低温下ABA调控冬小麦糖代谢及抗寒基因表达的研究
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摘要
东农冬麦1号是我国首能例能在黑龙江省（最低温可达-30℃）严寒条件下越冬的强抗寒冬小麦新品种，返青率大于85%;济麦22是在我国华北地区广泛种植的冬小麦品种，也具有较强的抗寒能力，但不能在黑龙江省越冬。本研究以这两种冬小麦为试验材料，在苗期低温下及越冬期探讨了外源ABA对糖代谢及抗寒基因表达的影响，以期了解外源ABA调控冬小麦糖代谢及抗寒基因表达的生理及分子基础，为揭示外源ABA提高冬小麦抗寒性提供新的研究思路。本研究所用材料东农冬麦1号为黑龙江省高寒地区唯一可以种植的冬小麦品种，具有典型性和代表性；国内外对冬小麦抗寒性的研究大多采用室内不同低温处理小麦幼苗，本研究采用室内模拟低温和大田严寒情况下相结合，更接近生产实际；本研究采用生理生化技术与分子生物学技术相结合的实验方法，首次系统全面地从糖代谢及抗寒基因表达角度研究ABA调控冬小麦抗寒性,从总体上把握糖类物质的来源和去路,阐明ABA对冷胁迫下植株能量流向、渗透水平,及在信号作用下能量代谢及抗寒基因调节过程的影响,为揭示冬小麦抗冻机理提供理论依据。最终为北方高寒地区栽培冬小麦的改良做出重要贡献。本文的主要研究
结果如下：
1．苗期低温下外源ABA对冬小麦糖代谢的影响
苗期低温胁迫时，外源ABA促进了抗寒性强的东农冬麦1号叶片中蔗糖、果糖的积累，从而提高了可溶性糖的含量，在地下茎中则在0℃以上时促进糖类的积累；而外源ABA没有促进济麦22的叶片和地下茎中糖类的积累；淀粉的积累在两品种小麦的叶片和地下茎中则受到抑制或与对照差异不显著。外源ABA对东农冬麦1号两种器官中TaRBC、TaFBP、TaPRK、TaUGP、TaSAInv、TaSPS、TaPK、TaG6PDH、TaTPT和TaSuT的表达有不同程度的促进作用，这些基因分别参与碳的固定、蔗糖的合成和分解、糖酵解、戊糖磷酸途径以及糖的运输，但在东农冬麦1号中，外源ABA处理后也有些基因如TaAGP、TaPD和TaHxK下调表达；在济麦22中，外源ABA处理后大部分糖代谢酶基因的表达都受到了抑制。
2．越冬期外源ABA对冬小麦糖代谢的影响
越冬期低温时，两品种冬小麦ABA处理组的叶片和分蘖节中都积累了更多的可溶性总糖、蔗糖和果糖，尤其是在越冬器官分蘖节中，在抗寒性强的东农冬麦1号中比抗寒性弱的济麦22中积累了更多的碳水化合物，外源ABA促进了东农冬麦1号两种器官和济麦22叶片中淀粉的积累，但对济麦22分蘖节中淀粉积累的影响却不明显。外源ABA不同程度地促进了东农冬麦1号和济麦22叶片和分蘖节中TaRBC、TaFBP、TaPRK、TaUGP、TaAGP、TaSAInv、TaSS、TaSPS、TaPK、TaH×K、TaPFK、TaPD、TaG6PDH和TaSuT基因的表达，但在东农冬麦1号中的促进程度大于济麦22，外源ABA显著地促进了TaPRK, TaUGP, TaSPS, TaSAInv, TaPD和TaPFK基因在两品种冬小麦叶片中的表达，而TaTPT仅在东农冬麦1号的叶片中上调表达。外源ABA促进蔗糖积累的同时也促进了蔗糖的分解，但积累大于分解，而蔗糖磷酸合成酶（SPS）与蔗糖合成酶（SS）在蔗糖积累方面起作用，尿苷二磷酸-葡萄糖焦磷酸化酶（UGP）对蔗糖的积累和分解同时起作用，酸性转化酶和碱性转化酶则在蔗糖的分解方向起作用；ABA处理不同程度地提高了两个品种小麦叶片及分蘖节中糖酵解代谢酶丙酮酸激酶、己糖激酶和磷酸果糖激酶的活性，尤其在最低温-25℃时明显抑制这几种酶活性的降低，从而减缓温度下降给冬小麦带来的伤害，但这种影响在抗寒性强的东农冬麦1号中表现得更明显，而在冷敏感的济麦22中相对不明显，丙酮酸激酶活性在济麦22分蘖节中甚至受到ABA的抑制。
3．糖含量与糖代谢关键酶基因表达以及酶活性间的相关分析
ABA处理组的糖含量与糖代谢酶基因表达之间的相关性更为普遍，外源ABA对糖含量与糖代谢酶基因的调控具有一定的一致性，但在苗期和越冬期，对照和ABA处理组中糖含量与糖代谢酶基因之间的相关性并不完全一致，表明糖类物质的积累与糖代谢酶基因的表达之间具有复杂的关系。越冬期通过对糖含量与糖代谢酶活性间的相关分析表明，ABA处理后也使糖含量与酶活性间的相关更为普遍，两小麦的叶片中糖含量与糖代谢酶活性的相关程度都高于糖含量与糖代谢酶基因表达之间的相关程度，外源ABA通过调控低温下冬小麦蔗糖代谢相关酶的活性和呼吸代谢相关酶的活性而影响着蔗糖代谢及呼吸代谢，蔗糖代谢酶之间，以及呼吸代谢酶之间相互影响、相互协调，错综复杂地调控蔗糖代谢和呼吸代谢的进行。
4．低温胁迫时外源ABA对冬小麦抗寒基因表达的影响
外源ABA主要通过依赖于ABA的信号途径影响冬小麦抗寒基因的表达，并且这种影响在抗寒性强的东农冬麦1号中比冷敏感的济麦22中更明显。苗期低温时，在东农冬麦1号叶片中外源ABA在低温胁迫8h前促进了依赖于ABA的TaCBFIVd-D22和TaMYB80基因的表达，在低温胁迫2h前促进了依赖于ABA的TaWabi5和TaWrab17基因的表达，在东农冬麦1号地下茎中，外源ABA在低温胁迫时促进了TaMYB80、TaWabi5和TaWrab17基因的表达，低温胁迫8h后促进TaCBFIVd-D22基因的表达；在济麦22叶片中，外源ABA在低温胁迫0.5h前促进了TaWabi5基因的表达，在低温胁迫8h前促进了TaCBFIVd-D22和TaMYB80基因的表达，但在济麦22的地下茎中，外源ABA则抑制了这四种基因的表达。在越冬期时，在东农冬麦1号叶片中，外源ABA促进了TaMYB80、TaWabi5和TaWrab17基因的表达，在0℃及以下低温时也促进了TaCBFIVd-D22基因的表达，在济麦22叶片中，0℃以上温度时促进了TaCBFIVd-D22、TaMYB80和TaWrab17基因的表达，在-10℃以上温度时也促进了TaWabi5基因的表达；在东农冬麦1号和济麦22的分蘖节中，外源ABA促进了TaCBFIVd-D22基因的表达，在0℃以上温度时也促进了TaWabi5基因的表达，在东农冬麦1号分蘖节中，外源ABA抑制了TaMYB80和TaWrab17基因的表达，在济麦22分蘖节中0℃以上温度时促进了TaWrab17基因的表达，而TaMYB80基因的表达也受到抑制。低温时外源ABA对不依赖于ABA的信号途径中的抗寒基因TaCBFII-5.2、TaWcor15、TaWcs120和TaCOR39的表达也有一定的影响，也能在一定程度上促进这些基因的表达，但多数情况下影响不明显或抑制这些基因的表达。
The Triticum aestivum Dongnongdongmai1cultivar has an85%winter survival rate and isthe only winter wheat variety grown in Heilongjiang Province (minimum temperature up to-30℃).The T. Aestivum Jimai22cultivar is widely planted in northern China, also has a strong coldresistance, but cannot survive in the cold winter temperatures in Heilongjiang Province. In thisstudy, both winter wheat were used as test materials, and the effects of exogenous ABA on sugarmetabolism and expression of antifreeze genes at low temperature stress were discussed in theseedling stage and wintering stage, and to understand the physiological and molecular basis forexogenous ABA regulating winter wheat sugar metabolism and expressions of antifreeze genes,and it will provide new ideas for revealing exogenous ABA improving cold hardiness of winterwheat. Dongnongdongmai1is the only winter wheat varieties that can be grown in the alpineregion of Heilongjiang Province, and it is typical and representative; Low temperature treatingwheat seedlings in room was adopted in studying cold resistance of winter wheat at home andabroad, but in this study, stimulated low temperature and natural low temperature in the field werecombined, and it is closer to the actual production; In this study, physiological and biochemicaltechnology and molecular biology techniques experimental method were used to study the ABAregulating winter wheat cold tolerance from the point of sugar metabolism and cold tolerance geneexpression at first time to overall grasp the source and path of the carbohydrate, and clarify theeffects of the ABA on plant energy flows, the penetration level, the energy metabolism undersignal effect and cold tolerance gene regulation process, and will provide theoretical basis forantifreeze mechanism of winter wheat, and make an important contribution to the improvement ofthe cultivation of winter wheat in the northern alpine region in the end. The results are as follows:
1. Effect of exogenous ABA on winter sugar metabolism in seedling stage at low temperatures
Exogenous ABA promoted the accumulation of soluble sugar、sucrose content and fructosecontent to improve the soluble sugar content in the leaves of Dongnongdongmai1with strong coldresistance in seedling stages at low temperatures, and exogenous ABA promoted the accumulatonof sugars in the underground stems above0℃; Accumulation of sugars in the leaves andunderground stems of Jimai22was not promoted by exogenous ABA. Starch accumulation wassuppressed by exogenous ABA in the leaves and underground stems of the two wheat or the differences between control and ABA treatment was not significant. Exogenous ABA promoted theexpression levels of TaRBC、TaFBP、TaPRK、TaUGP、TaSAInv、TaSPS、TaPK、TaG6PDH、TaTPT and TaSuT in the leaves and underground stems of Dongnongdongmai1to different extent,and these genes are involved in the carbon fixed, synthesis and degradation of sucrose、glycolysis、pentose phosphate pathway and sugar transport. But, some genes such as TaAGP, TaPDand TaHxK were down-regulated by exogenous ABA in Dongnongdongmai1; Most key enzymesgenes of sugar metabolism were suppressed by exogenous ABA in Jimai22.
2. Effect of exogenous ABA on winter sugar metabolism in wintering stage at low temperatures
In wintering period, more soluble sugars, sucrose and fructose were accumulated in leaves andtillering nodes of ABA treated seedlings of the two wheat cultivars, especially in the winteringorgan tillering nodes. More carbohydrates were accumulated in cold-resistant wheat cultivarDongnongdongmai1than in cold-sensitive wheat cultivar Jimai22. Exogenous ABA improved theaccumulation of starch in the two tissues of Dongnongdongmai1and the leaves of Jimai22, butthe accumulation in the tilleringnodes of Jimai22was not obviously effected by exogenous ABA.Exogenous ABA promoted the expression of TaRBC、TaFBP、TaPRK、TaUGP、TaAGP、TaSAInv、TaSS、TaSPS、TaPK、TaH xK、TaPFK、TaPD、TaG6PDH and TaSuT gene expression in the leavesand tillering nodes of Dongnongdongmai1and Jimai22to different extent, but a greater extentwas showed in the Dongnongdongmai1than Jimai22. Exogenous ABA significantly promotedexpression of TaPRK, TaUGP, TaSPS, TaSAInv, TaPD, and TaPFK gene in leaves of two varietiesof winter wheat, and TaTPT only upregulated in the leaves of Dongnongdongmai1. ExogenousABA promoted sucrose accumulation and also promoted the decomposition of sucrose, but theaccumulation was greater than decomposition, while the SPS and SS played a role in terms ofsucrose accumulation, and UGP played important role in sucrose accumulation and decompositionat the same time, acid invertase and alkaline invertase enzymes played roles in the decompositionof sucrose; ABA treatment improved the glycolytic metabolic enzyme pyruvate kinase, hexokinaseand phosphofructokinase activity to varing degrees in the leaves and tillering nodes of the twovarieties of wheat, especially in the minimum temperatures-25℃,enzyme activities decreaseswere inhibited by exogenous ABA, there by slowing the injury of temperature drops to winterwheat, but this effect was even more marked in cold-resistant wheat cultivar Dongnongdongmai1than in cold-sensitive wheat cultivar Jimai22, and the acetoneacid kinase activity was inhibited byABAin the tillering nodes of Jimai22.
3. Correlation analysis between the sugar content and the expression of key enzyme genes of andenzyme activity of sugar metabolism
The correlation between sugar content and gene expression of sugar metabolism was more prevalent in ABA treated group. There was a certain consistency regulation on sugar content andsugar metabolism gene expression by exogenous ABA. But the correlation between the sugarcontents and sugar metabolism gene expression in control and ABA treated group were not entirelyconsistent at seedling stage and wintering period, this indicated that there was a complexrelationship between the accumulation of carbohydrate and sugar metabolism gene expression.Correlation analysis between the sugar content and the activities of sugar metabolism in winteringstage indicated that the correlation was more prevalent in ABA treated group.The degree ofcorrelaton between sugar content and sugar metabolic enzymes activites was higher than thecorrelation between sugar content and sugar metabolism gene expression in leaves of the twowheat cultivars. Exogenous ABA affected sucrose metabolism and respiratory metabolism troughregulating the sucrose metabolism enzyme activities and respiratory metabolism related enzymesactivities in winter wheat at low temperatures. Surose metabolism enzymes as well as respiratoryMetablosm enzymes were affected and coordinated each other and regulated the sucrosemetabolism and respiratory metabolism intricately.
4. Effect of exogenous ABA on expression of antifreeze genes at low temperatures
Exogenous ABA affected antifreeze genes expressions of winter wheat by ABA-dependentsignaling pathways, and these effects were more obvious in in cold-resistant wheat cultivarDongnongdongmai1than in cold-sensitive wheat cultivar Jimai22. In seedling stage, at lowtemperatures, exogenous ABA promoted the expressions of TaCBFIVd-D22and TaMYB80genesbefore8h low temperature stress, and promoted the expressions of TaWabi5and TaWrab17genesin leaves of Dongnongdongmai1, and exogenous ABA promoted the expression of TaMYB80、TaWabi5and TaWrab17genes, and promoted the expression of TaCBFIVd-D22genes before8hlow temperature stress in underground stems of Dongnongdongmai1; In the leaves of Jimai22, theexpression of TaWabi5was upregulated by exogenous ABA at0.5h of low temperature stress, andthe expressions of TaCBFIVd-D22and TaMYB80were upregulated before8h low temperaturestress, the expression of these four genes were all inhibited by exogenous ABA in undergroundstems of Jimai22. In wintering stage, exogenous ABA promoted the expressions of TaMYB80、TaWabi5and TaWrab17genes and promoted the expression of TaCBFIVd-D22above0℃in theleaves of Dongnongdongmai1, and the expressions of TaCBFIVd-D22、TaMYB80and TaWrab17genes were promoted by exogenous ABA and the expression of TaWabi5gene was promoted byexogenous ABA as well in leaves of Jimai22; In tillering nodes of Dongnongdongmai1and Jimai22, exogenous ABA promoted the expression of TaCBFIVd-D22, and also pomoted the expressionof TaWabi5gene above0℃, and the expressions of TaMYB80and TaWrab17were all inhibitedby exogenous ABA in tillering nodes of Dongnongdongmai1, while in tillering nodes of Jimai22,exogenous ABA promoted the expression of TaWrab17, and inhibited the expression of TaMYB80. There were some impacts of exogenous ABA on cold resistance genes in the ABA-independentsignaling pathways as well, and expressions of these genes were promoted by exogenous ABA to acertain extent.
结果如下：
1．苗期低温下外源ABA对冬小麦糖代谢的影响
苗期低温胁迫时，外源ABA促进了抗寒性强的东农冬麦1号叶片中蔗糖、果糖的积累，从而提高了可溶性糖的含量，在地下茎中则在0℃以上时促进糖类的积累；而外源ABA没有促进济麦22的叶片和地下茎中糖类的积累；淀粉的积累在两品种小麦的叶片和地下茎中则受到抑制或与对照差异不显著。外源ABA对东农冬麦1号两种器官中TaRBC、TaFBP、TaPRK、TaUGP、TaSAInv、TaSPS、TaPK、TaG6PDH、TaTPT和TaSuT的表达有不同程度的促进作用，这些基因分别参与碳的固定、蔗糖的合成和分解、糖酵解、戊糖磷酸途径以及糖的运输，但在东农冬麦1号中，外源ABA处理后也有些基因如TaAGP、TaPD和TaHxK下调表达；在济麦22中，外源ABA处理后大部分糖代谢酶基因的表达都受到了抑制。
2．越冬期外源ABA对冬小麦糖代谢的影响
越冬期低温时，两品种冬小麦ABA处理组的叶片和分蘖节中都积累了更多的可溶性总糖、蔗糖和果糖，尤其是在越冬器官分蘖节中，在抗寒性强的东农冬麦1号中比抗寒性弱的济麦22中积累了更多的碳水化合物，外源ABA促进了东农冬麦1号两种器官和济麦22叶片中淀粉的积累，但对济麦22分蘖节中淀粉积累的影响却不明显。外源ABA不同程度地促进了东农冬麦1号和济麦22叶片和分蘖节中TaRBC、TaFBP、TaPRK、TaUGP、TaAGP、TaSAInv、TaSS、TaSPS、TaPK、TaH×K、TaPFK、TaPD、TaG6PDH和TaSuT基因的表达，但在东农冬麦1号中的促进程度大于济麦22，外源ABA显著地促进了TaPRK, TaUGP, TaSPS, TaSAInv, TaPD和TaPFK基因在两品种冬小麦叶片中的表达，而TaTPT仅在东农冬麦1号的叶片中上调表达。外源ABA促进蔗糖积累的同时也促进了蔗糖的分解，但积累大于分解，而蔗糖磷酸合成酶（SPS）与蔗糖合成酶（SS）在蔗糖积累方面起作用，尿苷二磷酸-葡萄糖焦磷酸化酶（UGP）对蔗糖的积累和分解同时起作用，酸性转化酶和碱性转化酶则在蔗糖的分解方向起作用；ABA处理不同程度地提高了两个品种小麦叶片及分蘖节中糖酵解代谢酶丙酮酸激酶、己糖激酶和磷酸果糖激酶的活性，尤其在最低温-25℃时明显抑制这几种酶活性的降低，从而减缓温度下降给冬小麦带来的伤害，但这种影响在抗寒性强的东农冬麦1号中表现得更明显，而在冷敏感的济麦22中相对不明显，丙酮酸激酶活性在济麦22分蘖节中甚至受到ABA的抑制。
3．糖含量与糖代谢关键酶基因表达以及酶活性间的相关分析
ABA处理组的糖含量与糖代谢酶基因表达之间的相关性更为普遍，外源ABA对糖含量与糖代谢酶基因的调控具有一定的一致性，但在苗期和越冬期，对照和ABA处理组中糖含量与糖代谢酶基因之间的相关性并不完全一致，表明糖类物质的积累与糖代谢酶基因的表达之间具有复杂的关系。越冬期通过对糖含量与糖代谢酶活性间的相关分析表明，ABA处理后也使糖含量与酶活性间的相关更为普遍，两小麦的叶片中糖含量与糖代谢酶活性的相关程度都高于糖含量与糖代谢酶基因表达之间的相关程度，外源ABA通过调控低温下冬小麦蔗糖代谢相关酶的活性和呼吸代谢相关酶的活性而影响着蔗糖代谢及呼吸代谢，蔗糖代谢酶之间，以及呼吸代谢酶之间相互影响、相互协调，错综复杂地调控蔗糖代谢和呼吸代谢的进行。
4．低温胁迫时外源ABA对冬小麦抗寒基因表达的影响
外源ABA主要通过依赖于ABA的信号途径影响冬小麦抗寒基因的表达，并且这种影响在抗寒性强的东农冬麦1号中比冷敏感的济麦22中更明显。苗期低温时，在东农冬麦1号叶片中外源ABA在低温胁迫8h前促进了依赖于ABA的TaCBFIVd-D22和TaMYB80基因的表达，在低温胁迫2h前促进了依赖于ABA的TaWabi5和TaWrab17基因的表达，在东农冬麦1号地下茎中，外源ABA在低温胁迫时促进了TaMYB80、TaWabi5和TaWrab17基因的表达，低温胁迫8h后促进TaCBFIVd-D22基因的表达；在济麦22叶片中，外源ABA在低温胁迫0.5h前促进了TaWabi5基因的表达，在低温胁迫8h前促进了TaCBFIVd-D22和TaMYB80基因的表达，但在济麦22的地下茎中，外源ABA则抑制了这四种基因的表达。在越冬期时，在东农冬麦1号叶片中，外源ABA促进了TaMYB80、TaWabi5和TaWrab17基因的表达，在0℃及以下低温时也促进了TaCBFIVd-D22基因的表达，在济麦22叶片中，0℃以上温度时促进了TaCBFIVd-D22、TaMYB80和TaWrab17基因的表达，在-10℃以上温度时也促进了TaWabi5基因的表达；在东农冬麦1号和济麦22的分蘖节中，外源ABA促进了TaCBFIVd-D22基因的表达，在0℃以上温度时也促进了TaWabi5基因的表达，在东农冬麦1号分蘖节中，外源ABA抑制了TaMYB80和TaWrab17基因的表达，在济麦22分蘖节中0℃以上温度时促进了TaWrab17基因的表达，而TaMYB80基因的表达也受到抑制。低温时外源ABA对不依赖于ABA的信号途径中的抗寒基因TaCBFII-5.2、TaWcor15、TaWcs120和TaCOR39的表达也有一定的影响，也能在一定程度上促进这些基因的表达，但多数情况下影响不明显或抑制这些基因的表达。
The Triticum aestivum Dongnongdongmai1cultivar has an85%winter survival rate and isthe only winter wheat variety grown in Heilongjiang Province (minimum temperature up to-30℃).The T. Aestivum Jimai22cultivar is widely planted in northern China, also has a strong coldresistance, but cannot survive in the cold winter temperatures in Heilongjiang Province. In thisstudy, both winter wheat were used as test materials, and the effects of exogenous ABA on sugarmetabolism and expression of antifreeze genes at low temperature stress were discussed in theseedling stage and wintering stage, and to understand the physiological and molecular basis forexogenous ABA regulating winter wheat sugar metabolism and expressions of antifreeze genes,and it will provide new ideas for revealing exogenous ABA improving cold hardiness of winterwheat. Dongnongdongmai1is the only winter wheat varieties that can be grown in the alpineregion of Heilongjiang Province, and it is typical and representative; Low temperature treatingwheat seedlings in room was adopted in studying cold resistance of winter wheat at home andabroad, but in this study, stimulated low temperature and natural low temperature in the field werecombined, and it is closer to the actual production; In this study, physiological and biochemicaltechnology and molecular biology techniques experimental method were used to study the ABAregulating winter wheat cold tolerance from the point of sugar metabolism and cold tolerance geneexpression at first time to overall grasp the source and path of the carbohydrate, and clarify theeffects of the ABA on plant energy flows, the penetration level, the energy metabolism undersignal effect and cold tolerance gene regulation process, and will provide theoretical basis forantifreeze mechanism of winter wheat, and make an important contribution to the improvement ofthe cultivation of winter wheat in the northern alpine region in the end. The results are as follows:
1. Effect of exogenous ABA on winter sugar metabolism in seedling stage at low temperatures
Exogenous ABA promoted the accumulation of soluble sugar、sucrose content and fructosecontent to improve the soluble sugar content in the leaves of Dongnongdongmai1with strong coldresistance in seedling stages at low temperatures, and exogenous ABA promoted the accumulatonof sugars in the underground stems above0℃; Accumulation of sugars in the leaves andunderground stems of Jimai22was not promoted by exogenous ABA. Starch accumulation wassuppressed by exogenous ABA in the leaves and underground stems of the two wheat or the differences between control and ABA treatment was not significant. Exogenous ABA promoted theexpression levels of TaRBC、TaFBP、TaPRK、TaUGP、TaSAInv、TaSPS、TaPK、TaG6PDH、TaTPT and TaSuT in the leaves and underground stems of Dongnongdongmai1to different extent,and these genes are involved in the carbon fixed, synthesis and degradation of sucrose、glycolysis、pentose phosphate pathway and sugar transport. But, some genes such as TaAGP, TaPDand TaHxK were down-regulated by exogenous ABA in Dongnongdongmai1; Most key enzymesgenes of sugar metabolism were suppressed by exogenous ABA in Jimai22.
2. Effect of exogenous ABA on winter sugar metabolism in wintering stage at low temperatures
In wintering period, more soluble sugars, sucrose and fructose were accumulated in leaves andtillering nodes of ABA treated seedlings of the two wheat cultivars, especially in the winteringorgan tillering nodes. More carbohydrates were accumulated in cold-resistant wheat cultivarDongnongdongmai1than in cold-sensitive wheat cultivar Jimai22. Exogenous ABA improved theaccumulation of starch in the two tissues of Dongnongdongmai1and the leaves of Jimai22, butthe accumulation in the tilleringnodes of Jimai22was not obviously effected by exogenous ABA.Exogenous ABA promoted the expression of TaRBC、TaFBP、TaPRK、TaUGP、TaAGP、TaSAInv、TaSS、TaSPS、TaPK、TaH xK、TaPFK、TaPD、TaG6PDH and TaSuT gene expression in the leavesand tillering nodes of Dongnongdongmai1and Jimai22to different extent, but a greater extentwas showed in the Dongnongdongmai1than Jimai22. Exogenous ABA significantly promotedexpression of TaPRK, TaUGP, TaSPS, TaSAInv, TaPD, and TaPFK gene in leaves of two varietiesof winter wheat, and TaTPT only upregulated in the leaves of Dongnongdongmai1. ExogenousABA promoted sucrose accumulation and also promoted the decomposition of sucrose, but theaccumulation was greater than decomposition, while the SPS and SS played a role in terms ofsucrose accumulation, and UGP played important role in sucrose accumulation and decompositionat the same time, acid invertase and alkaline invertase enzymes played roles in the decompositionof sucrose; ABA treatment improved the glycolytic metabolic enzyme pyruvate kinase, hexokinaseand phosphofructokinase activity to varing degrees in the leaves and tillering nodes of the twovarieties of wheat, especially in the minimum temperatures-25℃,enzyme activities decreaseswere inhibited by exogenous ABA, there by slowing the injury of temperature drops to winterwheat, but this effect was even more marked in cold-resistant wheat cultivar Dongnongdongmai1than in cold-sensitive wheat cultivar Jimai22, and the acetoneacid kinase activity was inhibited byABAin the tillering nodes of Jimai22.
3. Correlation analysis between the sugar content and the expression of key enzyme genes of andenzyme activity of sugar metabolism
The correlation between sugar content and gene expression of sugar metabolism was more prevalent in ABA treated group. There was a certain consistency regulation on sugar content andsugar metabolism gene expression by exogenous ABA. But the correlation between the sugarcontents and sugar metabolism gene expression in control and ABA treated group were not entirelyconsistent at seedling stage and wintering period, this indicated that there was a complexrelationship between the accumulation of carbohydrate and sugar metabolism gene expression.Correlation analysis between the sugar content and the activities of sugar metabolism in winteringstage indicated that the correlation was more prevalent in ABA treated group.The degree ofcorrelaton between sugar content and sugar metabolic enzymes activites was higher than thecorrelation between sugar content and sugar metabolism gene expression in leaves of the twowheat cultivars. Exogenous ABA affected sucrose metabolism and respiratory metabolism troughregulating the sucrose metabolism enzyme activities and respiratory metabolism related enzymesactivities in winter wheat at low temperatures. Surose metabolism enzymes as well as respiratoryMetablosm enzymes were affected and coordinated each other and regulated the sucrosemetabolism and respiratory metabolism intricately.
4. Effect of exogenous ABA on expression of antifreeze genes at low temperatures
Exogenous ABA affected antifreeze genes expressions of winter wheat by ABA-dependentsignaling pathways, and these effects were more obvious in in cold-resistant wheat cultivarDongnongdongmai1than in cold-sensitive wheat cultivar Jimai22. In seedling stage, at lowtemperatures, exogenous ABA promoted the expressions of TaCBFIVd-D22and TaMYB80genesbefore8h low temperature stress, and promoted the expressions of TaWabi5and TaWrab17genesin leaves of Dongnongdongmai1, and exogenous ABA promoted the expression of TaMYB80、TaWabi5and TaWrab17genes, and promoted the expression of TaCBFIVd-D22genes before8hlow temperature stress in underground stems of Dongnongdongmai1; In the leaves of Jimai22, theexpression of TaWabi5was upregulated by exogenous ABA at0.5h of low temperature stress, andthe expressions of TaCBFIVd-D22and TaMYB80were upregulated before8h low temperaturestress, the expression of these four genes were all inhibited by exogenous ABA in undergroundstems of Jimai22. In wintering stage, exogenous ABA promoted the expressions of TaMYB80、TaWabi5and TaWrab17genes and promoted the expression of TaCBFIVd-D22above0℃in theleaves of Dongnongdongmai1, and the expressions of TaCBFIVd-D22、TaMYB80and TaWrab17genes were promoted by exogenous ABA and the expression of TaWabi5gene was promoted byexogenous ABA as well in leaves of Jimai22; In tillering nodes of Dongnongdongmai1and Jimai22, exogenous ABA promoted the expression of TaCBFIVd-D22, and also pomoted the expressionof TaWabi5gene above0℃, and the expressions of TaMYB80and TaWrab17were all inhibitedby exogenous ABA in tillering nodes of Dongnongdongmai1, while in tillering nodes of Jimai22,exogenous ABA promoted the expression of TaWrab17, and inhibited the expression of TaMYB80. There were some impacts of exogenous ABA on cold resistance genes in the ABA-independentsignaling pathways as well, and expressions of these genes were promoted by exogenous ABA to acertain extent.
引文
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