寒地水稻对碱胁迫的响应及鉴定指标评价
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摘要
试验于2010-2011年在黑龙江省农业科学院佳木斯水稻研究所内进行。共选择173份国内外适合黑龙江省种植的水稻种质资源，进行芽期、苗期和成熟期的耐碱研究和鉴定指标评价。试验从对水稻各个生育期的生理形态、产量和超微结构观察等入手，探讨了寒地水稻对苏打碱胁迫的响应机理，对部分参试品种进行了苗期耐碱分级，从微观角度阐述了胁迫后水稻根细胞的形态变化，揭示了不同基因型品种叶片细胞超微结构变化差异和不同伤害机制的形成，通过试验提出了以下结论和观点：
1.芽期碱胁迫，胚根反应最敏感。芽鞘变化与耐碱性无关。影响耐碱级别判定的主要指标为相对芽长、相对胚根长、相对根条数、相对芽干重、相对根干重，依其建立胁迫7天的耐碱级别判定方程为：Y=11.251-3.6747X1-1.641X2-1.179X3-3.900X4-1.973X5，其中相对芽长、相对芽干重和相对根干重是水稻芽期耐碱的三大指标，依其建立的耐碱级别判定方程为：Y=9.234-4.506X1-2.167X2-2.621X3。
2.0.20%混合碱处理后，苗长和根系长度均有下降趋势，对根系的伸长生长抑制作用最显著，但地上和地下部位对碱胁迫敏感性不相关。碱胁迫延迟水稻的叶龄进程，降低叶面积、绿叶数、根系干质量、冠层干物质积累及根冠比，刺激水稻分蘖提前发生，以及主茎同一叶龄叶片伸长、宽度下降、叶面积增大。单株叶面积的变化、绿叶数、根冠比和根冠长度比的相对值可作为水稻苗期耐碱性鉴定的形态指标；碱处理对水稻叶片色素合成影响显著，影响最大的是叶绿素A，其次是叶绿素B，类胡萝卜素受到的影响相对最小。各色素成分之间存在极显著相关性。胁迫后，茎叶脯氨酸含量相对根系脯氨酸含量增加显著。胁迫10天能够激发部分品种SOD保护酶的活性，也会造成部分品种SOD合成系统的功能受阻或者酶本身的活性受阻；碱胁迫会提高水稻苗期叶片POD活性。
3.碱胁迫致使水稻根细胞排列的整齐性和紧凑性降低，抑制水稻根毛的生长、根细胞纵向伸长、大维管束的形成，胁迫会产生部分未发育完全的维管束，同时促进根系木质化和枝根形成；皮层薄壁细胞、中柱鞘部分细胞会因胁迫坏死，表皮细胞相对对照脱落早，立体感会因胁迫而降低。通过对4个基因型品种的超微结构观察，发现碱胁迫对不同基因型水稻品种伤害机制有差异。龙粳27主要伤害叶绿体和线粒体，同时淀粉体数量变少，体积变小；绥粳5号线粒体和叶绿体伤害相对较小，淀粉体体积变大；藤系138叶绿体所受影响较大，线粒体受影响较小，内含淀粉体颗粒减少；越光叶绿体结构变化明显，线粒体数量明显下降，结构模糊，内脊不清晰，淀粉体数量增多，体积明显增大。处理后所有品种嗜锇颗粒明显增多，且程度较深。
4.随处理浓度增加水稻株高和顶端第一片展开叶相对含水量呈先增后降趋势；叶片色素含量、叶绿素A/B值、SOD活性、维生素C含量、伤流量均随处理浓度升高呈直线下降趋势，回归曲线为一次方程；脯氨酸含量、可溶性糖含量、过氧化物酶（POD）活性、过氧化氢酶(CAT)活性随处理浓度呈先降后升或逐渐升高的趋势，与处理浓度之间均存在二次回归关系；丙二醛（MDA）含量随处理浓度升高呈直线上升趋势，可溶性糖含量随脯氨酸含量的增加亦呈逐渐上升趋势；POD与SOD活性呈线性负相关；随碱处理浓度升高，水稻叶片相对电导率呈上升趋势，两者呈显著二次回归关系。
5.每穴收获穗数和结实率随处理浓度呈先升高后降低趋势。低浓度处理增加有效穗数和结实率，处理浓度达到和超过0.10%，水稻有效穗数和结实率会显著降低。处理浓度增加，穗颍花量、千粒重、根干质量、茎叶干质量、穗干质量与单株干质量呈显著下降趋势。在一定浓度范围内，茎叶物质转化均优先向穗部积累。低浓度处理对水稻穗长影响不显著，处理浓度达0.15%，穗长显著降低。碱胁迫对水稻二次枝梗抑制作用明显大于一次枝梗，但空秕率无显著差异。高浓度处理显著降低籽粒饱满度。0.05%碱处理对一次枝梗千粒重影响小，高于此浓度对枝梗数的影响较大。二次枝梗千粒重与二次枝梗数比值变化则与之相反。敏感系数受浓度干扰因素较低，可将其作为水稻耐碱鉴定指标之一，经济系数、生物耐碱系数、耐碱系数和耐碱指数等指标可作为参考指标。水稻营养生长期和生殖生长期耐碱性会有差异。
The experiment is carried out in Jiamusi rice research institute of Heilongjiang academyof agricultural sciences in2010-2011.173varieties which are suitable for the cultivation ofHeilongjiang Province are collected from domestic and foreign country. Research of Alkaliresistance and assessment of identification index are proceed in buds period, seeding stageand mature stage. In this paper the response mechanism of soda alkali stress and classificationof some test material are discussed from aspect of morphology, physiology, yield trait andultrastructure. Changes of root-cells-morphological and ultrastructure and formation of thedifferent damage mechanisms among different genotype varieties are evaluated after stressfrom the microscopic point of view. The main conclusion and views are as follow:
1. The radicle is the most sensitive to the alkali stress at germinating stage. There is nocorrelation between the changes of coleoptiles and alkali resistance. The main indexes whichdefined alkali resistance level are relative shoot length, relative radicle length, relative radiclenumber, relative shoot dry weight and relative radicle dry weight. Based on that, thedetermine equation of alkali resistance isY=11.251-3.6747X1-1.641X2-1.179X3-3.900X4-1.973X5during that7days, and thedetermine equation of alkali resistance is Y=9.234-4.506X1-2.167X2-2.621X3which is builtbased on three key index, relative shoot length, relative shoot dry weight and relative rootsdry weight.
2. After0.20%mixed alkali treatment, the length of shoot and root length are showed adownward trend and significant inhibition to root elongation and growth， but there is nocorrelation in susceptibility for alkali stress between aboveground and underground parts.Many physiological indicators changed during the alkali stress treatment such as the leaf ageprocess delayed, the reduction of leaf area, green leaf number, root dry weight, canopy drymatter accumulation and root-shoot ratio, meanwhile the leaf length and area are increasedand the leaf width is declined. Some indexes can serve as indicator of alkali resistance at ricegerminating stage, such as the changes of leaf area per plant and relative value of green leafnumber, root shoot ratio and root shoot length ratio. A significant effect is occurred on the leafpigment synthesis, descending order is chlorophyll A, chlorophyll B and carotenoid minimally.The proline content in stems and leaves are increased significantly than that in root underalkali stress. Alkali stress could simulate the activity of protective enzyme of SOD in somevarieties, and also block the functions of SOD synthetic system or the activity of SOD insome varieties. Alkali stress could improve the activity of leaves POD in rice at germinatingstage.
3. The regularity and tenseness of root cell is descended, the longitudinal elongation ofroot cell and formation of great vascular bundles are restrained by alkali stress, and also someimmature vascular bundles are formed and roots lignification and branch roots formation arepromoted. Cortex parenchyma cells and part of pericycle cells became necrosis because ofalkali stress; epidermic cells dropped earlier than CK, the stereoscopic impression wouldlower because of the alkali stress. The damage mechanism of alkali stress to rice varieties isdifference based on the observation of the4varieties’ ultrastructure. The damage inchloroplasts and mitochondria are serious and the quantity and size of starch descended InLongjing27, however Suijing5is on the contrary part. Tengxi138’s chloroplasts are moresensitive to Alkali Stress than mitochondria, and the quantity of starch became less. The effects on Koshihikari are complex: the structure of chloroplasts changed obviously, thequantity of anyloplast descended significantly, the structure indistincted, and the inner ridge isnot clear, the quantity of anyloplast became more, and the volume became bigger. Under thealkali stress, the granular osmophilic materials (GOM) of all varieties became more anddarker.
4. The effects of alkali stress on plant height and the relative water content of the firstunfold leaf present a trend of fluctuations (increase first then descend) with the treatmentconcentration increasing. A linear downward trend is present at leave pigments content, thevalue of Chlorophyll A/Chlorophyll B, the activity of SOD, the content of vitamin C and thebleeding, regression curve is first-order equation. There are quadratic regression relationshipbetween treatment concentration and many physiological indexes that is the content of proline,soluble sugar, the activity of SOD and CAT. The tendency showed firstly descend thenincrease trend with the treatment concentration increasing. The content of MDA is present astraight line upward trend within the increasing of treatment concentration. The increment ofthe soluble sugar is present gradual upward trend within the increasing of proline. The activityof POD and SOD shows negative linear correlation. With the increasing of alkaliconcentration, relative conductivity in rice leaf present a significant upward trend, besides, asignificant quadratic regression appeared between them.
5. The number of panicles per hole and seed setting rate present a floating trend with thechanging of concentration. The number of panicles per hole and seed setting rate increasingwhen treatment are in low concentration level, once the concentration reach or exceed0.10%,the two indicators begin to decrease significantly. The root dry weights, stems and leaves,panicles and plant of rice begin to significantly descend with the concentration increasing.The substance of stems and leaves is primary accumulated in panicles within a certainconcentration range. There is no significant effect when treatment of concentration at lowlevel. However, panicle length descended significantly when the concentration rose to0.15%.The inhibitory action of alkali stress on secondary branches is more effective than that of onprimary branches. However, fail to detect significant difference on grain empty rate.Plumpness of grains descend significantly when treatment of concentration at high level. Thelittle effect is present in thousand grains weight of primary branches when treatmentconcentration of alkali is lower than0.05%. However, a greater impact is present once theconcentration beyond0.05%. The changes of thousand grains weight ratio of secondarybranches and number of secondary branches are on the contrary part. Interference ofsensitivity coefficient is low to the concentration which could be used for evaluating the alkaliresistance of rice. The other index could be considered as reference, such as economiccoefficient, biological alkali resistance coefficient, alkali resistance index and alkali resistancecoefficient. There would be difference in alkali resistance of rice between vegetative growthstage and reproductive growth stage.
1.芽期碱胁迫，胚根反应最敏感。芽鞘变化与耐碱性无关。影响耐碱级别判定的主要指标为相对芽长、相对胚根长、相对根条数、相对芽干重、相对根干重，依其建立胁迫7天的耐碱级别判定方程为：Y=11.251-3.6747X1-1.641X2-1.179X3-3.900X4-1.973X5，其中相对芽长、相对芽干重和相对根干重是水稻芽期耐碱的三大指标，依其建立的耐碱级别判定方程为：Y=9.234-4.506X1-2.167X2-2.621X3。
2.0.20%混合碱处理后，苗长和根系长度均有下降趋势，对根系的伸长生长抑制作用最显著，但地上和地下部位对碱胁迫敏感性不相关。碱胁迫延迟水稻的叶龄进程，降低叶面积、绿叶数、根系干质量、冠层干物质积累及根冠比，刺激水稻分蘖提前发生，以及主茎同一叶龄叶片伸长、宽度下降、叶面积增大。单株叶面积的变化、绿叶数、根冠比和根冠长度比的相对值可作为水稻苗期耐碱性鉴定的形态指标；碱处理对水稻叶片色素合成影响显著，影响最大的是叶绿素A，其次是叶绿素B，类胡萝卜素受到的影响相对最小。各色素成分之间存在极显著相关性。胁迫后，茎叶脯氨酸含量相对根系脯氨酸含量增加显著。胁迫10天能够激发部分品种SOD保护酶的活性，也会造成部分品种SOD合成系统的功能受阻或者酶本身的活性受阻；碱胁迫会提高水稻苗期叶片POD活性。
3.碱胁迫致使水稻根细胞排列的整齐性和紧凑性降低，抑制水稻根毛的生长、根细胞纵向伸长、大维管束的形成，胁迫会产生部分未发育完全的维管束，同时促进根系木质化和枝根形成；皮层薄壁细胞、中柱鞘部分细胞会因胁迫坏死，表皮细胞相对对照脱落早，立体感会因胁迫而降低。通过对4个基因型品种的超微结构观察，发现碱胁迫对不同基因型水稻品种伤害机制有差异。龙粳27主要伤害叶绿体和线粒体，同时淀粉体数量变少，体积变小；绥粳5号线粒体和叶绿体伤害相对较小，淀粉体体积变大；藤系138叶绿体所受影响较大，线粒体受影响较小，内含淀粉体颗粒减少；越光叶绿体结构变化明显，线粒体数量明显下降，结构模糊，内脊不清晰，淀粉体数量增多，体积明显增大。处理后所有品种嗜锇颗粒明显增多，且程度较深。
4.随处理浓度增加水稻株高和顶端第一片展开叶相对含水量呈先增后降趋势；叶片色素含量、叶绿素A/B值、SOD活性、维生素C含量、伤流量均随处理浓度升高呈直线下降趋势，回归曲线为一次方程；脯氨酸含量、可溶性糖含量、过氧化物酶（POD）活性、过氧化氢酶(CAT)活性随处理浓度呈先降后升或逐渐升高的趋势，与处理浓度之间均存在二次回归关系；丙二醛（MDA）含量随处理浓度升高呈直线上升趋势，可溶性糖含量随脯氨酸含量的增加亦呈逐渐上升趋势；POD与SOD活性呈线性负相关；随碱处理浓度升高，水稻叶片相对电导率呈上升趋势，两者呈显著二次回归关系。
5.每穴收获穗数和结实率随处理浓度呈先升高后降低趋势。低浓度处理增加有效穗数和结实率，处理浓度达到和超过0.10%，水稻有效穗数和结实率会显著降低。处理浓度增加，穗颍花量、千粒重、根干质量、茎叶干质量、穗干质量与单株干质量呈显著下降趋势。在一定浓度范围内，茎叶物质转化均优先向穗部积累。低浓度处理对水稻穗长影响不显著，处理浓度达0.15%，穗长显著降低。碱胁迫对水稻二次枝梗抑制作用明显大于一次枝梗，但空秕率无显著差异。高浓度处理显著降低籽粒饱满度。0.05%碱处理对一次枝梗千粒重影响小，高于此浓度对枝梗数的影响较大。二次枝梗千粒重与二次枝梗数比值变化则与之相反。敏感系数受浓度干扰因素较低，可将其作为水稻耐碱鉴定指标之一，经济系数、生物耐碱系数、耐碱系数和耐碱指数等指标可作为参考指标。水稻营养生长期和生殖生长期耐碱性会有差异。
The experiment is carried out in Jiamusi rice research institute of Heilongjiang academyof agricultural sciences in2010-2011.173varieties which are suitable for the cultivation ofHeilongjiang Province are collected from domestic and foreign country. Research of Alkaliresistance and assessment of identification index are proceed in buds period, seeding stageand mature stage. In this paper the response mechanism of soda alkali stress and classificationof some test material are discussed from aspect of morphology, physiology, yield trait andultrastructure. Changes of root-cells-morphological and ultrastructure and formation of thedifferent damage mechanisms among different genotype varieties are evaluated after stressfrom the microscopic point of view. The main conclusion and views are as follow:
1. The radicle is the most sensitive to the alkali stress at germinating stage. There is nocorrelation between the changes of coleoptiles and alkali resistance. The main indexes whichdefined alkali resistance level are relative shoot length, relative radicle length, relative radiclenumber, relative shoot dry weight and relative radicle dry weight. Based on that, thedetermine equation of alkali resistance isY=11.251-3.6747X1-1.641X2-1.179X3-3.900X4-1.973X5during that7days, and thedetermine equation of alkali resistance is Y=9.234-4.506X1-2.167X2-2.621X3which is builtbased on three key index, relative shoot length, relative shoot dry weight and relative rootsdry weight.
2. After0.20%mixed alkali treatment, the length of shoot and root length are showed adownward trend and significant inhibition to root elongation and growth， but there is nocorrelation in susceptibility for alkali stress between aboveground and underground parts.Many physiological indicators changed during the alkali stress treatment such as the leaf ageprocess delayed, the reduction of leaf area, green leaf number, root dry weight, canopy drymatter accumulation and root-shoot ratio, meanwhile the leaf length and area are increasedand the leaf width is declined. Some indexes can serve as indicator of alkali resistance at ricegerminating stage, such as the changes of leaf area per plant and relative value of green leafnumber, root shoot ratio and root shoot length ratio. A significant effect is occurred on the leafpigment synthesis, descending order is chlorophyll A, chlorophyll B and carotenoid minimally.The proline content in stems and leaves are increased significantly than that in root underalkali stress. Alkali stress could simulate the activity of protective enzyme of SOD in somevarieties, and also block the functions of SOD synthetic system or the activity of SOD insome varieties. Alkali stress could improve the activity of leaves POD in rice at germinatingstage.
3. The regularity and tenseness of root cell is descended, the longitudinal elongation ofroot cell and formation of great vascular bundles are restrained by alkali stress, and also someimmature vascular bundles are formed and roots lignification and branch roots formation arepromoted. Cortex parenchyma cells and part of pericycle cells became necrosis because ofalkali stress; epidermic cells dropped earlier than CK, the stereoscopic impression wouldlower because of the alkali stress. The damage mechanism of alkali stress to rice varieties isdifference based on the observation of the4varieties’ ultrastructure. The damage inchloroplasts and mitochondria are serious and the quantity and size of starch descended InLongjing27, however Suijing5is on the contrary part. Tengxi138’s chloroplasts are moresensitive to Alkali Stress than mitochondria, and the quantity of starch became less. The effects on Koshihikari are complex: the structure of chloroplasts changed obviously, thequantity of anyloplast descended significantly, the structure indistincted, and the inner ridge isnot clear, the quantity of anyloplast became more, and the volume became bigger. Under thealkali stress, the granular osmophilic materials (GOM) of all varieties became more anddarker.
4. The effects of alkali stress on plant height and the relative water content of the firstunfold leaf present a trend of fluctuations (increase first then descend) with the treatmentconcentration increasing. A linear downward trend is present at leave pigments content, thevalue of Chlorophyll A/Chlorophyll B, the activity of SOD, the content of vitamin C and thebleeding, regression curve is first-order equation. There are quadratic regression relationshipbetween treatment concentration and many physiological indexes that is the content of proline,soluble sugar, the activity of SOD and CAT. The tendency showed firstly descend thenincrease trend with the treatment concentration increasing. The content of MDA is present astraight line upward trend within the increasing of treatment concentration. The increment ofthe soluble sugar is present gradual upward trend within the increasing of proline. The activityof POD and SOD shows negative linear correlation. With the increasing of alkaliconcentration, relative conductivity in rice leaf present a significant upward trend, besides, asignificant quadratic regression appeared between them.
5. The number of panicles per hole and seed setting rate present a floating trend with thechanging of concentration. The number of panicles per hole and seed setting rate increasingwhen treatment are in low concentration level, once the concentration reach or exceed0.10%,the two indicators begin to decrease significantly. The root dry weights, stems and leaves,panicles and plant of rice begin to significantly descend with the concentration increasing.The substance of stems and leaves is primary accumulated in panicles within a certainconcentration range. There is no significant effect when treatment of concentration at lowlevel. However, panicle length descended significantly when the concentration rose to0.15%.The inhibitory action of alkali stress on secondary branches is more effective than that of onprimary branches. However, fail to detect significant difference on grain empty rate.Plumpness of grains descend significantly when treatment of concentration at high level. Thelittle effect is present in thousand grains weight of primary branches when treatmentconcentration of alkali is lower than0.05%. However, a greater impact is present once theconcentration beyond0.05%. The changes of thousand grains weight ratio of secondarybranches and number of secondary branches are on the contrary part. Interference ofsensitivity coefficient is low to the concentration which could be used for evaluating the alkaliresistance of rice. The other index could be considered as reference, such as economiccoefficient, biological alkali resistance coefficient, alkali resistance index and alkali resistancecoefficient. There would be difference in alkali resistance of rice between vegetative growthstage and reproductive growth stage.
引文
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