【摘要】：本研究調(diào)查了寧夏鹽土叢枝菌根(arbuscular mycorrhizas,AM)真菌的多樣性,并探明了影響鹽土中AM真菌群落變化的主要土壤因子,闡述了AM真菌群落、土壤因子及植物種類之間的關(guān)系。在此基礎(chǔ)上,從鹽脅迫條件下AM真菌對枸杞激素調(diào)節(jié)、光合系統(tǒng)、抗氧化和滲透調(diào)節(jié)、葉超微結(jié)構(gòu)和葉營養(yǎng)成分的影響等方面,研究了接種AM真菌提高枸杞耐鹽性的生理生化機(jī)制。評估了鹽脅迫條件下接種AM真菌作為生物肥料,提高枸杞生長以及改善枸杞產(chǎn)品品質(zhì)的潛力。主要研究結(jié)論如下:1.寧夏鹽土AM真菌群落多樣性在寧夏鹽土枸杞(Lycium barbarum L.)和沙棗(Elaeagnus angustifolia L.)2種植物根際鹽土中鑒定出33種AM真菌,隸屬于7個科10個屬。其中,球囊霉科(Glomeraceae)的AM真菌數(shù)目最多,包含13種,Acaulospora和Glomus在屬水平上豐度最高,分別包含10種和9種,Septoglomus constrictum是豐度最高的種,2種植物共有種是S.constrictum和Funneliformis coronatum,表明寧夏鹽土中具有很高的AM真菌群落多樣性。枸杞與沙棗根際的AM真菌群落顯著不同(R=0.26,p0.05),枸杞根際分離出27種AM真菌,遠(yuǎn)高于沙棗的12種,枸杞根際AM真菌群落的Shannon、Simpson和Evenness等指數(shù)也顯著高于沙棗,表明宿主會影響AM真菌群落結(jié)構(gòu)。2.土壤因子對寧夏鹽土AM真菌群落多樣性的影響鹽脅迫程度與AM真菌多樣性指數(shù)成顯著負(fù)相關(guān),表明鹽脅迫會降低AM真菌多樣性。但是,土壤肥力指標(biāo)(速效鉀、速效磷、速效氮和全磷)與AM真菌多樣性成顯著正相關(guān),表明土壤肥力提升會增加AM真菌多樣性。土壤pH與孢子豐度成正相關(guān),表明偏堿性土壤有利于AM真菌產(chǎn)孢。速效磷與AM真菌均一度成正相關(guān),說明土壤磷含量能提高AM真菌種間均衡度。由此可見,土壤因子能顯著影響AM真菌群落結(jié)構(gòu)。3.鹽脅迫條件下AM真菌對枸杞激素調(diào)節(jié)及生長的影響在鹽脅迫條件下,接種Glomus versiforme的枸杞葉和根部IAA含量分別提高59.0%和39.1%,葉ABA含量提高13.8%。協(xié)方差分析表明,G.versiforme直接影響了枸杞的IAA和ABA含量,并且通過參與這2種激素的調(diào)控,使枸杞的生物量提高1.75倍,根系總吸收和活躍吸收面積分別增加2.17和2.66倍。接種G.versiforme的枸杞株高、地徑和單株葉片數(shù)等形態(tài)指標(biāo)分別提高93.6%、57.8%和86.6%,表明接種G.versiforme的枸杞長勢更好。由此可見,接種G.versiforme通過對枸杞激素調(diào)節(jié),顯著緩解了鹽脅迫對枸杞生長的抑制作用,特別是促進(jìn)了枸杞根系的生長發(fā)育,保證了枸杞對水分和養(yǎng)分的吸收功能。4.鹽脅迫條件下AM真菌對枸杞光合系統(tǒng)的影響在鹽脅迫條件下,研究了接種G.versiforme對枸杞葉綠素、光合作用和葉綠素?zé)晒獾挠绊�。結(jié)果顯示,G.versiforme在非鹽脅迫下使枸杞的葉綠素含量提高25.7%。枸杞的葉綠素含量未受到鹽脅迫的影響,說明枸杞具有較強(qiáng)的耐鹽能力。在鹽脅迫條件下枸杞的光合作用顯著降低,葉綠素?zé)晒庖彩艿胶艽笠种�。但�?接種G.versiforme的枸杞在100 mM鹽濃度下,其光合速率和蒸騰速率分別比對照提高56.0%和28.7%,胞間CO2濃度降低9.0%。在200 mM鹽濃度下,接種G.versiforme的枸杞的電子傳遞效率(ΦPSⅡ)、光化學(xué)猝滅系數(shù)(qP)及非光化學(xué)猝滅系數(shù)(qN)均顯著高于對照,表明G.versiforme能提高枸杞的光能捕獲能力,并協(xié)助其合理分配能量,以保證枸杞在鹽脅迫下的光合作用以及葉的光合系統(tǒng)免受光氧化損傷。5.鹽脅迫條件下AM真菌對枸杞抗氧化和滲透調(diào)節(jié)的影響本試驗研究了鹽脅迫下接種G.versiforme對枸杞抗氧化酶活性、滲透調(diào)節(jié)物質(zhì)以及水勢的作用。結(jié)果表明,接種AM真菌枸杞的超氧化物歧化酶(SOD)和過氧化物酶(POD)活性分別提高11.0%和7.6%,過氧化氫酶(CAT)活性在100 mM鹽脅迫下提高2.85倍,表明G.versiforme能提高枸杞的抗氧化酶活性,降低膜系統(tǒng)的損傷。同時,G.versiforme提高了枸杞的可溶性糖(32.3%)和還原糖(33.6%)含量,降低了淀粉含量(23.8%),表明G.versiforme能促進(jìn)枸杞淀粉水解成可溶性糖和還原糖,進(jìn)而提高枸杞的水勢(32.6%)。以上結(jié)果表明,G.versiforme能在鹽脅迫下提高枸杞抗氧化酶活性和滲透調(diào)節(jié)能力。6.鹽脅迫條件下AM真菌對枸杞葉超微結(jié)構(gòu)的影響本研究評估了接種G.versiforme對鹽脅迫下枸杞葉細(xì)胞超微結(jié)構(gòu)的影響。鹽脅迫嚴(yán)重?fù)p傷了對照枸杞葉細(xì)胞膜結(jié)構(gòu),導(dǎo)致質(zhì)壁分離并破壞了葉綠體類囊體和基質(zhì)片層。與對照相比,接種G.versiforme的枸杞葉肉細(xì)胞質(zhì)壁分離程度較輕,葉綠體相對完整且積累的油滴數(shù)較少,基粒和類囊體完好有序。這些結(jié)果表明,G.versiforme在鹽脅迫下有效保護(hù)了枸杞葉細(xì)胞的結(jié)構(gòu)。7.鹽脅迫條件下接種AM真菌對枸杞葉營養(yǎng)成分的影響本研究在施加鹽脅迫條件下接種Rhizophagus irregularis于菜用枸杞,測定R.irregularis對枸杞葉營養(yǎng)成分的影響。結(jié)果顯示,R.irregularis在非脅迫與鹽脅迫條件下,使枸杞芽的再生速率分別提高了79.0%和113.0%。與對照相比,在鹽脅迫條件下,接種R.irregularis使2次收獲的枸杞葉蘆丁含量分別提高了96.1%和77.5%,枸杞葉酸性多糖分別增加了66.7%和103.1%。此外,R.irregularis還提高了2次收獲枸杞葉的多糖含量,增加幅度分別為9.9%和32.5%。結(jié)果表明,R.irregularis在鹽脅迫下能顯著提高枸杞芽的再生速率,增加枸杞葉的營養(yǎng)成分。
[Abstract]:In this study, the diversity of arbuscular mycorrhizas (AM) fungi in Ningxia saline soil was investigated, and the main soil factors affecting the changes of AM Fungi Community in saline soil were identified. The relationships among AM fungi community, soil factors and plant species were expounded. Physiological and biochemical mechanisms of AM fungi inoculation on improving salt tolerance of Lycium barbarum were studied. The potential of AM fungi inoculation as biofertilizer to improve the growth and quality of Lycium barbarum under salt stress was evaluated. The main conclusions are as follows: 1. The diversity of AM Fungi Community in summer saline soil was identified in the rhizosphere saline soil of Lycium barbarum L. and Elaeagnus angustifolia L. in Ningxia. 33 species of AM fungi belonging to 10 genera and 7 families were identified. Among them, the number of AM fungi in Glomeraceae was the largest, including 13 species, Aclospora and Glomusau were the richest. Septoglomus constrictum was the highest abundant species, and S. constrictum and Funneliformis coronatum were the common species, indicating that there was a high diversity of AM fungal communities in Ningxia saline soil. The Shannon, Simpson and Evenness indices of AM Fungi Community in Lycium barbarum rhizosphere were also significantly higher than those in Elaeagnus angustifolia L. in 12 species of Elaeagnus angustifolia, suggesting that host could affect the structure of AM fungi community. 2. The effect of soil factors on the diversity of AM Fungi Community in Ningxia saline soil was negatively correlated with the diversity index of AM fungi, suggesting that salt Diversity. However, soil fertility indices (available potassium, available phosphorus, available nitrogen and total phosphorus) were positively correlated with AM fungi diversity, indicating that soil fertility increased AM fungi diversity. Soil pH was positively correlated with spore abundance, indicating that alkaline soil was conducive to AM fungi sporulation. The results showed that soil factors could significantly affect the structure of AM fungi community. 3. The effects of AM fungi on hormone regulation and growth of Lycium barbarum L. Under salt stress, IAA content in leaves and roots of Lycium barbarum inoculated with Glomus versiform increased by 59.0% and 39.1%, respectively, and ABA content in leaves increased by 13.8%. Covariance analysis showed that G. versiform directly affected the contents of IAA and ABA in Lycium barbarum. By participating in the regulation of these two hormones, the biomass of Lycium barbarum was increased by 1.75 times, the total absorption and active absorption area of root system were increased by 2.17 and 2.66 times, respectively. 3.6%, 57.8% and 86.6% showed that the growth of Lycium barbarum inoculated with G.versiform was better. Therefore, G.versiform significantly alleviated the inhibition of salt stress on the growth of Lycium barbarum, especially promoted the growth and development of Lycium barum roots, and ensured the absorption of water and nutrients of Lycium barum. 4. AM was true under salt stress. The effects of G. versiform on chlorophyll, photosynthesis and chlorophyll fluorescence of Lycium barbarum L. were studied under salt stress. The results showed that G. versiform increased the chlorophyll content of Lycium barbarum L. by 25.7% under non-salt stress. The chlorophyll content of Lycium barum L. was not affected by salt stress, indicating that Lycium barbarum L. had stronger chlorophyll content. The photosynthesis and chlorophyll fluorescence of Lycium barbarum L. were significantly decreased under salt stress. However, the photosynthetic rate and transpiration rate of Lycium barbarum L. inoculated with G. versiform at 100 mM increased by 56.0% and 28.7% respectively, and the intercellular CO2 concentration decreased by 9.0%. Electron transfer efficiency (PSII), photochemical quenching coefficient (qP) and non-photochemical quenching coefficient (qN) of Lycium barbarum L. were significantly higher than those of the control, indicating that G. versiform could improve the photosynthetic capture ability of Lycium barbarum L. and help it distribute energy reasonably, so as to ensure the photosynthesis of Lycium barum L. under salt stress and protect the photosynthetic system from photooxidation damage.5. Effects of AM fungi on antioxidant and osmotic regulation of Lycium barbarum L. Under salt stress, the effects of G. versiform on the activities of antioxidant enzymes, osmotic regulators and water potential of Lycium barum L. were studied. The results showed that the activities of superoxide dismutase (SOD) and peroxidase (POD) of Lycium barum L. inoculated with AM fungi increased by 11.0% and 7.6% respectively. Catalase (CAT) activity increased by 2.85 times under 100 mM salt stress, indicating that G. versiform could increase the activity of antioxidant enzymes and reduce the damage of membrane system in Lycium barbarum. At the same time, G. versiform increased the content of soluble sugar (32.3%) and reducing sugar (33.6%) and decreased the content of starch (23.8%) in Lycium barbarum, suggesting that G. versiform could promote the hydrolysis of starch in Lycium barum barbarum. The above results showed that G. versiform could increase the activity of antioxidant enzymes and osmotic regulation of Lycium barbarum L. under salt stress. 6. The effect of AM fungi on ultrastructure of Lycium barbarum L. leaves under salt stress was evaluated in this study. The chloroplast thylakoids and stroma lamellae were separated and destroyed seriously by salt stress. Compared with the control, the isolation degree of mesophyll cytoplasmic wall of Lycium barbarum inoculated with G. versiform was lighter, the chloroplast was relatively intact and accumulated less oil droplets, and the matrix and thylakoids were intact and orderly. The results showed that G. versiform effectively protected the structure of Lycium barbarum leaf cells under salt stress. 7. Effects of AM fungi inoculation on the nutritional components of Lycium barbarum leaves under salt stress. Rhizophagus irregularis was inoculated in Lycium barbarum under salt stress. The effects of R. irregularis on the nutritional components of Lycium barbarum leaves were determined. Irregularis increased the regeneration rate of Lycium barbarum buds by 79.0% and 113.0% under non-stress and salt stress, respectively. Compared with the control, inoculation with R. irregularis increased the rutin content of Lycium barbarum leaves by 96.1% and 77.5%, and the acid polysaccharide content of Lycium barum leaves by 66.7% and 103.1% respectively under salt stress. The results showed that R. irregularis could significantly increase the regeneration rate of Lycium barbarum buds and the nutritional components of Lycium barbarum leaves under salt stress.
【學(xué)位授予單位】：西北農(nóng)林科技大學(xué)
【學(xué)位級別】：博士
【學(xué)位授予年份】：2016
【分類號】：S567.19

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