Organic acids, such as malate, citrate and oxalate, have been proposed to be involved in many processes operating in the rhizosphere, including nutrient acquisition and metal detoxification, alleviation of anaerobic stress in roots, mineral weathering and pathogen attraction. A full assessment of their role in these processes, however, cannot be determined unless the exact mechanisms of plant organic acid release and the fate of these compounds in the soil are more fully understood. This review therefore includes information on organic acid levels in plants (concentrations, compartmentalisation, spatial aspects, synthesis), plant efflux (passive versus active transport, theoretical versus experimental considerations), soil reactions (soil solution concentrations, sorption) and microbial considerations (mineralization). In summary, the release of organic acids from roots can operate by multiple mechanisms in response to a number of well-defined environmental stresses (e.g., Al, P and Fe stress, anoxia): These responses, however, are highly stress- and plant-species specific. In addition, this review indicates that the sorption of organic acids to the mineral phase and mineralisation by the soil's microbial biomass are critical to determining the effectiveness of organic acids in most rhizosphere processes.

Organic acids in the rhizosphere: a critical review

Vesicular-arbuscular mycorrhizal fungi can affect the water balance of both amply watered and droughted host plants. This review summarizes these effects and possible causal mechanisms. Also discussed are host drought resistance and the influence of soil drying on the fungi.

Water relations, drought and vesicular-arbuscular mycorrhizal symbiosis

Contents

 Summary

I Introduction
II Variability of N : P ratios in response to nutrient  supply
III Critical N : P ratios as indicators of nutrient  limitation
IV Interspecific variation in N : P ratios
V Vegetation properties in relation to N : P ratios
VI Implications of N : P ratios for human impacts  on ecosystems
VII Conclusions

 Acknowledgements


 References



Summary
Nitrogen (N) and phosphorus (P) availability limit plant growth in most terrestrial ecosystems. This review examines how variation in the relative availability of N and P, as reflected by N : P ratios of plant biomass, influences vegetation composition and functioning. Plastic responses of plants to N and P supply cause up to 50-fold variation in biomass N : P ratios, associated with differences in root allocation, nutrient uptake, biomass turnover and reproductive output. Optimal N : P ratios – those of plants whose growth is equally limited by N and P – depend on species, growth rate, plant age and plant parts. At vegetation level, N : P ratios 20 often (not always) correspond to N- and P-limited biomass production, as shown by short-term fertilization experiments; however long-term effects of fertilization or effects on individual species can be different. N : P ratios are on average higher in graminoids than in forbs, and in stress-tolerant species compared with ruderals; they correlate negatively with the maximal relative growth rates of species and with their N-indicator values. At vegetation level, N : P ratios often correlate negatively with biomass production; high N : P ratios promote graminoids and stress tolerators relative to other species, whereas relationships with species richness are not consistent. N : P ratios are influenced by global change, increased atmospheric N deposition, and conservation managment.

https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/j.1469-8137.2004.01192.x

N : P ratios in terrestrial plants: variation and functional significance

A great diversity of plants and fungi engage in mycorrhizal associations. In natural habitats, and in an ecologically meaningful time span, these associations have evolved to improve the fitness of both plant and fungal symbionts. In systems managed by humans, mycorrhizal associations often improve plant productivity, but this is not always the case. Mycorrhizal fungi might be considered to be parasitic on plants when net cost of the symbiosis exceeds net benefits. Parasitism can be developmentally induced, environmentally induced, or possibly genotypically induced. Morphological, phenological, and physiological characteristics of the symbionts influence the functioning of mycorrhizas at an individual scale. Biotic and abiotic factors at the rhizosphere, community, and ecosystem scales further mediate mycorrhizal functioning. Despite the complexity of mycorrhizal associations, it might be possible to construct predictive models of mycorrhizal functioning. These models will need to incorporate variables and parameters that account for differences in plant responses to, and control of, mycorrhizal fungi, and differences in fungal effects on, and responses to, the plant. Developing and testing quantitative models of mycorrhizal functioning in the real world requires creative experimental manipulations and measurements. This work will be facilitated by recent advances in molecular and biochemical techniques. A greater understanding of how mycorrhizas function in complex natural systems is a prerequisite to managing them in agriculture, forestry, and restoration.

http://max2.ese.u-psud.fr/epc/conservation/PDFs/HIPE/Johnson1997.pdf

Functioning of mycorrhizal associations along the mutualism–parasitism continuum*

Publisher Summary This chapter discusses a search for physiological causes and ecological consequence in reference with the variation in growth rate between higher plants. When grown under optimum conditions, plant species from fertile, productive habitats tend to have inherently higher relative growth rates (RGR) than species from less favorable environments. Under these conditions, fast-growing species produce relatively more leaf area and less root mass, which greatly contributes to their larger carbon gain per unit plant weight. Fast-growing species also have higher respiration rates per unit organ weight, due to demands of a higher RGR and higher rate of nutrient uptake. Fast-growing species have a greater capacity to acquire nutrients, which is likely to be a consequence, rather than the cause, of their higher RGR. This chapter analyses variations in morphological, physiological, chemical, and allocation characteristics underlying variation in RGR, to arrive at an appraisal of its ecological significance. The lower specific leaf area (SLA) of slow-growing species is because of the relatively high concentration of cell-wall material and quantitative secondary compounds, which may protect against detrimental abiotic and biotic factors. This chapter concludes that it is likely that there are trade-offs between growth potential and performance under adverse conditions, however, the current ecophysiological information explaining variation in RGR is too limited to support this contention quantitatively.

/pdf/inherent-variation-in-growth-rate-between-higher-plants-a-26yywnhi7b.pdf

Inherent Variation in Growth Rate Between Higher Plants: A Search for Physiological Causes and Ecological Consequences

SUMMARY
The mechanism responsible for inhibition of the establishment of mycorrhizal associations in Sorghum vulgare Pers. (herbaceous monocot) and Citrus aurantium L. (woody dicot) under high levels of soil phosphorus (P) was studied. Plants were grown on low fertility loamy sand (4.5 ppm P), receiving superphosphate [Ca (H2PO4)2H2O] at 0, 6, 28, 56, 228 and 556 ppm P along with all the other necessary nutrients. The percentage P content of root tissue was correlated with the amount of P added to the soil. Root exudation, measured in terms of the net leakage of soluble amino acids and reducing sugars from the roots within a 17–h period, was significantly higher under low P levels (0, 6 and 28 ppm P) than under high P levels (56, 228 and 556 ppm P). The amount of exudation was correlated with a P-induced decrease in phospholipid levels and associated changes in permeability properties of root membranes, rather than with changes in the root content of sugars and amino nitrogen. The hypothesis is proposed that phosphorus inhibition of mycorrhizal symbiosis is associated with a membrane-mediated decrease in root exudation.

https://nph.onlinelibrary.wiley.com/doi/pdf/10.1111/j.1469-8137.1978.tb01627.x

Root exudation in relation to supply of phosphorus and its possible relevance to mycorrhizal formation

SUMMARY
Using a ‘split root’ technique, it was found that phosphorus fertilization of half of the root system of sudangrass could significantly reduce the number of chlamydospores of the mycorrhizal fungus Glomus fasciculatus in the unfertilized half of the root system. In a second experiment, vials of soil containing different concentrations of phosphorus were inserted into pots of sudangrass which were fertilized with different amounts of phosphorus and inoculated with G. fasciculatus. The numbers of chlamydospores, vesicles and arbuscles and the amount of hyphae produced by G. fasciculatus on roots within the vials were not influenced by the soil P in the vials but were inversely correlated with the concentration of P in roots outside the vial. All evidence indicates that it is the concentration of P within the plant and not the soil P which leads to a reduction in colonization, infection, and spore production of G. fasciculatus.

https://nph.onlinelibrary.wiley.com/doi/pdf/10.1111/j.1469-8137.1978.tb01589.x

Phosphorus concentrations in plants responsible for inhibition of mycorrhizal infection

SUMMARY
Six citrus cultivars were grown with and without the mycorrhizal fungus Glomus fasciculatus under three fertilizer regimes all without phosphorus. The average mycorrhizal dependency (Gerdemann 1975) of Rough lemon and Brazilian sour orange were greater than the mycorrhizal dependecies of Alemow, Troyer citrange, Bessie sweet orange and Trifoliate orange. The mycorrhizal dependency of each cultivar, except Bessie sweet and Trifoliate orange, was substantially altered by at least one of the fertilizer regimes, and therefore the order of mycorrhizal dependency was different at all three fertilizer regimes. On the average, citrus rootstocks exhibited the greatest mycorrhizal dependency with the least fertilization. The average percentage phosphorus in non-mycorrhizal leaf tissues was inversely correlated with the mycorrhizal dependency of citrus cultivars at the medium fertilizer regime. An inverse correlation was observed between the dry weights of non-mycorrhizal roots of the citrus cultivars and the mycorrhizal dependency of the citrus cultivars.

https://nph.onlinelibrary.wiley.com/doi/pdf/10.1111/j.1469-8137.1978.tb01628.x

Mycorrhizal dependency of several citrus cultivars under three nutrient regimes

SUMMARY
The hypothesis was tested that the amount of external hyphae of a vesicular-arbuscular mycorrhizal (VAM) fungus extending from roots out into soil is not always proportional to the extent of colonization of the root cortex. Growth enhancement and amount of external hyphae were compared for eight isolates of five Glomus spp. that differed in their geographic origin and capacity to enhance growth of Troyer citrange, but were similar in their capacity to extensively colonize Troyer citrange roots. In general, isolates from California increased growth in a P-deficient (9.8 mg kg−1) California soil more than did non-native isolates from Florida soils. The difference between the capacity of California and Florida isolates to enhance growth was not a function of the degree to which they colonized the roots since all had colonized over 95% of the root length by the time of harvest. Differences in growth enhancement did appear, however, to be a function of the amount of external hyphae that had developed as estimated by the weight of soil they had bound into aggregates. This study suggests that isolates of VA mycorrhizal fungi may differ in their capacity to develop an external hyphal system independent of their capacity to colonize the root cortex, and that we cannot assume that high levels of colonization will necessarily mean the fungus has also developed the mycelium in the soil necessary to transport nutrients responsible for plant growth enhancement.

https://nph.onlinelibrary.wiley.com/doi/pdf/10.1111/j.1469-8137.1982.tb03304.x

Development of external hyphae by different isolates of mycorrhizal glomus spp. in relation to root colonization and growth of troyer citrange

SUMMARY

Larger populations of bacteria and actinomycetes were recovered from the rhizospheres of tomato plants inoculated with the mycorrhizal fungus Glomus fasciculatus and Azotobacter chroococcum, either individually or together, than from those of non-inoculated plants. Plants inoculated with both Glomus fasciculatus and Azotobacter chroococcum had greater numbers of bacteria and actinomycetes in the rhizosphere than plants inoculated with either Glomus fasciculatus or Azotobacter chroococcum alone. The fungal population of the rhizosphere was unaffected by inoculation with Glomus fasciculatus but reduced by inoculation with Azotobacter chroococcum. Inoculation of tomato with Glomus fasciculatus increased the Azotobacter chroococcum population in the rhizosphere which was maintained at a high level for a longer time. Inoculation of tomato roots with A. chroococcum enhanced infection and spore production by Glomus fasciculatus. The dry weights of tomato plants inoculated with both G. fasciculatus and Azotobacter chroococcum were significantly (62%) greater than non-inoculated plants. These results suggest a synergistic or additive interaction between Glomus fasciculatus and Azotobacter chroococcum.

https://nph.onlinelibrary.wiley.com/doi/pdf/10.1111/j.1469-8137.1978.tb01588.x

John A. Menge

Papers

Root exudation in relation to supply of phosphorus and its possible relevance to mycorrhizal formation

Phosphorus concentrations in plants responsible for inhibition of mycorrhizal infection

Mycorrhizal dependency of several citrus cultivars under three nutrient regimes

Development of external hyphae by different isolates of mycorrhizal glomus spp. in relation to root colonization and growth of troyer citrange

Interaction between a va mycorrhiza and azotobacter and their effects on rhizosphere microflora and plant growth