土壤水分对寒地冬小麦越冬性的影响
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摘要
本研究于2009-2010年在黑龙江省哈尔滨市进行,以抗寒性存在差异的两个冬小麦品种为试验材料,并根据常年土壤水分含量与返青率的关系,设置了不同的水分梯度处理,对越冬期间冬小麦植株的光合特性、生理生化指标、以及蛋白质组等方面展开研究,以明确寒地冬小麦安全越冬的适宜土壤水分,探讨品种抗寒性鉴定的指标和方法,并从蛋白表达影响上揭示寒地冬小麦安全越冬的机制,为寒地冬小麦的生产提供理论依据。得到的主要结论如下：
1.从返青情况看,不同土壤水分处理下的济麦22均没有越冬返青；东农冬麦1号在不同水分处理下均能返青,其返青率在水分处理间差异极显著,表现为CK(正常水分管理)>30%±2%>45%±2%>15%±2%,其返青率分别为：83.3%、49.2%、23.3%、2.5%。随着取样日期的延后,两个品种叶片、叶鞘的死细胞比例逐渐加重。进入封冻期20d以后品种间细胞活力差异明显,表现为东农冬麦1号受损程度轻于济麦22。不同土壤水分处理下,CK处理(正常水分管理)下受损最轻；叶片受损重于叶鞘。从封冻期以后主茎生长锥发育情况上看,东农冬麦1号茎生长锥分化进程慢,且受水分影响较小分化进程主要集中在伸长期；济麦22分化进程快,分化进程集中在伸长期和单棱期,45%±2%水分处理下分化进程快。
2.两个品种叶片、叶鞘总水分含量随取样时间变化表现为先降低后升高的趋势,两个品种的15%±2%土壤水分处理下的叶片、叶鞘总含水量较低,济麦22高于东农冬麦1号,叶鞘高于叶片。取样期间15%±2%土壤水分处理叶片、叶鞘自由水含量较低,而45%±2%土壤水分处理叶片、叶鞘自由水含量较高。东农冬麦1号的叶鞘在取样期间保持着较高的束缚水含量,而济麦22多数水分处理下的束缚水含量随取样日期表现为双峰曲线变化。
3.随着取样日期的延后,两个品种各水分处理可溶性蛋白表现为增加趋势；取样期间过度干旱和水分过多条件均有助于可溶性蛋白含量升高,且品种间差异不明显。
4.随取样日期延后,两个品种叶片、叶鞘的相对电导率逐渐增大,前期上升快速,后期趋于平稳；CK处理(正常水分管理)下最低,15%±2%水分处理最高。品种间相比,东农冬麦1号的相对电导率小于济麦22。取样部位间,叶鞘大于叶片。在MDA含量上,两品种MDA含量随取样日期变化都呈现出先降低后升高的趋势；水分间,两品种15%±2%土壤水分处理的MDA含量相对较大,而CK处理(正常水分管理)相对较小30%±2%土壤水分与45%±2%土壤水分处理居中,即干旱和水分过多都会导致膜脂过氧化程度加重。
5.在SOD活性上,随取样日期延后两个品种的SOD活性都表现为波动变化,东衣冬麦1号水分处理间大小表现为15%±2%>30%±2%>45%±2%>CK(正常水分管理)：济麦22品种在15%±2%处理上具有较高的SOD活性。POD活性上,随取样日期变化有增大趋势；品种间相比,东农冬麦1号POD活性较高；水分间相比,东农冬麦1号低温驯化期15%±2%土壤水分处理较高,封冻期以后45%±2%土壤水分处理较高；越冬期间济麦22的15%±2%土壤水分处理具有较高的POD活性。GSH含量上,两品种GSH含量随取样日期变化都表现为降低的趋势；东农冬麦1号高于济麦22；东农冬麦1号在30%±2%水分处理上GSH含量较高,而济麦22则在45%±2%处理上较高。抗坏血酸(AsA)含量上,东农冬麦1号随取样日期变化表现为先增大后减小的特点；两品种在低温驯化期CK处理(正常水分管理)具有较小值；封冻期以后东农冬麦1号CK处理AsA含量较高,而济麦22各水分处理下都波动剧烈。
6.两个品种各水分处理的初始荧光(Fo)随取样日期都呈双峰曲线变化,干旱和水分过多的土壤条件均可引起Fo的增大。在封冻期以前两个品种的各水分处理下最大荧光(Fm)迅速增大,封冻后又迅速降低；进入封冻期20d以后随日期变化相对平稳,此时CK处理(正常水分管理)Fm具有最大值。从低温驯化期至封冻期东农冬麦1号各水分处理下的最大量子产量(Fv/Fm)和实际量子产量(Yield)均呈平稳变化或略有增加的趋势,而济麦22都表现为降低趋势。封冻期以后,两品种的Fv/Fm和Yield都逐渐降低；封冻期以后,两品种的CK(正常水分管理)处理的Fv/Fm和Yield都明显高于其他水分亏缺或水分过多处理。东农冬麦1号不同土壤水分处理下光化学淬灭系数(qP)随取样日期变化逐渐降低,而济麦22则呈波动变化,整体表现为降低趋势。两品种非光化学淬灭系数(qN)随取样日期呈现波动变化,整体表现为增大趋势。两个品种的实时荧光产量(Ft)在取样日期间均差异极显著；低温驯化期,土壤适当干旱可引起Ft的上升,封冻初期,土壤干旱条件反而会造成Ft的下降。随取样日期的延后,两个品种光合电子传递速率(ETR)有逐渐降低的趋势,封冻期后急剧下降；适宜和过量的土壤水分条件其光合电子传递速率要好于干旱缺水的条件；品种间比较,在低温驯化期东农冬麦1号发生光抑制重于济麦22,封冻期后则相反。东农冬麦1号品种的瞬时最大量子产量(Fm')大于济麦22品种,在各光化光下两个品种间均差异显著或极显著；封冻初期CK处理(正常水分管理)Fm'具有较小值,12月2日以后,CK处理值相对较大；在低温驯化期和封冻初期,两个品种Fm'随光化光都呈波动变化,封冻期以后,则呈平稳变化。
7.两个品种15%±2%、30%±2%和45%±2%水分处理下在初始封冻期和封冻后期共有32个差异蛋白点,其中26个与小麦蛋白匹配度较高,3个与水稻蛋白匹配度较高,1个与玉米蛋白匹配度较高,1个与大麦蛋白匹配度较高,还有1个蛋白点与黄瓜蛋白匹配度高。这些蛋白点参与能量代谢的占12.5%、物质合成的占25%、分子伴侣的占9.38%、抗性蛋白的占31.25%,光合反应的占6.25%、其他类蛋白占15.62%。封冻初期抗寒品种与不抗寒品种共同表达的蛋白点有9个,这9个蛋白参与渗透调节物质的形成、参与清除对细胞伤害的活性物质的蛋白、抗性蛋白和参与光合作用的蛋。区别是抗寒品种参与能量代谢的差异蛋白点较不抗寒品种高；封冻后期抗寒品种与不抗寒品种没有共同表达的差异蛋白,抗寒品种差异表达的蛋白参与细胞壁的加厚、能量代谢和抗性蛋白,而不抗寒品种差异表达的蛋白参与细胞膜的降解和光合作用。重度缺水(15%±2%)和过湿(45%±2%)土壤水分下差异表达的蛋白高,轻度干旱的土壤水分条件(30%±2%)下差异表达的蛋白少。
8.抗寒性强的品种具有如下的特征：越冬期间组织受损轻,活细胞数量多,茎生长锥分化进程慢；越冬期间叶鞘总含水量处于较低水平；越冬期间相对电导率低,质膜透性小；低温驯化期叶鞘的丙二醛含量处于较低水平；低温驯化期至封冻初期,SOD活性、POD活性高；整个越冬期间的GSH含量高；在低温驯化期qP、qN值大；封冻期以后,Fv/Fm、Yield值高；进入封冻期20d以后,Fo值低；整个越冬期间Fm值高。可以采用植株活力观察、茎生长锥分化观察、相对电导率、丙二醛含量、叶绿素荧光参数(Fo、Fm、Fv/Fm、Yield、qP、qN)、SOD活性、POD活性和GSH含量这些指标对寒地冬小麦的抗寒性能进行评价。
9.试验中春季返青率,越冬期间的植株活力、相对电导率、丙二醛含量,进入封冻期20d以后的最大荧光、瞬时最大量子产量、封冻期以后的最大量子产量和实际量子产量,越冬期间的光合电子传递速率这些指标与抗寒性关系密切,且受土壤水分含量影响较大。依据这些指标与抗寒性的关系及随土壤水分的变化情况,最终得出入冬前土壤绝对含水量为35%-40%时,更利于冬小麦的越冬生长。
The trial for the effects of the soil moisture on the overwintering survival rates of winter wheat was carried out in Harbin,Heilongjiang prvince,China,which located in the frigid region from 2009 to 2010.Two winter wheat cultivars different in the cold hardiness were used in this trial.Different soil water content gradients were set up based on the relation between the soil water contents and the overwinter survival rate of winter wheat.The photosynthetic preperties,physiological and biochemical indices and proteomics and so on in winter wheat plants during winter season were studied,and furthermore,the optical soil water content for the safe overwintering of winter wheat was definited,the indices and methods in the identification for the cold tolerance of wheat were explored and the mechanism in the safe overwintering of winter wheat in frigid region was revealed through the influence of proteins expression,and the theoretic fundation was provided for the production of winter wheat in the frigid area.The main conclusions were as follows:
1.On the overwintering cases none of Jimai 22 but all of Dongnongdongmai 1 was able to be overwinterin in the all water treatments.It was very significant in the difference in the overwinter survival rate of Dongnongdongmai 1 among the soil water treatments,appearing CK(normal soil water management)>30%±2%>45%±2%>15%±2% in the soil moisture,and was 83.3%,49.2%,23.3%,2.5%,in the survival rate respectively.The ratio of the dead cells in the leaf and leaf sheath of the two cultivars was gradually enhanced with the delay of the sampling date.And the obvious difference in the cell vigor between cultivars occurred at the 20th day after frozen date in winter and the injury in Dongnongdongmai 1 was less than that in Jimai 22.The damage of plants under CK (normal soil water management) in soil water content were slightest.The extent of damage in the leaf was more serious than that in the leaf sheath.The change of the development status of the stem apical point was very slow and less affected by the soil water content after entering the frozen period and the profile of the apical point almost was in the elongation stage.On the contrary/the transition of the apical point in Jimai 22 was fast and quicker in the soil water treatment of 45%±2% and the form of the part was in the elongation and single ridge.
2.The water content in the leaf and leaf sheath of the 2 cultivars performed downward and then upward as the sampling dates increased and lower in 15%±2% of soil water treatment. This trait in Jimai 22 was higher than that in Dongnongdongmai 1 and the one in leaf sheath higher than that in leaf.The free water content in samples was lower in 15%±2% and higher in 45%±2% in the soil water content treatment.The bound water content in the leaf sheath of Dongnongdongmai 1 remained higher and that of Jimai22 performed bimodal curve in the whole sampling period.
3.The soluble protein content of the 2 cultivars increased in all soil water theatments and would be facilited to a higher level both under drought and hyperhydration condition during the sampling period.No significant difference between the 2 cultivars was observed in this trait.
4.The relative electric conductivity in the leafs and leaf sheaths of the 2 cultivars elevated gradually during the sampling period and quickly in the early days and smoothly in the late days and lowest in CK (normal soil water management) and highest in 15%±2% of soil water treatment.The conductivity of Dongnongdongmai 1 was lower than that of Jimai22 and one in the leaf sheath higher than that in leaf.MDA of both cultivars appeared downward in the early days and upward in the late days in the sampling period.The content of MDA in the cultivars was higher relatively in 15%±2% and lower in CK(normal soil water management) and media both in 30%±2% and 45%±2% in the soil water treatment.The level of membrane lipid overoxidation was found to be worsened both in dought and overmoisture.
5.The SOD activity of cultivars behaved in fluctuation during the sampling period.The order in the trait of Dongnongdongmai 1 among the soil water treatments showed as 15%±2%>30%±2%>45%±2%>CK (normal soil water management) and of Jimai 22 the highest in 15%±2%.And the POD activity appeared enhancing trands during the same period and higher in Dongnongdongmai 1 than in Jimai 22.The higher POD activity of Dongnongdongmai 1 presented in 15%±2% of soil water content in cold acclimation (?) in 45%±2% after frozen date and of Jimai 22 in 15%±2% covering winter,respectively.The GSH contents of 2 cultivars were downward during the sampling period and higher in Dongnongdongmai 1 than in Jimai 22 and of Dongnongdongmai 1 in 30%±2% while of Jimai 22 in 45%±2%..The As A contents of 2 cultivars showed upward in the early stage of the sampling period and then downward in the late stage.The lower value of As A was in CK(normal soil water management) of soil water content during cold acclimation phase. After entering frozen period the higher content of AsA was found in CK of soil water content in Dongnongdongmai 1,but the values of AsA in Jimai 22 waved remarkably in all soil water treatments.
6.The change of the initial fluorescence values (Fo) of both cultivars in the soil water treatments presented the bimodal curv in the whole sampling period.The Fo value would increase under the dought and water logging conditions.The Maximum fluorescence value (Fm) of the 2 cultivars enhanced rapidly in all soil water treatments before the frozen date and decreased fast after the stage.After 20 days entering the frozen stage the change of Fm was smooth while showed highest in CK (normal soil water management).The maximal quantum yield (Fv/Fm) and the actual quantum yield (Yield) of Dongnongdongmai 1 appeared to be smooth or a slightly increasing trend in all soil water treatments from the cold acclimation period to the frozen stage and of Jimai 22 downward.The Fv/Fm and Yield of the cultivars reduced gradually and those values in CK (normal soil water management) treatment were higher obviously than those in other soil water contents after entering into the frozen stage.The photochemical quenching parameter (qP) of Dongnongdongmai 1 in all soil water treatments dropped gradually during the sampling period and of Jiami 22 fluctuated downward.The non-photochemical quenching parameter (qN) of both cultivars appeared to be fluctuating upward.The difference in the actual fluorescence yield (Ft) of both cultivars was very significant among the sampling dates.The value of Ft could be induced to incease in the cold acclimation period but to reduce in the early stage of frozen perion by slightly drought.The photosynthetic electron transfering rate (ETR) of both cultivars decreased gradually in the whole sampling period and sharply in the frozen period and was higher under the optical and some more soil water contents than that under drought condition. The photoinhibition in Dongnongdongmai 1 was heavier than that in Jiami 22 in the cold acclimation period and more seriously in Jimai 22 in the frozen period.The transient maximal quantum yield (Fm) of Dongnongdongmai 1 was higher than that of Jimai 22 and the significant difference in Fm between cultivars was observed in all actinic light treatments.The Fm in CK (normal soil water management) was lower in the early stage of the frozen period and became higher after the date of Dec 2.The change of Fm with the actinic light intensives of the cultivars was fluctuating in the cold acclimation period and the early stage of the frozen period and became smooth in the whole frozen period.
7.The number of the different protein spots was found to be 32 in the early and late stages of the frozen period in the soil water contents of 15%±2%,30%±2% and 45%±2% in both cultivars.Of which 26 spots matched highly with proteins found in wheat and 3 with in rice,1 with in maize,1 with in barley and 1 with in cucumber.12.5% of the protein spots participated in the process of energy metablism,and 25% of material synthesis,9.3% belonged to molecular chapron,31.25% to resistant proteins,6.25% of photosynthetic reaction and 15.62% to other proteins.There were 9 protein spots appearing both in the cold tolerant cultivar and cold sensitive cultivar in the early stage of the frozen period and all participated in the process of the osmotic matters synthesis,scavenging of the active matters damaging cell.stress resistance and photosynthesis.The different protein spots participating energy metablism in the cold tolerant cultivar were more than that in the cold sensitive cultivar. There were no different protein spots both expressing in the cold-tolerant and -sensitive cultivars in the late stage of the frozen period.The different expressing protein spots jioned thicking cell wall,energy metablism and stress resistance in the cold tolerant cultivar and in the cold sensitive cultivar participated dissolving cell membrane and photosynthesis.The more of the different expression protein spots occurred in the soil water treatments of dought (15%±2%) and more moisture (45%±2%) and less of slight drought (30%±2%).
8.The strong cold resistant cultivar has characterestics as follows:slight damage of tissues in the frozen period,more alive cells,low progress of apical point development,low level of water content in leaf sheathes in the overwinter period,high activity of SOD,POD in the cold acclimation period and the early stage of the frozen period,high content of GSH in the overwinter period,high in qP,qN in the cold acclimation period,high in Fv/Fm,Yield after the date of frozen,low in Fo after 20 days entering into the frozen period,high in Fm in the whole frozen period.The evaluation for the cold tolerance of winter wheat in the frigid region could be carried into execution by the indices of paints vigor,apical point status,relative conductivity,MDA,chlorophyll fluorescence parameters (Fo,Fm,Fv/Fm,Yield,qP,qN),activity of SOD,POD and GSH content.
9.The close relationship was found between the cold tolerance of winter wheat and the indices of the overwinter survival rate in spring,plant vigor,conductivity and MDAin winter,the maximal fluorescence and transient maximal quantum yield after 20 days entering into the frozen period,maximal quantum yield and actual quantum yield in the frozen period,ETR in winter. All of the indices were affected obviously by the soil water content.The soil water content of 35%-40% before winter season was in favor of the safe overwintering of winter wheat according to the relationship between the indices and the cold tolerance of winter wheat and its changes with the soil water content levels.
1.从返青情况看,不同土壤水分处理下的济麦22均没有越冬返青；东农冬麦1号在不同水分处理下均能返青,其返青率在水分处理间差异极显著,表现为CK(正常水分管理)>30%±2%>45%±2%>15%±2%,其返青率分别为：83.3%、49.2%、23.3%、2.5%。随着取样日期的延后,两个品种叶片、叶鞘的死细胞比例逐渐加重。进入封冻期20d以后品种间细胞活力差异明显,表现为东农冬麦1号受损程度轻于济麦22。不同土壤水分处理下,CK处理(正常水分管理)下受损最轻；叶片受损重于叶鞘。从封冻期以后主茎生长锥发育情况上看,东农冬麦1号茎生长锥分化进程慢,且受水分影响较小分化进程主要集中在伸长期；济麦22分化进程快,分化进程集中在伸长期和单棱期,45%±2%水分处理下分化进程快。
2.两个品种叶片、叶鞘总水分含量随取样时间变化表现为先降低后升高的趋势,两个品种的15%±2%土壤水分处理下的叶片、叶鞘总含水量较低,济麦22高于东农冬麦1号,叶鞘高于叶片。取样期间15%±2%土壤水分处理叶片、叶鞘自由水含量较低,而45%±2%土壤水分处理叶片、叶鞘自由水含量较高。东农冬麦1号的叶鞘在取样期间保持着较高的束缚水含量,而济麦22多数水分处理下的束缚水含量随取样日期表现为双峰曲线变化。
3.随着取样日期的延后,两个品种各水分处理可溶性蛋白表现为增加趋势；取样期间过度干旱和水分过多条件均有助于可溶性蛋白含量升高,且品种间差异不明显。
4.随取样日期延后,两个品种叶片、叶鞘的相对电导率逐渐增大,前期上升快速,后期趋于平稳；CK处理(正常水分管理)下最低,15%±2%水分处理最高。品种间相比,东农冬麦1号的相对电导率小于济麦22。取样部位间,叶鞘大于叶片。在MDA含量上,两品种MDA含量随取样日期变化都呈现出先降低后升高的趋势；水分间,两品种15%±2%土壤水分处理的MDA含量相对较大,而CK处理(正常水分管理)相对较小30%±2%土壤水分与45%±2%土壤水分处理居中,即干旱和水分过多都会导致膜脂过氧化程度加重。
5.在SOD活性上,随取样日期延后两个品种的SOD活性都表现为波动变化,东衣冬麦1号水分处理间大小表现为15%±2%>30%±2%>45%±2%>CK(正常水分管理)：济麦22品种在15%±2%处理上具有较高的SOD活性。POD活性上,随取样日期变化有增大趋势；品种间相比,东农冬麦1号POD活性较高；水分间相比,东农冬麦1号低温驯化期15%±2%土壤水分处理较高,封冻期以后45%±2%土壤水分处理较高；越冬期间济麦22的15%±2%土壤水分处理具有较高的POD活性。GSH含量上,两品种GSH含量随取样日期变化都表现为降低的趋势；东农冬麦1号高于济麦22；东农冬麦1号在30%±2%水分处理上GSH含量较高,而济麦22则在45%±2%处理上较高。抗坏血酸(AsA)含量上,东农冬麦1号随取样日期变化表现为先增大后减小的特点；两品种在低温驯化期CK处理(正常水分管理)具有较小值；封冻期以后东农冬麦1号CK处理AsA含量较高,而济麦22各水分处理下都波动剧烈。
6.两个品种各水分处理的初始荧光(Fo)随取样日期都呈双峰曲线变化,干旱和水分过多的土壤条件均可引起Fo的增大。在封冻期以前两个品种的各水分处理下最大荧光(Fm)迅速增大,封冻后又迅速降低；进入封冻期20d以后随日期变化相对平稳,此时CK处理(正常水分管理)Fm具有最大值。从低温驯化期至封冻期东农冬麦1号各水分处理下的最大量子产量(Fv/Fm)和实际量子产量(Yield)均呈平稳变化或略有增加的趋势,而济麦22都表现为降低趋势。封冻期以后,两品种的Fv/Fm和Yield都逐渐降低；封冻期以后,两品种的CK(正常水分管理)处理的Fv/Fm和Yield都明显高于其他水分亏缺或水分过多处理。东农冬麦1号不同土壤水分处理下光化学淬灭系数(qP)随取样日期变化逐渐降低,而济麦22则呈波动变化,整体表现为降低趋势。两品种非光化学淬灭系数(qN)随取样日期呈现波动变化,整体表现为增大趋势。两个品种的实时荧光产量(Ft)在取样日期间均差异极显著；低温驯化期,土壤适当干旱可引起Ft的上升,封冻初期,土壤干旱条件反而会造成Ft的下降。随取样日期的延后,两个品种光合电子传递速率(ETR)有逐渐降低的趋势,封冻期后急剧下降；适宜和过量的土壤水分条件其光合电子传递速率要好于干旱缺水的条件；品种间比较,在低温驯化期东农冬麦1号发生光抑制重于济麦22,封冻期后则相反。东农冬麦1号品种的瞬时最大量子产量(Fm')大于济麦22品种,在各光化光下两个品种间均差异显著或极显著；封冻初期CK处理(正常水分管理)Fm'具有较小值,12月2日以后,CK处理值相对较大；在低温驯化期和封冻初期,两个品种Fm'随光化光都呈波动变化,封冻期以后,则呈平稳变化。
7.两个品种15%±2%、30%±2%和45%±2%水分处理下在初始封冻期和封冻后期共有32个差异蛋白点,其中26个与小麦蛋白匹配度较高,3个与水稻蛋白匹配度较高,1个与玉米蛋白匹配度较高,1个与大麦蛋白匹配度较高,还有1个蛋白点与黄瓜蛋白匹配度高。这些蛋白点参与能量代谢的占12.5%、物质合成的占25%、分子伴侣的占9.38%、抗性蛋白的占31.25%,光合反应的占6.25%、其他类蛋白占15.62%。封冻初期抗寒品种与不抗寒品种共同表达的蛋白点有9个,这9个蛋白参与渗透调节物质的形成、参与清除对细胞伤害的活性物质的蛋白、抗性蛋白和参与光合作用的蛋。区别是抗寒品种参与能量代谢的差异蛋白点较不抗寒品种高；封冻后期抗寒品种与不抗寒品种没有共同表达的差异蛋白,抗寒品种差异表达的蛋白参与细胞壁的加厚、能量代谢和抗性蛋白,而不抗寒品种差异表达的蛋白参与细胞膜的降解和光合作用。重度缺水(15%±2%)和过湿(45%±2%)土壤水分下差异表达的蛋白高,轻度干旱的土壤水分条件(30%±2%)下差异表达的蛋白少。
8.抗寒性强的品种具有如下的特征：越冬期间组织受损轻,活细胞数量多,茎生长锥分化进程慢；越冬期间叶鞘总含水量处于较低水平；越冬期间相对电导率低,质膜透性小；低温驯化期叶鞘的丙二醛含量处于较低水平；低温驯化期至封冻初期,SOD活性、POD活性高；整个越冬期间的GSH含量高；在低温驯化期qP、qN值大；封冻期以后,Fv/Fm、Yield值高；进入封冻期20d以后,Fo值低；整个越冬期间Fm值高。可以采用植株活力观察、茎生长锥分化观察、相对电导率、丙二醛含量、叶绿素荧光参数(Fo、Fm、Fv/Fm、Yield、qP、qN)、SOD活性、POD活性和GSH含量这些指标对寒地冬小麦的抗寒性能进行评价。
9.试验中春季返青率,越冬期间的植株活力、相对电导率、丙二醛含量,进入封冻期20d以后的最大荧光、瞬时最大量子产量、封冻期以后的最大量子产量和实际量子产量,越冬期间的光合电子传递速率这些指标与抗寒性关系密切,且受土壤水分含量影响较大。依据这些指标与抗寒性的关系及随土壤水分的变化情况,最终得出入冬前土壤绝对含水量为35%-40%时,更利于冬小麦的越冬生长。
The trial for the effects of the soil moisture on the overwintering survival rates of winter wheat was carried out in Harbin,Heilongjiang prvince,China,which located in the frigid region from 2009 to 2010.Two winter wheat cultivars different in the cold hardiness were used in this trial.Different soil water content gradients were set up based on the relation between the soil water contents and the overwinter survival rate of winter wheat.The photosynthetic preperties,physiological and biochemical indices and proteomics and so on in winter wheat plants during winter season were studied,and furthermore,the optical soil water content for the safe overwintering of winter wheat was definited,the indices and methods in the identification for the cold tolerance of wheat were explored and the mechanism in the safe overwintering of winter wheat in frigid region was revealed through the influence of proteins expression,and the theoretic fundation was provided for the production of winter wheat in the frigid area.The main conclusions were as follows:
1.On the overwintering cases none of Jimai 22 but all of Dongnongdongmai 1 was able to be overwinterin in the all water treatments.It was very significant in the difference in the overwinter survival rate of Dongnongdongmai 1 among the soil water treatments,appearing CK(normal soil water management)>30%±2%>45%±2%>15%±2% in the soil moisture,and was 83.3%,49.2%,23.3%,2.5%,in the survival rate respectively.The ratio of the dead cells in the leaf and leaf sheath of the two cultivars was gradually enhanced with the delay of the sampling date.And the obvious difference in the cell vigor between cultivars occurred at the 20th day after frozen date in winter and the injury in Dongnongdongmai 1 was less than that in Jimai 22.The damage of plants under CK (normal soil water management) in soil water content were slightest.The extent of damage in the leaf was more serious than that in the leaf sheath.The change of the development status of the stem apical point was very slow and less affected by the soil water content after entering the frozen period and the profile of the apical point almost was in the elongation stage.On the contrary/the transition of the apical point in Jimai 22 was fast and quicker in the soil water treatment of 45%±2% and the form of the part was in the elongation and single ridge.
2.The water content in the leaf and leaf sheath of the 2 cultivars performed downward and then upward as the sampling dates increased and lower in 15%±2% of soil water treatment. This trait in Jimai 22 was higher than that in Dongnongdongmai 1 and the one in leaf sheath higher than that in leaf.The free water content in samples was lower in 15%±2% and higher in 45%±2% in the soil water content treatment.The bound water content in the leaf sheath of Dongnongdongmai 1 remained higher and that of Jimai22 performed bimodal curve in the whole sampling period.
3.The soluble protein content of the 2 cultivars increased in all soil water theatments and would be facilited to a higher level both under drought and hyperhydration condition during the sampling period.No significant difference between the 2 cultivars was observed in this trait.
4.The relative electric conductivity in the leafs and leaf sheaths of the 2 cultivars elevated gradually during the sampling period and quickly in the early days and smoothly in the late days and lowest in CK (normal soil water management) and highest in 15%±2% of soil water treatment.The conductivity of Dongnongdongmai 1 was lower than that of Jimai22 and one in the leaf sheath higher than that in leaf.MDA of both cultivars appeared downward in the early days and upward in the late days in the sampling period.The content of MDA in the cultivars was higher relatively in 15%±2% and lower in CK(normal soil water management) and media both in 30%±2% and 45%±2% in the soil water treatment.The level of membrane lipid overoxidation was found to be worsened both in dought and overmoisture.
5.The SOD activity of cultivars behaved in fluctuation during the sampling period.The order in the trait of Dongnongdongmai 1 among the soil water treatments showed as 15%±2%>30%±2%>45%±2%>CK (normal soil water management) and of Jimai 22 the highest in 15%±2%.And the POD activity appeared enhancing trands during the same period and higher in Dongnongdongmai 1 than in Jimai 22.The higher POD activity of Dongnongdongmai 1 presented in 15%±2% of soil water content in cold acclimation (?) in 45%±2% after frozen date and of Jimai 22 in 15%±2% covering winter,respectively.The GSH contents of 2 cultivars were downward during the sampling period and higher in Dongnongdongmai 1 than in Jimai 22 and of Dongnongdongmai 1 in 30%±2% while of Jimai 22 in 45%±2%..The As A contents of 2 cultivars showed upward in the early stage of the sampling period and then downward in the late stage.The lower value of As A was in CK(normal soil water management) of soil water content during cold acclimation phase. After entering frozen period the higher content of AsA was found in CK of soil water content in Dongnongdongmai 1,but the values of AsA in Jimai 22 waved remarkably in all soil water treatments.
6.The change of the initial fluorescence values (Fo) of both cultivars in the soil water treatments presented the bimodal curv in the whole sampling period.The Fo value would increase under the dought and water logging conditions.The Maximum fluorescence value (Fm) of the 2 cultivars enhanced rapidly in all soil water treatments before the frozen date and decreased fast after the stage.After 20 days entering the frozen stage the change of Fm was smooth while showed highest in CK (normal soil water management).The maximal quantum yield (Fv/Fm) and the actual quantum yield (Yield) of Dongnongdongmai 1 appeared to be smooth or a slightly increasing trend in all soil water treatments from the cold acclimation period to the frozen stage and of Jimai 22 downward.The Fv/Fm and Yield of the cultivars reduced gradually and those values in CK (normal soil water management) treatment were higher obviously than those in other soil water contents after entering into the frozen stage.The photochemical quenching parameter (qP) of Dongnongdongmai 1 in all soil water treatments dropped gradually during the sampling period and of Jiami 22 fluctuated downward.The non-photochemical quenching parameter (qN) of both cultivars appeared to be fluctuating upward.The difference in the actual fluorescence yield (Ft) of both cultivars was very significant among the sampling dates.The value of Ft could be induced to incease in the cold acclimation period but to reduce in the early stage of frozen perion by slightly drought.The photosynthetic electron transfering rate (ETR) of both cultivars decreased gradually in the whole sampling period and sharply in the frozen period and was higher under the optical and some more soil water contents than that under drought condition. The photoinhibition in Dongnongdongmai 1 was heavier than that in Jiami 22 in the cold acclimation period and more seriously in Jimai 22 in the frozen period.The transient maximal quantum yield (Fm) of Dongnongdongmai 1 was higher than that of Jimai 22 and the significant difference in Fm between cultivars was observed in all actinic light treatments.The Fm in CK (normal soil water management) was lower in the early stage of the frozen period and became higher after the date of Dec 2.The change of Fm with the actinic light intensives of the cultivars was fluctuating in the cold acclimation period and the early stage of the frozen period and became smooth in the whole frozen period.
7.The number of the different protein spots was found to be 32 in the early and late stages of the frozen period in the soil water contents of 15%±2%,30%±2% and 45%±2% in both cultivars.Of which 26 spots matched highly with proteins found in wheat and 3 with in rice,1 with in maize,1 with in barley and 1 with in cucumber.12.5% of the protein spots participated in the process of energy metablism,and 25% of material synthesis,9.3% belonged to molecular chapron,31.25% to resistant proteins,6.25% of photosynthetic reaction and 15.62% to other proteins.There were 9 protein spots appearing both in the cold tolerant cultivar and cold sensitive cultivar in the early stage of the frozen period and all participated in the process of the osmotic matters synthesis,scavenging of the active matters damaging cell.stress resistance and photosynthesis.The different protein spots participating energy metablism in the cold tolerant cultivar were more than that in the cold sensitive cultivar. There were no different protein spots both expressing in the cold-tolerant and -sensitive cultivars in the late stage of the frozen period.The different expressing protein spots jioned thicking cell wall,energy metablism and stress resistance in the cold tolerant cultivar and in the cold sensitive cultivar participated dissolving cell membrane and photosynthesis.The more of the different expression protein spots occurred in the soil water treatments of dought (15%±2%) and more moisture (45%±2%) and less of slight drought (30%±2%).
8.The strong cold resistant cultivar has characterestics as follows:slight damage of tissues in the frozen period,more alive cells,low progress of apical point development,low level of water content in leaf sheathes in the overwinter period,high activity of SOD,POD in the cold acclimation period and the early stage of the frozen period,high content of GSH in the overwinter period,high in qP,qN in the cold acclimation period,high in Fv/Fm,Yield after the date of frozen,low in Fo after 20 days entering into the frozen period,high in Fm in the whole frozen period.The evaluation for the cold tolerance of winter wheat in the frigid region could be carried into execution by the indices of paints vigor,apical point status,relative conductivity,MDA,chlorophyll fluorescence parameters (Fo,Fm,Fv/Fm,Yield,qP,qN),activity of SOD,POD and GSH content.
9.The close relationship was found between the cold tolerance of winter wheat and the indices of the overwinter survival rate in spring,plant vigor,conductivity and MDAin winter,the maximal fluorescence and transient maximal quantum yield after 20 days entering into the frozen period,maximal quantum yield and actual quantum yield in the frozen period,ETR in winter. All of the indices were affected obviously by the soil water content.The soil water content of 35%-40% before winter season was in favor of the safe overwintering of winter wheat according to the relationship between the indices and the cold tolerance of winter wheat and its changes with the soil water content levels.
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