Mycorrhizal dynamics under elevated CO2 and nitrogen fertilization in a warm temperate forest

We examined the response of mycorrhizal fungi to free-air CO₂ enrichment (FACE) and nitrogen (N) fertilization in a warm temperate forest to better understand potential influences over plant nutrient uptake and soil carbon (C) storage. In particular, we hypothesized that mycorrhizal fungi and glomalin would become more prevalent under elevated CO₂ but decrease under N fertilization. In addition, we predicted that N fertilization would mitigate any positive effects of elevated CO₂ on mycorrhizal abundance. Overall, we observed a 14% increase in ectomycorrhizal (ECM) root colonization under CO₂ enrichment, which implies that elevated CO₂ results in greater C investments in these fungi. Arbuscular mycorrhizal (AM) hyphal length and glomalin stocks did not respond substantially to CO₂ enrichment, and effects of CO₂ on AM root colonization varied by date. Nitrogen effects on AM fungi were not consistent with our hypothesis, as we found an increase in AM colonization under N fertilization. Lastly, neither glomalin concentrations nor ECM colonization responded significantly to N fertilization or to an N-by-CO₂ interaction. A longer duration of N fertilization may be required to detect effects on these parameters.     

Response of soil biota to elevated atmospheric CO2 in poplar model systems

We tested the hypotheses that increased belowground allocation of carbon by hybrid poplar saplings grown under elevated atmospheric CO₂ would increase mass or turnover of soil biota in bulk but not in rhizosphere soil. Hybrid poplar saplings (Populus×euramericana cv. Eugenei) were grown for 5 months in open-bottom root boxes at the University of Michigan Biological Station in northern, lower Michigan. The experimental design was a randomized-block design with factorial combinations of high or low soil N and ambient (34 Pa) or elevated (69 Pa) CO₂ in five blocks. Rhizosphere microbial biomass carbon was 1.7 times greater in high-than in low-N soil, and did not respond to elevated CO₂. The density of protozoa did not respond to soil N but increased marginally (P < 0.06) under elevated CO₂. Only in high-N soil did arbuscular mycorrhizal fungi and microarthropods respond to CO₂. In high-N soil, arbuscular mycorrhizal root mass was twice as great, and extramatrical hyphae were 11% longer in elevated than in ambient CO₂ treatments. Microarthropod density and activity were determined in situ using minirhizotrons. Microarthropod density did not change in response to elevated CO₂, but in high-N soil, microarthropods were more strongly associated with fine roots under elevated than ambient treatments. Overall, in contrast to the hypotheses, the strongest response to elevated atmospheric CO₂ was in the rhizosphere where (1) unchanged microbial biomass and greater numbers of protozoa (P < 0.06) suggested faster bacterial turnover, (2) arbuscular mycorrhizal root length increased, and (3) the number of microarthropods observed on fine roots rose. Received: 18 March 1997 / Accepted: 5 August 1997     

13C and 15N in microarthropods reveal little response of Douglas-fir ecosystems to climate change

Understanding ecosystem carbon (C) and nitrogen (N) cycling under global change requires experiments maintaining natural interactions among soil structure, soil communities, nutrient availability, and plant growth. In model Douglas-fir ecosystems maintained for five growing seasons, elevated temperature and carbon dioxide (CO₂) increased photosynthesis and increased C storage belowground but not aboveground. We hypothesized that interactions between N cycling and C fluxes through two main groups of microbes, mycorrhizal fungi (symbiotic with plants) and saprotrophic fungi (free-living), mediated ecosystem C storage. To quantify proportions of mycorrhizal and saprotrophic fungi, we measured stable isotopes in fungivorous microarthropods that efficiently censused the fungal community. Fungivorous microarthropods consumed on average 35% mycorrhizal fungi and 65% saprotrophic fungi. Elevated temperature decreased C flux through mycorrhizal fungi by 7%, whereas elevated CO₂ increased it by 4%. The dietary proportion of mycorrhizal fungi correlated across treatments with total plant biomass (n= 4, r²= 0.96, P= 0.021), but not with root biomass. This suggests that belowground allocation increased with increasing plant biomass, but that mycorrhizal fungi were stronger sinks for recent photosynthate than roots. Low N content of needles (0.8–1.1%) and A horizon soil (0.11%) coupled with high C : N ratios of A horizon soil (25–26) and litter (36–48) indicated severe N limitation. Elevated temperature treatments increased the saprotrophic decomposition of litter and lowered litter C : N ratios. Because of low N availability of this litter, its decomposition presumably increased N immobilization belowground, thereby restricting soil N availability for both mycorrhizal fungi and plant growth. Although increased photosynthesis with elevated CO₂ increased allocation of C to ectomycorrhizal fungi, it did not benefit plant N status. Most N for plants and soil storage was derived from litter decomposition. N sequestration by mycorrhizal fungi and limited N release during litter decomposition by saprotrophic fungi restricted N supply to plants, thereby constraining plant growth response to the different treatments.     

Role of nutrient level and defoliation on symbiotic function: experimental evidence by tracing 14C/15N exchange in mycorrhizal birch seedlings

High nutrient availability and defoliation generally reduce ectomycorrhizal colonization levels in trees, but it is not known how this affects the functional aspects of mycorrhizal symbiosis. It was therefore investigated whether (1) defoliation or increasing substrate N availability reduce C allocation from the plant to the fungus and N allocation from the fungus to the plant (symbiotic resource exchange), (2) symbiotic resource exchange depends on relative N and P availability, and (3) fungal N translocation to plant and plant C allocation to fungus are interdependent. Birch (Betula pendula) seedlings were grown in symbiosis with the ectomycorrhizal fungus Paxillus involutus at five times excess N, or at five times excess N and P for 6 weeks. One-half of the plants were defoliated and the plant shoots were allowed to photosynthesize ¹⁴CO₂ while the fungal compartment was exposed to ¹⁴NH₄. After 3 days, the ¹⁴C of plant origin in fungal tissues and ¹⁵N of fungal origin in plant tissues were quantified. Nutrient availability had no observable effect on symbiotic resource exchange in non-defoliated systems. Defoliation reduced symbiotic N acquisition by plants at all levels of nutrient availability, with the reduction being most marked at higher N availability, indicating an increased tendency in the symbiotic system to discontinue resource exchange after defoliation at higher fertility levels. The concentration of ¹⁴C in extramatrical mycelium correlated significantly with the concentration of ¹⁵N in birch shoots. The results support the assumption that N delivery to the host by the mycorrhizal fungus is dependent on C flow from the plant to the fungus, and that exchanges between the partners are reciprocal. No significant reductions in root ¹⁴C content as a response to defoliation were observed, indicating that defoliation specifically reduced allocation to fungus, but not markedly to roots.     

Nitrogen nutrition,photosynthesis and carbon allocation in ectomycorrhizal pine

Summary Studies examined net photosynthesis (Pn) and dry matter production of mycorrhizal and nonmycorrhizalPinus taeda at 6 intervals over a 10-month period. Pn rates of mycorrhizal plants were consistently greater than nonmycorrhizal plants, and at 10 months were 2.1-fold greater. Partitioning of current photosynthate was examined by pulse-labelling with¹⁴CO₂ at each of the six time intervals. Mycorrhizal plants assimilated more¹⁴CO₂, allocated a greater percentage of assimilated¹⁴C to the root systems, and lost a greater percentage of¹⁴C by root respiration than did nonmycorrhizal plants. At 10 months, the quantity of¹⁴CO₂ respired by roots per unit root weight was 3.6-fold greater by mycorrhizal than nonmycorrhizal plants. Although the stimulation of photosynthesis and translocation of current photosynthate to the root system by mycorrhiza formation was consistent with the source-sink concept of sink demand, foliar N and P concentrations were also greater in mycorrhizal plants.Further studies examined Pn and dry matter production ofPinus contorta in response to various combinations of N fertilization (3, 62, 248 ppm), irradiance and mycorrhizal fungi inoculation. At 16 weeks of age, 6 weeks following inoculation with eitherPisolithus tinctorius orSuillus granulatus, Pn rates and biomass were significantly greater in mycorrhizal than nonmycorrhizal plants. Mycorrhizal plants had significantly greater foliar %P, but not %N, than did nonmycorrhizal plants. Fertilization with 62 ppm N resulted in greater mycorrhiza formation than either 3 or 248 ppm. Increased irradiance resulted in increased mycorrhiza formation.     

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.     

Populus tremuloides photosynthesis and crown architecture in response to elevated CO2 and soil N availability

We tested the hypothesis that elevated CO₂ would stimulate proportionally higher photosynthesis in the lower crown of Populus trees due to less N retranslocation, compared to tree crowns in ambient CO₂. Such a response could increase belowground C allocation, particularly in trees with an indeterminate growth pattern such as Populus tremuloides. Rooted cuttings of P. tremuloides were grown in ambient and twice ambient (elevated) CO₂ and in low and high soil N availability (89 ± 7 and 333 ± 16 ng N g⁻¹ day⁻¹ net mineralization, respectively) for 95 days using open-top chambers and open-bottom root boxes. Elevated CO₂ resulted in significantly higher maximum leaf photosynthesis (A _max) at both soil N levels. A _max was higher at high N than at low N soil in elevated, but not ambient CO₂. Photosynthetic N use efficiency was higher at elevated than ambient CO₂ in both soil types. Elevated CO₂ resulted in proportionally higher whole leaf A in the lower three-quarters to one-half of the crown for both soil types. At elevated CO₂ and high N availability, lower crown leaves had significantly lower ratios of carboxylation capacity to electron transport capacity (V _cmax/J _max) than at ambient CO₂ and/or low N availability. From the top to the bottom of the tree crowns, V _cmax/J _max increased in ambient CO₂, but it decreased in elevated CO₂ indicating a greater relative investment of N into light harvesting for the lower crown. Only the mid-crown leaves at both N levels exhibited photosynthetic down regulation to elevated CO₂. Stem biomass segments (consisting of three nodes and internodes) were compared to the total A _leaf for each segment. This analysis indicated that increased A _leaf at elevated CO₂ did not result in a proportional increase in local stem segment mass, suggesting that C allocation to sinks other than the local stem segment increased disproportionally. Since C allocated to roots in young Populus trees is primarily assimilated by leaves in the lower crown, the results of this study suggest a mechanism by which C allocation to roots in young trees may increase in elevated CO₂. Received: 12 August 1996 / Accepted: 12 November 1996     

Nutrient relations in calcareous grassland under elevated CO2

Plant nutrient responses to 4 years of CO₂ enrichment were investigated in situ in calcareous grassland. Beginning in year 2, plant aboveground C:N ratios were increased by 9% to 22% at elevated CO₂ (P < 0.01), depending on year. Total amounts of N removed in biomass harvests during the first 4 years were not affected by elevated CO₂ (19.9 ± 1.3 and 21.1 ± 1.3 g N m⁻² at ambient and elevated CO₂), indicating that the observed plant biomass increases were solely attained by dilution of nutrients. Total aboveground P and tissue N:P ratios also were not altered by CO₂ enrichment (12.5 ± 2 g N g⁻¹ P in both treatments). In contrast to non-legumes (>98% of community aboveground biomass), legume C/N was not reduced at elevated CO₂ and legume N:P was slightly increased. We attribute the less reduced N concentration in legumes at elevated CO₂ to the fact that virtually all legume N originated from symbiotic N₂ fixation (%N_dfa ≈ 90%), and thus legume growth was not limited by soil N. While total plant N was not affected by elevated CO₂, microbial N pools increased by +18% under CO₂ enrichment (P = 0.04) and plant available soil N decreased. Hence, there was a net increase in the overall biotic N pool, largely due increases in the microbial N pool. In order to assess the effects of legumes for ecosystem CO₂ responses and to estimate the degree to which plant growth was P-limited, two greenhouse experiments were conducted, using firstly undisturbed grassland monoliths from the field site, and secondly designed `microcosm' communities on natural soil. Half the microcosms were planted with legumes and half were planted without. Both monoliths and microcosms were exposed to elevated CO₂ and P fertilization in a factored design. After two seasons, plant N pools in both unfertilized monoliths and microcosm communities were unaffected by CO₂ enrichment, similar to what was found in the field. However, when P was added total plant N pools increased at elevated CO₂. This community-level effect originated almost solely from legume stimulation. The results suggest a complex interaction between atmospheric CO₂ concentrations, N and P supply. Overall ecosystem productivity is N-limited, whereas CO₂ effects on legume growth and their N₂ fixation are limited by P. Received: 12 July 1997 / Accepted: 15 April 1998     

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.     

Elevated atmospheric CO2 and feedback between carbon and nitrogen cycles

We tested a conceptual model describing the influence of elevated atmospheric CO₂ on plant production, soil microorganisms, and the cycling of C and N in the plant-soil system. Our model is based on the observation that in nutrient-poor soils, plants (C₃) grown in an elevated CO₂ atmosphere often increase production and allocation to belowground structures. We predicted that greater belowground C inputs at elevated CO₂ should elicit an increase in soil microbial biomass and increased rates of organic matter turnover and nitrogen availability. We measured photosynthesis, biomass production, and C allocation of Populus grandidentata Michx. grown in nutrient-poor soil for one field season at ambient and twice-ambient (i.e., elevated) atmospheric CO₂ concentrations. Plants were grown in a sandy subsurface soil i) at ambient CO₂ with no open top chamber, ii) at ambient CO₂ in an open top chamber, and iii) at twice-ambient CO₂ in an open top chamber. Plants were fertilized with 4.5 g N m⁻² over a 47 d period midway through the growing season. Following 152 d of growth, we quantified microbial biomass and the availabilities of C and N in rhizosphere and bulk soil. We tested for a significant CO₂ effect on plant growth and soil C and N dynamics by comparing the means of the chambered ambient and chambered elevated CO₂ treatments. Rates of photosynthesis in plants grown at elevated CO₂ were significantly greater than those measured under ambient conditions. The number of roots, root length, and root length increment were also substantially greater at elevated CO₂. Total and belowground biomass were significantly greater at elevated CO₂. Under N-limited conditions, plants allocated 50–70% of their biomass to roots. Labile C in the rhizosphere of elevated-grown plants was significantly greater than that measured in the ambient treatments; there were no significant differences between labile C pools in the bulk soil of ambient and elevated-grown plants. Microbial biomass C was significantly greater in the rhizosphere and bulk soil of plants grown at elevated CO₂ compared to that in the ambient treatment. Moreover, a short-term laboratory assay of N mineralization indicated that N availability was significantly greater in the bulk soil of the elevated-grown plants. Our results suggest that elevated atmospheric CO₂ concentrations can have a positive feedback effect on soil C and N dynamics producing greater N availability. Experiments conducted for longer periods of time will be necessary to test the potential for negative feedback due to altered leaf litter chemistry. ei]{gnH}{fnLambers} ei]{gnA C}{fnBorstlap}     

Combined effects of elevated CO2 and soil drought on carbon and nitrogen allocation of the desert shrub Caragana intermedia

Impacts of either elevated CO₂ or drought stress on plant growth have been studied extensively, but interactive effects of these on plant carbon and nitrogen allocation is inadequately understood yet. In this study the response of the dominant desert shrub, Caragana intermedia Kuanget H.c.Fu, to the interaction of elevated CO₂ (700 ± 20 μmol mol⁻¹) and soil drought were determined in two large environmental growth chambers (18 m²). Elevated CO₂ increased the allocation of biomass and carbon into roots and the ratio of carbon to nitrogen (C:N) as well as the leaf soluble sugar content, but decreased the allocation of biomass and carbon into leaves, leaf nitrogen and leaf soluble protein concentrations. Elevated CO₂ significantly decreased the partitioning of nitrogen into leaves, but increased that into roots, especially under soil drought. Elevated CO₂ significantly decreased the carbon isotope discrimination (Δ) in leaves, but increased them in roots, and the ratio of Δ values between root and leaf, indicating an increased allocation into below-ground parts. It is concluded that stimulation of plant growth by CO₂ enrichment may be negated under soil drought, and under the future environment, elevated CO₂ may partially offset the negative effects of enhanced drought by regulating the partitioning of carbon and nitrogen.     

Direct evidence that symbiotic N2 fixation in fertile grassland is an important trait for a strong response of plants to elevated atmospheric CO2

Although legumes showed a clearly superior yield response to elevated atmospheric pCO₂ compared to nonlegumes in a variety of field experiments, the extent to which this is due to symbiotic N₂ fixation per se has yet to be determined. Thus, effectively and ineffectively nodulating lucerne (Medicago sativa L.) plants with a very similar genetic background were grown in competition with each other on fertile soil in the Swiss FACE experiment in order to monitor their CO₂ response. Under elevated atmospheric pCO₂, effectively nodulating lucerne, thus capable of symbiotically fixing N₂, strongly increased the harvestable biomass and the N yield, independent of N fertilization. In contrast, the harvestable biomass and N yield of ineffectively nodulating plants were affected negatively by elevated atmospheric pCO₂ when N fertilization was low. Large amounts of N fertilizer enabled the plants to respond more favourably to elevated atmospheric pCO₂, although not as strongly as effectively nodulating plants. The CO₂‐induced increase in N yield of the effectively nodulating plants was attributed solely to an increase in symbiotic N₂ fixation of 50–175%, depending on the N fertilization treatment. N yield derived from the uptake of mineral N from the soil was, however, not affected by elevated pCO₂. This result demonstrates that, in fertile soil and under temperate climatic conditions, symbiotic N₂ fixation per se is responsible for the considerably greater amount of above‐ground biomass and the higher N yield under elevated atmospheric pCO₂. This supports the assumption that symbiotic N₂ fixation plays a key role in maintaining the C/N balance in terrestrial ecosystems in a CO₂‐rich world.     

Root respiratory quotient and nitrate uptake in hydroponically grown non-mycorrhizal and mycorrhizal wheat

 Oxygen and CO₂ fluxes were measured in hydroponically grown mycorrhizal and non-mycorrhizal Triticum aestivum L. cv. Hano roots. The NO₃ ^– uptake of the plants was used to estimate the amount of root respiration attributable to ion uptake. Plants were grown at 4 mM N and 10 μM P, where a total and viable mycorrhizal root colonisation of 48% and 18%, respectively, by Glomus mosseae (Nicol. and Gerd.) Gerd. and Trappe (BEG 107) was observed. The O₂ consumption and NO₃ ^– uptake rates were similar and the CO₂ release was higher in mycorrhizal than in non-mycorrhizal wheat. This resulted in a significantly higher respiratory quotient (RQ, mol CO₂ mol^–1 O₂) in mycorrhizal (1.27±0.13) than in non-mycorrhizal (0.79±0.05) wheat. As the biomass and N and P concentrations in mycorrhizal and non-mycorrhizal wheat were the same, the higher RQ resulted from the mycorrhizal colonisation and not differences in nutrition per se. Accepted: 26 March 1999     

Mineralization of carbon and nitrogen in soil samples taken from three fertilized pine stands: Long-term effects

Seven years after fertilization the rate of CO₂ production in the soil samples taken from the organic horizons of a poor pine forest site (Calluna vulgaris site type), treated with urea or ammonium nitrate with lime, was lower than that in the unfertilized soil. The same trend was also observed in samples of theEmpetrum-Calluna site type 14 years after fertilization. In the more fertileVaccinium myrtillus site type these rapidly-soluble N fertilizers had a long-term enhancing effect on the production of CO₂. Apatite and biotite eliminated the decreasing effect of urea on the production of CO₂. One reason for this might be the long-term increase in soil pH caused by apatite and biotite, or their constituents (Ca, Mg, K, P). Nitroform (a slow-releasing N fertilizer) had no statistically significant effect on the production of CO₂ in soil samples from any of the forest types. Despite the high N mineralization in the samples from nitroform fertilized soils there was no nitrification, and the high content of total N indicated that after nitroform fertilization the losses of N were low.The correlation between the net mineralization values for C (CO₂ production) and N was poor. However, multiple linear regression analysis, which also took into account the effect of nutrients and pH, indicated that there was a link between the mineralization of C and N.     

Elevated CO2 and temperature alter nitrogen allocation in Douglas-fir

The effects of elevated CO₂ and temperature on principal carbon constituents (PCC) and C and N allocation between needle, woody (stem and branches) and root tissue of Pseudotsuga menziesii Mirb. Franco seedlings were determined. The seedlings were grown in sun‐lit controlled‐environment chambers that contained a native soil. Chambers were controlled to reproduce ambient or ambient +180 ppm CO₂ and either ambient temperature or ambient +3.5 °C for 4 years. There were no significant CO₂ × temperature interactions; consequently the data are presented for the CO₂ and temperature effects. At the final harvest, elevated CO₂ decreased the nonpolar fraction of the PCC and increased the polar fraction and amount of sugars in the needles. In contrast, elevated temperature increased the nonpolar fraction of the PCC and decreased sugars in needles. There were no CO₂ or temperature effects on the PCC fractions in the woody tissue or root tissue. Elevated CO₂ and temperature had no significant effects on the C content of any of the plant tissues or fractions. In contrast, the foliar N content declined under elevated CO₂ and increased under elevated temperature; there were no significant effects in other tissues. The changes in the foliar N concentrations were in the cellulose and lignin fractions, the fractions, which contain protein, and are the consequences of changes in N allocation under the treatments. These results indicate reallocation of N among plant organs to optimize C assimilation, which is mediated via changes in the selectivity of Rubisco and carbohydrate modulation of gene expression.     

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.     

Metabolite analysis of the effects of elevated CO2 and nitrogen fertilization on the association between tall fescue (Schedonorus arundinaceus) and its fungal symbiont Neotyphodium coenophialum

Atmospheric CO₂ is expected to increase to between 550 ppm and 1000 ppm in the next century. CO₂‐induced changes in plant physiology can have ecosystem‐wide implications and may alter plant‐plant, plant‐herbivore and plant‐symbiont interactions. We examined the effects of three concentrations of CO₂ (390, 800 and 1000 ppm) and two concentrations of nitrogen fertilizer (0.004 g N/week versus 0.2 g N/week) on the physiological response of Neotyphodium fungal endophyte‐infected and uninfected tall fescue plants. We used quantitative PCR to estimate the concentration of endophyte under altered CO₂ and N conditions. We found that elevated CO₂ increased the concentration of water‐soluble carbohydrates and decreased the concentration of plant total amino acids in plants. Fungal‐derived alkaloids decreased in response to elevated CO₂ and increased in response to nitrogen fertilization. Endophyte concentration (expressed as the number of copies of an endophyte‐specific gene per total genomic DNA) increased under elevated CO₂ and nitrogen fertilization. The correlation between endophyte concentration and alkaloid production observed at ambient conditions was not observed under elevated CO₂. These results suggest that nutrient exchange dynamics important for maintaining the symbiotic relationship between fungal endophytes and their grass hosts may be altered by changes in environmental variables such as CO₂ and nitrogen fertilization.     

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     

Forest fine-root production and nitrogen use under elevated CO2: contrasting responses in evergreen and deciduous trees explained by a common principle

Despite the importance of nitrogen (N) limitation of forest carbon (C) sequestration at rising atmospheric CO₂ concentration, the mechanisms responsible are not well understood. To elucidate the interactive effects of elevated CO₂ (eCO₂) and soil N availability on forest productivity and C allocation, we hypothesized that (1) trees maximize fitness by allocating N and C to maximize their net growth and (2) that N uptake is controlled by soil N availability and root exploration for soil N. We tested this model using data collected in Free‐Air CO₂ Enrichment sites dominated by evergreen (Pinus taeda; Duke Forest) and deciduous [Liquidambar styraciflua; Oak Ridge National Laboratory (ORNL)] trees. The model explained 80–95% of variation in productivity and N‐uptake data among eCO₂, N fertilization and control treatments over 6 years. The model explains why fine‐root production increased, and why N uptake increased despite reduced soil N availability under eCO₂ at ORNL and Duke. In agreement with observations at other sites, the model predicts that soil N availability reduced below a critical level diminishes all eCO₂ responses. At Duke, a negative feedback between reduced soil N availability and N uptake prevented progressive reduction in soil N availability at eCO₂. At ORNL, soil N availability progressively decreased because it did not trigger reductions in N uptake; N uptake was maintained at ORNL through a large increase in the production of fast turnover fine roots. This implies that species with fast root turnover could be more prone to progressive N limitation of carbon sequestration in woody biomass than species with slow root turnover, such as evergreens. However, longer term data are necessary for a thorough evaluation of this hypothesis. The success of the model suggests that the principle of maximization of net growth to control growth and allocation could serve as a basis for simplification and generalization of larger scale forest and ecosystem models, for example by removing the need to specify parameters for relative foliage/stem/root allocation.     

Modifications of the carbon and nitrogen allocations in the plant (Triticum aestivum L.) soil system in response to increased atmospheric CO2 concentration

The aim of this work was to examine the response of wheat plants to a doubling of the atmospheric CO₂ concentration on: (1) carbon and nitrogen partitioning in the plant; (2) carbon release by the roots; and (3) the subsequent N uptake by the plants. The experiment was performed in controlled laboratory conditions by exposing fast-growing spring wheat plants, during 28 days, to a ¹⁴CO₂ concentration of 350 or 700 L L^–1 at two levels of soil nitrogen fertilization. Doubling CO₂ availability increased total plant production by 34% for both N treatment. In the N-fertilized soil, the CO₂ enrichment resulted in an increase in dry mass production of 41% in the shoots and 23% in the roots; without N fertilization this figure was 33% and 37%, respectively. In the N-fertilized soil, the CO₂ increase enhanced the total N uptake by 14% and lowered the N concentration in the shoots by 23%. The N concentration in the roots was unchanged. In the N-fertilized soil, doubling CO₂ availability increased N uptake by 32% but did not change the N concentrations, in either shoots or roots. The CO₂ enrichment increased total root-derived carbon by 12% with N fertilization, and by 24% without N fertilization. Between 85 and 90% of the total root derived-¹⁴C came from respiration, leaving only 10 to 15% in the soil as organic ¹⁴C. However, when total root-derived ¹⁴C was expressed as a function of root dry weight, these differences were only slightly significant. Thus, it appears that the enhanced carbon release from the living roots in response to increased atmospheric CO₂, is not due to a modification of the activity of the roots, but is a result of the increased size of the root system. The increase of root dry mass also resulted in a stimulation of the soil N mineralization related to the doubling atmospheric CO₂ concentration. The discussion is focused on the interactions between the carbon and nitrogen allocation, especially to the root system, and the implications for the acquisition of nutrients by plants in response to CO₂ increase.Abbreviations N soil fertilization without nitrogen - N soil fertilization with nitrogen