Arbuscular mycorrhizal fungi benefit from 7 years of free air CO2 enrichment in well-fertilized grass and legume monocultures

Rising atmospheric carbon dioxide partial pressure (pCO₂) and nitrogen (N) deposition are important components of global environmental change. In the Swiss free air carbon dioxide enrichment (FACE) experiment, the effect of altered atmospheric pCO₂ (35 vs. 60 Pa) and the influence of two different N‐fertilization regimes (14 vs. 56 g N m^?2 a^?1) on root colonization by arbuscular mycorrhizal fungi (AMF) and other fungi (non‐AMF) of Lolium perenne and Trifolium repens were studied. Plants were grown in permanent monoculture plots, and fumigated during the growth period for 7 years. At elevated pCO₂ AMF and non‐AMF root colonization was generally increased in both plant species, with significant effects on colonization intensity and on hyphal and non‐AMF colonization. The CO₂ effect on arbuscules was marginally significant (P=0.076). Moreover, the number of small AMF spores (≤100 μm) in the soils of monocultures (at low‐N fertilization) of both plant species was significantly increased, whereas that of large spores (>100 μm) was increased only in L. perenne plots. N fertilization resulted in a significant decrease of root colonization in L. perenne, including the AMF parameters, hyphae, arbuscules, vesicles and intensity, but not in T. repens. This phenomenon was probably caused by different C‐sink limitations of grass and legume. Lacking effects of CO₂ fumigation on intraradical AMF structures (under high‐N fertilization) and no response to N fertilization of arbuscules, vesicles and colonization intensity suggest that the function of AMF in T. repens was non‐nutritional. In L. perenne, however, AM symbiosis may have amended N nutrition, because all root colonization parameters were significantly increased under low‐N fertilization, whereas under high‐N fertilization only vesicle colonization was increased. Commonly observed P‐nutritional benefits from AMF appeared to be absent under the phosphorus‐rich soil conditions of our field experiment. We hypothesize that in well‐fertilized agricultural ecosystems, grasses benefit from improved N nutrition and legumes benefit from increased protection against pathogens and/or herbivores. This is different from what is expected in nutritionally limited plant communities.     

Mycorrhizas improve nitrogen nutrition of Trifolium repens after 8 yr of selection under elevated atmospheric CO2 partial pressure

Altered environmental conditions may change populations of arbuscular mycorrhizal fungi and thereby affect mycorrhizal functioning. We investigated whether 8 yr of free-air CO2 enrichment has selected fungi that differently influence the nutrition and growth of host plants. In a controlled pot experiment, two sets of seven randomly picked single spore isolates, originating from field plots of elevated (60 Pa) or ambient CO2 partial pressure (pCO2), were inoculated on nodulated Trifolium repens (white clover) plants. Fungal isolates belonged to the Glomus claroideum or Glomus intraradices species complex, and host plants were clonal micropropagates derived from nine genets. Total nitrogen (N) concentration was increased in leaves of plants inoculated with fungal isolates from elevated-pCO2 plots. These isolates took up nearly twice as much N from the soil as isolates from ambient-pCO2 plots and showed much greater stimulation of biological N2 fixation. The morpho-species identity of isolates had a more pronounced effect on N2 fixation and on root length colonized than isolate identity. We conclude that rising atmospheric pCO2 may select for fungal strains that will help their host plants to meet increased N demands.     

Growth in Elevated CO2 Can Both Increase and Decrease Photochemistry and Photoinhibition of Photosynthesis in a Predictable Manner. Dactylis glomerata Grown in Two Levels of Nitrogen Nutrition

Biochemically based models of C(3) photosynthesis can be used to predict that when photosynthesis is limited by the amount of Rubisco, increasing atmospheric CO(2) partial pressure (pCO(2)) will increase light-saturated linear electron flow through photosystem II (J(t)). This is because the stimulation of electron flow to the photosynthetic carbon reduction cycle (J(c)) will be greater than the competitive suppression of electron flow to the photorespiratory carbon oxidation cycle (J(o)). Where elevated pCO(2) increases J(t), then the ratio of absorbed energy dissipated photochemically to that dissipated non-photochemically will rise. These predictions were tested on Dactylis glomerata grown in fully controlled environments, at either ambient (35 Pa) or elevated (65 Pa) pCO(2), and at two levels of nitrogen nutrition. As was predicted, for D. glomerata grown in high nitrogen, J(t) was significantly higher in plants grown and measured at elevated pCO(2) than for plants grown and measured at ambient pCO(2). This was due to a significant increase in J(c) exceeding any suppression of J(o). This increase in photochemistry at elevated pCO(2) protected against photoinhibition at high light. For plants grown at low nitrogen, J(t) was significantly lower in plants grown and measured at elevated pCO(2) than for plants grown and measured at ambient pCO(2). Elevated pCO(2) again suppressed J(o); however growth in elevated pCO(2) resulted in an acclimatory decrease in leaf Rubisco content that removed any stimulation of J(c). Consistent with decreased photochemistry, for leaves grown at low nitrogen, the recovery from a 3-h photoinhibitory treatment was slower at elevated pCO(2).     

Arbuscular mycorrhizal fungi are necessary for the induced response to herbivores by Cucumis sativus

Aims Although ecological interactions are often conceptualized and studied in a pairwise framework, ecologists recognize that the outcomes of these interactions are influenced by other members of the community. Interactions (i) between plants and insect herbivores and (ii) between plants and mycorrhizal fungi are ubiquitous in terrestrial ecosystems and may be linked via common host plants. Previous studies suggest that colonization by arbuscular mycorrhizal fungi (AMF) can modify plants' induced responses to herbivore attack, but these indirect effects of fungal symbionts are poorly understood. I investigated the role of AMF in induced plant response to a generalist herbivore.Methods I manipulated AMF status and herbivory in Cucumis sativus L. (cucumber, Cucurbitaceae) in a greenhouse to investigate induced responses in the presence and absence of the mycorrhizal fungus Glomus intraradices (Glomeraceae). Spodoptera exigua Hübner (Noctuidae) were used to manipulate prior damage and later as assay caterpillars. I also measured G. intraradices and herbivory effects on plant N and effects on plant growth.Important findings AMF status affected the induced response of C. sativus, underscoring the importance of incorporating the roles of plant symbionts into plant defense theory. Assay caterpillars ate significantly more leaf tissue only on mycorrhizal plants that had experienced prior damage. Despite more consumption, biomass change in these caterpillars did not differ from those feeding on plants with other treatment combinations. Leaf N content was reduced by G. intraradices but unaffected by herbivory treatments, suggesting that the observed differences in assay caterpillar feeding were due to changes in defensive chemistry that depended on AMF.     

Is resource allocation and grain yield of rice altered by inoculation with arbuscular mycorrhizal fungi?

Aims Our study quantified the combined effects of fertilization and inoculation with arbuscular mycorrhizal fungi (AMF) on grain yield and allocation of biomass and nutrients in field-grown rice (Oryza sativa L.).Methods A two-factor experiment was conducted at a field site in northeast of China (in Shuangcheng, Heilongjiang Province, Songhua River basin): six nitrogen–phosphorus–potassium fertilizer levels were provided (0, 20, 40, 60, 80 and 100% of the local norm of fertilizer supply), with or without inoculation with Glomus mosseae. At maturity, we quantified the percentage of root length colonization by AMF, grain yield, shoot:root ratios, shoot N and P contents and nutrients allocated to panicles, leaves and stems.Important findings As expected, inoculation resulted in greatly increased AMF colonization, which in turn led to higher shoot:root ratios and greater shoot N contents. Shoot:root ratios of inoculated rice increased with increasing fertilization while there was a significant interaction between fertilization and inoculation on shoot:root ratio. Additionally, AMF inoculation increased panicle:shoot ratios, panicle N:shoot N ratios and panicle P:shoot P ratios, especially in plants grown at low fertilizer levels. Importantly, inoculated rice exhibited higher grain yield, with the maximum improvement (near 62%) at the lower fertilizer end. Our results showed that (i) AMF-inoculated plants conform to the functional equilibrium theory, albeit to a reduced extent compared to non-inoculated plants and (ii) AMF inoculation resulted in greater allocation of shoot biomass to panicles and increased grain yield by stimulating N and P redistribution to panicles.     

Stimulation of Symbiotic N2 Fixation in Trifolium repens L. under Elevated Atmospheric pCO2 in a Grassland Ecosystem

Symbiotic N2 fixation is one of the main processes that introduces N into terrestrial ecosystems. As such, it may be crucial for the sequestration of the extra C available in a world of continuously increasing atmospheric CO2 partial pressure (pCO2). The effect of elevated pCO2 (60 Pa) on symbiotic N2 fixation (15N-isotope dilution method) was investigated using Free-Air-CO2-Enrichment technology over a period of 3 years. Trifolium repens was cultivated either alone or together with Lolium perenne (a nonfixing reference crop) in mixed swards. Two different N fertilization levels and defoliation frequencies were applied. The total N yield increased consistently and the percentage of plant N derived from symbiotic N2 fixation increased significantly in T. repens under elevated pCO2. All additionally assimilated N was derived from symbiotic N2 fixation, not from the soil. In the mixtures exposed to elevated pCO2, an increased amount of symbiotically fixed N (+7.8, 8.2, and 6.2 g m-2 a-1 in 1993, 1994, and 1995, respectively) was introduced into the system. Increased N2 fixation is a competitive advantage for T. repens in mixed swards with pasture grasses and may be a crucial factor in maintaining the C:N ratio in the ecosystem as a whole.     

Elevated pCO(2 )favours nitrate reduction in the roots of wild-type tobacco (Nicotiana tabacum cv. Gat.) and significantly alters N-metabolism in transformants lacking functional nitrate reductase in the roots

The impact of elevated pCO(2 )on N-metabolism of hydroponically grown wild-type and transformed tobacco plants lacking root nitrate reduction was studied in order to elucidate the effects on (i) nitrate uptake, (ii) long-distance transport of N, (iii) nitrate reduction with emphasis on root-NR, and (iv) the allocation of N between the root and shoot. The findings were related to alterations of growth rates. At elevated pCO(2 )the wild type exhibited higher growth rates, which were accompanied by an increase of NO(3)(-)-uptake per plant, due to a higher root:shoot ratio. Furthermore, elevated pCO(2 )enhanced nitrate reduction in the roots of the wild type, resulting in enhanced xylem-loading of organic N (amino-N) to supply the shoot with sufficient nitrogen, and decreased phloem-transport of organic N in a basipetal direction. Transformed tobacco plants lacking root nitrate reduction were smaller than the wild type and exhibited lower growth rates. Nitrate uptake per plant was decreased in transformed plants as a consequence of an impeded root growth and, thus, a significantly decreased root:shoot ratio. Surprisingly, transformed plants showed an altered allocation of amino-N between the root and the shoot, with an increase of amino-N in the root and a substantial decrease of amino-N in the shoot. In transformed plants, xylem-loading of nitrate was increased and the roots were supplied with organic N via phloem transport. Elevated pCO(2 )increased shoot-NR, but only slightly affected the growth rates of transformed plants, whereas carbohydrates accumulated at elevated pCO(2 )as indicated by a significant increase of the C/N ratio in the leaves of transformed plants. Unexpectedly, the C/N balance and the functional equilibrium between root and shoot growth was disturbed dramatically by the loss of nitrate reduction in the root.     

Differential Effects of Pseudomonas mendocina and Glomus intraradices on Lettuce Plants Physiological Response and Aquaporin PIP2 Gene Expression Under Elevated Atmospheric CO2 and Drought

Arbuscular mycorrhizal (AM) symbiosis and plant-growth-promoting rhizobacterium (PGPR) can alleviate the effects of water stress in plants, but it is unknown whether these benefits can be maintained at elevated CO₂. Therefore, we carried out a study where seedlings of Lactuca sativa were inoculated with the AM fungus (AMF) Glomus intraradices N.C. Schenk & G.S. Sm. or the PGPR Pseudomonas mendocina Palleroni and subjected to two levels of watering and two levels of atmospheric CO₂ to ascertain their effects on plant physiological parameters and gene expression of one PIP aquaporin in roots. The inoculation with PGPR produced the greatest growth in lettuce plants under all assayed treatments as well as the highest foliar potassium concentration and leaf relative water content under elevated [CO₂] and drought. However, under such conditions, the PIP2 gene expression remained almost unchanged. G. intraradices increased significantly the AMF colonization, foliar phosphorus concentration and leaf relative water content in plants grown under drought and elevated [CO₂]. Under drought and elevated [CO₂], the plants inoculated with G. intraradices showed enhanced expression of the PIP2 gene as compared to P. mendocina or control plants. Our results suggest that both microbial inoculation treatments could help to alleviate drought at elevated [CO₂]. However, the PIP2 gene expression was increased only by the AMF but not by the PGPR under these conditions.     

Phosphorus supply and arbuscular mycorrhizas increase growth and net gas exchange responses of two Citrus spp. grown at elevated [CO2]

We hypothesized that greater photosynthate supply at elevated [CO₂] could compensate for increased below-ground C demands of arbuscular mycorrhizas. Therefore, we investigated plant growth, mineral nutrition, starch, and net gas exchange responses of two Citrus spp. to phosphorus (P) nutrition and mycorrhizas at elevated atmospheric [CO₂]. Half of the seedlings of sour orange (C. aurantium L.) and ‘Ridge Pineapple’ sweet orange (C. sinensis L. Osbeck) were inoculated with the arbuscular mycorrhizal (AM) fungus, Glomus intraradices Schenck and Smith and half were non-mycorrhizal (NM). Plants were grown at ambient or 2X ambient [CO₂] in unshaded greenhouses for 11 weeks and fertilized daily with nutrient solution either without added P or with 2 mM P in a low-P soil. High P supply reduced AM colonization whereas elevated [CO₂] counteracted the depressive effect of P on intraradical colonization and vesicle development. Seedlings grown at either elevated [CO₂], high P or with G. intraradices had greater growth, net assimilation of CO₂ (A _CO2) in leaves, leaf water-use efficiency, leaf dry wt/area, leaf starch and carbon/nitrogen (C/N) ratio. Root/whole plant dry wt ratio was decreased by elevated [CO₂], P, and AM colonization. Mycorrhizal seedlings had higher leaf-P status but lower leaf N and K concentrations than nonmycorrhizal seedlings which was due to growth dilution effects. Starch in fibrous roots was increased by elevated [CO₂] but reduced by G. intraradices, especially at low-P supply. In fibrous roots, elevated [CO₂] had no effect on C/N, but AM colonization decreased C/N in both Citrus spp. grown at low-P supply. Overall, there were no species differences in growth or A _CO2. Mycorrhizas did not increase plant growth at ambient [CO₂]. At elevated [CO₂], however, mycorrhizas stimulated growth at both P levels in sour orange, the more mycorrhiza-dependent species, but only at low-P in sweet orange, the less dependent species. At low-P and elevated [CO₂], colonization by the AM fungus increased A _CO2 in both species but more so in sour orange than in sweet orange. Leaf P and root N concentrations were increased more and root starch level was decreased less by AM in sour orange than in sweet orange. Thus, the additional [CO₂] availability to mycorrhizal plants increased CO₂ assimilation, growth and nutrient uptake over that of NM plants especially in sour orange under P limitation. This revised version was published online in June 2006 with corrections to the Cover Date.     

Cadmium accumulation in sunflower plants influenced by arbuscular mycorrhiza

In order to investigate the cadmium (Cd) accumulation patterns and possible alleviation of Cd stress by mycorrhization, sunflower plants (Helianthus annuus L.) were grown in the presence or absence of Cd (20 micromol L(-1)) and inoculated or not inoculated with the arbuscular mycorrhizal fungus (AMF) Glomus intraradices. No visual symptoms of Cd phytotoxicity were observed; nevertheless, in non-mycorrhizal plants the presence of Cd decreased plant growth. The addition of Cd had no significant effect on either mycorrhizal colonization or the amount of extra-radical mycelia that was produced by the AMF. Cd accumulated mainly in roots; only 22% of the total Cd absorbed was translocated to the shoots, where it accumulated to an average of 228 mg Cd kg(-1). Although the shoot-to-root ratio of Cd was similar in both the AMF inoculated and non-inoculated plants, the total absorbed Cd was 23% higher in mycorrhizal plants. Cd concentration in AMF extra-radical mycelium was 728 microg g(-1) dry weight. Despite the greater absorption of Cd, mycorrhizal plants showed higher photosynthetic pigment concentrations and shoot P contents. Cd also influenced mineral nutrition, leading to decreased Ca and Cu shoot concentrations; N, Fe and Cu shoot contents; and increased S and K shoot concentrations. Cd induced guaiacol peroxidase activity in roots in both mycorrhizal and non-mycorrhizal plants, but this increase was much more accentuated in non-mycorrhizal roots. In conclusion, sunflower plants associated with G. intraradices were less sensitive to Cd stress than non-mycorrhizal plants. Mycorrhizal sunflowers showed enhanced Cd accumulation and some tolerance to excessive Cd concentrations in plant tissues.     

Yield response of Lolium perenne swards to free air CO2 enrichment increased over six years in a high N input system on fertile soil

After a step increase in the atmospheric partial pressure of CO₂ (pCO₂), the availability of mineral N may be insufficient to meet the plant's increased demand for N. Over time, however, the ecosystem may adapt to the new conditions, and a new equilibrium may be established in the fluxes of C and N. This would result in a higher dry mass (DM) yield response of the plants to elevated pCO₂. The effect of elevated atmospheric pCO₂ (60 Pa pCO₂) was studied in Lolium perenne L. swards with two N fertilization treatments (14 and 56 g m^?2 y^?1) in a six‐year FACE (Free Air Carbon dioxide Enrichment) experiment. In the high N treatment, the input of N with fertilizer considerably exceeded the export of N with the harvested plant material in both CO₂ treatments leading to an apparent net input of N into the ecosystem. Accordingly, the proportion of harvested N derived from ¹⁵N labelled fertilizer N, applied throughout the experiment (< 6 years), increased over the years. Under these high N conditions, the annual DM yield response of the Lolium perenne sward to elevated pCO₂ increased (from 7% in 1993 to 25% in 1998). In parallel, the response of N yield to elevated pCO₂ increased, and the initially negative effect of elevated pCO₂ on specific leaf area (SLA) disappeared. The high N input system seemed to overcome in part an initially limiting effect of N on the yield response to elevated pCO₂ within a few years. In contrast, there was no apparent net input of N into the ecosystem in the low N treatment, because N fertilization just compensated the export of N with the harvested plant material. Accordingly, the proportion of harvested N yield, derived from fertilizer N, which was applied throughout the experiment, remained low. At low N, the availability of mineral N strongly limited plant growth and yield production in both CO₂ treatments; the low yields of DM and N, the low concentration of N in the plant material, and the low SLA reflected this. Although the plants grew under the same environmental conditions and the same management treatment as plants in the high N treatment, the response of DM yields to elevated pCO₂ in the low N treatment remained weak throughout the experiment (5% in 1993 and 9% in 1998). The results are discussed in the context of the sizes of the different N pools in the soil, the allocation of N within the plant and the possible effects on temporal immobilization, and the availability of mineral N for yield production as affected by elevated pCO₂ and N fertilization.     

Nitrogen-fixing bacteria alleviates competition between arbuscular mycorrhizal fungi and Solidago canadensis for nutrients under nitrogen limitation

丛枝菌根真菌(AMF)能与大多数陆生植物的根系形成共生体, 有助于宿主植物吸收养分。但营养胁迫下, 根系微生物对AMF与宿主植物间关系的影响少见报道。该研究假设: 在营养极度匮乏(如氮胁迫)环境下, AMF与宿主植物可能产生营养竞争, 而固氮菌的介入能够缓解两者对营养的竞争关系。为了验证这一假设, 该文探究了加拿大一枝黄花(Solidago canadensis)生长受限的氮浓度, 并在氮受限条件下检验了AMF、加拿大一枝黄花及固氮菌三者间的关系。结果表明: 低氮处理明显抑制了加拿大一枝黄花的地上生物量和总生物量, 尤其以0.025 mmol·L^-1 N的氨态氮对加拿大一枝黄花的负影响更甚。在此氮浓度下, 单独添加AMF总体上都进一步抑制了加拿大一枝黄花的生长, 而固氮菌的添加在一定程度上提高了氮受限条件下AMF对宿主的根部侵染率及宿主植物生物量。这表明固氮菌能够缓和氮受限条件下AMF和加拿大一枝黄花间的营养竞争关系。研究结果加深了对外来植物在极度营养胁迫环境下与多种微生物互作的入侵机制的理解。     

固氮菌缓解氮限制环境中丛枝菌根真菌对加拿大一枝黄花的营养竞争

丛枝菌根真菌(AMF)能与大多数陆生植物的根系形成共生体, 有助于宿主植物吸收养分。但营养胁迫下, 根系微生物对AMF与宿主植物间关系的影响少见报道。该研究假设: 在营养极度匮乏(如氮胁迫)环境下, AMF与宿主植物可能产生营养竞争, 而固氮菌的介入能够缓解两者对营养的竞争关系。为了验证这一假设, 该文探究了加拿大一枝黄花(Solidago canadensis)生长受限的氮浓度, 并在氮受限条件下检验了AMF、加拿大一枝黄花及固氮菌三者间的关系。结果表明: 低氮处理明显抑制了加拿大一枝黄花的地上生物量和总生物量, 尤其以0.025 mmol·L^-1 N的氨态氮对加拿大一枝黄花的负影响更甚。在此氮浓度下, 单独添加AMF总体上都进一步抑制了加拿大一枝黄花的生长, 而固氮菌的添加在一定程度上提高了氮受限条件下AMF对宿主的根部侵染率及宿主植物生物量。这表明固氮菌能够缓和氮受限条件下AMF和加拿大一枝黄花间的营养竞争关系。研究结果加深了对外来植物在极度营养胁迫环境下与多种微生物互作的入侵机制的理解。     

Interaction between Earthworms and Arbuscular Mycorrhizal Fungi in Plants: A Review

Different kinds of soil animals and microorganisms inhabit the plant rhizosphere, which function closely to plant roots. Of them, arbuscular mycorrhizal fungi (AMF) and earthworms play a critical role in sustaining the soil-plant health. Earthworms and AMF belong to the soil community and are soil beneficial organisms at different trophic levels. Both of them improve soil fertility and structural development, collectively promoting plant growth and nutrient acquisition capacity. Earthworm activities redistribute mycorrhizal fungi spores and give diversified effects on root mycorrhizal fungal colonization. Dual inoculation with both earthworms and AMF strongly magnifies the response on plant growth through increased soil enzyme activities and changes in soil nutrient availability, collectively mitigating the negative effects of heavy metal pollution in plants and soils. This thus enhances phytoremediation and plant disease resistance. This review simply outlines the effects of earthworms and AMF on the soil-plant relationship. The effects of earthworms on root AMF colonization and activities are also analyzed. This paper also summarizes the interaction between earthworms and AMF on plants along with suggested future research.     

Influence of a Bacillus sp. on physiological activities of two arbuscular mycorrhizal fungi and on plant responses to PEG-induced drought stress

The effects of bacterial inoculation (Bacillus sp.) on the development and physiology of the symbiosis between lettuce and the arbuscular mycorrhizal (AM) fungi Glomus mosseae (Nicol. and Gerd.) Gerd. and Trappe and Glomus intraradices (Schenck and Smith) were investigated. Plant growth, mineral nutrition and gas-exchange values in response to bacterial inoculation after PEG-induced drought stress were also evaluated. In AM plants, inoculation with Bacillus sp. enhanced fungal development and metabolism, measured as succinate dehydrogenase (SDH) and alkaline phosphatase (ALP) activities, more than plant growth. Under non-stressed conditions, G. intraradices colonization increased all plant physiological values to a higher extent when in dual inoculation with the bacterium. Under stress conditions, the bacterium had an important stimulatory effect on G. intraradices development. Under such conditions, the effects of the bacterium on photosynthetic rate, water use efficiency (WUE) and stomatal conductance of lettuce plants differed with the fungus species. Plant-gas exchange was enhanced in G. intraradices- and reduced in G. mosseae-colonized plants when co-inoculated with Bacillus sp. Thus, the effects of each fungus on plant physiology were modulated by the bacterium. Stress was detrimental, particularly in G. intraradices-colonized plants without the bacterium, reducing intra and extraradical mycelium growth and vitality (SDH), as well as plant-gas exchange. Nevertheless, Bacillus sp. inoculation improved all these plant and fungal parameters to the same level as in non-stressed plants. The highest amount of alive and active AM mycelium for both fungi was obtained after co-inoculation with Bacillus sp. These results suggest that selected free-living bacteria and AM fungi should be co-inoculated to optimize the formation and functioning of the AM symbiosis in both normal and adverse environments.     

Effects of co-inoculation with arbuscular mycorrhizal fungi and rhizobia on soybean growth as related to root architecture and availability of N and P

Soybean plants can form tripartite symbiotic associations with rhizobia and arbuscular mycorrhizal (AM) fungi, but little is known about effects of co-inoculation with rhizobia and AM fungi on plant growth, or their relationships to root architecture as well as nitrogen (N) and phosphorus (P) availability. In the present study, two soybean genotypes contrasting in root architecture were grown in a field experiment to evaluate relationships among soybean root architecture, AMF colonization, and nodulation under natural conditions. Additionally, a soil pot experiment in greenhouse was conducted to investigate the effects of co-inoculation with rhizobia and AM fungi on soybean growth, and uptake of N and P. Our results indicated that there was a complementary relationship between root architecture and AMF colonization in the field. The deep root soybean genotype had greater AMF colonization at low P, but better nodulation with high P supply than the shallow root genotype. A synergistic relationship dependent on N and P status exists between rhizobia and AM fungi on soybean growth. Co-inoculation with rhizobia and AM fungi significantly increased soybean growth under low P and/or low N conditions as indicated by increased shoot dry weight, along with plant N and P content. There were no significant effects of inoculation under adequate N and P conditions. Furthermore, the effects of co-inoculation were related to root architecture. The deep root genotype, HN112, benefited more from co-inoculation than the shallow root genotype, HN89. Our results elucidate new insights into the relationship between rhizobia, AM fungi, and plant growth under limitation of multiple nutrients, and thereby provides a theoretical basis for application of co-inoculation in field-grown soybean.     

Nitrogen supply affects arbuscular mycorrhizal colonization of Artemisia vulgaris in a phosphate-polluted field site

Root colonization by arbuscular mycorrhizal fungi (AMF) was investigated in industrially polluted grassland characterized by exceptionally high phosphorus levels (up to 120 g kg(-1) soil). Along a pollution-induced nitrogen gradient, soil and tissue element concentrations of Artemisia vulgaris plants and their mycorrhizal status were determined. Additionally, we compared mycorrhization rates and above-ground biomass of A. vulgaris at N-fertilized and control plots in the N-poor area. Despite high soil and tissue P concentrations, plants from N-deficient plots, which were characterized by low tissue N concentrations and N : P ratios, were strongly colonized by AMF, whereas at a plot with comparable P levels, but higher soil and plant N concentrations and N : P ratios, mycorrhization rates were significantly lower. Correlation analyses revealed a negative relationship between percentage root colonization of A. vulgaris by AMF and both tissue N concentration and N : P ratio. Accordingly, in the fertilization experiment, control plants had higher mycorrhization rates than N-fertilized plants, whereas the species attained higher biomass at N-fertilized plots. The results suggest that N deficiency stimulates root colonization by AMF in this extraordinarily P-rich field site.