Glycine-Glomus-Rhizobium Symbiosis : VI. Photosynthesis in Nodulated, Mycorrhizal, or N- and P-Fertilized Soybean Plants

Soybean (Glycine max [L.] Merr. cv Hobbit) plants were grown in a growth chamber for 56 days in a phosphorus- and nitrogen-deficient soil and were colonized by the vesicular-arbuscular mycorrhizal (VAM) fungus Glomus mosseae (Nicol. & Gerd) Gerd. and Trappe and Rhizobium japonicum strain USDA 136, or by either organism alone, or by neither. Non-VAM plants received supplemental phosphorus and nonnodulated plants supplemental nitrogen to achieve the same rate of growth in all treatments. Plants of all four treatments had the same (P > 0.05) dry weights at harvest, but VAM plants had higher rates of CO₂ exchange (CER, P < 0.05) and lower leaf P concentrations (P < 0.01). Leaf nitrogen concentrations were lower in nodulated than in nitrogen-supplemented plants (P < 0.01) while starch concentrations were higher (P < 0.01). There was a significant negative relationship between nitrogen and starch (r = −0.989). Statistical evaluation of the data showed that some parameters (CER, leaf area and phosphorus content) were associated with phosphorus nutrition (or the presence of the VAM fungus), others (leaf fresh weight and root dry weight) with nitrogen nutrition (or the presence of Rhizobium), and some (leaf nitrogen and starch content) by both factors. The development of microsymbiont structures and nodule activity were significantly lower in the tripartite association than in plants colonized by one endophyte only. The findings suggest that endophyte effects go beyond those of simple nutrition and associated source-sink relationships.     

Glycine-Glomus-Rhizobium Symbiosis: V. Effects of Mycorrhiza on Nodule Activity and Transpiration in Soybeans under Drought Stress

Soybean (Glycine max [L.] Merr.) plants were nodulated (Bradyrhizobium japonicum) and either inoculated with the vesicular-arbuscular mycorrhizal (VAM) fungus Glomus mosseae (Nicol. & Gerd.) Gerd. and Trappe or left uncolonized. All plants were grown unstressed for 21 days initially. After this period, some VAM and non-VAM plants were exposed to four 8-day drought cycles while others were kept well watered. Drought cycles were terminated by rewatering when soil moisture potentials reached −1.2 megapascal. Nodule development and activity, transpiration, leaf conductance, leaf and root parameters including fresh and dry weight, and N and P nutrition of VAM plants and of non-VAM, P-fed plants grown under the same controlled conditions were compared. All parameters, except N content, were greater in VAM plants than in P-fed, non-VAM plants when under stress. The opposite was generally true in the unstressed comparisons. Transpiration and leaf conductance were significantly greater in stressed VAM than in non-VAM plants during the first half of the final stress cycle. Values for both VAM and non-VAM plants decreased linearly with time during the cycle and converged at a high level of stress (−1.2 megapascal). Effects of VAM fungi on the consequences of drought stress relative to P nutrition and leaf gas exchange are discussed in the light of these findings and those reported in the literature.     

Light effects in mycorrhizal soybeans

Soybean (Glycine max. L. Merr.) plants were grown in an experiment with a 3 × 3 factorial design using different levels of light (170, 350, and 700 μE·m⁻²·s⁻¹) and P as factors. Plants were grown in a greenhouse in pot cultures using a soil low in plant-available P under three P regimes: no additional P, P added as KH₂PO₄, or P uptake enhanced by colonization of the host plant with the vesicular-arbuscular mycorrhizal (VAM) fungus Glomus fasciculatum (Thaxt. sensu Gerd.) Gerd. and Trappe. Development of the VAM fungal endophyte and of plants under all three P regimes was depressed by limiting light. However, the growth response of VAM plants to increasing light relative to non-VAM plants in the absence of additional P increased while the response relative to non-VAM plants with additional P decreased slightly. The highly significant interaction between the factors (P < 0.001) of the experiment was due to differences in the magnitude and direction of simple effects of the factors. The implications of these differences in terms of source-sink relationships of the symbionts and the value of different non-VAM controls in interpreting VAM effects are discussed.     

The Glycine–Glomus–Rhizobium symbiosis. IV. Interactions between the mycorrhizal and nitrogen-fixing endophytes

Abstract Soybean (Glycine max (L.) Men) plants were grown under controlled conditions in an experiment designed as a 4 × 4 factorial. The factors were N or P nutrition, with different strains of Rhizobium japonicum or N-fertilization as levels of the first factor and different species of vesicular-arbuscular mycorrhizal (VAM) fungi or P fertilization as levels of the other. Organisms used were R. japonicum strains USDA 110, USDA 136, and 61A118, and the VAM fungi Glomus versiforme (Karst.) Berch, Glomus fasciculatum (Thaxt. sensu Gerd.) Gerd. and Trappe, and Glomus mosseae (Nicol. & Gerd.) Gerd. and Trappe. There were 16 treatments: nine Rhizobium + Glomus combinations, three Rhizobium + V and three Glomus+ N combinations, and one non-symbiotic set of plants supplied with N + P. The tripartite symbioses were evaluated by analysis of variance against the Rhizobium + P and Glomus + N comparison treatments for effects on root and leaf dry mass, root N and P content, nodule mass and activity, and VAM colinization. Significant to highly significant main effects and interactions were found in virtually all evaluations due to both Rhizobium strain and VAM–fungal species. We conclude that different endophyte isolates affect not only the host plant, but also the development and function of their co-endophytes. These findings establish the existence of inter-endophyte compatibility, an important consideration when selecting or engineering for desirable endophyte traits.     

The Glycine-Glomus-Bradyrhizobium symbiosis. IX. Nutritional, morphological and physiological responses of nodulated soybean to geographic isolates of the mycorrhizal fungus Glomus mosseae.

The objective of the work was to determine differences in plant response to geographic isolates of a vesicular-arbuscular mycorrhizal (VAM) fungus, and to demonstrate the need for such determinations in the selection of desirable host-endophyte combinations for practical applications. Soybean ( Glycine max (L.) Merr.) plants were inoculated with Bradyrhizobium japonicum and isolates of the VAM-fungal morphospecies Glomus mosseae (Nicol. & Gerd.) Gerd. and Trappe, collected from an arid (AR), semiarid (SA) or mesic (ME) area. Inoculum potentials of the VAM-fungal isolates were determined and the inocula equalized, achieving the same level of root colonization (41%, P >0.05) at harvest (50 days). Plants of the three VAM treatments (AR, SA and ME) were evaluated against von VAM controls. Significant differences in plant response to colonization were found in dry mass, leaf K, N and P concentrations, and in root/shoot, nodule/root, root length/leaf area and root length/root mass ratios. The differences were most pronounced and consistent between the AR and all other treatments. Photosynthesis and nodule activity were higher ( P <0.05) in all VAM treatments, but only the AR plants had higher ( P <0.05) photosynthetic water-use efficiency than the controls. Nodule activity, evaluated by H₂ evolution and C₂H₂ reduction, differed significantly between treatments. The results are discussed in terms of nutritional and non-nutritional effects of VAM colonization on the development and physiology of the tripartite soybean association in the light of intraspecific variability within the fungal endophyte.     

The Glycine-Glomus-Rhizobium Symbiosis : VII. Photosynthetic Nutrient-Use Efficiency in Nodulated, Mycorrhizal Soybeans

Four consecutive trifoliate leaves of 56-day-old symbiotic or nonsymbiotic soybean plants were evaluated individually for CO₂ exchange rates (CER), leaf area and dry weight, and leaf N, P, and starch concentrations. Plants had been inoculated with the vesicular-arbuscular mycorrhizal (VAM) fungus Glomus mosseae and Rhizobium japonicum, with either of the endophytes alone, or with neither at time of planting. Plants lacking one or both endophytes received N and/or P fertilizers to produce plants of equal total leaf dry weight in all four treatments. Photosynthetic P-use efficiency (CER per unit leaf P) was higher in the leaves of VAM plants than in P-fertilized plants regardless of the N source (N₂ fixation or combined N). Photosynthetic N-use efficiency was also higher in VAM than in non-VAM plants, but it was affected by the N source, with higher CER in the nodulated plants. The greatest differences in CER, starch accumulation and leaf area were found between the nonsymbiotic plants and those with both endophytes. Statistical evaluations of leaf parameters for treatment or nutrient concentration (N and P) effects between the tri-partite and the nonsymbiotic treatments showed significant changes in concentration of P, but not N, with decreasing leaf age. Both endophytes apparently enhance CO₂ fixation at N and/or P concentrations lower than those of the nonsymbiotic plants. The effects of the endophytes on CO₂ fixation were additive.     

Interactions between Nitrogen Fixation, Mycorrhizal Colonization, and Host-Plant Growth in the Phaseolus-Rhizobium-Glomus Symbiosis

Bean (Phaseolus vulgaris L. cv. Dwarf) roots were inoculated with Rhizobium phaseoli and colonized by the vesicular-arbuscular mycorrhizal (VAM) fungus Glomus fasciculatum Gerd. and Trappe or left uncolonized as controls. The symbiotic associations were grown in an inert substrate using 0, 25, 50, 100, or 200 milligrams hydroxyapatite (HAP) (Ca₁₀[PO₄]₆[OH]₂) per pot as a P amendment. Plant and nodule dry weights and nodule activity increased for both VAM and control plants with increasing P availability, but values for VAM plants were significantly lower in all parameters than for controls. Inhibition of growth and of N₂ fixation in VAM plants was greatest at the lowest and highest P regimes. It was smallest at 50 milligrams HAP, where available P at harvest (7 weeks after planting) was 5 micrograms P per gram substrate. At this level of P availability, the association apparently benefited from increased P uptake by the fungal endophyte. Percent P values for shoots, roots, and nodules did not differ significantly (p > 0.05) between VAM and control plants. The extent of colonization, fungal biomass, and the fungus/association dry weight ratio increased several fold as HAP was increased from 0 to 200 milligrams. It is concluded that intersymbiont competition for P and photosynthate was the primary cause for the inhibition of growth, nodulation, and nodule activity in VAM plants. Impaired N₂ fixation resulted in N stress which contributed to inhibition of host plant growth at all levels of P availability.     

Alteration of growth ofSolanum opacum andPlantago drummondii and inhibition of regrowth of hyphae of vesicular-arbuscular mycorrhizal fungi from dried root pieces by manganese

Addition of MnSO₄ or MnCl₂ to a fine sandy soil from South Australia had a negative effect on shoot growth and root elongation ofSolanum opacum in the absence of significant presence of vesicular-arbuscular mycorrhiza (VAM). VAM ameliorated the reduction of plant growth by Mn, even though mycorrhizal development was decreased. Mn inhibited infection of roots by a fine endophyte less than that by some coarse endophytes. High concentrations of available Mn inhibited growth of hyphae of VAM fungi from dried root pieces, a significant source of infection by mycorrhizal fungi in the soil used.     

Glycine-Glomus-Bradyrhizobium Symbiosis : X. Relationships between Leaf Gas Exchange and Plant and Soil Water Status in Nodulated, Mycorrhizal Soybean under Drought Stress

Soybean (Glycine max [L.] Merr.) plants were colonized by the vesicular-arbuscular mycorrhizal (VAM) fungus Glomus mosseae (Nicol. and Gerd.) Gerd. and Trappe (VAM plants) or fertilized with KH₂PO₄ (nonVAM plants) and grown for 50 days under controlled conditions. Plants were harvested over a 4-day period during which the soil was permitted to dry slowly. The harvest was terminated when leaf gas exchange was no longer measurable due to drought stress. Significantly different effects in shoot water content, but not in shoot water potential, were found in VAM and nonVAM plants in response to drought stress. Leaf conductances of the two treatments showed similar response patterns to changes in soil water and shoot water potential but were significantly different in magnitude and trend relative to shoot water content. The relationships between transpiration, CO₂ exchange and water-use efficiency (WUE) were the same in VAM and nonVAM plants in response to decreasing soil water and shoot water potential. As a function of shoot water content, however, WUE showed different response patterns in VAM and nonVAM plants.     

The Glycine-Glomus-Bradyrhizobiun symbiosis. XI. Nodule gas exchange and efficiency as a function of soil and root water status in mycorrhizal soybean

Soybean [ Glycine max (L.) Merr. cv. Hobbit] plants were inoculated with a HUP− strain of Bradyrhizobium japonicum (Nitragin 61A118) and either colonized by the vesicular-arbuscular mycorrhizal (VAM) fungus Glomus mosseae (Nicol & Gerd.) Gerd. and Trappe or fertilized with KH₂PO₄ (nonVAM). They were grown for 50 days in a growth chamber and harvested over a 4-day drought period during which available soil water decreased to 0. Nodule P concentrations and P-use efficiency declined linearly with soil and root water content during the harvest period in both VAM and nonVAM plants. Nitrogenase activity, estimated from H₂ evolution and C₂H₂ reduction data, was also a linear function of declining nodule P concentrations and CO₂-exchange rates and showed simular patterns in both treatments. Hydrogen evolution and the relative efficiency of N₂ fixation, on the other hand, reacted differently to increasing drought in VAM and nonVAM plants. Differences in the responses of nodule activity in VAM and nonVAM plants to drought are interpreted in terms of demand for nodule P and carbohydrates and of the effects of dehydration on O₂ diffusion through nodule tissue.     

Effects of drought on host and endophyte development in mycorrhizal soybeans in relation to water use and phosphate uptake

Bethlenfalvay, G. J., Brown, M. S., Ames, R. N. and Thomas, R. S. 1988. Effects of drought on host and endophyte development in mycorrhizal soybeans in relation to water use and phosphate uptake. - Physiol. Plant. 72: 565–571.
Soybean [ Glycine max (L.) Merr.] plants were grown in pot cultures and inoculated with the vesicular-arbuscular mycorrhizal (VAM) fungus Glomus mosseae (Nicol. & Gerd.) Gerd. and Trappe or provided with P fertilizer (non-VAM plants). After an initial growth period (21 days), plants were exposed to cycles of severe, moderate or no drought stress over a subsequent 28-day period by rewatering at soil water potentials of -1.0, -0.3 or -0.05 MPa. Dry weights of VAM plants were greater at severe stress and smaller at no stress than those of non-VAM plants. Phosphorus fertilization was applied to produce VAM and non-VAM plants of the same size at moderate stress. Root and leaf P concentrations were higher in non-VAM plants at all stress levels. All plants were stressed to permanent wilting prior to harvest. VAM plants had lower soil moisture content at harvest than non-VAM plants. Colonization of roots by G. mosseae did not vary with stress, but the biomass and length of the extraradical mycelium was greater in severely stressed than in non-stressed plants. Growth enhancement of VAM plants relative to P-fertilized non-VAM plants under severe stress was attributed to increased uptake of water as well as to more efficient P uptake. The ability of VAM plants to deplete soil water to a greater extent than non-VAM plants suggests lower permanent wilting potentials for the former.     

Plant nitrogen acquisition and interactions under elevated carbon dioxide: impact of endophytes and mycorrhizae

Both endophytic and mycorrhizal fungi interact with plants to form symbiosis in which the fungal partners rely on, and sometimes compete for, carbon (C) sources from their hosts. Changes in photosynthesis in host plants caused by atmospheric carbon dioxide (CO₂) enrichment may, therefore, influence those mutualistic interactions, potentially modifying plant nutrient acquisition and interactions with other coexisting plant species. However, few studies have so far examined the interactive controls of endophytes and mycorrhizae over plant responses to atmospheric CO₂ enrichment. Using Festuca arundinacea Schreb and Plantago lanceolata L. as model plants, we examined the effects of elevated CO₂ on mycorrhizae and endophyte (Neotyphodium coenophialum) and plant nitrogen (N) acquisition in two microcosm experiments, and determined whether and how mycorrhizae and endophytes mediate interactions between their host plant species. Endophyte‐free and endophyte‐infected F. arundinacea varieties, P. lanceolata L., and their combination with or without mycorrhizal inocula were grown under ambient (400 μmol mol⁻¹) and elevated CO₂ (ambient + 330 μmol mol⁻¹). A ¹⁵N isotope tracer was used to quantify the mycorrhiza‐mediated plant acquisition of N from soil. Elevated CO₂ stimulated the growth of P. lanceolata greater than F. arundinacea, increasing the shoot biomass ratio of P. lanceolata to F. arundinacea in all the mixtures. Elevated CO₂ also increased mycorrhizal root colonization of P. lanceolata, but had no impact on that of F. arundinacea. Mycorrhizae increased the shoot biomass ratio of P. lanceolata to F. arundinacea under elevated CO₂. In the absence of endophytes, both elevated CO₂ and mycorrhizae enhanced ¹⁵N and total N uptake of P. lanceolata but had either no or even negative effects on N acquisition of F. arundinacea, altering N distribution between these two species in the mixture. The presence of endophytes in F. arundinacea, however, reduced the CO₂ effect on N acquisition in P. lanceolata, although it did not affect growth responses of their host plants to elevated CO₂. These results suggest that mycorrhizal fungi and endophytes might interactively affect the responses of their host plants and their coexisting species to elevated CO₂.     

Parasitic and mutualistic associations between a mycorrhizal fungus and soybean: The effect of phosphorus on host plant-endophyte interactions

Soybean [ Glycine max (L.) Merr. cv. Kent] plants were colonized by the vesicular-arbuscular mycorrhizal (VAM) fungus Glomus fasciculatum (Thaxt. sensu Gerd.) Gerd. and Trappe in pot cultures using an inert medium and a nutrient solution. Phosphorus was provided initially as 0, 25,50, 100 or 200 mg hydroxyapatite [HAP, Ca₁₀(PO₄)₆(OH)₂] per pot. Under the low (0 mg HAP) and high (100 and 200 mg HAP) P regimes, VAM plants showed 20, 25 and 38% growth retardation, respectively, relative to non-colonized controls. At 50 mg HAP, VAM plant growth was significantly enhanced (14%). Dry weight and P content of both VAM and control plants increased with increased P availability throughout the HAP gradient. Intraradical VAM fungal biomass increased linearly with increasing P availability. Extraradical VAM fungal biomass was smaller than the intraradical component of the fungus at the lowest and highest levels of P addition in the growth medium. The ratio of extra- to intraradical mycelium, a suggested index of VAM fungal effectiveness, was greatest for the 50 mg HAP treatment, coinciding with growth enhancement of the host plant. This enhanced growth of the host at an intermediate P level was apparently a result of increased P uptake by the endophyte.     

Root Respiration and Growth in Plantago major as Affected by Vesicular-Arbuscular Mycorrhizal Infection

Effects of vesicular-arbuscular mycorrhizal (VAM) infection and P on root respiration and dry matter allocation were studied in Plantago major L. ssp. pleiosperma (Pilger). By applying P, the relative growth rate of non-VAM controls and plants colonized by Glomus fasciculatum (Thaxt. sensu Gerdemann) Gerdemann and Trappe was increased to a similar extent (55-67%). However, leaf area ratio was increased more and net assimilation rate per unit leaf area was increased less by VAM infection than by P addition. The lower net assimilation rate could be related to a 20 to 30% higher root respiration rate per unit leaf area of VAM plants. Root respiration per unit dry matter and specific net uptake rates of N and P were increased more by VAM infection than by P addition. Neither the contribution of the alternative respiratory path nor the relative growth rate could account for the differences in root respiration rate between VAM and non-VAM plants. It was estimated that increased fungal respiration (87%) and ion uptake rate (13%) contributed to the higher respiratory activity of VAM roots of P. major.     

甘肃马先蒿寄生对禾草内生真菌共生体光合特性的影响

[目的]甘肃马先蒿与感染内生真菌的禾草(紫花针茅和麦宾草)建立根寄生关系,有关内生真菌对根寄生危害禾草光合作用调控方面的研究较少。[方法]本研究以紫花针茅和麦宾草带菌(E+)、不带菌(E-)植株为研究对象,研究甘肃马先蒿寄生和未寄生处理对紫花针茅和麦宾草E+、E-植株不同生长阶段光合特性影响的动态变化。[结果]甘肃马先蒿寄生显著降低紫花针茅和麦宾草的净光合速率、蒸腾速率和气孔导度,而胞间二氧化碳浓度和水分利用率却显著增加,这与禾草是否感染内生真菌无关。甘肃马先蒿寄生后E+紫花针茅的净光合速率、气孔导度和蒸腾速率高于E-植株,而麦宾草E+植株的净光合速率、气孔导度和蒸腾速率却低于E-植株;同时,根寄生条件下E+紫花针茅的胞间二氧化碳浓度和水分利用率低于E-植株;而E-麦宾草植株的胞间二氧化碳浓度和水分利用率却低于E+植株。[结论]内生真菌侵染影响甘肃马先蒿根寄生危害禾草的光合作用;甘肃马先蒿和内生真菌同时成为禾草营养消耗库时,内生真菌与禾草的共生关系处于一种互惠共生和相互拮抗的动态变化。     

Fungal Endophyte (Epichlo? festucae) Alters the Nutrient Content of Festuca rubra Regardless of Water Availability

Festuca rubra plants maintain associations with the vertically transmitted fungal endophyte Epichloë festucae. A high prevalence of infected host plants in semiarid grasslands suggests that this association could be mutualistic. We investigated if the Epichloë-endophyte affects the growth and nutrient content of F. rubra plants subjected to drought. Endophyte-infected (E+) and non-infected (E−) plants of two half-sib lines (PEN and RAB) were subjected to three water availability treatments. Shoot and root biomass, nutrient content, proline, phenolic compounds and fungal alkaloids were measured after the treatments. The effect of the endophyte on shoot and root biomass and dead leaves depended on the plant line. In the PEN line, E+ plants had a greater S:R ratio than E-, but the opposite occurred in RAB. In both plant lines and all water treatments, endophyte-infected plants had greater concentrations of N, P and Zn in shoots and Ca, Mg and Zn in roots than E- plants. On average, E+ plants contained in their shoots more P (62%), Zn (58%) and N (19%) than E- plants. While the proline in shoots increased in response to water stress, the endophyte did not affect this response. A multivariate analysis showed that endophyte status and plant line impose stronger differences in the performance of the plants than the water stress treatments. Furthermore, differences between PEN and RAB lines seemed to be greater in E- than in E+ plants, suggesting that E+ plants of both lines are more similar than those of their non-infected version. This is probably due to the endophyte producing a similar effect in both plant lines, such as the increase in N, P and Zn in shoots. The remarkable effect of the endophyte in the nutrient balance of the plants could help to explain the high prevalence of infected plants in natural grasslands.     

Nutrient transfer between the root zones of soybean and maize plants connected by a common mycorrhizal mycelium

The objective of the study was to determine whether nutrient fluxes mediated by hyphae of vesicular-arbuscular mycorrhizal (VAM) fungi between the root zones of grass and legume plants differ with the legume's mode of N nutrition. The plants, nodulating or nonnodulating isolines of soybean [ Glycine max (L.) Merr.], were grown in association with a dwarf maize ( Zea mays L.) cultivar in containers which interposed a 6-cm-wide root-free soil bridge between legume and grass container compartments. The bridge was delimited by screens (44 μm) which permitted the passage of hyphae, but not of roots and minimized non VAM interactions between the plants. All plants were colonized by the VAM fungus Glomus mosseae (Nicol. & Gerd.) Gerd. and Trappe. The effects of N input to N-sufficient soybean plants through N₂-fixation or N-fertilization on associated maize-plant growth and nutrition were compared to those of an N-deficient (nonnodulating, unfertilized) soybean control. Maize, when associated with the N-fertilized soybean, increased 19% in biomass, 67% in N content and 77% in leaf N concentration relative to the maize plants of the N-deficient association. When maize was grown with nodulated soybean, maize N content increased by 22%, biomass did not change, but P content declined by 16%. Spore production by the VAM fungus was greatest in the soils of both plants of the N-fertilized treatment. The patterns of N and P distribution, as well as those of the other essential elements, indicated that association with the N-fertilized soybean plants was more advantageous to maize than was association with the N₂-fixing ones.     

Leaky ribosomal scanning in mammalian genomes: significance of histone H4 alternative translation in vivo

Like alternative splicing, leaky ribosomal scanning (LRS), which occurs at suboptimal translational initiation codons, increases the physiological flexibility of the genome by allowing alternative translation. Comprehensive analysis of 22208 human mRNAs indicates that, although the most important positions relative to the first nucleotide of the initiation codon, −3 and +4, are usually such that support initiation (A₋₃ = 42%, G₋₃ = 36% and G₊₄ = 47%), only 37.4% of the genes adhere to the purine (R)₋₃/G₊₄ rule at both positions simultaneously, suggesting that LRS may occur in some of the remaining (62.6%) genes. Moreover, 12.5% of the genes lack both R₋₃ and G₊₄, potentially leading to sLRS. Compared with 11 genes known to undergo LRS, 10 genes with experimental evidence for high fidelity A₊₁T₊₂G₊₃ initiation codons adhered much more strongly to the R₋₃/G₊₄ rule. Among the intron-less histone genes, only the H3 genes adhere to the R₋₃/G₊₄ rule, while the H1, H2A, H2B and H4 genes usually lack either R₋₃ or G₊₄. To address in vivo the significance of the previously described LRS of H4 mRNAs, which results in alternative translation of the osteogenic growth peptide, transgenic mice were engineered that ubiquitously and constitutively express a mutant H4 mRNA with an A₊₁→T₊₁ mutation. These transgenic mice, in particular the females, have a high bone mass phenotype, attributable to increased bone formation. These data suggest that many genes may fulfill cryptic functions by LRS.     

Increased Sporulation of Vesicular-Arbuscular Mycorrhizal Fungi by Manipulation of Nutrient Regimens

Adjustment of pot culture nutrient solutions increased root colonization and sporulation of vesicular-arbuscular mycorrhizal (VAM) fungi. Paspalum notatum Flugge and VAM fungi were grown in a sandy soil low in N and available P. Hoagland nutrient solution without P enhanced sporulation in soil and root colonization of Acaulospora longula, Scutellospora heterogama, Gigaspora margarita, and a wide range of other VAM fungi over levels produced by a tap water control or nutrient solutions containing P. However, Glomus intraradices produced significantly more spores in plant roots in the tap water control treatment. The effect of the nutrient solutions was not due solely to N nutrition, because the addition of NH₄NO₃ decreased both colonization and sporulation by G. margarita relative to levels produced by Hoagland solution without P.     

Hyphal contribution to water uptake in mycorrhizal plants as affected by the fungal species and water status

Vesicular-arbuscular mycorrhizae may increase resistance of plants to drought by a number of mechanisms, such as increased root hydraulic conductivity, stomatal regulation, hyphal water uptake and osmotic adjustment. However, a substantial contribution of vesicular-arbuscular mycorrhizal (VAM) hyphae to water uptake has not been demonstrated unequivocally. The objective of this investigation was to examine the contribution of hyphae from two VAM fungi to water uptake and transport by the host plant. Lettuce (Lactuca sativa L.) plants were grown in a container divided by a screen into two compartments. One was occupied by roots, the other only by VAM hyphae, which the screen permitted to pass. Roots were colonized by the VAM fungi Glomus deserticola or Glomus fasciculatum, or were left uninoculated but P-supplemented. Water was supplied to the hyphal compartment at a distance of 10 cm from the screen (root). CO₂ exchange rate, water-use efficiency, transpiration, stomatal conductance and photosynthetic phosphorus-use efficiency of VAM or P-amended control plants were evaluated at three levels of water application in the hyphal compartment. Results indicate that much of the water was taken up by the hyphae in VAM plants. VAM plants, which had access to the hyphal compartment, had higher water and nutrient contents. G. deserticola functioned efficiently under water limitation and mycelium from G. fasciculatum-colonized plants was very sensitive to water in the medium. This discrepancy in VAM behaviour reflects the various abilities of each fungus according to soil water levels. Different abilities of specific mycelia were also expressed in terms of nutritional and leaf gas-exchange parameters. G. fasciculatum caused a significant increase in net photosynthesis and rate of water use efficiency compared to G. deserticola and P-fertilized plants. In contrast, the G. deserticola treatment was the most efficient affecting N, P and K nutrition, leaf conductance and transpiration. Since no differences in the intra- and extra-radical hyphal extension of the two endophytes were found, the results demonstrate that mycorrhizal hyphae can take up water and that there are considerable variations in both the behaviour of these two VAM fungi and in the mechanisms involved in their effects on plant water relations.