木本植物低温胁迫生理及分子机制研究进展
목본식물저온협박생리급분자궤제연구진전
Research Progress on the Physiological and Molecular Mechanisms of Woody Plants Under Low Temperature Stress
  • 万方数据
    林业科学 임업과학
    Scientia Silvae Sinicae
    2015年 7期 116-128 ,共13页

    乌凤章%王贺新%徐国辉%张自川

    오봉장%왕하신%서국휘%장자천

    木本植物%低温胁迫%抗冷性%抗冻性%低温锻炼 木本植物%低溫脅迫%抗冷性%抗凍性%低溫鍛煉
    목본식물%저온협박%항랭성%항동성%저온단련
    woody plants%low temperature stress%chilling resistance%freezing tolerance%low temperature acclimation
    低温胁迫限制了许多野生植物和作物的地理分布，降低了生产率。植物在低温胁迫下生存能力差别很大，来源于热带和亚热带的冷敏感植物在温度高于冰点的低温条件下会发生不可逆的伤害，而温带地区的植物能够承受极端的冰冻条件。近年来有关冷敏感植物和温带植物低温胁迫下生理生化响应特征，转录组学、蛋白质组学和代谢组学分析取得了较快进展，从不同层次阐明了木本植物调节低温反应和抗冷(冻)性的生理和分子机制，为利用抗冷(冻)相关基因进行林木抗(冷)冻分子育种提供了重要参考。冷敏感树木在1～10℃的低温胁迫下，会出现水分状况、矿质营养、光合作用、呼吸作用和新陈代谢等生理过程的紊乱，造成冷害甚至死亡。钙信号途径是低温应答过程中重要的信号转导途径。ABA 通过 ABA 依赖的转录因子的转录激活参与植物胁迫响应基因的调节。低温响应转录调控分为 CBF 途径和非 CBF 途径。CBF 调控下游基因的表达过程：植物细胞膜相关受体首先感知环境信号并传送到细胞核，通过 Ca2+和 MAPKs 等感知和传导信号，诱导转录因子 CBF 的表达，从而激活冷响应基因的启动子，触发冷响应机制(基因激活)，转录的mRNA 被翻译成不同的蛋白质。这些基因产物参与改变膜质成分组成、抗氧化酶活性、渗透物质含量等生理过程，从而提高抗冷性。另外，CBF 基因表达也受到上游一些转录因子的调节，包括钙离子信号途径和 ICE1－CBF －寒冷响应途径。树木抗冷锻炼和抗冻锻炼的机制与代谢途径有许多相似之处，但后者可能更复杂，因为它需要面对极端的低温和不正常的温度波动。抗冻锻炼期间低温诱导的抗冻蛋白具有较强的抑制冰晶重结晶的活性；脱水蛋白和胚胎后期丰富蛋白( LEA)通过渗透调节保护细胞；热休克蛋白( HSPs)调节蛋白质的折叠和运输，恢复钝化酶的活性；抗氧化酶系统清除氧自由基和过氧化氢；早期光诱导蛋白参与高光胁迫适应过程，这些蛋白在树木抗冻机制中具有重要作用。在树木抗冷(冻)领域，未来应加强对控制抗冷(冻)生理变化的转录因子及关键功能基因的全面解析，深入探讨光、温度等环境信号诱导抗冻性形成机制，使用蛋白质组学方法与其他技术相结合阐明低温胁迫下重要蛋白质表达机制和功能，完整地揭示木本植物在低温逆境下的生存机制。
    저온협박한제료허다야생식물화작물적지리분포，강저료생산솔。식물재저온협박하생존능력차별흔대，래원우열대화아열대적랭민감식물재온도고우빙점적저온조건하회발생불가역적상해，이온대지구적식물능구승수겁단적빙동조건。근년래유관랭민감식물화온대식물저온협박하생리생화향응특정，전록조학、단백질조학화대사조학분석취득료교쾌진전，종불동층차천명료목본식물조절저온반응화항랭(동)성적생리화분자궤제，위이용항랭(동)상관기인진행림목항(랭)동분자육충제공료중요삼고。랭민감수목재1～10℃적저온협박하，회출현수분상황、광질영양、광합작용、호흡작용화신진대사등생리과정적문란，조성냉해심지사망。개신호도경시저온응답과정중중요적신호전도도경。ABA 통과 ABA 의뢰적전록인자적전록격활삼여식물협박향응기인적조절。저온향응전록조공분위 CBF 도경화비 CBF 도경。CBF 조공하유기인적표체과정：식물세포막상관수체수선감지배경신호병전송도세포핵，통과 Ca2+화 MAPKs 등감지화전도신호，유도전록인자 CBF 적표체，종이격활랭향응기인적계동자，촉발랭향응궤제(기인격활)，전록적mRNA 피번역성불동적단백질。저사기인산물삼여개변막질성분조성、항양화매활성、삼투물질함량등생리과정，종이제고항랭성。령외，CBF 기인표체야수도상유일사전록인자적조절，포괄개리자신호도경화 ICE1－CBF －한랭향응도경。수목항랭단련화항동단련적궤제여대사도경유허다상사지처，단후자가능경복잡，인위타수요면대겁단적저온화불정상적온도파동。항동단련기간저온유도적항동단백구유교강적억제빙정중결정적활성；탈수단백화배태후기봉부단백( LEA)통과삼투조절보호세포；열휴극단백( HSPs)조절단백질적절첩화운수，회복둔화매적활성；항양화매계통청제양자유기화과양화경；조기광유도단백삼여고광협박괄응과정，저사단백재수목항동궤제중구유중요작용。재수목항랭(동)영역，미래응가강대공제항랭(동)생리변화적전록인자급관건공능기인적전면해석，심입탐토광、온도등배경신호유도항동성형성궤제，사용단백질조학방법여기타기술상결합천명저온협박하중요단백질표체궤제화공능，완정지게시목본식물재저온역경하적생존궤제。
    Low temperature stress limits the geographical distribution of many wild plant and crops,and reduces their productivity. There are huge differences in survivability of trees to low temperature stress. Chilling sensitive trees from tropics and subtropics are irreversibly damaged at temperature condition higher than freezing point,while those species originated from temperate-zone can withstand extreme freezing conditions. In recent years,the analyses of physiological and biochemical response characteristics, transcriptome, proteome and metabolome of chilling sensitive trees and temperate-zone trees under low temperature stress have been rapidly developed,which from various levels have elucidated the physiological and molecular mechanisms that woody plant regulating low-temperature response and freezing tolerance. These analyses provide important reference for molecular breeding of freezing-tolerant woody plant by using anti-freezing related genes. Under the low temperature stress of 1—10 ℃,chilling sensitive trees would appear disorders physiological processes in water status,mineral nutrition,photosynthesis,respiration and metabolism,and hence lead to chilling damage,and even death. Calcium signal transduction pathway is the important signal transduction pathway during low temperature response process. ABA is involved in the regulation of plant stress response gene by transcriptional activation of ABA-dependent transcription factor. Regulation of low temperature response transcription can be divided into CBF pathway and non-CBF pathway. CBF regulates the expression processes of downstream target genes: related receptors of plant cell membrane first perceive environmental signal and transmit it to cell nucleus. The signal is perceived and transducted through Ca2 + and MAPK and induces expression of CBF,which will activate the promoter of cold-response gene and trigger cold response mechanism ( gene activation ) , and the transcribed mRNA is translated into various proteins. These gene products participate in physiological processes of changing the composition of membrane,antioxidant enzyme activities,content of osmotic substances,which will improve cold resistance. On the other hand,CBF gene expression is also regulated by some upstream transcriptional factors,including Ca2 + signal way and ICE1-CBF cold response way. Mechanism and metabolic pathway of chilling acclimation in trees are much similar to freezing acclimation in trees,but the latter may be more complicated,because it needs to face extreme and abnormal temperature variance. The antifreeze protein induced by cold during freezing acclimation has strong activity to inhibit ice recrystallization;Dehydrin and LEA protein protect cellular by osmoregulation; Heat shock protein ( HSP ) regulates protein folding and transportation,and restores activity of modifying enzyme; antioxidant enzyme system scavenges oxygen free radical and hydrogen peroxide; early light-induced protein ( ELIP ) takes parts in the high light stress acclimation process,which plays important roles for antifreeze mechanism of trees. In the field of tree's freezing tolerance,comprehensive analysis of the transcriptional factor and functional genes shall be emphasized in the future; the development mechanism of freezing tolerance induced by environmental signal such as light,temperature,etc. shall be more thoroughly study; expression mechanism and function of important proteins under low temperature stress shall be explained by combination of proteomics methods and other technology to integrally reveal the surviving mechanism of woody plants under low temperature stress.
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