鱼腥草单萜次生代谢研究
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摘要
鱼腥草是一种传统中药,为三白草科(Saururaceae)蕺菜属(Houttuynia)植物蕺菜(Houttuynia cordata Thunb.)。单萜类化合物是鱼腥草挥发油的主要成分之一,具有抗菌、抗病毒和抗氧化等多种生物活性。根据单萜的生态学意义,本试验首先用UV-B胁迫诱导了鱼腥草单萜次生代谢防御反应,然后以单萜合酶基因为研究对象进一步挖掘了其涉及的信号传导途径,进而采用信号分子诱导策略调控鱼腥草单萜含量,辅以优化的钾肥供应水平,以期实现鱼腥草药用品质和生物产量的双重提升。其主要研究结果如下：
1.首先评估了UV-B辐射对鱼腥草的损伤以及抗氧化酶和单萜次生代谢物质的防御响应。结果表明鱼腥草H202含量和脂质过氧化水平会迅速上升,即UV-B诱导产生了氧化胁迫。不过,遭受氧化损伤的植株在2d后能逐渐恢复。其中,抗氧化酶和单萜类化合物对UV-B诱导的氧化胁迫可能提供了快速的防御响应,而且都表现为多段式防御策略。过氧化氢酶、超氧化物歧化酶和过氧化物酶分别在第1d、2d和3d达到最大活性。单萜在短时间内得到大量地积累。其中,4-萜品醇含量在第1d达到最大值,β-罗勒烯、γ-萜品烯和D-柠檬烯在第2d达到最大值,而其余4种单萜成分在72h一直持续增加。单萜富含不饱和双键,作为抗氧化因子参与防御UV-B诱导的氧化胁迫。这种多段式防御策略可能是由于多重信号传导途径调控的。这为后续挖掘调控鱼腥草单萜合酶基因的信号传导途径奠定了基础。
2.采用cDNA末端快速扩增(RACE)从鱼腥草中分离到两个单萜合酶基因HcTPS1和HcTPS2的全长cDNA。HcTPSl和HcTPS2分别长1886bp和1962bp,最大开放阅读框分别编码558AA和584AA,分别属于萜类合酶亚家族TPS-g和TPS-b。氨基酸序列均以疏水区域为主,包含一段信号肽、DDxxD和RR保守区。HcTPS1和HcTPS2被发现存在单核苷酸多态性(SNP)、插入或缺失突变、内含子滞留和外显子丢失四种形式的变异。HcTPS1和HcTPS2分别发现44个和33个SNP位点。HcTPS1和HcTPS2均存在单碱基插入或缺失的现象,翻译提前终止。HcTPS2内含子ⅢI和Ⅷ的部分序列滞留在cDNA序列上。HcTPS2基因第4和第6个外显子出现丢失。这是首次发现单萜合酶如此奇特而频繁地变异,也暗示了突变在鱼腥草单萜合酶基因中的普遍性。
3.探讨了两种化学型鱼腥草单萜含量与HcTPSl和HcTPS2表达量的关系。首先通过避光和损伤组合处理,解决了鱼腥草叶片离体繁殖过程中易褐化和难分化的两大技术性难题,建立了鱼腥草叶高频离体快繁体系。其再生频率达每外植体20.64±5.94个不定芽。以此技术培养了生长状态基本一致的月桂烯型鱼腥草w01-99、w01-46和w01-71和癸醛型鱼腥草w01-4、w01-86和x10-1组培苗。qPCR分析表明癸醛型鱼腥草HcTPSl和HcTPS2表达水平显著高于月桂烯型鱼腥草。然而,SPME-GC/MS分析表明月桂烯型鱼腥草总单萜含量较癸醛型鱼腥草高,其中x10-1不含单萜。因此,月桂烯型鱼腥草HcTPSl和HcTPS2的表达与单萜含量呈一定的正相关性,而癸醛型鱼腥草的单萜含量和HcTPSl和HcTPS2表达量呈现严重不对称关系。上述单萜合酶基因突变可能是导致癸醛型鱼腥草的单萜含量低的重要原因。
4.采用信号分子抑制剂处理的方式挖掘了涉及UV-B诱导的单萜合酶基因响应的信号传导途径。将乙烯、水杨酸、茉莉酸、NO、H2O2和Ca2+信号抑制剂预处理的植株经UV-B辐射。形态学观察和丙二醛含量的变化表明UV-B剂量在鱼腥草可接受范围之内。除乙烯抑制剂AgNO_3外,其它信号分子的抑制剂预处理未对植株构成胁迫。qPCR分析表明水杨酸、茉莉酸、NO、H2O2和Ca2+抑制剂处理降低或推迟了UV-B诱导的HcTPSl和HcTPS2表达。同时,单萜含量也有所降低,特别是在72h以内。这表明上述5种信号分子可能参与介导了UV-B诱导的单萜合酶基因表达。这为外源施用信号分子促进鱼腥草单萜生物合成提供了理论依据。由于AgNO_3胁迫诱导了单萜次生代谢防御反应,因此本试验未能确定其是否涉及乙烯信号传导途径。
5.评价了信号物质外源施用对鱼腥草单萜含量和脂质过氧化水平的影响。将水杨酸、茉莉酸甲酯、乙烯利(乙烯供体)、硝普钠(NO供体)和CaC1_2叶面喷施于鱼腥草,测定其单萜含量和脂质过氧化水平。结果表明,0.01mmol/L水杨酸不对植株造成氧化胁迫,能提高单萜含量,8d时达到峰值；0.1mmol/L茉莉酸甲酯不对植株造成胁迫,能提高单萜含量,4d时达到峰值;0.1mmol/L乙烯利能显著提高鱼腥草单萜含量,不构成酸胁迫。0.1mmol/L硝普钠可以提高鱼腥草抗性,使脂质过氧化维持在更低的水平,能提高单萜含量,有2d和8d两个单萜积累高峰。1mmol/L CaC1_2不对鱼腥草造成胁迫,总单萜含量在8d时达到最大值。该信号分子诱导策略为鱼腥草药用品质的提升提供了有益的指导。
6.优化了鱼腥草钾供应水平,以为信号分子诱导策略提供茁壮的植株。结果表明1.28mmol/L钾供应水平获得干重、株高、根长和根数最高值,是鱼腥草最适生长浓度。在该浓度下,鱼腥草具有最大净光合速率,最高CO2利用率,这与叶绿素含量直接相关。钾过量供应虽可以提高鱼腥草含钾量,却会降低生物产量。钾饥饿和高钾水平会提高蒸腾速率,降低水分吸收能力,导致植物过度失水。同时会过量积累H2O2,对植株造成氧化胁迫。而最优的钾离子浓度可以提高抗氧化酶活性,维持最低的H2O2水平。过量的钾可以提高单萜含量,然而伴随而来的胁迫效应严重影响植物正常生长。虽然最优的钾浓度未能促进单萜生物合成,但可以使鱼腥草维持在最佳的生长状态。因此,在大田生产中,信号分子诱导策略辅以最优的钾供应浓度也许可以实现单萜含量和生物产量的双重提升。
7.从抗菌和抗氧化活性两方面评估了单萜含量变化对鱼腥草挥发油药用品质的影响。平板扩散法分析表明单萜成分如β-蒎烯、萜品油烯、γ-萜品烯、D-柠檬烯和4-萜品醇都具较强的抗细菌活性,对鱼腥草挥发油的抑菌能力有重要贡献。ABTS~((?)+)法和β-胡萝卜素漂白法分析表明鱼腥草单萜成分具有较强的自由基清除能力和抗脂质过氧化能力。总单萜含量越高,鱼腥草挥发油抗氧化活性越强,其相关性达到显著水平。受测试的单萜个体抗氧化能力均较强。当然,不同单萜个体之间抗氧化活性有一定的差异。因此,本试验结果初步表明诱导单萜生物合成对鱼腥草挥发油药用品质的提升有重要意义。
Honttuynia cordata Thunb. is a species of the genus Houttuynia in ancient Saururaceae family. The plants used as traditional Chinese medicine, are rich in monoterpenes which possess a variety of pharmacological activities, such as antioxidant, anti-bacteria, and anti-viruses. Based on the ecological role, a defense response of monoterpenes were induced by UV-B radiation, and related signaling pathways, which regulating the gene of monoterpene synthases, were investigated. Then, signal compounds were applied to accumulate monoterpenes in H. cordata. In addition, optimization of potassium for proper growth was also tested. Therefore, this study was conducted to promote monoterpene production and biomass together. The main results of this study are as following:
Oxidative damage, antioxidant enzymes and monoterpenes were evaluated in H. cordata after exposure with elevated UV-B radiation. Results showed that the shock of UV-B induced oxidative damage according to the increase of H_2O_2and malondialdehyde. However, a recovery could be found after two days. Catalase, superoxide dismutase, and peroxidase were stimulated to their maximum activities on the first, second, and third day, respectively. Monoterpenes showed rapid accumulation and might play a considerable role as antioxidant agents. Nevertheless, only four individuals mainly contributed to the substantial increase of total monoterpene contents,4-terpineol was quickly induced to the peak on the first day, and β-ocimene, y-terpinene and D-limonene reached the maximum on the second day. This clearly suggests that not only antioxidant enzymes but also monoterpenes play their role in rapid and multiple-phase defense strategy against UV-B-induced oxidative stress in H. cordata. Therefore, multiple signalling pathways may be involved in the regulation of monoterpene biosynthesis under stresses.
Mono-TPS, HcTPSl and HcTPS2were isolated from H. cordata by using rapid amplification of cDNA ends (RACE). Full length cDNA of HcTPSl and HcTPS2are1886bp and1962bp, and encode558AA and584AA, respectively. Both amino-acid sequences are dominated by hydrophobic regions, and contain the transit peptide, the highly conserved DDxxD motif and RR (degenerated RRx_8W) motif. HcTPS1and (?)TPS2belong to TPS-g andTPS-b subfamilies, respectively. Multiple clones of HcTPS1and HcTPS2were isolated from H. cordata accessions w01-99, w01-86, and x10-1. Various mutations, including single nucleotide polymorphisms (SNP), insertion/deletion, intron detention, and exon loss are observed in HcTPS1and HcTPS2.44and33SNP sites are noted in HcTPS1and HcTPS2, respectively. HcTPS1and HcTPS2have insertion and deletion insertion and deletion mutations, respectively, and premature stop codon block the protein translation. Because the change of cleavage site, partial intron sequences are observed in the cDNA of HcTPS2. Two exons are lost in different clones of HcTPS2. Therefore, the mutations of10-TPS might be extensive in H. cordata.
Monoterpene contents and the expression levels of HcTPSl and HcTPS2in two chemotypes of H. cordata were investigated. The Through4week pre-culture in darkness and wounding after1week pre-culture, the browning rate of leaf explants decreased significantly and resulted in efficient regeneration (20.64±5.94adventitious buds per explant) on the induction medium. In vitro plantlets of chemotype myrcene accessions w01-99, w01-46, w01-71, and chemotype decanal accessions w01-4, w01-86, and x10-1were obtained by using the high-frequency induction technique. GC-MS analysis indicates that total monoterpene contents of w01-99, w01-46and w01-71are higher than that of w01-86, and x10-1. Interestingly, no terpenoids were detected. However, the relative transcript levels of HcTPSl and HcTPS2in w01-86and x10-1were significantly higher than that of w01-99, w01-46, w01-71. This suggests that some other targets, such as gene mutation, carbon flux and posttranscriptional processes might play an important role in regulation of monoterpene biosynthesis in H. cordata.
Multiple signalling pathways mediating UV-B-induced monoterpene biosynthesis in H. cordata were investigated by employing the inhibitors of signal transduction pathways. Before exposure with elevated UV-B radiation, the plants were sprayed with the inhibitors of ethylene, salicylic acid, jasmonate, nitric oxide, hydrogen peroxide and calcium ion. Results showed that the dosage of UV-B radiation was within the capacity of H. cordata according to the changes of morphology and malondialdehyde content. foliar application of the inhibitors of salicylic acid, jasmonate, nitric oxide, hydrogen peroxide and calcium ion choud reduce the induction of UV-B-induced HcTPSl expression. Therefore these signalling pathways might be involved in regulation of monoterpene biosynthesis in H. cordata. But ethylene inhibitor, AgNO3resulted in oxidative damage to the plants, and induced the defense response of monoterpenes. Thus it is not well known whether ethylene pathway is involved in regulation of UV-B-induced monoterpene biosynthesis in H. cordata.
Effects of the foliar application of signal molecules or donors, salicylic acid, methyl jasmonate, ethephon (an ethylene-releasing compound), sodium nitroprusside (a nitric oxide-releasing compound) and CaC1_2on monoterpene biosynthesis and lipid peroxidation in H. cordata were evaluated. Results showed that the optimizing concentrations of salicylic acid, methyl jasmonate, ethephon, sodium nitroprusside, and CaC1_2for monoterpene production are0.01,0.1,0.1,0.1, and1mM because of the maximum yield of monoterpenes and the little adverse effect on H. cordata plants, even some treatments could increase the resistance against oxidative stress. The maximum yield of monoterpenes were generally observed on4d or8d after foliar application.
Optimization of potassium for proper growth and physiological response of H. cordata were investigated. Sterile plantlets were cultured in media with different potassium levels, and parameters related to growth, foliar potassium, water and chlorophyll contents, photosynthesis, transpiration, H_2O_2contents and antioxidative enzyme activities were determined after a month. Results showed that1.28mM potassium was the optimum for H. cordata as highest values of dry weight, shoot height, root length and number were obtained at this concentration. The optimum potassium concentration resulted in the maximum net photosynthetic rate which could be associated with the highest chlorophyll content rather than limited stomatal conductance. The supply of surplus potassium resulted in higher content of foliar potassium, but negatively correlated with the biomass. Both potassium starvation (0mM) and high potassium (>1.28mM) could lead to water loss through high transpiration rate and low water absorption, respectively, and resulted in H_2O_2accumulation and increased activities of catalase and peroxidase, which suggested induction of oxidative stress. Moreover, H. cordata showed the minimum of H_2O_2content and the maximum of superoxide dismutase activitiy on1.28mM potassium, implying its role in inducing tolerance against oxidative stress. Total monoterpene content only had a minor share (9.85%) of essential oil in the control treatment (0mM), while it showed a remarkable increase in the treatments supplemented with potassium and10.26mM K resulted in the maximum total monoterpenes. The optimum potassium concentration resulted in lower total monoterpene conent. However, the optimum potassium could make sure the optimal growth of plants for signal strategies in the commercial production of H. cordata
The role of monoterpenes in the essential oils of H. cordata was assessed by antimicrobial activitiy and antioxidant activity assays. Disc diffusion method was employed for the determination of antimicrobial activities. Results indicated that monoterpene components, such as β-pinene, terpinolene, y-terpinene, D-limonene, and4-terpinenol had great potential against bacteria tested, and partly contributed to the antimicrobial capability of essential oils. ABTS~+and β-Carotene bleaching assays indicate that the essential oils have hydrogen-donating capacities and acceptable activities against lipid peroxidation, respectively, which may be significantly associated with total monoterpenes according to correlation analysis. Although differential antioxidant capacities are noted, monoterpene individuals observed in the essential oils generally show great antioxidant activities. This suggests that the induction of monoterpene biosynthesis in H. cordata may be very important for medicinal uses.
1.首先评估了UV-B辐射对鱼腥草的损伤以及抗氧化酶和单萜次生代谢物质的防御响应。结果表明鱼腥草H202含量和脂质过氧化水平会迅速上升,即UV-B诱导产生了氧化胁迫。不过,遭受氧化损伤的植株在2d后能逐渐恢复。其中,抗氧化酶和单萜类化合物对UV-B诱导的氧化胁迫可能提供了快速的防御响应,而且都表现为多段式防御策略。过氧化氢酶、超氧化物歧化酶和过氧化物酶分别在第1d、2d和3d达到最大活性。单萜在短时间内得到大量地积累。其中,4-萜品醇含量在第1d达到最大值,β-罗勒烯、γ-萜品烯和D-柠檬烯在第2d达到最大值,而其余4种单萜成分在72h一直持续增加。单萜富含不饱和双键,作为抗氧化因子参与防御UV-B诱导的氧化胁迫。这种多段式防御策略可能是由于多重信号传导途径调控的。这为后续挖掘调控鱼腥草单萜合酶基因的信号传导途径奠定了基础。
2.采用cDNA末端快速扩增(RACE)从鱼腥草中分离到两个单萜合酶基因HcTPS1和HcTPS2的全长cDNA。HcTPSl和HcTPS2分别长1886bp和1962bp,最大开放阅读框分别编码558AA和584AA,分别属于萜类合酶亚家族TPS-g和TPS-b。氨基酸序列均以疏水区域为主,包含一段信号肽、DDxxD和RR保守区。HcTPS1和HcTPS2被发现存在单核苷酸多态性(SNP)、插入或缺失突变、内含子滞留和外显子丢失四种形式的变异。HcTPS1和HcTPS2分别发现44个和33个SNP位点。HcTPS1和HcTPS2均存在单碱基插入或缺失的现象,翻译提前终止。HcTPS2内含子ⅢI和Ⅷ的部分序列滞留在cDNA序列上。HcTPS2基因第4和第6个外显子出现丢失。这是首次发现单萜合酶如此奇特而频繁地变异,也暗示了突变在鱼腥草单萜合酶基因中的普遍性。
3.探讨了两种化学型鱼腥草单萜含量与HcTPSl和HcTPS2表达量的关系。首先通过避光和损伤组合处理,解决了鱼腥草叶片离体繁殖过程中易褐化和难分化的两大技术性难题,建立了鱼腥草叶高频离体快繁体系。其再生频率达每外植体20.64±5.94个不定芽。以此技术培养了生长状态基本一致的月桂烯型鱼腥草w01-99、w01-46和w01-71和癸醛型鱼腥草w01-4、w01-86和x10-1组培苗。qPCR分析表明癸醛型鱼腥草HcTPSl和HcTPS2表达水平显著高于月桂烯型鱼腥草。然而,SPME-GC/MS分析表明月桂烯型鱼腥草总单萜含量较癸醛型鱼腥草高,其中x10-1不含单萜。因此,月桂烯型鱼腥草HcTPSl和HcTPS2的表达与单萜含量呈一定的正相关性,而癸醛型鱼腥草的单萜含量和HcTPSl和HcTPS2表达量呈现严重不对称关系。上述单萜合酶基因突变可能是导致癸醛型鱼腥草的单萜含量低的重要原因。
4.采用信号分子抑制剂处理的方式挖掘了涉及UV-B诱导的单萜合酶基因响应的信号传导途径。将乙烯、水杨酸、茉莉酸、NO、H2O2和Ca2+信号抑制剂预处理的植株经UV-B辐射。形态学观察和丙二醛含量的变化表明UV-B剂量在鱼腥草可接受范围之内。除乙烯抑制剂AgNO_3外,其它信号分子的抑制剂预处理未对植株构成胁迫。qPCR分析表明水杨酸、茉莉酸、NO、H2O2和Ca2+抑制剂处理降低或推迟了UV-B诱导的HcTPSl和HcTPS2表达。同时,单萜含量也有所降低,特别是在72h以内。这表明上述5种信号分子可能参与介导了UV-B诱导的单萜合酶基因表达。这为外源施用信号分子促进鱼腥草单萜生物合成提供了理论依据。由于AgNO_3胁迫诱导了单萜次生代谢防御反应,因此本试验未能确定其是否涉及乙烯信号传导途径。
5.评价了信号物质外源施用对鱼腥草单萜含量和脂质过氧化水平的影响。将水杨酸、茉莉酸甲酯、乙烯利(乙烯供体)、硝普钠(NO供体)和CaC1_2叶面喷施于鱼腥草,测定其单萜含量和脂质过氧化水平。结果表明,0.01mmol/L水杨酸不对植株造成氧化胁迫,能提高单萜含量,8d时达到峰值；0.1mmol/L茉莉酸甲酯不对植株造成胁迫,能提高单萜含量,4d时达到峰值;0.1mmol/L乙烯利能显著提高鱼腥草单萜含量,不构成酸胁迫。0.1mmol/L硝普钠可以提高鱼腥草抗性,使脂质过氧化维持在更低的水平,能提高单萜含量,有2d和8d两个单萜积累高峰。1mmol/L CaC1_2不对鱼腥草造成胁迫,总单萜含量在8d时达到最大值。该信号分子诱导策略为鱼腥草药用品质的提升提供了有益的指导。
6.优化了鱼腥草钾供应水平,以为信号分子诱导策略提供茁壮的植株。结果表明1.28mmol/L钾供应水平获得干重、株高、根长和根数最高值,是鱼腥草最适生长浓度。在该浓度下,鱼腥草具有最大净光合速率,最高CO2利用率,这与叶绿素含量直接相关。钾过量供应虽可以提高鱼腥草含钾量,却会降低生物产量。钾饥饿和高钾水平会提高蒸腾速率,降低水分吸收能力,导致植物过度失水。同时会过量积累H2O2,对植株造成氧化胁迫。而最优的钾离子浓度可以提高抗氧化酶活性,维持最低的H2O2水平。过量的钾可以提高单萜含量,然而伴随而来的胁迫效应严重影响植物正常生长。虽然最优的钾浓度未能促进单萜生物合成,但可以使鱼腥草维持在最佳的生长状态。因此,在大田生产中,信号分子诱导策略辅以最优的钾供应浓度也许可以实现单萜含量和生物产量的双重提升。
7.从抗菌和抗氧化活性两方面评估了单萜含量变化对鱼腥草挥发油药用品质的影响。平板扩散法分析表明单萜成分如β-蒎烯、萜品油烯、γ-萜品烯、D-柠檬烯和4-萜品醇都具较强的抗细菌活性,对鱼腥草挥发油的抑菌能力有重要贡献。ABTS~((?)+)法和β-胡萝卜素漂白法分析表明鱼腥草单萜成分具有较强的自由基清除能力和抗脂质过氧化能力。总单萜含量越高,鱼腥草挥发油抗氧化活性越强,其相关性达到显著水平。受测试的单萜个体抗氧化能力均较强。当然,不同单萜个体之间抗氧化活性有一定的差异。因此,本试验结果初步表明诱导单萜生物合成对鱼腥草挥发油药用品质的提升有重要意义。
Honttuynia cordata Thunb. is a species of the genus Houttuynia in ancient Saururaceae family. The plants used as traditional Chinese medicine, are rich in monoterpenes which possess a variety of pharmacological activities, such as antioxidant, anti-bacteria, and anti-viruses. Based on the ecological role, a defense response of monoterpenes were induced by UV-B radiation, and related signaling pathways, which regulating the gene of monoterpene synthases, were investigated. Then, signal compounds were applied to accumulate monoterpenes in H. cordata. In addition, optimization of potassium for proper growth was also tested. Therefore, this study was conducted to promote monoterpene production and biomass together. The main results of this study are as following:
Oxidative damage, antioxidant enzymes and monoterpenes were evaluated in H. cordata after exposure with elevated UV-B radiation. Results showed that the shock of UV-B induced oxidative damage according to the increase of H_2O_2and malondialdehyde. However, a recovery could be found after two days. Catalase, superoxide dismutase, and peroxidase were stimulated to their maximum activities on the first, second, and third day, respectively. Monoterpenes showed rapid accumulation and might play a considerable role as antioxidant agents. Nevertheless, only four individuals mainly contributed to the substantial increase of total monoterpene contents,4-terpineol was quickly induced to the peak on the first day, and β-ocimene, y-terpinene and D-limonene reached the maximum on the second day. This clearly suggests that not only antioxidant enzymes but also monoterpenes play their role in rapid and multiple-phase defense strategy against UV-B-induced oxidative stress in H. cordata. Therefore, multiple signalling pathways may be involved in the regulation of monoterpene biosynthesis under stresses.
Mono-TPS, HcTPSl and HcTPS2were isolated from H. cordata by using rapid amplification of cDNA ends (RACE). Full length cDNA of HcTPSl and HcTPS2are1886bp and1962bp, and encode558AA and584AA, respectively. Both amino-acid sequences are dominated by hydrophobic regions, and contain the transit peptide, the highly conserved DDxxD motif and RR (degenerated RRx_8W) motif. HcTPS1and (?)TPS2belong to TPS-g andTPS-b subfamilies, respectively. Multiple clones of HcTPS1and HcTPS2were isolated from H. cordata accessions w01-99, w01-86, and x10-1. Various mutations, including single nucleotide polymorphisms (SNP), insertion/deletion, intron detention, and exon loss are observed in HcTPS1and HcTPS2.44and33SNP sites are noted in HcTPS1and HcTPS2, respectively. HcTPS1and HcTPS2have insertion and deletion insertion and deletion mutations, respectively, and premature stop codon block the protein translation. Because the change of cleavage site, partial intron sequences are observed in the cDNA of HcTPS2. Two exons are lost in different clones of HcTPS2. Therefore, the mutations of10-TPS might be extensive in H. cordata.
Monoterpene contents and the expression levels of HcTPSl and HcTPS2in two chemotypes of H. cordata were investigated. The Through4week pre-culture in darkness and wounding after1week pre-culture, the browning rate of leaf explants decreased significantly and resulted in efficient regeneration (20.64±5.94adventitious buds per explant) on the induction medium. In vitro plantlets of chemotype myrcene accessions w01-99, w01-46, w01-71, and chemotype decanal accessions w01-4, w01-86, and x10-1were obtained by using the high-frequency induction technique. GC-MS analysis indicates that total monoterpene contents of w01-99, w01-46and w01-71are higher than that of w01-86, and x10-1. Interestingly, no terpenoids were detected. However, the relative transcript levels of HcTPSl and HcTPS2in w01-86and x10-1were significantly higher than that of w01-99, w01-46, w01-71. This suggests that some other targets, such as gene mutation, carbon flux and posttranscriptional processes might play an important role in regulation of monoterpene biosynthesis in H. cordata.
Multiple signalling pathways mediating UV-B-induced monoterpene biosynthesis in H. cordata were investigated by employing the inhibitors of signal transduction pathways. Before exposure with elevated UV-B radiation, the plants were sprayed with the inhibitors of ethylene, salicylic acid, jasmonate, nitric oxide, hydrogen peroxide and calcium ion. Results showed that the dosage of UV-B radiation was within the capacity of H. cordata according to the changes of morphology and malondialdehyde content. foliar application of the inhibitors of salicylic acid, jasmonate, nitric oxide, hydrogen peroxide and calcium ion choud reduce the induction of UV-B-induced HcTPSl expression. Therefore these signalling pathways might be involved in regulation of monoterpene biosynthesis in H. cordata. But ethylene inhibitor, AgNO3resulted in oxidative damage to the plants, and induced the defense response of monoterpenes. Thus it is not well known whether ethylene pathway is involved in regulation of UV-B-induced monoterpene biosynthesis in H. cordata.
Effects of the foliar application of signal molecules or donors, salicylic acid, methyl jasmonate, ethephon (an ethylene-releasing compound), sodium nitroprusside (a nitric oxide-releasing compound) and CaC1_2on monoterpene biosynthesis and lipid peroxidation in H. cordata were evaluated. Results showed that the optimizing concentrations of salicylic acid, methyl jasmonate, ethephon, sodium nitroprusside, and CaC1_2for monoterpene production are0.01,0.1,0.1,0.1, and1mM because of the maximum yield of monoterpenes and the little adverse effect on H. cordata plants, even some treatments could increase the resistance against oxidative stress. The maximum yield of monoterpenes were generally observed on4d or8d after foliar application.
Optimization of potassium for proper growth and physiological response of H. cordata were investigated. Sterile plantlets were cultured in media with different potassium levels, and parameters related to growth, foliar potassium, water and chlorophyll contents, photosynthesis, transpiration, H_2O_2contents and antioxidative enzyme activities were determined after a month. Results showed that1.28mM potassium was the optimum for H. cordata as highest values of dry weight, shoot height, root length and number were obtained at this concentration. The optimum potassium concentration resulted in the maximum net photosynthetic rate which could be associated with the highest chlorophyll content rather than limited stomatal conductance. The supply of surplus potassium resulted in higher content of foliar potassium, but negatively correlated with the biomass. Both potassium starvation (0mM) and high potassium (>1.28mM) could lead to water loss through high transpiration rate and low water absorption, respectively, and resulted in H_2O_2accumulation and increased activities of catalase and peroxidase, which suggested induction of oxidative stress. Moreover, H. cordata showed the minimum of H_2O_2content and the maximum of superoxide dismutase activitiy on1.28mM potassium, implying its role in inducing tolerance against oxidative stress. Total monoterpene content only had a minor share (9.85%) of essential oil in the control treatment (0mM), while it showed a remarkable increase in the treatments supplemented with potassium and10.26mM K resulted in the maximum total monoterpenes. The optimum potassium concentration resulted in lower total monoterpene conent. However, the optimum potassium could make sure the optimal growth of plants for signal strategies in the commercial production of H. cordata
The role of monoterpenes in the essential oils of H. cordata was assessed by antimicrobial activitiy and antioxidant activity assays. Disc diffusion method was employed for the determination of antimicrobial activities. Results indicated that monoterpene components, such as β-pinene, terpinolene, y-terpinene, D-limonene, and4-terpinenol had great potential against bacteria tested, and partly contributed to the antimicrobial capability of essential oils. ABTS~+and β-Carotene bleaching assays indicate that the essential oils have hydrogen-donating capacities and acceptable activities against lipid peroxidation, respectively, which may be significantly associated with total monoterpenes according to correlation analysis. Although differential antioxidant capacities are noted, monoterpene individuals observed in the essential oils generally show great antioxidant activities. This suggests that the induction of monoterpene biosynthesis in H. cordata may be very important for medicinal uses.
引文
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