根瘤菌在苜蓿植株体内的运移及影响因素
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摘要
根瘤菌与豆科植物的共生固氮作用在改良土壤肥力、提高牧草和作物产量、改善生态环境等方面有着十分重要的意义和作用。前期的工作表明内生根瘤菌普遍存在于豆科植物的种子及植株体内，但对于根瘤菌自植株根系向种子的运移过程和途径还未有直接的证据。为明确内生根瘤菌在苜蓿体内的运移通道、运移规律及影响内生根瘤菌寄主体内运移的因素，本研究利用三亲本杂交技术将cfp荧光蛋白导入苜蓿根瘤菌S.12531(苜蓿根瘤菌标准菌)和R. GN5(分离自甘农5号紫花苜蓿种子的内生根瘤菌)中，得到外源荧光标记根瘤菌S.12531f和内源荧光标记根瘤菌R. GNf。针对芽苗期、幼苗期及栽培1a越冬后返青的甘农5号紫花苜蓿田间植株，在接种方法、接种部位和外源干扰物质对内生根瘤菌和外源侵入根瘤菌在植株体内运移和转运规律的影响方面进行了试验和探讨。结果表明：
1.内源菌R. GNf和外源菌S.12531f对宿主芽苗的侵染和在芽苗体内的运移过程存在相似性和异质性。相似性在于两种标记菌都能侵染芽苗根系并进入茎内。异质性表现为内源菌在植株茎和根内的菌体密度差异小，并能进入芽苗的子叶，而外源菌24h内不能进入芽苗子叶，且菌体密度在宿主植株根内和茎内差异很大。表明根瘤菌在芽苗内由根系向茎和子叶运移的过程存在选择性屏障，第一个选择性屏障出现在芽苗的根茎之间，可降低外源菌的通过数量而对内源菌没有影响，第二个屏障出现在茎与子叶之间，能够阻断外源菌向子叶的运移。
2.标记菌通过茎中部的表面创口侵入幼苗和田间植株后，能向下运移至茎和根，但不能由接种部位向上运移，也不能长期定殖在接种部位至茎基的运移通道内，因此形成了侵入标记菌在植物组织中的不连续分布。外源标记菌由茎表创口进入植株并向根系运移的速率高于由茎杆中髓部向根系运移的速率，而内源菌相反。对田间植株而言，由茎表伤口侵入植株有利于外源菌的运移，而由茎杆髓部向根系的运移通道(注射处理)更适宜内源菌的转运。对苜蓿幼苗不同部位叶片进行菌液涂抹后发现，茎杆划伤接种及叶片涂抹接种都能使标记根瘤菌进入植株并向下运移，但并不经由同一路径；涂抹于不同部位叶片的标记菌都能向下转运至茎和根，但不能进入其他部位的叶片，也不能向上运移。在10-12d龄的芽苗中，由真叶叶片向下至根系的根瘤菌运移通道尚未形成，涂抹于划伤页面的菌体不能侵入植株。
3.由根系侵入的标记根瘤菌在植株生长各个阶段都有自根向地上部分运移的通道。芽苗、幼苗和田间植株在菌液浸根处理后，各检出组织部位的标记菌密度存在很大差异，但两种标记菌都能进入苜蓿植株的根并运移至茎内。内源标记菌R.GNf还能进入芽苗的子叶、幼苗的下部叶片和田间植株的上部叶片，而无外源物质作用时外源菌S.12531f只能由根向上运移到植株茎内。对根系进行切根(芽苗)或切除原有根瘤(田间植株)处理后，受损伤的根系形成了更多的根瘤菌侵染路径，能显著增加标记菌在植株根组织内的数量，促进芽苗中两种标记菌和田间植株中的内源菌向植株茎的运移，能促使内源菌运移至芽苗子叶，但抑制了内源菌向田间植株上部叶片的运移。
4.低浓度的LaCl3、IAA及相异胞外多糖均能不同程度的提高宿主芽苗和幼苗根系中标记菌的分布密度，但LaCl3会显著降低外源菌S.12531f在宿主茎内的菌体密度，并抑制内源菌向茎和子叶内运移。IAA能提高外源菌在茎内的菌体数量，但会降低内源菌在茎和子叶中的菌体密度。不同作用机理的外源物质对标记菌的侵入和运移会产生不同影响，仅仅依靠破坏根系、改变或破坏胞壁构象的外源物质，只能促使标记菌进入根系，但对进一步向茎和子叶的运移没有积极的作用，使用内源菌的胞外多糖可以降低芽苗体内根-茎运移屏障对外源菌的选择压力，但不能使外源菌通过茎叶之间的选择性运移屏障；外源菌胞外多糖会增加内源菌在幼苗和芽苗茎内的分布密度。
5.常温下根瘤菌能够在2d内由根部外环境侵入幼苗根系并运移至茎，少量的内源标记菌能进入下部叶片。幼苗的茎部环境对菌种具有选择性，外源菌可以存在于茎内，但数量和在总检出内生菌中的比例均显著低于相同部位的内源菌；4℃低温抑制根瘤菌的侵入和在宿主体内的运移，且对外源根瘤菌的抑制作用强于内源根瘤菌。营养供给的均衡性能促进根瘤菌侵入植株根系，而缺乏氮素营养的微环境则有利于标记菌向幼苗茎部运移。
6.内源及外源标记根瘤菌侵入根系后都能持续向植株茎部转运，2d时到达植株下部茎，5d时内源菌能进入植株上部叶，而外源菌只到达植株上部茎，此时标记菌菌体密度在根以外的检出部位组织中达到最高，6-7d内茎内标记菌的密度逐渐降低但分布部位不变。
7.田间营养生长初期的植株菌液浸根接种40d后，标记菌仍由根系向植株地上部分运移，且只有内源菌能进入植株上部叶片。表明返青后的田间植株在营养生长初期，体内存在根瘤菌向上运移的通道，但标记菌在运移通道中的分布是不连续的，在部分通道组织内能只能选择性允许标记菌通过，不能长期定殖。并指出外源菌在宿主根和茎内能够存在40d且菌体密度接近内源菌，但不能像内源菌一样进入植株上部叶片，证明苜蓿植株体内存在菌体运移的选择性屏障。
The symbiotic azotification between Rhizobia and leguminous forages plays an importantrole in ameliorating soil fertilization, enhancing the productivity of forage grass and crop, as wellas improving ecological environment. Previous work indicated that endogenous Rhizobia widelyexisted in the seeds and plants of leguminous forages, while no direct proof was found to provethe migration and pathway of Rhizobia between roots and seeds. In order to make clear themigrational pathway and rules of endogenous rhizobia in alfalfa plants, as well as the factors thatinfluence the migration of endogenous rhizobia in host plants, triparental mating technology wasused to transfer cfp fluorescent protein into Sinorhizobium meliloti.12531(standard alfalfa rhizobia)and Rhizobium meliloti. GN5(isolated from the endogenous rhizobia in seeds of Medicago sativaGannong NO.5) to obtain exogenous fluorescence marked rhizobia S.12531f and endogenousfluorescence marked rhizobia R.GNf. Inoculation methods, inoculation positions and the effect ofexogenous interfering substances on the migration and transferring rules of endogenous rhizobiaand exogenous invasive rhizobia in plants were tested based on the bud stage, seedling stage andreviving Gannong NO.5(1a after cultivation) that cultivated in the open fields. The results showedthat:
1. Both similarity and heterogeneity were found for endogenous R.GNf and exogenousS.12531f in terms of host seedlings infection and transportational process. For the similarity, boththe two marked bacteria could infect the root of seedling and enter the stem. For the heterogeneity,endogenous bacteria had relatively smaller differnece in bacterial density in stems and roots, andcould enter the cotyledon of seedling, while exogenous bacteria could not enter the cotyledon ofseeding within24h, and the bacterial densities in roots and stems of the host plants was relativelygreater, indicating that in the seedlings, a selective barrier exists in the transportational process ofrhizobia from root to stem and cotyledon, the1stselective barried occurred between root and stem,which could possibly decrease the quantity of exogenous bacteria while play no effect onendogenous, and the2ndselective barrier occurred between stem and cotyledon, which could cutoff the migration of exogenous bacteria to cotyledon.
2. After infecting into seedlings and field plants, the marked bacteria could be migrated toroot and stem, but could not be migrated upward from inoculation position, or colonized in thepathway between the inoculation position and stem base, and therefore formed a discontinuousdistribution of marked bacteria in the plant tissues. The enter speed of exogenous bacteria from the scar to root was found faster than that from the medulla to root, whereas endogenous bacteria hadreversed situation. To field plants, the scar on the stem was good for the migration of exogenousbacteria, while the pathway between medulla and root was found more appropriate for themigration of endogenous (injected treatments). The daub tests conducted on young leaves ofdifferent parts indicate that both stem-scratching inoculation and leaf-daub inoculation could makethe marked bacteria migrated downward but in different pathways; the marked bacteria thatdaubed on different parts of leaves could move downward to stem and root, but could neither enterthe leaves in other positions nor move upward. In10-12d seedlings, the rhizobia migrationalpathway from euphylla to root hadn't been formed, and the bacteria daubed onto the scratchedleaves could not infect the plants.
3. In all stages of growth, the infected marked bacteria had the pathway to be migratedupward to the aboveground part. After the roots of bud seedling, young seedling and field plantsbeing treated with bacterial solution, the detected marked bacteria in different positions werefound with significant difference, but both the two marked bacteria could enter the root of alfalfaand been migrated to stem. Endogenous marked bacteria R.GNf could enter cotyledon of buds,lower leaves of seedlings and upper leaves of field plants, whereas exogenous S.12531f could onlymove upward to stems when exogenous substances were absent. When the root (for bud seedlings)or the original nodules (for field plants) been cut off, the injured roots gave more ways to rhizobiainfection, which significantly increased the quantity of marked bacteria in root tissues, andpromoted the migration of the two bacteria in bud seedlings and endogenous bacteria in plants tomake more endogenous bacteria moved to cotyledon of seedlings, but which also inhibited theupward movement of endogenous bacteria in field plants.
4. Low concentraion LaCl3, IAA and alien extracellular polysaccharides could inordinatelyincrease the distribution density of marked bacteria that existed in the root of host bud seedlingsand young seedlings, while LaCl3significantly decreased the distribution density of S.12531inhost stem, and also inhibited the migration of endogenous bacteria to stem and cotyledon. IAAcould increase the quantity of exogenous bacteria that existed in stems, but decreased the bacterialdensity of endogenous bacteria that existed in stem and cotyledon. Exogenous substances withdifferent mechanisms like root damage, the change or damage of cell wall conformation, playdifferent effects on the infection and migration of marked bacteria, but could only urge the markedbacteria to enter the root, while could not play an positive role in the further transportation of the marked bacteria to stem and cotyledon. The usage of extracelluar polysaccharides of endogenousbacteria could decrease the selective pressure from root-stem barrier in bud seedlings, but couldnot going through the selective barrier between leaf and stem by using exogenous bacteria;exogenous extracellular polysaccharides could increase the distributional density of endogenousbacteria in stems of bud seedlings and young seedlings.
5. On condition of normal temperature, rhizobia could infected the young seedlings fromoutside and be migrated from root to stem, few endogenous marked bacteria could move to thelower leaves. The stems of young seedlings were selective to bacteria stains, exogenous bacteriacould exist in stem, but the detected quantity takes only a very low proportion in the totalendogenous bacteria, which was significantly lower than other endogenous bacteria existed in thesame part. Low temperature at4℃could inhibit the infection and migration of rhizobia, and theinhibitory effect of which on exogenous rhizobia was found stronger than that on endogenousrhizobia. Balanced nutrition supply could promote rhizobia infecting plant roots, while themicro-environment which is nitrogen deficient was beneficial for the migration of marked rhizobiato seedlings stem.
6. Both endogenous and exogenous marked rhizobia could keep the continuity of migratingto stems after the infection into roots, and arrived at lower stem2days later,5days later,endogenous bacteria could come into upper leaves, while exogenous bacteria could only arrievedat upper stem, at this time, bacterial density in plant tissues except in root were found the highest,on6-7thday, the density decreased progressively without change in distributional position.
7. For plants that grown for40d after root-soaked inoculation treatment and that were in theprimary stages of vegetative growth in the fields, the marked bacteria of which could still bedetected move upward to the aboveground part, but only endogenous bacteria could enter upperleaves of plants, indicating that the reviving plants which were in the primary stages of vegetativegrowth had the upward pathway for rhizobia migration, while the distribution of marked bacteriawas discontinued, some pathway and tissue were selective to marked bacteria and could makethem colonized indefinitely. The exogenous bacteria could exist about40d in the stems and rootsof host plants with approximately the same density of endogenous bacteria, but could not enter theupper leaves like endogenous, indicating that selective barriers for rhizobia migration are exist inalfalfa plants.
1.内源菌R. GNf和外源菌S.12531f对宿主芽苗的侵染和在芽苗体内的运移过程存在相似性和异质性。相似性在于两种标记菌都能侵染芽苗根系并进入茎内。异质性表现为内源菌在植株茎和根内的菌体密度差异小，并能进入芽苗的子叶，而外源菌24h内不能进入芽苗子叶，且菌体密度在宿主植株根内和茎内差异很大。表明根瘤菌在芽苗内由根系向茎和子叶运移的过程存在选择性屏障，第一个选择性屏障出现在芽苗的根茎之间，可降低外源菌的通过数量而对内源菌没有影响，第二个屏障出现在茎与子叶之间，能够阻断外源菌向子叶的运移。
2.标记菌通过茎中部的表面创口侵入幼苗和田间植株后，能向下运移至茎和根，但不能由接种部位向上运移，也不能长期定殖在接种部位至茎基的运移通道内，因此形成了侵入标记菌在植物组织中的不连续分布。外源标记菌由茎表创口进入植株并向根系运移的速率高于由茎杆中髓部向根系运移的速率，而内源菌相反。对田间植株而言，由茎表伤口侵入植株有利于外源菌的运移，而由茎杆髓部向根系的运移通道(注射处理)更适宜内源菌的转运。对苜蓿幼苗不同部位叶片进行菌液涂抹后发现，茎杆划伤接种及叶片涂抹接种都能使标记根瘤菌进入植株并向下运移，但并不经由同一路径；涂抹于不同部位叶片的标记菌都能向下转运至茎和根，但不能进入其他部位的叶片，也不能向上运移。在10-12d龄的芽苗中，由真叶叶片向下至根系的根瘤菌运移通道尚未形成，涂抹于划伤页面的菌体不能侵入植株。
3.由根系侵入的标记根瘤菌在植株生长各个阶段都有自根向地上部分运移的通道。芽苗、幼苗和田间植株在菌液浸根处理后，各检出组织部位的标记菌密度存在很大差异，但两种标记菌都能进入苜蓿植株的根并运移至茎内。内源标记菌R.GNf还能进入芽苗的子叶、幼苗的下部叶片和田间植株的上部叶片，而无外源物质作用时外源菌S.12531f只能由根向上运移到植株茎内。对根系进行切根(芽苗)或切除原有根瘤(田间植株)处理后，受损伤的根系形成了更多的根瘤菌侵染路径，能显著增加标记菌在植株根组织内的数量，促进芽苗中两种标记菌和田间植株中的内源菌向植株茎的运移，能促使内源菌运移至芽苗子叶，但抑制了内源菌向田间植株上部叶片的运移。
4.低浓度的LaCl3、IAA及相异胞外多糖均能不同程度的提高宿主芽苗和幼苗根系中标记菌的分布密度，但LaCl3会显著降低外源菌S.12531f在宿主茎内的菌体密度，并抑制内源菌向茎和子叶内运移。IAA能提高外源菌在茎内的菌体数量，但会降低内源菌在茎和子叶中的菌体密度。不同作用机理的外源物质对标记菌的侵入和运移会产生不同影响，仅仅依靠破坏根系、改变或破坏胞壁构象的外源物质，只能促使标记菌进入根系，但对进一步向茎和子叶的运移没有积极的作用，使用内源菌的胞外多糖可以降低芽苗体内根-茎运移屏障对外源菌的选择压力，但不能使外源菌通过茎叶之间的选择性运移屏障；外源菌胞外多糖会增加内源菌在幼苗和芽苗茎内的分布密度。
5.常温下根瘤菌能够在2d内由根部外环境侵入幼苗根系并运移至茎，少量的内源标记菌能进入下部叶片。幼苗的茎部环境对菌种具有选择性，外源菌可以存在于茎内，但数量和在总检出内生菌中的比例均显著低于相同部位的内源菌；4℃低温抑制根瘤菌的侵入和在宿主体内的运移，且对外源根瘤菌的抑制作用强于内源根瘤菌。营养供给的均衡性能促进根瘤菌侵入植株根系，而缺乏氮素营养的微环境则有利于标记菌向幼苗茎部运移。
6.内源及外源标记根瘤菌侵入根系后都能持续向植株茎部转运，2d时到达植株下部茎，5d时内源菌能进入植株上部叶，而外源菌只到达植株上部茎，此时标记菌菌体密度在根以外的检出部位组织中达到最高，6-7d内茎内标记菌的密度逐渐降低但分布部位不变。
7.田间营养生长初期的植株菌液浸根接种40d后，标记菌仍由根系向植株地上部分运移，且只有内源菌能进入植株上部叶片。表明返青后的田间植株在营养生长初期，体内存在根瘤菌向上运移的通道，但标记菌在运移通道中的分布是不连续的，在部分通道组织内能只能选择性允许标记菌通过，不能长期定殖。并指出外源菌在宿主根和茎内能够存在40d且菌体密度接近内源菌，但不能像内源菌一样进入植株上部叶片，证明苜蓿植株体内存在菌体运移的选择性屏障。
The symbiotic azotification between Rhizobia and leguminous forages plays an importantrole in ameliorating soil fertilization, enhancing the productivity of forage grass and crop, as wellas improving ecological environment. Previous work indicated that endogenous Rhizobia widelyexisted in the seeds and plants of leguminous forages, while no direct proof was found to provethe migration and pathway of Rhizobia between roots and seeds. In order to make clear themigrational pathway and rules of endogenous rhizobia in alfalfa plants, as well as the factors thatinfluence the migration of endogenous rhizobia in host plants, triparental mating technology wasused to transfer cfp fluorescent protein into Sinorhizobium meliloti.12531(standard alfalfa rhizobia)and Rhizobium meliloti. GN5(isolated from the endogenous rhizobia in seeds of Medicago sativaGannong NO.5) to obtain exogenous fluorescence marked rhizobia S.12531f and endogenousfluorescence marked rhizobia R.GNf. Inoculation methods, inoculation positions and the effect ofexogenous interfering substances on the migration and transferring rules of endogenous rhizobiaand exogenous invasive rhizobia in plants were tested based on the bud stage, seedling stage andreviving Gannong NO.5(1a after cultivation) that cultivated in the open fields. The results showedthat:
1. Both similarity and heterogeneity were found for endogenous R.GNf and exogenousS.12531f in terms of host seedlings infection and transportational process. For the similarity, boththe two marked bacteria could infect the root of seedling and enter the stem. For the heterogeneity,endogenous bacteria had relatively smaller differnece in bacterial density in stems and roots, andcould enter the cotyledon of seedling, while exogenous bacteria could not enter the cotyledon ofseeding within24h, and the bacterial densities in roots and stems of the host plants was relativelygreater, indicating that in the seedlings, a selective barrier exists in the transportational process ofrhizobia from root to stem and cotyledon, the1stselective barried occurred between root and stem,which could possibly decrease the quantity of exogenous bacteria while play no effect onendogenous, and the2ndselective barrier occurred between stem and cotyledon, which could cutoff the migration of exogenous bacteria to cotyledon.
2. After infecting into seedlings and field plants, the marked bacteria could be migrated toroot and stem, but could not be migrated upward from inoculation position, or colonized in thepathway between the inoculation position and stem base, and therefore formed a discontinuousdistribution of marked bacteria in the plant tissues. The enter speed of exogenous bacteria from the scar to root was found faster than that from the medulla to root, whereas endogenous bacteria hadreversed situation. To field plants, the scar on the stem was good for the migration of exogenousbacteria, while the pathway between medulla and root was found more appropriate for themigration of endogenous (injected treatments). The daub tests conducted on young leaves ofdifferent parts indicate that both stem-scratching inoculation and leaf-daub inoculation could makethe marked bacteria migrated downward but in different pathways; the marked bacteria thatdaubed on different parts of leaves could move downward to stem and root, but could neither enterthe leaves in other positions nor move upward. In10-12d seedlings, the rhizobia migrationalpathway from euphylla to root hadn't been formed, and the bacteria daubed onto the scratchedleaves could not infect the plants.
3. In all stages of growth, the infected marked bacteria had the pathway to be migratedupward to the aboveground part. After the roots of bud seedling, young seedling and field plantsbeing treated with bacterial solution, the detected marked bacteria in different positions werefound with significant difference, but both the two marked bacteria could enter the root of alfalfaand been migrated to stem. Endogenous marked bacteria R.GNf could enter cotyledon of buds,lower leaves of seedlings and upper leaves of field plants, whereas exogenous S.12531f could onlymove upward to stems when exogenous substances were absent. When the root (for bud seedlings)or the original nodules (for field plants) been cut off, the injured roots gave more ways to rhizobiainfection, which significantly increased the quantity of marked bacteria in root tissues, andpromoted the migration of the two bacteria in bud seedlings and endogenous bacteria in plants tomake more endogenous bacteria moved to cotyledon of seedlings, but which also inhibited theupward movement of endogenous bacteria in field plants.
4. Low concentraion LaCl3, IAA and alien extracellular polysaccharides could inordinatelyincrease the distribution density of marked bacteria that existed in the root of host bud seedlingsand young seedlings, while LaCl3significantly decreased the distribution density of S.12531inhost stem, and also inhibited the migration of endogenous bacteria to stem and cotyledon. IAAcould increase the quantity of exogenous bacteria that existed in stems, but decreased the bacterialdensity of endogenous bacteria that existed in stem and cotyledon. Exogenous substances withdifferent mechanisms like root damage, the change or damage of cell wall conformation, playdifferent effects on the infection and migration of marked bacteria, but could only urge the markedbacteria to enter the root, while could not play an positive role in the further transportation of the marked bacteria to stem and cotyledon. The usage of extracelluar polysaccharides of endogenousbacteria could decrease the selective pressure from root-stem barrier in bud seedlings, but couldnot going through the selective barrier between leaf and stem by using exogenous bacteria;exogenous extracellular polysaccharides could increase the distributional density of endogenousbacteria in stems of bud seedlings and young seedlings.
5. On condition of normal temperature, rhizobia could infected the young seedlings fromoutside and be migrated from root to stem, few endogenous marked bacteria could move to thelower leaves. The stems of young seedlings were selective to bacteria stains, exogenous bacteriacould exist in stem, but the detected quantity takes only a very low proportion in the totalendogenous bacteria, which was significantly lower than other endogenous bacteria existed in thesame part. Low temperature at4℃could inhibit the infection and migration of rhizobia, and theinhibitory effect of which on exogenous rhizobia was found stronger than that on endogenousrhizobia. Balanced nutrition supply could promote rhizobia infecting plant roots, while themicro-environment which is nitrogen deficient was beneficial for the migration of marked rhizobiato seedlings stem.
6. Both endogenous and exogenous marked rhizobia could keep the continuity of migratingto stems after the infection into roots, and arrived at lower stem2days later,5days later,endogenous bacteria could come into upper leaves, while exogenous bacteria could only arrievedat upper stem, at this time, bacterial density in plant tissues except in root were found the highest,on6-7thday, the density decreased progressively without change in distributional position.
7. For plants that grown for40d after root-soaked inoculation treatment and that were in theprimary stages of vegetative growth in the fields, the marked bacteria of which could still bedetected move upward to the aboveground part, but only endogenous bacteria could enter upperleaves of plants, indicating that the reviving plants which were in the primary stages of vegetativegrowth had the upward pathway for rhizobia migration, while the distribution of marked bacteriawas discontinued, some pathway and tissue were selective to marked bacteria and could makethem colonized indefinitely. The exogenous bacteria could exist about40d in the stems and rootsof host plants with approximately the same density of endogenous bacteria, but could not enter theupper leaves like endogenous, indicating that selective barriers for rhizobia migration are exist inalfalfa plants.
引文
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