油用向日葵对Na~+盐胁迫的生理生化响应及农艺性状表现研究
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摘要
本研究从盐胁迫对油用向日葵种子萌发、农艺性状、生物量的影响及生理生化响应等方面入手,初步明确了油用向日葵耐盐适应性范围和部分耐盐机理。
1 NaCl、NaHCO_3、NaHCO_3+NaCl与温度对发芽势、发芽率的胁迫与协同作用:随温度升高油用向日葵种子发芽势和发芽率显著升高,随盐浓度增加发芽势和发芽率明显降低。NaCl盐浓度达到15.5g/kg,温度在15℃、25℃和35℃时,发芽势分别较对照降低86.1%、75.0%、33.3%;不同温度油用向日葵种子发芽势和发芽率受盐胁迫的降幅表现为15℃＞25℃＞35℃;高盐和高温互作加剧了对油用向日葵胚根和子叶的伤害。盐分对油用向日葵种子发芽势、发芽率的影响依次为NaCl＞NaHCO_3+NaCl＞NaHCO_3。盐胁迫对油用向日葵种子萌发的伤害主要是离子毒害。油用向日葵种子在盐环境中适宜的高温可有效增强种子的发芽势,以此可选择适宜播期,通过缩短种子萌发时间,有效减轻盐胁迫。
2盐胁迫下不同时期不同器官及农艺性状表现:盐胁迫延长了生育期,对生育过程的影响表现为营养生长阶段大于生育生长阶段,成株期对地上部分的影响大于地下部分。随土壤含盐量的增大,油用向日葵的株高、叶面积、茎粗、盘径和粒重显著下降。当盐浓度大于5.5g/kg时根冠比、根系活力急剧下降,根系活跃吸收面积/总吸收面积减小。高盐胁迫改变了油用向日葵的库源关系,最终表现为对经济产量的影响程度大于生物产量;对干物质积累影响为灌浆期＞成熟期＞开花期＞现蕾期＞苗期;当盐浓度超过6.0g/kg时,油用向日葵的生长发育就会受到严重抑制,脱盐效果无经济意义。
3不同NaCl+NaHCO_3浓度和生育阶段的生理生化响应:细胞膜透性苗期高于现蕾期和花期;在盐浓度大于6.0g/kg时膜透性显著增强。幼苗叶片本身的SOD活性很强,在盐浓度7.5g/kg时达最大值。幼苗根系POD酶活性在11.5g/kg时达到最大值,叶片在盐浓度9.5g/kg时达最大值。在盐胁迫下叶绿素含量表现为苗期＞现蕾期＞开花期,在含盐量为6.5g/kg时达到峰值。脯氨酸含量表现为现蕾期＞苗期＞开花期。现蕾期当盐浓度达到8.5g/kg时,脯氨酸的含量从不到100μg/g猛增到1000μg/g,是苗期和花期的十倍。苗期丙二醛的含量在在盐浓度为6.5g/kg时出现最小值后显著升高;现蕾期和花期丙二醛含量在盐浓度为6.0g/kg时出现最小值后显著升高。综合不同盐分区间细胞膜透性、叶绿素含量的出现的最大值和对应脯氨酸、丙二醛含量出现的最小值,可以初步确认:供试油用向日葵品种生理生化指标响应最敏感的土壤盐含量在6.5g/kg左右。
4盐胁迫对含油率及脂肪酸组成的影响:盐胁迫显著抑制了子实和籽仁油份积累,对籽仁含油率的影响大于子实含油率。随土壤含盐量的增加,抑制油酸的生成,促进亚油酸的生成;含盐量对脂肪酸组成的影响表现为棕榈酸＞油酸＞亚油酸＞硬脂酸＞亚麻酸。
5生理生化响应与抗盐机制:随盐浓度的增加细胞膜透性显著增强,半透性功能受损,造成叶绿素外渗,植株中Na~+含量增加、K~+含量减少、K~+/Na~+值降低、离子毒害加重,脯氨酸和丙二醛含量显著增加,相关正常代谢平衡与生理生化功能被破坏,产生盐分的各种次生伤害。油用向日葵在盐胁迫下受活性氧伤害后的自我修复能力很强,这主要是油用向日葵本身含有能消除盐胁迫过程中产生的有害物质-氧自由基、从而对细胞起保护作用、增强油用向日葵抗盐功能的SOD和POD酶活性比较强,特别是各时期SOD的酶活性强是提高供试油用向日葵品种抗盐能力的关键。
6粗灰分含量与脱盐效果:植株中Na~+含量是根系Na~+含量的2-6倍,而且这种差值随盐浓度的增加进一步在拉大,说明油葵对土壤盐分的携出主要是通过植株带走的。不同生育期植株中粗灰分含量从高到低依次为苗期＞现蕾期＞开花期＞成熟期＞灌浆期。当盐浓度超过6.0g/kg时,油用向日葵的生长发育就会受到严重抑制,脱盐效果无经济意义。所以,要想有一个好的脱盐效果就必须保证其地上部有一定的生物量。
In order to define the adaptability range of salt tolerance and oil-sunflower mechanism, study the effects of salt stress on germination of seeds,main economic character, agronomical character,biological yield,and physiological and biochemical indexes of oil-sunflower.
1.NaCl、NaHCO_3、NaHCO_3+NaCl and the temperature exert the stress and collaborated effect on the germination trend and germination rate of oil-sunflower: With the temperature increased,the seed germination trend and germination rate significantly increased,and with the increase of concentrations the seed germination trend and germination rate significantly dropped.The germination trend decreased by 86.1%, 75.0%and 33.3%compared with the contrast while the temperature was 15℃,25℃and 35℃.The germination trend and germination rate of oil-flower seed under the different temperature were harmed by the salt stress,while the order of the temperature was 15℃＞25℃＞35℃.The high temperature combined high salt concentrations increased the harm of the radicel length and fresh weight of cotyledon.The germination trend and germination rate of oil-flower seed were harmed under the salt stress,and the sequence of damage degree was NaCl＞NaHCO_3+NaCl＞NaHCO_3.The main reason of harm by salt stress on the germination of oil-sunflower seed was ion toxicity.Under salt environment,the suitable temperature can effectively increase the germination trend.Thus,the suitable sowing time can be chosed and alleviate the salt stress through shortening the germination time.
2.The effects of the salt stress on different organs in different period:The salt stress significantly prolonged procreation period,the asexuality growing stage had more bigger effects on procreation course than on the sexual reproduction growing stage,and the salt stress had higher effect on the ground parts than on the under ground parts in adult period. With the increase of salt concentrations,plant height,leaf area,stem wide,branches diameters,and grain weight significantly decreased.When the salt concentrations exceeded 5.5g/kg,the ration of the root weight to the shoot weight and the activity of roots began to drop significantly,and ratio of living absorbance area to the total absorbance area of roots reduced.The high salt stress could change the sink-source relation,and more significantly affected the economical yield than the biomass yield.The high salt stress can also differently affected the dry matter at different growing stage,and the sequence of growing stage was filling>maturity>flowering>budding>seedling stage.When the soil salt content exceeded 6.0g/kg,the growth of oil-sunflower could be restrained seriously.
3.The effect of NaCl+NaHCO_3 dealt with physiological and biochemical indexes under different salt range at different procreation:The cell membrane permeability in seedling stage was higher than that in squaring and flowering period,and when the salt concentrations exceed 6.0g/kg,the cell membrane permeability significantly increased.The SOD activity of leaf in oil-sunflower seedling was strong,reaching maximum at the salt concentration of 7.5 g/kg.The salt stress could influence the POD activity of root and leaf differently,and the POD activity of root peaked at the salt concentrations of 9.5g/kg,and the POD activity peaked at the salt concentrations of 11.5g/kg.The salt stress can differently affect the chlorophyll content of sprout at different growing stage,and the sequence of growing stage was seedling>budding>flowering period,and the chlorophyll content peaked at salt concentration of 6.5 g/kg.The salt stress can differently affect proline content at different growing stage,and the sequence of growing stage was budding>seedling>flowering.The proline content quickly increased from 100μg/g to 1000μg/g at the salt concentrations of 8.5g/kg in squaring period,which is ten times that of in seedling and flower period.In the salt content of 6.5g/kg,the MDA content was minimum in seedling period,then began to significantly increase;however,in squaring and flowering period,the MDA content was minimum at the content of 6.0g/kg,and then began to significantly increase.
Based on a comprehensive analysis of the maximum value of cell membrane permeability,chlorophyll content,the corresponding minimum value of proline,and malonaldehyde content corresponding in different salinity zone,we can conclude that the most sensitive range soil salt content of oil-sunflower was about 6.5g/kg.
4.Effects of salt stress on oil content and fatty acid of oil-sunflower:The salt stress significantly restrained the accumulation of seed and kernel;moreover,the influence to oil content of kernel was greater than did oil content of seed.With the increase of salt concentrations,the production of oleic acid was restrained,and linoleic acid was improved. The sequence of influence of soil salt content on fatty acid was:palmitic acid>oleic acid> linoleic acid> Itearic acid> linolenic acid.
5.The effects of salt stress on the main physiological and biochemical of oil-sunflower:
Cell membrane permeability significantly increased with the concentration of salt increasing,and the translucency function was so damaged as to make chlorophyll extravasated,the content of Na~+ increased,K~+ decreased,metabolic balance damaged,ion toxicity aggravated,and diversified secondary injuries induced,oil-sunflower self-restoration capability increased under salt stress,mainly because oil-flower contained oxygen free radical which could eliminate harmful substance during salt stress period so that the substance had a protective effect on the cells,and increased SOD and POD activity, especially in each stage.
6.The crude ash content of plant and desalting efficiency of oil-sunflower:
The content of Na~+ in plant was 2～6 times than that of in the root systems and the difference enlarged with the increase of salt concentration,which showed that the loss of soil salt was mainly through oil-sunflower plant absorbed.The sequence of the ash content of oil-sunflower at different growing stage was seedling period＞budding＞flowering＞maturity＞filling period.When the soil salt concentration exceeded 6.0g/kg,the oil-sunflower growth could be significantly restricted,and the effect of desalting had no economical meaning.So if we want to obtain better desalting effects,we must ensure a certain number of biomass-yield on the ground part of oil-sunflower.
1 NaCl、NaHCO_3、NaHCO_3+NaCl与温度对发芽势、发芽率的胁迫与协同作用:随温度升高油用向日葵种子发芽势和发芽率显著升高,随盐浓度增加发芽势和发芽率明显降低。NaCl盐浓度达到15.5g/kg,温度在15℃、25℃和35℃时,发芽势分别较对照降低86.1%、75.0%、33.3%;不同温度油用向日葵种子发芽势和发芽率受盐胁迫的降幅表现为15℃＞25℃＞35℃;高盐和高温互作加剧了对油用向日葵胚根和子叶的伤害。盐分对油用向日葵种子发芽势、发芽率的影响依次为NaCl＞NaHCO_3+NaCl＞NaHCO_3。盐胁迫对油用向日葵种子萌发的伤害主要是离子毒害。油用向日葵种子在盐环境中适宜的高温可有效增强种子的发芽势,以此可选择适宜播期,通过缩短种子萌发时间,有效减轻盐胁迫。
2盐胁迫下不同时期不同器官及农艺性状表现:盐胁迫延长了生育期,对生育过程的影响表现为营养生长阶段大于生育生长阶段,成株期对地上部分的影响大于地下部分。随土壤含盐量的增大,油用向日葵的株高、叶面积、茎粗、盘径和粒重显著下降。当盐浓度大于5.5g/kg时根冠比、根系活力急剧下降,根系活跃吸收面积/总吸收面积减小。高盐胁迫改变了油用向日葵的库源关系,最终表现为对经济产量的影响程度大于生物产量;对干物质积累影响为灌浆期＞成熟期＞开花期＞现蕾期＞苗期;当盐浓度超过6.0g/kg时,油用向日葵的生长发育就会受到严重抑制,脱盐效果无经济意义。
3不同NaCl+NaHCO_3浓度和生育阶段的生理生化响应:细胞膜透性苗期高于现蕾期和花期;在盐浓度大于6.0g/kg时膜透性显著增强。幼苗叶片本身的SOD活性很强,在盐浓度7.5g/kg时达最大值。幼苗根系POD酶活性在11.5g/kg时达到最大值,叶片在盐浓度9.5g/kg时达最大值。在盐胁迫下叶绿素含量表现为苗期＞现蕾期＞开花期,在含盐量为6.5g/kg时达到峰值。脯氨酸含量表现为现蕾期＞苗期＞开花期。现蕾期当盐浓度达到8.5g/kg时,脯氨酸的含量从不到100μg/g猛增到1000μg/g,是苗期和花期的十倍。苗期丙二醛的含量在在盐浓度为6.5g/kg时出现最小值后显著升高;现蕾期和花期丙二醛含量在盐浓度为6.0g/kg时出现最小值后显著升高。综合不同盐分区间细胞膜透性、叶绿素含量的出现的最大值和对应脯氨酸、丙二醛含量出现的最小值,可以初步确认:供试油用向日葵品种生理生化指标响应最敏感的土壤盐含量在6.5g/kg左右。
4盐胁迫对含油率及脂肪酸组成的影响:盐胁迫显著抑制了子实和籽仁油份积累,对籽仁含油率的影响大于子实含油率。随土壤含盐量的增加,抑制油酸的生成,促进亚油酸的生成;含盐量对脂肪酸组成的影响表现为棕榈酸＞油酸＞亚油酸＞硬脂酸＞亚麻酸。
5生理生化响应与抗盐机制:随盐浓度的增加细胞膜透性显著增强,半透性功能受损,造成叶绿素外渗,植株中Na~+含量增加、K~+含量减少、K~+/Na~+值降低、离子毒害加重,脯氨酸和丙二醛含量显著增加,相关正常代谢平衡与生理生化功能被破坏,产生盐分的各种次生伤害。油用向日葵在盐胁迫下受活性氧伤害后的自我修复能力很强,这主要是油用向日葵本身含有能消除盐胁迫过程中产生的有害物质-氧自由基、从而对细胞起保护作用、增强油用向日葵抗盐功能的SOD和POD酶活性比较强,特别是各时期SOD的酶活性强是提高供试油用向日葵品种抗盐能力的关键。
6粗灰分含量与脱盐效果:植株中Na~+含量是根系Na~+含量的2-6倍,而且这种差值随盐浓度的增加进一步在拉大,说明油葵对土壤盐分的携出主要是通过植株带走的。不同生育期植株中粗灰分含量从高到低依次为苗期＞现蕾期＞开花期＞成熟期＞灌浆期。当盐浓度超过6.0g/kg时,油用向日葵的生长发育就会受到严重抑制,脱盐效果无经济意义。所以,要想有一个好的脱盐效果就必须保证其地上部有一定的生物量。
In order to define the adaptability range of salt tolerance and oil-sunflower mechanism, study the effects of salt stress on germination of seeds,main economic character, agronomical character,biological yield,and physiological and biochemical indexes of oil-sunflower.
1.NaCl、NaHCO_3、NaHCO_3+NaCl and the temperature exert the stress and collaborated effect on the germination trend and germination rate of oil-sunflower: With the temperature increased,the seed germination trend and germination rate significantly increased,and with the increase of concentrations the seed germination trend and germination rate significantly dropped.The germination trend decreased by 86.1%, 75.0%and 33.3%compared with the contrast while the temperature was 15℃,25℃and 35℃.The germination trend and germination rate of oil-flower seed under the different temperature were harmed by the salt stress,while the order of the temperature was 15℃＞25℃＞35℃.The high temperature combined high salt concentrations increased the harm of the radicel length and fresh weight of cotyledon.The germination trend and germination rate of oil-flower seed were harmed under the salt stress,and the sequence of damage degree was NaCl＞NaHCO_3+NaCl＞NaHCO_3.The main reason of harm by salt stress on the germination of oil-sunflower seed was ion toxicity.Under salt environment,the suitable temperature can effectively increase the germination trend.Thus,the suitable sowing time can be chosed and alleviate the salt stress through shortening the germination time.
2.The effects of the salt stress on different organs in different period:The salt stress significantly prolonged procreation period,the asexuality growing stage had more bigger effects on procreation course than on the sexual reproduction growing stage,and the salt stress had higher effect on the ground parts than on the under ground parts in adult period. With the increase of salt concentrations,plant height,leaf area,stem wide,branches diameters,and grain weight significantly decreased.When the salt concentrations exceeded 5.5g/kg,the ration of the root weight to the shoot weight and the activity of roots began to drop significantly,and ratio of living absorbance area to the total absorbance area of roots reduced.The high salt stress could change the sink-source relation,and more significantly affected the economical yield than the biomass yield.The high salt stress can also differently affected the dry matter at different growing stage,and the sequence of growing stage was filling>maturity>flowering>budding>seedling stage.When the soil salt content exceeded 6.0g/kg,the growth of oil-sunflower could be restrained seriously.
3.The effect of NaCl+NaHCO_3 dealt with physiological and biochemical indexes under different salt range at different procreation:The cell membrane permeability in seedling stage was higher than that in squaring and flowering period,and when the salt concentrations exceed 6.0g/kg,the cell membrane permeability significantly increased.The SOD activity of leaf in oil-sunflower seedling was strong,reaching maximum at the salt concentration of 7.5 g/kg.The salt stress could influence the POD activity of root and leaf differently,and the POD activity of root peaked at the salt concentrations of 9.5g/kg,and the POD activity peaked at the salt concentrations of 11.5g/kg.The salt stress can differently affect the chlorophyll content of sprout at different growing stage,and the sequence of growing stage was seedling>budding>flowering period,and the chlorophyll content peaked at salt concentration of 6.5 g/kg.The salt stress can differently affect proline content at different growing stage,and the sequence of growing stage was budding>seedling>flowering.The proline content quickly increased from 100μg/g to 1000μg/g at the salt concentrations of 8.5g/kg in squaring period,which is ten times that of in seedling and flower period.In the salt content of 6.5g/kg,the MDA content was minimum in seedling period,then began to significantly increase;however,in squaring and flowering period,the MDA content was minimum at the content of 6.0g/kg,and then began to significantly increase.
Based on a comprehensive analysis of the maximum value of cell membrane permeability,chlorophyll content,the corresponding minimum value of proline,and malonaldehyde content corresponding in different salinity zone,we can conclude that the most sensitive range soil salt content of oil-sunflower was about 6.5g/kg.
4.Effects of salt stress on oil content and fatty acid of oil-sunflower:The salt stress significantly restrained the accumulation of seed and kernel;moreover,the influence to oil content of kernel was greater than did oil content of seed.With the increase of salt concentrations,the production of oleic acid was restrained,and linoleic acid was improved. The sequence of influence of soil salt content on fatty acid was:palmitic acid>oleic acid> linoleic acid> Itearic acid> linolenic acid.
5.The effects of salt stress on the main physiological and biochemical of oil-sunflower:
Cell membrane permeability significantly increased with the concentration of salt increasing,and the translucency function was so damaged as to make chlorophyll extravasated,the content of Na~+ increased,K~+ decreased,metabolic balance damaged,ion toxicity aggravated,and diversified secondary injuries induced,oil-sunflower self-restoration capability increased under salt stress,mainly because oil-flower contained oxygen free radical which could eliminate harmful substance during salt stress period so that the substance had a protective effect on the cells,and increased SOD and POD activity, especially in each stage.
6.The crude ash content of plant and desalting efficiency of oil-sunflower:
The content of Na~+ in plant was 2～6 times than that of in the root systems and the difference enlarged with the increase of salt concentration,which showed that the loss of soil salt was mainly through oil-sunflower plant absorbed.The sequence of the ash content of oil-sunflower at different growing stage was seedling period＞budding＞flowering＞maturity＞filling period.When the soil salt concentration exceeded 6.0g/kg,the oil-sunflower growth could be significantly restricted,and the effect of desalting had no economical meaning.So if we want to obtain better desalting effects,we must ensure a certain number of biomass-yield on the ground part of oil-sunflower.
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