黄土高原枸杞根际微生态特征及其共生真菌调控宿主生长与耐旱响应机制
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摘要
本研究采用巢式PCR-DGGE、克隆测序、实时定量PCR、BIOLOG和叶绿素荧光等技术研究了黄土高原旱区不同品种枸杞根系丛枝菌根真菌(AMF)与深色有隔内生真菌(DSE)定殖特征,不同品种枸杞根际土壤球囊霉素含量和AMF群落结构多样性季相变化特征,不同生态条件下枸杞根际土壤粒径分布与细菌和真菌群落多样性的关系,枸杞共生DSE菌株生物学特性和菌丝体对人工模拟干旱胁迫的响应,接种DSE对枸杞生长、根际细菌群落功能与遗传多样性的影响及干旱胁迫条件下接种AMF和DSE对枸杞叶片形态、光合参数和根系结构的影响。获得以下主要结果:
1.黄土高原枸杞丛枝菌根真菌(AMF)与深色有隔内生真菌(DSE)定殖特征
对柠杞1号(NQ-1)、柠杞5号(NQ-5)和柠杞菜1号(NQC-1)根系丛枝菌根真菌(AMF)与深色有隔内生真菌(DSE)定殖特征进行了调查研究,结果表明,枸杞形成P-型AM菌根。NQ-1丛枝菌根侵染率最高。不同品种枸杞的AMF侵染率不同,且呈现季节差异。同时,3种枸杞根系均存在DSE定殖。微菌核、深色(棕色)有隔菌丝主要分布在皮层细胞中。8月份DSE镜空菌丝侵染率最高,12月份最低。镜空菌丝和微菌核中存在大量脂类物质,这些脂类物质可能扮演着协助宿主渡过干旱逆境的碳源能量物质的角色。
2.枸杞根际球囊霉素含量及丛枝菌根真菌(AMF)群落结构
采用巢式PCR-DGGE和克隆测序技术研究了枸杞根际土壤AMF群落结构和球囊霉素含量,结果表明,不同品种枸杞根际土壤中易提取球囊霉素含量存在显著性差异,且呈现季节变化,8月份球囊霉素含量最高,12月份最低。总球囊霉素与易提取球囊霉素含量呈显著正相关。巢式PCR-DGGE图谱表明不同品种枸杞根际AMF群落不同,季节和品种对AMF群落丰富度指数(S)和多样性指数(H)影响显著,12月份丰富度指数显著低于8月份和10月份,但是两者交互作用不显著。冗余分析表明,枸杞根际土壤AMF群落多样性与土壤环境因子之间存在密切关系。克隆测序结果表明17条序列均属于球囊霉属(Glomus)。
3.不同生态条件下枸杞土壤粒径与微生物群落多样性
固原、同心、银川和中宁生态区枸杞根际土壤粒径最多分布在20~50μm范围之内,土壤粒径分别占总粒径的37.21±0.21%、33.25±0.21%、38.35±0.18%和42.33±0.45%。惠农地区有40.21±0.26%分布在100~200μm范围。巢式PCR-DGGE结果表明,5个生态区枸杞根际细菌群落丰富度指数为:同心>中宁>固原=银川>惠农;真菌群落丰富度指数为:银川>同心>惠农>中宁>固原,细菌和真菌群落多样性指数之间相关性不显著。主成分分析表明,固原典型变量值的变异(离散)最小,而银川细菌典型变量值的变异(离散)最大,说明银川枸杞根际细菌群落最不稳定。固原与同心真菌群落多样性差异不显著。克隆测序结果表明枸杞根际细菌和真菌均为不可培养微生物类群。
4.枸杞DSE的分离、鉴定及其生物学特性
从枸杞根系上分离DSE菌株,对3株能与枸杞形成共生关系的菌株利用分子生物学技术(rDNA-ITS)进行鉴定。结果显示,3株DSE分别为Phoma chrysanthemicola,Cladosporium cladosporioides LBF3和C. cladosporioides LBF6。3种DSE对不同单一碳源和氮源物质利用有显著差异,P. chrysanthemicola最佳利用碳源和氮源为葡萄糖和甘氨酸,C. cladosporioides LBF3最佳利用碳源和氮源为甘露醇和脯氨酸,C. cladosporioidesLBF6最佳利用碳源和氮源为木糖和甘氨酸。模拟干旱胁迫结果发现P. chrysanthemicola的菌落形态、菌丝形态、菌丝蛋白质表达发生变化,傅里叶红外光谱分析表明模拟干旱引起菌丝中蛋白质结构发生改变。
5. DSE对枸杞根际细菌群落功能和遗传多样性的影响
为了探索接种DSE对枸杞根际微生态的影响,盆栽实验结果表明,接种3种DSE可以改变枸杞幼苗根际土壤细菌数量,接种DSE真菌60 d后枸杞根际土壤细菌代谢活性(BIOLOG)、代谢多样性均高于不接菌的对照。不同DSE菌株的作用有差异。主成分分析表明,接种深色有隔内生真菌后枸杞根际细菌群落对碳源的利用种类和程度均高于对照,细菌群落功能多样性发生了较大变化,接种DSE使得细菌群落变的更稳定;同时,巢式PCR-DGGE结果显示,DSE对细菌群落遗传多样性产生影响。
6. DSE对枸杞生长的影响
为了探索DSE与枸杞幼苗互作机制,将分离自枸杞根系的LBF-2菌株接种到枸杞幼苗根部,在实验室控制条件下研究了接种DSE对枸杞幼苗生长的影响,35天后检测结果发现深色有隔菌丝侵入根系皮层细胞,形成微菌核。首次采用巢式PCR-DGGE技术证明了DSE菌丝在植物组织中的定位和分布,结果认为DSE菌丝分布在枸杞根系、茎和叶片组织中。接种LBF-2可以提高枸杞幼苗生物量28%,接种幼苗叶绿素荧光参数显著高于不接种对照,叶绿素a和总叶绿素含量提高16%和19%。同时,接种植株黄酮物质含量比不接种高26%。结果表明LBF-2可能在枸杞育苗和栽培中发挥作用。
7.接种AMF和DSE枸杞苗叶片与根系对干旱胁迫的响应
为了探索接种AMF和DSE枸杞对干旱胁迫的响应机制,盆栽实验条件下,研究了接种AMF和DSE枸杞苗叶片与根系对干旱胁迫的响应。结果表明,接种AMF和DSE可以改变根系形态和叶片形态,显著降低叶片电解质透出率,提高叶片气孔导度和净光合速率。接种共生真菌能够有效地促进枸杞根系发育,通过提高枸杞叶片气孔导度和光合速率来减轻干旱胁迫的毒害作用。
In this work, nested PCR-DGGE, clone sequence, RT-PCR, BIOLOG and chlorophyll fluorescence techniques were employed to explore the arbuscular mycorrhizal fungal and dark septate endophytes colonization status, arbuscular mycorrhizal fungal (AMF) community structure diversity and glomalin content associated with the Lycium barbarum from arid and semi-arid Loess Plateau, China. Whilst, we investigated the soil particle size distributions and diversity of soil microbial communities in the rhizosphere of L. barbarum growing in different ecological environmental conditions. We also isolated several DSE strains from the roots of L. barbarum, identified these fungal species by molecular method as rDNA-ITS, and determined the effects of carbon and nitrogen sources on the growth of three dark septate endophytes. After that, we examined the responses of DSE under drought stress conditions. Finally, we examined the effects of inoculation with AMF and DSE on the growth and drought stress alleviation of L. barbarum seedlings by determination of leaf and root structures as well as photosynthetic parameters. The main points of this dissertation are as follows:
1. Arbuscular mycorrhizas and dark septate endophytes colonization status in L. barbarum L. growing on Loess Plateau. The AM and DSE fungal colonization status in three different L. barbarum cultivars (NQ-1, NQ-5, NQC-1) in arid northwestern China were investigated. The results showed that the three cultivars were simultaneously colonized by Paris-type AM and DSE associations. The highest colonization by AM was found in L. barbarum Ningqi No.1. The significant“month”and“cultivar”indicates that the AM colonization varied among the months within the cultivar. Meanwhile, roots of the three cultivars were heavily colonized by DSE fungi. Melanized hyphae were frequently observed. Microsclerotia of varied shapes were also found in the cortex cells of L. barbarum. Hyaline hyphae were most abundant in August but their occurrence decreased in December. Lipid contents were abundant in hyaline hyphae, and changed with month. Lipid may act as energy material reserves to sustain the DSE-host symbioses under drought conditions.
2. Arbuscular mycorrhizal fungal community and glomalin content associated with the rhizosphere of L. barbarum. We aimed to examine the AM fungal community diversity using nested PCR-DGGE and clone sequencing techniques, and to quantify the glomalin content associated with different cultivars of L. barbarum. The results showed that the significant“month”and“cultivar”indicates that easily extractable glomalin content varied among the months within the cultivar, the highest content was found in August, while the lowest was in December. The significant positive corelation was observed between total glomalin and easily extractable glomalin content. According to the DGGE fingerprints, the significant“month”and“cultivar”indicates that Species richness (S) and Shannon’s diversity (H) index of AMF community varied among the months within the cultivar, the Species richness (S) in December was lower than August and October, the interaction effect was no significant. Redundancy analysis (RDA) showed the significant relationship between AMF community diversity and soil environmental factors. The results of clone sequence showed that 17 sequences were belonged to Glomus.
3. The soil particle size distributions and diversity of soil microbial communities associated with the rhizosphere of L. barbarum from different ecological conditions were investigated. The results showed that the soils from Guyuan, Tongxin, Yinchuan, and Zhongning were most distributed from 20~50μm, the percentages were 37.21±0.21%, 33.25±0.21%, 38.35±0.18%, and 42.33±0.45%, respectively. In Huinong area, there was 40.21±0.26% soils distributed from 100~200μm. Nested PCR-DGGE suggested that the bacterial species richness of the five ecological regions was ordered as: Tongxin>Zhongning>Guyuan=Yinchuan>Huinong, whilst, fungal species richness was ordered as: Yinchuan>Tongxin>Huinong>Zhongning>Guyuan. There was no significant correlation between bacterial and fungal species richness diversity. Principle Component Analyses (PCA) revealed that bacterial community in Yinchuan was instability, because the community variation was lowest in Guyuan and highest in Yinchuan, no significant fungal community diversity was observed between Guyuan and Tongxin regions. The results of clone sequencing suggested that the bacterial and fungal bands were belonged to uncultured microbial community, respectively.
4. Isolation, identification, and biological characteristics of dark septate endophytes isolates were determined. The results showed that three DSE isolates were belonged to Phoma chrysanthemicola,Cladosporium cladosporioides LBF3, and C. cladosporioides LBF6 based on rDNA-ITS analyses. There were different responses of P. chrysanthemicola, C. cladosporioides LBF3, and C. cladosporioides LBF6 grown on the same carbon and nitrogen medium, and differences in patterns of growth on the different substrates were also found. Glucose, mannitol and xylose were the best carbon sources; glycine and proline were the best nitrogen sources for the growth of P. chrysanthemicola, C. cladosporioides LBF3 and C. cladosporioides LBF6, respectively. Clone and mycelial structures, and protein expression were changed when P. chrysanthemicola exposed to water stress, whilst, fourier transform infrared spectroscopy showed that protein structues were changed under drought stress conditions.
5. To explore the effects of dark septate endophytes inoculation on microecosystem associated with the rhizosphere of L. barbarum under pot culture conditions, bacterial abundance, functional and genetic diversity were determined. The results showed that bacterial abundance, functional and genetic diversity were affected by dark septate endophytes inoculation. DSE inoculations have higher bacterial metabolic activity (BIOLOG), and metabolic diversity than that of control, and varied among different DSE isolates. Principle Component Analyses (PCA) of BIOLOG data revealed that the application of DSE significantly in?uenced bacterial functional diversity in the rhizosphere of L. barbarum seedlings. A wider range of sole carbon sources were utilized by the bacterial community in the rhizosphere of inoculated seedlings. Whilst, nested PCR-DGGE showed that inoculation with DSE influenced bacterial genetic community diversity in the rhizosphere of L. barbarum seedlings.
6. To explore the mechanism of the interaction between DSE and L. barbarum. Effect of a dark septate endophyte fungus on the growth of L. barbarum seedlings were examined under the lab conditions. LBF-2, isolated from the roots of L. barbarum, was inoculated onto the roots of plants, which were grown for 35 d. The results showed that melanized septate hyphae and microsclerotia were observed in the root cortical cells of L. barbarum. Nested PCR-denaturing gradient gel electrophoresis showed, for the first time, that LBF-2 colonizes L. barbarum systemically, with LBF-2 being detected in the root, stem and leaf tissues of L. barbarum. Inoculation with LBF-2 increased the total biomass by 28% and also enhanced chlorophyll fluorescence. Inoculation increased the concentration of total chlorophyll was by 19% and of chlorophyll a by 16%, relative to uninoculated controls. LBF-2 also increased the flavonoid concentration of L. barbarum seedlings by 26%. These data indicate that LBF-2 might be used to facilitate the cultivation of L. barbarum, which has medicinal applications.
7. To examine the response of L. barbarum seedlings to inoculation with AMF and DSE under drought stress conditions, the pot cultured experiment was assayed. The results showed that root and leaf structues were changed with inoculation process. AM and DSE inoculation increased stomatal conductance. Inoculation with symbiotic fungal species was effective for L. barbarum seedlings as an acceptable and ecofriendly technology to improve root and leaf performance and development, and alleviate the damage of drought stress by improving stomatal conductance and efficiency of photosystem II.
1.黄土高原枸杞丛枝菌根真菌(AMF)与深色有隔内生真菌(DSE)定殖特征
对柠杞1号(NQ-1)、柠杞5号(NQ-5)和柠杞菜1号(NQC-1)根系丛枝菌根真菌(AMF)与深色有隔内生真菌(DSE)定殖特征进行了调查研究,结果表明,枸杞形成P-型AM菌根。NQ-1丛枝菌根侵染率最高。不同品种枸杞的AMF侵染率不同,且呈现季节差异。同时,3种枸杞根系均存在DSE定殖。微菌核、深色(棕色)有隔菌丝主要分布在皮层细胞中。8月份DSE镜空菌丝侵染率最高,12月份最低。镜空菌丝和微菌核中存在大量脂类物质,这些脂类物质可能扮演着协助宿主渡过干旱逆境的碳源能量物质的角色。
2.枸杞根际球囊霉素含量及丛枝菌根真菌(AMF)群落结构
采用巢式PCR-DGGE和克隆测序技术研究了枸杞根际土壤AMF群落结构和球囊霉素含量,结果表明,不同品种枸杞根际土壤中易提取球囊霉素含量存在显著性差异,且呈现季节变化,8月份球囊霉素含量最高,12月份最低。总球囊霉素与易提取球囊霉素含量呈显著正相关。巢式PCR-DGGE图谱表明不同品种枸杞根际AMF群落不同,季节和品种对AMF群落丰富度指数(S)和多样性指数(H)影响显著,12月份丰富度指数显著低于8月份和10月份,但是两者交互作用不显著。冗余分析表明,枸杞根际土壤AMF群落多样性与土壤环境因子之间存在密切关系。克隆测序结果表明17条序列均属于球囊霉属(Glomus)。
3.不同生态条件下枸杞土壤粒径与微生物群落多样性
固原、同心、银川和中宁生态区枸杞根际土壤粒径最多分布在20~50μm范围之内,土壤粒径分别占总粒径的37.21±0.21%、33.25±0.21%、38.35±0.18%和42.33±0.45%。惠农地区有40.21±0.26%分布在100~200μm范围。巢式PCR-DGGE结果表明,5个生态区枸杞根际细菌群落丰富度指数为:同心>中宁>固原=银川>惠农;真菌群落丰富度指数为:银川>同心>惠农>中宁>固原,细菌和真菌群落多样性指数之间相关性不显著。主成分分析表明,固原典型变量值的变异(离散)最小,而银川细菌典型变量值的变异(离散)最大,说明银川枸杞根际细菌群落最不稳定。固原与同心真菌群落多样性差异不显著。克隆测序结果表明枸杞根际细菌和真菌均为不可培养微生物类群。
4.枸杞DSE的分离、鉴定及其生物学特性
从枸杞根系上分离DSE菌株,对3株能与枸杞形成共生关系的菌株利用分子生物学技术(rDNA-ITS)进行鉴定。结果显示,3株DSE分别为Phoma chrysanthemicola,Cladosporium cladosporioides LBF3和C. cladosporioides LBF6。3种DSE对不同单一碳源和氮源物质利用有显著差异,P. chrysanthemicola最佳利用碳源和氮源为葡萄糖和甘氨酸,C. cladosporioides LBF3最佳利用碳源和氮源为甘露醇和脯氨酸,C. cladosporioidesLBF6最佳利用碳源和氮源为木糖和甘氨酸。模拟干旱胁迫结果发现P. chrysanthemicola的菌落形态、菌丝形态、菌丝蛋白质表达发生变化,傅里叶红外光谱分析表明模拟干旱引起菌丝中蛋白质结构发生改变。
5. DSE对枸杞根际细菌群落功能和遗传多样性的影响
为了探索接种DSE对枸杞根际微生态的影响,盆栽实验结果表明,接种3种DSE可以改变枸杞幼苗根际土壤细菌数量,接种DSE真菌60 d后枸杞根际土壤细菌代谢活性(BIOLOG)、代谢多样性均高于不接菌的对照。不同DSE菌株的作用有差异。主成分分析表明,接种深色有隔内生真菌后枸杞根际细菌群落对碳源的利用种类和程度均高于对照,细菌群落功能多样性发生了较大变化,接种DSE使得细菌群落变的更稳定;同时,巢式PCR-DGGE结果显示,DSE对细菌群落遗传多样性产生影响。
6. DSE对枸杞生长的影响
为了探索DSE与枸杞幼苗互作机制,将分离自枸杞根系的LBF-2菌株接种到枸杞幼苗根部,在实验室控制条件下研究了接种DSE对枸杞幼苗生长的影响,35天后检测结果发现深色有隔菌丝侵入根系皮层细胞,形成微菌核。首次采用巢式PCR-DGGE技术证明了DSE菌丝在植物组织中的定位和分布,结果认为DSE菌丝分布在枸杞根系、茎和叶片组织中。接种LBF-2可以提高枸杞幼苗生物量28%,接种幼苗叶绿素荧光参数显著高于不接种对照,叶绿素a和总叶绿素含量提高16%和19%。同时,接种植株黄酮物质含量比不接种高26%。结果表明LBF-2可能在枸杞育苗和栽培中发挥作用。
7.接种AMF和DSE枸杞苗叶片与根系对干旱胁迫的响应
为了探索接种AMF和DSE枸杞对干旱胁迫的响应机制,盆栽实验条件下,研究了接种AMF和DSE枸杞苗叶片与根系对干旱胁迫的响应。结果表明,接种AMF和DSE可以改变根系形态和叶片形态,显著降低叶片电解质透出率,提高叶片气孔导度和净光合速率。接种共生真菌能够有效地促进枸杞根系发育,通过提高枸杞叶片气孔导度和光合速率来减轻干旱胁迫的毒害作用。
In this work, nested PCR-DGGE, clone sequence, RT-PCR, BIOLOG and chlorophyll fluorescence techniques were employed to explore the arbuscular mycorrhizal fungal and dark septate endophytes colonization status, arbuscular mycorrhizal fungal (AMF) community structure diversity and glomalin content associated with the Lycium barbarum from arid and semi-arid Loess Plateau, China. Whilst, we investigated the soil particle size distributions and diversity of soil microbial communities in the rhizosphere of L. barbarum growing in different ecological environmental conditions. We also isolated several DSE strains from the roots of L. barbarum, identified these fungal species by molecular method as rDNA-ITS, and determined the effects of carbon and nitrogen sources on the growth of three dark septate endophytes. After that, we examined the responses of DSE under drought stress conditions. Finally, we examined the effects of inoculation with AMF and DSE on the growth and drought stress alleviation of L. barbarum seedlings by determination of leaf and root structures as well as photosynthetic parameters. The main points of this dissertation are as follows:
1. Arbuscular mycorrhizas and dark septate endophytes colonization status in L. barbarum L. growing on Loess Plateau. The AM and DSE fungal colonization status in three different L. barbarum cultivars (NQ-1, NQ-5, NQC-1) in arid northwestern China were investigated. The results showed that the three cultivars were simultaneously colonized by Paris-type AM and DSE associations. The highest colonization by AM was found in L. barbarum Ningqi No.1. The significant“month”and“cultivar”indicates that the AM colonization varied among the months within the cultivar. Meanwhile, roots of the three cultivars were heavily colonized by DSE fungi. Melanized hyphae were frequently observed. Microsclerotia of varied shapes were also found in the cortex cells of L. barbarum. Hyaline hyphae were most abundant in August but their occurrence decreased in December. Lipid contents were abundant in hyaline hyphae, and changed with month. Lipid may act as energy material reserves to sustain the DSE-host symbioses under drought conditions.
2. Arbuscular mycorrhizal fungal community and glomalin content associated with the rhizosphere of L. barbarum. We aimed to examine the AM fungal community diversity using nested PCR-DGGE and clone sequencing techniques, and to quantify the glomalin content associated with different cultivars of L. barbarum. The results showed that the significant“month”and“cultivar”indicates that easily extractable glomalin content varied among the months within the cultivar, the highest content was found in August, while the lowest was in December. The significant positive corelation was observed between total glomalin and easily extractable glomalin content. According to the DGGE fingerprints, the significant“month”and“cultivar”indicates that Species richness (S) and Shannon’s diversity (H) index of AMF community varied among the months within the cultivar, the Species richness (S) in December was lower than August and October, the interaction effect was no significant. Redundancy analysis (RDA) showed the significant relationship between AMF community diversity and soil environmental factors. The results of clone sequence showed that 17 sequences were belonged to Glomus.
3. The soil particle size distributions and diversity of soil microbial communities associated with the rhizosphere of L. barbarum from different ecological conditions were investigated. The results showed that the soils from Guyuan, Tongxin, Yinchuan, and Zhongning were most distributed from 20~50μm, the percentages were 37.21±0.21%, 33.25±0.21%, 38.35±0.18%, and 42.33±0.45%, respectively. In Huinong area, there was 40.21±0.26% soils distributed from 100~200μm. Nested PCR-DGGE suggested that the bacterial species richness of the five ecological regions was ordered as: Tongxin>Zhongning>Guyuan=Yinchuan>Huinong, whilst, fungal species richness was ordered as: Yinchuan>Tongxin>Huinong>Zhongning>Guyuan. There was no significant correlation between bacterial and fungal species richness diversity. Principle Component Analyses (PCA) revealed that bacterial community in Yinchuan was instability, because the community variation was lowest in Guyuan and highest in Yinchuan, no significant fungal community diversity was observed between Guyuan and Tongxin regions. The results of clone sequencing suggested that the bacterial and fungal bands were belonged to uncultured microbial community, respectively.
4. Isolation, identification, and biological characteristics of dark septate endophytes isolates were determined. The results showed that three DSE isolates were belonged to Phoma chrysanthemicola,Cladosporium cladosporioides LBF3, and C. cladosporioides LBF6 based on rDNA-ITS analyses. There were different responses of P. chrysanthemicola, C. cladosporioides LBF3, and C. cladosporioides LBF6 grown on the same carbon and nitrogen medium, and differences in patterns of growth on the different substrates were also found. Glucose, mannitol and xylose were the best carbon sources; glycine and proline were the best nitrogen sources for the growth of P. chrysanthemicola, C. cladosporioides LBF3 and C. cladosporioides LBF6, respectively. Clone and mycelial structures, and protein expression were changed when P. chrysanthemicola exposed to water stress, whilst, fourier transform infrared spectroscopy showed that protein structues were changed under drought stress conditions.
5. To explore the effects of dark septate endophytes inoculation on microecosystem associated with the rhizosphere of L. barbarum under pot culture conditions, bacterial abundance, functional and genetic diversity were determined. The results showed that bacterial abundance, functional and genetic diversity were affected by dark septate endophytes inoculation. DSE inoculations have higher bacterial metabolic activity (BIOLOG), and metabolic diversity than that of control, and varied among different DSE isolates. Principle Component Analyses (PCA) of BIOLOG data revealed that the application of DSE significantly in?uenced bacterial functional diversity in the rhizosphere of L. barbarum seedlings. A wider range of sole carbon sources were utilized by the bacterial community in the rhizosphere of inoculated seedlings. Whilst, nested PCR-DGGE showed that inoculation with DSE influenced bacterial genetic community diversity in the rhizosphere of L. barbarum seedlings.
6. To explore the mechanism of the interaction between DSE and L. barbarum. Effect of a dark septate endophyte fungus on the growth of L. barbarum seedlings were examined under the lab conditions. LBF-2, isolated from the roots of L. barbarum, was inoculated onto the roots of plants, which were grown for 35 d. The results showed that melanized septate hyphae and microsclerotia were observed in the root cortical cells of L. barbarum. Nested PCR-denaturing gradient gel electrophoresis showed, for the first time, that LBF-2 colonizes L. barbarum systemically, with LBF-2 being detected in the root, stem and leaf tissues of L. barbarum. Inoculation with LBF-2 increased the total biomass by 28% and also enhanced chlorophyll fluorescence. Inoculation increased the concentration of total chlorophyll was by 19% and of chlorophyll a by 16%, relative to uninoculated controls. LBF-2 also increased the flavonoid concentration of L. barbarum seedlings by 26%. These data indicate that LBF-2 might be used to facilitate the cultivation of L. barbarum, which has medicinal applications.
7. To examine the response of L. barbarum seedlings to inoculation with AMF and DSE under drought stress conditions, the pot cultured experiment was assayed. The results showed that root and leaf structues were changed with inoculation process. AM and DSE inoculation increased stomatal conductance. Inoculation with symbiotic fungal species was effective for L. barbarum seedlings as an acceptable and ecofriendly technology to improve root and leaf performance and development, and alleviate the damage of drought stress by improving stomatal conductance and efficiency of photosystem II.
引文
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