外生菌根菌丝桥在板栗幼苗间传递磷的效应

采用^32P示踪和4室根箱方法研究了外生菌根菌丝桥对板栗磷营养和植株间磷素传递作用的效应。给一株板栗幼苗(供体)接种外生菌根真菌美味牛肝菌(Boletus edulis)、褐环乳牛肝菌(Suillus luteus),菌根真菌在侵染供体植物以后其根外菌丝继续生长并侵染邻近的另外一株板栗植株(受体)。同位素示踪试验表明,供体板栗体内的^32P可通过菌丝桥传递给受体板栗,受体植株不仅根中^32P放射性强度高于对照,而且茎中^32P强度也显著高于对照。说明外生菌根真菌在不同板栗植株间形成了菌丝桥,但是菌丝桥传递的磷的数量很有限,仅占供体植株体内总磷量的5％-8％。美味牛肝菌和褐环乳牛肝菌侵染供体板栗植株以后,使植株含磷量、总吸磷量和生物量较对照明显增加。受体板栗幼苗在菌丝桥建立以后其植株含磷量和总吸磷量显著高于对照,但生物量与对照没有显著差别。     

菌丝桥在日本落叶松幼苗间磷传递和植株生长中的作用

 应用四室隔网系统研究了菌丝桥在日本落叶松(Larix kaempferi)幼苗间传递磷的作用。结果表明,供体接种卷缘桩菇(Paxillus involutus)和彩色豆马勃（Pisolithus tinctorius）后,其外延菌丝可以穿过隔离层侵染受体落叶松,在供体和受体落叶松间形成了菌丝桥。供体植株接种菌根真菌后生物量明显增加,但是对受体植株没有显著的影响。菌根真菌侵染的供体和受体植株的根、地上部吸磷量均分别显著高于对照,而且供体植株根、地上部吸磷量增加的程度明显高于受体。被卷缘桩菇和彩色豆马勃侵染的受体植株体内32P的放射性强度分别是对照的10倍和6倍,两者形成菌丝桥后传递到受体植株的32P分别为供体植株体内32P的1.10%和0.22%。供体植株吸收的32P可以通过菌丝桥传递给受体,但是绝对数量十分有限,对受体植株磷营养没有产生显著的影响,但P. involutus和P. tinctorius侵染受体植株后,促进了受体落叶松对磷的吸收,这是菌丝桥形成后,真菌帮助受体植株吸收磷引起的。     

三叶草根间菌丝桥传递衰亡根系中磷的作用

应用五室方法研究了三叶草根间菌线桥传递衰亡根系中磷的作用。三叶草生长至１０周切除供体地上部让根系衰亡,１１周收收获样品进行分析测定。结果表明：菌线桥可以在植株间传递＾３２Ｐ,从而使受体三叶草地上部磷营养状况得到改善;供体植株地上部切除后有利于＾３２Ｐ通过菌丝桥从衰亡根系向受体植株的转移,表现为受体植株含磷量有所增加,但对植株的生长影响不大。     

丛枝菌根菌丝桥介导的番茄植株根系间抗病信号的传递

菌根菌丝桥是植物间在地下进行物质交流的通道, 但它能否作为植物间地下化学通讯的通道来传递抗病信号则缺乏研究. 本文利用丛枝菌根真菌（AMF）摩西球囊霉在供体与受体番茄植株间建立菌丝桥, 对供体植株接种早疫病病原菌茄链格孢菌, 研究供体与受体番茄植株根系间是否存在抗病信号的传递. 荧光定量PCR检测表明, AMF侵染后的供体番茄植株再接种病原菌, 其根系中苯丙氨酸解氨酶基因（PAL)、脂氧合酶基因（LOX）和几丁质酶基因（PR₃）的转录水平显著高于仅接种病原菌、未接种病原菌和AMF, 以及只接种AMF的番茄植株. 更重要的是, 与供体有菌丝桥连接的受体番茄根系中PAL、LOX和PR₃的基因的表达量也显著高于无菌丝桥连接、菌丝桥连接被阻断以及有菌丝桥连接但供体植物未接种病原菌的处理,3个基因最高转录水平达到无菌丝桥连接对照受体植物的4.2、4.5和3.5倍. 此外, 供体植株根系启动防御反应的时间（18和65 h）比受体（100和140 h）早. 表明病原菌诱导番茄供体根系产生的抗病信号可以通过菌丝桥传递到受体根系.     

外生菌根真菌菌丝对马尾松苗氮素传递的影响

为探究外生菌根真菌菌丝对马尾松Pinus massoniana幼苗氮素的吸收、传递及分配规律的影响,本研究应用¹⁵N示踪技术,采用四室隔网系统进行盆栽试验。研究表明：供体接种外生菌根真菌后,根外菌丝能够穿过粘土层和隔离层到达受体室,在供、受体植株之间形成菌丝桥。接种不同外生菌根真菌均显著增加供体植株苗高、地径及生物量(P<0.05),但对受体植株的影响有差异,其中接种橙黄硬皮马勃Scleroderma citrinum (Sc)对植株苗高、地径及生物量具有显著的促进作用(P<0.05)。接种对供、受体植株全氮含量无显著性的规律(P>0.05),只有接种东方豆马勃Pisolithus orientalis (Po)的植株全氮含量显著高于其他处理。接种能够显著提高供体植株根、茎、叶的吸氮量,且叶>根>茎,受体植株中,接种Po、Sc和褐环乳牛肝菌Suillus luteus (Sl)的植株总吸氮量较不接种(NE)显著增加(P<0.05),而接种混合菌(Sm)的植株与NE无显著差异(P>0.05)。接种Sl的植株¹⁵N传递率最高(75.86%),受体植株对¹⁵N吸收量最大(91.68 μg/pot),Po次之。供体植株中,除接种Sl的植株¹⁵N主要分配在叶外,其他处理的¹⁵N分配中心均在根部,在受体植株中,接种Po、Sl、Sm的植株¹⁵N主要分配在根部,而NE和接种Sc的植株¹⁵N主要分配在茎。综上,在马尾松供、受体苗木间能形成菌丝桥,接种外生菌根真菌能够促进马尾松供体植株苗高、地径的生长及生物量的积累,提高植株吸氮量,对于受体植株不同菌种的作用有差异,其中接种Sc的苗木表现最好。此外,不同的菌种菌丝对植株的全氮含量、¹⁵N吸收量、传递率及分配率的影响也不同,其中,以接种Po和Sl的苗木表现较好。     

丛枝菌根根外菌丝网络形成过程中的时间效应及植物介导作用

以豆科植物紫花苜蓿为试验材料,应用三室(供体室-间隔室-受体室)培养系统,研究在供体和受体紫花苜蓿根系之间菌丝网络形成的时间效应以及间隔室中不同植物对菌丝网络建成的介导作用.第一个试验在供体和受体植物生长8、10、12、14周之后进行收获以检验菌丝网络形成的时间效应;第二个试验则在间隔室分别种植紫花苜蓿、羊草和独行菜,以考察菌根依赖性不同的植物对菌丝网络形成的介导作用.试验结果显示:(1)接种丛枝菌根真菌的供体紫花苜蓿根系能够形成良好的菌根共生,其外延菌丝可穿过尼龙网和间隔室侵染受体植物根系;植物生长8周后,在受体植物根系检测到菌根侵染,证实供体和受体植物间形成了根间菌丝网络;10周后,尽管供体室和受体室植物的侵染率已无差异,但二者的生物量和地上部磷浓度差异却加大,表现出菌丝网络对植物种内竞争影响的不对称性.(2)试验条件下,不同介导植物对受体植物的菌根侵染及生物量均无明显影响,但显著降低了供体植物生物量和地上部磷浓度;间隔室无介导植物或种植独行菜时,受体植物地上部和根系生物量显著低于供体植物,而当介导植物为紫花苜蓿和羊草时,受体和供体植物生物量无显著差异.研究表明,植物根间菌丝网络的形成受时间和介导植物的影响,同时也具有调节植物间资源分配和植物相互作用的功能.     

菌丝桥介入枳根系间抗溃疡病的信号物质响应

对两室根箱试验装置中一分室(供体室)定植的枳Poncirus trifoliata实生苗接种丛枝菌根(AM)真菌Paraglomus occultum,另一分室(受体室)不作任何处理。定植17周后,将溃疡病病菌Xanthomonas axonopodis(Xac)接种于供体植株,研究不同处理受体植株间的信号物质响应差异,揭示菌丝桥的功能。与不接种处理相比,单接种AM真菌、预先接种AM真菌后再接种Xac均显著提高了供体根系一氧化氮(NO)含量,单接种AM真菌只显著提高了受体根系NO含量;单接种Xac显著降低了供体根系脱落酸(ABA)含量,而单接种AM真菌、双接种(AM真菌+Xac)均增加了供体根系ABA含量。双接种处理显著提高了受体ABA含量;与不接种对照相比,接种处理均降低了供体根系玉米素核苷(ZR)含量,同时单接种AM真菌、双接种处理也降低了受体根系ZR含量;AM真菌显著降低了供体和受体根系茉莉酸(JA)含量,但双接种却显著提高了供体和受体根系JA含量;与不接种对照相比,单接种AM真菌或Xac、双接种处理均显著下调供体根系JA合成基因(PtLOX、PtAOS和PtAOC)的表达,单接种Xac显著下调了受体根系PtLOX和Pt AOC的表达,单接种AM真菌显著下调了受体根系PtAOC的表达,而双接种显著上调了受体根系PtAOS基因的表达。表明菌根真菌的菌丝桥传导JA并诱导受体植株中NO、ABA和ZR含量改变以抵御溃疡病菌危害。     

混栽杨树-刺槐间磷素养分转移途径的研究

应用^32P同位素示踪法,人工接种VA菌根菌(Glomus mosseae),对种植在根箱中的杨树(Populus euramericana cv.‘I-214')和刺槐(Robinia pseudoacia)苗木进行了树种间P素养分转移途径的研究．结果表明,5室根箱中,接种VA菌根菌的杨树一例,菌根侵染率为34％,根箱隔网另一侧的刺槐根系侵染率为26％,而对照的杨树和刺槐根系侵染率均为零．在杨树一例施放射性同位累^32P,施用后第14—27天,刺槐一例的放射性同位素值,处理显著高于对照(P＜0．05)、在根箱中的杨树和刺槐根系间观察到菌丝连接,表明人工接种VA菌根菌能在杨树和刺槐根系间产生菌丝桥,菌丝桥可以在杨树和刺槐根系间传递P素．养分转移定量分析表明,根系接触和根系分泌物是树种间P素转移的主要途径,其转移P素量占转移总量的62％;菌根菌等微生物活动及其与根系接触和根系分泌物两种途径的交互作用占38％;菌丝桥通过隔网发挥的作用仅表现为一种趋势．     

大豆柑桔间作系统中磷的分配和迁移规律研究

用微区试验和³²P同位素示踪技术,比较研究了大豆、柑桔间作和单作条件下,P在大豆和柑桔体中的分配、转移及其在土壤中的迁移规律.结果表明,间作大豆的吸P量和各部位累积P量显著地低于单作大豆;³²P肥料浅施,间作大豆吸收的³²P量显著低于单作大豆;³²P肥料深施,间作大豆吸收的³²P量显著高于单作大豆,但间作不影响P和³²P在各部位的转移和分配.间作柑桔吸收的³²P量显著低于单作柑桔.柑桔新吸收的³²P可快速向地上部分输送,并优先供应生长活跃部位.间作不影响³²P在柑桔各部位的转移和分配,但是P肥深施使柑桔吸收的³²P向地上部分和生长活跃部位的转移速率减慢.间作使土壤中P的生物移动性增强,可促进土壤深层P向土壤浅层迁移.试验结果表明,大豆柑桔间作磷肥的施用深度以保持在20cm以内为佳.     

丛枝菌根菌丝桥介导刺槐-魔芋间碳转运和磷吸收

明确间作和接种丛枝菌根真菌(AMF)对刺槐(Robinia pseudoacacia)与魔芋(Amorphophallus konjac)碳转运和磷吸收的影响,可为揭示间作刺槐对魔芋的防病促生机制和推广刺槐林下魔芋绿色高产栽培技术提供科学依据。该研究使用孔径25μm尼龙网设置两室根箱隔网系统,分为A室(刺槐不接种/接种AMF)和B室(单作刺槐/刺槐间作魔芋);采用¹³C稳定性同位素技术对A室刺槐叶片进行标记,研究其同化的碳与B室魔芋之间的传递,以及AMF定植对两种作物的农艺性状、¹³C丰度和磷含量的影响。结果表明:(1)在接种条件下,菌丝桥对B室刺槐和魔芋的侵染率分别达到47.1%和60.4%;其中,刺槐侵染率与单作直接接种处理相比降低14.1%。在间作体系中,接种AMF提高了魔芋的生物量,其地上干质量和地下根系干质量分别较未接种处理显著增加9.7%和36.2%。(2)与单作未接种处理相比,间作未接种、单作接种和间作接种的刺槐同化的碳被更多地分配到根系和根际土壤(A室),并以根系分泌物的形式穿越尼龙网到达邻近作物根际(B室)。(3)与各自未接...     

A comparison of phosphorus and nitrogen transfer between plants of different phosphorus status

Summary This study has two objections: (1) to compare transfers of phosphorus (³²P) with nitrogen (¹⁵N) from undefoliated and defoliated mycorrhizal P-rich plants to an adjacent mycorrhizal plant and (2) to determine whether the improved nutrient status of a plant growing with a nutrient-rich plant is due primarily to movement of nutrients from roots of its nutrient-rich neighbor (= nutrient transfer) or to reduced nutrient uptake by its nutrient-rich neighbor (=shift in competition). Two plants of Plantago lanceolata were grown in a three-pot unit in which each of their root systems were split, with part in the central shared pot and part by themselves in an outside pot. There were three treatments: (1) no added P; (2) P added in the outer pot to only plant, termed the donor plant, since it might provide P to the companion plant, acting as a receiver; and (3) as in the previous treatment but the P-fertilized donor plant was also clipped. To encourage the formation of hyphal links between roots of the different plants, transfers were determined when root length densities were high (90 to 130 cm cm^-3 soil) and when 56 to 85% of the root length was infected with vesicular-arbuscular mycorrhizae. Phosphorus fertilization enhanced P but not N movement within donor plants. Regardless of treatment, N transfer from donor to receiver plants was an order of magnitude greater than P transfer and in amounts that could potentially affect plant nutrition in very infertile soils. Phosphorus transfer was very small in any of the treatments. Although P fertilization and clipping improved P status of receiver plants, P transfer was not indicated as the main reason for the improved nutrition. A shift in competition between donor and receiver plants was likely the major factor in the shift in nutrition of the receiver plants.     

Contribution of the VA mycorrhizal hyphae in acquisition of phosphorus and zinc by maize grown in a calcareous soil

An investigation was carried out to test whether the mechanism of increased zinc (Zn) uptake by mycorrhizal plants is similar to that of increased phosphorus (P) acquisition. Maize (Zea mays L.) was grown in pots containing sterilised calcareous soil either inoculated with a mycorrhizal fungus Glomus mosseae (Nicol. and Gerd.) Gerdemann and Trappe or with a mixture of mycorrhizal fungi, or remaining non-inoculated as non-mycorrhizal control. The pots had three compartments, a central one for root growth and two outer ones for hyphal growth. The compartmentalization was done using a 30-m nylon net. The root compartment received low or high levels of P (50 or 100 mg kg^–1 soil) in combination with low or high levels of P and micronutrients (2 or 10 mg kg^–1 Fe, Zn and Cu) in the hyphal compartments.Mycorrhizal fungus inoculation did not influence shoot dry weight, but reduced root dry weight when low P levels were supplied to the root compartment. Irrespective of the P levels in the root compartment, shoots and roots of mycorrhizal plants had on average 95 and 115% higher P concentrations, and 164 and 22% higher Zn concentrations, respectively, compared to non-mycorrhizal plants. These higher concentrations could be attributed to a substantial translocation of P and Zn from hyphal compartments to the plant via the mycorrhizal hyphae. Mycorrhizal inoculation also enhanced copper concentration in roots (135%) but not in shoots. In contrast, manganese (Mn) concentrations in shoots and roots of mycorrhizal plants were distinctly lower, especially in plants inoculated with the mixture of mycorrhizal fungi.The results demonstrate that VA mycorrhizal hyphae uptake and translocation to the host is an important component of increased acquisition of P and Zn by mycorrhizal plants. The minimal hyphae contribution (delivery by the hyphae from the outer compartments) to the total plant acquisition ranged from 13 to 20% for P and from 16 to 25% for Zn.     

Changes in soil hyphal abundance and viability can alter the patterns of hydraulic redistribution by plant roots

Background and aims

We conducted a mesocosm study to investigate the extent to which the process of hydraulic redistribution of soil water by plant roots is affected by mycorrhizosphere disturbance.

Methods

We used deuterium-labeled water to track the transfer of hydraulically lifted water (HLW) from well-hydrated donor oaks (Quercus agrifolia Nee.) to drought-stressed receiver seedlings growing together in mycorrhizal or fungicide-treated mesocosms. We hypothesized that the transfer of HLW from donor to receiver plants would be enhanced in undisturbed (non-fungicide-treated) mesocosms where an intact mycorrhizal hyphal network was present.

Results

Contrary to expectations, both upper soil and receiver seedlings contained significantly greater proportions of HLW in mesocosms where the abundance of mycorrhizal hyphal links between donor and receiver roots had been sharply reduced by fungicide application. Reduced soil hyphal density and viability likely hampered soil moisture retention properties in fungicide-treated mesocosms, thus leading to faster soil water depletion in upper compartments. The resulting steeper soil water potential gradient between taproot and upper compartments enhanced hydraulic redistribution in fungicide-treated mesocosms.

Conclusions

Belowground disturbances that reduce soil hyphal density and viability in the mycorrhizosphere can alter the patterns of hydraulic redistribution by roots through effects on soil hydraulic properties.     

Interspecific nutrient transfer in a tallgrass prairie plant community

Interplant nutrient transfer may be an important ecological process in grasslands, and may significantly influence plant neighborhood interactions. We investigated the potential for phosphorus transfer between the dominant grass Andropogon gerardii and several neighboring plant species in tallgrass prairie via a field ³²PO₄ labelling experiment. The mean amount of ³²P received from donor shoots differed significantly among neighboring species and decreased with increasing distance from the donor. In general, forbs and cool-season C₃ grasses received more labelled ³²P than warm-season C₄ grasses. Phosphorus transfer occurred over distances up to 0.5 m. The effects of species and distance on movement of phosphorus changed with increasing time after labelling. The relative mass of receiver and donor shoots did not affect amounts of ³²P transfer. A benomyl fungicide treatment, applied to suppress mycorrhizal activity, likely did not affect existing vegetative hyphae and did not affect the amount of ³²P transferred. These studies demonstrate that: (1) phosphorus is transferred among neighboring species in tallgrass prairie plant communities, (2) phosphorus may be transferred over significantly greater distances than reported in other grasslands, and (3) there is differential transfer among co-occurring species. Hypothesized mechanisms accounting for these patterns in tallgrass prairie include mycorrhizal hyphal interconnections and/or extensive and differential root and rhizosphere overlap among neighboring species.     

Absence of carbon transfer between Medicago truncatula plants linked by a mycorrhizal network, demonstrated in an experimental microcosm

Carbon transfer between plants via a common extraradical network of arbuscular mycorrhizal (AM) fungal hyphae has been investigated abundantly, but the results remain equivocal. We studied the transfer of carbon through this fungal network, from a Medicago truncatula donor plant to a receiver (1) M. truncatula plant growing under decreased light conditions and (2) M. truncatula seedling. Autotrophic plants were grown in bicompartmented Petri plates, with their root systems physically separated, but linked by the extraradical network of Glomus intraradices. A control Myc-/Nod- M. truncatula plant was inserted in the same compartment as the receiver plant. Following labeling of the donor plant with 13CO2, 13C was recovered in the donor plant shoots and roots, in the extraradical mycelium and in the receiver plant roots. Fatty acid analysis of the receiver's roots further demonstrated 13C enrichment in the fungal-specific lipids, while almost no label was detected in the plant-specific compounds. We conclude that carbon was transferred from the donor to the receiver plant via the AM fungal network, but that the transferred carbon remained within the intraradical AM fungal structures of the receiver's root and was not transferred to the receiver's plant tissues.     

Nitrogen and phosphorus acquisition by the mycelium of the ectomycorrhizal fungus Paxillus involutus and its effect on host nutrition

The contribution of the extramatrical mycelium to N and P nutrition of mycorrhizal Norway spruce ( Picea abies (L.) Karst.) was investigated. Seedlings either inoculated with Paxillus involutus (Batsch) Fr. or non-mycorrhizal were grown in a two compartment sand culture system where hyphae were separated from roots by a 45 μm nylon net. Nutrient solution of the hyphal compartment contained either 1.8 m m NH₄⁺ and 0.18 m m H₂PO₄⁻ or no N and P. Aluminium added to the hyphal compartment as a tracer of mass flow was not detected in the plant compartment, indicating that measurements of N and P transfer by the mycelium were not biased by solute movement across the nylon net.
The addition of N and P to the hyphal compartment markedly increased dry weight, N and P concentration and N and P content of mycorrhizal plants. Calculating uptake from the difference in input and output of nutrient in solution confirmed a hyphal contribution of 73% and 76% to total N and P uptake, respectively. Hyphal growth was increased at the site of nutrient solution input.     

Endomycorrhizal fungi in nitrogen transfer from soybean to maize

Using ¹⁵N as a tracer, interspecific N-transfer was studied during the course of plant development. The use of barriers of differing permeabilities between donor and receiver plants allowed separation of the effect of mycorrhizal colonization, root or hyphal contact and interplant hyphal bridging, on ¹⁵N-transfer from soybean (Glycine max (L.) Merrill) to maize (Zea mays L.). More transfer was measured between mycorrhizal plants, but transport of ¹⁵N from the labelled host plant to Glomus versiforme (Karsten) Berch did not seem to occur at the symbiotic interface, suggesting that the fungus is independent of its host for its N-nutrition, and that the role of hyphal bridges in N-transfer between plants, is not significant. Uptake by the receiver plant of the N excreted by the donor plant root system appears to be the mechanism of N-transfer between plants. The factor most affecting ¹⁵N-transfer between plants was found to be the extent of the contact between plant root systems. The presence of the endomycorrhizal fungus in plant roots reduced ¹⁵N-loss from soybean, but at the same time, its extensive hyphal network improved the efficiency of the maize root system for the recovery of the ¹⁵N excreted by soybeans. The net result was a better conservation of the N resource within the plant system. The transfer of N between mycorrhizal plants was particularly enhanced by the death of the soybean.