Mycorrhizal responses to nitrogen fertilization in boreal ecosystems: potential consequences for soil carbon storage

Mycorrhizal fungi can contribute to soil carbon sequestration by immobilizing carbon in living fungal tissues and by producing recalcitrant compounds that remain in the soil following fungal senescence. We hypothesized that nitrogen (N) fertilization would decrease these carbon stocks, because plants should reduce investment of carbon in mycorrhizal fungi when N availability is high. We measured the abundance of two major groups of mycorrhizal fungi, arbuscular mycorrhizal (AM) and ectomycorrhizal (ECM) fungi, in the top 10 cm of soil in control and N-fertilized plots within three Alaskan boreal ecosystems that represented different recovery stages following severe fire. Pools of mycorrhizal carbon included root-associated AM and ECM structures; soil-associated AM hyphae; and glomalin, a glycoprotein produced by AM fungi. Total mycorrhizal carbon pools decreased by approximately 50 g C m⁻² in the youngest site under N fertilization, and this reduction was driven mostly by glomalin. Total mycorrhizal carbon did not change significantly in the other sites. Root-associated AM structures were more abundant under N fertilization across all sites, and root-associated ECM structures increased marginally significantly. We found no significant N effects on AM hyphae. Carbon sequestered within living mycorrhizal structures (0.051–0.21 g m⁻²) was modest compared with that of glomalin (33–203 g m⁻²). We conclude that our hypothesis was only supported in relation to glomalin stocks within one of the three study sites. As N effects on glomalin were inconsistent among sites, an understanding of the mechanisms underlying this variation would improve our ability to predict ecosystem feedbacks to global change.     

Fungal root colonization responses in natural grasslands after long-term exposure to elevated atmospheric CO2

Arbuscular mycorrhizae, ubiquitous mutualistic symbioses between plant roots and fungi in the order Glomales, are believed to be important controllers of plant responses to global change, in particular to elevated atmospheric CO₂. In order to test if any effects on the symbiosis can persist after long-term treatment, we examined root colonization by arbuscular mycorrhizal (AM) and other fungi of several plant species from two grassland communities after continuous exposure to elevated atmospheric CO₂ for six growing seasons in the field. For plant species from both a sandstone and a serpentine annual grassland there was evidence for changes in fungal root colonization, with changes occurring as a function of plant host species. We documented decreases in percentage nonmycorrhizal fungal root colonization in elevated CO₂ for several plant species. Total AM root colonization (%) only increased significantly for one out of the five plant species in each grassland. However, when dividing AM fungal hyphae into two groups of hyphae (fine endophyte and coarse endophyte), we could document significant responses of AM fungi that were hidden when only total percentage colonization was measured. We also documented changes in elevated CO₂ in the percentage of root colonized by both AM hyphal types simultaneously. Our results demonstrate that changes in fungal root colonization can occur after long-term CO₂ enrichment, and that the level of resolution of the study of AM fungal responses may have to be increased to uncover significant changes to the CO₂ treatment. This study is also one of the first to document compositional changes in the AM fungi colonizing roots of plants grown in elevated CO₂. Although it is difficult to relate the structural data directly to functional changes, possible implications of the observed changes for plant communities are discussed.     

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.     

Nitrogen input mediates the effect of free-air CO2 enrichment on mycorrhizal fungal abundance

Plots containing Lolium perenne L., Trifolium repens L. or a mixture of both plant species were exposed to elevated atmospheric CO₂ (eCO₂) for 10 consecutive seasons using free‐air CO₂ enrichment technology at ETH Zürich, Switzerland. The CO₂ treatment was crossed with a two‐level nitrogen (N) fertilization treatment. In the tenth year, soil samples were collected on three occasions through the growing season to assess the impact of eCO₂ and N fertilization on mycorrhizal fungal abundance. Soil moisture content, which varied with harvest date, was linked to the vegetation type and was higher under eCO₂. Root weight density was affected by vegetation type: lower for clover, higher for grass. Root weight density was stimulated by eCO₂ and decreased by high N fertilization. The percent root length colonized by mycorrhizal fungi was lowest in the clover plots and highest in the grass plots. High N significantly decreased root length colonized. There was no overall effect of eCO₂ on root length colonized; however, there was a significant eCO₂× N interaction: eCO₂ increased root length colonized at high N, but decreased root length colonized at low N. Extraradical mycorrhizal hyphal density was linked to soil moisture content. Extraradical mycorrhizal hyphal density was not affected by eCO₂ or high N individually, but as for root length colonized, there was a significant eCO₂× N interaction: eCO₂ increased extraradical mycorrhizal hyphal density at low N but not at high N. These environmental effects on root colonization and external mycorrhizal hyphae were independent of soil moisture content and root weight density. This field study demonstrated a significant mediating effect of N fertilization on the response of arbuscular mycorrhizal fungi to eCO₂ irrespective of any change in root biomass.     

Responses of fungal root colonization,plant cover and leaf nutrients to long‐term exposure to elevated atmospheric CO2 and warming in a subarctic birch forest understory

Responses of the mycorrhizal fungal community in terrestrial ecosystems to global change factors are not well understood. However, virtually all land plants form symbiotic associations with mycorrhizal fungi, with approximately 20% of the plants' net primary production transported down to the fungal symbionts. In this study, we investigated how ericoid mycorrhiza (ErM), fine endophytes (FE) and dark septate endophytes (DSE) in roots responded to elevated atmospheric CO₂ concentrations and warming in the dwarf shrub understory of a birch forest in the subarctic region of northern Sweden. To place the belowground results into an ecosystem context we also investigated how plant cover and nutrient concentrations in leaves responded to elevated atmospheric CO₂ concentrations and warming. The ErM colonization in ericaceous dwarf shrubs increased under elevated atmospheric CO₂ concentrations, but did not respond to warming following 6 years of treatment. This suggests that the higher ErM colonization under elevated CO₂ might be due to increased transport of carbon belowground to acquire limiting resources such as N, which was diluted in leaves of ericaceous plants under enhanced CO₂. The elevated CO₂ did not affect total plant cover but the plant cover was increased under warming, which might be due to increased N availability in soil. FE colonization in grass roots decreased under enhanced CO₂ and under warming, which might be due to increased root growth, to which the FE fungi could not keep up, resulting in proportionally lower colonization. However, no responses in aboveground cover of Deschampsia flexuosa were seen. DSE hyphal colonization in grass roots significantly increased under warmer conditions, but did not respond to elevated CO₂. This complex set of responses by mycorrhizal and other root‐associated fungi to global change factors of all the fungal types studied could have broad implications for plant community structure and biogeochemistry of subarctic ecosystems.     

Plant species-specific changes in root-inhabiting fungi in a California annual grassland: responses to elevated CO2 and nutrients

Five co-occurring plant species from an annual mediterranean grassland were grown in monoculture for 4 months in pots inside open-top chambers at the Jasper Ridge Biological Preserve (San Mateo County, California). The plants were exposed to elevated atmospheric CO₂ and soil nutrient enrichment in a complete factorial experiment. The response of root-inhabiting non-mycorrhizal and arbuscular mycorrhizal fungi to the altered resource base depended strongly on the plant species. Elevated CO₂ and fertilization altered the ratio of non-mycorrhizal to mycorrhizal fungal colonization for some plant species, but not for others. Percent root infection by non-mycorrhizal fungi increased by over 500% for Linanthus parviflorus in elevated CO₂, but decreased by over 80% for Bromus hordeaceus. By contrast, the mean percent infection by mycorrhizal fungi increased in response to elevated CO₂ for all species, but significantly only for Avena barbata and B. hordeaceus. Percent infection by mycorrhizal fungi increased, decreased, or remained unchanged for different plant hosts in response to fertilization. There was evidence of a strong interaction between the two treatments for some plant species and non-mycorrhizal and mycorrhizal fungi. This study demonstrated plant species- and soil fertility-dependent shifts in below-ground plant resource allocation to different morpho-groups of fungal symbionts. This may have consequences for plant community responses to elevated CO₂ in this California grassland ecosystem. Received: 2 June 1997 / Accepted: 22 August 1997     

Interactions among mycorrhizae, atmospheric CO2 and soil N impact plant community composition

We examined plant community responses to interactions between arbuscular mycorrhizal (AM) fungi and availability of atmospheric CO₂ and soil N. Communities of 14 plant species were grown in mesocosms containing living or killed AM fungal inoculum, ambient or elevated atmospheric CO₂ and low or enriched soil N. After one growing season, significantly different plant communities existed in the different treatments. Plant species richness was lowest in +N mesocosms and highest in +AM + CO₂ mesocosms. At ambient CO₂, AM fungi reduced richness but at elevated CO₂ they increased it. This was caused by changes in mortality rates of several C₃ forbs and may suggest that CO₂ enrichment ameliorates the carbon cost of some AM symbioses. Soil moisture was higher in +CO₂ mesocosms but +AM counteracted this effect. These results suggest that AM symbioses may be important mediators of plant community responses to anthropogenic CO₂ and N enrichment.     

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.     

Expression of genes involved in symbiotic carbon and nitrogen transport in Pinus taeda mycorrhizal roots exposed to CO2 enrichment and nitrogen fertilization

As atmospheric carbon dioxide (CO₂) concentrations rise, one important mechanism by which plants can gain greater access to necessary soil nutrients is through greater investment in their mycorrhizal symbionts. In this study, we tested the hypotheses that (1) plants increase C allocation to ectomycorrhizal fungi (EMF) under elevated CO₂ conditions, (2) N fertilization decreases C allocation to EMF, and (3) EMF activity at the site of symbiotic C and nutrient exchange is enhanced with CO₂ enrichment. To test these hypotheses, we examined expression levels of Pinus taeda genes encoding monosaccharide transport (MST) and ammonium transport (AMT) proteins thought to be involved in symbiotic C and N movement, respectively, from mycorrhizal root tips exposed to CO₂ and N fertilization. We also examined EMF ribosomal RNA expression (18S rRNA) to determine EMF activity. There was a trend toward lower relative MST expression with increased CO₂. AMT expression levels showed no significant differences between control and treatment plots. EMF 18S rRNA expression was increased in CO₂-enriched plots and there was a marginally significant positive interactive effect of CO₂ and N fertilization on expression (p = 0.09 and 0.10, respectively). These results are consistent with greater C allocation to EMF and greater EMF metabolic activity under elevated CO₂ conditions, although selective allocation of C to particular EMF species and greater fungal biomass on roots are plausible alternative hypotheses.     

Species of plants and associated arbuscular mycorrhizal fungi mediate mycorrhizal responses to CO2 enrichment

It has been suggested that enrichment of atmospheric CO₂ should alter mycorrhizal function by simultaneously increasing nutrient‐uptake benefits and decreasing net C costs for host plants. However, this hypothesis has not been sufficiently tested. We conducted three experiments to examine the impacts of CO₂ enrichment on the function of different combinations of plants and arbuscular mycorrhizal (AM) fungi grown under high and low soil nutrient availability. Across the three experiments, AM function was measured in 14 plant species, including forbs, C₃ and C₄ grasses, and plant species that are typically nonmycorrhizal. Five different AM fungal communities were used for inoculum, including mixtures of Glomus spp. and mixtures of Gigasporaceae (i.e. Gigaspora and Scutellospora spp.). Our results do not support the hypothesis that CO₂ enrichment should consistently increase plant growth benefits from AM fungi, but rather, we found CO₂ enrichment frequently reduced AM benefits. Furthermore, we did not find consistent evidence that enrichment of soil nutrients increases plant growth responses to CO₂ enrichment and decreases plant growth responses to AM fungi. Our results show that the strength of AM mutualisms vary significantly among fungal and plant taxa, and that CO₂ levels further mediate AM function. In general, when CO₂ enrichment interacted with AM fungal taxa to affect host plant dry weight, it increased the beneficial effects of Gigasporaceae and reduced the benefits of Glomus spp. Future studies are necessary to assess the importance of temperature, irradiance, and ambient soil fertility in this response. We conclude that the affects of CO₂ enrichment on AM function varies with plant and fungal taxa, and when making predictions about mycorrhizal function, it is unwise to generalize findings based on a narrow range of plant hosts, AM fungi, and environmental conditions.     

Shifts in symbiotic associations in plants capable of forming multiple root symbioses across a long‐term soil chronosequence

Changes in soil nutrient availability during long‐term ecosystem development influence the relative abundances of plant species with different nutrient‐acquisition strategies. These changes in strategies are observed at the community level, but whether they also occur within individual species remains unknown. Plant species forming multiple root symbioses with arbuscular mycorrhizal (AM) fungi, ectomycorrhizal (ECM) fungi, and nitrogen‐(N) fixing microorganisms provide valuable model systems to examine edaphic controls on symbioses related to nutrient acquisition, while simultaneously controlling for plant host identity. We grew two co‐occurring species, Acacia rostellifera (N₂‐fixing and dual AM and ECM symbioses) and Melaleuca systena (AM and ECM dual symbioses), in three soils of contrasting ages (c. 0.1, 1, and 120 ka) collected along a long‐term dune chronosequence in southwestern Australia. The soils differ in the type and strength of nutrient limitation, with primary productivity being limited by N (0.1 ka), co‐limited by N and phosphorus (P) (1 ka), and by P (120 ka). We hypothesized that (i) within‐species root colonization shifts from AM to ECM with increasing soil age, and that (ii) nodulation declines with increasing soil age, reflecting the shift from N to P limitation along the chronosequence. In both species, we observed a shift from AM to ECM root colonization with increasing soil age. In addition, nodulation in A. rostellifera declined with increasing soil age, consistent with a shift from N to P limitation. Shifts from AM to ECM root colonization reflect strengthening P limitation and an increasing proportion of total soil P in organic forms in older soils. This might occur because ECM fungi can access organic P via extracellular phosphatases, while AM fungi do not use organic P. Our results show that plants can shift their resource allocation to different root symbionts depending on nutrient availability during ecosystem development.     

Arbuscular mycorrhizae respond to plants exposed to elevated atmospheric CO2 as a function of soil depth

The importance of arbuscular mycorrhizae (AM) in plant and ecosystem responses to global changes, e.g. elevated atmospheric CO₂, is widely acknowledged. Frequently, increases in AM root colonization occur in response to increased CO₂, but also the lack of significant changes has been reported. The goal of this study was to test whether arbuscular mycorrhizae (root colonization and composition of root colonization) respond to plants grown in elevated CO₂ as a function of soil depth. We grew Bromus hordeaceus L. and Lotus wrangelianus Fischer & C. Meyer monocultures in large pots with a synthetic serpentine soil profile for 4 yr in an experiment, in which CO₂ concentration was crossed factorially with NPK fertilization. When analyzing root infection separately for topsoil (0–15 cm) and subsoil (15–45 cm), we found large (e.g., about 5-fold) increases of AM fungal root colonization in the subsoil in response to CO₂, but no significant changes in the corresponding topsoil of Bromus. Only the coarse endophyte AM fungi, not the fine endophyte AM fungi, were responsible for the observed increase in the bottom soil layer, indicating a depth-dependent shift in the AM community colonizing the roots, even at this coarse morphological level. Other response variables also had significant soil layer ^* CO₂ interaction terms. The subsoil response would have been hidden in an unstratified assessment of the total root system, since most of the root length was concentrated in the top soil layer. The increased presence of mycorrhizae in roots deeper in the soil should be considered in sampling protocols, as it may be indicative of changed patterns of nutrient acquisition and carbon sequestration.     

Arbuscular mycorrhiza improve growth,nitrogen uptake,and nitrogen use efficiency in wheat grown under elevated CO<Subscript>2</Subscript>

Effects of the arbuscular mycorrhizal (AM) fungus Rhizophagus irregularis on plant growth, carbon (C) and nitrogen (N) accumulation, and partitioning was investigated in Triticum aestivum L. plants grown under elevated CO₂ in a pot experiment. Wheat plants inoculated or not inoculated with the AM fungus were grown in two glasshouse cells with different CO₂ concentrations (400 and 700 ppm) for 10 weeks. A ¹⁵N isotope labeling technique was used to trace plant N uptake. Results showed that elevated CO₂ increased AM fungal colonization. Under CO₂ elevation, AM plants had higher C concentration and higher plant biomass than the non-AM plants. CO₂ elevation did not affect C and N partitioning in plant organs, while AM symbiosis increased C and N allocation into the roots. In addition, plant C and N accumulation, ¹⁵N recovery rate, and N use efficiency (NUE) were significantly higher in AM plants than in non-AM controls under CO₂ enrichment. It is concluded that AM symbiosis favors C and N partitioning in roots, increases C accumulation and N uptake, and leads to greater NUE in wheat plants grown at elevated CO₂.     

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.     

毛乌素沙地沙打旺根围AM真菌的空间分布

对毛乌素沙地5个不同生态条件下沙打旺(Astragalus adsurgens)根围土壤0～50 cm土层 AM真菌空间分布进行研究.结果表明：不同样地和采样深度下AM真菌孢子密度和定殖率存在显著差异,最高定殖率和最大孢子密度均出现在10～30 cm土层; 定边样地孢子密度最高,并显著高于其他样地;塔湾样地菌丝和总定殖率最高,宁条梁和塔湾样地泡囊定殖率高于其他样地.总球囊霉素(TG)和易提取球囊霉素(EEG)含量均在塔湾样地最高,分别为1.18 mg·g^-1和0.65 mg·g^-1.土壤pH对孢子密度、泡囊和丛枝定殖率、TG和EEG均有极显著正效应,TG和EEG分别与孢子密度、土壤有机C、碱解N和速效P呈极显著正相关.沙漠土壤球囊霉素占有机C的比例高于农田土壤,可能是沙漠土壤有机C的主要来源之一.球囊霉素可作为评估沙漠土壤AM真菌活动和土壤生态状况的有效指标.     

Net root growth and nutrient acquisition in response to predicted climate change in two contrasting heathland species

Background and aims

Accurate predictions of nutrient acquisition by plant roots and mycorrhizas are critical in modelling plant responses to climate change.

Methods

We conducted a field experiment with the aim to investigate root nutrient uptake in a future climate and studied root production by ingrowth cores, mycorrhizal colonization, and fine root N and P uptake by root assay of Deschampsia flexuosa and Calluna vulgaris.

Results

Net root growth increased under elevated CO₂, warming and drought, with additive effects among the factors. Arbuscular mycorrhizal colonization increased in response to elevated CO₂, while ericoid mycorrhizal colonization was unchanged. The uptake of N and P was not increased proportionally with root growth after 5 years of treatment.

Conclusions

While aboveground biomass was unchanged, the root growth was increased under elevated CO₂. The results suggest that plant production may be limited by N (but not P) when exposed to elevated CO₂. The species-specific response to the treatments suggests different sensitivity to global change factors, which could result in changed plant competitive interactions and belowground nutrient pool sizes in response to future climate change.     

Nitrogen fertilization alters the functioning of arbuscular mycorrhizas at two semiarid grasslands

The effects of nitrogen (N) fertilization on arbuscular mycorrhizas were studied at two semiarid grasslands with different soil properties and N-enrichment history (Shortgrass Steppe in Colorado, and Sevilleta National Wildlife Refuge in New Mexico). These sites are part of the National Science Foundation's Long-Term Ecological Research Network. The experimental plots at Shortgrass Steppe were fertilized with ammonium nitrate (NH₄NO₃) from 1971 to 1975, and have not received additional N since then. The experimental plots at Sevilleta were also fertilized with NH₄NO₃, but were established in 1995, 2 years before the soils were used for this study. Greenhouse experiments were conducted to compare the growth response of local grasses to arbuscular mycorrhizal (AM) fungi from fertilized (FERT) and unfertilized (UNFERT) field soils, at each site. Two species per site were chosen, Bouteloua gracilis and Elymus elymoides from Shortgrass Steppe, and B. gracilis and B. eriopoda from Sevilleta. Plants were grown for 3 months at HIGH N and LOW N levels, with FERT or UNFERT soil inoculum and in a non-mycorrhizal condition. Fertilization with N altered the functioning of AM fungi at both sites. Grasses inoculated with AM fungi from UNFERT soils had the most tillers, greatest biomass and highest relative growth rates. There were no significant differences in the growth response of plants inoculated with AM fungi from FERT soils and the non-mycorrhizal controls. These results were consistent across sites and species except for the plants grown at LOW N in Sevilleta soils. These plants were deficient in N and phosphorus (P) and did not show growth enhancement in response to AM inoculation with either FERT or UNFERT soils. Percent root length colonized by AM fungi was not directly related to plant performance. However, enrichment with N consistently decreased root colonization by AM fungi in the grasses grown in soils from Shortgrass Steppe with high P availability (18.4 mg kg^–1), but not in the grasses grown in Sevilleta soils with low P availability (6.6 mg kg^–1). Our study supports the hypotheses that (1) fertilization with N alters the balance between costs and benefits in mycorrhizal symbioses and (2) AM fungal communities from N fertilized soils are less beneficial mutualists than those from unfertilized soils.     

Does carbon partitioning in ectomycorrhizal pine seedlings under elevated CO2 vary with fungal species?

Enhanced soil respiration in response to elevated atmospheric CO₂ has been demonstrated, and ectomycorrhizal (ECM) fungi are of particular interest since they partition host-derived photoassimilates belowground. Although a strong response of ECM fungi to elevated CO₂ has been shown, little is still known about the functional diversity among species. We studied carbon (C) partitioning in mycorrhizal Scots pine seedlings in response to short-term CO₂ enrichment, using seven ECM species with different ecological strategies. Mycorrhizal associations were synthesised and seedlings grown in large Petri dishes containing peat:vermiculite and nutrient solution for 10–15 weeks, after which half of the microcosms were exposed to elevated CO₂ treatment (710 ppm) for 15 days and the other half were kept in ambient CO₂ treatment. Partitioning of C was quantified by pulse labelling the seedlings with ¹⁴CO₂ and examining the distribution of labelled assimilates in shoot, root and extraradical mycelial compartments by destructive harvest and liquid scintillation counting. Fungal biomass was determined with PLFA analysis. The respiratory loss of ¹⁴CO₂ was on average greater in the elevated CO₂ treatment for most species compared to the ambient CO₂ treatment. More label was retrieved in the shoots in the ambient CO₂ treatment compared to elevated CO₂ (significant for P. involutus and P. fallax). Greater amounts of label were found in the extraradical mycelial compartment in all species (except P. involutus) in elevated CO₂ compared to ambient CO₂ (significant for L. bicolor, P. byssinum, P. fallax and R. roseolus). Fungal biomass production increased significantly with elevated CO₂ for two species (H. velutipes and A. muscaria); three species (P. fallax, P. involutus and R. roseolus) showed a similar but non-significant trend, whereas L. bicolor and P. byssinum produced less biomass in elevated CO₂ compared to ambient CO₂. When ¹⁴C in the mycelial compartment and respiration was expressed per unit fungal PLFA the difference between CO₂ treatments disappeared. We demonstrated that different ECM fungal isolates respond differently in C partitioning in response to CO₂ enrichment. These results suggest that under certain growth conditions, when nutrients are not limiting, ECM fungi respond rapidly to increasing C-availability through changed biomass production and respiration.     

Elevated carbon dioxide and irrigation effects on water stable aggregates in a Sorghum field: a possible role for arbuscular mycorrhizal fungi

While soil biota and processes are becoming increasingly appreciated as important parameters for consideration in global change studies, the fundamental characteristic of soil structure is a neglected area of research. In a sorghum [Sorghum bicolor (L.) Moench] field experiment in which CO₂[supplied using free‐air CO₂ enrichment (FACE) technology] was crossed factorially with an irrigation treatment, soil aggregate (1–2 mm) water stability increased in response to elevated CO₂. Aggregate water stability was increased by 40% and 20% in response to CO₂, at ample and limited water supply treatments, respectively. Soil hyphal lengths of arbuscular mycorrhizal fungi (AMF) increased strongly (with a threefold increase in the dry treatment) in response to CO₂, and the concentrations of one fraction (easily extractable glomalin, EEG) of the AMF‐produced protein glomalin were also increased. Two fractions of glomalin, and AMF hyphal lengths were all positively correlated with soil aggregate water stability. The present results further support the hypothesis that AMF can become important in global change scenarios. Although in this field study a causal relationship between hyphal length, glomalin and aggregate stability cannot be demonstrated, the present data do suggest that AMF could mediate changes in soil structure under elevated CO₂. This could be of great importance in agricultural systems threatened by erosional soil loss.     

Interaction between C4 barnyard grass and C3 upland rice under elevated CO2: Impact of mycorrhizae

Atmospheric CO₂ enrichment may impact arbuscular mycorrhizae (AM) development and function, which could have subsequent effects on host plant species interactions by differentially affecting plant nutrient acquisition. However, direct evidence illustrating this scenario is limited. We examined how elevated CO₂ affects plant growth and whether mycorrhizae mediate interactions between C₄ barnyard grass (Echinochloa crusgalli (L.) Beauv.) and C₃ upland rice (Oryza sativa L.) in a low nutrient soil. The monocultures and combinations with or without mycorrhizal inoculation were grown at ambient (400 ± 20 μmol mol^?1) and elevated CO₂ (700 ± 20 μmol mol^?1) levels. The ¹⁵N isotope tracer was introduced to quantify the mycorrhizally mediated N acquisition of plants. Elevated CO₂ stimulated the growth of C₃ upland rice but not that of C₄ barnyard grass under monoculture. Elevated CO₂ also increased mycorrhizal colonization of C₄ barnyard grass but did not affect mycorrhizal colonization of C₃ upland rice. Mycorrhizal inoculation increased the shoot biomass ratio of C₄ barnyard grass to C₃ upland rice under both CO₂ concentrations but had a greater impact under the elevated than ambient CO₂ level. Mycorrhizae decreased relative interaction index (RII) of C₃ plants under both ambient and elevated CO₂, but mycorrhizae increased RII of C₄ plants only under elevated CO₂. Elevated CO₂ and mycorrhizal inoculation enhanced ¹⁵N and total N and P uptake of C₄ barnyard grass in mixture but had no effects on N and P acquisition of C₃ upland rice, thus altering the distribution of N and P between the species in mixture. These results implied that CO₂ stimulation of mycorrhizae and their nutrient acquisition may impact competitive interaction of C₄ barnyard grass and C₃ upland rice under future CO₂ scenarios.