干扰与竞争条件下丛枝菌根菌和数种作物的互作
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摘要
丛枝菌根菌(Arbuscular mycorrhizal fungi, AM)广泛存在于农业系统中,其可与80%的维管植物共生形成内生菌根,提高宿主植物对营养元素和水分的吸收,增加植物产量和抗逆性,进而影响系统中植物群落的组成和演替。本论文在甘肃省庆阳黄土高原于2004年和2005年连续两年研究了传统耕作、免耕、传统耕作+秸杆覆盖和免耕+秸杆覆盖等耕作方式对丛枝菌根菌的影响。在此基础上,以小麦(Triticum aestivum)和野生型(A17)蒺藜苜蓿(Medicago truncatula var. Jemalong)为轮作作物,在温室条件下模拟了黄土高原耕作方式下,土壤干扰和秸秆覆盖对根内球囊霉(Glomus intraradices)和珍珠巨孢囊霉(Gigaspora margarita)等两种丛枝菌根菌和蒺藜苜蓿互作的影响；并以野生型(A17)和突变体(dmi1)蒺藜苜蓿为竞争组合,采用同位素32P标记,进一步研究了丛枝菌根菌在植物竞争中的作用机理。主要结果如下：
1、田间条件下,丛枝菌根菌与玉米(Zea mays)、冬小麦、大豆(Glycine max)等三种轮作作物在苗期均建立了共生关系,但丛枝菌根菌对3种作物的侵染率均较低,分别为玉米11%～30%,冬小麦8%～31%,大豆3%～30%。耕作方式对丛枝菌根菌侵染率的影响因作物、年份和生长阶段而异。与传统耕作相比,免耕和耕作+秸秆覆盖对3种作物菌根菌侵染率平均提高了10%～15%。3种作物在盛花(结荚)期的菌根菌侵染率均高于苗期,2005年的菌根菌侵染率总体高于2004年。耕作方式对作物产量、地上生物量和作物P吸收未产生显著影响,但三者变化趋势较为一致。
2、温室内模拟黄土高原耕作方式,土壤干扰和秸秆覆盖未影响丛枝菌根菌对蒺藜苜蓿的侵染率,亦未影响菌丝室(hyphal compartment, HCs)和根室(root hyphal compartment, RHCs)土壤中的丛枝菌根菌的菌丝长度。根内球囊霉和珍珠巨孢囊霉单独或混合接种,在A17出苗4周后的侵染率分别达到70%、30%和60%,混合接种效应介于单独接种效应之间。7周时,A17单独接种的侵染率均为60%,混合接种的侵染率为25%,混合接种效应表现为对侵入位点的竞争。土壤干扰通过对丛枝菌根菌的作用而影响A17的P吸收和生物量,未接种处理未对干扰表现出任何响应。干扰显著降低了接种丛枝菌根菌A17的P含量和生物量。两种丛枝菌根菌单独接种或混合接种表现出不同的效应,混合接种和珍珠巨孢囊霉使4周龄A17地上部分组织P含量分别降低37.64%和69.67%；但未影响接种根内球囊霉A17地上组织P含量,在7周龄时,干扰使接种根内球囊霉的A17地上组织P含量降低了30.29%,但未影响其他接种处理A17的地上组织P含量。秸秆覆盖仅提高了干扰条件下HCs土壤接种根内球囊霉和RHC土壤混合接种的菌丝长度,或降低了未干扰条件下RHCs土壤菌丝长度(P＜0.05),但未对丛枝菌根菌和A17互作产生显著影响。土壤干扰、秸秆覆盖和丛枝菌根菌接种亦影响了土壤水稳性团聚体,但因土壤团聚体颗粒直径而异,土壤干扰降低了1～2 mm和0.25～0.5mm直径土壤团聚体,秸秆覆盖提高了1～2 mm和0.25～0.5 mm直径土壤团聚体,丛枝菌根菌仅提高了直径＞2 mm土壤团聚体形成(P＜0.05)。
3、筛选出了对丛枝菌根菌生长正效应的野生型蒺藜苜蓿A17和无反应型蒺藜苜蓿突变体dmil,并用于不同基因型蒺藜苜蓿生长竞争试验。A17对dmi1的竞争优势主要是通过其与丛枝菌根菌建立共生体实现,A17中通过丛枝菌根菌吸收的32P含量为dmi1的100～200倍。在不接种丛枝菌根菌的条件下,A17和dmi1对P的吸收和生物量基本相同。接种丛枝菌根菌后,提高了A17在A17+dmi1和A17+17组合中的竞争能力,尤其是在低P水平下,A17的P含量和生物量可达dmi1的1～2倍。丛枝菌根菌-植物组合对竞争反应存在多样性。A17在接种根内球囊霉后,其在不同竞争组合中的侵染率较一致,不施P处理下约为75%,施P处理下约为40%；但接种珍珠巨孢囊霉后,其在A17+A17中的侵染率高于在A17+dmi1中的侵染率。dmi1在dmi1+dmi1组合中,仅个别植株检测到共生体结构,且侵染率极低(＜2%),但在A17+dmi1中,个别植株侵染率可高达15%。
Arbuscular mycorrhizal (AM) fungi widely exist in agricultural ecosystems and can form symbiosis with approximately 80% of vascular plants. The symbiosis usually enhances plant mineral nutrient uptake, improves water stress tolerance and disease resistance. It plays an important role in agricultural production, vegetation components and succession. A two-year field survey was undertaken in 2004 and 2005 to investigate the effects of various tillage systems on AM fungi infection in the Loess Plateau of Gansu China. These were conventional tillage (CT), no tillage (NT), conventional tillage with stubble retention (TS) and no tillage with stubble retention (NTS). The rotation crops were maize (Zea mays), winter wheat (Triticum aestivum) and soybean(Glycine max). To further understand the effect of soil disturbance and residue application on the interactions of plant and AM fungi, a greenhouse experiment was conducted using wheat and a wild variety (A17) of medic (Medicago truncatula var. Jemalong) to simulate a single rotation of wheat and medic. Soils were disturbed or non-disturbed to simulate tillage or no tillage, with or without application of residue to simulate stubble retention. The two AM fungi used in the trial were Glomus intraradices and Gigaspora margarita. The role of AM fungi in medic growth competition was also studied by using the wild (A17) and mutant (dmil) variety of medic as competition combinations.32P labelled soil was employed to measure the direct P uptake by AM fungi. The results were as the follows:
1. AM colonisation of the crops was well established at seedling stage and varied with agricultural practices and growth stage. All crops had higher AM colonisation at the flowering/podding stage than that of the seedling stage. Percentage of root colonisation was 11%-30% for maize,8%-31% for winter wheat and 3%-30% for soybean. In general, NT and TS increased AM colonisation of the three crops by 10%-15% compared with CT and NTS. There was no effect of residue application or tillage treatment on grain yield, biomass and P uptake of the three crops in either year, however the trends were similar for grain yield, biomass and P uptake under different agricultural practices.
2. In the greenhouse, the two stage experiment was successful in simulating the wheat/medic rotation. Soil disturbance and residue application did not affect AM colonisation in medic nor did it affect hyphal length density in hyphal compartments (HCs) and root hyphal compartments (RHCs). Four weeks after seedling emergence, AM colonies in the medic were well established from the inoculation. The mean colonisation rate was 70%,30% and 60%, by G intraradices, Gi. margarita and mixtures of both species respectively. Percentages of AM colonisation from mixed inoculation was in between inoculation of a single species, however the infection rate by single inoculation of G. intraradices and Gi. margarita was 60%, whereas the rate by mixed inoculation was 25% 7 weeks after emergence. Plants with AM fungi showed an advantage in competition to infection site and growth space in roots. Non mycorrhizal medic did not show any response to soil disturbance, but medic with AM treatment was significantly affected by soil disturbance, which suggests that the effect of soil disturbance on medic was via its effects on AM fungi. Soil disturbance significantly decreased medic P uptake and biomass. Compared with the undisturbed treatment, soil disturbance decreased P uptake of medic inoculated with the mix of the two AM fungi by 37.64% or with Gi. margarita by 69.67% while it had no effect on the G intraradices inoculation. Soil disturbance only decrease P uptake of medic colonized by G intraracides by 30.29% compared to the undisturbed treatment at 7 weeks after emergence. Residue application had little effect on the interactions of medic and AM fungi, it only increased hyphal length density in HCs or RHCs in disturbed and undisturbed treatment. The effects of treatments on size distribution of water stable aggregates was an increased number of particles>2 mm in all inoculated treatments compared with NM treatments. Soil disturbance decreased the number of particles at size of 1-2 mm and 0.25-5 mm where as residue application increased numbers of particles of 1-2 mm and 0.25-0.5 mm.
3. Medic plants of wild type A17 and mutant dimil were chosen as a positive and non response plant to AM fungi in the growth competition experiment. The results showed that A17 had an advantage compared to dmil in P absorbance and that the P extracted via AM fungi was transferred into biomass of A 17 at both A17+A17 and A17+dmil combinations. The biomass of A17 was 1-2 times greater than dim1 and the 32P uptake by A17 was about 100-200 times to dmi1. There is function diversity among plant-AM fungi combination in competition situations. The symbiosis of A17 and AM fungi was well established in the range of 40%-80%. The colonisation of A17 was lower at high P than that in the low P treatment. A 17 also had higher AM colonisation in A17+A17 than in A 17+dmil. The dmi1 showed different competition strategy to AM colonisation, it had higher AM colonisation at A17+dmi1 (15%) than in dmi1+dmi1(2%).
1、田间条件下,丛枝菌根菌与玉米(Zea mays)、冬小麦、大豆(Glycine max)等三种轮作作物在苗期均建立了共生关系,但丛枝菌根菌对3种作物的侵染率均较低,分别为玉米11%～30%,冬小麦8%～31%,大豆3%～30%。耕作方式对丛枝菌根菌侵染率的影响因作物、年份和生长阶段而异。与传统耕作相比,免耕和耕作+秸秆覆盖对3种作物菌根菌侵染率平均提高了10%～15%。3种作物在盛花(结荚)期的菌根菌侵染率均高于苗期,2005年的菌根菌侵染率总体高于2004年。耕作方式对作物产量、地上生物量和作物P吸收未产生显著影响,但三者变化趋势较为一致。
2、温室内模拟黄土高原耕作方式,土壤干扰和秸秆覆盖未影响丛枝菌根菌对蒺藜苜蓿的侵染率,亦未影响菌丝室(hyphal compartment, HCs)和根室(root hyphal compartment, RHCs)土壤中的丛枝菌根菌的菌丝长度。根内球囊霉和珍珠巨孢囊霉单独或混合接种,在A17出苗4周后的侵染率分别达到70%、30%和60%,混合接种效应介于单独接种效应之间。7周时,A17单独接种的侵染率均为60%,混合接种的侵染率为25%,混合接种效应表现为对侵入位点的竞争。土壤干扰通过对丛枝菌根菌的作用而影响A17的P吸收和生物量,未接种处理未对干扰表现出任何响应。干扰显著降低了接种丛枝菌根菌A17的P含量和生物量。两种丛枝菌根菌单独接种或混合接种表现出不同的效应,混合接种和珍珠巨孢囊霉使4周龄A17地上部分组织P含量分别降低37.64%和69.67%；但未影响接种根内球囊霉A17地上组织P含量,在7周龄时,干扰使接种根内球囊霉的A17地上组织P含量降低了30.29%,但未影响其他接种处理A17的地上组织P含量。秸秆覆盖仅提高了干扰条件下HCs土壤接种根内球囊霉和RHC土壤混合接种的菌丝长度,或降低了未干扰条件下RHCs土壤菌丝长度(P＜0.05),但未对丛枝菌根菌和A17互作产生显著影响。土壤干扰、秸秆覆盖和丛枝菌根菌接种亦影响了土壤水稳性团聚体,但因土壤团聚体颗粒直径而异,土壤干扰降低了1～2 mm和0.25～0.5mm直径土壤团聚体,秸秆覆盖提高了1～2 mm和0.25～0.5 mm直径土壤团聚体,丛枝菌根菌仅提高了直径＞2 mm土壤团聚体形成(P＜0.05)。
3、筛选出了对丛枝菌根菌生长正效应的野生型蒺藜苜蓿A17和无反应型蒺藜苜蓿突变体dmil,并用于不同基因型蒺藜苜蓿生长竞争试验。A17对dmi1的竞争优势主要是通过其与丛枝菌根菌建立共生体实现,A17中通过丛枝菌根菌吸收的32P含量为dmi1的100～200倍。在不接种丛枝菌根菌的条件下,A17和dmi1对P的吸收和生物量基本相同。接种丛枝菌根菌后,提高了A17在A17+dmi1和A17+17组合中的竞争能力,尤其是在低P水平下,A17的P含量和生物量可达dmi1的1～2倍。丛枝菌根菌-植物组合对竞争反应存在多样性。A17在接种根内球囊霉后,其在不同竞争组合中的侵染率较一致,不施P处理下约为75%,施P处理下约为40%；但接种珍珠巨孢囊霉后,其在A17+A17中的侵染率高于在A17+dmi1中的侵染率。dmi1在dmi1+dmi1组合中,仅个别植株检测到共生体结构,且侵染率极低(＜2%),但在A17+dmi1中,个别植株侵染率可高达15%。
Arbuscular mycorrhizal (AM) fungi widely exist in agricultural ecosystems and can form symbiosis with approximately 80% of vascular plants. The symbiosis usually enhances plant mineral nutrient uptake, improves water stress tolerance and disease resistance. It plays an important role in agricultural production, vegetation components and succession. A two-year field survey was undertaken in 2004 and 2005 to investigate the effects of various tillage systems on AM fungi infection in the Loess Plateau of Gansu China. These were conventional tillage (CT), no tillage (NT), conventional tillage with stubble retention (TS) and no tillage with stubble retention (NTS). The rotation crops were maize (Zea mays), winter wheat (Triticum aestivum) and soybean(Glycine max). To further understand the effect of soil disturbance and residue application on the interactions of plant and AM fungi, a greenhouse experiment was conducted using wheat and a wild variety (A17) of medic (Medicago truncatula var. Jemalong) to simulate a single rotation of wheat and medic. Soils were disturbed or non-disturbed to simulate tillage or no tillage, with or without application of residue to simulate stubble retention. The two AM fungi used in the trial were Glomus intraradices and Gigaspora margarita. The role of AM fungi in medic growth competition was also studied by using the wild (A17) and mutant (dmil) variety of medic as competition combinations.32P labelled soil was employed to measure the direct P uptake by AM fungi. The results were as the follows:
1. AM colonisation of the crops was well established at seedling stage and varied with agricultural practices and growth stage. All crops had higher AM colonisation at the flowering/podding stage than that of the seedling stage. Percentage of root colonisation was 11%-30% for maize,8%-31% for winter wheat and 3%-30% for soybean. In general, NT and TS increased AM colonisation of the three crops by 10%-15% compared with CT and NTS. There was no effect of residue application or tillage treatment on grain yield, biomass and P uptake of the three crops in either year, however the trends were similar for grain yield, biomass and P uptake under different agricultural practices.
2. In the greenhouse, the two stage experiment was successful in simulating the wheat/medic rotation. Soil disturbance and residue application did not affect AM colonisation in medic nor did it affect hyphal length density in hyphal compartments (HCs) and root hyphal compartments (RHCs). Four weeks after seedling emergence, AM colonies in the medic were well established from the inoculation. The mean colonisation rate was 70%,30% and 60%, by G intraradices, Gi. margarita and mixtures of both species respectively. Percentages of AM colonisation from mixed inoculation was in between inoculation of a single species, however the infection rate by single inoculation of G. intraradices and Gi. margarita was 60%, whereas the rate by mixed inoculation was 25% 7 weeks after emergence. Plants with AM fungi showed an advantage in competition to infection site and growth space in roots. Non mycorrhizal medic did not show any response to soil disturbance, but medic with AM treatment was significantly affected by soil disturbance, which suggests that the effect of soil disturbance on medic was via its effects on AM fungi. Soil disturbance significantly decreased medic P uptake and biomass. Compared with the undisturbed treatment, soil disturbance decreased P uptake of medic inoculated with the mix of the two AM fungi by 37.64% or with Gi. margarita by 69.67% while it had no effect on the G intraradices inoculation. Soil disturbance only decrease P uptake of medic colonized by G intraracides by 30.29% compared to the undisturbed treatment at 7 weeks after emergence. Residue application had little effect on the interactions of medic and AM fungi, it only increased hyphal length density in HCs or RHCs in disturbed and undisturbed treatment. The effects of treatments on size distribution of water stable aggregates was an increased number of particles>2 mm in all inoculated treatments compared with NM treatments. Soil disturbance decreased the number of particles at size of 1-2 mm and 0.25-5 mm where as residue application increased numbers of particles of 1-2 mm and 0.25-0.5 mm.
3. Medic plants of wild type A17 and mutant dimil were chosen as a positive and non response plant to AM fungi in the growth competition experiment. The results showed that A17 had an advantage compared to dmil in P absorbance and that the P extracted via AM fungi was transferred into biomass of A 17 at both A17+A17 and A17+dmil combinations. The biomass of A17 was 1-2 times greater than dim1 and the 32P uptake by A17 was about 100-200 times to dmi1. There is function diversity among plant-AM fungi combination in competition situations. The symbiosis of A17 and AM fungi was well established in the range of 40%-80%. The colonisation of A17 was lower at high P than that in the low P treatment. A 17 also had higher AM colonisation in A17+A17 than in A 17+dmil. The dmi1 showed different competition strategy to AM colonisation, it had higher AM colonisation at A17+dmi1 (15%) than in dmi1+dmi1(2%).
引文
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