甜高粱苗期对苏打盐碱胁迫的适应性机制及差异基因表达分析
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摘要
土壤盐碱化制约作物生长并导致减产，甜高粱是耐盐碱、生物产量高、可食用和饲用，具有发展潜力的作物。为探讨甜高粱对苏打盐碱胁迫的适应机制，本试验使用NaHCO_3和Na_2CO_3按5：1摩尔比混合的苏打盐碱胁迫筛选甜高粱芽期及苗期敏感性及耐性品种，然后以耐性强的M-81E和耐性弱的314B两个品种为试材，分析了不同甜高粱品种在苏打盐碱胁迫后生长发育、渗透调节、活性氧清除、Na~+吸收与分配、有机酸变化与分泌、叶片解剖结构、叶绿体及线粒体超微结构等方面的变化及差异，较全面的阐述了甜高粱苗期对苏打盐碱胁迫的适应机制，为盐碱土壤甜高粱种植种质的筛选和栽培提供参考。同时，通过SSH方法获得一批甜高粱抗盐碱相关基因的ESTs，可为甜高粱及禾本科植物耐盐碱相关基因克隆或抗盐碱基因工程育种提供理论依据和基因资源。主要研究结果如下：
1.苏打盐碱胁迫下，随浓度升高，甜高粱发芽率逐渐降低，盐害率升高，各品种间存在较大差异。盐碱胁迫对根芽生长的抑制显著高于对发芽的影响，且对胚根生长的抑制尤其严重。根据多个发芽相关指标用隶属函数法对甜高粱品种进行耐盐碱性分析，7个品种的种子耐盐碱性由强到弱的综合排序为： MN3739>M-81E>贝利>Rio>ES725>3222>雷伊。
2.经盐碱筛选发现甜高粱各品种在芽期和苗期对苏打盐碱胁迫的耐受性不同。进一步苗期试验表明，苏打盐碱胁迫可降低幼苗植株总重量、株高、根长、根系活力和总叶绿素含量，增加质膜相对透性和丙二醛的含量。但胁迫后根冠比在不同品种中或升或降。综合分析后认为，品种M-81E具有较强的耐受性，314B表现出较弱的耐受性。
3.植物可通过产生渗透调节物质来缓解盐碱胁迫造成的渗透胁迫，还能提高自身抗氧化酶系统活力清除活性氧。苏打盐碱胁迫可增加供试的两个品种甜高粱幼苗的可溶性糖、可溶性蛋白、脯氨酸和甜菜碱等4个渗透调节物质含量，耐性弱的314B产生更多的渗透调节物质。胁迫还使POD、CAT和GSH-Px酶活性增加，SOD在品种间变化不同，有增有降。耐性强的M-81E的抗性酶系统活力均比314B高，体现出更强的活性氧清除能力。甜高粱幼苗地上部的NO含量在胁迫后升高，而地下部则下降。
4.盐碱胁迫增加了甜高粱幼苗整株、叶、叶鞘及根中的Na~+含量，但Na~+较多积累在根及叶鞘中；胁迫使幼苗各部分的K~+和Ca~(2+)含量降低，导致幼苗叶、叶鞘及根中K~+/Na~+值和Ca2+/Na+值降低，但离子选择性运输系数TS_(K，Na)和TS_(Ca，Na)明显升高，说明甜高粱从根向叶中运输K~+和Ca~(2+)的选择性强，而运输Na~+的选择性弱。
5.苏打盐碱胁迫降低了甜高粱幼苗根液泡膜上PPase酶的水解和质子泵活力，而增
强了ATPase酶的水解和质子泵活力。同时，胁迫使H~+/Na~+反向运输蛋白活力显著增强（P<0.05），但高浓度会使该运输体活力显著下降（P<0.05），但仍明显高于对照。
6.苏打盐碱胁迫下，甜高粱幼苗叶片具有非常微弱的排Na~+能力，因此，幼苗叶片的泌Na~+行为不是其抵抗盐碱胁迫的主要方式。幼苗根及叶鞘的拒Na~+作用以及根液泡中Na~+的区域化是甜高粱适应苏打盐碱胁迫的主要方式。与品种314B相比，M-81E品种根和叶鞘的拒Na~+能力强，叶片中Na+含量少而受盐碱胁迫的影响较小；根液泡膜上ATPase酶和H~+/Na~+反向运输蛋白活力强，PPase酶活力下降程度小，使得液泡中Na+区域化程度高；同时体内K+含量高将进一步提高其耐盐性。
7.甜高粱在受到苏打盐碱胁迫后，地上部及根部各类有机酸及有机酸总量均下降，且未发现主效有机酸积累。PEPC酶在低浓度胁迫时，活力升高，而在高浓度时则下降。盐碱胁迫下甜高粱有机酸含量与PEPC酶活力不存在直接相关性，但PEPC酶活力升高，有机酸含量下降幅度小，若PEPC酶活力降低，有机酸含量就会大幅下降。甜高粱在受到苏打盐碱胁迫时，能迅速降低根外pH值，且调节能力较强。但根系分泌的有机酸量很少，推测甜高粱利用有机酸降低根外pH值的作用很小。M-81E无论地上部还是根部，其各有机酸量及总酸量均比314B高；其PEPC酶活力在低浓度胁迫时升高幅度大，而在高浓度胁迫时下降幅度小。M-81E降低根外pH值的能力更强，可将培养液的pH值从9.34降低到8.81，而314B则只降到9.10。
8.苏打盐碱胁迫影响了甜高粱幼苗的生长发育，使幼苗的株高和茎基周长显著降低；使植株基部叶片及叶尖变黄；并使叶片发育滞后，在相同的生长时间内，叶片数变少。胁迫后，甜高粱叶片厚度、中脉厚度和最大导管直径都极显著变小（P<0.01），叶片上下角质层的厚度均极显著增加（P<0.01），下表皮厚度也显著增加。品种314B在盐碱胁迫后叶片变黄面积大，中脉厚度、株高和茎基周长下降的程度均比M-81E大。
9.苏打盐碱胁迫使叶绿体膨胀，被膜和基质分离，类囊体膜破坏，基粒片层松散肿胀，有些基粒片层还发生扭曲变形，淀粉粒积累，且形成较多嗜锇滴颗粒；线粒体的数量增加，且分布在叶绿体附近，外膜断裂、消解，多数线粒体内膜上嵴数量减少，甚至消失，结构紊乱。314B叶绿体的超微结构受损伤较严重，嗜锇滴颗粒出现数量多，尤其是叶绿体被膜和基质发生的类似质壁分离现象更明显。
10.以耐苏打盐碱胁迫甜高粱品种M-81E为材料，使用SSH方法成功构建了苏打盐碱胁迫下差异表达基因的cDNA文库。经Blast比对分析，103个ESTs（81.1%）可以找到已知的同源序列，另有24个ESTs（18.9%）未获得同源匹配。其中只有48个ESTs（占46.6%）有功能注释，而55个ESTs（占53.4%）功能未知或者为假定蛋白。且在推测为假定蛋白的ESTs中，发现大多数ESTs与高粱逆境胁迫后构建cDNA文库中的ESTs序列有较高的相似性。
11.根据数据库中的功能注释，结合GO和COG分类发现，在盐碱胁迫过程中，多种基因被诱导表达，涉及植物的光合作用、物质（包括碳水化合物、脂肪、氨基酸、辅酶和无机离子等）和能量代谢、细胞壁和细胞膜的组成、水通道、信号转导及转录调控因子等。
Large amount areas of saline and alkaline soils limit crop growth and reduce agriculturalproductivity. Sorghum (Sorghum bicolor L. Moench) crop has been considered relativelymore salt tolerant than other cereals and has the potential as a grain and fodder crop in salineand alkaline soils. In order to study the adaptive mechanisms of sweet sorghum seedlingsresponded to saline-alkali stress (molar ratio of NaHCO_3:Na_2CO_3=5:1), based on thescreening results of different varieties resisting the stress, the two contrasting varieties,314B(the sensitive) and M-81E (the resistant) were selected for the actual study. Undersaline-alkali stress, a series of issues were analyzed, including growth and development,osmoregulation, scaenging of reactive oxygen species (ROS), sodium absorption anddistribution, changes and secretion of organic acid, ultrastructure of chloroplast andmitochondria, and leaf anatomical structure. Comparison of the changes and difference ofabove matters between two varieties, the adaptive mechanisms of sweet sorghum seedlingresponded to saline-alkali stress were generally stated. These results will attribute to screenresistant genotypes of sweet sorghum and formulate reasonable measures for planting crops atsoda saline-alkaline lands. Meanwhile, using SSH (suppression subtractive hybridization) kits,a group of ESTs of genes related to enduring the saline-alkali stress were obtained, that willprovide partial sequences to further acquire the overall length cDNA of some important genesand scientific basis on genetic improvement of sweet sorghum and gramineous plants for salttolerance. The major results were as follows:
1. Under saline-alkali stress, the germination rate of sweet sorghum increased withincreasing concentration of treatment solutions, but salt toxicity rate was the opposite. Therewere differences among varieties. The degree of restraining the growth of radicle and germ,which come from saline-alkali stress was more serious than germination. On the basis of thegermination rate, germination percentage, germination index, relative-vigor index,average-root length, average-germination percentage and salt toxicity rate of7varieties, thetolerance of sweet sorghum was assessed with the method of subordinate function valuesanalysis. The results showed that the7varieties of sweet sorghum were ordered in sequencefrom the strong to weak: MN3739> M-81E> Berry> Rio> ES725>3222>Wray.
2. After screening the tolerances in buds period and seedling stage of sweet sorghum, therewere various tolerant to saline-alkali stress among varieties. Then, the growth changes anddamage of plasma membrane in the six varieties of sweet sorghum seedlings was determinedwith condition of saline-alkali stress. The results showed that compared with the respectivecontrol group, under saline-alkali stress, the sweet sorghum seedlings was decreased the totalweight of plant, height, root vigor and total chlorophyll content, but increased the relativepermeability of plasma membranes and content of malonaldehyde. However, root-shoot ratioswere showed higher or lower among the different varieties. The assessment of the sweet sorghum tolerance to saline-alkali stress with the method of subordinate function valuesshowed that M-81was more tolerant to saline-alkali stress than314B.
3. The osmotic regulation substances, antoxidant enzymes and NO contents of sweetsorghum seedlings treated with alkali solutions were detected. The results showed that undersaline-alkali stress, the sweet sorghum seedlings increased the content of soluble sugar,soluble protein, proline and betaine, also, the activities of peroxidase, catalase, andglutathione peroxidase were improved. But, the activities of superoxide were showed higheror lower between the two varieties. The osmotic regulation substances produced from314Bwere more than M-81E, yet, the activities of antoxidant enzymes of M-81E were high than314B. Therefore,314B possessed higher osmotic adjustment abilities, and M-81E ownedstronger scavenging ROS abilities. Under saline-alkali stress, the NO contents in shoots ofsweet sorghum were increased, but, the ones in roots were decreased.
4. After sweet sorghum seedlings treated for3days with soda alkali solutions, the Na~+contents of whole plant, leaves, sheath and roots in two genotypes almost entirely increased,yet, major Na~+were accumulated in roots and sheath. Under saline-alkali stress, the contentsof K~+and Ca~(2+)in whole plant, leaves, sheath and roots decreased universally in twogenotypes. Consequently, the K~+/Na~+and Ca~(2+)/Na~+ratio decreased completely in eachtissue of two varieties, but, the K~+/Na~+transport selectivity (TSK，Na) and the Ca~(2~+)/Na~+transport selectivity (TSCa，Na) in roots were obviously risen, that from roots to leaves, theselective abilities of sweet sorghum to transport K~+and Ca~(2+)were strong, and to transport Na~+were weak.
5. Both hydrolytic activities and proton transporting activities of V-H~+-ATPase of rootswere stimulated under soda alkali stress, yet V-H~+-PPase activities were decreased followingincreasing concentration of solutions. Na~+/H~+exchange activities were intensively promoted.When treated with high concentration solution, the Na~+/H~+exchange activities weremarkedly restrained, but the activities were still higher than control.
6. The Na~+secretion from leaves wasn’t main way for sweet sorghum to resist soda alkali,because under saline-alkali stress, the Na~+concentration expelled from leaves was very little.It played an important role in sweet sorghum adapting to saline-alkali stress that the main Na~+exclusion localization was in roots and sheath. Meanwhile, vacuolar Na~+compartment wasalso major adaptive mechanism of sorghum to alkali stress. In roots, M-81E accumulatedmajor Na~+, but314B in leaves. M-81E had a higher ability to vacuolar Na~+compartment inroot cells that was in view of M-81E had higher activities of V-H~+-ATPase and Na~+/H~+exchanger, the depressed degree of V-H~+_-PPase activities was lesser than314B. And M-81Emaintained a higher level of K~+uptake, which resulted in higher K~+/Na~+transport selectivitythan314B. These result also demonstrated that the most of Na~+accumulated in root and vacuolar Na~+compartment in root cells were conductive to sweet sorghum tolerance to sodaalkali stress.
7. The organic acid contents in shoots and roots of sweet sorghum after treated soda alkalisolutions were decreased generally. Some organic acids which played important role inresisting stress weren’t accumulated. When treated with low concentration solutions, PEPCactivities were enhanced, but with high concentration solutions, the ones were decreased.Under saline-alkali stress, the organic acid contents had no direct correlation with PEPCactivities, but, the fall of organic acid was small with increasing PEPC activities, and the fallwas declined largely following with decreasing PEPC activities. When grown in soda alkalisolutions, sweet sorghum could rapidly cut down pH value of culture solution. Yet, theorganic acid secreted from root system was very little, thus, we concluded that the organicacid played a little part in regulating pH value outside roots. Both all kinds of organic acidsand total organic acid in shoots or roots were higher of M-81E than314B. Compare with314B, the degree of increasing PEPC activities was greater under low concentration solutions,and the degree of decreasing PEPC activities was littler under high concentration solutions.The pH value was from9.34to9.10in314B culture solution, yet, the value was from9.34to8.81in M-81E culture solution and M-81E showed stronger ability to regulate pH value ofoutside roots than314B.
8. The growth and development of sweet sorghum seedlings were suppressed on conditionof alkali stress. The plant height and stem base perimeter were reduced dramatically (P<0.01).The leaves located stem base and blade tips were flavescent, the leaf developments werelagged, and the leaf numbers become less. The leaf thickness, midrib thickness and biggervessels diameter of the two sweet sorghum seedlings were significantly decreased undersaline-alkali stress, while the upper stratum corneum thickness and lower stratum corneumthickness were significantly increased compared with the control group (P<0.01). The upperepidermis thickness stayed mainly the same, but lower epidermis thickness were significantlyenhanced (P<0.05). After treated with alkali solutions, the etiolated areas of leaves, the fall ofmidrib thickness plant height and stem base perimeter in314B were larger than in M-81E.
9. The effects of saline-alkali stress on ultrastructure in chloroplast and mitochondria werestudied. The results showed that the damages appeared such as expansion of chloroplast, thematrix separated from the chloroplast envelope, dissociation of thylakoid membrane,loosening and expanding of grana lamella, and even deformation of some grana lamella. Thestarch grain and osmophilic globule number in chloroplast were significantly increased(P<0.01). The mitochondria number become more and distributed largely near chloroplast. Inthe mitochondria, the number of ridge in the inner coat appeared to decrease and evendisappear. The chloroplast of314B seemed to be injured more serious than M-81E, for themore osmophilic globule number and greater degree of separated between matrix and the envelope.
10. A suppression subtractive hybridization cDNA library was constructed which wasM-81E treated with soda alkali. Similarity analysis based on Blast software in GenBank wasperformed.81.1%(103) of the ESTs could be found encoding putative proteins.18.9%(24)of the ESTs were found no significant similarity with ESTs collected in GenBank. Functionof48ESTs (46.6%) were known, and55(53.4%) were unknown. Most ESTs of guessedputative proteins were found having similarity with ESTs from cDNA libraries which wereconstructed about the different expression of sweet sorghum grown in various stresses.
11. According to the annotated function, classifying with GO and COGs, we found that agood deal of genes were induced to express excessively, involving related withphotosynthesis, the metabolisms of substances (such as carbohydrate, fat, amino acid,cozymaze and inorganic irons) and energy, structure of cell wall and membrane, aquaporin,signal transduction and transcription regulation.
1.苏打盐碱胁迫下，随浓度升高，甜高粱发芽率逐渐降低，盐害率升高，各品种间存在较大差异。盐碱胁迫对根芽生长的抑制显著高于对发芽的影响，且对胚根生长的抑制尤其严重。根据多个发芽相关指标用隶属函数法对甜高粱品种进行耐盐碱性分析，7个品种的种子耐盐碱性由强到弱的综合排序为： MN3739>M-81E>贝利>Rio>ES725>3222>雷伊。
2.经盐碱筛选发现甜高粱各品种在芽期和苗期对苏打盐碱胁迫的耐受性不同。进一步苗期试验表明，苏打盐碱胁迫可降低幼苗植株总重量、株高、根长、根系活力和总叶绿素含量，增加质膜相对透性和丙二醛的含量。但胁迫后根冠比在不同品种中或升或降。综合分析后认为，品种M-81E具有较强的耐受性，314B表现出较弱的耐受性。
3.植物可通过产生渗透调节物质来缓解盐碱胁迫造成的渗透胁迫，还能提高自身抗氧化酶系统活力清除活性氧。苏打盐碱胁迫可增加供试的两个品种甜高粱幼苗的可溶性糖、可溶性蛋白、脯氨酸和甜菜碱等4个渗透调节物质含量，耐性弱的314B产生更多的渗透调节物质。胁迫还使POD、CAT和GSH-Px酶活性增加，SOD在品种间变化不同，有增有降。耐性强的M-81E的抗性酶系统活力均比314B高，体现出更强的活性氧清除能力。甜高粱幼苗地上部的NO含量在胁迫后升高，而地下部则下降。
4.盐碱胁迫增加了甜高粱幼苗整株、叶、叶鞘及根中的Na~+含量，但Na~+较多积累在根及叶鞘中；胁迫使幼苗各部分的K~+和Ca~(2+)含量降低，导致幼苗叶、叶鞘及根中K~+/Na~+值和Ca2+/Na+值降低，但离子选择性运输系数TS_(K，Na)和TS_(Ca，Na)明显升高，说明甜高粱从根向叶中运输K~+和Ca~(2+)的选择性强，而运输Na~+的选择性弱。
5.苏打盐碱胁迫降低了甜高粱幼苗根液泡膜上PPase酶的水解和质子泵活力，而增
强了ATPase酶的水解和质子泵活力。同时，胁迫使H~+/Na~+反向运输蛋白活力显著增强（P<0.05），但高浓度会使该运输体活力显著下降（P<0.05），但仍明显高于对照。
6.苏打盐碱胁迫下，甜高粱幼苗叶片具有非常微弱的排Na~+能力，因此，幼苗叶片的泌Na~+行为不是其抵抗盐碱胁迫的主要方式。幼苗根及叶鞘的拒Na~+作用以及根液泡中Na~+的区域化是甜高粱适应苏打盐碱胁迫的主要方式。与品种314B相比，M-81E品种根和叶鞘的拒Na~+能力强，叶片中Na+含量少而受盐碱胁迫的影响较小；根液泡膜上ATPase酶和H~+/Na~+反向运输蛋白活力强，PPase酶活力下降程度小，使得液泡中Na+区域化程度高；同时体内K+含量高将进一步提高其耐盐性。
7.甜高粱在受到苏打盐碱胁迫后，地上部及根部各类有机酸及有机酸总量均下降，且未发现主效有机酸积累。PEPC酶在低浓度胁迫时，活力升高，而在高浓度时则下降。盐碱胁迫下甜高粱有机酸含量与PEPC酶活力不存在直接相关性，但PEPC酶活力升高，有机酸含量下降幅度小，若PEPC酶活力降低，有机酸含量就会大幅下降。甜高粱在受到苏打盐碱胁迫时，能迅速降低根外pH值，且调节能力较强。但根系分泌的有机酸量很少，推测甜高粱利用有机酸降低根外pH值的作用很小。M-81E无论地上部还是根部，其各有机酸量及总酸量均比314B高；其PEPC酶活力在低浓度胁迫时升高幅度大，而在高浓度胁迫时下降幅度小。M-81E降低根外pH值的能力更强，可将培养液的pH值从9.34降低到8.81，而314B则只降到9.10。
8.苏打盐碱胁迫影响了甜高粱幼苗的生长发育，使幼苗的株高和茎基周长显著降低；使植株基部叶片及叶尖变黄；并使叶片发育滞后，在相同的生长时间内，叶片数变少。胁迫后，甜高粱叶片厚度、中脉厚度和最大导管直径都极显著变小（P<0.01），叶片上下角质层的厚度均极显著增加（P<0.01），下表皮厚度也显著增加。品种314B在盐碱胁迫后叶片变黄面积大，中脉厚度、株高和茎基周长下降的程度均比M-81E大。
9.苏打盐碱胁迫使叶绿体膨胀，被膜和基质分离，类囊体膜破坏，基粒片层松散肿胀，有些基粒片层还发生扭曲变形，淀粉粒积累，且形成较多嗜锇滴颗粒；线粒体的数量增加，且分布在叶绿体附近，外膜断裂、消解，多数线粒体内膜上嵴数量减少，甚至消失，结构紊乱。314B叶绿体的超微结构受损伤较严重，嗜锇滴颗粒出现数量多，尤其是叶绿体被膜和基质发生的类似质壁分离现象更明显。
10.以耐苏打盐碱胁迫甜高粱品种M-81E为材料，使用SSH方法成功构建了苏打盐碱胁迫下差异表达基因的cDNA文库。经Blast比对分析，103个ESTs（81.1%）可以找到已知的同源序列，另有24个ESTs（18.9%）未获得同源匹配。其中只有48个ESTs（占46.6%）有功能注释，而55个ESTs（占53.4%）功能未知或者为假定蛋白。且在推测为假定蛋白的ESTs中，发现大多数ESTs与高粱逆境胁迫后构建cDNA文库中的ESTs序列有较高的相似性。
11.根据数据库中的功能注释，结合GO和COG分类发现，在盐碱胁迫过程中，多种基因被诱导表达，涉及植物的光合作用、物质（包括碳水化合物、脂肪、氨基酸、辅酶和无机离子等）和能量代谢、细胞壁和细胞膜的组成、水通道、信号转导及转录调控因子等。
Large amount areas of saline and alkaline soils limit crop growth and reduce agriculturalproductivity. Sorghum (Sorghum bicolor L. Moench) crop has been considered relativelymore salt tolerant than other cereals and has the potential as a grain and fodder crop in salineand alkaline soils. In order to study the adaptive mechanisms of sweet sorghum seedlingsresponded to saline-alkali stress (molar ratio of NaHCO_3:Na_2CO_3=5:1), based on thescreening results of different varieties resisting the stress, the two contrasting varieties,314B(the sensitive) and M-81E (the resistant) were selected for the actual study. Undersaline-alkali stress, a series of issues were analyzed, including growth and development,osmoregulation, scaenging of reactive oxygen species (ROS), sodium absorption anddistribution, changes and secretion of organic acid, ultrastructure of chloroplast andmitochondria, and leaf anatomical structure. Comparison of the changes and difference ofabove matters between two varieties, the adaptive mechanisms of sweet sorghum seedlingresponded to saline-alkali stress were generally stated. These results will attribute to screenresistant genotypes of sweet sorghum and formulate reasonable measures for planting crops atsoda saline-alkaline lands. Meanwhile, using SSH (suppression subtractive hybridization) kits,a group of ESTs of genes related to enduring the saline-alkali stress were obtained, that willprovide partial sequences to further acquire the overall length cDNA of some important genesand scientific basis on genetic improvement of sweet sorghum and gramineous plants for salttolerance. The major results were as follows:
1. Under saline-alkali stress, the germination rate of sweet sorghum increased withincreasing concentration of treatment solutions, but salt toxicity rate was the opposite. Therewere differences among varieties. The degree of restraining the growth of radicle and germ,which come from saline-alkali stress was more serious than germination. On the basis of thegermination rate, germination percentage, germination index, relative-vigor index,average-root length, average-germination percentage and salt toxicity rate of7varieties, thetolerance of sweet sorghum was assessed with the method of subordinate function valuesanalysis. The results showed that the7varieties of sweet sorghum were ordered in sequencefrom the strong to weak: MN3739> M-81E> Berry> Rio> ES725>3222>Wray.
2. After screening the tolerances in buds period and seedling stage of sweet sorghum, therewere various tolerant to saline-alkali stress among varieties. Then, the growth changes anddamage of plasma membrane in the six varieties of sweet sorghum seedlings was determinedwith condition of saline-alkali stress. The results showed that compared with the respectivecontrol group, under saline-alkali stress, the sweet sorghum seedlings was decreased the totalweight of plant, height, root vigor and total chlorophyll content, but increased the relativepermeability of plasma membranes and content of malonaldehyde. However, root-shoot ratioswere showed higher or lower among the different varieties. The assessment of the sweet sorghum tolerance to saline-alkali stress with the method of subordinate function valuesshowed that M-81was more tolerant to saline-alkali stress than314B.
3. The osmotic regulation substances, antoxidant enzymes and NO contents of sweetsorghum seedlings treated with alkali solutions were detected. The results showed that undersaline-alkali stress, the sweet sorghum seedlings increased the content of soluble sugar,soluble protein, proline and betaine, also, the activities of peroxidase, catalase, andglutathione peroxidase were improved. But, the activities of superoxide were showed higheror lower between the two varieties. The osmotic regulation substances produced from314Bwere more than M-81E, yet, the activities of antoxidant enzymes of M-81E were high than314B. Therefore,314B possessed higher osmotic adjustment abilities, and M-81E ownedstronger scavenging ROS abilities. Under saline-alkali stress, the NO contents in shoots ofsweet sorghum were increased, but, the ones in roots were decreased.
4. After sweet sorghum seedlings treated for3days with soda alkali solutions, the Na~+contents of whole plant, leaves, sheath and roots in two genotypes almost entirely increased,yet, major Na~+were accumulated in roots and sheath. Under saline-alkali stress, the contentsof K~+and Ca~(2+)in whole plant, leaves, sheath and roots decreased universally in twogenotypes. Consequently, the K~+/Na~+and Ca~(2+)/Na~+ratio decreased completely in eachtissue of two varieties, but, the K~+/Na~+transport selectivity (TSK，Na) and the Ca~(2~+)/Na~+transport selectivity (TSCa，Na) in roots were obviously risen, that from roots to leaves, theselective abilities of sweet sorghum to transport K~+and Ca~(2+)were strong, and to transport Na~+were weak.
5. Both hydrolytic activities and proton transporting activities of V-H~+-ATPase of rootswere stimulated under soda alkali stress, yet V-H~+-PPase activities were decreased followingincreasing concentration of solutions. Na~+/H~+exchange activities were intensively promoted.When treated with high concentration solution, the Na~+/H~+exchange activities weremarkedly restrained, but the activities were still higher than control.
6. The Na~+secretion from leaves wasn’t main way for sweet sorghum to resist soda alkali,because under saline-alkali stress, the Na~+concentration expelled from leaves was very little.It played an important role in sweet sorghum adapting to saline-alkali stress that the main Na~+exclusion localization was in roots and sheath. Meanwhile, vacuolar Na~+compartment wasalso major adaptive mechanism of sorghum to alkali stress. In roots, M-81E accumulatedmajor Na~+, but314B in leaves. M-81E had a higher ability to vacuolar Na~+compartment inroot cells that was in view of M-81E had higher activities of V-H~+-ATPase and Na~+/H~+exchanger, the depressed degree of V-H~+_-PPase activities was lesser than314B. And M-81Emaintained a higher level of K~+uptake, which resulted in higher K~+/Na~+transport selectivitythan314B. These result also demonstrated that the most of Na~+accumulated in root and vacuolar Na~+compartment in root cells were conductive to sweet sorghum tolerance to sodaalkali stress.
7. The organic acid contents in shoots and roots of sweet sorghum after treated soda alkalisolutions were decreased generally. Some organic acids which played important role inresisting stress weren’t accumulated. When treated with low concentration solutions, PEPCactivities were enhanced, but with high concentration solutions, the ones were decreased.Under saline-alkali stress, the organic acid contents had no direct correlation with PEPCactivities, but, the fall of organic acid was small with increasing PEPC activities, and the fallwas declined largely following with decreasing PEPC activities. When grown in soda alkalisolutions, sweet sorghum could rapidly cut down pH value of culture solution. Yet, theorganic acid secreted from root system was very little, thus, we concluded that the organicacid played a little part in regulating pH value outside roots. Both all kinds of organic acidsand total organic acid in shoots or roots were higher of M-81E than314B. Compare with314B, the degree of increasing PEPC activities was greater under low concentration solutions,and the degree of decreasing PEPC activities was littler under high concentration solutions.The pH value was from9.34to9.10in314B culture solution, yet, the value was from9.34to8.81in M-81E culture solution and M-81E showed stronger ability to regulate pH value ofoutside roots than314B.
8. The growth and development of sweet sorghum seedlings were suppressed on conditionof alkali stress. The plant height and stem base perimeter were reduced dramatically (P<0.01).The leaves located stem base and blade tips were flavescent, the leaf developments werelagged, and the leaf numbers become less. The leaf thickness, midrib thickness and biggervessels diameter of the two sweet sorghum seedlings were significantly decreased undersaline-alkali stress, while the upper stratum corneum thickness and lower stratum corneumthickness were significantly increased compared with the control group (P<0.01). The upperepidermis thickness stayed mainly the same, but lower epidermis thickness were significantlyenhanced (P<0.05). After treated with alkali solutions, the etiolated areas of leaves, the fall ofmidrib thickness plant height and stem base perimeter in314B were larger than in M-81E.
9. The effects of saline-alkali stress on ultrastructure in chloroplast and mitochondria werestudied. The results showed that the damages appeared such as expansion of chloroplast, thematrix separated from the chloroplast envelope, dissociation of thylakoid membrane,loosening and expanding of grana lamella, and even deformation of some grana lamella. Thestarch grain and osmophilic globule number in chloroplast were significantly increased(P<0.01). The mitochondria number become more and distributed largely near chloroplast. Inthe mitochondria, the number of ridge in the inner coat appeared to decrease and evendisappear. The chloroplast of314B seemed to be injured more serious than M-81E, for themore osmophilic globule number and greater degree of separated between matrix and the envelope.
10. A suppression subtractive hybridization cDNA library was constructed which wasM-81E treated with soda alkali. Similarity analysis based on Blast software in GenBank wasperformed.81.1%(103) of the ESTs could be found encoding putative proteins.18.9%(24)of the ESTs were found no significant similarity with ESTs collected in GenBank. Functionof48ESTs (46.6%) were known, and55(53.4%) were unknown. Most ESTs of guessedputative proteins were found having similarity with ESTs from cDNA libraries which wereconstructed about the different expression of sweet sorghum grown in various stresses.
11. According to the annotated function, classifying with GO and COGs, we found that agood deal of genes were induced to express excessively, involving related withphotosynthesis, the metabolisms of substances (such as carbohydrate, fat, amino acid,cozymaze and inorganic irons) and energy, structure of cell wall and membrane, aquaporin,signal transduction and transcription regulation.
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