高粱抗旱机制及评价指标的研究
摘要
本试验于2011～2012年在沈阳农业大学农学院试验基地进行,采用人工气候箱培养,PEG-6000溶液模拟干旱胁迫环境,在萌发期以4个不同浓度PEG-6000水溶液处理31个高粱品种,旨在根据高粱品种萌发期对不同干旱胁迫程度的响应,筛选出具有抗旱能力的高粱品种并探讨高粱萌发期抗旱性鉴定的方法。室外盆栽条件下,以4个不同抗旱性高粱品种为对象,研究农艺和生理生化指标与高粱抗旱性的关系,从多方面揭示高粱的抗旱机制,并对高粱的抗旱性进行鉴定和指标筛选,以期提供简单易行的农艺和生理生化指标对高粱抗旱性进行有效的评价,同时为丰富高粱抗旱性的研究内容、建立高粱抗旱鉴定指标体系及为抗旱育种提供理论依据和方法。本试验主要研究结果如下：
1、采用人工气候箱内培养,PEG-6000溶液模拟干旱胁迫环境,在萌发期以80g/L、120g/L、150g/L、175g/L PEG-6000水溶液处理31个高粱品种,旨在根据高粱品种萌发期对不同干旱胁迫程度的响应,筛选出具有抗旱能力的高粱品种并探讨高粱萌发期抗旱性鉴定的方法。通过主成分分析法(PCA)和神经网络自组织映射(SOM)聚类分析法对各品种进行抗旱性综合分析与评定。PCA结果表明,相对芽长、相对根长和相对萌发抗旱指数载荷量最大,将其作为萌发期高粱抗旱性筛选的主要评价指标,对31个高粱品种抗旱性进行了排序。最后通过SOM聚类分析将31个高粱品种按抗旱性强弱分为5类,吉杂305等4个品种为高度抗旱品种,HL5等4个品种为抗旱品种,辽杂10等8个品种为中等抗旱品种,锦杂103等7个品种为干旱敏感品种,锦杂93等8个品种为高度干旱敏感品种。研究认为,相对芽长、相对根长和相对萌发抗旱指数等指标可以作为高粱品种抗旱性鉴定的重要指标；SOM聚类分析可作为品种抗旱性分类的重要方法。
2、干旱胁迫下,高粱叶片自由水含量、束缚水含量、相对含水量和离体叶片失水速率等水分状况在不同生育时期均呈下降趋势,吉杂305(高度抗旱品种)的下降幅度显著低于吉杂127(高度干旱敏感品种),锦杂106(中等抗旱品种)和锦杂103(干旱敏感品种)介于二者之间。抗旱性品种具有较高的过氧化物酶(POD)、超氧化物歧化酶(SOD)和过氧化氢酶(CAT)活性,而丙二醛(MDA)含量少,具有较好的清除自由基的能力,脯氨酸、可溶性糖和可溶性蛋白含量较高,细胞水势较低。相关性分析表明,叶片温度可以直接或间接的反映干旱胁迫下高粱植株的水分状况及叶片保护性酶和渗透调节物质变化状况,能够体现出高粱抗旱性的强弱。叶温差可以作为高粱抗旱性筛选的一个重要指标,应用远红外热成像技术在高粱抗旱性的鉴定上具有可行性。
3、干旱胁迫下,高粱叶片最大光合效率(Fv/Fm)、表观电子传递速率(ETR)、实际光合效率(ΦPSⅡ)、光化学淬灭系数(qL)和净光合速率(Pn)在不同生育时期均呈下降趋势,而Fo和NPQ则呈升高趋势,作物叶片光系统PSⅡ反应中心遭到破坏,但与干旱敏感性品种(锦杂103和吉杂127)相比,抗旱性品种(吉杂305和锦杂106)表现出较高的Pn、ETR和ΦPSⅡ和qL。干旱胁迫下,抗旱性高粱品种叶片具有较高的净光合速率和电子传递速率是其适应干旱胁迫的主要光合生理基础。4个高粱品种的叶片正面和背面气孔密度、长度、宽度和长宽比,在3个时期对干旱胁迫的响应不同,但均为抗旱性品种气孔状况对干旱胁迫不敏感,而干旱敏感性品种受干旱胁迫影响较大。叶温差、Pn、气体交换参数和气孔性状相关性分析表明,Pn与气孔导度(Gs)、蒸腾速率(Tr)、正面气孔长度和宽度极显著正相关,与叶片背面气孔长度显著相关。叶温差与背面气孔密度极显著正相关,与正面气孔宽度显著正相关。
4、高粱在干旱胁迫下地上部分干重和根系均较对照有所降低,但根冠比升高,并且抗旱性品种升幅较大。与旱敏感性品种相比,在干旱胁迫下,抗旱性品种总根长、表面积、平均直径和根体积均升幅较大,籽粒产量受干旱胁迫影响较小。经灰色关联度分析,高粱叶片Pn、Tr和叶干重等指标与高粱抗旱性关系较密切。
The test was conducted at the experiment station of Shenyang Agricultural University during2011to2012. PEG-6000was used to simulate drought stress environment in artificial climate chamber for studying the effects of different concentrations of PEG-6000on the response of31sorghum cultivars at the seed germination stage. And the purpose of this dissertation is to filter sorghum cultivars with better drought-resistance based on responses of sorghum cultivars at the seed germination stage with different drought stress, and the same time to explore suitable identifying methods for sorghum drought-resistance. Under outdoor potted conditions,4sorghum cultivars with different drought-resistance were used to study relationships among agricultural technology, physiologic and biochemical index and sorghum cultivars drought-resistance, and to reveal drought-resistant mechanism in many aspects. Drought-resistance identification and indicator selection were also studied to supply simple and allowable agricultural technology and physiologic and biochemical index, which could evaluate sorghum drought-resistant efficiently. Meanwhile, the study established sorghum drought-resistant identification indicator system to enrich research content of sorghum drought-resistant. And the study also supplied theory and method of breeding for drought-resistant in sorghum. And the main results were as followed:
1. PEG-6000was used to simulate drought stress environment in artificial climate chamber for studying the effects of different concentrations of PEG-6000(80g/L,120g/L,150g/L,175g/L) on the response of31sorghum cultivars at the seed germination stage, with the objectives of screening drought-resistant cultivars and exploring suitable identifying methods for sorghum's drought-resistance. Through principal component analysis (PCA) and Self-Organizing-Map (SOM) cluster analysis, the drought-resistance of31sorghum cultivars was evaluated. PCA showed that the payloads of relative bud length, relative root length and drought-resistance index during germination had the maximum values, and were used as the main evaluating indexes to identify drought-resistance capability of sorghum, and based on which the drought-resistant capabilities of31sorghum cultivars were ranked. Five groups of drought-resistance capabilities of31sorghum cultivars were divided based on SOM cluster analysis:4cultivars including Jiza305as highly drought-resistant,4cultivars including HL8as drought-resistant,8cultivars including LiaozalO as medium drought-resistant,7cultivars including Jinza103as drought-sensitive and8cultivars including Jinza93as highly drought-sensitive. It was concluded that parameters such as relative bud length, relative root length and germination drought-resistance index can be used as criteria for identifying sorghum's drought-resistance, and that SOM cluster analysis is an important method for classifying drought-resistance.
2. Results from this research indicated that drought tolerant cultivar was comparatively resistant to water stress owing to the greater increased in peroxidase (POD), superoxide dismutase (SOD), and catalase (CAT) activities with lower malondialdehyde (MDA) content. Findings showed that the greater drought tolerant capacity was resulted from the superior ability to lower increase in free radicals along with higher increase in proline, water soluble sugar, and protein. Correlation analysis showed that leaf temperature serves an important role in influencing the sorghum water retention capacity, protective enzyme activity, as well as osmotic adjustment substance, resulting in the differential water stress responses between cultivars. This suggests that the leaf temperature might be useful as a critical criterion in selecting drought resistance sorghum. Additionally, there appears to be potential for using far infrared thermal imaging technology to appraise the capability of sorghum drought-resistant.
3. The maximum photosynthetic efficiency (Fv/Fm), the apparent electron transport rate(ETR), actual photosynthetic efficiency (ΦPSⅡ) photochemical quenching coefficient (qL) and net photosynthetic rate (Pn) of sorghum leaves at different growth stages showed a downward trend, while the initial fluorescence (Fo) and non-photochemical quenching (NPQ) showed an upward trend, Photosystem Ⅱ (PS Ⅱ) reaction center of leaf was destroyed, resulting in a reduction of energy conversion efficiency of primary light. The Pn, ETR,(ΦPSⅡ and relative electron transfer rate (Pm) of drought-resistant cultivars (Jiza305and Jinza106) were higher than those of drought-sensitive cultivars (Jinza103and Jizal27), and their abilities of conversion of light energy were greater. The chlorophyll fluorescence parameters of ΦPSⅡ and relative electron transfer rate (Pm) in sorghum leaves of drought-resistant cultivars are important physiological characteristics for the drought-resistant sorghum cultivars.
4. There had different trends of stomatal density, length, width and length/width on leaf ventral and back of4sorghum cultivars within drought stress at three different growth stages, but all that showed drought-resistant sorghum cultivars were less sensitive to drought stress, and drought sentitive sorghum cultivars were sensitive to drought stress. Correlation analysis indicated Pn was greatly positively correlated with stomatal conductance (Gs), transpiration rate (Tr), stomatal length and width on leaf ventral and back, and significantly positively correlated with length on the back. Meanwhile, leaf temperature different was greatly positively correlated with stomatal density on back and significantly positively correlated with width on ventral.
With drought stress, the dry weights of above-ground and root of sorghum were decreased slightly, but the ratio of that was increased and which was significant in drought-resistant sorghum cultivars. Comparing with drought-sensitive cultivars, there was largely increased in total root length, total surface area, average diameter and root volume of drought-resistant sorghum cultivars and no obvious changes in yield of that with drought stress. The results of grey correlation analysis showed that the physiological characteristics of leaf, such as Pn、Tr and leaf dry weight, were closely related to sorghum drought resistance.
1、采用人工气候箱内培养,PEG-6000溶液模拟干旱胁迫环境,在萌发期以80g/L、120g/L、150g/L、175g/L PEG-6000水溶液处理31个高粱品种,旨在根据高粱品种萌发期对不同干旱胁迫程度的响应,筛选出具有抗旱能力的高粱品种并探讨高粱萌发期抗旱性鉴定的方法。通过主成分分析法(PCA)和神经网络自组织映射(SOM)聚类分析法对各品种进行抗旱性综合分析与评定。PCA结果表明,相对芽长、相对根长和相对萌发抗旱指数载荷量最大,将其作为萌发期高粱抗旱性筛选的主要评价指标,对31个高粱品种抗旱性进行了排序。最后通过SOM聚类分析将31个高粱品种按抗旱性强弱分为5类,吉杂305等4个品种为高度抗旱品种,HL5等4个品种为抗旱品种,辽杂10等8个品种为中等抗旱品种,锦杂103等7个品种为干旱敏感品种,锦杂93等8个品种为高度干旱敏感品种。研究认为,相对芽长、相对根长和相对萌发抗旱指数等指标可以作为高粱品种抗旱性鉴定的重要指标；SOM聚类分析可作为品种抗旱性分类的重要方法。
2、干旱胁迫下,高粱叶片自由水含量、束缚水含量、相对含水量和离体叶片失水速率等水分状况在不同生育时期均呈下降趋势,吉杂305(高度抗旱品种)的下降幅度显著低于吉杂127(高度干旱敏感品种),锦杂106(中等抗旱品种)和锦杂103(干旱敏感品种)介于二者之间。抗旱性品种具有较高的过氧化物酶(POD)、超氧化物歧化酶(SOD)和过氧化氢酶(CAT)活性,而丙二醛(MDA)含量少,具有较好的清除自由基的能力,脯氨酸、可溶性糖和可溶性蛋白含量较高,细胞水势较低。相关性分析表明,叶片温度可以直接或间接的反映干旱胁迫下高粱植株的水分状况及叶片保护性酶和渗透调节物质变化状况,能够体现出高粱抗旱性的强弱。叶温差可以作为高粱抗旱性筛选的一个重要指标,应用远红外热成像技术在高粱抗旱性的鉴定上具有可行性。
3、干旱胁迫下,高粱叶片最大光合效率(Fv/Fm)、表观电子传递速率(ETR)、实际光合效率(ΦPSⅡ)、光化学淬灭系数(qL)和净光合速率(Pn)在不同生育时期均呈下降趋势,而Fo和NPQ则呈升高趋势,作物叶片光系统PSⅡ反应中心遭到破坏,但与干旱敏感性品种(锦杂103和吉杂127)相比,抗旱性品种(吉杂305和锦杂106)表现出较高的Pn、ETR和ΦPSⅡ和qL。干旱胁迫下,抗旱性高粱品种叶片具有较高的净光合速率和电子传递速率是其适应干旱胁迫的主要光合生理基础。4个高粱品种的叶片正面和背面气孔密度、长度、宽度和长宽比,在3个时期对干旱胁迫的响应不同,但均为抗旱性品种气孔状况对干旱胁迫不敏感,而干旱敏感性品种受干旱胁迫影响较大。叶温差、Pn、气体交换参数和气孔性状相关性分析表明,Pn与气孔导度(Gs)、蒸腾速率(Tr)、正面气孔长度和宽度极显著正相关,与叶片背面气孔长度显著相关。叶温差与背面气孔密度极显著正相关,与正面气孔宽度显著正相关。
4、高粱在干旱胁迫下地上部分干重和根系均较对照有所降低,但根冠比升高,并且抗旱性品种升幅较大。与旱敏感性品种相比,在干旱胁迫下,抗旱性品种总根长、表面积、平均直径和根体积均升幅较大,籽粒产量受干旱胁迫影响较小。经灰色关联度分析,高粱叶片Pn、Tr和叶干重等指标与高粱抗旱性关系较密切。
The test was conducted at the experiment station of Shenyang Agricultural University during2011to2012. PEG-6000was used to simulate drought stress environment in artificial climate chamber for studying the effects of different concentrations of PEG-6000on the response of31sorghum cultivars at the seed germination stage. And the purpose of this dissertation is to filter sorghum cultivars with better drought-resistance based on responses of sorghum cultivars at the seed germination stage with different drought stress, and the same time to explore suitable identifying methods for sorghum drought-resistance. Under outdoor potted conditions,4sorghum cultivars with different drought-resistance were used to study relationships among agricultural technology, physiologic and biochemical index and sorghum cultivars drought-resistance, and to reveal drought-resistant mechanism in many aspects. Drought-resistance identification and indicator selection were also studied to supply simple and allowable agricultural technology and physiologic and biochemical index, which could evaluate sorghum drought-resistant efficiently. Meanwhile, the study established sorghum drought-resistant identification indicator system to enrich research content of sorghum drought-resistant. And the study also supplied theory and method of breeding for drought-resistant in sorghum. And the main results were as followed:
1. PEG-6000was used to simulate drought stress environment in artificial climate chamber for studying the effects of different concentrations of PEG-6000(80g/L,120g/L,150g/L,175g/L) on the response of31sorghum cultivars at the seed germination stage, with the objectives of screening drought-resistant cultivars and exploring suitable identifying methods for sorghum's drought-resistance. Through principal component analysis (PCA) and Self-Organizing-Map (SOM) cluster analysis, the drought-resistance of31sorghum cultivars was evaluated. PCA showed that the payloads of relative bud length, relative root length and drought-resistance index during germination had the maximum values, and were used as the main evaluating indexes to identify drought-resistance capability of sorghum, and based on which the drought-resistant capabilities of31sorghum cultivars were ranked. Five groups of drought-resistance capabilities of31sorghum cultivars were divided based on SOM cluster analysis:4cultivars including Jiza305as highly drought-resistant,4cultivars including HL8as drought-resistant,8cultivars including LiaozalO as medium drought-resistant,7cultivars including Jinza103as drought-sensitive and8cultivars including Jinza93as highly drought-sensitive. It was concluded that parameters such as relative bud length, relative root length and germination drought-resistance index can be used as criteria for identifying sorghum's drought-resistance, and that SOM cluster analysis is an important method for classifying drought-resistance.
2. Results from this research indicated that drought tolerant cultivar was comparatively resistant to water stress owing to the greater increased in peroxidase (POD), superoxide dismutase (SOD), and catalase (CAT) activities with lower malondialdehyde (MDA) content. Findings showed that the greater drought tolerant capacity was resulted from the superior ability to lower increase in free radicals along with higher increase in proline, water soluble sugar, and protein. Correlation analysis showed that leaf temperature serves an important role in influencing the sorghum water retention capacity, protective enzyme activity, as well as osmotic adjustment substance, resulting in the differential water stress responses between cultivars. This suggests that the leaf temperature might be useful as a critical criterion in selecting drought resistance sorghum. Additionally, there appears to be potential for using far infrared thermal imaging technology to appraise the capability of sorghum drought-resistant.
3. The maximum photosynthetic efficiency (Fv/Fm), the apparent electron transport rate(ETR), actual photosynthetic efficiency (ΦPSⅡ) photochemical quenching coefficient (qL) and net photosynthetic rate (Pn) of sorghum leaves at different growth stages showed a downward trend, while the initial fluorescence (Fo) and non-photochemical quenching (NPQ) showed an upward trend, Photosystem Ⅱ (PS Ⅱ) reaction center of leaf was destroyed, resulting in a reduction of energy conversion efficiency of primary light. The Pn, ETR,(ΦPSⅡ and relative electron transfer rate (Pm) of drought-resistant cultivars (Jiza305and Jinza106) were higher than those of drought-sensitive cultivars (Jinza103and Jizal27), and their abilities of conversion of light energy were greater. The chlorophyll fluorescence parameters of ΦPSⅡ and relative electron transfer rate (Pm) in sorghum leaves of drought-resistant cultivars are important physiological characteristics for the drought-resistant sorghum cultivars.
4. There had different trends of stomatal density, length, width and length/width on leaf ventral and back of4sorghum cultivars within drought stress at three different growth stages, but all that showed drought-resistant sorghum cultivars were less sensitive to drought stress, and drought sentitive sorghum cultivars were sensitive to drought stress. Correlation analysis indicated Pn was greatly positively correlated with stomatal conductance (Gs), transpiration rate (Tr), stomatal length and width on leaf ventral and back, and significantly positively correlated with length on the back. Meanwhile, leaf temperature different was greatly positively correlated with stomatal density on back and significantly positively correlated with width on ventral.
With drought stress, the dry weights of above-ground and root of sorghum were decreased slightly, but the ratio of that was increased and which was significant in drought-resistant sorghum cultivars. Comparing with drought-sensitive cultivars, there was largely increased in total root length, total surface area, average diameter and root volume of drought-resistant sorghum cultivars and no obvious changes in yield of that with drought stress. The results of grey correlation analysis showed that the physiological characteristics of leaf, such as Pn、Tr and leaf dry weight, were closely related to sorghum drought resistance.
引文
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