向日葵对碱胁迫和盐胁迫适应机制比较
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摘要
土壤碱化问题日益突出,并且其对环境的破坏力远远大于土壤盐化。目前,不同层面的研究均表明碱胁迫和盐胁迫是两种性质截然不同的胁迫,这两种胁迫对植物的伤害作用不同,而且植物对二者的适应机制也明显不同。向日葵是土壤碱化后唯一可种植的油料作物,但是至今对其抗盐碱性,尤其是抗碱机制未见有系统的报道。本文采用了不同于以往的室内实验路线,以期更贴近生产实际。在人工模拟天然盐碱生境的条件下,播种抗碱性较强的向日葵种子白葵杂六号,通过分析盐胁迫、碱胁迫和盐碱混合胁迫下从种子萌发到幼苗形成及幼苗生长等不同生理阶段的生理响应,来探讨盐胁迫和碱胁迫对向日葵的作用机制以及向日葵对其适应机制。其主要实验结果及结论如下:
1盐、碱胁迫对向日葵种子萌发和幼苗形成的影响
从种子萌发到成苗阶段对植物适应盐碱环境至关重要,甚至在一定程度上决定着植物能否在盐碱生境定居和繁衍后代。我们的结果表明:在适度的胁迫强度下(≤120mM),盐胁迫仅延迟向日葵出苗时间,而并未影响出苗率和成苗率;但当胁迫强度超过120mM,盐胁迫则导致出苗率及成苗率急剧下降。碱胁迫并未明显延迟向日葵出苗时间和降低出苗率,但却导致成苗率急剧下降,甚至在60mM碱胁迫强度下,幼苗全部死亡。这说明碱胁迫对植物不同生理阶段的影响明显不同。外界高pH并不影响种子的吸水,但当胚根突破种皮后,高pH则会直接作用于根细胞,破坏其结构和功能,导致萌发的胚内有毒离子急剧增加,使其受害而难以长成正常的幼苗。这说明,碱胁迫所造成的高pH是限制成苗的主要因素。
2盐、碱胁迫对向日葵幼苗生长和光合作用的影响
虽然部分幼苗能够在盐胁迫和碱胁迫下存活,但光合和生长都不同程度地受到抑制,而且碱胁迫的抑制程度明显大于相同盐度的盐胁迫。碱胁迫下干重、叶面积、株高、净光合速率、气孔导度、蒸腾速率的下降程度都明显大于盐胁迫下。通过对光合电子传递过程(叶绿素荧光参数)深入分析发现,盐胁迫并未使光合系统Ⅱ受到损伤,而碱胁迫则明显破坏光合系统Ⅱ,降低光合电子传递效率,这可能就是向日葵净光合速率在碱胁迫下明显低于盐胁迫的主要原因。
3盐、碱胁迫对向日葵矿质营养的影响
盐、碱胁迫下向日葵体内矿质元素积累明显不同。盐胁迫下氮、磷、硫、铁等元素的含量非但没下降反而有所升高;而盐碱混合胁迫下,几乎所有矿质营养元素的含量均明显降低。碱胁迫下的高pH导致矿质营养水平的急剧下降可能是碱胁迫对生长和光合抑制作用甚于盐胁迫的主要原因之一。
4盐、碱胁迫下向日葵体内渗透调节及离子平衡机制
本文比较了各个溶质在胁迫条件下对渗透调节的贡献。结果表明K+是向日葵地上部分主要的无机渗透调节物质,而可溶性糖则是向日葵体内主要的有机渗透调节剂。我们推测向日葵根中可能有一个特殊的控制Na+、K+转运机制,抑制Na+向茎叶转移,而促进K+向上传输,使叶片保持一个高K+低Na+的稳态,这可能是向日葵相对耐盐碱性的主要原因。碱胁迫特别是高碱胁迫会导致细胞内离子严重失衡、pH不稳定,向日葵可能通过提高有机酸合成来维持体内离子平衡和pH稳定。
5盐、碱胁迫下向日葵体内DNA甲基化响应特点
实验结果表明,向日葵叶片的DNA甲基化水平明显高于根,并且根和叶的DNA甲基化位点都集中于CG/CHG。通过对根、叶DNA甲基化变异率分析发现,两种胁迫下,向日葵根中DNA甲基化变异率均明显高于叶,而且无论是根还是叶,甲基化变化都集中在CG/CHG位点。根系由于直接接触各种土壤胁迫,会率先感受并传递胁迫信号,并且会迅速做出适应性响应,根中DNA甲基化变异的模式不同于叶,其变异率也明显高于叶,说明DNA甲基化可能直接参与根中胁迫信号的产生和传递,同时也表明:盐碱胁迫下根部所进行的DNA水平上的调节过程不仅比地上部分更直接更迅速,也更复杂。此外,不论根还是叶,碱胁迫下DNA甲基化变异率明显大于盐胁迫,这从表观遗传学层面进一步证明盐胁迫和碱胁迫是两种性质不同的胁迫,不但作用机制不同,而且植物对其适应机制也明显不同。根的调节作用是向日葵抗碱的关键所在,DNA甲基化在这一调节过程中可能起重要作用,但具体机制有待进一步研究。
Soil alkalization is becoming increasingly prominent, and its destructive power is greater than soil salinization. Salt stress and alkali stress are two kinds of stresses not only in aspects of their effects on plant, but also in the response of plant to them. Sunflower is the only oil crop under alkaline condition, but the tolerance, especially, the mechanisms of alkali-tolerant have not been reported systematically. In this paper, our experiment formation is different from the previous laboratory experiment, to be closer to natural conditions. In this study, we firstly simulated saline and alkaline conditions, and then the sunflower seeds-baikuiza 6-were sowed and stressed under salt stress and alkali stress. We analyzed the effects of salt stress, alkali stress and salt-alkaline mixed stress on sunflower and eco-physiological adaptive mechanisms of sunflower to its salt-aklaline habitat. Major experiment results and conclusions were as follows:
1. Effects of salt stress and alkali stress on seed germination and seedling survival in sunflower
It is a critical statge of plant from seed germination and seedling growth to adapting to saline and alkaline environment, and to some exent which decide whether plant can live and reproduce or not. Our experiment showed the emergence time was delayed under salt stress when≤120mM,but the emergence rate and seedling survival rate did not decrease significantly. With increasing stress intensity, the emergence rate and seedling survival rate delayed signigicantly. Under alkali stress, the emergence rate and emergence time did not change, but the seedling survival rate decreased sharply, even when salinity was 60 mM,the seedlings were all died under alkali stress. This indicated the effects were significantly different among different growth stages under alkali stress. High pH did not interfere with seed water absorption, but was directly on the root cells when radicel borke through seed coat and destroyed their stucture and function, and led to increase in toxic ions, making it difficult to seedlings. This showed high pH caused by alkali stress was main factor to limiting seedlings.
2. Effects of salt stress and alkali stress on growth and photosynthesis in sunflower
Although some seedlings could survive under salt stress and alkali stress, photosynthesis and growth were inhibited, and the inhibition was greater under alkali stress than that under salt stress. The decreases of dry matter, leaf area, height, net photosynthetic rates, stomatal conductance and transpiration rates under alkali stress were greater than those under salt stress. By analyzing the photosynthetic electron transfer processes (chlorophyll fluorescence parameters), we found alkali stress damaged the photosynthetic systemⅡ, and reduced the efficiency of photosynthetic electron transport, but salt stress did not. These data indicated that high pH damaged the photosynthetic systemⅡand interferred photosynthetic electron transport, this might be the main reason that PN was lower under alkali stress than that under salt stress.
3. Effects of salt stress and alkali stress on mineral nutrition in sunflower
The experimental results indicated that the accumulation of mineral elements was markedly different under salt and alkali stresses. The contents of N, P, S and Fe did not decrease, but increased under salt stress; while contents of almost all mineral nutrients declined under salt-alkaline mixed stress. It was clear that mineral nutrition deficits was a main reason that the inhibitions of growth and photosynthesis under alkali stress were more severe than those under salt stress.
4. Effects of salt stress and alkali stress on osmotic adjustment and ion balance in sunflower
We compared the contributions of various osmolytes under stress. The results showed K+ was the main inorganic osmolyte in shoots, but the solute sugar was the main organic osmolyte in sunflower. At moderate stress, it may be a special mechanism which controls Na+,K+ transport in the roots of sunflower. Sunflower roots could control Na+ but promote K+ transfer to leaves, maintaining low Na+ high K+ status of leaves. This may be the main reason that sunflower is the relative salt and alkali-tolerant crop. Alkali stress especially high alkali stress could cause ion imbalance and pH instability, and sunflower synthesized organic acids to keep ion balance and adjust pH.
5. The response characteristics of DNA methylation in sunflower under salt stress and alkali stress
The results showed the DNA methylation in leaves were much higher than that in roots and the simultaneous CG/CHG methylation was the main pattern. Under two stresses, the roots had greater variation than that in leaves. Moreover, the methylation variation of roots and leaves were both concentrated in CG/CHG sites. Because the roots have contact with various soil stresses directly, and can feel and transport the signals, and then make the adaptive responses quickly. DNA methylation patterns in roots were different from in leaves, and the variation was significantly greater than that in leaves. Those proved that DNA methylation might be directly involved the signals generation and transminssion in roots and the regulation of the roots on DNA level was faster and more complex than that leaves. In addition, the DNA methylation variations in roots and leaves under alkali stress were both significantly greater than those under alkali stress, this proved salt stress and alkali stress were two kinds of stresses once again. The regulation of sunflower roots is a key in alkali-tolerant, and DNA methylation plays an important role in the regulation processes, but the mechanisms deserve further investigation.
1盐、碱胁迫对向日葵种子萌发和幼苗形成的影响
从种子萌发到成苗阶段对植物适应盐碱环境至关重要,甚至在一定程度上决定着植物能否在盐碱生境定居和繁衍后代。我们的结果表明:在适度的胁迫强度下(≤120mM),盐胁迫仅延迟向日葵出苗时间,而并未影响出苗率和成苗率;但当胁迫强度超过120mM,盐胁迫则导致出苗率及成苗率急剧下降。碱胁迫并未明显延迟向日葵出苗时间和降低出苗率,但却导致成苗率急剧下降,甚至在60mM碱胁迫强度下,幼苗全部死亡。这说明碱胁迫对植物不同生理阶段的影响明显不同。外界高pH并不影响种子的吸水,但当胚根突破种皮后,高pH则会直接作用于根细胞,破坏其结构和功能,导致萌发的胚内有毒离子急剧增加,使其受害而难以长成正常的幼苗。这说明,碱胁迫所造成的高pH是限制成苗的主要因素。
2盐、碱胁迫对向日葵幼苗生长和光合作用的影响
虽然部分幼苗能够在盐胁迫和碱胁迫下存活,但光合和生长都不同程度地受到抑制,而且碱胁迫的抑制程度明显大于相同盐度的盐胁迫。碱胁迫下干重、叶面积、株高、净光合速率、气孔导度、蒸腾速率的下降程度都明显大于盐胁迫下。通过对光合电子传递过程(叶绿素荧光参数)深入分析发现,盐胁迫并未使光合系统Ⅱ受到损伤,而碱胁迫则明显破坏光合系统Ⅱ,降低光合电子传递效率,这可能就是向日葵净光合速率在碱胁迫下明显低于盐胁迫的主要原因。
3盐、碱胁迫对向日葵矿质营养的影响
盐、碱胁迫下向日葵体内矿质元素积累明显不同。盐胁迫下氮、磷、硫、铁等元素的含量非但没下降反而有所升高;而盐碱混合胁迫下,几乎所有矿质营养元素的含量均明显降低。碱胁迫下的高pH导致矿质营养水平的急剧下降可能是碱胁迫对生长和光合抑制作用甚于盐胁迫的主要原因之一。
4盐、碱胁迫下向日葵体内渗透调节及离子平衡机制
本文比较了各个溶质在胁迫条件下对渗透调节的贡献。结果表明K+是向日葵地上部分主要的无机渗透调节物质,而可溶性糖则是向日葵体内主要的有机渗透调节剂。我们推测向日葵根中可能有一个特殊的控制Na+、K+转运机制,抑制Na+向茎叶转移,而促进K+向上传输,使叶片保持一个高K+低Na+的稳态,这可能是向日葵相对耐盐碱性的主要原因。碱胁迫特别是高碱胁迫会导致细胞内离子严重失衡、pH不稳定,向日葵可能通过提高有机酸合成来维持体内离子平衡和pH稳定。
5盐、碱胁迫下向日葵体内DNA甲基化响应特点
实验结果表明,向日葵叶片的DNA甲基化水平明显高于根,并且根和叶的DNA甲基化位点都集中于CG/CHG。通过对根、叶DNA甲基化变异率分析发现,两种胁迫下,向日葵根中DNA甲基化变异率均明显高于叶,而且无论是根还是叶,甲基化变化都集中在CG/CHG位点。根系由于直接接触各种土壤胁迫,会率先感受并传递胁迫信号,并且会迅速做出适应性响应,根中DNA甲基化变异的模式不同于叶,其变异率也明显高于叶,说明DNA甲基化可能直接参与根中胁迫信号的产生和传递,同时也表明:盐碱胁迫下根部所进行的DNA水平上的调节过程不仅比地上部分更直接更迅速,也更复杂。此外,不论根还是叶,碱胁迫下DNA甲基化变异率明显大于盐胁迫,这从表观遗传学层面进一步证明盐胁迫和碱胁迫是两种性质不同的胁迫,不但作用机制不同,而且植物对其适应机制也明显不同。根的调节作用是向日葵抗碱的关键所在,DNA甲基化在这一调节过程中可能起重要作用,但具体机制有待进一步研究。
Soil alkalization is becoming increasingly prominent, and its destructive power is greater than soil salinization. Salt stress and alkali stress are two kinds of stresses not only in aspects of their effects on plant, but also in the response of plant to them. Sunflower is the only oil crop under alkaline condition, but the tolerance, especially, the mechanisms of alkali-tolerant have not been reported systematically. In this paper, our experiment formation is different from the previous laboratory experiment, to be closer to natural conditions. In this study, we firstly simulated saline and alkaline conditions, and then the sunflower seeds-baikuiza 6-were sowed and stressed under salt stress and alkali stress. We analyzed the effects of salt stress, alkali stress and salt-alkaline mixed stress on sunflower and eco-physiological adaptive mechanisms of sunflower to its salt-aklaline habitat. Major experiment results and conclusions were as follows:
1. Effects of salt stress and alkali stress on seed germination and seedling survival in sunflower
It is a critical statge of plant from seed germination and seedling growth to adapting to saline and alkaline environment, and to some exent which decide whether plant can live and reproduce or not. Our experiment showed the emergence time was delayed under salt stress when≤120mM,but the emergence rate and seedling survival rate did not decrease significantly. With increasing stress intensity, the emergence rate and seedling survival rate delayed signigicantly. Under alkali stress, the emergence rate and emergence time did not change, but the seedling survival rate decreased sharply, even when salinity was 60 mM,the seedlings were all died under alkali stress. This indicated the effects were significantly different among different growth stages under alkali stress. High pH did not interfere with seed water absorption, but was directly on the root cells when radicel borke through seed coat and destroyed their stucture and function, and led to increase in toxic ions, making it difficult to seedlings. This showed high pH caused by alkali stress was main factor to limiting seedlings.
2. Effects of salt stress and alkali stress on growth and photosynthesis in sunflower
Although some seedlings could survive under salt stress and alkali stress, photosynthesis and growth were inhibited, and the inhibition was greater under alkali stress than that under salt stress. The decreases of dry matter, leaf area, height, net photosynthetic rates, stomatal conductance and transpiration rates under alkali stress were greater than those under salt stress. By analyzing the photosynthetic electron transfer processes (chlorophyll fluorescence parameters), we found alkali stress damaged the photosynthetic systemⅡ, and reduced the efficiency of photosynthetic electron transport, but salt stress did not. These data indicated that high pH damaged the photosynthetic systemⅡand interferred photosynthetic electron transport, this might be the main reason that PN was lower under alkali stress than that under salt stress.
3. Effects of salt stress and alkali stress on mineral nutrition in sunflower
The experimental results indicated that the accumulation of mineral elements was markedly different under salt and alkali stresses. The contents of N, P, S and Fe did not decrease, but increased under salt stress; while contents of almost all mineral nutrients declined under salt-alkaline mixed stress. It was clear that mineral nutrition deficits was a main reason that the inhibitions of growth and photosynthesis under alkali stress were more severe than those under salt stress.
4. Effects of salt stress and alkali stress on osmotic adjustment and ion balance in sunflower
We compared the contributions of various osmolytes under stress. The results showed K+ was the main inorganic osmolyte in shoots, but the solute sugar was the main organic osmolyte in sunflower. At moderate stress, it may be a special mechanism which controls Na+,K+ transport in the roots of sunflower. Sunflower roots could control Na+ but promote K+ transfer to leaves, maintaining low Na+ high K+ status of leaves. This may be the main reason that sunflower is the relative salt and alkali-tolerant crop. Alkali stress especially high alkali stress could cause ion imbalance and pH instability, and sunflower synthesized organic acids to keep ion balance and adjust pH.
5. The response characteristics of DNA methylation in sunflower under salt stress and alkali stress
The results showed the DNA methylation in leaves were much higher than that in roots and the simultaneous CG/CHG methylation was the main pattern. Under two stresses, the roots had greater variation than that in leaves. Moreover, the methylation variation of roots and leaves were both concentrated in CG/CHG sites. Because the roots have contact with various soil stresses directly, and can feel and transport the signals, and then make the adaptive responses quickly. DNA methylation patterns in roots were different from in leaves, and the variation was significantly greater than that in leaves. Those proved that DNA methylation might be directly involved the signals generation and transminssion in roots and the regulation of the roots on DNA level was faster and more complex than that leaves. In addition, the DNA methylation variations in roots and leaves under alkali stress were both significantly greater than those under alkali stress, this proved salt stress and alkali stress were two kinds of stresses once again. The regulation of sunflower roots is a key in alkali-tolerant, and DNA methylation plays an important role in the regulation processes, but the mechanisms deserve further investigation.
引文
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