乙酸及NaCl和Na_2CO_3胁迫对莱茵衣藻的影响
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摘要
目前，水体酸化已成为当今世界广为关注的环境问题之一，除无机酸外，有机酸也是引起环境酸化的重要原因，其中乙酸是有机酸的主要成分，广泛存在于自然界中，其含量可高达毫摩尔水平，对藻类植物的生长具有巨大的潜在威胁。除酸化外，干旱和半干旱地区的水体由于过度蒸发而存在不同程度的盐碱化，所以藻类植物还面临着盐碱化的影响。因此，鉴于酸化和盐碱化正日趋严重，日益影响着水域生态系统平衡与环境安全，本文以单细胞模式植物莱茵衣藻为实验材料，研究了乙酸、NaCl和Na_2CO_3胁迫对其生长的影响、与光呼吸代谢的相互关系，同时对乙酸诱导莱茵衣藻发生PCD进行了深入研究，并测定了此过程中释放的VOCs，探讨了其信号作用，此外还就NaCl和Na_2CO_3胁迫诱导VOCs释放以及NaCl胁迫诱导光呼吸途径产生ROS进行了研究。结果如下：
1. pH5.0乙酸、300mM NaCl以及25mM、50mM和150mM Na_2CO_3胁迫均会引起莱茵衣藻光合色素降解，并且此种降解作用与光照条件无关。除叶绿素降解外，乙酸、NaCl和Na_2CO_3胁迫还使莱茵衣藻的光合特性发生改变，Fv/Fm、Y(II)和ETR均发生了不同程度的降低；低强度胁迫使qP增强，而高强度胁迫则降低了qP；在一定时间内，所有胁迫处理均使NPQ增强。
2.乙酸、NaCl和Na_2CO_3胁迫均能诱导莱茵衣藻迅速产生大量的O_2ˉ.和H_2O_2，当乙酸胁迫10min时，两种ROS含量均达到最大值；当NaCl和Na_2CO_3胁迫30min时，ROS含量达到最大值。在ROS含量升高的同时，细胞内SOD、POD和CAT活性也发生相应变化，其中低强度胁迫使这些酶的活性得到了不同程度的提高。
3.莱茵衣藻在乙酸、NaCl和Na_2CO_3胁迫处理后，对照株系CW-Arg及SGAT敲减株系T和A的SGAT酶活性均显著增强，Ser和Gly含量升高，并且随NaCl胁迫强度增强，酶活性进一步增强、两种氨基酸含量进一步升高。虽然在正常以及乙酸、NaCl和Na_2CO_3胁迫条件下，T和A株系的生长状况均弱于CW-Arg，但是在胁迫条件下，T和A株系受到的影响明显大于对照株系。这表明藻类植物的光呼吸途径与其抗逆性密切相关，逆境胁迫促进藻类植物的光呼吸途径运转，而光呼吸途径受阻后，藻类植物抵抗逆境胁迫的能力明显降低。
4.采用抑制剂SHAM阻断乙醇酸-醌氧化还原酶系统或MOA阻断甘氨酸脱羧酶后，莱茵衣藻细胞内Ser含量明显降低，Gly含量明显升高，这表明其光呼吸途径的正常运转已受到明显影响。在NaCl胁迫条件下，经抑制剂处理的细胞所产生的ROS均明显少于未用抑制剂处理的细胞，其中SHAM处理后ROS含量最低，这表明SHAM阻断的反应即乙醇酸生成乙醛酸可能为光呼吸途径中ROS的产生位点。由于MOA处理阻断了甘氨酸向丝氨酸的转化，因此也影响了乙醇酸转化为乙醛酸，所以其ROS产量低于对照，但高于SHAM处理。采用NaCl胁迫SGAT敲减效果较强的A株系，由于丝氨酸向甘氨酸的转化被阻断，所以其ROS产生量也明显低于对照株系CW-Arg。这些结果表明，藻类植物的光呼吸途径也会导致ROS产生，其产生位点是叶绿体中乙醇酸经乙醇酸-醌氧化还原酶系统催化形成乙醛酸的反应。
5.莱茵衣藻不能直接吸收利用外源L-Ser和L-Lys，而是通过胞外脱氨氧化的方式进行利用，并且此种脱氨过程依赖于氮饥饿，同时乙酸能促进此过程的进行。莱茵衣藻可利用L-Cys作为唯一氮源，其吸收量随胞外浓度增加或饲喂时间延长而增加，并且其被动扩散吸收过程不影响细胞对L-Leu的被动扩散吸收。
6. pH5.0乙酸胁迫莱茵衣藻30min后，其DNA开始降解，在胁迫6h时DNA降解为150-350bp的片段。ZnSO_4处理对此降解过程具有明显的抑制作用，并且1mM ZnSO_4的抑制作用明显强于0.5mM。在pH5.0乙酸胁迫1h和2h时，分别有10%和95%的莱茵衣藻细胞内出现呈TUNEL阳性染色的细胞核，在胁迫4h时所有的细胞内均出现此细胞核，这表明pH5.0乙酸可诱导莱茵衣藻细胞发生PCD。
7.乙酸（34.3mM）在pH5.0时可诱导莱茵衣藻细胞发生PCD，而H_2SO_4、HCl、H_3PO_4、磷酸缓冲液和乳酸在pH_3.0时才能诱导细胞死亡，这表明溶液酸度不是诱导PCD的主要原因，而与乙酸自身的性质有关。当溶液中含有50mM乙酸，pH为6.0时，细胞死亡；然而当乙酸浓度增加至150mM，pH为7.0时，细胞则不死亡，在此情况下，溶液中的乙酸主要以乙酸根离子形式存在，因此，细胞死亡与溶液中的乙酸根离子浓度无关，而与溶液的pH值和乙酸分子浓度同时相关。
8.在pH5.0乙酸诱导莱茵衣藻发生PCD的过程中，VOCs大量释放，其中在2h时释放量最大，并且成分最多，主要包括有烷烃类、烯烃类、醛类、醇类、酮类、酯类和萜烯类等34种化合物。此外，300mM NaCl和150mM Na_2CO_3胁迫莱茵衣藻2h也可诱导其释放大量的VOCs。乙酸和NaCl胁迫均可诱导己醛（GLVs）释放，其峰面积分别为23.49×10~7和3.14×10~7；而Na_2CO_3胁迫则不能诱导己醛释放，其诱导的特异性成分是3,4-二甲基-己烷和5-甲基-2-庚烯。长叶烯是此三种胁迫诱导释放的主要萜烯类化合物，其峰面积分别为5.82×10~7（乙酸）、3.11×10~7（NaCl）和4.00×10~7（Na_2CO_3）。
9.将乙酸、NaCl和Na_2CO_3胁迫诱导的莱茵衣藻VOCs通入正常生长的莱茵衣藻溶液中，其细胞生长受到明显抑制，而其叶绿素含量、Fv/Fm、H_2O_2含量和抗氧化酶活性均不同程度增加，这表明非生物胁迫诱导的VOCs可能具有在藻类植物间进行信息传递的作用。
At present, water body acidification is one of the most dramaticallyenvironmental problems in the world. Besides inorganic acids, organic acids are otherreasons for the acidification, and acetic acid is the dominant component of organicacids. It widely spreads in nature, and its content can accumulate up to millimolarconcentrations, so it is a tremendous potential danger for the growth of algae. Besidesthat, water body in arid and semi-arid regions is becoming salinization mainly causedby high evaporation, so the algae often face this stress. As the acidification andsalinization are seriously affecting the ecosystem balance and environment safety inwater body, we investigated the effects of acetic acid, NaCl and Na_2CO_3stresses onthe growth of Chlamydomonas reinhardtii, a model organism of single cell, and therelationship between those stresses and photorespiration of the alga. We furtherstudied the induction of PCD by acetic acid in C. reinhardtii cells and release ofVOCs during the cell death. The possible role of VOCs as a signal was alsoinvestigated. Meanwhile, we investigated the VOCs from the cells stressed by NaCland Na_2CO_3, too. We finally tried to find whether ROS could be produced by theoperation of photorespiration under NaCl stress. The results are summarized asfollows:
1. The degradation of photosynthetic pigments in C. reinhardtii cells was foundunder the stress of acetic acid at pH5.0, of NaCl at300mM, and of Na_2CO_3at25mM,50mM or150mM, and the degradation was independent of light condition. Thephotosynthetic characters were also changed by acetic acid, NaCl and Na_2CO_3stresses, and the Fv/Fm, Y(II) and ETR were declined in some degree. The qP wasincreased under lower strength stresses, but was decreased under stronger stresses.The NPQ was increased under all stresses in a certain time.
2. The stresses of acetic acid, NaCl and Na_2CO_3induced the burst of O_2ˉ.andH_2O_2in C. reinhardtii cells, and the burst happened at10min under acetic acid stressand at30min under NaCl and Na_2CO_3stresses. The activities of SOD, POD and CATwere also changed with the increase of ROS content, and all raised in some degree under lower strength stresses.
3. In both the cell line of control (CW-Arg) and the SGAT gene knocked downcell lines of C. reinhardtii (line T and A), the SGAT activity, as well as the content ofSer and Gly were increased under salt stress, and the increase was accompanied withthe stress strengthening. The growth rate of T and A cell lines was lower than that ofCW-Arg (control) under normal and stress conditions, however, the stress effects ofacetic acid, NaCl and Na_2CO_3on the growth of T and A was stronger than that ofCW-Arg. Those indicated that alga photorespiration closely related with its capabilityof anti-stress. The stress promoted the photorespiration in algae, and their capabilityof stress resistance was markedly declined after the photorespiration was affected.
4. When the glycolate-quinone oxidoreductase system was inhibited by SHAMor glycine decarboxylase was inhibit by MOA, the content of Ser in C. reinhardtiicells was significantly reduced, but Gly was significantly increased, indicating thatthe photorespiration was markedly affected. Under NaCl stress, ROS content in thecells treated with the two inhibitors was dramatically lower than that in the untreatedcells, with the SHAM treated cells being the lowest. This suggested that the reactioninhibited by SHAM might be the ROS production site. Thus, the decrease in ROScontent after inhibition of glycine decarboxylase by MOA might also result from thereduction of the conversion of glycolate to glyoxylate, which was caused by theinhibition of its downstreem reaction, the conversion of glycine to serine. Because thephotorespiration pathway in A cell line was markedly inhibited in the conversion ofserine to glycine, its ROS content was also significantly lower than that in CW-Arg(cell line of control) under NaCl stress. Those results demonstrated that ROS was alsoproduced during the operation of photorespiration in this alga, and that the productionsite was the conversion of glycolate to glyoxylate catalyzed by glycolate-quinoneoxidoreductase system.
5. The usage of extracellular L-Ser and L-Lys by C. reinhardtii cells was not viadirect absorption, but via extracellular deamination. The deamination was dependenton nitrogen starvation, and acetic acid can promote this process. C. reinhardtii cellscan use L-Cys as the sole nitrogen source, and its intracellular content was increasedwith the increase of extracellular content and prolonging the feeding time, indicating that L-Cys was taken up via passive diffusion. The uptake of L-Cys was not affectedby Leu, indicating that their absorptions are independent.
6. Acetic acid at pH5.0induced DNA degradation after30min treatment, and thedegradation was strengthened gradually during the treatment and finally resulted infragments of about150-350bp after6h treatment. The degradation of DNA can bedelayed by ZnSO_4treatment, and the pretreatment with1mM had greater inhibitioneffect on DNA degradation than with0.5mM. The cells treated with pH5.0aceticacid had TUNEL positive nuclei, and the positive nuclei appeared in10%,95%and100%cells after1h,2h and4h treatment, respectively. Those results suggested thatpH5.0acetic acid induced PCD in C. reinhardtii cells.
7. Acetic acid (34.3mM) at pH5.0induced PCD in C. reinhardtii cells, whileH_2SO_4, HCl, H_3PO_4, phosphate buffer and propionic acid did not, but killed the cellsat pH_3.0, indicating that the induction of PCD did not mainly relate with the mediumacidity, but related with its chemical property of acetic acid. When the cells were keptin the culture medium containing50mM acetic acid and at pH6.0, they were alsokilled. However, the cells could survive in the medium containing150mM acetic acidand at pH7.0, at which most of the acetic acid molecules are in ionic form. Thisdemonstrated that there was no relationship between the cell death and the CH_3COOˉform in the medium. The results showed that PCD induced by acetic acid was mostprobably related with both pH and CH_3COOH form.
8. Lots of VOCs were released from the C. reinhardtii cells in PCD induced byacetic acid, and their components and content reached the highest level after2htreatment. At least34different compounds were determined and are among thealdehydes, alcohols, esters, ketones, alkanes, alkenes and terpenoids. Besides aceticacid,300mM NaCl or150mM Na_2CO_3stress also induced VOCs release from C.reinhardtii cells. Hexanal (GLVs) was induced by both acetic acid and NaCl stresses,with the peak areas of23.49×10~7and3.14×10~7, respectively. However, Na_2CO_3stressdid not induce hexanal, but induced special components of3,4-dimethyl-hexane and5-methyl-2-heptene. Longifolene was the main terpenoid under the3kinds of stresses,with peak areas of5.82×10~7,3.11×10~7and4.00×10~7, respectively, for acetic acid,NaCl and Na_2CO_3.
9. When normal C. reinhardtii cells were exposed to those VOCs from the cellsstressed by acetic acid, NaCl or Na_2CO_3, the cell growth was significantly inhibited,but the chlorophyll content, Fv/Fm, H_2O_2content and antioxidant enzyme activitieswere significantly increased, indicating that VOCs from algae under abiotic stress cantransfer information between algae.
1. pH5.0乙酸、300mM NaCl以及25mM、50mM和150mM Na_2CO_3胁迫均会引起莱茵衣藻光合色素降解，并且此种降解作用与光照条件无关。除叶绿素降解外，乙酸、NaCl和Na_2CO_3胁迫还使莱茵衣藻的光合特性发生改变，Fv/Fm、Y(II)和ETR均发生了不同程度的降低；低强度胁迫使qP增强，而高强度胁迫则降低了qP；在一定时间内，所有胁迫处理均使NPQ增强。
2.乙酸、NaCl和Na_2CO_3胁迫均能诱导莱茵衣藻迅速产生大量的O_2ˉ.和H_2O_2，当乙酸胁迫10min时，两种ROS含量均达到最大值；当NaCl和Na_2CO_3胁迫30min时，ROS含量达到最大值。在ROS含量升高的同时，细胞内SOD、POD和CAT活性也发生相应变化，其中低强度胁迫使这些酶的活性得到了不同程度的提高。
3.莱茵衣藻在乙酸、NaCl和Na_2CO_3胁迫处理后，对照株系CW-Arg及SGAT敲减株系T和A的SGAT酶活性均显著增强，Ser和Gly含量升高，并且随NaCl胁迫强度增强，酶活性进一步增强、两种氨基酸含量进一步升高。虽然在正常以及乙酸、NaCl和Na_2CO_3胁迫条件下，T和A株系的生长状况均弱于CW-Arg，但是在胁迫条件下，T和A株系受到的影响明显大于对照株系。这表明藻类植物的光呼吸途径与其抗逆性密切相关，逆境胁迫促进藻类植物的光呼吸途径运转，而光呼吸途径受阻后，藻类植物抵抗逆境胁迫的能力明显降低。
4.采用抑制剂SHAM阻断乙醇酸-醌氧化还原酶系统或MOA阻断甘氨酸脱羧酶后，莱茵衣藻细胞内Ser含量明显降低，Gly含量明显升高，这表明其光呼吸途径的正常运转已受到明显影响。在NaCl胁迫条件下，经抑制剂处理的细胞所产生的ROS均明显少于未用抑制剂处理的细胞，其中SHAM处理后ROS含量最低，这表明SHAM阻断的反应即乙醇酸生成乙醛酸可能为光呼吸途径中ROS的产生位点。由于MOA处理阻断了甘氨酸向丝氨酸的转化，因此也影响了乙醇酸转化为乙醛酸，所以其ROS产量低于对照，但高于SHAM处理。采用NaCl胁迫SGAT敲减效果较强的A株系，由于丝氨酸向甘氨酸的转化被阻断，所以其ROS产生量也明显低于对照株系CW-Arg。这些结果表明，藻类植物的光呼吸途径也会导致ROS产生，其产生位点是叶绿体中乙醇酸经乙醇酸-醌氧化还原酶系统催化形成乙醛酸的反应。
5.莱茵衣藻不能直接吸收利用外源L-Ser和L-Lys，而是通过胞外脱氨氧化的方式进行利用，并且此种脱氨过程依赖于氮饥饿，同时乙酸能促进此过程的进行。莱茵衣藻可利用L-Cys作为唯一氮源，其吸收量随胞外浓度增加或饲喂时间延长而增加，并且其被动扩散吸收过程不影响细胞对L-Leu的被动扩散吸收。
6. pH5.0乙酸胁迫莱茵衣藻30min后，其DNA开始降解，在胁迫6h时DNA降解为150-350bp的片段。ZnSO_4处理对此降解过程具有明显的抑制作用，并且1mM ZnSO_4的抑制作用明显强于0.5mM。在pH5.0乙酸胁迫1h和2h时，分别有10%和95%的莱茵衣藻细胞内出现呈TUNEL阳性染色的细胞核，在胁迫4h时所有的细胞内均出现此细胞核，这表明pH5.0乙酸可诱导莱茵衣藻细胞发生PCD。
7.乙酸（34.3mM）在pH5.0时可诱导莱茵衣藻细胞发生PCD，而H_2SO_4、HCl、H_3PO_4、磷酸缓冲液和乳酸在pH_3.0时才能诱导细胞死亡，这表明溶液酸度不是诱导PCD的主要原因，而与乙酸自身的性质有关。当溶液中含有50mM乙酸，pH为6.0时，细胞死亡；然而当乙酸浓度增加至150mM，pH为7.0时，细胞则不死亡，在此情况下，溶液中的乙酸主要以乙酸根离子形式存在，因此，细胞死亡与溶液中的乙酸根离子浓度无关，而与溶液的pH值和乙酸分子浓度同时相关。
8.在pH5.0乙酸诱导莱茵衣藻发生PCD的过程中，VOCs大量释放，其中在2h时释放量最大，并且成分最多，主要包括有烷烃类、烯烃类、醛类、醇类、酮类、酯类和萜烯类等34种化合物。此外，300mM NaCl和150mM Na_2CO_3胁迫莱茵衣藻2h也可诱导其释放大量的VOCs。乙酸和NaCl胁迫均可诱导己醛（GLVs）释放，其峰面积分别为23.49×10~7和3.14×10~7；而Na_2CO_3胁迫则不能诱导己醛释放，其诱导的特异性成分是3,4-二甲基-己烷和5-甲基-2-庚烯。长叶烯是此三种胁迫诱导释放的主要萜烯类化合物，其峰面积分别为5.82×10~7（乙酸）、3.11×10~7（NaCl）和4.00×10~7（Na_2CO_3）。
9.将乙酸、NaCl和Na_2CO_3胁迫诱导的莱茵衣藻VOCs通入正常生长的莱茵衣藻溶液中，其细胞生长受到明显抑制，而其叶绿素含量、Fv/Fm、H_2O_2含量和抗氧化酶活性均不同程度增加，这表明非生物胁迫诱导的VOCs可能具有在藻类植物间进行信息传递的作用。
At present, water body acidification is one of the most dramaticallyenvironmental problems in the world. Besides inorganic acids, organic acids are otherreasons for the acidification, and acetic acid is the dominant component of organicacids. It widely spreads in nature, and its content can accumulate up to millimolarconcentrations, so it is a tremendous potential danger for the growth of algae. Besidesthat, water body in arid and semi-arid regions is becoming salinization mainly causedby high evaporation, so the algae often face this stress. As the acidification andsalinization are seriously affecting the ecosystem balance and environment safety inwater body, we investigated the effects of acetic acid, NaCl and Na_2CO_3stresses onthe growth of Chlamydomonas reinhardtii, a model organism of single cell, and therelationship between those stresses and photorespiration of the alga. We furtherstudied the induction of PCD by acetic acid in C. reinhardtii cells and release ofVOCs during the cell death. The possible role of VOCs as a signal was alsoinvestigated. Meanwhile, we investigated the VOCs from the cells stressed by NaCland Na_2CO_3, too. We finally tried to find whether ROS could be produced by theoperation of photorespiration under NaCl stress. The results are summarized asfollows:
1. The degradation of photosynthetic pigments in C. reinhardtii cells was foundunder the stress of acetic acid at pH5.0, of NaCl at300mM, and of Na_2CO_3at25mM,50mM or150mM, and the degradation was independent of light condition. Thephotosynthetic characters were also changed by acetic acid, NaCl and Na_2CO_3stresses, and the Fv/Fm, Y(II) and ETR were declined in some degree. The qP wasincreased under lower strength stresses, but was decreased under stronger stresses.The NPQ was increased under all stresses in a certain time.
2. The stresses of acetic acid, NaCl and Na_2CO_3induced the burst of O_2ˉ.andH_2O_2in C. reinhardtii cells, and the burst happened at10min under acetic acid stressand at30min under NaCl and Na_2CO_3stresses. The activities of SOD, POD and CATwere also changed with the increase of ROS content, and all raised in some degree under lower strength stresses.
3. In both the cell line of control (CW-Arg) and the SGAT gene knocked downcell lines of C. reinhardtii (line T and A), the SGAT activity, as well as the content ofSer and Gly were increased under salt stress, and the increase was accompanied withthe stress strengthening. The growth rate of T and A cell lines was lower than that ofCW-Arg (control) under normal and stress conditions, however, the stress effects ofacetic acid, NaCl and Na_2CO_3on the growth of T and A was stronger than that ofCW-Arg. Those indicated that alga photorespiration closely related with its capabilityof anti-stress. The stress promoted the photorespiration in algae, and their capabilityof stress resistance was markedly declined after the photorespiration was affected.
4. When the glycolate-quinone oxidoreductase system was inhibited by SHAMor glycine decarboxylase was inhibit by MOA, the content of Ser in C. reinhardtiicells was significantly reduced, but Gly was significantly increased, indicating thatthe photorespiration was markedly affected. Under NaCl stress, ROS content in thecells treated with the two inhibitors was dramatically lower than that in the untreatedcells, with the SHAM treated cells being the lowest. This suggested that the reactioninhibited by SHAM might be the ROS production site. Thus, the decrease in ROScontent after inhibition of glycine decarboxylase by MOA might also result from thereduction of the conversion of glycolate to glyoxylate, which was caused by theinhibition of its downstreem reaction, the conversion of glycine to serine. Because thephotorespiration pathway in A cell line was markedly inhibited in the conversion ofserine to glycine, its ROS content was also significantly lower than that in CW-Arg(cell line of control) under NaCl stress. Those results demonstrated that ROS was alsoproduced during the operation of photorespiration in this alga, and that the productionsite was the conversion of glycolate to glyoxylate catalyzed by glycolate-quinoneoxidoreductase system.
5. The usage of extracellular L-Ser and L-Lys by C. reinhardtii cells was not viadirect absorption, but via extracellular deamination. The deamination was dependenton nitrogen starvation, and acetic acid can promote this process. C. reinhardtii cellscan use L-Cys as the sole nitrogen source, and its intracellular content was increasedwith the increase of extracellular content and prolonging the feeding time, indicating that L-Cys was taken up via passive diffusion. The uptake of L-Cys was not affectedby Leu, indicating that their absorptions are independent.
6. Acetic acid at pH5.0induced DNA degradation after30min treatment, and thedegradation was strengthened gradually during the treatment and finally resulted infragments of about150-350bp after6h treatment. The degradation of DNA can bedelayed by ZnSO_4treatment, and the pretreatment with1mM had greater inhibitioneffect on DNA degradation than with0.5mM. The cells treated with pH5.0aceticacid had TUNEL positive nuclei, and the positive nuclei appeared in10%,95%and100%cells after1h,2h and4h treatment, respectively. Those results suggested thatpH5.0acetic acid induced PCD in C. reinhardtii cells.
7. Acetic acid (34.3mM) at pH5.0induced PCD in C. reinhardtii cells, whileH_2SO_4, HCl, H_3PO_4, phosphate buffer and propionic acid did not, but killed the cellsat pH_3.0, indicating that the induction of PCD did not mainly relate with the mediumacidity, but related with its chemical property of acetic acid. When the cells were keptin the culture medium containing50mM acetic acid and at pH6.0, they were alsokilled. However, the cells could survive in the medium containing150mM acetic acidand at pH7.0, at which most of the acetic acid molecules are in ionic form. Thisdemonstrated that there was no relationship between the cell death and the CH_3COOˉform in the medium. The results showed that PCD induced by acetic acid was mostprobably related with both pH and CH_3COOH form.
8. Lots of VOCs were released from the C. reinhardtii cells in PCD induced byacetic acid, and their components and content reached the highest level after2htreatment. At least34different compounds were determined and are among thealdehydes, alcohols, esters, ketones, alkanes, alkenes and terpenoids. Besides aceticacid,300mM NaCl or150mM Na_2CO_3stress also induced VOCs release from C.reinhardtii cells. Hexanal (GLVs) was induced by both acetic acid and NaCl stresses,with the peak areas of23.49×10~7and3.14×10~7, respectively. However, Na_2CO_3stressdid not induce hexanal, but induced special components of3,4-dimethyl-hexane and5-methyl-2-heptene. Longifolene was the main terpenoid under the3kinds of stresses,with peak areas of5.82×10~7,3.11×10~7and4.00×10~7, respectively, for acetic acid,NaCl and Na_2CO_3.
9. When normal C. reinhardtii cells were exposed to those VOCs from the cellsstressed by acetic acid, NaCl or Na_2CO_3, the cell growth was significantly inhibited,but the chlorophyll content, Fv/Fm, H_2O_2content and antioxidant enzyme activitieswere significantly increased, indicating that VOCs from algae under abiotic stress cantransfer information between algae.
引文
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