盐芥重要生长发育期的生理生态机制研究
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摘要
盐生植物在其整个生长发育内会面临许多外界环境变化，如：土壤盐度和季节更替带来的光照、温度和水分的变化。盐生植物能否在重要生长发育期（如：种子萌发和开花繁殖期）适应这些外部环境的变化，直接决定着其种群生存和分布。本文针对盐生植物模式材料盐芥在重要生长发育期对外界环境的生理生态适应性及机制进行研究，旨在探究盐芥在重要生长发育对外界环境的适应机制，为阐明其分布和生存机理提供依据。主要研究结果如下：
在盐胁迫条件下种子能否萌发及高盐条件不能萌发的种子在足够降雨后能否萌发直接决定该物种能否在盐碱地形成群落及完成生活史。为此，我们通过与拟南芥的对比实验，用不同浓度盐胁迫对盐芥种子进行处理，探讨盐芥种子萌发期和幼苗生长期的耐盐机制。实验采用培养皿法，盐芥和拟南芥种子分别用0、30、60、90、120、200、300、400和500mmol L~(-1)NaCl处理液处理，处理后分别春化10天，取出观察其种子萌发情况，萌发的幼苗7天后测定生理指标。结果表明，NaCl处理对盐芥和拟南芥种子萌发均有一定的抑制作用。拟南芥和盐芥在萌发的最初两天在各个NaCl浓度的胁迫下，萌发率都在缓慢增加，3天后萌发趋于稳定。30mmol L~(-1)NaCl对拟南芥的种子萌发没有影响。在较高盐胁迫浓度，拟南芥和盐芥的萌发受到显著抑制，盐芥种子萌发受抑程度强于拟南芥。30、60、90和120mmol L~(-1)NaCl胁迫7天时，拟南芥种子的萌发率分别为对照的100%、99%、77%和28%，盐芥的种子萌发率分别为对照的91%、68%、26%和0%。当NaCl浓度超过200mmol/L，盐芥和拟南芥种子的萌发率均为0。但是，在高浓度NaCl处理抑制的盐芥种子在盐胁迫解除后比拟南芥种子表现出较好的萌发，与对照相比，在200、300、400、500mmol L~(-1)NaCl胁迫下盐芥种子萌发恢复率分别为105%、104%、106%和99%，但拟南芥在相同浓度的NaCl胁迫下种子萌发恢复率分别为94%、84%、82%、54%。可见盐胁迫促进了盐芥种子在盐胁迫解除后的萌发能力，而拟南芥在高盐胁迫下，种子萌发恢复率低说明发生了拟南芥种子在高盐胁迫下受到了一定程度的离子毒害。
在幼苗生长阶段，盐芥的幼苗比拟南芥表现出较好的耐盐性，幼苗长势好，根长、干鲜重、色素含量、脯氨酸含量和K+/Na+均高于拟南芥，MDA积累少。这些结果表明：盐芥在种子萌发和幼苗生长期对盐胁迫表现出了较强的适应机制，这是盐芥能够在盐碱地存活并形成群落的原因之一。
耐逆模式植物-盐芥是十字花科植物，自然条件下在秋天萌发，以莲座状幼苗度过冬季并完成春化作用，春天接受长日照完成光周期诱导并开花结果。但是，对于盐芥春化作用所需天数及个体发育阶段及光周期诱导参数了解很少，这极大地限制了对盐芥耐逆分子机理的研究。本实验对山东生态型盐芥种子春化和光周期进行了系统研究，旨在缩短盐芥春化和开花的时间，获得实验室条件完成生活史的最短时间，以方便对盐芥耐逆机理的研究。种子春化采用培养皿恒定4℃冰箱处理法，将准备好的盐芥种子，在实验室充分吸水后，置于4℃冰箱中，使种子春化的时间分别为：0、2、4、6、8、10、12、14、16、18、20、22、24、26、28、30天，处理结束后，将春化后的种子置于拟南芥培养箱中培养观察种子萌发情况，7天后将萌发幼苗移栽于盆中培养观察其开花情况。结果表明，盐芥种子春化促进了盐芥种子的萌发，随着种子春化时间的延长，盐芥种子萌发率逐渐升高，种子春化30天，盐芥种子萌发率为90%。萌发的种子定植于盆观察到盐芥种子春化后，可以开花，说明盐芥种子具有春化的能力。盐芥种子春化时间小于6天时，盐芥不开花，一直处于营养生殖状态。盐芥种子春化时间6-22天，盐芥开花植株数随着春化时间的增加呈S曲线上升，盐芥种子春化时间22-40天，盐芥开花植株数恒为100%。开花盐芥植株第一朵花开放时间被盐芥种子春化时间缩短，盐芥种子春化时间越长，盐芥植株第一朵花开时间越短。盐芥种子春化时间30-40天，盐芥植株第一朵花开放时间在春化结束后的第29天。种子经过30天的春化处理，开花最理想。种子春化能够促进盐芥开花，与其抑制ThFLC（开花抑制基因）的表达有关。盐芥种子未经春化，ThFLC的表达量为100%，随着种子春化时间的延长，盐芥相应植株内ThFLC的表达量逐渐减少，当盐芥种子春化30天时，ThFLC的表达量为24.3%。经过30天种子春化处理的盐芥植株在拟南芥培养箱的恒定温度、湿度和光照下，分别进行7、8、9、10、11、12、13、14小时的光照长度处理，结果表明：经过种子春化的盐芥盐芥种子经过30天春化长出植株在光周期为9到14小时，能够开花。如果光周期小于8小时，盐芥则不开花。随着光周期的延长，盐芥开花的时间缩短，开花数量和单株干鲜重增加。在14小时的光周期下，盐芥植株开花数量和单株干鲜重达到最大值。这些结果表明盐芥为典型的长日照植物，至少需要9小时的光周期才能开花。这些结果极大地方便了人们对盐芥分子机理的研究。其次，这些结果还表明盐芥对春化和对光周期的要求，与其对生存环境的适应有关。
山东生态型盐芥在黄河三角洲盐碱地在秋天萌发，以莲座状幼苗度过冬季，三月份开始返青生长、抽薹开花，五月份种子成熟。这个季节正是降雨少、温度回升快及返盐高峰期。盐芥为什么在这一时期完成其生活史，其生理生态机理是什么？值得深入研究。我们通过2012年一月至五月对在黄河三角洲盐碱地自然生长盐芥植株内的离子含量和某些生理生态指标变化的研究，揭示盐芥耐盐的生理生态基础。结果表明，土壤中Na~+、Ca~(2+)和Cl~-含量从2012年一月至五月呈逐渐上升趋势，与气温的上升正相关，与土壤水分含量呈负相关。说明气温和土壤含水量是影响土壤中Na+、Ca2+和Cl-浓度的主要原因。在气温较低的一月和二月盐芥地上部分Na+离子含量最高，随着植物的生长发育，整体呈下降趋势，而K~+、Ca~(2+)、Mg~(2+)、Cl~-、SO_4~(2-)则呈上升趋势，尤其是Cl~-上升更为明显。盐芥植株内这些离子的累积与其生长发育阶段有很大关系，离子的累积也有利于其进行渗透调节作用。在生长发育中后期根部呈下降趋势，可能是由于根部向地上部输送较多的营养物质。盐芥的生理生态指标变化表明，盐芥的生长受低温的影响要大于受土壤盐度变化的影响。在气温较低的一月和二月，盐芥表现出一定的受害症状，MDA累积，叶绿素含量低，但是其植株内较高的脯氨酸和可溶性糖含量使盐芥在低温条件下免受伤害，起到一定的渗透调节和保护作用。这些结果表明，盐芥植株组织中可溶性物质含量与春天温度有关，是其适应低温和高盐的机制之一。
人们对盐芥耐盐性进行了深入研究，可以在200mmolL~(-1)NaCl处理条件完成生活史，其耐盐机理主要是把过多的盐区域化到液泡中，这种特性与重金属超富集植物有类似的机理，而黄河三角洲土壤近年来重金属污染日趋严重，作为自然生长该地区盐碱地的植物是否具有重金属超富集特性及耐重金属机理值得研究，这些结果对于利用盐芥对遭受重金属污染的盐碱地进行修复具有重要意义。我们通过盆栽试验，比较分析了盐芥在Cd、Cu和Pb重金属处理下生长等生理生态特性。结果表明，随着Cd、Cu和Pb处理浓度的升高，盐芥的根长及干物质重均显著下降，MDA均显著升高。与Cu和Pb相比， Cd处理浓度较低，但受害症状明显，具体表现为：高浓度下，叶绿素含量显著下降，叶片黄化，MDA累积多，幼苗死亡率高等。Pb处理浓度最高，但在这三种重金属中Pb处理的受害症状表现不明显，说明盐芥对Pb具有一定的耐性。从盐芥对三种重金属的累积量来看，盐芥对Cd具有一定的累积作用，但在高浓度(130mg kg~(-1))下很难存活，在Cu和Pb胁迫下能够完成正常的生活周期，但对Cu和Pb并不累积。所以盐芥对Cd污染土壤的修复具有一定的潜力，但不适宜用于Cu和Pb污染土壤的修复。
本论文主要创新点：
1通过与拟南芥的比较研究发现，虽然盐芥种子萌发受到高浓度盐的强烈抑制，但能够在盐胁迫缓解时迅速且整齐萌发，甚至超过无盐胁迫时的萌发率。在幼苗生长期，盐芥幼苗在盐胁迫下，能够维持较高的叶绿素含量，K~+/Na~+和脯氨酸含量，累积较少的MDA，表现出良好的生长势。这些结果表明盐芥在种子萌发和幼苗生长期比拟南芥表现出较好的对盐的适应性。这可能是盐芥能够在盐碱地存活并形成群落的原因之一。
2本文首次研究了种子春化和光周期对盐芥开花的影响。研究结果表明种子春化促进了盐芥种子的萌发，种子春化后盐芥能够开花说明盐芥具有种子春化的能力。种子春化低于6天盐芥不能开花，一直处于营养生长阶段，种子春化22-40天，盐芥开花率可达100%。种子春化缩短了植株的开花时间，降低了开花抑制基因ThFLC的表达。山东生态型盐芥的开花依赖于春化作用，对春化作用的需求是必须的，而不是兼性的。山东生态型盐芥种子经过充分的春化后，只有在光照时间大于9小时的光周期下才能开花。而且随着光照时间的延长，盐芥开花数目增加，植株干重增大，开花时间缩短。当光周期为14小时，盐芥植株的干重和开花数达到最大值，开花时间最短。这些结果表明盐芥为典型的长日照植物，其临界日长为9小时。盐芥对春化和对光周期的要求，与其对生存环境的适应有关。对盐芥种子春化和光周期的研究大大方便了今后对盐芥耐逆分子机理的研究。
3自然生境下盐芥植株内离子含量、可溶性有机物、MDA和叶绿素含量的变化与生长阶段、土壤中的离子含量、降水量等外界环境变化有关。通过对盐芥生理生态的季节性研究结果表明：对盐芥生理生态起主要作用的是植物本身的生长发育阶段，其次是环境温度及土壤盐度等环境因子。盐芥在黄河三角洲盐碱地低温、干旱和盐度的逆境中通过生理生态的适应来得以生存并形成群落。
4在土壤Cd、Cu和Pb污染环境下，盐芥对Cd、Cu和Pb都有吸收，其中盐芥对Cd具有一定的累积作用，但在高浓度下(130mg kg~(-1))盐芥很难存活。盐芥在Cu和Pb胁迫下能够完成正常的生活周期，说明盐芥对Cu和Pb具有一定的耐性但对Cu和Pb并不累积。所以盐芥对Cd污染土壤的修复具有一定的潜力，但不适宜用于Cu和Pb污染土壤的修复。
Halophytes face many abiotic stressors such as salinity, light,temperature and water content caused by seasonal alteration during thewhole life span. Whether halophyte could adapt the changes at the keystages such as seed germination and flowering determined the surviral anddistribution of the species population. In this paper the eco-physiologicaladaptations of Thellungiella halphila to salinity, temperatures and seasonalchanges were studied with the aim to illustrate the adaptive mechanism andcertainty of its survival and distribution. The main results are as follows:
Salinity is one of the major abiotic stresses that adversely affect plantgrowth and development. The occurrence of halophytes in inland saltmarshes may depend on their tolerance to salt stress at different stages ofplant development. Germination is a key stage in the life cycle of plants insaline environments as it determines whether or not plants can establishsuccessfully in certain areas. So, in the first experiment, effects of salinityon germination and seedling growth of T. salsuginea and Arabdopsisthaliana were compared with the aim to explore the salinity-resistantmechanisms of T. salsuginea. The seeds of T. salsuginea and A. thalianawere treated with0(control),30,60,90,120,200,300,400and500mmol L~(-1)NaCl in discs after10days’ seed vernalization. Then the seedgermination and physiology indexes of seedlings grown for7days weredetermined.The results showed that salinity inhibited seed germination inboth species.The germination rate of both A. thaliana and T. salsugineaincreased gradually by all concentrations of NaCl in the initial2days,germination rate kept consitent in some extent after3days.30mmolL~(-1)NaCl treatments had no effect on seed germination rate of A. thaliana,but higher concentrations of NaCl markedly decreased seed germinationrate in both species. At30,60,90and120mmol L~(-1)NaCl, the germinationpercentage of A. thaliana was100%,99%,77%and28%of the control,however the germination percentage of T. salsuginea was91%、68%、26%and0%of the control. Seeds of both species did not germinate when theconcentration of NaCl was over200mmol L~(-1). Surprisingly, theungerminated seeds of both species quickly germinated. Moreover, the totalseed germination rate recovering from200,300,400and500mmol L~(-1)NaCl in T. salsuginea was105%,104%,106%and99%, but in A.thaliana was94%、84%、82%、54%, respectively. Therefore, salinitypromotes the seed germination of T. salsuginea during the water recoverystage. The ungerminated seeds of A. thaliana suffered ion toxicity in highsalinity to some degree as evidenced by the low final germination rate afterungerminated seeds pretreated with NaCl were transferred to distilledwater.
At the seedling growth stage, seedlings of T. salsuginea were more salttolerant than those of A. thaliana. For example, seedlings of T. salsugineahad better plant growth (root length, fresh and dry weight), higherchlorophyll content, less MDA content and higher proline content and K+/Na+ratio under salinity. These characteristics indicate that T. salsuginea ismore salt-tolerant to salinity than A. thaliana at both seed germination andseedling stages.
Generally, salt cress, Thellungiella salsuginea (Cruciferae), germinatesin autumn, is vernalized at rosette seedling in winter and flowers afterlong-day irradiance in spring in its natural habitat. But little was knowabout how many days did T. salsuginea finish its vernalization andphotoperiod requirement, which restricts the insight into the molecularmechanisms of abiotic stresses sach as salt tolerance in labotory. Tofacilitate research on T. salsuginea, the present study focuses on seedvernalization and photoperiod for flowering induction. Seeds of T.salsuginea were vernalized at4°C in discs for0,2,4,6,8,10,12,14,16,18,20,22,24,26,28and30days respectively. After vernalization, seedswere placed in a growth chamber to observe the seed germination. After7days’germination, the seedling were planted in the pots in the samechamber to observe the flowering.The results showed that seedvernalization promoted the seed germination. Seed germination rateincreased with the prolonged duration of vernalization. The seed germination rate was90%of the control when the seeds were vernalizedfor30days and flowered after the the seedlings grown under suitable lightperiod. These results suggested that T. salsuginea can finish itsvernalization at stage of imbibed seeds. T. salsuginea still stayed atvegetative stage and could not flower when seeds were vernalized for lessthan6days. The flowering plants increased when seeds were vernalized for6-22days. And100%of T. salsuginea flowered when seeds werevernalized for22-40days. The time when the first flower appeared wasshortened by seed vernalization. Longer seed vernalizaiton, earlier the firstflower emergence. The earliest emergence of the first flower was the29thday after seeds were vernalized for30-40days. Expression of ThFLC,which is a repressor of flowering, was reduced by vernalization. Relativeexpression of ThFLC dropped from100%of control to24.3%of30d ofseed vernalization. Plants growing from seeds that had been vernalized for30d did not flower when day length was <9h, and day lengths>9hpromoted flowering. These results will facilitate the utilization of T.salsuginea as a model plant of abiotic stress to study the molecularmechanisms. The results also suggest that the dependency on vernalizationand long-day photoperiod is critical for the adaptation of T. salsuginea tothe local environment.
T. salsuginea (Shangdong ecotype) germinates in autumn, vernalized asrosette seedlin in winter, flowers in March and ripen in May in the Yellow River Delta. This season is characteristic of low rainfall, quick temperaturerise and high salinity in the soil.Why does T. salsuginea complete its lifespan in this season and what is the adaptive mechanism? These questionsdeserve further research. In the third expreripent, seasonal ion content andeco-physiological characters variation in T. salsuginea in its habitat, YellowRiver delta saline and alkaline land, from January to May in2012were alsostudied. The results showed that Na~+, Ca~(2+) and Cl~-content in the soilincreased from January to May, and was closely related to temperature andwater content in the soil. Na+content in T. salsuginea was highest inJanuary and February and decreased with time. K~+、Ca~(2+)、Mg~(2+)、Cl~-、SO_4~(2-)content showed increasing tendency, in particular, Cl-content, with theincrease of temperature and the growth stage. The accumulation of K~+、Ca~(2+)、Mg~(2+)、Cl~-、SO_4~(2-) was beneficial to osmaregulation in T. salsugineaunder salinity. K~+, Mg~2+and Ca~(2+) content in the root were lower than that inthe shoot because nutritional ion was in priority distribution to leaves,flowers and pods from roots. The eco-physiological variation in T.salsuginea showed that the effect of temperature was larger than that of ioncontent changes in the soil. T. salsuginea showed retarded growth withhigher accumulation of MDA, lower chlorophyte content. But T. salsugineaadjusted eco-physiological characters to the low temperature such as higherproline and soluble sugar content which protected T.salsuginea from theharm of low temperature and ion variation. These results suggested that compatible solute content is closely related to temperature, which is one ofthe adaptive mechanisms of T. salsuginea to low temperature and highsalinity.
Based on the research of T. salsuginea, T. salsuginea could survive in200mmol L~(-1)NaCl. The main mechanism was compartmentalization ofexcessive salt ion into vacuoles, which is similar to heavy metalsuperaccumulator plant. Heavy metal pollution in Yellow River delta soil isgetting worse; T. salsuginea might be a potential heavy metalsuperaccumulator, which needs to study. In the fourth experiment, a potexperiment was used to investigate growth responses and eco-physiologicalcharacters of T. salsuginea under Cd, Cu and Pb treatments. The resultshowed that the root length and dry mass decreased and MDA accumulatedsignificantly with the increase of Cd, Cu and Pb concentrations. T.salsuginea showed apparent damage symptoms such as lower chlorophyllcontent, etiolated leaves, higher MDA accumulations and death ofseedlings under Cd treatment compared with Cu and Pb. T. salsuginea cangrow well under Pb treatment though Pb treatment concentration was thehighest. From the point of metal accumulation and BAF, T. salsugineaabsorbed Cd>Cu>Pb. These results indicate that T. salsuginea has certainpotential on remedying the soil polluted by Cd with low and middleconcentrations.
The innovations of this thesis were shown as follows:
1. In this study, the ungerminated seeds of T. salsuginea under highsalinity quickly germinated though the seed germination of T. salsugineawas a little more sensitive to salinity compared with A. thaliana. At theseedling stage, however, T. salsuginea showed better growth with higherchlorophyll content, less MDA content and higher proline content and K~+/Na~+ratio under salinity. These characteristics indicate that T. salsuginea ismore adaptive to salinity than A. thaliana at both seed germination andseedling stages and explain why A. thaliana is excuded from salinelocations and T. salsuginea can survive and form its populations in salinesoils.
2. The seed vernalization and photoperiod of T. salsuginea were studiedfor the first time. The results showed that seed vernalization promoted theseed germination and that T. salsuginea flowered after seed vernalizationwhich showed that T. salsuginea is capable of vernalization at stage ofimbibed seeds. T. salsuginea still stayed at vegetative stage and couldn’tflower when seeds were vernalized less than6days. And100%of T.salsuginea flowered when seeds were vernalized for22-40days. The timewhen the first flower appeared was shortened by seed vernalization.Expression of ThFLC, which is a repressor of flowering, was markedlyreduced by vernalization. The requirement for vernalization of T. salsuginea is obligate instead of facultative. We also observed here that T.salsuginea generated flowers only when day length exceeded9h. Thenumber of flowers and dry mass per plant also increased with an increaseof day length. Dry mass per plant and number of flowers per plant were thehighest when day length exceeded14h. These results therefore confirmthat T. salsuginea is a long-day plant with a critical day length requirementof approximately9h. In addition, the time required to produce the firstflower decreased as day length exceeded9h. These results indicated that T.salsuginea is a typical long-day plant with9h photoperiod. Therequirement of vernalization and long days by T. salsuginea (Shandongecotype) is at least partially the product of adaptation to the localenvironment. These results facilitate greatly the future research onsalt-tolerant mechanisms of T. salsuginea.
3. Soluble ion, proline, MDA and chlorophyll content of T. salsugineawere closely related to growth and development stages, ion content in thesoil and rainfall. For T. salsuginea, the effect of growth and developmentstages on the adaptation to stress environment was larger than that oftemperature and ion content changes in the soil. Under the natural location,T. salsuginea adjusted eco-physiological characters to low temperature,drought and salinity caused by the seasonal variances.
4. T. salsuginea accumulated Cd to some degree, but did not survive inhigher concentration of Cd (130mg kg~(-1)). T. salsuginea finished its life cycle under Cu and Pb stress. Therefore, T. salsuginea was tolerant to Cuand Pb but not accumulated them. Conclusively, T. salsuginea can be usedin remedying the soil polluted by Cd with low and middle concentrations.
在盐胁迫条件下种子能否萌发及高盐条件不能萌发的种子在足够降雨后能否萌发直接决定该物种能否在盐碱地形成群落及完成生活史。为此，我们通过与拟南芥的对比实验，用不同浓度盐胁迫对盐芥种子进行处理，探讨盐芥种子萌发期和幼苗生长期的耐盐机制。实验采用培养皿法，盐芥和拟南芥种子分别用0、30、60、90、120、200、300、400和500mmol L~(-1)NaCl处理液处理，处理后分别春化10天，取出观察其种子萌发情况，萌发的幼苗7天后测定生理指标。结果表明，NaCl处理对盐芥和拟南芥种子萌发均有一定的抑制作用。拟南芥和盐芥在萌发的最初两天在各个NaCl浓度的胁迫下，萌发率都在缓慢增加，3天后萌发趋于稳定。30mmol L~(-1)NaCl对拟南芥的种子萌发没有影响。在较高盐胁迫浓度，拟南芥和盐芥的萌发受到显著抑制，盐芥种子萌发受抑程度强于拟南芥。30、60、90和120mmol L~(-1)NaCl胁迫7天时，拟南芥种子的萌发率分别为对照的100%、99%、77%和28%，盐芥的种子萌发率分别为对照的91%、68%、26%和0%。当NaCl浓度超过200mmol/L，盐芥和拟南芥种子的萌发率均为0。但是，在高浓度NaCl处理抑制的盐芥种子在盐胁迫解除后比拟南芥种子表现出较好的萌发，与对照相比，在200、300、400、500mmol L~(-1)NaCl胁迫下盐芥种子萌发恢复率分别为105%、104%、106%和99%，但拟南芥在相同浓度的NaCl胁迫下种子萌发恢复率分别为94%、84%、82%、54%。可见盐胁迫促进了盐芥种子在盐胁迫解除后的萌发能力，而拟南芥在高盐胁迫下，种子萌发恢复率低说明发生了拟南芥种子在高盐胁迫下受到了一定程度的离子毒害。
在幼苗生长阶段，盐芥的幼苗比拟南芥表现出较好的耐盐性，幼苗长势好，根长、干鲜重、色素含量、脯氨酸含量和K+/Na+均高于拟南芥，MDA积累少。这些结果表明：盐芥在种子萌发和幼苗生长期对盐胁迫表现出了较强的适应机制，这是盐芥能够在盐碱地存活并形成群落的原因之一。
耐逆模式植物-盐芥是十字花科植物，自然条件下在秋天萌发，以莲座状幼苗度过冬季并完成春化作用，春天接受长日照完成光周期诱导并开花结果。但是，对于盐芥春化作用所需天数及个体发育阶段及光周期诱导参数了解很少，这极大地限制了对盐芥耐逆分子机理的研究。本实验对山东生态型盐芥种子春化和光周期进行了系统研究，旨在缩短盐芥春化和开花的时间，获得实验室条件完成生活史的最短时间，以方便对盐芥耐逆机理的研究。种子春化采用培养皿恒定4℃冰箱处理法，将准备好的盐芥种子，在实验室充分吸水后，置于4℃冰箱中，使种子春化的时间分别为：0、2、4、6、8、10、12、14、16、18、20、22、24、26、28、30天，处理结束后，将春化后的种子置于拟南芥培养箱中培养观察种子萌发情况，7天后将萌发幼苗移栽于盆中培养观察其开花情况。结果表明，盐芥种子春化促进了盐芥种子的萌发，随着种子春化时间的延长，盐芥种子萌发率逐渐升高，种子春化30天，盐芥种子萌发率为90%。萌发的种子定植于盆观察到盐芥种子春化后，可以开花，说明盐芥种子具有春化的能力。盐芥种子春化时间小于6天时，盐芥不开花，一直处于营养生殖状态。盐芥种子春化时间6-22天，盐芥开花植株数随着春化时间的增加呈S曲线上升，盐芥种子春化时间22-40天，盐芥开花植株数恒为100%。开花盐芥植株第一朵花开放时间被盐芥种子春化时间缩短，盐芥种子春化时间越长，盐芥植株第一朵花开时间越短。盐芥种子春化时间30-40天，盐芥植株第一朵花开放时间在春化结束后的第29天。种子经过30天的春化处理，开花最理想。种子春化能够促进盐芥开花，与其抑制ThFLC（开花抑制基因）的表达有关。盐芥种子未经春化，ThFLC的表达量为100%，随着种子春化时间的延长，盐芥相应植株内ThFLC的表达量逐渐减少，当盐芥种子春化30天时，ThFLC的表达量为24.3%。经过30天种子春化处理的盐芥植株在拟南芥培养箱的恒定温度、湿度和光照下，分别进行7、8、9、10、11、12、13、14小时的光照长度处理，结果表明：经过种子春化的盐芥盐芥种子经过30天春化长出植株在光周期为9到14小时，能够开花。如果光周期小于8小时，盐芥则不开花。随着光周期的延长，盐芥开花的时间缩短，开花数量和单株干鲜重增加。在14小时的光周期下，盐芥植株开花数量和单株干鲜重达到最大值。这些结果表明盐芥为典型的长日照植物，至少需要9小时的光周期才能开花。这些结果极大地方便了人们对盐芥分子机理的研究。其次，这些结果还表明盐芥对春化和对光周期的要求，与其对生存环境的适应有关。
山东生态型盐芥在黄河三角洲盐碱地在秋天萌发，以莲座状幼苗度过冬季，三月份开始返青生长、抽薹开花，五月份种子成熟。这个季节正是降雨少、温度回升快及返盐高峰期。盐芥为什么在这一时期完成其生活史，其生理生态机理是什么？值得深入研究。我们通过2012年一月至五月对在黄河三角洲盐碱地自然生长盐芥植株内的离子含量和某些生理生态指标变化的研究，揭示盐芥耐盐的生理生态基础。结果表明，土壤中Na~+、Ca~(2+)和Cl~-含量从2012年一月至五月呈逐渐上升趋势，与气温的上升正相关，与土壤水分含量呈负相关。说明气温和土壤含水量是影响土壤中Na+、Ca2+和Cl-浓度的主要原因。在气温较低的一月和二月盐芥地上部分Na+离子含量最高，随着植物的生长发育，整体呈下降趋势，而K~+、Ca~(2+)、Mg~(2+)、Cl~-、SO_4~(2-)则呈上升趋势，尤其是Cl~-上升更为明显。盐芥植株内这些离子的累积与其生长发育阶段有很大关系，离子的累积也有利于其进行渗透调节作用。在生长发育中后期根部呈下降趋势，可能是由于根部向地上部输送较多的营养物质。盐芥的生理生态指标变化表明，盐芥的生长受低温的影响要大于受土壤盐度变化的影响。在气温较低的一月和二月，盐芥表现出一定的受害症状，MDA累积，叶绿素含量低，但是其植株内较高的脯氨酸和可溶性糖含量使盐芥在低温条件下免受伤害，起到一定的渗透调节和保护作用。这些结果表明，盐芥植株组织中可溶性物质含量与春天温度有关，是其适应低温和高盐的机制之一。
人们对盐芥耐盐性进行了深入研究，可以在200mmolL~(-1)NaCl处理条件完成生活史，其耐盐机理主要是把过多的盐区域化到液泡中，这种特性与重金属超富集植物有类似的机理，而黄河三角洲土壤近年来重金属污染日趋严重，作为自然生长该地区盐碱地的植物是否具有重金属超富集特性及耐重金属机理值得研究，这些结果对于利用盐芥对遭受重金属污染的盐碱地进行修复具有重要意义。我们通过盆栽试验，比较分析了盐芥在Cd、Cu和Pb重金属处理下生长等生理生态特性。结果表明，随着Cd、Cu和Pb处理浓度的升高，盐芥的根长及干物质重均显著下降，MDA均显著升高。与Cu和Pb相比， Cd处理浓度较低，但受害症状明显，具体表现为：高浓度下，叶绿素含量显著下降，叶片黄化，MDA累积多，幼苗死亡率高等。Pb处理浓度最高，但在这三种重金属中Pb处理的受害症状表现不明显，说明盐芥对Pb具有一定的耐性。从盐芥对三种重金属的累积量来看，盐芥对Cd具有一定的累积作用，但在高浓度(130mg kg~(-1))下很难存活，在Cu和Pb胁迫下能够完成正常的生活周期，但对Cu和Pb并不累积。所以盐芥对Cd污染土壤的修复具有一定的潜力，但不适宜用于Cu和Pb污染土壤的修复。
本论文主要创新点：
1通过与拟南芥的比较研究发现，虽然盐芥种子萌发受到高浓度盐的强烈抑制，但能够在盐胁迫缓解时迅速且整齐萌发，甚至超过无盐胁迫时的萌发率。在幼苗生长期，盐芥幼苗在盐胁迫下，能够维持较高的叶绿素含量，K~+/Na~+和脯氨酸含量，累积较少的MDA，表现出良好的生长势。这些结果表明盐芥在种子萌发和幼苗生长期比拟南芥表现出较好的对盐的适应性。这可能是盐芥能够在盐碱地存活并形成群落的原因之一。
2本文首次研究了种子春化和光周期对盐芥开花的影响。研究结果表明种子春化促进了盐芥种子的萌发，种子春化后盐芥能够开花说明盐芥具有种子春化的能力。种子春化低于6天盐芥不能开花，一直处于营养生长阶段，种子春化22-40天，盐芥开花率可达100%。种子春化缩短了植株的开花时间，降低了开花抑制基因ThFLC的表达。山东生态型盐芥的开花依赖于春化作用，对春化作用的需求是必须的，而不是兼性的。山东生态型盐芥种子经过充分的春化后，只有在光照时间大于9小时的光周期下才能开花。而且随着光照时间的延长，盐芥开花数目增加，植株干重增大，开花时间缩短。当光周期为14小时，盐芥植株的干重和开花数达到最大值，开花时间最短。这些结果表明盐芥为典型的长日照植物，其临界日长为9小时。盐芥对春化和对光周期的要求，与其对生存环境的适应有关。对盐芥种子春化和光周期的研究大大方便了今后对盐芥耐逆分子机理的研究。
3自然生境下盐芥植株内离子含量、可溶性有机物、MDA和叶绿素含量的变化与生长阶段、土壤中的离子含量、降水量等外界环境变化有关。通过对盐芥生理生态的季节性研究结果表明：对盐芥生理生态起主要作用的是植物本身的生长发育阶段，其次是环境温度及土壤盐度等环境因子。盐芥在黄河三角洲盐碱地低温、干旱和盐度的逆境中通过生理生态的适应来得以生存并形成群落。
4在土壤Cd、Cu和Pb污染环境下，盐芥对Cd、Cu和Pb都有吸收，其中盐芥对Cd具有一定的累积作用，但在高浓度下(130mg kg~(-1))盐芥很难存活。盐芥在Cu和Pb胁迫下能够完成正常的生活周期，说明盐芥对Cu和Pb具有一定的耐性但对Cu和Pb并不累积。所以盐芥对Cd污染土壤的修复具有一定的潜力，但不适宜用于Cu和Pb污染土壤的修复。
Halophytes face many abiotic stressors such as salinity, light,temperature and water content caused by seasonal alteration during thewhole life span. Whether halophyte could adapt the changes at the keystages such as seed germination and flowering determined the surviral anddistribution of the species population. In this paper the eco-physiologicaladaptations of Thellungiella halphila to salinity, temperatures and seasonalchanges were studied with the aim to illustrate the adaptive mechanism andcertainty of its survival and distribution. The main results are as follows:
Salinity is one of the major abiotic stresses that adversely affect plantgrowth and development. The occurrence of halophytes in inland saltmarshes may depend on their tolerance to salt stress at different stages ofplant development. Germination is a key stage in the life cycle of plants insaline environments as it determines whether or not plants can establishsuccessfully in certain areas. So, in the first experiment, effects of salinityon germination and seedling growth of T. salsuginea and Arabdopsisthaliana were compared with the aim to explore the salinity-resistantmechanisms of T. salsuginea. The seeds of T. salsuginea and A. thalianawere treated with0(control),30,60,90,120,200,300,400and500mmol L~(-1)NaCl in discs after10days’ seed vernalization. Then the seedgermination and physiology indexes of seedlings grown for7days weredetermined.The results showed that salinity inhibited seed germination inboth species.The germination rate of both A. thaliana and T. salsugineaincreased gradually by all concentrations of NaCl in the initial2days,germination rate kept consitent in some extent after3days.30mmolL~(-1)NaCl treatments had no effect on seed germination rate of A. thaliana,but higher concentrations of NaCl markedly decreased seed germinationrate in both species. At30,60,90and120mmol L~(-1)NaCl, the germinationpercentage of A. thaliana was100%,99%,77%and28%of the control,however the germination percentage of T. salsuginea was91%、68%、26%and0%of the control. Seeds of both species did not germinate when theconcentration of NaCl was over200mmol L~(-1). Surprisingly, theungerminated seeds of both species quickly germinated. Moreover, the totalseed germination rate recovering from200,300,400and500mmol L~(-1)NaCl in T. salsuginea was105%,104%,106%and99%, but in A.thaliana was94%、84%、82%、54%, respectively. Therefore, salinitypromotes the seed germination of T. salsuginea during the water recoverystage. The ungerminated seeds of A. thaliana suffered ion toxicity in highsalinity to some degree as evidenced by the low final germination rate afterungerminated seeds pretreated with NaCl were transferred to distilledwater.
At the seedling growth stage, seedlings of T. salsuginea were more salttolerant than those of A. thaliana. For example, seedlings of T. salsugineahad better plant growth (root length, fresh and dry weight), higherchlorophyll content, less MDA content and higher proline content and K+/Na+ratio under salinity. These characteristics indicate that T. salsuginea ismore salt-tolerant to salinity than A. thaliana at both seed germination andseedling stages.
Generally, salt cress, Thellungiella salsuginea (Cruciferae), germinatesin autumn, is vernalized at rosette seedling in winter and flowers afterlong-day irradiance in spring in its natural habitat. But little was knowabout how many days did T. salsuginea finish its vernalization andphotoperiod requirement, which restricts the insight into the molecularmechanisms of abiotic stresses sach as salt tolerance in labotory. Tofacilitate research on T. salsuginea, the present study focuses on seedvernalization and photoperiod for flowering induction. Seeds of T.salsuginea were vernalized at4°C in discs for0,2,4,6,8,10,12,14,16,18,20,22,24,26,28and30days respectively. After vernalization, seedswere placed in a growth chamber to observe the seed germination. After7days’germination, the seedling were planted in the pots in the samechamber to observe the flowering.The results showed that seedvernalization promoted the seed germination. Seed germination rateincreased with the prolonged duration of vernalization. The seed germination rate was90%of the control when the seeds were vernalizedfor30days and flowered after the the seedlings grown under suitable lightperiod. These results suggested that T. salsuginea can finish itsvernalization at stage of imbibed seeds. T. salsuginea still stayed atvegetative stage and could not flower when seeds were vernalized for lessthan6days. The flowering plants increased when seeds were vernalized for6-22days. And100%of T. salsuginea flowered when seeds werevernalized for22-40days. The time when the first flower appeared wasshortened by seed vernalization. Longer seed vernalizaiton, earlier the firstflower emergence. The earliest emergence of the first flower was the29thday after seeds were vernalized for30-40days. Expression of ThFLC,which is a repressor of flowering, was reduced by vernalization. Relativeexpression of ThFLC dropped from100%of control to24.3%of30d ofseed vernalization. Plants growing from seeds that had been vernalized for30d did not flower when day length was <9h, and day lengths>9hpromoted flowering. These results will facilitate the utilization of T.salsuginea as a model plant of abiotic stress to study the molecularmechanisms. The results also suggest that the dependency on vernalizationand long-day photoperiod is critical for the adaptation of T. salsuginea tothe local environment.
T. salsuginea (Shangdong ecotype) germinates in autumn, vernalized asrosette seedlin in winter, flowers in March and ripen in May in the Yellow River Delta. This season is characteristic of low rainfall, quick temperaturerise and high salinity in the soil.Why does T. salsuginea complete its lifespan in this season and what is the adaptive mechanism? These questionsdeserve further research. In the third expreripent, seasonal ion content andeco-physiological characters variation in T. salsuginea in its habitat, YellowRiver delta saline and alkaline land, from January to May in2012were alsostudied. The results showed that Na~+, Ca~(2+) and Cl~-content in the soilincreased from January to May, and was closely related to temperature andwater content in the soil. Na+content in T. salsuginea was highest inJanuary and February and decreased with time. K~+、Ca~(2+)、Mg~(2+)、Cl~-、SO_4~(2-)content showed increasing tendency, in particular, Cl-content, with theincrease of temperature and the growth stage. The accumulation of K~+、Ca~(2+)、Mg~(2+)、Cl~-、SO_4~(2-) was beneficial to osmaregulation in T. salsugineaunder salinity. K~+, Mg~2+and Ca~(2+) content in the root were lower than that inthe shoot because nutritional ion was in priority distribution to leaves,flowers and pods from roots. The eco-physiological variation in T.salsuginea showed that the effect of temperature was larger than that of ioncontent changes in the soil. T. salsuginea showed retarded growth withhigher accumulation of MDA, lower chlorophyte content. But T. salsugineaadjusted eco-physiological characters to the low temperature such as higherproline and soluble sugar content which protected T.salsuginea from theharm of low temperature and ion variation. These results suggested that compatible solute content is closely related to temperature, which is one ofthe adaptive mechanisms of T. salsuginea to low temperature and highsalinity.
Based on the research of T. salsuginea, T. salsuginea could survive in200mmol L~(-1)NaCl. The main mechanism was compartmentalization ofexcessive salt ion into vacuoles, which is similar to heavy metalsuperaccumulator plant. Heavy metal pollution in Yellow River delta soil isgetting worse; T. salsuginea might be a potential heavy metalsuperaccumulator, which needs to study. In the fourth experiment, a potexperiment was used to investigate growth responses and eco-physiologicalcharacters of T. salsuginea under Cd, Cu and Pb treatments. The resultshowed that the root length and dry mass decreased and MDA accumulatedsignificantly with the increase of Cd, Cu and Pb concentrations. T.salsuginea showed apparent damage symptoms such as lower chlorophyllcontent, etiolated leaves, higher MDA accumulations and death ofseedlings under Cd treatment compared with Cu and Pb. T. salsuginea cangrow well under Pb treatment though Pb treatment concentration was thehighest. From the point of metal accumulation and BAF, T. salsugineaabsorbed Cd>Cu>Pb. These results indicate that T. salsuginea has certainpotential on remedying the soil polluted by Cd with low and middleconcentrations.
The innovations of this thesis were shown as follows:
1. In this study, the ungerminated seeds of T. salsuginea under highsalinity quickly germinated though the seed germination of T. salsugineawas a little more sensitive to salinity compared with A. thaliana. At theseedling stage, however, T. salsuginea showed better growth with higherchlorophyll content, less MDA content and higher proline content and K~+/Na~+ratio under salinity. These characteristics indicate that T. salsuginea ismore adaptive to salinity than A. thaliana at both seed germination andseedling stages and explain why A. thaliana is excuded from salinelocations and T. salsuginea can survive and form its populations in salinesoils.
2. The seed vernalization and photoperiod of T. salsuginea were studiedfor the first time. The results showed that seed vernalization promoted theseed germination and that T. salsuginea flowered after seed vernalizationwhich showed that T. salsuginea is capable of vernalization at stage ofimbibed seeds. T. salsuginea still stayed at vegetative stage and couldn’tflower when seeds were vernalized less than6days. And100%of T.salsuginea flowered when seeds were vernalized for22-40days. The timewhen the first flower appeared was shortened by seed vernalization.Expression of ThFLC, which is a repressor of flowering, was markedlyreduced by vernalization. The requirement for vernalization of T. salsuginea is obligate instead of facultative. We also observed here that T.salsuginea generated flowers only when day length exceeded9h. Thenumber of flowers and dry mass per plant also increased with an increaseof day length. Dry mass per plant and number of flowers per plant were thehighest when day length exceeded14h. These results therefore confirmthat T. salsuginea is a long-day plant with a critical day length requirementof approximately9h. In addition, the time required to produce the firstflower decreased as day length exceeded9h. These results indicated that T.salsuginea is a typical long-day plant with9h photoperiod. Therequirement of vernalization and long days by T. salsuginea (Shandongecotype) is at least partially the product of adaptation to the localenvironment. These results facilitate greatly the future research onsalt-tolerant mechanisms of T. salsuginea.
3. Soluble ion, proline, MDA and chlorophyll content of T. salsugineawere closely related to growth and development stages, ion content in thesoil and rainfall. For T. salsuginea, the effect of growth and developmentstages on the adaptation to stress environment was larger than that oftemperature and ion content changes in the soil. Under the natural location,T. salsuginea adjusted eco-physiological characters to low temperature,drought and salinity caused by the seasonal variances.
4. T. salsuginea accumulated Cd to some degree, but did not survive inhigher concentration of Cd (130mg kg~(-1)). T. salsuginea finished its life cycle under Cu and Pb stress. Therefore, T. salsuginea was tolerant to Cuand Pb but not accumulated them. Conclusively, T. salsuginea can be usedin remedying the soil polluted by Cd with low and middle concentrations.
引文
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