Investigations on phosphate uptake and polyphosphate metabolism by mycorrhized and nonmycorrhized roots of beech and pine as investigated by in vivo 31P-NMR

Comparative in vivo ³¹P-NMR studies of mycorrhized and nonmycorrhized roots of Fagus sylvatica and Pinus sylvestris and of the fungus Suillus bovinus in pure culture have produced interesting new data. With respect to intracellular compartments and pH, ³¹P-NMR spectroscopy showed that the spectrum of the mycorrhiza results from simple superimposition of the spectra of its symbionts. A special method of cyclic phosphate supply followed by block averaging of the NMR spectra was used to determine the kinetic behaviour of phosphate uptake and storage and its incorporation into polyphosphate at a constant external pH of 5.5. Mycorrhized roots and pure fungus showed transformation of accumulated inorganic phosphate into mobile polyphosphate with a medium chain length. Transformation of mobile into immobile polyphosphate either with a long chain length or in a granular state was also observed. Thus, two different types of fungal polyphosphate could be verified. Deficiency of external phosphate initiated the mobilization of internal phosphate, transforming stored polyphosphate into phosphate. It could be shown that a high fungal mass renders mycorrhizal phosphate metabolism less sensitive to external variation in nutrient concentration. The central role of the fungus in regulating mycorrhizal phosphate metabolism is discussed.     

31P NMR for the study of P metabolism and translocation in arbuscular mycorrhizal fungi

³¹P nuclear magnetic resonance (NMR) spectroscopy was used to study phosphate (P) metabolism in mycorrhizal and nonmycorrhizal roots of cucumber (Cucumis sativus L) and in external mycelium of the arbuscular mycorrhizal (AM) fungus Glomus intraradices Schenck & Smith. The in vivo NMR method allows biological systems to be studied non-invasively and non-destructively. ³¹P NMR experiments provide information about cytoplasmic and vacuolar pH, based on the pH-dependent chemical shifts of the signals arising from the inorganic P (P_i) located in the two compartments. Similarly, the resonances arising from α, β and γ phosphates of nucleoside triphosphates (NTP) and nucleoside diphosphates (NDP) supply knowledge about the metabolic activity and the energetic status of the tissue. In addition, the kinetic behaviour of P uptake and storage can be determined with this method. The ³¹P NMR spectra of excised AM fungi and mycorrhizal roots contained signals from polyphosphate (PolyP), which were absent in the spectra of nonmycorrhizal roots. This demonstrated that the P_i taken up by the fungus was transformed into PolyP with a short chain length. The spectra of excised AM fungi revealed only a small signal from the cytoplasmic P_i, suggesting a low cytoplasmic volume in this AM fungus. This revised version was published online in June 2006 with corrections to the Cover Date.     

A 31P nuclear magnetic resonance study of phosphate levels in roots of ectomycorrhizal and nonmycorrhizal plants of Castanea sativa Mill.

 ³¹P-Nuclear Magnetic Resonance (NMR) was used to assess phosphate distribution in ectomycorrhizal and nonmycorrhizal roots of Castanea sativa Mill. as well as in the mycorrhizal fungus Pisolithus tinctorius in order to gain insight into phosphate trafficking in these systems. The fungus P. tinctorius accumulated high levels of polyphosphates during the rapid phase of growth. Mycorrhizal and nonmycorrhizal roots accumulate orthophosphate. Only mycorrhizal roots presented polyphosphates. The content in polyphosphates increased along the 3 months of mycorrhiza formation. In mycorrhizal roots of plants cultured under axenic conditions, the orthophosphate pool decreased along the culture time. In nonmycorrhizal roots the decrease in the orthophosphate content was less pronounced. The level of orthophosphate in mycorrhizal roots was significantly lower than in nonmycorrhizal ones, which indicates that this system relies upon the fungal polyphosphates as a major source of phosphate. Received: 28 July 1998 / Accepted: 21 October 1998     

Effects of aluminium on polyphosphate mobilization of the ectomycorrhizal fungus Suillus bovinus

Aluminium toxicity may be an important factor in the decline in vitality of many forest trees and the associated ectomycorrhizal fungal flora. In this study, comparative in vivo ³¹P NMR investigations on Al-adapted and non-Aladapted fungus of Suillus bovinus in pure culture have produced interesting new data. With respect to intracellular compartments, ³¹P NMR spectroscopy showed the spectra to differ in a peak-6 ppm appearing in the spectra of the A1-adapted fungus indicating terminal phosphate groups of mobile polyphosphate. Thus, in the Al-adapted fungus the average chain length of mobile polyphosphate is considerably shorter than in the non-Al-adapted fungus. A special method of cyclic phosphate supply followed by block averaging of the NMR spectra was used to determine the kinetic behaviour of phosphate uptake, storage and incorporation into polyphosphate at a constant external pH 3.5. While the Al-adapted fungus showed resistance to Al, an irreversible break-down in phosphate metabolism of the non-Al-adapted fungus by exposure to Al was caused. In comparison with the non-Al-adapted fungus supplied by nutrient solutions omitting Al, the Al-adapted fungus showed higher levels in both phosphate uptake and mobile polyphosphate concentration. As a consequence of these results a de-toxification of freely mobile Al-ions into a stable and insoluble complex in the Al-adapted fungus is considered to be due to a capture of intracellular Al by mobile polyphosphate of shorter chain length.     

Behavior of mercury in a soil–plant system as affected by inoculation with the arbuscular mycorrhizal fungus Glomus mosseae

Effects of inoculation with the arbuscular mycorrhizal (AM) fungus Glomus mosseae on the behavior of Hg in soil–plant system were investigated using an artificially contaminated soil at the concentrations of 0, 1.0, 2.0, and 4.0 mg Hg kg⁻¹. Mercury accumulation was lower in mycorrhizal roots than in nonmycorrhizal roots when Hg was added at the rates of 2.0 and 4.0 mg kg⁻¹, while no obvious difference in shoot Hg concentration was found between mycorrhizal and nonmycorrhizal treatments. Mycorrhizal inoculation significantly decreased the total and extractable Hg concentrations in soil as well as the ratio of extractable to total Hg in soil. Equilibration sorption of Hg by soil was investigated, and the results indicated that mycorrhizal treatment enhanced Hg sorption on soil. The uptake of Hg was lower by mycorrhizal roots than by nonmycorrhizal roots. These experiments provide further evidence for the role of mycorrhizal inoculation in increasing immobilization of Hg in soil and reducing the uptake of Hg by roots. Calculation on mass balance of Hg in soil suggests the presence of Hg loss from soil presumably through evaporation, and AM inoculation enhanced Hg evaporation. This was evidenced by a chamber study to detect the Hg evaporated from soil.     

Phosphorus Distribution in Red Pine Roots and the Ectomycorrhizal Fungus Hebeloma arenosa

Red pines (Pinus resinosa Ait.) were grown in a pasteurized sandy loam either unamended with phosphate or fertilized with one of two levels of phosphate (34 or 136 mg/kg) as superphosphate, and with and without addition of Hebeloma arenosa inoculum. Shoot and total dry weights of mycorrhizal seedlings grown in soil unamended with P were greater than those for nonmycorrhizal seedlings grown in the same soil, but less than the dry weights of seedlings grown in soil amended with middle to high levels of P. Mycorrhizal infection was inhibited at the highest level of P amendment. ³¹P nuclear magnetic resonance spectra of intact mycorrhizal roots showed the presence of two dominant peaks, orthophosphate (Pi) and polyphosphate (polyP). The polyP peak was absent in spectra of nonmycorrhizal roots. The ratio for areas under the two peaks, Pi/polyP, was 1.8 for mycorrhizal roots grown in both unamended soil and soil that had received middle levels of superphosphate. Apparently, the fungus strongly mediates the supply of phosphate to the tree through the production of polyP, even at growth-limiting levels of soil P, and regulates compartmentalization of P in the mycorrhizal roots.     

The effects of heavy metals,content of nutrients and inoculation with mycorrhizal fungi on the level of terpenoids in roots of <Emphasis Type="Italic">Pinus sylvestris</Emphasis> seedlings

The effect of two substrates differing in the amount of toxic metals and nutrients on the content of volatile organic compounds in roots of Pinus sylvestris seedlings inoculated with Paxillus involutus, Laccaria laccata, Suillus luteus and a strain of the ectendomycorrhizal fungus Mrg X (Ascomycotina) in pot experiment was investigated. Volatile compounds extracted with a supercritical fluid extraction were primarily terpenes and sesquiterpenes, and they were qualitatively the same in roots of mycorrhizal and nonmycorrhizal plants grown on both substrates. The major monoterpenes were α-pinene, Δ³-carene and β-pinene. On the unpolluted substrate, inoculation of plants with the fungi resulted in a similar amount of the volatiles, while on the polluted substrate the mycorrhizal roots showed significantly lower content of terpenoids than nonmycorrhizal roots. The mycorrhizal fungi showed different effects on the concentrations of several terpenoids on both substrates.     

Increases in α-mannosidase activity in the arbuscular mycorrhizal symbiosis of Allium schoenoprasum

 Mycorrhizal and nonmycorrhizal roots of Allium schoenoprasum were tested for activities of α-mannosidase, β-glucosidase and arabinosidase. Mannosidase activity was higher by a factor of two in mycorrhizal than in nonmycorrhizal root extracts. The apparent molecular weight of the enzyme was 152 kDa and its K_M was 1.25 mM in colonized roots and 1.85 mM in uncolonized roots. α-Mannosidase activity was further characterized by an acid pH optimum and Zn²⁺ dependency. No significant differences could be found between mycorrhizal and nonmycorrhizal roots for β-glucosidase and arabinosidase activities. Accepted: 28 August 1995     

The mycorrhizal fungus Paxillus involutus transports magnesium to Norway spruce seedlings. Evidence from stable isotope labeling

Although it is well established that ectomycorrhizas improve the mineral nutrition of forest trees, there has been little evidence that they mediate uptake of divalent cations such as Mg. We grew nonmycorrhizal seedlings and seedlings mycorrhizal with Paxillus involutus Batsch in a sand culture system with two compartments separated by a 45-μm Nylon mesh. Hyphae, but not roots, can penetrate this net. Labeling the compartment only accessible to hyphae with ²⁵Mg showed that hyphae of the ectomycorrhizal fungus Paxillus involutus transported Mg to their host plant. No label was found in nonmycorrhizal control plants. Our data support the idea that ectomycorrhizas are important for the Mg nutrition of forest trees. This revised version was published online in June 2006 with corrections to the Cover Date.     

Improved Tolerance of <Emphasis Type="Italic">Acacia nilotica</Emphasis> to Salt Stress by Arbuscular Mycorrhiza, <Emphasis Type="Italic">Glomus fasciculatum</Emphasis> may be Partly Related to Elevated K/Na Ratios in Root and Shoot Tissues

A pot experiment was conducted to examine the effect of arbuscular mycorrhizal fungus, Glomus fasciculatum, and salinity on the growth of Acacia nilotica. Plants were grown in soil under different salinity levels (1.2, 4.0, 6.5, and 9.5 dS m⁻¹). In saline soil, mycorrhizal colonization was higher at 1.2, 4.0, and 6.5 dS m⁻¹ salinity levels in AM-inoculated plants, which decreased as salinity levels further increased (9.5 dS m⁻¹). Mycorrhizal plants maintained greater root and shoot biomass at all salinity levels compared to nonmycorrhizal plants. AM-inoculated plants had higher P, Zn, and Cu concentrations than uninoculated plants. In mycorrhizal plants, nutrient concentrations decreased with the increasing levels of salinity, but were higher than those of the nonmycorrhizal plants. Mycorrhizal plants had greater Na concentration at low salinity levels (1.2, 4.0 dS m⁻¹), which lowered as salinity levels increased (6.5, 9.5 dS m⁻¹), whereas Na concentration increased in control plants. Mycorrhizal plants accumulated a higher concentration of K at all salinity levels. Unlike Na, the uptake of K increased in shoot tissues of mycorrhizal plants with the increasing levels of salinity. Our results indicate that mycorrhizal fungus alleviates deleterious effects of saline soils on plant growth that could be primarily related to improved P nutrition. The improved K/Na ratios in root and shoot tissues of mycorrhizal plants may help in protecting disruption of K-mediated enzymatic processes under salt stress conditions.     

Concentrations of terpenes in mycorrhizal roots of Scots pine (Pinus sylvestris L.) seedlings grown in vitro

The effect of Paxillus involutus, Laccaria laccata, Suillus luteus, S. bovinus, Hebeloma crustuliniforme and a strain of the ectendomycorrhizal fungus Mrg X (Ascomycotina) on the content of volatile organic compounds in roots of Pinus sylvestris seedlings grown in vitro was investigated. Volatile compounds extracted with a supercritical fluid extraction were primarily terpenes and sesquiterpenes and qualitatively were the same in roots of mycorrhizal and nonmycorrhizal plants. The major monoterpenes were α-pinene, Δ³-carene and β-pinene. Inoculation of plants with the fungi resulted in statistically non-significant increases in the total amount of the volatiles. The mycorrhizal fungi showed diversified effect on the concentrations of several terpenoids.     

Effects of monovalent cations on phosphate accumulation and storage of the ectomycorrhizal fungus Suillus bovinus

Comparative in vivo ³¹P-NMR studies of the fungus Suillus bovinus (L.: Fr.) O. Kuntze in pure culture have produced interesting new data. To investigate the response of phosphate metabolism to a change in external monovalent cations, samples were exposed to a Hoagland solution containing different monovalent cations Li⁺, Na⁺, K⁺, or Rb⁺ at 10 mM concentration. A method of nutrient cycling during analysis where the cation was changed and the phosphate kept constant allowed us to determine the kinetics of phosphate accumulation, storage and incorporation into polyphosphate following exposure to the range of test cations. Different external monovalent cations had different effects upon changes in the content of both phosphate and polyphosphate. Treatment with Li⁺, Na⁺, or Rb⁺ resulted in a change in phosphate accumulation to 60, 73, and 107% and in content of the intracellular mobile polyphosphate (polyP) to 119, 112, and 94%, respectively, compared with the control taken as 100%. The effect of each cation is related to its position in the periodic table. Reversing this process, i.e., exchanging with K⁺, returned phosphate metabolism to normal. Although, the increase in depolarization of the cell membrane should affect the internal pH, fungal metabolism using energy requiring mechanisms appeared necessary to maintain the intracellular pH. Thus, increasing contents of mobile polyP were the consequence of an increasing energy demand. On the other hand, the increasing depolarization of the cell membrane following the sequence Rb⁺ < K⁺ < Na⁺ < Li⁺ inhibited the net P_i accumulation. Furthermore, it is postulated that the P_i accumulation was also regulated by the intracellular content in polyP.     

Contribution of an arbuscular mycorrhizal fungus to the uptake of cadmium and nickel in bean and maize plants

Two experiments were carried out in pots with three compartments, a central one for root and hyphal growth and two outer ones which were accessible only for hyphae of the arbuscular mycorrhizal fungus, Glomus mosseae ([Nicol. and Gerd.] Gerdemann and Trappe). In the first experiment, mycorrhizal and nonmycorrhizal bean (Phaseolus vulgaris L.) plants were grown in two soils with high geogenic cadmium (Cd) or nickel (Ni) contents. In the second experiment, mycorrhizal and nonmycorrhizal maize (Zea mays L.) or bean plants were grown in a non-contaminated soil in the central compartment, and either the Cd- or Ni-rich soil in the outer compartments. In additional pots, mycorrhizal plants were grown without hyphal access to the outer compartments. Root and shoot dry weight was not influenced by mycorrhizal inoculation, but plant uptake of metals was significantly different between mycorrhizal and nonmycorrhizal plants. In the first experiment, the contribution of mycorrhizal fungi to plant uptake accounted for up to 37% of the total Cd uptake by bean plants, for up to 33% of the total copper (Cu) uptake and up to 44% of the total zinc (Zn) uptake. In contrast, Ni uptake in shoots and roots was not increased by mycorrhizal inoculation. In the second experiment, up to 24% of the total Cd uptake and also up to 24% of the total Cu uptake by bean could be attributed to mycorrhizal colonisation and delivery by hyphae from the outer compartments. In maize, the mycorrhizal colonisation and delivery by hyphae accounted for up to 41% of the total Cd uptake and 19% of the total Cu uptake. Again, mycorrhizal colonisation did not contribute to Ni uptake by bean or maize. The results demonstrate that the arbuscular mycorrhizal fungus contributed substantially not only to Cu and Zn uptake, but also to uptake of Cd (but not Ni) by plants from soils rich in these metal cations. Deceased 21 September 1996 Deceased 21 September 1996     

Nitrogen limitation in mycorrhizal Norway spruce (Picea abies) seedlings induced mycelial foraging for ammonium: implications for Ca and Mg uptake

Although many studies support the importance of the external mycelium for nutrient acquisition of ectomycorrhizal plants, direct evidence for a significant contribution to host nitrogen nutrition is still scarce. We grew nonmycorrhizal seedlings and seedlings mycorrhizal with Paxillus involutus (Batsch) Fr. in a sand culture system with two compartments separated by a 45-m Nylon mesh. Hyphae, but not roots, can penetrate this net. Nutrient solutions were designed to limit seedling growth by nitrogen. Hyphal density in the hyphal compartment, host N status and shoot growth of mycorrhizal seedlings significantly increased in response to NH₄ ⁺ addition to the hyphal compartment. Labeling the compartment only accessible to hyphae with ¹⁵NH₄ ⁺ showed that the increase in N uptake in the mycorrhizal seedlings was a result of hyphal N acquisition from the hyphal compartment. These results indicate that hyphae of P. involutus may actively forage into N-rich patches and improve host N status and growth. In the mycorrhizal seedlings, which received additional NH₄ ⁺ via their external mycelium, the increase in NH₄ ⁺ supply less negatively affected Ca and Mg uptake than in nonmycorrhizal seedlings, where the additional NH₄ ⁺ was directly supplied to the roots. This was most likely due to the close link of NH₄ ⁺ uptake and H⁺ extrusion, which, in the nonmycorrhizal seedlings, lead to a strong acidification in the root compartment, and subsequently reduced Ca and Mg uptake, whereas in the mycorrhizal seedlings the site of intensive NH₄ ⁺ uptake and acidification was in the hyphal and not in the root compartment. Our data support the idea that the ectomycorrhizal mycelium connected to an N-deficient host may actively forage for N. The mycelium may also be important as a biological buffer system ameliorating negative influence of high NH₄ ⁺ supply on cation uptake.     

Accumulation of sesquiterpenoid cyclohexenone derivatives induced by an arbuscular mycorrhizal fungus in members of the Poaceae

Sixty one members of the Poaceae, including various cereals, were grown in defined nutrient media with and without the arbuscular mycorrhizal (AM) fungus, Glomus intraradices Schenk & Smith. The roots of all species investigated were colonized by the AM fungus, however, to different degrees and independent of their systematic position. High-performance liquid chromatographic analyses of methanolic extracts from the roots of mycorrhizal and nonmycorrhizal species revealed dramatic changes in the patterns of UV-detectable products along with a widespread occurrence of AM-fungus-induced accumulation of sesquiterpenoid cyclohexenone derivatives. The latter occur most often in the tribes Poeae, Triticeae and Aveneae. Some additional control experiments on plant infection with pathogens (Gaeumannomyces graminis) and Drechslera sp.) or an endophyte (Fusarium sp.), as well as application of abiotic stress, proved that the metabolism of these terpenoids is part of a response pattern of many gramineous roots in their specific reaction to AM fungal colonization. Received: 23 October 1996 / Accepted 11 December 1996     

Effects of liming and mycorrhizal colonization on soil phosphate depletion and phosphate uptake by maize (Zea mays L.) and soybean (Glycine max L.) grown in two tropical acid soils

The effects of liming and inoculation with the arbuscular mycorrhizal fungus, Glomus intraradices Schenck and Smith on the uptake of phosphate (P) by maize (Zea mays L.) and soybean (Glycine max [L.] Merr.) and on depletion of inorganic phosphate fractions in rhizosphere soil (Al-P, Fe-P, and Ca-P) were studied in flat plastic containers using two acid soils, an Oxisol and an Ultisol, from Indonesia. The bulk soil pH was adjusted in both soils to 4.7, 5.6, and 6.4 by liming with different amounts of CaCO₃.In both soils, liming increased shoot dry weight, total root length, and mycorrhizal colonization of roots in the two plant species. Mycorrhizal inoculation significantly increased root dry weight in some cases, but much more markedly increased shoot dry weight and P concentration in shoot and roots, and also the calculated P uptake per unit root length. In the rhizosphere soil of mycorrhizal and non-mycorrhizal plants, the depletion of Al-P, Fe-P, and Ca-P depended in some cases on the soil pH. At all pH levels, the extent of P depletion in the rhizosphere soil was greater in mycorrhizal than in non-mycorrhizal plants. Despite these quantitative differences in exploitation of soil P, mycorrhizal roots used the same inorganic P sources as non-mycorrhizal roots. These results do not suggest that mycorrhizal roots have specific properties for P solubilization. Rather, the efficient P uptake from soil solution by the roots determines the effectiveness of the use of the different soil P sources. The results indicate also that both liming and mycorrhizal colonization are important for enhancing P uptake and plant growth in tropical acid soils.     

Zinc Phosphate Transformations by the Paxillus involutus/Pine Ectomycorrhizal Association

In this research, we investigate zinc phosphate transformations by Paxillus involutus/pine ectomycorrhizas using zinc-resistant and zinc-sensitive strains of the ectomycorrhizal fungus under high- and low-phosphorus conditions to further understand fungal roles in the transformation of toxic metal minerals in the mycorrhizosphere. Mesocosm experiments with ectomycorrhizas were performed under sterile conditions with zinc phosphate localized in cellophane bags: zinc and phosphorus mobilization and uptake by the ectomycorrhizal biomass were analyzed. In the presence of a phosphorus source, an ectomycorrhizal association with a zinc-resistant strain accumulated the least zinc compared to a zinc-sensitive ectomycorrhizal association and non-mycorrhizal plants. Under low-phosphorus conditions, mycorrhizal seedlings infected with the zinc-resistant strain increased the dissolution of zinc phosphate and zinc accumulation by the plant. Extended X-ray absorption fine structure analysis of both mycorrhizal and nonmycorrhizal roots showed octahedral coordination of zinc by oxygen-containing ligands such as carboxylates or phosphate. We conclude that zinc phosphate solubilization and zinc and phosphorus uptake by the association depend on ectomycorrhizal infection, strain of the mycobiont, and the phosphorus status of the matrix.     

Carbon Uptake and the Metabolism and Transport of Lipids in an Arbuscular Mycorrhiza

Both the plant and the fungus benefit nutritionally in the arbuscular mycorrhizal symbiosis: The host plant enjoys enhanced mineral uptake and the fungus receives fixed carbon. In this exchange the uptake, metabolism, and translocation of carbon by the fungal partner are poorly understood. We therefore analyzed the fate of isotopically labeled substrates in an arbuscular mycorrhiza (in vitro cultures of Ri T-DNA-transformed carrot [Daucus carota] roots colonized by Glomus intraradices) using nuclear magnetic resonance spectroscopy. Labeling patterns observed in lipids and carbohydrates after substrates were supplied to the mycorrhizal roots or the extraradical mycelium indicated that: (a) ¹³C-labeled glucose and fructose (but not mannitol or succinate) are effectively taken up by the fungus within the root and are metabolized to yield labeled carbohydrates and lipids; (b) the extraradical mycelium does not use exogenous sugars for catabolism, storage, or transfer to the host; (c) the fungus converts sugars taken up in the root compartment into lipids that are then translocated to the extraradical mycelium (there being little or no lipid synthesis in the external mycelium); and (d) hexose in fungal tissue undergoes substantially higher fluxes through an oxidative pentose phosphate pathway than does hexose in the host plant.     

Local measurements of nitrate and potassium fluxes along roots of maritime pine. Effects of ectomycorrhizal symbiosis

In order to assess the actual role of ectomycorrhizae in ion uptake by the ectomycorrhizal root system, we used a microelectrode ion flux estimation methodology that provided access to local values of net fluxes. This made it possible to investigate the heterogeneity of ion fluxes along the different types of roots of Pinus pinaster associated or not with ectomycorrhizal species. We compared two fungi able to grow with nitrate in pure culture, Rhizopogon roseolus and Hebeloma cylindrosporum, the former having a positive effect on host tree shoot growth (c. +30%) and the latter a negative effect (c.? 30%). In non‐mycorrhizal plants (control), NO₃^– was taken up at higher rates by the short roots than by the long ones, whereas K⁺ uptake occurred mainly in growing apices of long roots. In mycorrhizal plants, H. cylindrosporum did not modify K⁺ uptake and even decreased NO₃^– uptake at the level of ectomycorrhizal short roots, whereas R. roseolus strongly increased K⁺ and NO₃^– fluxes at the level of ectomycorrhizal short roots without any modification of the fluxes measured along the fungus‐free long roots. The measurement of ion influxes at the surface of the ectomycorrhizal roots can provide a way to reveal actual effects of mycorrhizal association on ion transport in relation to mycorrhizal efficiency in natural conditions.     

BIOMASS ALLOCATION IN AN AGROPYRON SMITHII-GLOMUS SYMBIOSIS

The type of response the steppe grass Agropyron smithii Rydb. exhibits to mycorrhizal fungi is dependent upon where along the resource gradient the experiment is conducted. At the lower end of a soil-phosphorus (P) gradient (2 μg P g^-1 soil) mycorrhizal A. smithii biomass was 35% greater than its nonmycorrhizal counterpart; while at the high end of the gradient (20 μg P g^-1 soil) no difference in host biomass production was evident. Mycorrhizal plants were shorter in stature (P < 0.002) and produced more tillers (P < 0.0001) than nonmycorrhizal plants. The stature and tiller response was more pronounced at the high end of the soil-P gradient. The partitioning of the mycorrhizal fungus was influenced by the soil-P gradient. At the lower end of the gradient the majority of the fungus was produced extramatrically, where over 5,000 cm of hyphae per plant were encountered after 10 wk growth. While at the high end of the gradient, the majority of fungus was produced internally. The highest mycorrhizal dependency values were measured at the lower end of the soil-P gradient where cost to the host (as measured by fungus biomass expressed as a percentage of the root system) was also highest. Increasing shoot and root-P concentrations were correlated with decreasing external fungus biomass. A positive association between tiller production and internal fungus biomass also existed. The significance of these findings, as well as the need to consider characteristics which measure fitness of a host in evaluating the mycorrhizal association of grasses, are discussed.