高寒地区嫁接桑光合特性及其对低温的响应
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摘要
本文以实生桑“泰来”和嫁接桑树(砧木：泰来桑；接穗：龙桑)为材料,在自然条件下、低温锻炼、低温胁迫等环境下,测定了光合参数、荧光参数、抗氧化酶活性等指标,研究了实生桑和嫁接桑光系统低温环境下的响应特性,以及适应能力,主要结果如下：
(1)自然生长条件下,通过嫁接桑和实生桑光响应曲线以及二氧化碳响应曲线显示,嫁接桑与实生桑的净光合速率均呈上升的趋势,嫁接桑各个叶位的Pn值都高于实生桑,差异显著。嫁接桑利用光能的效率高于实生桑,光合机构的能力及有机物的合成效率也要高于实生桑。嫁接桑气孔导度(Gs)各个叶位光合活性要高于实生桑,在光合及蒸腾等生理活动中的能力很强,体内的代谢情况也优于实生桑。
(2)自然生长条件下,在同一枝条不同叶位以及不同叶位上嫁接桑的实际化学效率(ΦPSⅡ)电子传递速率(ETR)、光化学猝灭(qP)均高于实生桑树,非光化学淬灭(NPQ)低于实生桑。从而说明嫁接桑PSⅡ反应中心开放比例和光化学效率以及电子传递速率、吸收光能用于光合作用合成有机物能力高于实生桑,易于有机物的大量合成。
(3)以桑树品种秋雨为试验材料,研究了桑树幼苗在低温锻炼、冷胁迫和常温恢复期间的光合作用和抗氧化酶活性的适应性变化。结果表明,12℃3d低温锻炼明显地诱导了桑树幼苗的抗冷性。3℃3d冷胁迫下,12℃3d低温锻炼后的桑树幼苗叶片净光合速率(Pn)、气孔导度(Gs)和(PSⅡ)最大光化学效率(Fv／Fm)明显高于未经低温锻炼的桑树幼苗,而且在常温下的恢复快于未经低温锻炼的桑树幼苗。在12℃3d低温锻炼和3℃3d冷胁迫期间,桑树幼苗叶片脯氨酸和可溶性糖含量明显增加,而经低温锻炼的叶片丙二醛(MDA)含量明显低于未经低温锻炼的桑树幼苗。经低温锻炼的桑树幼苗叶片抗坏血酸氧化酶(APX)活性明显高于未经低温锻炼的桑树幼苗。渗透调节物质含量增加和APX活性提高在低温锻炼诱导桑树幼苗的抗冷性上发挥重要的作用。
(4)以实生桑和嫁接桑为材料,利用叶绿素荧光技术,研究了低温胁迫对两种桑树样本的叶片叶绿素荧光参数的影响以及抗氧化酶的响应。结果表明：在低温胁迫下,2个桑树品种叶绿素荧光参数发生了明显变化,嫁接桑和实生桑的实际光化学效率(ΦPSⅡ)、表观电子传递速率(ETR)、PSⅡ最大光化学效率(Fv/Fm)、光化学猝灭系数(qP)均下降,初始荧光(F0)与非光化学猝灭系数(NPQ)上升,但嫁接桑的F0上升幅度比实生桑树小,实生桑树NPQ上升幅度比嫁接桑树大,干旱胁迫不仅直接引发植物光合结构的变化,而且也影响光合电子的传递。嫁接桑的SOD、POD保护酶的活性下降幅度略小于实生桑,MDA和可溶性糖上升的幅度略高,从而嫁接桑体现出了较强的抗低温能力。
This study use own-root mulberry and grafted mulberry (stock:Tailai mulberry; Scion:Long mulberry) as materials, under the natural conditions, cold hardening, low temperature stress and so on, measured photosynthetic parameters, fluorescence,antioxidant activity and other indicators, research own-root mulberry and grafted mulberry photosynthetic system response and adaptability under low temperature environment, all the main results as follows:
(1) Under natural conditions, grafting mulberry and own-root mulberry light response curve and the carbon dioxide response curves showed that grafted mulberry and own-root mulberry net photosynthetic rate showed an upward trend, all grafted mulberry leaf were higher than the own-root mulberry obvious. grafted mulberry the efficient of using light energy than own-root mulberry, the ability of photosynthetic institutions and the efficiency of organic synthesis have higher than own-root mulberry. Stomatal conductance (Gs) of grafted mulberry is higher than own-root mulberry in each leaves, photosynthesis and transpiration in the physiological activities are strong, the metabolism is better than own-root mulberry.
(2) Under natural conditions, in different leaf positions of the same branch and different position of leaves,grafted mulberry is higher than own-root mulberry on the actual photochemical efficiency of PSⅡ, electron transport rate (ETR), the photochemical quenching coefficient (qP) and non photochemical quenching (NPQ) is lower than grafted mulberry. It is show that the open ratio of PSⅡreaction centers of grafted mulberry and photochemical efficiency and electron transport rate, synthesis of organic compounds absorb light energy for photosynthesis capacity than own-root mulberry, and composing a large number of organic synthesis easily.
(3) Cold temperature damage is a common problem for mulberry production in Northeast China. Many attempts have been made to improve cold tolerance in mulberry plants, and one of the methods tested is cold hardening. The objectives of this research in this paper are to investigate cold tolerance in mulberry seedlings induced by cold hardening to identify mechanisms of chilling tolerance. The seedlings of mulberry variety 'Qiuyu' were exposed to 12℃(cold hardened) or 25℃(unhardened) for 3 d under 220～250μmol·m-2·s-1 PDF. All seedlings were then transported to cold stress at 3℃for 3 d and allowed to recover at 25℃for 2 d. The results showed that resistance to cold stress in leaves of mulberry seedlings was induced by cold hardening at 12℃for 3 d. Net photosynthetic rate (Pn), stomatal conductance (Gs) and the maximum photochemical efficiency of photosystemⅡ(Fv/Fm) in cold hardened leaves of mulberry seedlings under cold stress at 3℃for 3 d were higher than that of unhardened leaves, and cold hardened leaves of mulberry seedlings also recovered faster from cold stress than unhardened leaves. Proline contents and soluble sugar contents in leaves of mulberry seedlings during cold hardening, and proline contents and soluble sugar contents as cell osmolytes in cold hardened leaves of mulberry seedlings during cold stress and recovery were higher significantly than that of unhardened leaves. Malondialdehyde (MDA) contents in cold hardened leaves of mulberry seedlings during cold stress and recovery were lower significantly than that of unhardened leaves. Ascorbate peroxidase (APX, EC1.11.1.11) activities in cold hardened leaves of mulberry seedlings during cold hardening, cold stress and recovery were higher significantly than that of unhardened leaves. It concluded that osmolytes and APX activities in leaves of cold hardened leaves of mulberry seedlings played important role in cold tolerance induced by cold hardening.
(4) Using the own-root mulberry and grafts mulberry as the test material,the propose was to study on the effects of low temperature stress on chlorophyll fluorescence parameters of mulberry.The results showed that under the low temperature stress,the chlorophyll fluorescence in own-root mulberry and grafts mulberry showed obviously change. Both of them actual photochemical efficiency of PSⅡ(ΦPSⅡ),the electron transport rate (ETR),the diurnal changes of the maximum PSⅡphotochemical efficiency(Fv/Fm) and the photochemical quenching coefficient (qP) were decrease intensity, Minimal fluores-cence (Fo) and non photochemical quenching (NPQ) was risen, while the rising extent of NPQ in own-root mulberry was higher than that in grafts mulberry,It is shown that the ability of PSⅡresisting low temperature stress in grafts mulberry than that in own-root mulberry. Low temperature stress not only caused the change of photosynthetic structure in mulberry directly, but also affect the transformation of photosynthetic electron.The descend content of SOD and POD in grafts mulberry smaller than own-root mulberry,and the ascend content of MDA and oluble sugar contents in grafts mulberry higher than own-root mulberry, it is shown that grafts mulberry have better ability of cold tolerance.
(1)自然生长条件下,通过嫁接桑和实生桑光响应曲线以及二氧化碳响应曲线显示,嫁接桑与实生桑的净光合速率均呈上升的趋势,嫁接桑各个叶位的Pn值都高于实生桑,差异显著。嫁接桑利用光能的效率高于实生桑,光合机构的能力及有机物的合成效率也要高于实生桑。嫁接桑气孔导度(Gs)各个叶位光合活性要高于实生桑,在光合及蒸腾等生理活动中的能力很强,体内的代谢情况也优于实生桑。
(2)自然生长条件下,在同一枝条不同叶位以及不同叶位上嫁接桑的实际化学效率(ΦPSⅡ)电子传递速率(ETR)、光化学猝灭(qP)均高于实生桑树,非光化学淬灭(NPQ)低于实生桑。从而说明嫁接桑PSⅡ反应中心开放比例和光化学效率以及电子传递速率、吸收光能用于光合作用合成有机物能力高于实生桑,易于有机物的大量合成。
(3)以桑树品种秋雨为试验材料,研究了桑树幼苗在低温锻炼、冷胁迫和常温恢复期间的光合作用和抗氧化酶活性的适应性变化。结果表明,12℃3d低温锻炼明显地诱导了桑树幼苗的抗冷性。3℃3d冷胁迫下,12℃3d低温锻炼后的桑树幼苗叶片净光合速率(Pn)、气孔导度(Gs)和(PSⅡ)最大光化学效率(Fv／Fm)明显高于未经低温锻炼的桑树幼苗,而且在常温下的恢复快于未经低温锻炼的桑树幼苗。在12℃3d低温锻炼和3℃3d冷胁迫期间,桑树幼苗叶片脯氨酸和可溶性糖含量明显增加,而经低温锻炼的叶片丙二醛(MDA)含量明显低于未经低温锻炼的桑树幼苗。经低温锻炼的桑树幼苗叶片抗坏血酸氧化酶(APX)活性明显高于未经低温锻炼的桑树幼苗。渗透调节物质含量增加和APX活性提高在低温锻炼诱导桑树幼苗的抗冷性上发挥重要的作用。
(4)以实生桑和嫁接桑为材料,利用叶绿素荧光技术,研究了低温胁迫对两种桑树样本的叶片叶绿素荧光参数的影响以及抗氧化酶的响应。结果表明：在低温胁迫下,2个桑树品种叶绿素荧光参数发生了明显变化,嫁接桑和实生桑的实际光化学效率(ΦPSⅡ)、表观电子传递速率(ETR)、PSⅡ最大光化学效率(Fv/Fm)、光化学猝灭系数(qP)均下降,初始荧光(F0)与非光化学猝灭系数(NPQ)上升,但嫁接桑的F0上升幅度比实生桑树小,实生桑树NPQ上升幅度比嫁接桑树大,干旱胁迫不仅直接引发植物光合结构的变化,而且也影响光合电子的传递。嫁接桑的SOD、POD保护酶的活性下降幅度略小于实生桑,MDA和可溶性糖上升的幅度略高,从而嫁接桑体现出了较强的抗低温能力。
This study use own-root mulberry and grafted mulberry (stock:Tailai mulberry; Scion:Long mulberry) as materials, under the natural conditions, cold hardening, low temperature stress and so on, measured photosynthetic parameters, fluorescence,antioxidant activity and other indicators, research own-root mulberry and grafted mulberry photosynthetic system response and adaptability under low temperature environment, all the main results as follows:
(1) Under natural conditions, grafting mulberry and own-root mulberry light response curve and the carbon dioxide response curves showed that grafted mulberry and own-root mulberry net photosynthetic rate showed an upward trend, all grafted mulberry leaf were higher than the own-root mulberry obvious. grafted mulberry the efficient of using light energy than own-root mulberry, the ability of photosynthetic institutions and the efficiency of organic synthesis have higher than own-root mulberry. Stomatal conductance (Gs) of grafted mulberry is higher than own-root mulberry in each leaves, photosynthesis and transpiration in the physiological activities are strong, the metabolism is better than own-root mulberry.
(2) Under natural conditions, in different leaf positions of the same branch and different position of leaves,grafted mulberry is higher than own-root mulberry on the actual photochemical efficiency of PSⅡ, electron transport rate (ETR), the photochemical quenching coefficient (qP) and non photochemical quenching (NPQ) is lower than grafted mulberry. It is show that the open ratio of PSⅡreaction centers of grafted mulberry and photochemical efficiency and electron transport rate, synthesis of organic compounds absorb light energy for photosynthesis capacity than own-root mulberry, and composing a large number of organic synthesis easily.
(3) Cold temperature damage is a common problem for mulberry production in Northeast China. Many attempts have been made to improve cold tolerance in mulberry plants, and one of the methods tested is cold hardening. The objectives of this research in this paper are to investigate cold tolerance in mulberry seedlings induced by cold hardening to identify mechanisms of chilling tolerance. The seedlings of mulberry variety 'Qiuyu' were exposed to 12℃(cold hardened) or 25℃(unhardened) for 3 d under 220～250μmol·m-2·s-1 PDF. All seedlings were then transported to cold stress at 3℃for 3 d and allowed to recover at 25℃for 2 d. The results showed that resistance to cold stress in leaves of mulberry seedlings was induced by cold hardening at 12℃for 3 d. Net photosynthetic rate (Pn), stomatal conductance (Gs) and the maximum photochemical efficiency of photosystemⅡ(Fv/Fm) in cold hardened leaves of mulberry seedlings under cold stress at 3℃for 3 d were higher than that of unhardened leaves, and cold hardened leaves of mulberry seedlings also recovered faster from cold stress than unhardened leaves. Proline contents and soluble sugar contents in leaves of mulberry seedlings during cold hardening, and proline contents and soluble sugar contents as cell osmolytes in cold hardened leaves of mulberry seedlings during cold stress and recovery were higher significantly than that of unhardened leaves. Malondialdehyde (MDA) contents in cold hardened leaves of mulberry seedlings during cold stress and recovery were lower significantly than that of unhardened leaves. Ascorbate peroxidase (APX, EC1.11.1.11) activities in cold hardened leaves of mulberry seedlings during cold hardening, cold stress and recovery were higher significantly than that of unhardened leaves. It concluded that osmolytes and APX activities in leaves of cold hardened leaves of mulberry seedlings played important role in cold tolerance induced by cold hardening.
(4) Using the own-root mulberry and grafts mulberry as the test material,the propose was to study on the effects of low temperature stress on chlorophyll fluorescence parameters of mulberry.The results showed that under the low temperature stress,the chlorophyll fluorescence in own-root mulberry and grafts mulberry showed obviously change. Both of them actual photochemical efficiency of PSⅡ(ΦPSⅡ),the electron transport rate (ETR),the diurnal changes of the maximum PSⅡphotochemical efficiency(Fv/Fm) and the photochemical quenching coefficient (qP) were decrease intensity, Minimal fluores-cence (Fo) and non photochemical quenching (NPQ) was risen, while the rising extent of NPQ in own-root mulberry was higher than that in grafts mulberry,It is shown that the ability of PSⅡresisting low temperature stress in grafts mulberry than that in own-root mulberry. Low temperature stress not only caused the change of photosynthetic structure in mulberry directly, but also affect the transformation of photosynthetic electron.The descend content of SOD and POD in grafts mulberry smaller than own-root mulberry,and the ascend content of MDA and oluble sugar contents in grafts mulberry higher than own-root mulberry, it is shown that grafts mulberry have better ability of cold tolerance.
引文
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