Avena fatua L. seed and seedling nutrient dynamics as influenced by mycorrhizal infection of the maternal generation

Abstract. The objective of this study was to determine how mycorrhizal infection of one generation of plants influences the nutrient dynamics of seeds and seedlings comprising the subsequent generation. We showed that, for Avena fatua L., seeds produced by mycorrhizal (M) plants consistently contained significantly more phosphorus (particularly the phytate P and residual P fractions) than seeds produced by non-mycorrhizal (NM) plants. We also followed the development of spikelets produced by M and NM plants. The rates of increase in spikelet dry weight and nitrogen content were largely unaffected by mycorrhizal infection. However, the rate of P accumulation into spikelets was significantly increased by mycorrhizal infection. Greater endosperm P reserves in seeds produced by M plants were associated with greater rates of P accumulation in resultant seedlings. Moreover, offspring plants (all NM) produced by M mother plants had significantly higher root and rhizosphere phosphatase, ATPase and phytase activities than offspring plants produced by NM mother plants. This persistent maternal effect has never before been described. Our results suggest that mycorrhizal infection of one generation of plants may have substantial positive effects on the offspring generation, and thus, may influence plant population dynamics.     

Growth response and phosphorus uptake of rye with long and short root hairs: Interactions with mycorrhizal infection

Plant growth and phosphorus (P) uptake of two selections of rye (Secale cereale L.) differing in length of root hairs, in response to mycorrhizal infection were investigated. Rye plants with short root hairs (SRH) had a greater length of root infected by Glomus intraradices (up to 32 m pot^–1) than those with long root hairs (LRH) (up to 10 m pot^–1). Application of P decreased the percentage of root length infected in both selections. In low-P soil, mycorrhizal infection increased shoot and root P concentration, especially in LRH plants. Generally, LRH had higher shoot dry weight than SRH plants. P uptake was increased both by LRH and by mycorrhizal infection. Differences in specific P uptake and P utilization efficiency between SRH and LRH plants were observed in non-mycorrhizal plants. With low P supply, P utilization efficiency (dry matter yield per unit of P taken up) of LRH plants increased with time. However, mycorrhizal infection reduced P utilization efficiency, particularly of SRH plants. SRH plants, which were agronomically less efficient (i.e. low dry matter yield at low P supply) were more responsive to either mycorrhizal infection or P addition than the LRH plants. No interaction was observed between mycorrhizal infection and root hair length.     

Species-specific trade-offs between regrowth and mycorrhizas in the face of defoliation and phosphorus addition

Arbuscular mycorrhizal fungi (AMF) enhance nutrient provision in exchange for carbon. However, a shift from nutrient to carbon limitation, induced by repeated or intense defoliation, can represent a trade-off between plant regrowth and the maintenance of mycorrhiza. The combined effects of AMF, phosphorus and defoliation on growth of Agropyron elongatum (C₃ grass, low mycorrhizal responsiveness) and Brachiaria brizantha (C₄ grass, high mycorrhizal responsiveness) were explored. Each species was subjected to a manipulative experiment with AMF inoculation (non-inoculated, inoculated), soluble P supply (low, high) and defoliation (non-defoliated, 60% defoliated). In A. elongatum, at low P supply, mycorrhizal plants showed increased growth rates following defoliation without substantial changes in AMF colonization. At high P supply instead, we found a clear trade-off between regrowth and the maintenance of mycorrhiza evidenced by growth depression (biomass and tillers) and lower AMF activity (reduction of arbuscules). In contrast, in B. brizantha, defoliation effects on plant regrowth were independent from AMF at any P supply (no trade-off). This indicates that cost-benefit relationship in defoliated plants is highly context-dependent and may vary with mycorrhizal responsiveness of species. This variation of responses can play a decisive role on plant recovery in pastures and natural grasslands subjected to herbivory.     

Effect of phosphorus source and rate of application on VAM fungal infection and growth of maize (Zea mays L.)

The effects of two phosphorus (P) sources (triple superphosphate and Ghafsa phosphate rock), applied at rates equivalent to 44kg ha^-1 and 22 kg ha^-1, on vesicular-arbuscular mycorrhizal (VAM) fungal infection in roots, dry matter yield and nutrient content of maize grown in an oxisol and an alfisol, were investigated in a growth cabinet. The application of 44 kg P ha ^-1 resulted in root infection by VAM fungi not was significantly different (P<-0.01) from when no P was applied. Root infection was significantly greater when P was applied as triple superphosphate at the rate of 22 kg ha^-1 the higher rate. Phosphate rock treatments at both rates of application resulted in significantly greater root infection than in controls with no P or when triple superphosphate was applied at 44 kg ha^-1. Plant P uptake increased in all soils with the different P treatments compared with the control. No direct effects of the treatments on the aluminium and zinc contents of maize plants were observed. In the gleyic alfisol, reduced Mn uptake as a result of increased infection of plants with the superphosphate treatments was observed. Higher Mn was also found in plants with the higher rate of superphosphate treatment than with the phosphate rock treatments in the haplustox, although infection rates in plants with the latter treatments were higher. With the exception of plants with the phosphate rock treatment applied at 22kg ha^-1, dry matter yields of plants with all P sources were significantly greater than the controls.     

不同浓度和形态磷处理下内生真菌感染对高羊茅的影响

以感染内生真菌(endophyte-infected,EI)和不感染内生真菌(endophyte-free,EF)的高羊茅(Festuca arundinacea Schreb.)为材料,在温室沙培条件下研究内生真菌对高羊茅适应缺磷及利用不同形态磷肥的影响。结果表明,1)缺磷条件下,高羊茅EI和EF植株生长差异不显著;正常供磷条件下,高羊茅EI植株拥有更多分蘖数和绿叶数。说明正常供磷条件下内生真菌改善了宿主高羊茅的生长。2)与水溶性磷相比,高羊茅根有机酸和酸性磷酸酶(acid phosphatase,APase)活性在难溶性磷条件下显著增加,而根总酚含量无显著变化。在水溶性磷条件下,高羊茅EI植株根总酚含量显著高于EF植株,此时EI植株比EF植株拥有更多分蘖数和绿叶数,说明在水溶性磷条件下内生真菌对宿主地上部生长具有一定贡献。在难溶性磷条件下,虽然高羊茅EI植株根总酚含量仍然高于EF植株,但同时EI植株根有机酸含量显著低于EF植株,因此内生真菌感染只是增大了宿主植物的根冠比,而对分蘖数和绿叶数等无显著影响,说明内生真菌对宿主利用难溶性磷贡献不大。可见,内生真菌对宿主植物的生长在水溶性磷条件下更有利。     

Vesicular-arbuscular mycorrhiza of Calamagrostis villosa supplied with organic and inorganic phosphorus

Plants of Calamagrostis villosa were cultivated in nutrient solution alone or in association with a vesicular-arbuscular mycorrhizal (VAM) fungus Glomus etunicatum. They were supplied with two levels of inorganic phosphate (Pi; 0.1 or 1 mM) and with or without organic phosphate (dinatriumphenylphosphate, Po; 1 mM). Depression of growth and enhancement of root respiration of mycorrhizal plants in comparison with non-mycorrhizal plants were observed after 12 weeks of cultivation in a growth cabinet. Root colonization was not influenced by the higher phosphorus availability in contrast to the extraradical mycelium (ERM). The lengths of ERM hyphae both attached to the root surface and in the substrate were decreased substantially by higher phosphorus supply, irrespectively of its form. This revised version was published online in July 2006 with corrections to the Cover Date.     

Influence of arbuscular mycorrhizae and Rhizobium on nutrient content and water relations in drought stressed alfalfa

The objective of this research was to study the effect of drought on nutrient content and leaf water status in alfalfa (Medicago sativa L. cv Aragón) plants inoculated with a mycorrhizal fungus and/or Rhizobium compared with noninoculated ones. The four treatments were: a) plants inoculated with Glomus fasciculatum and Rhizobium meliloti 102 F51 strain, (MR); b) plants inoculated with R. meliloti only (R); c) plants with G. fasciculatum only (M); and d) noninoculated plants (N). Nonmycorrhizal plants were supplemented with phosphorus and nonnodulated ones with nitrogen to achieve similar size and nutrient content in all treatments. Plants were drought stressed using two cycles of moisture stress and recovery. The components of total leaf water potential (osmotic and pressure potentials at full turgor), percentage of apoplastic water volume and the bulk modulus of elasticity of leaf tissue were determined. Macronutrient (N, P, K, Ca, S and Mg) and micronutrient (Co, Mo, Zn, Mn, Cu, Na, Fe and B) content per plant were also measured. Leaves of N and R plants had decreased osmotic potentials and increased pressure potentials at full turgor, with no changes either in the bulk modulus of elasticity or the percentage of apoplastic water upon drought conditions. By contrast, M and MR leaves did not vary in osmotic and turgor potentials under drought stress but had increased apoplastic water volume and cell elasticity (lowering bulk modulus). Drought stress decreased nutrient content of leaves and roots of noninoculated plants. R plants showed a decrease in nutrient content of leaves but maintained some micronutrients in roots. Leaves of M plants were similar in content of nutrients to N plants. However, roots of M and MR plants had significantly lower nutrient content. Results indicate an enhancement of nutrient content in mycorrhizal alfalfa plants during drought that affected leaf water relations during drought stress.     

A new model of carbon and phosphorus transfers in arbuscular mycorrhizas

Existing models of nutrient transfer in arbuscular mycorrhizal (AM) symbioses are inadequate as they do not explain the range of real responses seen experimentally. A computer simulation model was used to evaluate the novel hypotheses that mycorrhizal nutrient transfers were based solely on symbionts' internal needs, and that carbon and phosphorus transfers were quantitatively unlinked. To be plausible, simulated mycorrhizal plants would show a +/-50% variation in weight vs nonmycorrhizal controls, with a normal response distribution (mimicking a real data set). One plant and one arbuscular mycorrhizal fungus (AMF) growing in a soil volume were simulated, using C, P and nitrogen nutrient cycling and stoichiometry. C- and P-exchange rates were independent and could be varied at will. The model was tested at realistic nutrient concentrations and a full range of nutrient exchange rates. The model showed -20% to +55% range in mycorrhizal plant weight distributed close to normal, suggesting that the hypotheses were plausible. The model suggests that theoretical assumptions about mycorrhizas should be reassessed. The model worked only because the symbionts possessed incomplete information on their partner and environmental conditions. Conventional cost-benefit models do not work under these circumstances, but both mutualistic and parasitic interactions were successfully simulated.     

菌根真菌在红壤中对柑桔吸收磷肥的研究

接种一种球囊霉 Glomus sp.编号 CC-1于红壤中,使酸桔和枳实生苗的根系感染形成泡囊丛枝内生菌根。应用放射性同位素~(32)P 标记的可溶性磷肥和难溶性磷肥来研究菌根真菌对柑桔吸收磷肥的作用。放射性宏观自显影和显微自显影表明菌根真菌加强了柑桔对两类磷肥的吸收。放射性测量和化学分析证明有菌根柑桔实生苗的地上部含磷量以及从红壤中和从肥料中吸收的磷量比无菌根的增多,有菌根实生苗的生长反应也较好。     

接种AM真菌对玉米和油菜种间竞争及土壤无机磷组分的影响

在温室盆栽条件下,分别模拟单作、间作和尼龙网分隔种植,比较接种丛枝菌根(arbuscular mycorrhizal, AM)真菌Glomus intraradices和Glomus mosseae对菌根植物玉米和非菌根植物油菜生长和磷吸收状况的影响,并分析土壤中各无机磷组分的变化。结果发现,接种AM真菌可以促进土壤中难溶性磷(Ca₁₀-P和O-P)向有效态磷转化,并显著降低总无机磷含量 (P<0.05),显著提高菌根植物玉米的生物量和磷吸收量(P<0.05),特别是在间作体系中使玉米的磷营养竞争比率显著提高了45.0%-104.1% (P<0.05),显著降低了油菜的生物量和磷吸收量(P<0.05),从而增强了了菌根植物的竞争优势,降低了非菌根植物与菌根植物的共存能力。揭示了石灰性土壤中AM真菌对植物物种多样性的影响,有助于更加全面地理解AM真菌在农业生态系统中的作用。     

Adaptation of selected trees and grasses to low availability of phosphorus

High-nutrient-adapted and low-nutrient-adapted species of New Zealand tussock grasses (Chionochloa), barley (Hordeum), and several taiga trees were grown at three rates of phosphorus supply. Low-nutrient-adapted species in each group of species had similar (grasses) or lower (trees) capacities for phosphate absorption, were less efficient in producing biomass (i.e. had higher nutrient concentrations), and grew more slowly than high-nutrient-adapted species. I conclude that the major adaptation to low nutrient availability in each of these comparisons is a slow growth rate that reduces the annual nutrient requirement.     

Carbon Cost of the Fungal Symbiont Relative to Net Leaf P Accumulation in a Split-Root VA Mycorrhizal Symbiosis

Translocation of ¹⁴C-photosynthates to mycorrhizal (+ +), half mycorrhizal (0+), and nonmycorrhizal (00) split-root systems was compared to P accumulation in leaves of the host plant. Carrizo citrange seedlings (Poncirus trifoliata [L.] Raf. × Citrus sinensis [L.] Osbeck) were inoculated with the vesicular-arbuscular mycorrhizal fungus Glomus intraradices Schenck and Smith. Plants were exposed to ¹⁴ CO₂ for 10 minutes and ambient air for 2 hours. Three to 4% of recently labeled photosynthate was allocated to metabolism of the mycorrhiza in each inoculated root half independent of shoot P concentration, growth response, and whether one or both root halves were colonized. Nonmycorrhizal roots respired more of the label translocated to them than did mycorrhizal roots. Label recovered in the potting medium due to exudation or transport into extraradical hyphae was 5 to 6 times greater for (+ +) versus (00) plants. In low nutrient media, roots of (0+) and (+ +) plants transported more P to leaves per root weight than roots of (00) plants. However, when C translocated to roots utilized for respiration, exudation, etc., as well as growth is considered, (00) plant roots were at least as efficient at P uptake (benefit) per C utilized (cost) as (0+) and (+ +) plants. Root systems of (+ +) plants did not supply more P to leaves than (0+) plants in higher nutrient media, yet they still allocated twice the ¹⁴C-photosynthate to the mycorrhiza as did (0+) root systems. This indicates there is an optimal level of mycorrhizal colonization above which the plant receives no enhanced P uptake yet continues to partition photosynthates to metabolism of the mycorrhiza.     

Effect of mycorrhizal-enhanced leaf phosphate status on carbon partitioning, translocation and photosynthesis in cucumber

An assessment of the effects of arbuscular mycorrhizal (AM) infection on photosynthesis, carbon (C) allocation, translocation and biomass production of cucumber, grown in sand culture, was made using a previously determined phosphorus (P) supply (0·13 mol m^?3 P) which had a significant impact on AM infection. Separation of a direct effect of AM infection from an indirect one due to an enhanced leaf P status was achieved using a comparable non‐mycorrhizal treatment (NAM + P) supplemented with extra P (0·19 mol m^?3 P). Total leaf P concentration, specific leaf mass, photosynthetic capacity, and incorporation of ¹⁴C into non‐structural carbohydrate pools were dependent on leaf age. Both maximum and ambient photosynthetic rates were significantly higher in the youngest fully expanded leaves from AM and NAM + P plants which also had the higher leaf P concentrations. There were no differences in the total concentrations of starch, sucrose, raffinose or stachyose in young or old leaves among AM, non‐mycorrhizal (NAM) and NAM + P treatments. However, younger leaves of NAM plants showed a shift in ¹⁴C‐partitioning from stachyose and raffinose synthesis to starch accumulation. Determination of ADP‐glucose pyrophosphorylase (AGPase), sucrose synthase and sucrose phosphate synthase enzyme activities revealed that only AGPase activity was correlated with the increased incorporation rate of ¹⁴C into starch in young leaves of NAM plants. Although there were significant AM‐specific effects on C translocation to the root system, AM plants had similar rate of photosynthesis to NAM + P plants. These results suggest that the increase in photosynthetic rate in leaves of AM‐infected cucumber was due to an increased P status, rather than a consequence of a mycorrhizal ‘sink’ for assimilates.     

The effect of vesicular arbuscular mycorrhizal associations on growth of cereals

The growth of wheat seedlings which were already mycorrhizal when transplanted to a field deficient in phosphorus was improved compared with non-mycorrhizal controls, and grain yield was increased three-fold by the fungus, indicating that Endogone stimulated growth and increased yield. Differences between mycorrhizal and non-mycorrhizal wheat were eliminated by the application of phosphate fertilizer, indicating that the fungus does not enhance wheat growth in soils containing enough available phosphate. It is probable that the mycorrhizal effect is primarily to improve the supply of phosphate. There were clear relationships between spore number in the soil and mycorrhizal development and between the extent of root infection and increased growth. The extent of root infection was greatest in mycorrhizal plants in soil not supplemented with phosphate and it decreased in inoculated plants in the plot supplemented with superphosphate. The non-centrospermous and non-zygophyllaceous weeds growing on the experimental field had typical vesicular arbuscular infection and indigenous Endogone spores in their rhizospheres. The centrospermous plants were non-mycorrhizal and had no Endogone spores in their rhizospheres.     

Enhancement of nitrogen fixation in lentil,faba bean,and soybean by dual inoculation with Rhizobia and mycorrhizae

The combined effect of Vesicular Arbuscular Mycorrhizae (VAM) and Rhizobium on the cold season legumes, lentil and faba bean, as well as on summer legume, soybean, were studied in soils with low indeginous VA mycorrhizal spores. Inoculation of the plant with VA mycorrhizal fungi increased the level of mycorrhizal root infection of lentil, faba bean and soybean. The inoculation with Rhizobium had no significant effect on VA mycorrhizal infection percent, but VA mycorrhizal inoculation increased nodulation of the three legumes. The inoculation with Rhizobium alone significantly increased plant dry weight and N content of lentil and faba bean as well as seed yield of soybean. VA mycorrhizal inoculation also significantly increased plant dry weight and phosphorus content of the plants as did fertilization with superphosphate. Rock phosphate fertilization, however, had no significant effect on plant growth or phosphorus uptake. The addition of rock phosphate in combination with VA mycorrhizal inoculation significantly increased plant dry weight and P uptake of the plants. The dual inoculation with both rhizobia and mycorrhizae induced more significant increases in plant dry weight, N and P content of lentil and faba bean as well as seed yield of soybean than inoculation with either VA mycorrhizae or Rhizobium alone.     

菌根真菌与植物共生营养交换机制研究进展

菌根是陆地生态系统普遍存在的、由土壤中的菌根真菌侵染宿主植物根系形成的联合共生体.菌根的建立是以共生体双方的营养交换为基础的:菌根真菌从土壤中吸收氮、磷等营养物质并转运给宿主植物,供其生长;作为交换,植物则以脂质或糖的形式向菌根真菌提供其生长所必需的碳水化合物.近年来,菌根真菌与宿主植物间的营养交换机制一直是研究的热点,国内外对菌根真菌介导的植物营养物质吸收和转运机制的研究也取得了巨大进展.本文综述了丛枝和外生两种菌根真菌与宿主植物间营养交换的最新研究进展,尤其是碳、氮、磷等几种重要营养物质的吸收与双向转运机制,以及营养交换在菌根形成中的潜在调控作用,并对目前存在的关键问题和未来研究方向进行了分析和展望,这对菌根模型的建立及菌根效益的优化具有重要意义.     

Effect of arbuscular mycorrhizal colonization and two levels of compost supply on nutrient uptake and flowering of pelargonium plants

Two challenges frequently encountered in the production of ornamental plants in organic horticulture are: (1) the rate of mineralization of phosphorus (P) and nitrogen (N) from organic fertilizers can be too slow to meet the high nutrient demand of young plants, and (2) the exclusive use of peat as a substrate for pot-based plant culture is discouraged in organic production systems. In this situation, the use of beneficial soil microorganisms in combination with high quality compost substrates can contribute to adequate plant growth and flower development. In this study, we examined possible alternatives to highly soluble fertilizers and pure peat substrates using pelargonium (Pelargonium peltatum L’Her.) as a test plant. Plants were grown on a peat-based substrate with two rates of compost addition and with and without arbuscular mycorrhizal (AM) fungi. Inoculation with three different commercial AM inocula resulted in colonization rates of up to 36% of the total root length, whereas non-inoculated plants remained free of root colonization. Increasing the rate of compost addition increased shoot dry weight and shoot nutrient concentrations, but the supply of compost did not always completely meet plant nutrient demand. Mycorrhizal colonization increased the number of buds and flowers, as well as shoot P and potassium (K) concentrations, but did not significantly affect shoot dry matter or shoot N concentration. We conclude that addition of compost in combination with mycorrhizal inoculation can improve nutrient status and flower development of plants grown on peat-based substrates.     

Plant–fungus competition for nitrogen erases mycorrhizal growth benefits of Andropogon gerardii under limited nitrogen supply

Considered to play an important role in plant mineral nutrition, arbuscular mycorrhizal (AM) symbiosis is a common relationship between the roots of a great majority of plant species and glomeromycotan fungi. Its effects on the plant host are highly context dependent, with the greatest benefits often observed in phosphorus (P)‐limited environments. Mycorrhizal contribution to plant nitrogen (N) nutrition is probably less important under most conditions. Moreover, inasmuch as both plant and fungi require substantial quantities of N for their growth, competition for N could potentially reduce net mycorrhizal benefits to the plant under conditions of limited N supply. Further compounded by increased belowground carbon (C) drain, the mycorrhizal costs could outweigh the benefits under severe N limitation. Using a field AM fungal community or a laboratory culture of Rhizophagus irregularis as mycorrhizal inoculants, we tested the contribution of mycorrhizal symbiosis to the growth, C allocation, and mineral nutrition of Andropogon gerardii growing in a nutrient‐poor substrate under variable N and P supplies. The plants unambiguously competed with the fungi for N when its supply was low, resulting in no or negative mycorrhizal growth and N‐uptake responses under such conditions. The field AM fungal communities manifested their potential to improve plant P nutrition only upon N fertilization, whereas the R. irregularis slightly yet significantly increased P uptake of its plant host (but not the host's growth) even without N supply. Coincident with increasing levels of root colonization by the AM fungal structures, both inoculants invariably increased nutritional and growth benefits to the host with increasing N supply. This, in turn, resulted in relieving plant P deficiency, which was persistent in non‐mycorrhizal plants across the entire range of nutrient supplies.     

The involvement of mycorrhizas in assessment of genetically dependent efficiency of nutrient uptake and use

This article summarises the way in which mycorrhizal infection of roots affects the mineral nutrition of plants and how the symbiosis may interact with the evaluation of efficiency of nutrient uptake and use by plants. A brief account of the processes of infection and the way they are affected by host genotype and environmental conditions is given and the relationships between this and mineral nutrition (especially phosphate nutrition) are outlined.The interactions between mycorrhizal infection and P efficiency are considered at two levels. Mycorrhizas may act as general modifiers of efficiency regardless of the extent to which the plants are infected and in some mycorrhiza-dependent plants infection may change the ranking of genotypes. The extent of infection is also under genetic control and shows considerable variability between genotypes in some species. This variation could be used in programs to select varieties in which infection is rapid and nutrient uptake from nutrient deficient or low input systems is, in consequence, increased.     

Alteration of nutrient allocation and transporter genes expression in rice under N,P, K,and Mg deficiencies

Nitrogen (N), phosphorus (P), potassium (K), and magnesium (Mg) have essential physiological functions in plants. Their interactions in plants are not fully understood especially at the molecular level. In this study, we detected the physiological and molecular responses of rice plants at the vegetative growth phase to N, P, K, and Mg starvations. Deficiencies of N and P resulted in accumulation of soluble sugar and starch in the leaves. The root to shoot ratio increased under N and P deficiencies, but decreased under K and Mg deficiencies. In addition, deficiency of either K or Mg resulted in accumulation of the other cation in shoots. Moreover, K starvation decreased both K and soluble sugar contents in the roots pronouncedly. RT-PCR analysis showed that several sugar transporter genes in the leaves orchestrated with sugar accumulation induced by the nutrient shortages. Expression of a high affinity K transporter gene (OsHAK1) and a putative Mg transporter gene (OsMGT) showed opposite down- and up-regulation in the roots by K starvation. These findings suggest that deficiencies of the major nutrients suppressed the export of carbohydrates from source leaves. The regulated sugar and nutrient transporter genes investigated in this study could be used for elucidating the molecular mechanism of plants in their adaptation to varied nutrient supply.