Å©ÒÕѧÂÛÎÄ

Äúµ±Ç°µÄλÖãºÑ§ÊõÌà > ũѧÂÛÎÄ > Å©ÒÕѧÂÛÎÄ >

Ñо¿É¸Ñ¡Ä͵͵ªÄÜÁ¦Ç¿ºÍµªÐ§ÂʸߵÄ×÷Îï»ùÒòÐÍ

À´Ô´£ººËũѧ±¨ ×÷ÕߣºÖÓ˼ÈÙ;¹¨Ë¿Óê;ÕÅÊÀ´¨
·¢²¼ÓÚ£º2018-07-03 ¹²15019×Ö
¡¡¡¡ÕªÒª£ºÉ¸Ñ¡Ä͵͵ªÄÜÁ¦Ç¿¡¢µªÐ§ÂʸߵÄ×÷Îï»ùÒòÐÍÊÇÌá¸ßµªËØÀûÓÃЧÂÊ£¬ ½ÚÔ¼¿óÖÊÓªÑø×ÊÔ´£¬ ¼õÉÙ»·¾³ÎÛȾµÄÒ»ÖÖÓÐЧ;¾¶¡£±¾ÎÄͨ¹ýÂÛÊö×÷ÎïÄ͵͵ªÐԺ͵ªÐ§ÂʵĸÅÄɸѡʱÆÚ¡¢ÆÀ¼ÛÖ¸±ê£¬ ·ÖÎöÁ½ÕßµÄÒìͬµã£¬ ·¢ÏÖ×÷ÎïÄ͵͵ªÐԺ͵ªÐ§ÂÊÁ½ÕßÆÀ¼ÛµÄÖ¸±êºÍʱÆÚ»ù±¾ÀàËÆ¡£Í¬Ê±£¬ ±È½ÏÁ˵ªËØÓªÑø»®·ÖµÄ²»Í¬ÀàÐÍ£¬ ÈÏΪ½«²»Í¬»ùÒòÐÍ»®·Ö³ÉµÍЧ²»ÏìÓ¦ÐÍ¡¢µÍЧÏìÓ¦ÐÍ¡¢¸ßЧ²»ÏìÓ¦ÐÍ¡¢¸ßЧÏìÓ¦Ð͸üºÏÀí£¬ ÄܸüÇåÎú±íÏÖ³ö²»Í¬Æ·ÖÖ¶ÔµªËصÄÀûÓÃ×´¿ö¡£ºÏÀí¡¢È«ÃæÆÀ¼Û×÷Îﲻͬ»ùÒòÐͶԵªËصÄÀûÓÃ×´¿ö£¬ ¼ÓÇ¿¶Ô×÷ÎïÄ͵͵ªÐԺ͵ªÐ§ÂʵÄÉúÀí»úÖƺͷÖ×ÓÉúÎïѧ»úÀíÑо¿£¬ ¶ÔÓÚ×÷ÎïÓýÖÖ¡¢ÔÔÅ༰·¢Õ¹½ÚÔ¼¸ßЧµÄÂÌÉ«Å©Òµ¾ßÓÐÖØÒªÒâÒå¡£
¡¡¡¡
¡¡¡¡¹Ø¼ü´Ê£º×÷Î Ä͵͵ªÐÔ£» µªÐ§ÂÊ£»
¡¡¡¡
¡¡¡¡Research Progress on Low Nitrogen Tolerance and Nitrogen Efficiency in Crop Plants
¡¡¡¡
¡¡¡¡Abstract:Screening of crop genotypes with strong tolerance to low nitrogen £¨TLN£© and high nitrogen efficiency £¨HNE£© is an effective way to improve nitrogen use efficiency, save mineral resources and reduce environmental pollution. This paper discussed the concepts, screening periods, evaluation indicators of crop tolerance to low nitrogen and nitrogen efficiency, analyzed the similarities and differences between the TLN and HNE. The results showed that the evaluation index and screening period of low nitrogen resistance and nitrogen efficiency was basically similar. Through compared with the different types of nitrogen nutrition division, it was reasonably concluded that different varieties divided into inefficient nonresponder, inefficient responder, efficient nonresponder and efficient responder, which could clearly show the different crop genotypes nitrogen use efficiency situation. Reasonable and comprehensive evaluating the nitrogen utilization in different crop genotypes, strengthen the physiological and molecular biological mechanism of crop low nitrogen tolerance and nitrogen efficiency will make a big impact to crop breeding, cultivation and the development of efficient green agriculture.
¡¡¡¡
¡¡¡¡Keyword:crop; resistance to low nitrogen; nitrogen efficiency;

Å©×÷Îï
¡¡¡¡
¡¡¡¡µªËØÊÇÓ°Ïì×÷ÎïÉú³¤·¢Óý¡¢²úÁ¿¼°Æ·ÖÊÐγɵÄÖØÒªÓªÑøÔªËØ[1-4], Ò²ÊÇÍÁÈÀÖÐÒ×±»ºÄ¾¡µÄÔªËØ[5].´ÓÂÌÉ«¸ïÃü¿ªÊ¼£¬ ¹¤ÒµºÏ³ÉµÄµª·Ê¶ÔÌá¸ß×÷ÎïÉú²úÁ¦Æð×ÅÖØÒª×÷Ó㬠ÔÚÒ»¶¨³Ì¶ÈÉÏ»º½âÁËÓÉÓÚÈ«ÇòÈË¿ÚÔö³¤Òý·¢µÄÁ¸Ê³°²È«ÎÊÌâ[6].ÿÄêµÄµª·ÊÊ©ÓÃÁ¿ÔÚ²»¶ÏÔö³¤£¬ µ«µª·ÊÀûÓÃЧÂÊÖð½¥½µµÍ[7], ¾Ýͳ¼Æµª·ÊÊ©ÈëÍÁÈÀºó£¬ ½öÓÐ30%~35%±»×÷ÎïÎüÊÕ[8].´óÁ¿Í¶Èë°º¹óµÄ»¯Ê¯ÄÜÔ´·ÊÁÏ£¬ ²»½öÔì³É×ÊÔ´ÀË·Ñ£¬ ¾­¼ÃЧÒæ½µµÍ£¬ »¹µ¼ÖÂÅ©ÌïÉú̬ϵͳÔâµ½ÎÛȾºÍÆÆ»µ[9-10].ÎÒ¹úÊÇÅ©Òµ´ó¹ú£¬ ¸ûµØÃæ»ýλ¾ÓÊÀ½çÇ°ÁУ¬ µª·ÊͶÈëÒ²ÔÚÖðÄêµÝÔö£¬ µ«µª·ÊÀûÓÃÂʵÍÓÚÊÀ½çƽ¾ùˮƽ[11-12].Òò´Ë£¬ Ñ¡Óýµª¸ßЧ»òÄ͵͵ªµÄ×÷ÎïÆ·ÖÖ£¬ ¶Ô¼õÉÙµª·ÊͶÈëÁ¿£¬ ά³Ö×÷Îï²úÁ¿Îȶ¨[13], ·¢Õ¹½ÚÔ¼¸ßЧ¡¢»·¾³ÓѺÃÐÍÅ©Òµ¾ßÓÐÖØÒªÒâÒå[14].²»Í¬×÷Îͬһ×÷ÎﲻͬƷÖÖ¼äµÄÄ͵͵ªÄÜÁ¦ºÍµªÐ§ÂÊ´æÔÚÒ»¶¨²îÒì[15-17], ͨ¹ýÑ¡Óý¸ß²úÓÅÖʵÄÄ͵͵ª¡¢µªÐ§ÂʸߵÄ×÷ÎïÆ·ÖÖ£¬ ³ä·ÖÍÚ¾ò×÷Îï×ÔÉíµÄ»ùÒòDZÁ¦£¬ ÊÇÌá¸ßµªËØÀûÓÃЧÂʵÄÓÐЧ;¾¶¡£×Ô20ÊÀ¼Í30Äê´úHarvey[18]Ê״ᨵÀ²»Í¬ÓñÃ×Æ·ÖÖÔÚµªËØÓªÑøÉÏ´æÔÚ²îÒ죬 ½ØÖÁÄ¿Ç°£¬ ¶Ô²»Í¬×÷ÎïÆ·ÖÖµªËØÀûÓòîÒìµÄÑо¿ÒÑÔÚË®µ¾[19-20]¡¢Ð¡Âó[21]¡¢ÓñÃ×[22-23]¡¢Ñ̲Ý[24]¡¢ÓͲË[25-26]¡¢´óÂó[27]µÈ¶àÖÖÅ©×÷ÎïÉÏ¿ªÕ¹¡£Ä¿Ç°£¬ ¶Ôͬһ×÷ÎﲻͬƷÖÖɸѡµªËظßЧÀûÓõÄÑо¿Ö÷Òª¼¯ÖÐÔÚÄ͵͵ªÐԺ͵ªÐ§ÂÊÁ½·½Ã棬 ¶ø¹ØÓÚÄ͵͵ªÐԺ͵ªÐ§ÂʵÄÒìͬµãµÄÑо¿ÉÐÏʼû±¨µÀ¡£±¾ÎÄͨ¹ý¶Ô×÷ÎïµÄÄ͵͵ªÐԺ͵ªÐ§ÂʽøÐбȽϷÖÎö£¬ ̽¾¿Á½ÕßµÄÒìͬµã£¬ ÒÔÆڿ͹ÛÆÀ¼Û×÷Îﲻͬ»ùÒòÐ͵ĵªËØÓªÑøÌØÐÔ¡£
¡¡¡¡
¡¡¡¡1¡¢×÷ÎïÄ͵͵ªÐԺ͵ªÐ§ÂʵĸÅÄî¼°ÆÀ¼Û·½·¨
¡¡¡¡
¡¡¡¡RehmanµÈ[28]ÈÏΪ´ÓÉúÎïѧ½Ç¶È¿¼ÂÇ£¬ ѹÁ¦ÊÇÓëÖ²ÎïÉú³¤µÄÀíÏëÌõ¼þµÄÏÔ×ÅÆ«²î£¬ ×è°­ÁËËüÃÇÔÚÉú³¤¡¢·¢ÓýºÍ·±Ö³Öгä·Ö±í´ïÆäÒÅ´«Ç±ÄÜ¡£¶øµÍµªÐ²Æȱ»ÈÏΪÊÇÒ»ÖÖ·ÇÉúÎïµÄÍâ½çѹÁ¦£¬ Òò´ËÄ͵͵ªÐÔÊÇÖ¸×÷ÎïÔÚµªËØÑø·ÖŨ¶ÈµÍʱÈÔÈ»¾ßÓб£³ÖÕý³£Éú³¤µÄÄÜÁ¦£¬ ÇÒÉúÎïѧ²úÁ¿ÓëÔÚÕý³£µªÊ±µÄ½Ó½ü[29].Ò»°ãÓõ͵ªÓëÕý³£µªÆÀ¼ÛÖ¸±êµÄÏà¶ÔÖµ±íʾÆäÄ͵͵ªÄÜÁ¦£¬ ÊÇÆ·ÖÖÖ®¼äÖ¸±êÏà¶ÔÖµµÄ±È½Ï£¬ ¾Ý´Ë¿ÉÒÔ·ÖΪÄ͵͵ªÐÍ¡¢ÖмäÐÍ¡¢µÍµªÃô¸ÐÐÍ3Àà¡£µªÐ§ÂÊÊÇÖ¸Ìض¨¹©µªÌõ¼þÏÂ×÷Îï²úÁ¿µÄ¸ßµÍ£¬ ¿ÉÒÔ±íʾΪͬµÈ¹©µªÌõ¼þÏÂÎüµªÁ¿µÄ´óС£¬ Ò²¿ÉÒÔÓõ¥Î»ÎüÊÕµªËØËù²úÉúµÄ¸ÉÎïÖʶàÉÙ±íʾ[30], ÊÇÆ·ÖÖÖ®¼äÆÀ¼ÛÖ¸±ê¾ø¶ÔÖµµÄ±È½Ï£¬ ÔÚÏàͬµªËØˮƽ¿ÉÒÔ·ÖΪµª¸ßЧºÍµªµÍЧÁ½Àà¡£Ä͵͵ªÆ·ÖÖµÄÔÔÅ࣬ ÄܼõÉÙÅ©Ìﵪ·ÊµÄÊ©È룬 ½ÚÔ¼µª·Ê×ÊÔ´£¬ ´Ù½ø½ÚÔ¼ÐÍÅ©ÒµµÄ·¢Õ¹¡£ÔÔÅ൪ЧÂʸߵÄÆ·ÖÖ£¬ ÔÚͬµÈÊ©µª·ÊµÄÌõ¼þÏ£¬ ¾ßÓвúÉú¸ü¸ß²úÁ¿»òÉúÎïÁ¿µÄDZÁ¦£¬ ÓÐÀûÓڸ߲úÅ©ÒµµÄ·¢Õ¹¡£
¡¡¡¡
¡¡¡¡Ñ§ÕßÃǶÔÄ͵͵ªÐԵĶ¨ÒåºÍÀí½â½ÏΪͳһ£¬ ¶ø¶ÔµªÐ§ÂʵĶ¨ÒåºÍÀí½âÈ´¹Ûµã²»Ò»¡£Ò»°ãÈÏΪÔÚ²»Ê©µª»òµªÅ¨¶ÈµÍµÄÇé¿öÏÂÓÐÀûÓÚÄ͵͵ªÖÖÖʵÄɸѡ[15], Ä͵͵ªÐÔÖ÷ÒªÒԵ͵ªÌõ¼þÏÂÓëÕý³£µªÌõ¼þÐÔ×´Ö¸±ê²îÒìÇé¿öÀ´ÆÀ¼Û¡£Ä¿Ç°£¬ ±È½ÏÈϿɵÄÊÇMollµÈ[31]µÄµªÐ§ÂÊÀíÂÛ£¬ ¼´µªÐ§ÂÊ¿ÉÒÔ·ÖΪµªÎüÊÕЧÂʺ͵ªÀûÓÃЧÂÊÁ½²¿·Ö£¬ Çҵ͵ªÌõ¼þϵĵªÐ§ÂʲîÒìÖ÷ÒªÓɵªÀûÓÃЧÂÊËù¾ö¶¨£¬ ¶øÔڸߵªÌõ¼þÏ£¬ µªÎüÊÕЧÂÊÆð¾ö¶¨×÷Óá£Ò²ÓÐÑо¿ÈÏΪËùÓеªË®Æ½ÏÂÎüÊÕЧÂʶԵªÐ§ÂʵÄÖ±½Ó×÷Óþù´óÓÚÀûÓÃЧÂÊ£¬ ÊǵªÐ§ÂÊÖ÷ÒªÀ´Ô´[32-33].FotymaµÈ[34]ÔòÈÏΪ×÷Îï¶ÔµªËØÀûÓÃ×´¿ö¿ÉÒÔ·Ö½âΪµªÊÕ»ñÖ¸Êý¡¢Å©ÒÕЧÂÊ¡¢ÉúÀíЧÂÊ¡¢µª·ÊÔö²úÂÊ¡¢µªÒƶ¯Ö¸ÊýºÍµªÏìÓ¦¶ÈµÈÖ¸±ê¡£ÓÉÓÚÑо¿µÄ²àÖصãºÍÄ¿µÄ²»Í¬£¬ µªÐ§ÂʵÄÆÀ¼Û·½·¨½Ï¶à £¨±í1£© .ÆäÖеªËØÀûÓÃЧÂÊ £¨nitrogen use efficiency, NUE£© ±íʾÍÁÈÀÖе¥Î»µªËØËù²úÉúµÄ×ÑÁ£²úÁ¿£¬ Ëü¿ÉÒÔ·ÖΪµªÎüÊÕЧÂÊ £¨uptake efficiency, Up E£© ºÍµªÀûÓÃЧÂÊ £¨utilization efficiency, Ut E£© , ·Ö±ð·´Ó³Ö²Öê´ÓÍÁÈÀÖлñÈ¡µªËز¢»ýÀÛµÄÄÜÁ¦ºÍÖ²ÎォÎüÊյĵª×ª»¯Îª²úÁ¿µÄÄÜÁ¦£»µªËØũѧÀûÓÃЧÂÊ £¨agronomic efficiency, AE£© ±íʾʩ·Ê¶Ô×÷Îï²úÁ¿µÄÔö¼ÓЧӦ£»µª·Ê±í¹ÛÀûÓÃÂÊ £¨apparent nitrogen recovery, AR£© ·´Ó³Ê©·Ê¶ÔÖ²Ö굪ËØÀÛ»ýÁ¿µÄÔö¼ÓЧӦ£»µª·ÊÉúÀíЧÂÊ £¨physiological efficiency, PE£© ±íʾÓÉÊ©·ÊÒýÆðµÄÖ²Ö굪ËØÀÛ»ýÁ¿µÄÔö¼Ó²¿·Öת»¯Îª²úÁ¿µÄÄÜÁ¦¡£Ä¿Ç°£¬ ʹÓý϶àµÄÊDZàºÅΪ1¡¢2¡¢3µÄÆÀ¼Û·½·¨£¬ ÓÐЩÑо¿Ò²½«¹«Ê½ÖÐ×ÑÁ£²úÁ¿ÓÃÉúÎïÁ¿´úÌæ[33], ±àºÅΪ4¡¢5¡¢6µÄÆÀ¼Û·½·¨Ö÷Òª·´Ó³Ê©·Ê¶Ô²úÁ¿¼°ÎüµªÁ¿µÄÔö¼ÓЧ¹û¡£Ä͵͵ªÄÜÁ¦µÄÆÀ¼Û·½·¨½ÏΪͳһ£¬ Ö÷ÒªÒԵ͵ªºÍÕý³£µªÌõ¼þϵÄÐÔ×´Ö¸±êÏà¶ÔÖµ×÷ΪÆÀ¼Û·½·¨£¬ ÓÉÓÚ½ö¿¼ÂÇÖ¸±êÏà¶ÔÖµ £¨±àºÅ7¡¢9£© ÈÝÒ׳öÏÖÆäÏà¶ÔÖµ½Ó½ü1 £¨Ä͵͵ªÄÜÁ¦Ç¿£© , µ«²»Í¬µªË®Æ½ÏÂÖ¸±ê¾ø¶ÔÖµ¾ù½ÏµÍ £¨µªÐ§Âʽϵͣ© µÄÇé¿ö£¬ ÕâÖÖÀàÐ͵ÄÆ·ÖÖÏÔÈ»²»ÀûÓÚÉú²úʵ¼ÊÓ¦Ó㬠¶ø¸Ä½øºóµÄÆÀ¼Û·½·¨ £¨±àºÅ8¡¢10£© ¿Ë·þÁËÉÏÊö²»×㣬 ¿ÉÒÔÈ·±£Ñ¡ÓýµÄÆ·ÖÖͬʱ¾ßÓиߵªÐ§ÂʺÍÄ͵͵ªÐÔ¡£

 µªÐ§ÂʺÍÄ͵͵ªÐÔÖ÷ÒªÆÀ¼Û·½·¨¼°¼ÆË㹫ʽ
¡¡¡¡
¡¡¡¡2¡¢×÷ÎïÄ͵͵ªÐԺ͵ªÐ§ÂʵÄÒìͬµã
¡¡¡¡
¡¡¡¡2.1 ²»Í¬»ùÒòÐÍÄ͵͵ªÐԺ͵ªÐ§ÂÊɸѡʱÆÚÏàËÆ
¡¡¡¡
¡¡¡¡Í¬Ò»×÷ÎïµÄ²»Í¬ÉúÓýʱÆÚ£¬ ¶ÔµªËØÓªÑøÏìÓ¦µÄÇé¿ö²¢²»Ò»Ö£¬ Ñ¡ÔñºÏÊÊʱÆÚ¶Ô×÷ÎïÄ͵͵ªÐԺ͵ªÐ§ÂʽøÐÐɸѡÖÁ¹ØÖØÒª¡£É¸Ñ¡Ê±ÆÚÐèÒª¿¼ÂÇÒÔϼ¸¸öÒòËØ£º1£© ɸѡʱÆڵļø¶¨¹¤×÷µÄÖ¸±ê±äÒìϵÊýÒª¾¡Á¿´ó£¬ ÓÐÀûÓÚ²»Í¬»ùÒòÐͲîÒìµÄÏÔÏÖ£»2£© ɸѡʱÆڵĵªËØÓªÑøÏìӦ״¿öÄÜ»ù±¾´ú±íÕû¸öÉúÓýÆڵĵªËØÓªÑøÌØÐÔ£»3£© ɸѡʱÆڵļø¶¨¹¤×÷Ó¦¾¡Á¿¼ò½Ý¸ßЧ¡¢ÊÜ»·¾³Ó°ÏìС¡£
¡¡¡¡
¡¡¡¡Ä͵͵ªÆ·ÖÖɸѡһ°ãÑ¡ÔñÔÚ³ÉÊìÊÕ»ñÆÚ[41-42], ³ÉÊìÊÕ»ñÆÚ¿ÉÒÔµ÷²éÖÖÖ²Õ߱ȽϹØÐĵIJúÁ¿Ö¸±ê£¬ µ«ÊÔÑéËùÐèʱ¼ä½Ï³¤£¬ ×÷Îï²ÄÁÏÒÔ´óÌïÖÖֲΪÖ÷£¬ ÆäÖе͵ªÐ²ÆÈ £¨Ò»°ãÈ˹¤²»Ê©µª·Ê£¬ µªÔ´ÒÀ¿¿ÍÁÈÀÖвÐÁôµÈ£© ³ÖÐøÕû¸ö×÷ÎïÉúÓýÆÚ£¬ ¶øʵ¼ÊÉÏÃçÆÚ¿ÉÄܲ»È±µª£¬ Ëæ×Å×÷ÎïÉú³¤¶øÖð½¥¼Ó¾çÁ˵ªÐ²ÆÈ¡£Ò²ÓÐÑо¿ÈÏΪӦÔÚÃçÆÚ½øÐÐÄ͵͵ªÆ·ÖÖµÄɸѡ[39,43], ×÷ÎïÃçÆڵĵªËØÓªÑøÌØÐÔÓë´óÌïÆÚÓÐÒ»¶¨µÄÏàËÆÐÔ[33], Òò´ËÃçÆÚɸѡ¾ßÓÐÒ»¶¨µÄÒâÒå¡£×÷ÎïÃçÆÚÐ赪Á¿½ÏÉÙ£¬ ɸѡһ°ã²ÉÓÃÎÞÍÁÔÔÅ༼Êõ £¨Ë®ÅࡢɳÅàµÈ£© .ÃçÆÚ¼ø¶¨¾ßÓкÄʱ¶Ì¡¢ÈÝÁ¿´ó¡¢Öظ´ÐÔÇ¿¡¢Ò×ÓÚ»îÌå¼ø¶¨ºÍ»·¾³Ó°ÏìСµÈÓŵã[39].ɸѡ¸ßµªÐ§ÂʵÄÆ·ÖÖÒ»°ãÒ²ÔÚ³ÉÊìÆÚ[44-45]ºÍÃçÆÚ[46-47].½­Á¢¸ýµÈ[45]ÈÏΪˮµ¾¸ÉÎïÖÊÉú²úЧÂÊÔÚ·ÖÞÁÆÚ±äÒìϵÊý×îС£¬ ³éËëÆÚ±äÒìϵÊý×î´ó£»¶øµªËØÀÛ»ýÁ¿ÔÚ³ÉÊìÆÚ±äÒìϵÊý×î´ó£¬ ÔÚ³éËëÆÚ±äÒìϵÊý×îС¡£Ñо¿±íÃ÷£¬ ÆÀ¼ÛË®µ¾²»Í¬»ùÒòÐ͵ªÐ§ÂʵÄʱÆÚ²»Ò˹ýÔ磬 ³éËëÆÚÊÊÒËÆÀ¼Û¸ÉÎïÖÊÉú²úЧÂÊ£¬ ³ÉÊìÆÚÊÊÒËÆÀ¼ÛµªËØÀÛ»ýÄÜÁ¦¡£³ÉÊìÆÚɸѡ´æÔÚºÄʱ³¤µÈȱµã£¬ Òò´ËÓÐÑо¿Ìá³ö“Á½²½É¸Ñ¡·¨”À´¼ø¶¨µªÐ§ÂÊ£¬ ¼´ÏȽøÐÐÃçÆÚɸѡ£¬ ³õ²½µÃµ½±È½ÏµäÐ͵IJÄÁÏ£¬ È»ºóÊʵ±µÄËõСȺÌ壬 ÔÙ½øÐÐÈ«ÉúÓýÆڵļø¶¨ÊÔÑ飬 ×îÖÕͨ¹ý»ñµÃ²úÁ¿µÄ¸ßµÍÀ´È·¶¨µª¸ßЧºÍµªµÍЧ»ùÒòÐÍ[48].×÷ÎïÄ͵͵ªÐԺ͵ªÐ§ÂʵÄɸѡʱÆÚ¾ßÓÐÏàËÆÐÔ£¬ ´ó²¿·ÖÑо¿¶¼Ñ¡ÔÚ×÷Îï³ÉÊìÆÚ»òÃçÆÚ½øÐС£
¡¡¡¡
¡¡¡¡2.2 ²»Í¬»ùÒòÐÍÄ͵͵ªÐԺ͵ªÐ§ÂÊɸѡָ±êÏàËÆ
¡¡¡¡
¡¡¡¡²»Í¬Ö¸±ê¶ÔµªËصÄÃô¸ÐÐÔ²»Í¬£¬ ¶øÇÒ¶Ôͬһ×÷Îﲻͬ»ùÒòÐͲîÒìµÄÏìÓ¦Ò²ÓÐÇø±ð¡£Ñ¡ÔñÊÊÒ˵ÄÖ¸±êÀ´É¸Ñ¡Ä͵͵ªÐԺ͵ªÐ§ÂÊÐ迼ÂÇÈýµã£º1£© Ñ¡ÓÃÖ¸±êÔÚ²»Í¬µªËØˮƽÐè¾ßÓÐÒ»¶¨Ãô¸ÐÐÔ£¬ ¼´Ö¸±ê¶Ô²»Í¬µªËØˮƽµÄ±íÏÖ¾ßÓвîÒ죻2£© Ñ¡ÓÃÖ¸±êÔÚͬһµªËØˮƽ²»Í¬»ùÒòÐͼäµÄ±äÒìϵÊýÓ¦½Ï´ó£¬ ÒÔÀûÓÚÏÔÏÖ²»Í¬»ùÒòÐÍÖ®¼äµÄ²îÒì¡£3£© ×îÖÕ²úÁ¿ÓëÕâЩָ±êÐÔ×´µÄÒÅ´«Ïà¹ØÐԸߡ£
¡¡¡¡
¡¡¡¡Ä¿Ç°£¬ ¹ØÓÚÑ°ÕÒÊÊÒ˵Äɸѡָ±êÀ´ÆÀ¼Û×÷ÎïÄ͵͵ªÐԺ͵ªÐ§ÂʵÄÑо¿½Ï¶à£¬ µ«Æù½ñΪֹδ´ï³ÉÒ»Ö¡£¿É×÷ΪÄ͵͵ª»ùÒòÐÍɸѡµÄÖ¸±ê½Ï¶à£¬ Ö÷Òª°üÀ¨²úÁ¿¡¢µØÉϲ¿ÉúÎïÁ¿¡¢Ö²Ö굪ÎüÊÕ×ÜÁ¿¡¢µ¥Öê¹ÈÖØ¡¢µªËØ·´Ó¦Ö¸Êý¡¢È«ÖêÉúÎïÁ¿¡¢µØÉϲ¿µª»ýÀÛÁ¿¡¢ÉúÎïÁ¿Ôö¼ÓÖµ¡¢SPADÖµ¡¢Ò¶Ãæ»ý¡¢¸ù¹Ú±È¡¢¸ùÌå»ý¡¢¸ùϵÉúÎïÁ¿µÈ[49-52].²»Í¬»ùÒòÐÍÄ͵͵ªÐÔɸѡµÄÖ¸±ê±È½ÏÁé»î£¬ °üÀ¨²úÁ¿Ö¸±ê¡¢µªÓªÑøÖ¸±êÒÔ¼°Ò»Ð©ÐÎ̬ѧָ±ê¡£ÆÀ¼Û²»Í¬»ùÒòÐ͵ªÐ§ÂʵÄÖ¸±êÖ÷ÒªÓвúÁ¿¡¢ÉúÎïÁ¿¡¢µªËØ»ýÀÛÁ¿¡¢¾¥Ò¶µªËØÀÛ»ýÁ¿¡¢¾¥Ò¶º¬µªÁ¿¡¢µØÉϲ¿ÉúÎïÁ¿¡¢¸ùÉúÎïÁ¿µÈ[53-58].µªÐ§ÂÊÆÀ¼ÛµÄÖ¸±êÏà¶Ô½Ï¼¯ÖУ¬ Ö÷ҪΪ²úÁ¿Ö¸±ê¼°µªÓªÑøÖ¸±ê¡£×÷ÎïµÄ²úÁ¿ÊÇÖÖÖ²ÕߺÍÓýÖÖÕßÖ±½Ó×·ÇóµÄÖ¸±ê£¬ ÓÃÀ´ÆÀ¼Û²»Í¬»ùÒòÐ͵ÄÄ͵͵ªÐԺ͵ªÐ§ÂÊÒ²×î¾ßÓÐ˵·þÁ¦£¬ µ«²úÁ¿ÆÀ¼Û±ØÐëÔÚ³ÉÊìÊÕ»ñʱÆÚ½øÐУ¬ Òò´ËÃçÆÚɸѡËùÑ¡ÓõÄÖ¸±êÒ»°ãΪÉúÎïÁ¿¡¢µªÀÛ»ýÁ¿µÈ¡£ÎªÁËÉîÈëÁ˽â×÷Îﲻͬ»ùÒòÐͶԵªËØÓªÑøÏìÓ¦¼°ÀûÓÃ×´¿öµÄ»úÀí£¬ ³ýÁ˱íÐÍÌØÕ÷Ö¸±êÍ⣬ ÓÐЩÑо¿»¹ÒýÈ븨Öúɸѡָ±ê£¬ Ö÷ÒªÊÇһЩÉúÀíÖ¸±êÈçÒ¶ÂÌËغ¬Á¿¡¢¸ùϵ»îÁ¦¡¢¹âºÏËÙÂÊ¡¢ÏõËỹԭø»îÐÔ £¨nitrate reductase, NR£© [59]¡¢¹È°±õ£°·ºÏ³Éø»îÐÔ £¨glutamine synthetase, GS£© [60-61]µÈ¡£ÉúÀíÖ¸±êµÄ²â¶¨ËäÈ»ÓÐÀûÓÚ½ÒʾÄÚÔڵĵªËØÎüÊÕÀûÓûúÖÆ£¬ µ«²â¶¨¹ý³Ì·ÑʱºÄÁ¦¡£×ÜÌåÀ´Ëµ£¬ ²»Í¬»ùÒòÐÍÄ͵͵ªÐԺ͵ªÐ§ÂÊɸѡָ±ê»ù±¾ÀàËÆ£¬ ´ó²¿·ÖÑо¿Ö÷ÒªÒÔ²úÁ¿¡¢ÉúÎïÁ¿¡¢µªËØÀÛ»ýÁ¿×÷ΪÖ÷Ҫɸѡָ±ê¡£
¡¡¡¡
¡¡¡¡2.3 Ä͵͵ªÐԺ͵ªÐ§ÂÊɸѡÊÔÑéµÄ²îÒì
¡¡¡¡
¡¡¡¡Í¬Ò»»ùÒòÐÍÔÚ²»Í¬µÄµªË®Æ½Ìõ¼þϵĵªÐ§ÂÊ´æÔڽϴó²îÒ죬 ¼´µªË®Æ½ºÍ»ùÒòÐͼä´æÔÚÏÔ×Å»¥×÷[33], Òò´Ë²»Í¬»ùÒòÐ͵ªÐ§ÂÊÑо¿µÄÊÔÑé·½·¨Ò²ÉèÖöà¸öµªË®Æ½£¬ Ò»°ãΪµÍµªºÍ¸ßµª2¸ö´¦Àí[44], »òµÍµª¡¢Öеª¡¢¸ßµª3¸ö´¦Àí[46], ÊÔÑé·½°¸ÓëÄ͵͵ª»ùÒòÐÍɸѡµÄÊÔÑé·½°¸½ÏΪÏàËÆ£¬ Á½ÕßɸѡʱÆÚºÍÖ¸±êÒ²»ù±¾ÀàËÆ£¬ Òò´Ëµ¼ÖÂÐí¶àÈËÈÏΪµªÐ§ÂÊÓëÄ͵͵ªÐÔûÓÐÇø±ð£¬ ¶øÊÂʵÉÏÁ½Õß´æÔڽϴó²îÒ죺һÊÇÊÔÑé·½°¸²»Í¬¡£Ä͵͵ª»ùÒòÐÍɸѡÊÔÑ飬 Ö»ÉèÖÃ2¸öµªË®Æ½£¬ ·Ö±ðÊǵ͵ªºÍÕý³£µª £¨»ò×îÊʵª£© , µÍµªË®Æ½ÊǵªÐ²ÆÈˮƽ£¬ Õý³£µªË®Æ½±íʾ¸ÃµªË®Æ½ÊÇ×îÊʺÏ×÷ÎïÉú³¤¼°·±Ö³µÄÐèÒªµÄ £¨Ðè¶à´ÎÊÔÑéÈ·¶¨£© , ÓëÖеªË®Æ½¡¢¸ßµªË®Æ½´æÔÚÒ»¶¨Çø±ð£¬ µ«Êµ¼ÊÊÔÑé²Ù×÷ÉÏÕý³£µªË®Æ½ÓëÖеªË®Æ½µÄŨ¶È²îÒì²¢²»´ó£»¶þÊÇÁ½Õ߸ÅÄîºÍÊÔÑéÄ¿²»Í¬¡£µªÐ§ÂʱíʾµÄÊǵªËØͶÈëÓë»ñµÃ²ú³öµÄ±ÈÀý£¬ Ñо¿Ä¿µÄÊÇͶÈ뵪Ëز»±ä»òÔöÊ©µªËØÄÜ»ñµÃ¸ü¸ßµÄ²ú³ö£»Ä͵͵ªÐÔÔò±íʾµÍµªÐ²ÆÈʱÓëÕý³£µªÌõ¼þÏÂ×÷ÎïµÄ²úÁ¿»òÆäËûÖ¸±êµÄ½Ó½ü³Ì¶È£¬ Ñо¿Ä¿µÄÊǼõÉÙµªËصÄͶÈëÄÜ»ñµÃÓëÕý³£Ê©µª²ú³öÏà½ü»ò½µµÍ½ÏÉÙ¡£Á½ÕߵĸÅÄî¼°Ä¿µÄ²»Í¬£¬ Òò´Ë¼ÆËã·½·¨Ò²²»Í¬¡£ÈýÊÇÑо¿±³¾°²»Í¬¡£µªÐ§ÂÊÊÔÑéÊÇ×·Çó²úÁ¿£¬ ÉõÖÁ²»Ï§¹ýÁ¿Ê©Óõª·ÊÀ´´ïµ½Ä¿µÄ£¬ ËäÈ»¶Ô»·¾³Ôì³ÉÒ»¶¨ÆÆ»µ£¬ µ«¶Ô½â¾öÁ¸Ê³°²È«ÎÊÌâ×÷³öÁ˹±Ïס£¶øÄ͵͵ªÐÔÑо¿ÊÇ»ùÓÚÁ¸Ê³ÒѾ­»ù±¾Âú×ãÈËÃǵÄÐèÇó £¨Í¬Ò»Æ·ÖÖ¼õÉÙµª·ÊµÄͶÈ룬 ²úÁ¿¼¸ºõ²»¿ÉÄÜ»áÔö¼Ó£© , ÈËÃǶԻ·¾³¼°Éú̬µÄ±£»¤ÒâʶÖð½¥ÔöÇ¿£¬ ÒÔ¼°µª·Ê×ÊÔ´ÈÕÒæ°º¹óµÈ±³¾°ÏÂÌá³öµÄ¡£
¡¡¡¡
¡¡¡¡3¡¢²»Í¬Æ·ÖÖÄ͵͵ªÐÔ¡¢µªÐ§ÂʵÄÀàÐÍ»®·Ö
¡¡¡¡
¡¡¡¡¸ù¾ÝÆ·ÖÖ¶ÔµªËØÀûÓõÄ×´¿ö½«²»Í¬µÄÆ·ÖÖ»®·Ö³É²»Í¬µÄÀàÐÍ£¬ ÓÐÀûÓÚÆ·ÖÖ¼äµÄ±È½Ï¡£Àµ¤µÈ[38]ͨ¹ýÑо¿32¸öСÂóÆ·ÖÖÄ͵͵ªÐ²ÆȵĻùÒòÐͲîÒ죬 ½«Ð¡ÂóÆ·ÖÖ·ÖΪµÍµªÃô¸ÐÐÍ¡¢ÖмäÐÍ¡¢Ä͵͵ªÐÍ3À࣬ ²¢Ö¸³öµÍµªÃô¸ÐÐͶàΪÔçÆÚÆ·ÖÖ £¨20ÊÀ¼Í50~70Äê´ú£© , ¶ø²¿·Ö½ÏÍíÆ·ÖÖ £¨20ÊÀ¼Í80~90Äê´ú£© ¾ßÓÐÄ͵͵ªµÄÓÅÊÆ¡£»ÆÓÀÀ¼µÈ[56]ºÍ³Â¶þÓ°µÈ[47]·Ö±ð¶ÔË®µ¾ºÍ¹È×ÓÖÖÖʵªËØÎüÊÕÓëÀûÓõÈ×´¿ö½øÐÐÁË·ÖÎö£¬ ½«Æ·ÖÖÀàÐÍ·Ö³ÉË«¸ßЧÐÍ £¨µÍµª¸ßЧ¸ßµª¸ßЧ£© ¡¢¸ßµª¸ßЧÐÍ¡¢µÍµª¸ßЧÐͺÍË«µÍЧÐÍ4Àà¡£ÆäÖУ¬ »ÆÓÀÀ¼µÈ[56]ÈÏΪ˫¸ßЧÐͺ͸ߵª¸ßЧÐÍÊܵª·ÊÓ°Ïì´ó£¬ ÊôÓÚµª¸ßЧÐÍ£»µÍµª¸ßЧÐÍÊܵª·ÊÓ°ÏìС£¬ ÊôÓÚÄ͵͵ªÐÍ¡£ÉÏÊöÑо¿±íÃ÷£¬ Ä͵͵ªÐԺ͵ªÐ§ÂÊÊÇ×÷Îï¶ÔµªËØÓªÑøÏìӦ״¿ö²»Í¬·½ÃæµÄÆÀ¼Û£¬ Á½ÕßÖ®¼äÓÐÏàËÆÐÔ£¬ Ò²´æÔÚ²îÒìÐÔ¡£ÎªÈ«ÃæÆÀ¼Û×÷ÎïµÄµªËØÓªÑøÀûÓÃ×´¿ö£¬ ½ö·ÖÎöÄ͵͵ªÐÔ»òµªÐ§ÂÊÊÇƬÃæµÄ£¬ Ó¦¸Ã½«Á½Õß½áºÏÆðÀ´£¬ ¿Í¹ÛÆÀ¼Û×÷Îﲻͬ»ùÒòÐ͵ĵªËØÓªÑøÏìÓ¦²îÒì¡£²ÜÃô½¨µÈ[62]ͨ¹ý½«Ä͵͵ªÐÔÓë²úÁ¿ £¨µªÐ§ÂʵÄÌåÏÖ£© Ïà½áºÏ°ÑÓñÃ×ÔÓ½»Æ·ÖÖ·ÖΪÒÔϼ¸¸öÀàÐÍ£º¸ß²ú²»Ãô¸ÐÐÍ £¨½ÏÇ¿Ä͵͵ªÐÔ£¬ µÍµªºÍ¸ßµª²úÁ¿¾ù½Ï¸ß£© ¡¢µÍ²ú²»Ãô¸ÐÐÍ £¨½ÏÇ¿Ä͵͵ªÐÔ¡¢¸ßµª²úÁ¿Öеȣ© ¡¢¸ß²úÃô¸ÐÐÍ £¨Ä͵͵ªÐԲ ¸ßµª²úÁ¿¸ß£¬ µÍµª²úÁ¿µÍ£© ¡¢µÍ²úÃô¸ÐÐÍ £¨Ä͵͵ªÐԲ ¸ßµª²úÁ¿²»¸ß£¬ µÍµª²úÁ¿µÍ£© , ²¢Ö¸³ö²úÁ¿ÓëÖ²ÖêȱµªÖ¢×´µÄÑÏÖس̶ÈÓйء£´ÞÎÄ·¼µÈ[63]ÒÔÄ͵͵ªÐ²ÆÈÖ¸ÊýÊÇ·ñ´óÓÚ0.9ºÍ¸ßµªÏ²úÁ¿ÊÇ·ñ´óÓÚƽ¾ùÖµ½«27·ÝÓñÃ××Ô½»²ÄÁÏ»®·Ö³É4ÖÖµªÐ§ÂÊÀàÐÍ£¬ ¼´¸ß²úµª¸ßЧÐÍ £¨Ä͵͵ªÐ²ÆÈÖ¸Êý´óÓÚ0.9, ¸ßµªÏ²úÁ¿¸ßÓÚƽ¾ùÖµ£© ¡¢µÍ²úµª¸ßЧÐÍ¡¢¸ß²úµªµÍЧÐÍ¡¢µÍ²úµªµÍЧÐÍ£¬ ²¢ÈÏΪѡÓýÄ͵͵ªÐÔÆ·ÖÖʱӦÖØÊÓµªÀûÓÃЧÂÊ¡£ÓÉͼ1¿ÉÖª£¬ Gerloff[64]ºÍBlair[65]½«²»Í¬Æ·ÖÖÔÚµÍÑø·ÖʱºÍÊ©µªÁ¿Ôö¼ÓʱµÄ²úÁ¿ÏìÓ¦Çé¿ö»®·ÖΪµÍЧ²»ÏìÓ¦ÐÍ¡¢µÍЧÏìÓ¦ÐÍ¡¢¸ßЧ²»ÏìÓ¦ÐÍ¡¢¸ßЧÏìÓ¦ÐÍ4ÖÖÀàÐÍ¡£¸Ã»®·Ö·½·¨Óë²ÜÃô½¨µÈ[62]¡¢´ÞÎÄ·¼µÈ[63]µÄ·½·¨Ïà±È£¬ ÄܸüÇå³þµÄ·´Ó³×÷Îï¶ÔµªËصÄÀûÓÃ×´¿ö£¬ µÍЧ²»ÏìÓ¦ÐͱíÏÖΪ˫µÍЧÐÍ¡¢Ä͵͵ªÐÍ£»µÍЧÏìÓ¦ÐͱíÏÖΪ¸ßµª¸ßЧÐÍ¡¢µÍµªÃô¸ÐÐÍ£»¸ßЧ²»ÏìÓ¦ÐͱíÏÖΪµÍµª¸ßЧÐÍ¡¢Ä͵͵ªÐÍ£»¸ßЧÏìÓ¦ÐͱíÏÖΪ˫¸ßЧÐÍ¡¢µÍµªÃô¸ÐÐÍ¡£±íÃ÷£¬ ¸ßЧÏìÓ¦ÐÍ¿ÉÒÔ×÷Ϊµª¸ßЧµÄÓÅÐãÆ·ÖÖ£¬ ¶ø¸ßЧ²»ÏìÓ¦ÐÍ¿ÉÒÔ×÷ΪÄ͵͵ªµÄÓÅÐãÆ·ÖÖ¡£MaiaµÈ[66]ÈÏΪÄ͵͵ªÐԺ͵ªÐ§ÂÊÊÇÓɲ»Í¬»ùÒò×é¿ØÖƵģ¬ Èç¹ûÁ½ÕßÖ®¼äûÓоºÕù£¬ ͬʱѡÔñÄ͵͵ªÐԺ͵ªÐ§ÂÊÊÇ¿ÉÐеġ£Ëæ×ÅÅ©ÒµÖð½¥Ïò¸ß²ú¡¢Éú̬µÄ·½Ïò·¢Õ¹£¬ Ñ¡Ôñ¼ÈµªÐ§ÂʸßÓÖÄ͵͵ªµÄ×÷ÎïÆ·ÖÖ³ÉΪÇ÷ÊÆ£¬ Òò´ËÄܹ»ÇåÎúÌåÏÖ×÷ÎﵪËØÀûÓÃ×´¿ö £¨ÄÜͬʱ±íÃ÷Ä͵͵ªÐԺ͵ªÐ§ÂÊ£© µÄ»®·Ö¸ü¼ÓÖØÒª¡£

²»Í¬Æ·ÖÖ²úÁ¿ÔÚÓªÑøˮƽ±ä»¯ÉϵIJîÒì
¡¡¡¡
¡¡¡¡4¡¢×÷Îﲻͬ»ùÒòÐ͵ªËØÀûÓòîÒìµÄÔ­Òò¼°»úÖÆ
¡¡¡¡
¡¡¡¡²»Í¬Æ·ÖÖÖ®¼äµÄµªËØÀûÓòîÒìÊÇÓÉÖ²ÎïѧÐÎ̬¡¢ÉúÀí¼°·Ö×Ó±í´ïµÈ¶à·½Ã湲ͬ×÷ÓõĽá¹û¡£¼Þ½Ó¿ÉÒÔÖ±¹Û·´Ó³Ö²ÎïµØÉϲ¿ºÍµØϲ¿·ÖµÄÖØÒªÐÔ£¬ ÷·¼[67]ÒÔ²»Í¬µÄµªÐ§ÂʵÄÑ̲ÝÆ·ÖÖ½øÐм޽ӣ¬ ·¢ÏÖ¼Þ½ÓÃçµÄµªÐ§ÂÊÒÀ´ÎΪµª¸ßЧÂÊÆ·ÖÖ>µªµÍЧÂÊÆ·ÖÖ½ÓËë/µª¸ßЧÂÊÆ·ÖÖÕèľ>µª¸ßЧÂÊÆ·ÖÖ½ÓËë/µªµÍЧÂÊÆ·ÖÖÕèľ>µªµÍЧÂÊÆ·ÖÖ¡£RuizµÈ[68]Ñо¿·¢ÏÖÒԸߵªÐ§ÂʵÄÕèľ×÷Ϊ¼Þ½Ó·½Ê½ÊÇÌá¸ßÑ̲ݵªËØÀûÓÃЧÂʵÄÒ»ÖÖ¿ìËÙ¶øÓÐЧµÄ·½Ê½¡£ÐíÈ«±¦[69]ÒÔ²»Í¬Ä͵͵ªÐԵĻƹÏÆ·ÖÖ½øÐм޽ӣ¬ ·¢ÏÖ²»Í¬¹©µªË®Æ½ÏÂÄ͵͵ªµÄ»Æ¹Ï×÷ΪÕèľʱ±È½ÓËë¿ÉÒÔÏÔ×ÅÌá¸ß»Æ¹Ï¶ÔµªËصÄÎüÊÕºÍÀûÓÃÄÜÁ¦¡£ËµÃ÷×÷ÎïµÄµØϲ¿·Ö £¨¸ùϵ£© ÊDz»Í¬Æ·ÖÖµªÎüÊÕÀûÓõÄÖ÷Òª²îÒìËùÔÚ¡£Òò´Ëµª¸ßЧƷÖÖÍùÍù¾ßÓнϴóµÄ¸ùϵÌå»ý¡¢¸ùϵ±íÃæ»ý¡¢¸ù³¤¡¢¸ùÊýµÈÌØÕ÷[70].×÷ÎïÉúÀí·½ÃæµÄ²îÒìÖ÷Òª±íÏÖÔÚ¸ùϵ»îÁ¦¡¢Ò¶ÂÌËغ¬Á¿¡¢¹âºÏËÙÂÊ¡¢µª´úл¹Ø¼üø[Ö÷ҪΪNR¡¢GS¡¢¹È°±ËáºÏ³Éø £¨glutamate synthase, GOGTA£© µÈ]¡¢ÄÍвÆÈÉúÀíÖ¸±êµÈ£¬ ÌرðÊÇGS±»ÈÏΪÓë×÷ÎïµÄµªÐ§ÂÊ[71]ºÍÄ͵͵ªÐÔ[72]Óйء£Ö²Îï·Ö×ÓÉúÎïѧ·½ÃæµÄÑо¿Ö÷Òª°üÀ¨£º1£© µªËØÎüÊÕתÔ˵ÄÔØÌå»ùÒò£¬ ÈçÏõËáÑÎÎüÊÕϵͳĿǰ°üÀ¨NRT1ºÍNRT2»ùÒò¼Ò×å[73];°±×ªÔ˵°°×£¬ ¿ÉÒÔ·ÖΪAMT1ºÍAMT2»ùÒò¼Ò×å[74].2£© µªËØÀûÓÃͬ»¯»ùÒò£¬ ÈçNR»ùÒò£¬ ÄâÄϽæÖзֱðΪNia1ºÍNia2[75], ÆäÖÐNia2ÆðÖ÷Òª×÷Óã»GS»ùÒòÔڸߵÈÖ²ÎïÖдæÔÚ2ÖÖÀàÐÍ£¬ ¼´GS1ºÍÖÊÌåÐÍGS2[76], GS1Ö÷ÒªÔÚ¸ù¾¥Öбí´ï£¬ ºÏ³É¹È°±õ£°·£¬ ²¢²ÎÓë°±»ùËáÑ­»·£»GS2Ö÷ÒªÔÚÂÌÉ«×éÖ¯µÄÒ¶ÂÌÌåµÈÖÊÌåÖбí´ï£¬ ͬ»¯¹âºôÎü²úÉúµÄ°±[77].Ö²ÎïÔÚµªÈ±·¦Ê±£¬ »á¼ÓËÙÌåÄÚµªÑ­»·¡£Ñо¿±íÃ÷£¬ GS»ùÒòÓ뵪µÄÔÙÒƶ¯ÓйØ[78].´ËÍ⣬ NRT1.7»ùÒòÒ²ÓÐÀûÓÚÏõ̬µªµÄÔÙÒƶ¯£¬ ÒÔÊÊÓ¦µªÐ²ÆÈ»·¾³[79].
¡¡¡¡
¡¡¡¡5¡¢Õ¹Íû
¡¡¡¡
¡¡¡¡ÎÒ¹úÅ©ÒµÏÖ´ú»¯µÄ½¨ÉèÓ¦»ùÓÚ“ÂÌÉ«¡¢Ñ­»·¡¢µÍ̼”µÄ·¢Õ¹ÀíÄ ×ß²ú³ö¸ßЧ¡¢×ÊÔ´½ÚÔ¼¡¢²úÆ·°²È«¡¢»·¾³ÓѺõĵÀ·[80].¶øÅ©Òµ×ÊÔ´ØÑ·¦£¬ »·¾³Ô¼Êø¼Ó¾çÒѾ­³ÉΪũҵ¿É³ÖÐø·¢Õ¹ËùÃæÁÙµÄÖØÒªÎÊÌ⣬ Lynch[81]ÈÏΪ“ÂÌÉ«¸ïÃü”±»ÅúÆÀµÄÒ»¸ö·½ÃæÊÇÐÂÆ·ÖÖÔÚÓªÑøÌõ¼þµÍµÄÇé¿öÏÂÆ䵪ЧÂÊÒ²µÍ£¬ Ö»ÊÇÌá¸ßÁËÆä¶Ô¸ßÓªÑøÎïÖÊͶÈëµÄ²úÁ¿ÏìÓ¦£¬ ¼´µÍЧÏìÓ¦ÐÍ£¬ ÈçË®µ¾Æ·ÖÖÔç89-01ºÍÔçôÌ152[56].LeµÈ[82]ÈÏΪÓÉÓÚ¾­¼ÃºÍÉú̬µÄÔ­Òò£¬ Å·ÖÞÅ©Òµ½«¿ÉÄÜ»áÏòµÍµªÍ¶Èë·½Ïò·¢Õ¹¡£Òò´ËÌá¸ß×÷ÎïµÄµªÐ§ÂʺÍÄ͵͵ªÐÔÊǼӿìÎÒ¹úÅ©ÒµÏÖ´ú»¯½ø³ÌµÄÖØҪ;¾¶£¬ Ñ¡ÓýµÄÆ·ÖÖÓ¦¼æ¹Ë¸ßµªÐ§ÂʺÍÄ͵͵ªÐÔ£¬ ÈçÑ̲ÝÆ·ÖÖ14P9[58]¡¢ÓñÃ×Æ·ÖÖ¾©¿Æ132ºÍÅ©´ó108[41].×÷ÎïµÄÄ͵͵ªÐԺ͵ªÐ§ÂÊÖ®¼ä¼ÈÓÐÁªÏµ£¬ ÓÖ´æÔڽϴó²îÒ죬 Ä͵͵ªÐÔÆ«ÖØÓÚµªÏìÓ¦¶ÈµÄ·´Ó³£¬ Ä͵͵ªÐÔÇ¿±íÏÖΪµÍµªÏÂijЩָ±êÓëÕý³£µª»ò¸ßµªÏ¸ü½Ó½ü£»µªÐ§ÂÊÔòÆ«ÖØÓÚijһµªË®Æ½²úÁ¿µÄ·´Ó³£¬ µª¸ßЧÔò±íÏÖΪµÍµªºÍ¸ßµªÏµIJúÁ¿¶¼¸ü¸ß¡£Ä͵͵ªÐԺ͵ªÐ§ÂÊÊǶÔ×÷ÎﵪËØÀûÓÃ×´¿ö¼°ÌØÐÔ´Ó²»Í¬·½ÃæµÄÆÀ¼Û£¬ Á½ÕßɸѡµÄʱÆÚºÍÖ¸±ê¾ßÓÐÏàËÆÐÔ¡£
¡¡¡¡
¡¡¡¡Ìá¸ß×÷ÎﵪЧÂʶԲúÁ¿Ìá¸ßÊÇÃ÷ÏԵģ¬ ÄÇô¸ÄÉÆ×÷ÎïµÄÄ͵͵ªÐÔ¶ÔÌá¸ß×÷ÎïµÄ²úÁ¿µÄ¹±Ï×ÔÚÓÚ£¬ TollenaarµÈ[83]Ñо¿±íÃ÷£¬ ÓñÃײúÁ¿µÄÌá¸ß½Ï´ó³Ì¶ÈÉÏÊÇÓÉÓÚÓñÃ׶ԵªÐ²ÆȵÄÄÍÊÜÐÔµÄÌá¸ß¡£ÓýÖÖÕß»òÖÖÖ²ÕßÒÔ×·Çó×îÖյIJúÁ¿ÎªÄ¿µÄ£¬ Òò´Ëɸѡָ±êÓ¦ÒÔ¾­¼Ã²úÁ¿ÎªÖ÷£¬ »òÓë²úÁ¿ÒÅ´«Ïà¹ØÐԸߵÄÖ¸±êΪ×î¼Ñ¡£¶øɸѡµÄʱÆÚÓ¦¸ÃÄÜ´ú±íÕû¸öÉúÓýÆÚµÄÇé¿ö£¬ ÌرðÊǶÔ×÷ÎïÄ͵͵ªÐÔµÄɸѡ£¬ Ó¦¸Ã¿¼Âǵ½×÷ÎïÔÚ²»Í¬µÄÉú³¤·¢ÓýʱÆÚÊܵ½Äæ¾³ £¨µª£© вÆȶÔ×îÖÕ²úÁ¿µÄÓ°ÏìÊÇÃ÷ÏÔ²»Í¬µÄ¡£ÈçÃçÆÚµªÐ²ÆÈ»áÓ°Ïìµ½Ö²ÖêµÄ¹âºÏ×÷ÓÃ[84] £¨Ö÷Ҫͨ¹ýÒ¶Ãæ»ý¡¢Ò¶ÂÌËغ¬Á¿µÈ£© ;¿ª»¨ÊÚ·ÛÆڼ䵪вÆÈ»áÓ°Ïìµ½¿ª»¨ÊýÁ¿¼°ÊÚ¾«³É¹¦ÂÊ£»¹ÈÎï¹à½¬ÆÚ¼äµÄµªÐ²ÆÈ»á¼ÓËÙҶƬ˥ÀÏ£¬ ¼õÉÙ¹âºÏ²úÎ ½µµÍǧÁ£Öصȡ£¶ø±ØÐëÐèÒª¿¼ÂǵÄÊÇ£¬ ×÷ÎïijһʱÆÚÔÚ×ÔÈ»»·¾³ÖÐÊܵ½µªÐ²ÆȵĸÅÂʽϸߣ¬ ÇÒÕâһʱÆÚµÄвÆȶÔ×îÖÕ²úÁ¿µÄÓ°Ïì½Ï´ó £¨ÃçÆÚµªÐ²ÆÈ¿ÉÄܶÔ×îÖÕ²úÁ¿Ó°Ïì²¢²»´ó£© , ÕâÑù²Å»áʹÑо¿¸ü¾ßÓÐʵÓÃÐÔ¡£
¡¡¡¡
¡¡¡¡ËäÈ»ÔÚ×÷ÎïÄ͵͵ªÐԺ͵ªÐ§ÂʵĸÅÄîºÍÆÀ¼Û·½·¨¡¢Ä͵͵ªÐԺ͵ªÐ§ÂʵÄɸѡʱÆÚ¡¢É¸Ñ¡Ö¸±êºÍÄ͵͵ªÐÔ¡¢µªÐ§ÂʵÄÀàÐÍ»®·ÖµÈ·½ÃæµÄÑо¿È¡µÃÁËÖØÒª½øÕ¹£¬ µ«ÓÉÓÚ¶Ô×÷ÎﵪÎüÊÕÀûÓõÄ×÷ÓûúÖÆÉв»Ã÷È·£¬ »¹ÄÑÒÔÓ¦ÓÃÓÚÉú²úʵ¼Ê¡£Òò´Ë£¬ ±ØÐë¼ÓÇ¿¶Ô×÷ÎïÄ͵͵ªÐԺ͵ªÐ§ÂʵÄÉúÀí»úÖƺͷÖ×ÓÉúÎïѧ»úÀíÑо¿£¬ Ϊɸѡ³öÄ͵͵ª¡¢¸ß²úµª¸ßЧÁ½Õß¼æ¹ËµÄÆ·ÖÖ£¬ ·¢Õ¹ÂÌÉ«¡¢¸ßЧũҵµì¶¨»ù´¡¡£
¡¡¡¡
¡¡¡¡²Î¿¼ÎÄÏ×
¡¡¡¡[1]Peng S, Buresh R J, Huang J, Huang J, Zhong X, Zou Y, Yang J, Wang G, Liu Y, Hu R, Tang Q, Cui K, Zhang F, Dobermann A.Improving nitrogen fertilization in rice by site-specific Nmanagement.A review[J].Agronomy for Sustainable Development, 2010, 30 £¨3£© :649-656
¡¡¡¡[2]Cheng X J, Roland J, Buresh R J, Wang Z Q, Zhang H, Liu L J, Yang J C, Zhang J H.Root and shoot traits for rice varieties with higher grain yield and higher nitrogen use efficiency at lower nitrogen rates application[J].Field Crops Research, 2015, 175 £¨4£© :47-55
¡¡¡¡[3]ÂÀ±ù£¬ ·¶ÖÙÇ䣬 ³£Ðñºç£¬ ÍõµÂ÷£¬ ÌÕ־ǿ£¬ ÑîÓñË«£¬ Õű£¾ü£¬ ÕÔ¹ã²Å¡£Ê©µªÁ¿¶Ô2¸öÁ£É«Ð¡Âó²úÁ¿¼°¼Ó¹¤Æ·ÖʵÄÓ°Ïì[J].ºËũѧ±¨£¬ 2017, 31 £¨6£© :1192-1199
¡¡¡¡[4]ÅËÓ££¬ ÕÅÒÇƽ£¬ ÖìÃô»Û£¬ Ò¶·á£¬ ÁÖÊËÐÛ£¬ »ÆÃɻۣ¬ ͯÔÙ¿µ£¬ ÕÅ¿¡ºì¡£¹âƤèëmiR393¼°Æä°Ð»ùÒòÔڵ͵ªÐ²ÆÈÖеıí´ï·ÖÎö[J].ºËũѧ±¨£¬ 2017, 31 £¨10£© :1921-1930
¡¡¡¡[5]Humbert S, Subedi S, Cohn J, Zeng B, Bi Y M, Chen X, Zhu T, Mc Nicholas P D, Rothstein S J.Genome-wide expression profiling of maize in response to individual and combined water and nitrogen stresses[J].BMC Genomics, 2013, 14 £¨1£© :3
¡¡¡¡[6]Ladha J K, Chakraborty D.Nitrogen and cereal production:Opportunities for enhanced efficiency and reduced N losses[C]//Solutions to Improve Nitrogen Use Efficiency for the World.Melbourne:Proceedings of the 2016 International Nitrogen Initiative Conference, 2016:4-8
¡¡¡¡[7]Raun W R, Solie J B, Johnson G V, Marvin L.Stone M L, Mullen R W, Freemana K W, Thomasona W E, Lukinaa E V.Improving nitrogen use efficiency in cereal grain production with optical sensing and variable rate application[J].Agronomy Journal, 2002, 94 £¨4£© :815-820
¡¡¡¡[8]Good A G, Shrawat A K, Muench D G.Can less yield more?Is reducing nutrient input into the environment compatible with maintaining crop production?[J].Trends in Plant Science, 2004, 9 £¨12£© :597-605
¡¡¡¡[9]Peoples M B, Brockwell J, Herridge D F, Rochester I J, Alves B JR, Urquiaga S, Boddey R M, Dakora F D, Bhattarai S, Maskey SL, Sampet C, Rerkasem B, Khan D F, Hauggaard-Nielsen H, Jensen E S.The contributions of nitrogen-fixing crop legumes to the productivity of agricultural systems[J].Symbiosis, 2009, 48 £¨1/2/3£© :1-17
¡¡¡¡[10]Schepers J S, Below F E.Influence of corn hybrids on nitrogen uptake and utilization efficiency[C]//Report of annual Corn and Sorghum Research Conference.USA:American Seed Trade Assoc, 1987:172-186
¡¡¡¡[11]Peng S, Tang Q, Zou Y.Current status and challenges of rice production in China[J].Plant Production Science, 2009, 12 £¨1£© :3-8
¡¡¡¡[12]Chen X P, Cui Z L, Fan M S, Vitousek P, Zhao M, Ma W Q, Wang Z L, Zhang W J, Yan X Y, Yang J C, Deng X P, Gao Q, Guo S W, Ren J, Li S Q, Ye Y L, Wang Z H, Huang J L, Tang QY, Sun Y X, Peng X L, Zhang J W, He M R, Zhu Y J, Xue J Q, Wang G L, An N, Wu L Q, Ma L, Zhang W F, Zhang F S.Producing more grain with lower environmental costs[J].Nature, 2014, 514 £¨7523£© :486-491
¡¡¡¡[13]Rao I M, Miles J W, Beebe S E, Horst W J.Root adaptations to soils with low fertility and aluminium toxicity[J].Annals of Botany, 2016, 118:593-605
¡¡¡¡[14]GirondéA, Poret M, Etienne P, Trouverie J, Bouchereau A, Cahérec L F, Leport L, Orsel M, Niogret M F, Deleu C, Avice JC.A profiling approach of the natural variability of foliar Nremobilization at the rosette stage gives clues to understand the limiting processes involved in the low N use efficiency of winter oilseed rape[J].Journal of Experimental Botany, 2015, 66 £¨9£© :2461-2473
¡¡¡¡[15]»ÆÔª¾¼£¬ ÕÅÉú½Ü£¬ ÂíÓÀ½¨£¬ ÑîÌúîÈ¡£²»Í¬¿¾ÑÌÆ·ÖÖ £¨»ùÒòÐÍ£© µªÐ§Âʼ°Ä͵͵ªÄÜÁ¦µÄ²îÒì[J].Ñ̲ݿƼ¼£¬ 2013 £¨4£© :71-77
¡¡¡¡[16]Inthapanya P, Sihavong P, Sihathep V, Chanphengsay M, Fukai S, Basnayake J.Genotype differences in nutrient uptake and utilisation for grain yield production of rainfed lowland rice under fertilised and non-fertilised conditions[J].Field Crops Research, 2000, 65 £¨1£© :57-68
¡¡¡¡[17]ñҹ⡣²»Í¬Ë®·Ö¡¢Ñø·ÖÀûÓÃЧÂÊË®µ¾Æ·ÖֵĸùϵÌØÕ÷¼°Æäµ÷¿Ø¼¼Êõ[D].ÑïÖÝ£ºÑïÖÝ´óѧ£¬ 2016
¡¡¡¡[18]Harvey P H.Hereditary variation in plant nutrition[J].Genetics, 1939, 24 £¨4£© :437
¡¡¡¡[19]Singh U, Ladha J K, Castillo E G, Punzalan G, Tirol-Padre A.Genotypic variation in nitrogen use efficiency in medium and longduration rice[J].Field Crops Research, 1998, 58 £¨1£© :35-53
¡¡¡¡[20]ÀîÃô£¬ Õźé³Ì£¬ ÑîÐÛ£¬ ¸ðÃÎæ¼£¬ ÂíȺ£¬ κº£Ñ࣬ ´÷Æä¸ù£¬ »ôÖÐÑó£¬ Ðíé𣬠²ÜÀûÇ¿£¬ ÎâºÆ¡£Ë®µ¾¸ß²úµª¸ßЧÐÍÆ·ÖֵĸùϵÐÎ̬ÉúÀíÌØÕ÷[J].×÷Îïѧ±¨£¬ 2012, 38 £¨4£© :648-656
¡¡¡¡[21]ÀîÊçÎÄ£¬ ÖÜÑåÕ䣬 Îĺê´ï£¬ ÀîÑãÃù£¬ Ф¿­¡£²»Í¬Ð¡ÂóÆ·ÖÖµªÐ§ÂʺͲúÁ¿ÐÔ×´µÄÑо¿[J].Ö²ÎïÒÅ´«×ÊԴѧ±¨£¬ 2006, 7 £¨2£© :204-208
¡¡¡¡[22]ÀîÇ¿£¬ ÂÞÑӺ꣬ Ì·½Ü£¬ ¿×·²ÀÚ£¬ ÑîÊÀÃñ£¬ Ô¬¼Ì³¬¡£ÓñÃ×ÔÓ½»ÖÖÃçÆÚÄ͵͵ªÖ¸±êµÄɸѡÓë×ÛºÏÆÀ¼Û[J].ÖйúÉú̬ũҵѧ±¨£¬ 2014, 22 £¨10£© :1190-1199
¡¡¡¡[23]²ÜÃô½¨£¬ ÒÂÓ¨£¬ Ù¡Õ¼²ý£¬ ¹¬¹ú°²¡£Ä͵͵ªÐ²ÆÈÓñÃ×µÄɸѡÓëÆÀ¼Û[J].ÓñÃ׿Æѧ£¬ 2000, 8 £¨4£© :64-69
¡¡¡¡[24]Áº¾°Ï¼£¬ Áº¿µåÉ£¬ ÆÃñ£¬ ÁÖÎÄÐÛ£¬ ³Â˳»Ô£¬ Çð¹óÊ¢¡£Ñ̲ݲ»Í¬»ùÒòÐÍÄ͵͵ªÄÜÁ¦²îÒìÆÀ¼Û[J].Ö²ÎïÒÅ´«×ÊԴѧ±¨£¬ 2007, 8 £¨4£© :451-455
¡¡¡¡[25]²ÜÀ¼ÇÛ£¬ ÎéÏþÃ÷£¬ ÀîÑǾü£¬ ³Â±ÌÔÆ£¬ ³ÂÏÊÄÝ£¬ ÌïÑôÑô£¬ ¸ßÑǾü¡£ÓͲ˵ªËØÎüÊÕЧÂʵĻùÒòÐͲîÒì¼°ÆäÓëÅ©ÒÕÐÔ×´µÄ¹Øϵ[J].ÖйúÓÍÁÏ×÷Îïѧ±¨£¬ 2010, 32 £¨2£© :270-278
¡¡¡¡[26]ÁõÇ¿£¬ Ëκ£ÐÇ£¬ ÈÙÏæÃñ£¬ Åí½¨Î°£¬ л¹ðÏÈ¡£²»Í¬Æ·ÖÖÓͲ˵ªÐ§ÂʲîÒì¼°ÆäÉúÀí»ù´¡Ñо¿[J].Ö²ÎïÓªÑøÓë·ÊÁÏѧ±¨£¬ 2008, 14 £¨1£© :113-119
¡¡¡¡[27]Ò¦Á¢ÈØ£¬ лÀÙÀÙ£¬ Ç¿ÐÀ£¬ Íô¾ü³É£¬ ˾¶þ¾²£¬ ÈÎÅÎÈÙ£¬ ÕÔÏòÌ ÀîÝá´º£¬ ÂíСÀÖ£¬ ÃÏÑÇÐÛ£¬ Íõ»¯¿¡¡£´óÂó²»Í¬µªÀûÓÃЧÂÊÆ·ÖÖɸѡ¼°GS2»ùÒòµÄ±í´ï·ÖÎö[J].ºËũѧ±¨£¬ 2017, 31 £¨7£© :1255-1262
¡¡¡¡[28]Rehman S, Harris P J C, Ashraf M.Stress environments and their impact on crop production[M]//Food Products Press.Abiotic Stresses:Plant Resistance through Breeding and Molecular Approaches.New York:Howarth, 2005:3-18
¡¡¡¡[29]Áõ»Ô£¬ ÕÔÖñÇà¡£Ö²ÎﵪӪÑø¸ßЧ»ùÒòÐÍɸѡָ±êÑо¿½øÕ¹[J].°²»ÕÅ©Òµ¿Æѧ£¬ 2006, 34 £¨14£© :3265-3267, 3272
¡¡¡¡[30]Íõ½ø¾ü£¬ »ÆÈ𶬡£ÓñÃ×µªÐ§Âʼ°ÆäÑо¿½øÕ¹[J].ÓñÃ׿Æѧ£¬ 2005, 13 £¨1£© :89-92
¡¡¡¡[31]Moll R H, Kamprath E J, Jackson W A.Analysis and interpretation of factors which contribute to efficiency of nitrogen utilization[J].Agronomy Journal, 1982, 74 £¨3£© :562-564
¡¡¡¡[32]³Â·¶¿¥£¬ Ã×¹ú»ª£¬ ÕŸ£Ëø£¬ ÍõÑÞ£¬ ÁõÏòÉú£¬ ´ºÁÁ¡£»ª±±Çø²¿·ÖÖ÷ÔÔÓñÃ×ÔÓ½»ÖֵĵªÐ§ÂÊ·ÖÎö[J].ÓñÃ׿Æѧ£¬ 2003, 11 £¨2£© :78-82
¡¡¡¡[33]ÖÓ˼ÈÙ£¬ ³ÂÈÊÏö£¬ ÌÕÑþ£¬ ¹¨Ë¿Ó꣬ ºÎ¿íÐÅ£¬ ÕÅÊÀ´¨£¬ ÕÅÆôÃ÷£¬ ÁõÆëÔª¡£²»Í¬Ñ̲ݻùÒòÐ͵ªËØÎüÊÕЧÂÊÓëÀûÓÃЧÂʲîÒì[J].ÖйúÑ̲ݿÆѧ£¬ 2017, 38 £¨4£© :58-63
¡¡¡¡[34]Fotyma E, Fotyma M.The agronomical and physiological efficiency of nitrogen applied for arable crops in Poland[J].Fertilizer Research, 1995, 43 £¨1/2/3£© :9-12
¡¡¡¡[35]ÁõÅô£¬ ½¹ÏþÑ࣬ ¶¡Óñ´¨£¬ Îä°®Á«£¬ Íõ¾¢ËÉ£¬ ¶­¶þ࣬ ¹ù¬B£¬ ÍõÁ¢¸ï¡£×÷ÎﵪËظßЧÀûÓÃÑо¿½øÕ¹[J].ɽÎ÷Å©Òµ¿Æѧ£¬ 2017, 45 £¨5£© :855-860
¡¡¡¡[36]Francisco M, Pangirayi T, John D.S1 selection of localmaize landraces for low soil nitrogen tolerance in Zambia[J].African Journal of Plant Science, 2010, 4:67-81
¡¡¡¡[37]Wu Y S, Liu W G, Li X H, Li M S, Zhang D G, Hao Z F, Weng J F, Xu Y B, Bai L, Zhang S H, Xie C X.Low-nitrogen stress tolerance and nitrogen agronomic efficiency among maize inbreds:comparison of multiple indices and evaluation of genetic variation[J].Euphytica, 2011, 180 £¨2£© :281-290
¡¡¡¡[38]Àµ¤£¬ ÌïÃÎÓ꣬ ´Þ껣¬ ´÷Í¢²¨£¬ ½ª¶«£¬ ¾£Æ棬 ²ÜÎÀÐÇ¡£Ð¡ÂóÃçÆÚÄ͵͵ªÐ²ÆȵĻùÒòÐͲîÒì[J].ÂóÀà×÷Îïѧ±¨£¬ 2009, 29 £¨2£© :222-227
¡¡¡¡[39]ÕÔ»¯Ì ÍõÈð·¼£¬ ÐíÔƷ壬 °²µ÷¹ý¡£Ð¡ÂóÃçÆÚÄ͵͵ª»ùÒòÐ͵ÄɸѡÓëÆÀ¼Û[J].ÖйúÉú̬ũҵѧ±¨£¬ 2011, 19 £¨5£© :1199-1204
¡¡¡¡[40]Fischer R A, Maurer R.Drought resistance in spring wheat cultivars.I.grain yield responses[J].Crop and Pasture Science, 1978, 29 £¨5£© :897-912
¡¡¡¡[41]Ïò´ºÑô£¬ ³£Ç¿£¬ ÂíÐËÁÖ£¬ ¹ØÒåУ¬ Áè±ÌÓ¨£¬ Õű¦Ê¯¡£ÓñÃײ»Í¬»ùÒòÐͶԵªÓªÑøвÆȵķ´Ó¦[J].ºÚÁú½­°Ëһũ¿Ñ´óѧѧ±¨£¬ 2002, 14 £¨4£© :5-7
¡¡¡¡[42]Ö£¼Ò¿ü£¬ ÎÄ´ºÑô£¬ ÕÅÌΣ¬ ÑîÀò£¬ ÑîǬ»ª£¬ Ö£½¨Ãô£¬ ÑîËÉÌΣ¬ Ìƽ­ÔÆ£¬ Ìï´ä¡£Ä͵͵ªË®µ¾²ÄÁÏɸѡ¼°É¸Ñ¡Ö¸±êÑо¿[J].°²»ÕÅ©Òµ¿Æѧ£¬ 2009, 37 £¨16£© :7361-7363, 7377
¡¡¡¡[43]ÑîÖÐÒ壬 ÕÅ·¢Ã÷£¬ ÀîÓÀÖÇ£¬ Ф־У¬ ÀîÓÀÁÁ£¬ ÁõÀòÀò£¬ ËÎÅô·É£¬ Öܼ½ºâ¡£²»Í¬¿¾ÑÌÆ·ÖÖ¶ÔµªËØвÆÈÏìÓ¦²îÒìµÄÑо¿[J].ÔÆÄÏÅ©Òµ´óѧѧ±¨ £¨×ÔÈ»¿Æѧ°æ£© , 2011, 26 £¨2£© :240-245
¡¡¡¡[44]»ÆÅ©ÈÙ£¬ ÖÓÐñ»ª£¬ Ö£º£²¨¡£Ë®µ¾µª¸ßЧ»ùÒòÐͼ°ÆäÆÀ¼ÛÖ¸±êµÄɸѡ[J].Öйúũѧͨ±¨£¬ 2006, 22 £¨6£© :29-34
¡¡¡¡[45]½­Á¢¸ý£¬ ´÷Í¢²¨£¬ ΤÉÆÇ壬 ¸ÊÐãÇÛ£¬ Ð콨ÔÆ£¬ ²ÜÎÀÐÇ¡£ÄÏ·½Ë®µ¾µªËØÎüÊÕÓëÀûÓÃЧÂʵĻùÒòÐͲîÒì¼°ÆÀ¼Û[J].Ö²ÎïÉú̬ѧ±¨£¬ 2003, 27 £¨4£© :466-471
¡¡¡¡[46]ÈîÐÂÃñ£¬ Ê©·üÖ¥£¬ ÂÞÖ¾Ï飬 Ùܵº졣ˮµ¾ÃçÆÚµª¸ßЧƷÖÖÆÀ¼ÛÓëɸѡµÄ³õ²½Ñо¿[J].Öйúµ¾Ã×£¬ 2010, 16 £¨2£© :22-25
¡¡¡¡[47]³Â¶þÓ°£¬ ÑîÑÓ±ø£¬ ÇØÁ룬 ÕÅ»ªÎÄ£¬ Áõ±ö£¬ Íõº£Á«£¬ ³Â¹ðÁᣬ ÓÚÊçæ㬠¹ÜÑÓ°²¡£¹È×ÓÃçÆÚµª¸ßЧƷÖÖɸѡ¼°Ïà¹ØÌØÐÔ·ÖÎö[J].ÖйúÅ©Òµ¿Æѧ£¬ 2016, 49 £¨17£© :3287-3297
¡¡¡¡[48]½ªË¬¡£²»Í¬µªÐ§ÂʻƹÏÆ·ÖÖɸѡ¼°Æä¸ùϵÉúÎïѧ²îÒì[D].¹þ¶û±õ£º¶«±±Å©Òµ´óѧ£¬ 2012
¡¡¡¡[49]½ªË¬£¬ Îâ·ïÖ¥£¬ ¹ØËÌÄÈ£¬ ÓÚÃôÈñ¡£Ä͵͵ªÐ²ÆȻƹÏÆ·ÖÖµÄɸѡ[J].ÖйúÊ߲ˣ¬ 2012 £¨8£© :51-56
¡¡¡¡[50]²Ü¹ðÀ¼£¬ ÕÅæÂ棬 ÆÓÖÓÔó£¬ º«ÁúÖ²¡£Ë®µ¾²»Í¬»ùÒòÐÍÄ͵͵ªÄÜÁ¦²îÒìÆÀ¼Û[J].Ö²ÎïÒÅ´«×ÊԴѧ±¨£¬ 2006, 7 £¨3£© :3316-3320
¡¡¡¡[51]ºÂÇàÄÏ£¬ Íõ³Ì£¬ ³ÂË®Á«£¬ ɳ°®»ª£¬ µ¥Ö¾»Û£¬ ³Âº£·å£¬ ÖÜÈØ£¬ ÖÜа²¡£´ó¶¹ÃçÆÚµª¸ßЧºÍµªÃô¸Ð×ÊÔ´µÄɸѡÑо¿[J].´ó¶¹¿Æѧ£¬ 2011, 30 £¨6£© :910-915, 920
¡¡¡¡[52]ÀîÁº£¬ »Æ½£»ª£¬ ³Â־࣬ ¶ÅÖ¾îÈ£¬ ¸ßÈóºì£¬ ×ÞÀÚ£¬ ºÎæá£×÷ÎïÄ͵͵ªµÄÏà¹ØÉúÎïѧÑо¿½øÕ¹[J].ÉϺ£Å©ÒµÑ§±¨£¬ 2012, 28 £¨2£© :117-122
¡¡¡¡[53]ÕÅÎýÖÞ£¬ ÑôÏÔ±ó£¬ ÀîÍ¢Ðù£¬ ÓຣӢ¡£Ð¡ÂóµªËØÀûÓÃЧÂʵĻùÒòÐͲîÒì[J].Ó¦ÓÃÉú̬ѧ±¨£¬ 2011, 22 £¨2£© :369-375
¡¡¡¡[54]ÕÔ´º²¨£¬ ËÎÊöÒ¢£¬ ÕÔ¾¸£¬ ÕÅѩ÷£¬ ÕÅÔ½£¬ ÕÅËÉæᣱ±·½µØÇø²»Í¬»Æ¹ÏÆ·ÖÖµªËØÎüÊÕÓëÀûÓÃЧÂʵIJîÒì[J].ÖйúÅ©Òµ¿Æѧ£¬ 2015, 48 £¨8£© :1569-1578
¡¡¡¡[55]ÕÔ¸¶½­£¬ Û§Àö¾ê£¬ ÕÅΡΡ£¬ ³Âѩƽ£¬ ÂÞ˫ϼ¡£ÇÑ×Óµª¸ßЧ»ùÒòÐÍÃçÆÚɸѡָ±êµÄÑо¿[J].»ª±±Å©Ñ§±¨£¬ 2015, 30 £¨S1£© :441-445
¡¡¡¡[56]»ÆÓÀÀ¼£¬ Àèë룬 «Ã÷£¬ Íò½¨ÁÖ£¬ ÁúÆðÕÁ£¬ Íõ»áÃñ£¬ ÌÆÐãÓ¢£¬ ·¶Ö¾½à¡£µª¸ßЧˮµ¾ÖÖÖÊ×ÊԴɸѡ¼°Ïà¹ØÌØÐÔ·ÖÎö[J].Ö²ÎïÒÅ´«×ÊԴѧ±¨£¬ 2015, 16 £¨1£© :87-93
¡¡¡¡[57]ÕÅ¿¡¹ú£¬ ÕÅÈýÔª£¬ Ñî´º¸Õ£¬ ¹ù¹ðÕ䣬 ËïÇ¿¡£Ë®µ¾µª¸ßЧƷÖÖ×ÊԴɸѡµÄ³õ²½Ñо¿¢ñ¡£²»Í¬Ê©µªË®Æ½ÏÂË®µ¾²úÁ¿ºÍ¸ÉÎïÖÊÉú²úµÄÆ·ÖÖ¼ä²îÒì[J].¼ªÁÖÅ©Òµ¿Æѧ£¬ 2008, 33 £¨6£© :7-10
¡¡¡¡[58]ÖÓ˼ÈÙ£¬ ³ÂÈÊÏö£¬ ÌÕÑþ£¬ ¹¨Ë¿Ó꣬ ºÎ¿íÐÅ£¬ ÕÅÆôÃ÷£¬ ÕÅÊÀ´¨£¬ ÁõÆëÔª¡£Ä͵͵ªÑ̲ݻùÒòÐ͵Äɸѡ¼°Æ䵪ЧÂÊÀàÐÍ[J].×÷Îïѧ±¨£¬ 2017, 43 £¨7£© :993-1002
¡¡¡¡[59]Zieserl J F, Rivenbark W L, Hageman R H.Nitrate reductase activity, protein content, and yield of four maize hybrids at varying plant populations[J].Crop Science, 1963, 3 £¨1£© :27-32
¡¡¡¡[60]Magalhaes J R, Purcino A A C, Machado A T, Maganavaca R, Fran9a G E, Magalh2es P C.Study of physiological parameters for the selection of maize genotypes efficient in ammonium uptake[J].Documentos-Empresa Capixaba de Pesquisa Agropecuária, 1990, 65:104
¡¡¡¡[61]Machado A T, Magalhaes J R, Magnavaca R, Silva MRe.Enzyme activities involved with nitrogen metabolism in different maize genotypes[J].Revista Brasileira de Fisiologia Vegetal £¨Brazil£© , 1992, 4 £¨1£© :45-47
¡¡¡¡[62]²ÜÃô½¨£¬ ÒÂÓ¨£¬ Ù¡Õ¼²ý£¬ ¹¬¹ú°²¡£Ä͵͵ªÐ²ÆÈÓñÃ×µÄɸѡÓëÆÀ¼Û[J].ÓñÃ׿Æѧ£¬ 2000, 8 £¨4£© :64-69
¡¡¡¡[63]´ÞÎÄ·¼£¬ ¸ß¾ÛÁÖ£¬ ÍõÖ¾¸Õ£¬ ´Þ³¬£¬ ºúÊ÷ƽ£¬ ÓÚÏþ·¼£¬ Ëï¼ÌÓ±£¬ ËÕÖξü¡£ÓñÃ××Ô½»ÏµµªÐ§ÂÊ»ùÒòÐͲîÒì·ÖÎö[J].ÓñÃ׿Æѧ£¬ 2013, 21 £¨3£© :6-12
¡¡¡¡[64]Gerloff G C.Intact-plant screening for tolerance of nutrientdeficiency stress[J].Plant and Soil, 1987, 99 £¨1£© :3-16
¡¡¡¡[65]Blair G.Nutrient efficiency-what do we really mean?[M]Randall P J, Delhaize E, Richards R A, Munns R.Genetic Aspects of Plant Mineral Nutrition.Berlin:Springer Netherlands, 1993:205-213
¡¡¡¡[66]Maia C, Do Vale J C, Fritsche-Neto R, Cavatte P C, Miranda G V.The difference between breeding for nutrient use efficiency and for nutrient stress tolerance[J].Crop Breeding and Applied Biotechnology, 2011, 11 £¨3£© :270-275
¡¡¡¡[67]÷·¼¡£¼Þ½Ó¶Ô¿¾Ñ̵ªËØÎüÊÕÀûÓõÄÓ°Ïì[D].Ö£ÖÝ£ººÓÄÏÅ©Òµ´óѧ£¬ 2013
¡¡¡¡[68]Ruiz J M, Rivero R M, Cervilla L M, Castellano R, Romero L.Grafting to improve nitrogen-use efficiency traits in tobacco plants[J].Journal of the Science of Food and Agriculture, 2006, 86 £¨6£© :1014-1021
¡¡¡¡[69]ÐíÈ«±¦¡£²»Í¬»Æ¹ÏÕèľµªËØÀûÓÃЧÂÊÆÀ¼Û¼°Õèľ¼Þ½Ó¶Ô»Æ¹ÏÓ×ÃçµÄÓ°Ïì[D].Î人£º»ªÖÐÅ©Òµ´óѧ£¬ 2011
¡¡¡¡[70]Mackay A D, Barber S A.Effect of nitrogen on root growth of two corn genotypes in the field[J].Agronomy Journal, 1986, 78 £¨4£© :699-703
¡¡¡¡[71]Kichey T, Heumez E, Pocholle D, Pageau K, Vanacker H, Dubois F, Le Gouis J, Hirel B.Combined agronomic and physiological aspects of nitrogen management in wheat highlight a central role for glutamine synthetase[J].New Phytologist, 2006, 169 £¨2£© :265-278
¡¡¡¡[72]Sun H, Huang Q M, Su J.Highly effective expression of glutamine synthetase genes GS1 and GS2 in transgenic rice plants increases nitrogen-deficiency tolerance[J].Journal of Plant Physiology and Molecular Biology, 2005, 31 £¨5£© :492-498
¡¡¡¡[73]Crawford N M, Glass A D M.Molecular and physiological aspects of nitrate uptake in plants[J].Trends in Plant Science, 1998, 3 £¨10£© :389-395
¡¡¡¡[74]Sohlenkamp C, Wood C C, Roeb G W, Udvardi M K.Characterization of Arabidopsis At AMT2, a high-affinity ammonium transporter of the plasma membrane[J].Plant Physiology, 2002, 130 £¨4£© :1788-1796
¡¡¡¡[75]Yu X, Sukumaran S, Márton L.Differential expression of the Arabidopsis Nia1 and Nia2 genes cytokinin-Induced nitrate reductase activity is correlated with increased Nia1 transcription and mRNAlevels[J].Plant Physiology, 1998, 116 £¨3£© :1091-1096
¡¡¡¡[76]Ochs G, Schock G, Trischler M, et al.Complexity and expression of the glutamine synthetase multigene family in the amphidiploid crop Brassica napus[J].Plant Molecular Biology, 1999, 39 £¨3£© :395-405
¡¡¡¡[77]Masclaux-Daubresse C, Daniel-Vedele F, Dechorgnat J, Chardon F, Gaufichon L, Suzuki A.Nitrogen uptake, assimilation andremobilization in plants:challenges for sustainable and productiveagriculture[J].Annals of Botany, 2010, 105 £¨7£© :1141-1157
¡¡¡¡[78]Gallais A, Hirel B.Anapproachof thegeneticsofnitrogen use efficiencyin maize[J].Journal of Experimental Botany, 2004, 55 £¨396£© :295-306
¡¡¡¡[79]Fan S C, Lin C S, Hsu P K, Lin S H, Tsay Y F.The Arabidopsis nitrate transporter NRT1.7, expressed inphloem is responsiblefor source-to-sink remobilization ofnitrate[J].The Plant Cell, 2009, 21 £¨9£© :2750-2761
¡¡¡¡[80]Òü²ý±ó£¬ ÕÔ¿¡Î°£¬ ÓÈ·É£¬ ÔøÏ͸գ¬ ³Â¸·¡£»ùÓÚÉú̬ÎÄÃ÷µÄÅ©ÒµÏÖ´ú»¯·¢Õ¹²ßÂÔÑо¿[J].Öйú¹¤³Ì¿Æѧ£¬ 2015, 17 £¨8£© :97-102
¡¡¡¡[81]Lynch J.The role of nutrient-efficient crops in modern agriculture[J].Journal of Crop Production, 1998, 1 £¨2£© :241-264
¡¡¡¡[82]Le G J, Béghin D, Heumez E, Pluchard P.Genetic differences for nitrogen uptake and nitrogen utilisation efficiencies in winter wheat[J].European Journal of Agronomy, 2000, 12 £¨3£© :163-173
¡¡¡¡[83]Tollenaar M, Wu J.Yield improvement in temperate maize is attributable to greater stress tolerance[J].Crop Science, 1999, 39 £¨6£© :1597-1604
¡¡¡¡[84]Sinclair T R, Vadez V.Physiological traits for crop yield improvement in low N and P environments[J].Plant&Soil, 2002, 245 £¨1£© :1-15
Ô­Îijö´¦£ºÖÓ˼ÈÙ,¹¨Ë¿Óê,ÕÅÊÀ´¨,³ÂÈÊÏö,ÁõÆëÔª,µÔСÇå.×÷Îﲻͬ»ùÒòÐÍÄ͵͵ªÐԺ͵ªÐ§ÂÊÑо¿½øÕ¹[J].ºËũѧ±¨,2018,32(08):1656-1663.
Ïà¹ØÄÚÈÝÍƼö
Ïà¹Ø±êÇ©£º
·µ»Ø£ºÅ©ÒÕѧÂÛÎÄ