Competition for nitrogen between Pinus sylvestris and ectomycorrhizal fungi generates potential for negative feedback under elevated CO2

We investigated fungal species-specific responses of ectomycorrhizal (ECM) Scots pine (Pinus sylvestris) seedlings on growth and nutrient acquisition together with mycelial development under ambient and elevated CO₂. Each seedling was associated with one of the following ECM species: Hebeloma cylindrosporum, Laccaria bicolor, Suillus bovinus, S. luteus, Piloderma croceum, Paxillus involutus, Boletus badius, or non-mycorrhizal, under ambient, and elevated CO₂ (350 or 700 μl l⁻¹ CO₂); each treatment contained six replicates. The trial lasted 156 days. During the final 28 days, the seedlings were labeled with ¹⁴CO₂. We measured hyphal length, plant biomass, ¹⁴C allocation, and plant nitrogen and phosphorus concentration. Almost all parameters were significantly affected by fungal species and/or CO₂. There were very few significant interactions. Elevated CO₂ decreased shoot-to-root ratio, most strongly so in species with the largest extraradical mycelium. Under elevated CO₂, ECM root growth increased significantly more than hyphal growth. Extraradical hyphal length was significantly negatively correlated with shoot biomass, shoot N content, and total plant N uptake. Root dry weight was significantly negatively correlated with root N and P concentration. Fungal sink strength for N strongly affected plant growth through N immobilization. Mycorrhizal fungal-induced progressive nitrogen limitation (PNL) has the potential to generate negative feedback with plant growth under elevated CO₂. Responsible Editor: Herbert Johannes Kronzucker     

Ectomycorrhizal fungi associated with Pinus sylvestris seedlings respond differently to increased carbon and nitrogen availability: implications for ecosystem responses to global change

The ectomycorrhizal (ECM) symbiosis can cause both positive and negative feedback with trees under elevated CO₂. Positive feedback arises if the additional carbon (C) increases both nutrient uptake by the fungus and nutrient transfer to the plant, whereas negative feedback results from increased nutrient uptake and immobilization by the fungus and reduced transfer to the plant. Because species of ECM fungi differ in their C and nitrogen (N) demand, understanding fungal species‐specific responses to variation in C and N supply is essential to predict impacts of global change. We investigated fungal species‐specific responses of ECM Scots pine (Pinus sylvestris) seedlings under ambient and elevated CO₂ (350 or 700 μL L⁻¹ CO₂) and under low and high mineral N availability. Each seedling was associated with one of the following ECM species: Hebeloma cylindrosporum, Laccaria bicolor and Suillus bovinus. The experiment lasted 103 days. During the final 27 days, seedlings were labeled with ¹⁴CO₂ and ¹⁵N. Most plant and fungal parameters were significantly affected by fungal species, CO₂ level and N supply. Interactions between fungal species and CO₂ were also regularly significant. At low N availability, elevated CO₂ had the smallest impact on the photosynthetic performance of seedlings inoculated with H. cylindrosporum and the largest impact on seedlings with S. bovinus. At ambient CO₂, increasing N supply had the smallest impact on seedlings inoculated with S. bovinus and the largest on seedlings inoculated with H. cylindrosporum. At low N availability, extraradical hyphal length increased after doubling CO₂ level, but this was significant only for L. bicolor. At ambient CO₂, increasing N levels reduced hyphal length for both H. cylindrosporum and S. bovinus, but not for L. bicolor. We discuss the potential interplay of two major elements of global change, elevated CO₂ and increased N availability, and their effects on plant growth. We conclude that increased N supply potentially relieves mycorrhiza‐induced progressive N limitation under elevated CO₂.     

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.     

Ectomycorrhizal fungal communities associated with Pinus thunbergii in the eastern coastal pine forests of Korea

We investigated the ectomycorrhizal (ECM) fungal colonization status of Pinus thunbergii mature trees and regenerating seedlings varying in age in coastal pine forests on the east coast of Korea. We established one 20 × 20-m plot at each of two study sites at P. thunbergii coastal forests in Samcheok. Fifty soil blocks (5 × 5 × 15 cm) were sampled at regular intervals, and ten P. thunbergii seedlings of age 0, 1–3, 3–5, and 5–10 years were sampled in each study plot. In total of 27 ECM fungal taxa, Cenococcum geophilum was dominant, followed by Russula sp., Sebacina sp., and unidentified Cortinuris sp. in mature trees. In 0-year-old seedlings, some fungal species such as Sebacina sp., C. geophilum, and unidentified Cortinarius sp. were dominant whereas only C. geophilum was dominant after 1 year, and there were no apparent succession patterns in ECM fungal compositions beyond a host age of 1 year. Most ECM fungal taxa that had colonized seedlings of each age class were also observed in roots of mature trees in each site. These taxa accounted for 86.7–100% and 96.4–98.4% of ECM abundance in seedlings and mature trees, respectively. The results indicate that the species composition of ECM fungal taxa colonizing seedlings of different age in forests is similar to that of surrounding mature trees. Our results also showed that C. geophilum is a common and dominant ECM fungus in P. thunbergii coastal forests and might play a significant role in their regeneration.     

Nitrogen dynamics and growth of seedlings of an N-fixing tree (Gliricidia sepium (Jacq.) Walp.) exposed to elevated atmospheric carbon dioxide

Summary Seeds of Gliricidia sepium (Jacq.) Walp., a tree native to seasonal tropical forests of Central America, were inoculated with N-fixing Rhizobium bacteria and grown in growth chambers for 71 days to investigate interactive effects of atmospheric CO₂ and plant N status on early seedling growth, nodulation, and N accretion. Seedlings were grown with CO₂ partial pressures of 350 and 650 bar (current ambient and a predicted partial pressure of the mid-21st century) and with plus N or minus N nutrient solutions to control soil N status. Of particular interest was seedling response to CO₂ when grown without available soil N, a condition in which seedlings initially experienced severe N deficiency because bacterial N-fixation was the sole source of N. Biomass of leaves, stems, and roots increased significantly with CO₂ enrichment (by 32%, 15% and 26%, respectively) provided seedlings were supplied with N fertilizer. Leaf biomass of N-deficient seedlings was increased 50% by CO₂ enrichment but there was little indication that photosynthate translocation from leaves to roots or that plant N (fixed by Rhizobium) was altered by elevated CO₂. In seedlings supplied with soil N, elevated CO₂ increased average nodule weight, total nodule weight per plant, and the amount of leaf nitrogen provided by N-fixation (as indicated by leaf ¹⁵N). While CO₂ enrichment reduced the N concentration of some plant tissues, whole plant N accretion increased. Results support the contention that increasing atmospheric CO₂ partial pressures will enhance productivity and N-fixing activity of N-fixing tree seedlings, but that the magnitude of early seedling response to CO₂ will depend greatly on plant and soil nutrient status.     

Elevated CO2 alleviates the effects of low P on the growth of N2-fixing Acacia auriculiformis and Acacia mangium

Nodulated seedlings of Acacia auriculiformis Cunn. ex Benth and Acacia mangium Willd were grown with different phosphorus (P) regimes for 90 days, and half of them were exposed to elevated CO₂ (800 μl l⁻¹) during the last 30 days. Under ambient CO₂, plant growth and the amount of N fixed symbiotically in N₂-fixing seedlings decreased with the decrease of supplied P; this relationship did not occur under elevated CO₂. The increase in plant biomass by elevated CO₂ at low P was accompanied by the increase in internal P use efficiency, the amount of N fixed symbiotically and N use efficiency. Elevated CO₂ recovered the low P-induced reduction in leaf dry matter per unit area or unit fresh weight, but it had no effect on the low P-induced increase in partitioning dry matter to roots. These results suggest that elevated CO₂ alleviates the low P effect mainly by increasing the use efficiency of internal P for plant growth and symbiotic N₂ fixation, and the source-sink relationship is possibly an important driving force for this effect of elevated CO₂ in A. auriculiformis and A. mangium.     

Growth,nutrient dynamics,and efficiency responses to carbon dioxide and phosphorus nutrition in soybean

Plant mineral nutrients such as phosphorus may exert major control on crop responses to the rising atmospheric carbon dioxide (CO₂) concentrations. To evaluate the growth, nutrient dynamics, and efficiency responses to CO₂ and phosphorus nutrition, soybean (Glycine max (L.) Merr.) was grown in controlled environment growth chambers with sufficient (0.50 mM) and deficient (0.10 and 0.01 mM) phosphate (Pi) supply under ambient and elevated CO₂ (aCO₂, 400 and eCO₂, 800 µmol mol^?1, respectively). The CO₂ × Pi interaction was detected for leaf area, leaf and stem dry weight, and total plant biomass. The severe decrease in plant biomass in Pi-deficient plants (10–76%) was associated with reduced leaf area and photosynthesis (P_net). The degree of growth stimulation (0–55% total biomass) by eCO₂ was dependent upon the severity of Pi deficiency and was closely associated with the increased phosphorus utilization efficiency. With the exception of leaf and root biomass, Pi deficiency decreased the biomass partitioning to other plant organs with the maximum decrease observed in seed weight (8–42%) across CO₂ levels. The increased tissue nitrogen (N) concentration in Pi-deficient plants was accredited to the lower biomass and increased nutrient uptake due to the larger root to shoot ratio. The tissue P and N concentration tended to be lower at eCO₂ versus aCO₂ and did not appear to be the main cause of the lack of CO₂ response of growth and P_net under severe Pi deficiency. The leaf N/P ratio of >16 was detrimental to soybean growth. The tissue P concentration needed to attain the maximum productivity for biomass and seed yield tended to be higher at eCO₂ versus aCO₂. Therefore, the eCO₂ is likely to increase the leaf critical P concentration for maximum biomass productivity and yield in soybean.     

Carbon for nutrient exchange between arbuscular mycorrhizal fungi and wheat varies according to cultivar and changes in atmospheric carbon dioxide concentration

Arbuscular mycorrhizal fungi (AMF) form symbioses with most crops, potentially improving their nutrient assimilation and growth. The effects of cultivar and atmospheric CO₂ concentration ([CO₂]) on wheat–AMF carbon‐for‐nutrient exchange remain critical knowledge gaps in the exploitation of AMF for future sustainable agricultural practices within the context of global climate change. We used stable and radioisotope tracers (¹⁵N, ³³P, ¹⁴C) to quantify AMF‐mediated nutrient uptake and fungal acquisition of plant carbon in three wheat (Triticum aestivum L.) cultivars. We grew plants under current ambient (440 ppm) and projected future atmospheric CO₂ concentrations (800 ppm). We found significant ¹⁵N transfer from fungus to plant in all cultivars, and cultivar‐specific differences in total N content. There was a trend for reduced N uptake under elevated atmospheric [CO₂]. Similarly, ³³P uptake via AMF was affected by cultivar and atmospheric [CO₂]. Total P uptake varied significantly among wheat cultivars and was greater at the future than current atmospheric [CO₂]. We found limited evidence of cultivar or atmospheric [CO₂] effects on plant‐fixed carbon transfer to the mycorrhizal fungi. Our results suggest that AMF will continue to provide a route for nutrient uptake by wheat in the future, despite predicted rises in atmospheric [CO₂]. Consideration should therefore be paid to cultivar‐specific AMF receptivity and function in the development of climate smart germplasm for the future.     

Effects of carbon dioxide enrichment on response of mycorrhizal pitch pine (Pinus rigida) to aluminum: growth and mineral nutrition

 Carbon dioxide enrichment may increase the Al tolerance of trees by increasing root growth, root exudation and/or mycorrhizal colonization. The effect of elevated CO₂ on the response of mycorrhizal pitch pine (Pinus rigida Mill.) seedlings to Al was determined in two experiments with different levels of nutrients, 0.1- or 0.2-strength Clark solution. During each experiment, seedlings inoculated with the ectomycorrhizal fungus Pisolithus tinctorius (Pers.) Coker & Couch were grown 13 weeks in sand irrigated with nutrient solution (pH 3.8) containing 0, 6.25, 12.5, or 25 mg/l Al (0, 232, 463, or 927 μM Al) in growth chambers fumigated with 350 (ambient) or 700 (elevated) μl/l CO₂. At ambient CO₂, in the absence of Al, mean total dry weights (DW) of seedlings at the high nutrient level were 164% higher than those at the low level. Total DW at elevated CO₂, in the absence of Al, was significantly greater than that in ambient CO₂ at the low (+34%) and high (+16%) nutrient levels. Root and shoot DW at both nutrient levels decreased with increasing Al concentrations with Al reducing root growth more than shoot growth. Although visible symptoms of Al toxicity in roots and needles were reduced by CO₂ enrichment, there were no significant CO₂ × Al interactions for shoot or root DW. The percentage of seedling roots that became mycorrhizal was negatively related to nutrient level and was greater at elevated than at ambient CO₂ levels. Generally, elevated CO₂ had little effect on concentration of mineral nutrients in roots and needles. Aluminum reduced concentrations of most nutrients by inhibiting uptake. Received: 18 June 1997 / Accepted: 8 December 1997     

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.     

The effects of enhanced ultraviolet-B and nitrogen supply on growth,photosynthesis and nutrient status of <Emphasis Type="Italic">Abies faxoniana</Emphasis> seedlings

Abies faxoniana is a key species in reforestation processes in the southeast of the Qinghai-Tibetan Plateau of China. The changes in growth, photosynthesis and nutrient status of A. faxoniana seedlings exposed to enhanced ultraviolet-B (UV-B), nitrogen supply and their combination were investigated. The experimental design included two levels of UV-B treatments (ambient UV-B, 11.02 KJ m⁻² day⁻¹; enhanced UV-B, 14.33 KJ m⁻² day⁻¹) and two nitrogen levels (0; 20 g N m⁻²). The results indicated that: (1) enhanced UV-B significantly caused a marked decline in growth parameters, net photosynthetic rate (Pn), photosynthetic pigments and F _v/F _m, (2) supplemental nitrogen supply increased the accumulation of total biomass, Pn, photosynthetic pigments and F _v/F _m under ambient UV-B, whereas supplemental nitrogen supply reduced Pn, and not affect biomass under enhanced UV-B, (3) enhanced UV-B or nitrogen supply changed the concentration of nutrient elements of various organs.     

Growth response of Afzelia africana Sm. seedlings to ectomycorrhizal inoculation in a nutrient-deficient soil

 The growth and mineral nutrition responses of seedlings of two provenances of Afzelia africana Sm. from Senegal and Burkina Faso, inoculated with four ectomycorrhizal (ECM) fungi (Scleroderma spp. and an unidentified isolate) from the same regions were assessed in a pot experiment in a savanna ECM-propagule-free soil deficient in NPK. There was little variation in the ability of the different fungal species to colonize roots of either provenance of A. africana or to produce external hyphal in soil. Root colonization by ECM fungi and their hyphal development were not related to mineral nutrition or ECM dependency. Differences in P, N, Mg and Ca concentrations in the leaves of inoculated and non-inoculated Afzelia seedlings were not always associated with production of biomass. Only leaf K concentration increased in both provenances after ECM inoculation. However, the Burkina Faso provenance responded better to inoculation with the two fungal isolates than the Senegal provenance in terms of biomass production. This was due to stimulation of root dry weight of the Burkina Faso provenance. Therefore, the hypothesis arises that non-nutritional rather than nutritional effects explain the contribution of ECM inoculation to the growth of A. africana seedlings. Accepted: 27 April 1999     

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     

Arbuscular Mycorrhizal Fungi Alter Fractal Dimension Characteristics of Robinia pseudoacacia L. Seedlings Through Regulating Plant Growth,Leaf Water Status,Photosynthesis, and Nutrient Concentration Under Drought Stress

The influence of arbuscular mycorrhizal fungi (AMF), Funneliformis mosseae and Rhizophagus intraradices, on plant growth, leaf water status, chlorophyll concentration, photosynthesis, nutrient concentration, and fractal dimension (FD) characteristics of black locust (Robinia pseudoacacia L.) seedlings was studied in pot culture under well-watered, moderate drought stress, and severe drought stress treatments. Mycorrhizal seedlings had higher dry biomass, leaf relative water content (RWC), and water use efficiency (WUE) compared with non-mycorrhizal seedlings. Under all treatments, AMF colonization notably enhanced net photosynthetic rate, stomatal conductance, and transpiration rate, but decreased intercellular CO₂ concentration. Leaf chlorophyll a and total chlorophyll concentrations were higher in AM seedlings than those in non-AM seedlings although there was no significant difference between AMF species. AMF colonization improved leaf C, N, and P concentrations, but decreased C:N, C:P, and N:P ratios. Mycorrhizal seedlings had a larger FD value than non-mycorrhizal seedlings. The FD value was positively and significantly correlated to the plant growth parameters, photosynthesis, RWC, WUE, and nutrient concentration but negatively correlated to leaf/stem ratio, C:N and C:P ratios, and intercellular CO₂ concentration. We conclude that AMF lead to an improvement of growth performance of black locust seedlings under all growth conditions, including drought stress via improving leaf water status, chlorophyll concentration, photosynthesis, and nutrient uptake. Moreover, FD technology proved to be a powerful non-destructive method to characterize the effect of AMF on the physiology of host plants during drought stress.     

Combined effects of CO2 concentration and nutrient status on the biomass production and nutrient uptake of birch seedlings (Betula pendula)

Birch seedlings (Betula pendula) were grown for four months in a greenhouse at three nutrient levels (fertilization of 0, 100 and 500 kg ha^-1 monthy) and at four CO₂ concentrations (350, 700, 1050 and 1400 ppm). The effect of CO₂ concentration on the biomass production depended on the nutrient status. When mineralization of the soil material was the only source of nutrients (0 kg ha^-1), CO₂ enhancement reduced the biomass production slightly, whereas the highest production increase occurred at a fertilization of 100 kg ha^-1, being over 100% between 350 and 700 ppm CO₂. At 500 kg ha^-1 the production increase was smaller, and the production decreased beyond a CO₂ concentration of 700 ppm. The CO₂ concentration had a slight effect on the biomass distribution, the leaves accounting for the highest proportion at the lowest CO₂ concentration (350 ppm). An increase in nutrient status led to a longer growth period and increased the nutrient concentrations in the plants, but the CO₂ concentration had no effect on the growth rhythm and higher CO₂ reduced the nutrient concentrations.     

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.     

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.     

Sensitivity to Cd or Zn of host and symbiont of ectomycorrhizal Pinus sylvestris L. (Scots pine) seedlings

Pinus sylvestris seedlings infected with either the ectomycorrhizal (ECM) fungus Paxillus involutus or Suillus variegatus were exposed to a range of Cd or Zn concentrations. This was done to investigate the relationship between the sensitivity of ECM fungi and their host plants over a wide range of concentrations. P. involutus ameliorated the toxicity of Cd and Zn to P. sylvestris with respect to root length, despite significant inhibition of ECM infection levels by Cd (Cd EC₅₀ [effective concentration which inhibits ECM infection by 50%] values were: P. involutus 3.7 μg g^-1 Cd; S. variegatus 2.3 μg g^-1 Cd). ECM infection by P. involutus also decreased Cd and Zn transport to the plant shoots at potentially toxic concentrations and also influenced the proportion of Zn transported to the roots and shoots, with a higher proportion retained in the roots of the seedlings. ECM infection did increase host biomass production, but this was not affected by the presence of Cd or Zn. Root and shoot biomass production by P. sylvestris, in both the presence and absence of ECM fungi, was unaffected by Cd and Zn at all concentrations tested. This revised version was published online in August 2006 with corrections to the Cover Date.     

Contrasting species responses to continued nitrogen and phosphorus addition in tropical montane forest tree seedlings

Global changes in nutrient deposition rates are likely to have profound effects on plant communities, particularly in the nutrient‐limited systems of the tropics. We studied the effects of increased nutrient availability on the seedlings of six tree species in montane forests of southern Ecuador in situ. After five years of continued N, P, or N+P addition, naturally grown seedlings of each of the two most common species at each elevation (1000, 2000, and 3000 m asl) were harvested for analyses of leaf morphology, nutrient content, herbivory, and tissue biomass allocation. Most species showed increased foliar N and P concentrations after addition of each respective element. Leaf tissue N:P ratios of >20 in the control plants of all species suggest that P is more growth‐limiting in these forests than N. Leaf morphological responses to nutrient addition were species and nutrient specific, with some species (Hedyosmum purparescens, Graffenrieda emarginata) exhibiting increased specific leaf area (SLA), and others (Graffenrieda harlingii) increased leaf area ratios (LAR). Pouteria torta (1000 m) had lower SLA and LAR after P addition. Increased herbivory was only evident in G. emarginata (after N and N+P addition). Only the species from 3000 m asl modified biomass allocation after nutrient addition. In general, N and N+P addition more strongly affected the species studied at the upper elevations, whereas P addition had a similar range of effects on the species at all elevations. We conclude that the responses of the studied tropical montane forest tree seedlings to chronic N and P addition are highly species‐specific and that successful adaptation to increased nutrient availability will depend on species‐specific morphological and physiological plasticity.     

Interactive effects of elevated CO2 and drought stress on leaf water potential and growth in Caragana intermedia

We studied the responses of leaf water potential (Ψ_w), morphology, biomass accumulation and allocation, and canopy productivity index (CPI) to the combined effects of elevated CO₂ and drought stress in Caragana intermedia seedlings. Seedlings were grown at two CO₂ concentrations (350 and 700 μmol mol⁻¹) interacted with three water regimes (60–70%, 45–55%, and 30–40% of field capacity of soil). Elevated CO₂ significantly increased Ψ_w, decreased specific leaf area (SLA) and leaf area ratio (LAR) of drought-stressed seedlings, and increased tree height, basal diameter, shoot biomass, root biomass as well as total biomass under the all the three water regimes. Growth responses to elevated CO₂ were greater in well-watered seedlings than in drought-stressed seedlings. CPI was significantly increased by elevated CO₂, and the increase in CPI became stronger as the level of drought stress increased. There were significant interactions between elevated CO₂ and drought stress on leaf water potential, basal diameter, leaf area, and biomass accumulation. Our results suggest that elevated CO₂ may enhance drought avoidance and improved water relations, thus weakening the effect of drought stress on growth of C. intermedia seedings.