转菠菜BADH小麦抗性生理与不同抗旱性小麦品种中BADH的表达研究
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摘要
在干旱、低温等胁迫环境中，植物细胞内通常会积累一些相容性物质（也叫渗调物质），从而减轻环境胁迫对植物造成的伤害，甜菜碱是主要的相容性物质之一。编码甜菜碱醛脱氢酶（BADH，催化甜菜碱醛合成甜菜碱）的基因在甜菜碱的生物合成过程中处于关键地位。本研究首先对抗旱性强的小麦品系旱丰9703与抗旱性弱的品种山农215953干旱胁迫下的渗透调节能力和光合作用进行了比较，分析了不同生育期叶片中BADH基因表达特性、BADH活性与甜菜碱含量，从生理与分子机制两方面对甜菜碱的生理功能进行了初步探讨。其次，研究测定了转菠菜BADH基因小麦低温及干旱胁迫下，不同生育期BADH活性、甜菜碱含量与光合作用等生理参数的变化，进一步研究了小麦胁迫耐性与甜菜碱含量、BADH活性之间的关系。考虑到菠菜与小麦BADH基因的保守性，我们从抗旱小麦新品系旱丰9703叶片中克隆了BADH cDNA序列，并构建了表达载体，对模式植物烟草进行了转化，旨在进一步研究小麦BADH基因在抗逆性中的作用，为其今后在小麦育种中的应用打下基础。主要结论如下：
1. 长期大田干旱胁迫下，旱丰9703表现出了较强抗性；三个生育期中旱丰9703的叶片相对含水量与渗透调节能力均高于山农215953，灌浆期的渗透调节能力最强，比乳熟期高0.284MPa；干旱胁迫下旱丰9703仍具有较高的净光合速率（Pn）、最大光化学效率（Fv/Fm）和实际光化学效率（φPSII），耐光抑制能力较强。Northern印迹杂交结果表明：灌浆期中度水分胁迫时BADH的表达水平显著增加，旱丰9703略高于山农215953；乳熟期较严重的土壤水分胁迫下抗旱性强的品种叶片中BADH表达量明显高于抗旱性弱的品种；三个生育期中，BADH活性和甜菜碱含量的变化与BADH基因的表达呈现相同的趋势，均是先增加后降低，旱丰9703的BADH活性与甜菜碱含量均高于山农215953，以灌浆期最为明显。
干旱胁迫下旱丰9703叶片中较高的BADH表达使其甜菜碱含量与渗透调节能力一直高于山农215953，其叶片的保水能力较强，维持了较高的净光合速率和PSII光化学活性；叶片中甜菜碱的积累量与小麦抗旱性成正相关，叶片中BADH活性与甜菜碱含量水平可以作为小麦抗性强弱的主要参考指标。
2. 研究发现，转菠菜BADH基因小麦植株在耐低温、干旱胁迫和抗光氧化方面的能力
较强，叶片中较高的甜菜碱含量可能对它们胁迫耐性的提高起着重要作用，结合农艺性状的改良与选择，株系99T4和99T6可以作为较好的抗逆种质资源在小麦抗性育种过程中应用。低温胁迫下转基因株系99T4和99T6的渗透调节能力、抗氧化酶活性、D1蛋白周转能力均高于4185，99T1、99T4和99T6叶片的净光合速率分别是对照植株的0.96、2.11和1.56倍；-7℃低温胁迫24h以后，株系99T4与99T6没有受到明显的低温伤害，表现出较强的低温胁迫耐性。
灌浆期不同程度的干旱胁迫表明，99T4与99T6耐干旱胁迫的能力要高于99T1与4185，能够保持相对较高的渗透调节能力与叶绿素含量；较严重干旱胁迫下， 99T4与99T6的Pn（约为7.5 μmolCO2·m-2·s-1）要高于4185（约为4.9 μmol CO2·m-2·s-1）；株系99T4与99T6在光抑制环境下PSII反应中心受到的损伤较小，具有较高的抗光氧化能力，99T4与99T6依赖叶黄素循环的热耗散在耗散过剩光能的过程中起到更大作用；
3. 通过5，-和3，-RACE，从抗旱性小麦旱丰9703叶片中克隆了BADH cDNA全序列，命名为wBADH，在Genebank中注册（注册号：AY 050316）。小麦BADH cDNA开放阅读框架含有1509bp，编码503个AA，将氨基酸序列与Genebank中已登录的几种植物的BADH氨基酸序列比较，与大麦、水稻的相应片断的同源性分别为98％、73％。进一步构建了小麦BADH基因的表达载体，利用农杆菌介导法侵染烟草，通过PCR和PCR-Southern鉴定证明我们已得到转基因烟草植株；低温处理后转基因植株中BADH的活性较低，最高的只有4.8 nmol·mg protein-1·min-1；这些研究为该基因在小麦抗性育种中的应用打下了基础。
Under environmental stress, plant cells usually accumulates some compatible solutes(also regarded as osmotin) in order to alleviate the injury derived from environmental stress. Glycine betaine (GB) is regarded as an extremely effective compatible solute, therefore researchers pay much attention to the bio-engineering of GB. The BADH encoding BADH which catalyzes betaine aldehyde (BA) into GB stands the essential position in the synthesis of GB. Firstly, we determined the OA and photosynthesis of flag leaves in strong and weak drought resistant wheat varieties, and then analyzed the expression level of BADH, BADH activity and GB content of flag leaves in two wheat varieties at different stage under drought stress and dissertated the physiological function of GB in molecular biology. Secondly, some physiological parameters such as the BADH activity, GB content and photosynthesis of transformed wheat plants with BADH from spinach under drought stress were studied; Finally, we cloned complete sequence of the BADH cDNA from the leaves of strong drought resistant cultivar Hanfeng9703, constructed the expression vector and acquired transgenic tobacco plants. The results are followed:
1. Two winter wheat(Triticum aestivum L.)varieties( Drought-resistant cultivar Hanfeng 9703 and Drought-sensitive cultivar Shannong 215953) were used to study the OA, Pn and photochemical activity of PSII complex as well as the expression level of BADH and GB content in flag leaves at different stage under long-term field drought stress. OA and RWC of Hanfeng 9703 were higher than those of Shannong 215953 at three stages, OA at
seed filling stage was the highest which increased 0.058and 0.284MPa comparing to that of flowering stage and milk ripening stage respectively. Drought stress resulted in the decrease of Pn, Fv/Fm and φPSII of two wheat varieties at three different stage; But Pn, Fv/Fm, φPSII of Hanfeng 9703 were higher than those of Shannong 215953 under drought stress. Drought stress aggravated the depression of Pn in midday in leaves of two wheat varieties at milk ripening stage, PSII photochemical activity of leaves of Hanfeng 9703 were higher than those of Shannong 215953; Higher osmotic adjustment of Hanfeng 9703 maintained Pn and PSII photochemical activity which were beneficial for the higher production, it was concluded that the higher GB content in Hanfeng9703 might play pivotal role in enhancement of stress resistance.
Northern blot results showed that continuous drought stress leads to the increase of the BADH expression at seed filling stage in two varieties, severe drought stress results in the decline of expression level at milk ripening stage, the BADH expression level of flag leaves in Hanfeng 9703 were obviously higher than that in Shannong 215953. The changes of BADH activity and GB content was similar to the change of BADH expression in two varieties. After moderate drought stress at seed filling stage, the higher BADH expression of leaves in Hanfeng 9703 leaded to the higher GB content in comparason to Shannong 215953 which was one of the major reasons of keeping stronger PSII photochemical efficiency in Hanfeng 9703 under drought stress. It can be concluded that the GB content and BADH activity in flag leaves could be used as the reference criterion for resistance appraisal of crop germplasm.
The tolerance to the low temperature, drought and anti-photoxidation stress in transgenic wheat plants (line-99T1,99T4,99T6) transformed with BADH from Atriplex hortensis were higher comparing to those of wild-type plants(4185); The stronger OA, higher anti-oxidation enzymes activity and faster turnover of D1 protein were main protective mechanism in 99T4 and 99T6, Furthermore, the higher GB content in leaves may play important role in the enhancement of stress tolerance.. The leaf electrolyte leakage in 99T4 and 99T6 were lower than those in 4185 and 99T1.Under low temperature conditions, compared to the wild-type plants, the osmotic adjustment in 99T4 and 99T6 were 17.3% and 8.2% h
1. 长期大田干旱胁迫下，旱丰9703表现出了较强抗性；三个生育期中旱丰9703的叶片相对含水量与渗透调节能力均高于山农215953，灌浆期的渗透调节能力最强，比乳熟期高0.284MPa；干旱胁迫下旱丰9703仍具有较高的净光合速率（Pn）、最大光化学效率（Fv/Fm）和实际光化学效率（φPSII），耐光抑制能力较强。Northern印迹杂交结果表明：灌浆期中度水分胁迫时BADH的表达水平显著增加，旱丰9703略高于山农215953；乳熟期较严重的土壤水分胁迫下抗旱性强的品种叶片中BADH表达量明显高于抗旱性弱的品种；三个生育期中，BADH活性和甜菜碱含量的变化与BADH基因的表达呈现相同的趋势，均是先增加后降低，旱丰9703的BADH活性与甜菜碱含量均高于山农215953，以灌浆期最为明显。
干旱胁迫下旱丰9703叶片中较高的BADH表达使其甜菜碱含量与渗透调节能力一直高于山农215953，其叶片的保水能力较强，维持了较高的净光合速率和PSII光化学活性；叶片中甜菜碱的积累量与小麦抗旱性成正相关，叶片中BADH活性与甜菜碱含量水平可以作为小麦抗性强弱的主要参考指标。
2. 研究发现，转菠菜BADH基因小麦植株在耐低温、干旱胁迫和抗光氧化方面的能力
较强，叶片中较高的甜菜碱含量可能对它们胁迫耐性的提高起着重要作用，结合农艺性状的改良与选择，株系99T4和99T6可以作为较好的抗逆种质资源在小麦抗性育种过程中应用。低温胁迫下转基因株系99T4和99T6的渗透调节能力、抗氧化酶活性、D1蛋白周转能力均高于4185，99T1、99T4和99T6叶片的净光合速率分别是对照植株的0.96、2.11和1.56倍；-7℃低温胁迫24h以后，株系99T4与99T6没有受到明显的低温伤害，表现出较强的低温胁迫耐性。
灌浆期不同程度的干旱胁迫表明，99T4与99T6耐干旱胁迫的能力要高于99T1与4185，能够保持相对较高的渗透调节能力与叶绿素含量；较严重干旱胁迫下， 99T4与99T6的Pn（约为7.5 μmolCO2·m-2·s-1）要高于4185（约为4.9 μmol CO2·m-2·s-1）；株系99T4与99T6在光抑制环境下PSII反应中心受到的损伤较小，具有较高的抗光氧化能力，99T4与99T6依赖叶黄素循环的热耗散在耗散过剩光能的过程中起到更大作用；
3. 通过5，-和3，-RACE，从抗旱性小麦旱丰9703叶片中克隆了BADH cDNA全序列，命名为wBADH，在Genebank中注册（注册号：AY 050316）。小麦BADH cDNA开放阅读框架含有1509bp，编码503个AA，将氨基酸序列与Genebank中已登录的几种植物的BADH氨基酸序列比较，与大麦、水稻的相应片断的同源性分别为98％、73％。进一步构建了小麦BADH基因的表达载体，利用农杆菌介导法侵染烟草，通过PCR和PCR-Southern鉴定证明我们已得到转基因烟草植株；低温处理后转基因植株中BADH的活性较低，最高的只有4.8 nmol·mg protein-1·min-1；这些研究为该基因在小麦抗性育种中的应用打下了基础。
Under environmental stress, plant cells usually accumulates some compatible solutes(also regarded as osmotin) in order to alleviate the injury derived from environmental stress. Glycine betaine (GB) is regarded as an extremely effective compatible solute, therefore researchers pay much attention to the bio-engineering of GB. The BADH encoding BADH which catalyzes betaine aldehyde (BA) into GB stands the essential position in the synthesis of GB. Firstly, we determined the OA and photosynthesis of flag leaves in strong and weak drought resistant wheat varieties, and then analyzed the expression level of BADH, BADH activity and GB content of flag leaves in two wheat varieties at different stage under drought stress and dissertated the physiological function of GB in molecular biology. Secondly, some physiological parameters such as the BADH activity, GB content and photosynthesis of transformed wheat plants with BADH from spinach under drought stress were studied; Finally, we cloned complete sequence of the BADH cDNA from the leaves of strong drought resistant cultivar Hanfeng9703, constructed the expression vector and acquired transgenic tobacco plants. The results are followed:
1. Two winter wheat(Triticum aestivum L.)varieties( Drought-resistant cultivar Hanfeng 9703 and Drought-sensitive cultivar Shannong 215953) were used to study the OA, Pn and photochemical activity of PSII complex as well as the expression level of BADH and GB content in flag leaves at different stage under long-term field drought stress. OA and RWC of Hanfeng 9703 were higher than those of Shannong 215953 at three stages, OA at
seed filling stage was the highest which increased 0.058and 0.284MPa comparing to that of flowering stage and milk ripening stage respectively. Drought stress resulted in the decrease of Pn, Fv/Fm and φPSII of two wheat varieties at three different stage; But Pn, Fv/Fm, φPSII of Hanfeng 9703 were higher than those of Shannong 215953 under drought stress. Drought stress aggravated the depression of Pn in midday in leaves of two wheat varieties at milk ripening stage, PSII photochemical activity of leaves of Hanfeng 9703 were higher than those of Shannong 215953; Higher osmotic adjustment of Hanfeng 9703 maintained Pn and PSII photochemical activity which were beneficial for the higher production, it was concluded that the higher GB content in Hanfeng9703 might play pivotal role in enhancement of stress resistance.
Northern blot results showed that continuous drought stress leads to the increase of the BADH expression at seed filling stage in two varieties, severe drought stress results in the decline of expression level at milk ripening stage, the BADH expression level of flag leaves in Hanfeng 9703 were obviously higher than that in Shannong 215953. The changes of BADH activity and GB content was similar to the change of BADH expression in two varieties. After moderate drought stress at seed filling stage, the higher BADH expression of leaves in Hanfeng 9703 leaded to the higher GB content in comparason to Shannong 215953 which was one of the major reasons of keeping stronger PSII photochemical efficiency in Hanfeng 9703 under drought stress. It can be concluded that the GB content and BADH activity in flag leaves could be used as the reference criterion for resistance appraisal of crop germplasm.
The tolerance to the low temperature, drought and anti-photoxidation stress in transgenic wheat plants (line-99T1,99T4,99T6) transformed with BADH from Atriplex hortensis were higher comparing to those of wild-type plants(4185); The stronger OA, higher anti-oxidation enzymes activity and faster turnover of D1 protein were main protective mechanism in 99T4 and 99T6, Furthermore, the higher GB content in leaves may play important role in the enhancement of stress tolerance.. The leaf electrolyte leakage in 99T4 and 99T6 were lower than those in 4185 and 99T1.Under low temperature conditions, compared to the wild-type plants, the osmotic adjustment in 99T4 and 99T6 were 17.3% and 8.2% h
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