇÕÀ» ÅëÇÑ Ãµ¿¬¹° ÀüÇÕ¼º ±â.. 15.04.30 12533
49 Á¤º¸ À¯º´Á¶ ¹ÙÀÌ¿ÀÀ̼ҺÎź¿Ã(Bio-isobutanol)ÀÇ »ý»ê±â¼ú°ú .. 15.04.30 17244
48 Á¤º¸ ±è¼±¿ø ¹Ì»ý¹° ¼¼Æ÷°øÀå ´ë»çÁ¶Àý ±â¼úÀÇ °³¹ßµ¿Çâ 15.03.19 12273
47 Á¤º¸ À̱չΠÁö´ÉÇü ¹ÙÀÌ¿À ½Ã½ºÅÛ¿¡ ÀÇÇÑ °íÇ°Áú Ç×ü ÀǾàÇ°.. 15.03.16 14986
46 Á¤º¸ ÀÌ»ó¿± ÇÕ¼º Á¶Àý sRNAs °³¹ß ¹× ´ë»ç°øÇп¡¼­ÀÇ ÀÀ¿ë¿¡ .. 15.03.16 13116
45 Á¤º¸ Á¶ÁÖÇö »ýü¹æ¾îÆéŸÀ̵åÀÇ ´ë·®»ý»ê ¹× ½Ç¿ëÈ­¿¡ ´ëÇÑ .. 15.02.25 19157
44 Á¤º¸ ±èµ¿¸³ ¹Ì»ý¹° Àü±â»ýÇÕ¼ºÀ» À§ÇÑ ÁöÁöü °³¹ß µ¿Çâ 15.02.25 11506
43 Á¤º¸ ±è±ÙÁß °íºÎ°¡°¡Ä¡ ´Ü¹éÁú ¼ÒÀ縦 ¾ÏȣȭÇÑ ³­¹ßÇö À¯Àü.. 15.01.27 15100
42 Á¤º¸ À̼º±¹ Evolutionary adaptation ¹× reverse engineering.. 15.01.27 13278
41 Á¤º¸ ±èÇÊ ±³ ÀΰøÀûÀÎ »êÈ­½ºÆ®·¹½º³»¼º ÄÚ¸®³×¹ÚÅ׸®¿ò 14.12.10 13358
40 Á¤º¸ ÇÑÁøÈñ Áø¼¼³ë»çÀ̵带 ÀÌ¿ëÇÑ ³úÀÎÁö±â´É Çâ»ó¿¡ ´ëÇÑ .. 14.12.09 15390
39 Á¤º¸ Á¤±âÁØ ÄÚ¸®³×¹ÚÅ׸®¿ò ±â¹Ý ÇÕ¼º»ý¹°ÇÐÀû ¿¬±¸ µ¿Çâ 14.11.21 16440
[1] [2] [3] [4] [5] [6] [7] [8]