表达二穗短柄草<em> CBF2</em>基因拟南芥抗旱性分析

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张彦妮, 焦孟月, 刘奕佳. 表达二穗短柄草 CBF2基因拟南芥抗旱性分析 . 草业科学, 2016,33(8):1518-1525
Zhang Yan-ni, Jiao Meng-yue, Liu Yi-jia. Drought Resistance of Arabidopsis thaliana expressing CBF2 gene in Brachypodium distachyon . Pratacultural Science,2016,33(8): 1518-1525 复制到剪切板

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《草业科学》编辑部

表达二穗短柄草 CBF2基因拟南芥抗旱性分析

张彦妮, 焦孟月, 刘奕佳

东北林业大学园林学院,黑龙江哈尔滨 150040

通讯作者:张彦妮(1974-),女,山西大同人,副教授,博士,主要从事园林植物繁殖栽培及育种。E-mail:zhangyanni808@126.com

收稿日期: 2016-06-05

基金: 中央高校基本科研业务费专项(DL12CA17); 黑龙江省留学归国基金(LC201410); 黑龙江省教育厅科研项目(12543012)

摘要

以表达二穗短柄草( Brachypodium distachyon) CBF2基因的拟南芥( Arabidopsis thaliana)为材料,通过模拟干旱胁迫分析其种子和幼苗对甘露醇渗透胁迫的响应和成苗对盆栽条件下水分胁迫的响应。结果表明,渗透压为-0.62 MPa时种子萌发率开始下降,转基因拟南芥种子萌发率略高于野生拟南芥,但差异不显著( P>0.05)。以根的增长量作为评价幼苗抗旱性的主要标准,当渗透压小于-0.37 MPa时,根长增长量随着胁迫强度的增加而明显减小,转基因株系根长增长量始终显著高于野生型拟南芥( P<0.05)。拟南芥成苗干旱胁迫至25 d,野生拟南芥成活率为11.7%,转基因植株平均成活率为41.6%;随着干旱胁迫时间的延长,转基因拟南芥过氧化物酶、超氧化物歧化酶活性均显著高于野生植株,丙二醛含量及相对电导率显著低于野生植株。结果表明,二穗短柄草 CBF2基因提高了拟南芥幼苗及成苗的抗旱性。

关键词: CBF2基因; 二穗短柄草; 拟南芥; 抗旱性

中图分类号:Q945.78 文献标志码:A 文章编号:1001-0629(2016)8-1518-08 doi: 10.11829/j.issn.1001-0629.2015-0542

Drought Resistance of Arabidopsis thaliana expressing CBF2 gene in Brachypodium distachyon

Zhang Yan-ni, Jiao Meng-yue, Liu Yi-jia

Landscape Architecture College of Northeast Forestry University, Harbin 150040, China

Corresponding author: Zhang Yan-ni E-mail:zhangyanni808@126.com

Abstract

The Arabidopsis thaliana expressing CBF2 gene in Brachypodium distachyon was used as the experimental material. The response of the germination rate of transgenic A. thaliana seeds and seedlings growth to drought was analyzed by simulated mannitol osmotic stress, and the drought resistance of the mature seedlings was detected by pot experiments. Results showed the seed germination rate declined with the increase of the osmotic pressure, and it become significant when it reached -0.62 MPa. The germination rate of transgenic seed was slightly higher than wild seeds, but the difference was not significant. When the osmotic pressure was higher than -0.37 MPa, the root length increment significantly reduced with the increase of stress intensity, and the root length increment of transgenic plants was significantly higher than wild A. thaliana. The survival rate of wild A. thaliana was 11.7%, while the average survival rate of transgenic plants was 41.6% when the A. thaliana seedlings was dealing with drought stress for 25 days. POD and SOD activity of transgenic A. thaliana was significantly higher than wild plants, and MDA content and relative electrical conductivity were significantly lower than wild plants with the extension of drought stress time. Our results suggested that the CBF2 gene in B. distachyon can significantly enhanced the drought resistance of transgenic A. thaliana seedlings and mature seedlings.

Keyword: CBF2 gene; Brachypodium distachyon; Arabidopsis thaliana; drought resistance

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干旱是影响植物生长的重要环境因子之一, 也是对全球影响最广泛的自然灾害^[1]。我国是一个干旱灾害频发的国家, 据统计每年我国干旱灾害造成的损失超过各种自然灾害损失总和的15%^{[2, 3]}。因此, 提高植物的抗旱性在育种工作中尤为重要。随着分子生物学与基因工程技术的不断发展, 基因工程技术成为改良植株抗旱性、培育抗旱新品种的重要途径之一。相比功能基因, 转录因子在一定条件下可以调控多个逆境相关基因的表达, 因此, 利用转录因子来提高植株的抗逆性将是一条更有效的途径^[4]。AP2、bZIP、NAC、MYB、Cys2/His2锌指等不同类型的转录因子能够调控植物应答干旱胁迫, 这些转录因子通过激活或抑制下游干旱应答基因的表达来调节植物的抗旱性, 且能获得持久的抗逆性, 具有广阔的应用前景^{[5, 6, 7, 8, 9]}。

DREB/CBF(dehydration responsive element binding protein/C-repeat binding factor)属于AP2类转录因子, 被称为低温反应转录因子。CBF基因家族目前发现编码6个相关的转录活化因子, 分别称为CBF1、CBF2、CBF3、CBF4、CBF5和CBF6^{[10, 11]}。当某些植物受到干旱、盐碱、冷胁迫时DREB/CBF上调表达, 可激活其它一系列依赖DRE/CRT顺式作用元件的抗逆功能基因的表达, 从而增强植物对干旱、低温及高盐碱等逆境的抗性, 例如在拟南芥(Arabidopsis thaliana)中超表达的CBF4激活了包含低温驯化及干旱适应的下游基因C-重复/脱水应答元件, 从而提高了植株的抗寒性、抗旱性^{[10, 12]}。目前, CBF家族转录因子抗寒功能及应用的研究比较广泛, 研究表明, DREB/CBF基因的表达可以被多种胁迫诱导, CBF1及CBF3多在盐胁迫下表达, 而CBF2主要在干旱及高盐胁迫下表达^{[13, 14]}。通过对干旱胁迫时CBF 4种基因在不同葡萄(Vitis vinifera)品种中表达模式的研究发现, 干旱条件下CBF2在叶片及根系中均表达, 且相比衰老组织, 幼嫩组织中的表达量较多^{[15, 16]}。此外, 受ABA的诱导, CBF2的表达量也会增加, 原因是植物对ABA的响应促进了CBF启动子活性的提高^{[12, 17, 18]}。CBF2在CBF1和CBF3表达上具有负的调控作用, 并且对拟南芥的耐胁迫性起着重要作用^[18]。

二穗短柄草(Brachypodium distachyon)是禾本科短穗草属一年生野生草本植物, 具有极强的抗寒性, 对干旱、盐碱也有一定的抗性^[19]。目前已证明, 二穗短柄草CBF3转录因子在提高植物的抗寒性方面具有重要作用^[20], 但对二穗短柄草CBF2转录因子抗性的研究较少。本研究以已转入二穗短柄草CBF2基因的拟南芥为研究材料, 通过对比干旱胁迫条件下转基因拟南芥与野生植株形态及生理指标的差异, 探讨该基因对干旱的抗性, 从而为该基因的抗旱性应用及基因作用机理等研究奠定基础, 同时也为培育抗旱新品种提供新思路。

1 材料与方法

1.1 材料

二穗短柄草CBF2转基因拟南芥T₃代种子T₃-3、T₃-6、T₃-9为试验材料, 以野生拟南芥(WT, 由加州大学戴维斯分校Blumwald Eduardo教授赠送的种子的后代)为对照。

1.2 方法

1.2.1 种子消毒取适量拟南芥种子(储存在4 ℃的冰箱中以保证发芽整齐)至离心管中, 先用75%酒精消毒30 s, 无菌水冲洗一次, 再用5%的次氯酸钠消毒12 min, 无菌水冲洗3次。

1.2.2 种子渗透胁迫处理将已消毒的转基因和野生拟南芥种子, 播于渗透压分别为0、-0.12、-0.25、-0.37、-0.49、-0.62、-0.74 MPa的1/2MS培养基上, 放入培养箱(23~24 ℃)中暗培养3 d, 后置于光照培养室培养, 每个处理重复3次, 20 d后统计不同浓度甘露醇下转基因和野生拟南芥种子的萌发率。

萌发率=萌发种子数/播种种子总数× 100%。

1.2.3 幼苗的渗透胁迫处理将在1/2MS培养基培养7 d的转基因与野生拟南芥幼苗移入渗透压分别为0、-0.12、-0.25、-0.37、-0.49、-0.62、-0.74 MPa的1/2MS培养基上, 每个处理重复3次, 垂直培养10 d后, 测量不同浓度甘露醇处理下转基因和野生幼苗的根长增长量。

根长增长量=垂直培养10 d后主根长度-垂直培养10 d前主根长度。

1.2.4 成苗的干旱胁迫处理当幼苗长出4片真叶时将其移入花盆中(培养基质为营养土:蛭石=1:1), 放入长日照培养室中培养。培养30 d后, 停止浇水, 每4 d观察记录转基因和野生拟南芥的形态变化, 24 d时统计其成活率。

成活率=成活植株株数/移栽植株总株数× 100%。

1.2.5 干旱胁迫处理转基因拟南芥成苗的生理指标测定分别取干旱处理0、4、8、12、20 d的转基因和野生拟南芥成苗叶片为待测样品, 每份待测样品取自于一株植物, 每个处理重复3次, 即每个处理取3株植物。采用氮蓝四唑(NBT)光化还原法测定超氧化物歧化酶(SOD)总活性, 采用愈创木酚法测定过氧化物酶(POD)总活性, 采用硫代巴比妥酸法测定丙二醛(MDA)含量, 采用电导仪法测定电导率^[21]。

1.3 数据分析

采用Excel整理数据, 使用SPSS软件对每个测定指标进行单因素方差分析, 然后对同一胁迫条件下不同株系间的数据进行多重比较。

2 结果与分析

2.1 渗透胁迫下转基因拟南芥种子的萌发

当渗透压为0、-0.12、-0.25、-0.37、-0.49 MPa时, 转基因和野生型拟南芥种子萌发率均为100%(图1)。当渗透压达-0.62 MPa时, 野生型拟南芥种子平均萌发率降至76.4%, 转基因拟南芥T₃-3、T₃-6、T₃-9的种子平均萌发率分别为89.3%、93.1%和95.7%(图1), 但转基因株系与野生型差异不显著(P> 0.05)。而此时已萌发的野生型拟南芥与转基因拟南芥株系表型差异明显, 野生型拟南芥幼苗较转基因幼苗更小, 且叶片数量少。转基因拟南芥株系T₃-3、T₃-6、T₃-9之间种子萌发率和表型均无明显差异。当渗透胁迫达-0.74 MPa时, 野生型拟南芥种子萌发率降为33.3%, 而转基因拟南芥株系T₃-3、T₃-6、T₃-9的种子萌发率分别为38.7%、37.1%和39.0%, 转基因与野生型相比差异仍不显著。因此, 转基因拟南芥在种子萌发阶段并未表现出明显强于野生型的抗旱能力。

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图1 甘露醇渗透胁迫对转基因和野生型拟南芥萌发率的影响
注:不同字母表示同一胁迫条件下不同植株材料间差异显著(P< 0.05)。WT, 野生型拟南芥; T₃-3, T₃-6和T₃-9为转基因型拟南芥。图2同。Fig.1 Germination percentage of transgenic and wild type A. thaliana with different osmotic pressure
Note: Different lower case letters under the same stress condition indicate significant difference among different plant materials at 0.05 level. WT, wild A. Tthaliana; T₃-3, T₃-6 and T₃-9 are transgenic A. thaliana. The same in Fig.2.

2.2 渗透胁迫下转基因拟南芥幼苗的形态指标分析

当渗透压为-0.25 MPa时, 转基因拟南芥幼苗的根长增长量与未干旱胁迫下幼苗相比差异不显著(P> 0.05), 野生型幼苗根长增长量明显下降, 其根长增长量显著低于同一渗透压下的转基因幼苗(P< 0.05)(图2)。当渗透压为-0.37、-0.49、-0.62、-0.74 MPa时, 根长增长量随着渗透胁迫的增强而明显减小, 转基因株系的根长增长量始终显著高于野生型拟南芥的(P< 0.05)(图2)。而在渗透压-0.49、-0.74 MPa时, 转基因株系T₃-3的根长增长量显著低于转基因株系T₃-6和T₃-9的(P< 0.05)。随着渗透胁迫的增强, 以未进行干旱胁迫的幼苗为对照, 转基因与野生拟南芥幼苗根增长量不断减少, 植株形态差异不明显。表明, CBF2基因在一定程度上增强了转基因拟南芥在幼苗期的抗旱能力, 且转基因株系T₃-6和T₃-9相对于株系T₃-3的基因表达性状更稳定。

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	图2 甘露醇渗透胁迫对转基因和野生拟南芥根长增长影响Fig.2 Root growth of transgenic and wild type A. thaliana with different osmotic pressure

2.3 转基因拟南芥成苗的抗旱性形态指标分析

水分胁迫结果表明, 干旱8 d时, 野生型拟南芥表型开始发生变化, 其植株叶片开始失绿、变黄, 而转基因拟南芥各株系正常生长(图3)。此后8-20 d, 随着处理时间的增长, 转基因与野生型拟南芥生长状态皆下降, 叶片渐枯、萎蔫, 至处理20 d时, 转基因与野生型拟南芥皆已出现死亡植株, 此时野生型拟南芥的枯黄现象更为严重, 与转基因各株系之间具有一定的表型差异(图3)。24 d时统计成活率, 野生型拟南芥的成活率仅为11.7%, 而转基因拟南芥各株系T₃-3、T₃-6和T₃-9株系成活率分别为38.3%、44.5%和41.9%。表明CBF2基因在增强转基因拟南芥植株抗旱性方面具有一定的作用。

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	图3 自然干旱处理8天和20天下转基因和野生型拟南芥的表型Fig.3 Phenotype of transgenic and wild type A. thaliana with drought for 8 and 20 days

2.4 干旱胁迫处理转基因拟南芥成苗的生理指标分析

2.4.1 干旱胁迫对超氧化物歧化酶(SOD)及过氧化物酶(POD)总活性的影响随着干旱胁迫时间的延长, 转基因各株系和野生型拟南芥的SOD、POD活性均呈现先升高后降低的趋势, 未处理时各株系间酶活性差异不显著(P> 0.05)(图4、5)。胁迫至8 d, 转基因与野生拟南芥SOD、POD的活性达到最高, 随后活性降低, 可能是干旱胁迫下拟南芥启动了其胁迫响应机制。与野生拟南芥相比, 转基因植株SOD、 POD活性始终略高于野生型植株内酶活性, 但差异不显著。

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图4 干旱胁迫对转基因和野生型拟南芥SOD的影响
注:CK1为野生型未处理植株, CK2为转基因株系T₃-9未处理植株。图5、6、7同。Fig.4 SOD activity of transgenic and wild type A. thaliana under drought stress
Note: CK₁ represent wild type A. thaliana of normal growth, CK₂ represent T₃-9 transgenic A. thaliana of normal growth. The same in Fig.5, Fig.6, Fig.7.

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	图5 干旱胁迫对转基因和野生型拟南芥POD的影响Fig.5 POD activity of transgenic and wild type A. thaliana under drought stress

2.4.2 干旱胁迫对丙二醛含量的影响随着干旱胁迫时间的不断延长, 转基因各株系和野生型拟南芥的丙二醛(MDA)含量均呈现不断升高的趋势(图6)。处理8-20 d内, 转基因拟南芥MDA含量始终显著低于野生型拟南芥植株(P< 0.05)。在干旱胁迫下, 转基因拟南芥的细胞膜质过氧化程度始终显著低于野生型拟南芥, 间接说明转基因植株内产生的超氧负离子更少, 干旱胁迫使其受害的程度更小。CBF2基因能够减缓植物呼吸平衡的破环, 即提高了植株的抗旱性。

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	图6 干旱胁迫对转基因和野生型拟南芥MDA的影响Fig.6 MDA content of transgenic and wild type A. thaliana under drought stress

2.4.3 干旱胁迫对电导率的影响随着干旱胁迫时间的延长, 转基因各株系和野生型拟南芥的相对外渗电导率均不断升高(图7)。胁迫8-20 d内, 转基因各株系电导率均显著低于野生型拟南芥(P< 0.05)。与野生拟南芥相比, 转基因拟南芥各株系的相对电导率上升趋势更平缓, 说明转基因拟南芥细胞膜结构稳定性更好, 膜结构受到的功能破坏更小, 即转基因拟南芥抗旱性提高。

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	图7 干旱胁迫对转基因和野生型拟南芥相对电导率的影响Fig.7 Relative electric conductivity of transgenic and wild type A. thaliana under drought stress

3 讨论与结论

在干旱环境下, 种子能否保持活力及幼苗能否继续生长是植物存活的关键, 目前对转基因植株种子萌发期的抗性研究较少, 而种子萌发期抗旱性鉴定是整体抗旱性评价的重要内容^{[22, 23, 24]}。渗透调节是植物适应水分胁迫的主要生理机制, 通过渗透调节植物在干旱条件下维持一定的膨压, 从而维持细胞生长、气孔开放和光合作用等生理过程的进行^[25]。本研究对种子及幼苗抗性的研究采用渗透调节模拟干旱的方法, 研究发现, 干旱胁迫下CBF2转基因拟南芥的种子萌发率略高于野生拟南芥的, 但差异不显著, CBF2基因在改善拟南芥种子的抗旱性上, 没有表现出明显优势。干旱胁迫下, 幼苗根的生长对植物幼苗生长发育具有十分重要的意义^[26]。因此, 本研究以根长增长量作为分析幼苗抗旱性的主要指标。当渗透压达-0.25 MPa时, 转基因拟南芥各株系根长增长量皆显著高于野生型(P< 0.05), 转基因拟南芥显示出较强的抗旱性, 即CBF2基因增强了植株幼苗时期的抗旱能力。

对转基因与野生拟南芥成苗进行相同时间的水分胁迫, 胁迫8和20 d时转基因拟南芥各株系与野生型拟南芥表型差异明显, 24 d时统计成活率, 野生拟南芥成活率为11.7%, 转基因不同株系的平均成活率为41.6%。胁迫条件下, 细胞及酶活性变化以及渗透调节物质累积情况需要依靠植物的生理生化指标进行研究^[27]。干旱胁迫打破了植物细胞中活性氧产生与清除之间的平衡, 导致活性氧积累, 引起膜脂过氧化作用^[28]。POD、SOD是植物体内清除活性氧的关键酶, 可以减轻或避免活性氧对细胞的伤害。本研究中, 转基因及野生各株系在干旱胁迫下POD、SOD活性均表现出先升高后下降的趋势, 胁迫至8 d, POD、SOD活性达到最高, 表明8 d后抗氧化酶防止膜脂过氧化和保护膜系统的能力降低^[29]。转基因各株系POD、SOD的活性在胁迫8-20 d内显著高于野生植株, 表明转基因植株中清除活性氧的能力较强。MDA含量及相对电导率表明逆境对植株的伤害程度, 有研究表明^{[30, 31]}MDA的含量、相对电导率的大小与植物抗旱系数呈线性负相关, 本研究中, 转基因株系的MDA含量与外渗电导率显著低于野生植株(P< 0.05)的, 表明转基因植株细胞膜遭受的破坏较小。综上所述, CBF2基因在提高拟南芥成苗抗旱能力上发挥了作用, 与李新玲等^[31]对CBF2基因研究结果类似。

此外, 转基因拟南芥不同株系之间抗旱性也存在差异, T₃-3株系的抗旱性较其它株系较弱。如干旱胁迫下, T₃-3的根长增长量明显低于转基因株系T₃-6和T₃-9的, 原因可能是转基因株系T₃-6和T₃-9相对于株系T₃-3基因表达性状更稳定。

对已转入二穗短柄草CBF2基因的拟南芥种子、幼苗及成苗进行干旱胁迫处理, 结果表明, 转基因拟南芥的抗旱性在幼苗及成苗期得到了提高, 即CBF2具有提高植株抗旱性的功能, 为研究CBF2基因其它功能奠定了基础, 也为利用转基因综合改良观赏植物的抗逆性提供了新思路。

The authors have declared that no competing interests exist.

参考文献

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2014

0.0

张强, 韩兰英, 张立阳, 王劲松. 论气候变暖背景下干旱和干旱灾害风险特征与管理策略. 地球科学进展, 2014, 29(1): 80-90.

Zhang

, Han L

, Zhang L

, Wang J

Analysis on the character and management strategy of drought disaster and risk under the climatic warming.

Advances in Earth Science, 2014, 29(1): 80-90. (in Chinese)

The drought is a most severe natural disaster worldwide,which leads to great risk in human being. The drought disaster and risk have more prominent because of obvious climatic warming in the last hundred years. At present,the understanding of the internal laws of the occurrence of drought and drought risk is not comprehensive,and the recognition of the characteristics of the drought and drought risk under climatic warming is obscure. In this paper,we summarized systematically the domestic and overseas research progress of the drought and drought disaster risk,introduced the principle of the drought disaster transfer process and the essential features of drought disaster,analyzed synthetically the main characteristics and interactions among the key factors of the drought disaster risk, discussed the effect of climatic warming on drought and drought disaster risk,and probed into the basic requirement of drought disaster risk management. Above all,we provide the main protective measurements of the drought disaster and the main strategy of drought disaster risk management.

干旱是全球影响最广泛的自然灾害,给人类带来了巨大的危害,近百年气候显著变暖使干旱灾害及其风险问题更加突出。目前,对干旱和干旱灾害风险的内在规律理解并不全面,对气候变暖背景下干旱和干旱灾害风险的表现特征认识也比较模糊。在系统总结国内外已有干旱和干旱灾害风险研究成果的基础上,归纳了干旱灾害传递过程的基本规律及干旱灾害的本质特征,综合分析了干旱灾害风险关键要素的主要特点及其相互作用关系,讨论了气候变暖对干旱和干旱灾害风险的影响特点,探讨了干旱灾害风险管理的基本要求。在此基础上,提出了干旱灾害防御的主要措施及干旱灾害风险管理的重点策略。

... 干旱是影响植物生长的重要环境因子之一,也是对全球影响最广泛的自然灾害^[1] ...

1944

0.0

... 我国是一个干旱灾害频发的国家,据统计每年我国干旱灾害造成的损失超过各种自然灾害损失总和的15%^[2,3] ...

2003

0.0

李茂松, 李森, 李育慧. 中国近50年来旱灾灾情分析. 中国农业气象, 2003, 24(1): 7-10.

Li M

, Li

, Li Y

Studies on drought in the past 50 years in China.

Agricultural Meteorology, 2003, 24(1): 7-10. (in Chinese)

20世纪后50年全国受旱面积和受旱成灾面积呈上升趋势,各年代旱灾成灾率也呈上升趋势;2000年是建国以来旱灾最严重的一年,全国受旱面积和成灾面积分别达4054万hm2和2678万hm2.黄淮海地区和长江中下游地区受旱面积和成灾面积占全国的60%以上,是中国的干旱重灾区.总结了有关干旱发展预测的研究,对未来一段时期旱灾的可能发生情况进行了预测,并提出了有关旱灾防御的几点建议.

... 我国是一个干旱灾害频发的国家,据统计每年我国干旱灾害造成的损失超过各种自然灾害损失总和的15%^[2,3] ...

2009

0.0

... 相比功能基因,转录因子在一定条件下可以调控多个逆境相关基因的表达,因此,利用转录因子来提高植株的抗逆性将是一条更有效的途径^[4] ...

2005

0.0

... AP2、bZIP、NAC、MYB、Cys2/His2锌指等不同类型的转录因子能够调控植物应答干旱胁迫,这些转录因子通过激活或抑制下游干旱应答基因的表达来调节植物的抗旱性,且能获得持久的抗逆性,具有广阔的应用前景^[5,6,7,8,9] ...

2005

0.0

2006

0.0

2005

0.0

2014

0.0

徐立明, 张振葆, 梁晓玲, 卢文, 张辰路, 黄凤珠, 王雷, 张素芝. 植物抗旱基因工程研究进展. 草业学报, 2014, 23(6): 293-303.

Xu L

, Zhang Z

, Liang X

, Lu

, Zhang C

, Huang F

, Wang

, Zhang S

Advances in genetic engineering for drought tolerance in plants.

Acta Prataculturae Sinica, 2014, 23(6): 293-303. (in Chinese)

Drought is a major abiotic stress that significantly reduces crop yield. The use of high throughput screening techniques has accelerated the identification of the gene series that are involved in drought tolerance. Some of the more important genes have already been transferred into plants by genetic transformation in order to activate specific or broad pathways related to drought tolerance. More recent advances have improved our understanding of drought-resistant regulation by controlling the expression pattern of drought-tolerant genes. We now better understand the complex regulatory network that regulates drought stress response and so are able to develop practical approaches for engineering drought tolerance in plants. This article reviews these advances in genetic engineering, including such factors as identifying the genes encoding signaling molecules, transcription factors, small RNA and the gene expressions involved in osmotic substances biosynthesis, polyamines biosynthesis and ROS scavenging. Current problems and future applications are also discussed.

1.Key Laboratory of Biology and Genetic Improvement of Maize in Southwest Region, Ministry of Agriculture, Maize Research Institute, Sichuan Agricultural University, Chengdu 611130,China; 2.Agricultural Bureau of Wulian District, Rizhao 262300, China

干旱是影响农作物产量的主要胁迫因素之一。高通量生物技术的使用促成了新的干旱胁迫相关基因的发现,一些重要基因已转化到植物中,通过专一性或广谱性地应答路径来调节其干旱耐受性。最近的一些研究进展,进一步加深了对植物通过调控干旱相关基因的表达而抵御干旱胁迫的理解,并对植物干旱胁迫的复杂调控网络有了更深刻的认识,同时逐渐探索出一些具体的植物抗旱基因工程研究的途径。本文主要综述了信号分子、转录因子、小RNA分子、渗透调节分子、多胺类分子,活性氧清除分子方面的基因工程研究进展,并对其研究中存在的问题及应用前景进行了讨论。

2002

0.0

... CBF基因家族目前发现编码6个相关的转录活化因子,分别称为CBF1、CBF2、CBF3、CBF4、CBF5和CBF6^[10,11] ...

... 当某些植物受到干旱、盐碱、冷胁迫时DREB/CBF上调表达,可激活其它一系列依赖DRE/CRT顺式作用元件的抗逆功能基因的表达,从而增强植物对干旱、低温及高盐碱等逆境的抗性,例如在拟南芥(Arabidopsis thaliana)中超表达的CBF4激活了包含低温驯化及干旱适应的下游基因C-重复/脱水应答元件,从而提高了植株的抗寒性、抗旱性^[10,12] ...

2002

0.0

... CBF基因家族目前发现编码6个相关的转录活化因子,分别称为CBF1、CBF2、CBF3、CBF4、CBF5和CBF6^[10,11] ...

2008

0.0

... 此外,受ABA的诱导,CBF2的表达量也会增加,原因是植物对ABA的响应促进了CBF启动子活性的提高^[12,17,18] ...

1998

0.0

... 目前,CBF家族转录因子抗寒功能及应用的研究比较广泛,研究表明,DREB/CBF基因的表达可以被多种胁迫诱导,CBF1及CBF3多在盐胁迫下表达,而CBF2主要在干旱及高盐胁迫下表达^[13,14] ...

2015

0.0

2012

0.0

... 通过对干旱胁迫时CBF 4种基因在不同葡萄(Vitis vinifera)品种中表达模式的研究发现,干旱条件下CBF2在叶片及根系中均表达,且相比衰老组织,幼嫩组织中的表达量较多^[15,16] ...

2002

0.0

2009

0.0

刘艳香, 董宽虎. 转录因子CBF及其抗寒作用机制. 草业科学, 2009, 26(5): 86-94.

Liu Y

, Dong K

Transcription factor CBF and its cold tolerance mechanism.

Pratacultural Science, 2009, 26(5): 86-94.

This paper summarized the discovery, classification, molecular mechanism in the process of coldtolerance and some associated molecular information regulating CBF and regulated by CBF, as well as the latest progress of CBF research. Low temperature is one of the most important environmental factors influencing plant growth and production performance. With the development of molecular biology, the molecular mechanism that plants survive in stress environmental conditions has become a hot field in the study of botany nowadays. Transcription factor is the accessory factor that RNA polymerase needs in transcription initiation. Eukaryotic RNA polymerase can not start its transcription unless the transcription factor (protein) is binding at its recognition sequence on DNA. Transcription factor CBF is an important regulator that exists extensively in the process of cold acclimation of all plants and it recognizes CRT／DRE（Crepeat/dehydrationresponsive element）in COR and starts its transcription.

从植物的抗寒方面综述了转录因子CBF（CRT／DRE—binding factor）的发现、分类、结构、在植物抗寒过程中发挥作用的分子机制，并介绍了调控CBF和受CBF调控的其他分子的信息，较为系统地归纳了CBF研究的最新进展。低温是影响植物生长、生产性能的重要因素之一。随着分子生物学的发展，植物抗逆分子机制成为当前植物学研究的一个热点。转录因子是转录起始过程中RNA聚合酶所需的辅助因子。真核生物RNA聚合酶自身无法启动基因转录，只有当转录因子结合在其识别的DNA序列上后，基因才开始转录。转录因子CBF广泛存在于各种植物中，可以识别COR基因中的CRT／DRE（Crepeat/dehydrationresponsive element）元件，从而启动COR基因转录，是植物抗逆过程中一个重要的调节因子。

... 此外,受ABA的诱导,CBF2的表达量也会增加,原因是植物对ABA的响应促进了CBF启动子活性的提高^[12,17,18] ...

2004

0.0

... 此外,受ABA的诱导,CBF2的表达量也会增加,原因是植物对ABA的响应促进了CBF启动子活性的提高^[12,17,18] ...

... CBF2在CBF1和CBF3表达上具有负的调控作用,并且对拟南芥的耐胁迫性起着重要作用^[18] ...

2002

0.0

... 二穗短柄草(Brachypodium distachyon)是禾本科短穗草属一年生野生草本植物,具有极强的抗寒性,对干旱、盐碱也有一定的抗性^[19] ...

2012

0.0

... 目前已证明,二穗短柄草CBF3转录因子在提高植物的抗寒性方面具有重要作用^[20],但对二穗短柄草CBF2转录因子抗性的研究较少 ...

2000

0.0

... 采用氮蓝四唑(NBT)光化还原法测定超氧化物歧化酶(SOD)总活性,采用愈创木酚法测定过氧化物酶(POD)总活性,采用硫代巴比妥酸法测定丙二醛(MDA)含量,采用电导仪法测定电导率^[21] ...

2010

0.0

秦文静, 梁宗锁. 四种豆科牧草萌发期对干旱胁迫的响应及抗旱性评价. 草业学报, 2010, 19(4): 61-70.

Qin W

, Liang Z

Response and drought resistance of four leguminous pastures to drought during seed germination.

Acta Prataculturae Sinica, 2010, 19(4): 61-70. (in Chinese)

对陕北黄土高原主要的４种豆科牧草毛苕子、沙打旺、白花草木樨、尖叶胡枝子种子生物学特性及干旱胁迫条件下种子萌发特性进行了研究。结果表明，１）４种植物种子生物学特性各异，沙打旺和毛苕子种子含水量、累计吸水率均大于其他２种，尖叶胡枝子和毛苕子的种子活力大于沙打旺和白花草木樨；２）随着干旱胁迫程度的加剧，４种植物种子总萌发率、幼苗鲜重、抗旱指数及胚根和胚芽的生长均呈下降趋势，且白花草木樨下降幅度最大；３）低浓度的ＰＥＧ６０００对尖叶胡枝子的种子萌发、沙打旺的胚芽生长以及它们的抗旱指数均具有一定的促进作用；４）采用隶属函数法对干旱胁迫下４种植物萌发期抗旱性进行了综合评价，结果显示，４种植物种子萌发期的抗旱性次序为：毛苕子＞尖叶胡枝子＞沙打旺＞白花草木樨。

... 3 讨论与结论在干旱环境下,种子能否保持活力及幼苗能否继续生长是植物存活的关键,目前对转基因植株种子萌发期的抗性研究较少,而种子萌发期抗旱性鉴定是整体抗旱性评价的重要内容^[22,23,24] ...

2013

0.0

2014

0.0

张彦妮, 李博, 何淼. PEG干旱胁迫对大花飞燕草幼苗生理特性的影响. 草业科学, 2014, 31(3): 446-450.

Zhang Y

, Li

, He

Physiological characteristics of Delphinium grand iflorum under drought stress.

Pratacultural Science, 2014, 31(3): 446-450. (in Chinese)

The experiment was conducted to estimate the drought tolerance of Delphinium grandiflorum seedling stressed by PEG with the concentrations of 0, 5%, 10%, 15%, 20% and 25%, respectively. Different physiology indices including cell membrane permeability, malondialdehyde (MDA), free proline (Pro), peroxidase (POD) activity, superoxide dismutase (SOD) activity and soluble protein were measured. The results showed that the cell membrane permeability, MDA and Pro concentration in seedling leaves of D.grandiflorum increased with drought stress increased. SOD and POD activity increased firstly and then decreased with drought stress increased. Soluble protein concentration had no difference between control and different stress treatments. All of these variation suggested that the seedling of D.grandiflorum had drought tolerance in some degree.

College of Landscape Architecture, Northeast Forestry University, Harbin 150040, China

为研究大花飞燕草( Delphinium grandiflorum )幼苗抗旱性, 采用不同浓度(0、5%、10%、15%、20%和25%)的PEG溶液对大花飞燕草幼苗进行模拟干旱胁迫, 测定了其细胞膜透性、丙二醛、脯氨酸、过氧化物酶、超氧化物歧化酶以及可溶性蛋白等生理生化指标。结果表明, 随着干旱胁迫的加剧, 大花飞燕草幼苗叶片细胞膜透性、丙二醛含量和脯氨酸总体呈上升趋势;超氧化物歧化酶和过氧化物酶活性总体呈先升后降的趋势, 可溶性蛋白在各个胁迫条件下差异不大;综合各指标表明大花飞燕草幼苗具有一定的耐旱性。

2010

0.0

万里强, 李向林, 石永红, 何峰, 贾亚雄. PEG胁迫下4个黑麦草品种生理生化指标响应与比较研究. 草业学报, 2010, 19(1): 83-88.

Wan L

, Li X

, Shi Y

, He

, Jia Y

A study on the response and on the comparison of physiological and biochemical indexes of four Lolium perenne varieties under PEG stress.

Acta Prataculturae Sinica, 2010, 19(1): 83-88. (in Chinese)

利用不同浓度ＰＥＧ-６０００溶液模拟干旱胁迫，比较研究４个多年生黑麦草品种叶片生理生化指标响应。结果表明，不同ＰＥＧ胁迫处理下，品种“欧必克”叶片相对含水量均最高且降幅最小，叶片电导率均最低，ＭＤＡ含量增幅较小，叶片细胞膜系统有较强的忍耐干旱的能力；而品种“凤凰”忍耐干旱的性能相对较弱。胁迫下各品种叶片游离脯氨酸含量显著增加，随渗透胁迫加剧，品种“欧必克”脯氨酸含量急剧上升。低浓度胁迫下（５％ＰＥＧ），各品种叶片ＳＯＤ活性均提高，胁迫加剧时ＳＯＤ活性则下降。ＰＥＧ渗透胁迫导致４个多年生黑麦草品种叶片相对含水量下降、细胞膜透性增大、ＭＤＡ含量增加、脯氨酸含量上升而ＳＯＤ活性下降，但４个品种对应指标的变化幅度存在差异，品种“欧必克”的抗旱性最强，而“凤凰”的抗旱性最弱。

... 渗透调节是植物适应水分胁迫的主要生理机制,通过渗透调节植物在干旱条件下维持一定的膨压,从而维持细胞生长、气孔开放和光合作用等生理过程的进行^[25] ...

2013

0.0

刘颖, 张巨明. 暖季型草坪草对干旱胁迫的反应. 草业科学, 2013, 30(11): 1732-1738.

Liu

, Zhang J

Review on the response of warm-season turfgrass to drought stress.

Pratacultural Science, 2013, 30(11): 1732-1738. (in Chinese)

Drought is one of the main environmental stress factors that influence the growth of warm-season turfgrass. Resent researches on morphological, physiological, biochemical and genetic response of stem, leaf and root of warm-season turfgrass to drought stress were reviewed in this study. Besides, a research direction of this field that, future studies should focus on genetic response of warm-season turfgrass to drought stress, then morphological, physiological and biochemical resposes, was came up.

College of Agriculture, South China Agricultural University, Guangzhou 510642, China

干旱胁迫是影响草坪草生长的主要环境胁迫因子之一。本文对国内外关于暖季型草坪草茎、叶及根组织器官对干旱胁迫的形态反应、生理生化反应和基因响应等方面的研究进行综述,指出了日后对干旱胁迫反应机制的研究以基因水平为主,形态水平、生理生化水平为辅的研究发展方向。

... 干旱胁迫下,幼苗根的生长对植物幼苗生长发育具有十分重要的意义^[26] ...

2013

0.0

李长慧, 李淑娟, 雷有升, 胡凯, 王佩羽. 4种高原乡土禾草的抗旱生理比较. 草业科学, 2013, 30(9): 1386-1393.

Li C

, Li S

, Lei Y

, Hu

, Wang P

Study on drought resistance of four native grass species.

Pratacultural Science, 2013, 30(9): 1386-1393. (in Chinese)

Drought resistance of four native grass species, including Roegneria thoroldiana, Poa pratensis, Elymus nutans and Festuca sinensis, in seedling stage was studied by pot culturing experiment. The results indicated that, the content of free proline(Fpro), soluble protein and soluble sugar, and the activity of catalase(CAT) and peroxidase(POD) increased with the increasing of drought stress. Beside, with the increasing of drought stress, decompose of chlorophyll, membrane permeability and electrolyte leakage rate were also increased, and the cellular tissues of grasses were damaged. Different grass spices had different changes on physiological and biochemical indexes under drought stress. The results of subordinate function analysis indicated that, the drought resistance ability of four grass species were R.thoroldiana>P.pratensis>E.nutans>F.sinensis.

采用盆栽法对梭罗草（Roegneria thoroldiana）、草地早熟禾（Poa pratensis）、垂穗披碱草（Elymus nutans）和中华羊茅（Festuca sinensis）4种高原乡土禾本科牧草幼苗的抗旱性进行了比较。结果表明，一定程度的干旱胁迫后，4种禾草的游离脯氨酸含量、可溶性蛋白含量、可溶性糖含量、过氧化氢酶活性、过氧化物酶活性等均有所增加，且叶绿素分解、细胞膜透性增大、电解质外渗率增加，牧草组织细胞受到伤害。不同牧草在干旱胁迫下生理生化指标的变化不同，综合各项指标，运用隶属函数分析表明，4种禾草抗旱性强弱的排序为梭罗草>草地早熟禾>垂穗披碱草>中华羊茅。

... 胁迫条件下,细胞及酶活性变化以及渗透调节物质累积情况需要依靠植物的生理生化指标进行研究^[27] ...

2014

0.0

林武星, 谷凌, 朱炜, 聂森. 干旱胁迫对台湾海桐生长和生理生化特性的影响. 草业科学, 2014, 31(10): 1915-1922.

Lin W

, Gu

, Zhu

, Nie

Effects of drought stress on growth and physiological and biochemical characteristics of Pittosporum pentand rum.

Pratacultural Science, 2014, 31(10): 1915-1922. (in Chinese)

In order to explore the mechanism of drought-tolerance of Pittosporum pentandrum , the growth and physiological and biochemical characteristics of 1-year-old P. pentandrum seedlings were analysis under four levels of drought stress including optimal water content(CK), mild drought stress(LD), moderate drought stress(MD)and heavy drought stress(HD) which were controlled by weighing the soil and pots. With the increase of drought stress, the heights and biomass of the seedlings decreased but the root/shoot ratio increased; net photosynthesis rate (P n ), transpiration rate (T r ), stomatal conductance(G s ), stomatal limitation(L s )and water use efficiency(WUE)decreased while intercellular CO 2 concentration (C i )increased; the contents of K + and Mg 2+ decreased while the content of Ca 2+ firstly increased and then decreased; the contents of soluble sugar and soluble protein increased while the content of proline firstly increased and then decreased; the contents of malondialdehyde and membrane permeability increased but the activity of catalase (CAT), peroxidase (POD), superoxide dismutase (SOD) and ascorbate peroxidase enzyme (APX) firstly increased and then decreased. These results indicated that the membrane lipid peroxidation increased and the growth was inhibited under drought stress. However, P. pentandrum protected the membrane structure and maintained normally physiological activity through improving the activities of the antioxidant enzymes system and accumulating osmotic regulation such as soluble sugar, proline ,soluble protein and Ca 2+ which improve its tolerance to the drought stress.

1.Fujian Academy of Forestry, Fuzhou 350012, China; 2.College of Forestry, Fujian Agriculture and Forestry University, Fuzhou 350003, China

以一年龄台湾海桐( Pittosporum pentandrum )盆栽苗为试材,采用人为称重控水方法,研究了正常供水(CK)、轻度干旱胁迫(LD)、中度干旱胁迫(MD)和重度干旱胁迫(HD)4种水分处理对台湾海桐生长和生理生化特性的影响,以探讨台湾海桐耐旱机制。结果显示,随干旱胁迫程度的加剧,台湾海桐幼苗高度和生物量下降,根冠比增加;净光合速率(P n )、蒸腾速率(T r )、气孔导度(G s )、气孔限制值(L s )和水分利用效率(WUE)降低,胞间CO 2 浓度(C i )增加;叶中K + 和Mg 2+ 含量降低,Ca 2+ 含量先增加后降低;叶中可溶性糖和可溶性蛋白质含量增加,叶中脯氨酸含量先增加后降低;叶中丙二醛(MDA)含量和质膜透性增加,过氧化氢酶(CAT)、过氧化物酶(POD)、超氧化物歧化酶(SOD)和抗坏血酸过氧化物酶(APX)活性先增加后降低。干旱胁迫下台湾海桐幼苗生长受到抑制和膜脂过氧化加剧,但其能通过对渗透调节物质的含量和抗氧化酶系统的活性调节来保护膜结构,保持正常的生理活动,表现出一定的抗旱能力。

... 干旱胁迫打破了植物细胞中活性氧产生与清除之间的平衡,导致活性氧积累,引起膜脂过氧化作用^[28] ...

2012

0.0

... 本研究中,转基因及野生各株系在干旱胁迫下POD、SOD活性均表现出先升高后下降的趋势,胁迫至8 d,POD、SOD活性达到最高,表明8 d后抗氧化酶防止膜脂过氧化和保护膜系统的能力降低^[29] ...

2009

0.0

张永峰, 殷波. 混合盐碱胁迫对苗期紫花苜蓿抗氧化酶活性及丙二醛含量的影响. 草业学报, 2009, 18(1): 46-50.

Zhang Y

, Yin

Influences of salt and alkali mixed stresses on anti oxidative activity and MDA content of Medicago sativa at seedling stage.

Acta Prataculturae Sinica, 2009, 18(1): 46-50. (in Chinese)

用不同浓度混合盐碱（ＮａＣｌ／Ｎａ２ＣＯ３）对苗期紫花苜蓿进行胁迫处理，测定其对抗氧化酶系统和丙二醛含量的影响。结果表明，随着混合盐碱处理浓度的增大，经处理的苗期紫花苜蓿叶片的超氧化物歧化酶（ＳＯＤ）、过氧化物酶（ＰＯＤ）、过氧化氢酶（ＣＡＴ）活性及丙二醛（ＭＤＡ）的含量呈增加趋势，并且均高于对照。

... MDA含量及相对电导率表明逆境对植株的伤害程度,有研究表明^[30,31]MDA的含量、相对电导率的大小与植物抗旱系数呈线性负相关,本研究中,转基因株系的MDA含量与外渗电导率显著低于野生植株(P#cod#x0003C ...

2010

0.0

李新玲, 徐香玲, 王全伟. 拟南芥CBF2基因的抗旱性研究. 生物技术通报, 2010(10): 96-103.

Li X

, Xu X

, Wang Q

Study on desiccation tolerance of CBF2 gene in Arabidopsis thaliana. Biotechnology Bulletin, 2010(10): 96-103. (in Chinese)

以拟南芥为材料,根据已报道的序列设计引物克隆了CBF2基因及rd29A基因启动子,并构建了植物表达载体pBI-rd-cbf,用以转化烟草。转基因烟草的Northern杂交检测显示,30%PEG诱导条件下CBF2基因在诱导30 min之后持续较高的表达水平,且表达水平受胁迫诱导程度的影响。抗旱生理指标测定显示,干旱条件下转基因烟草植株的脯氨酸含量迅速增高,远超于野生型;而丙二醛含量的增长幅度要比野生型植株小很多,且随着胁迫时间的延长,转基因植株体内的丙二醛含量逐步趋于稳定。试验证明,CBF2基因在提高农作物的抗旱性方面具有极大的利用价值。

... 综上所述,CBF2基因在提高拟南芥成苗抗旱能力上发挥了作用,与李新玲等^[31]对CBF2基因研究结果类似 ...