Plant species richness, elevated CO2, and atmospheric nitrogen deposition alter soil microbial community composition and function

We determined soil microbial community composition and function in a field experiment in which plant communities of increasing species richness were exposed to factorial elevated CO₂ and nitrogen (N) deposition treatments. Because elevated CO₂ and N deposition increased plant productivity to a greater extent in more diverse plant assemblages, it is plausible that heterotrophic microbial communities would experience greater substrate availability, potentially increasing microbial activity, and accelerating soil carbon (C) and N cycling. We, therefore, hypothesized that the response of microbial communities to elevated CO₂ and N deposition is contingent on the species richness of plant communities. Microbial community composition was determined by phospholipid fatty acid analysis, and function was measured using the activity of key extracellular enzymes involved in litter decomposition. Higher plant species richness, as a main effect, fostered greater microbial biomass, cellulolytic and chitinolytic capacity, as well as the abundance of saprophytic and arbuscular mycorrhizal (AM) fungi. Moreover, the effect of plant species richness on microbial communities was significantly modified by elevated CO₂ and N deposition. For instance, microbial biomass and fungal abundance increased with greater species richness, but only under combinations of elevated CO₂ and ambient N, or ambient CO₂ and N deposition. Cellobiohydrolase activity increased with higher plant species richness, and this trend was amplified by elevated CO₂. In most cases, the effect of plant species richness remained significant even after accounting for the influence of plant biomass. Taken together, our results demonstrate that plant species richness can directly regulate microbial activity and community composition, and that plant species richness is a significant determinant of microbial response to elevated CO₂ and N deposition. The strong positive effect of plant species richness on cellulolytic capacity and microbial biomass indicate that the rates of soil C cycling may decline with decreasing plant species richness.     

Aphid response to elevated ozone and CO2

Numerous reports indicate that pollution stress caused by sulphur dioxide (SO₂), oxies of nitrogen or fluorides promote aphid growth on herbaceous and woody plants. At SO₂ exposures, the response curve of aphids is bell-shaped having the peak at 100 ppb. This curvilinear response is related to physiological stress responses of host plants exposed to pollutants. On the other hand, observations of aphid performance on ozone-exposed (O₃) or elevated carbon dioxide-exposed (CO₂) plants have given very variable results. Depending on the duration and concentration of O₃ or elevated CO₂ exposure or the age of the exposed plants, aphid growth on the same plants either decreased or increased in comparison to growth on control plants grown in filtered air. The results of these studies suggest that there is no general air pollution-induced plant stress that triggers aphid outbreaks on plants. Plants grown in elevated CO₂ usually have higher C/N ratios than plants grown in current ambient CO₂ atmosphere. A reduced proportion of nitrogen in the plant foliage decreases growth of chewing herbivorous insects, but the few studies of elevated CO₂ effects on sucking insects such as aphids have not yielded similar consistent effects. The present paper reviews recent studies of elevated CO₂ effects on aphids and discusses the effects of combined elevated O₃ and CO₂ exposures on aphid performance on woody plants using pine and birch aphids as examples.     

Contribution of sediment fluxes and transformations to the summer nitrogen budget of an Upper Mississippi River backwater system

Routing nitrate through backwaters of regulated floodplain rivers to increase retention could decrease loading to nitrogen (N)-sensitive coastal regions. Sediment core determinations of N flux were combined with inflow–outflow fluxes to develop mass balance approximations of N uptake and transformations in a flow-controlled backwater of the Upper Mississippi River (USA). Inflow was the dominant nitrate source (>95%) versus nitrification and varied as a function of source water concentration since flow was constant. Nitrate uptake length increased linearly, while uptake velocity decreased linearly, with increasing inflow concentration to 2 mg l⁻¹, indicating limitation of N uptake by loading. N saturation at higher inflow concentration coincided with maximum uptake capacity, 40% uptake efficiency, and an uptake length 2 times greater than the length of the backwater. Nitrate diffusion and denitrification in sediment accounted for 27% of the backwater nitrate retention, indicating that assimilation by other biota or denitrification on other substrates were the dominant uptake mechanisms. Ammonium export from the backwater was driven by diffusive efflux from the sediment. Ammonium increased from near zero at the inflow to a maximum mid-lake, then declined slightly toward the outflow due to uptake during transport. Ammonium export was small compared to nitrate retention. Handling editor: J. Padisak     

大气CO2浓度升高对土壤碳库稳定性的影响

工业革命以来,大气CO₂浓度持续上升,升高的CO₂浓度会改变植物光合产物积累、土壤碳库的碳输入和碳输出过程,进而通过影响有机碳组成和周转特征来调控土壤碳库动态变化。土壤碳库是陆地生态系统碳库的重要组成部分,其碳储量的微小变化都会对大气CO₂浓度和气候变化产生巨大影响。但目前关于CO₂浓度升高对土壤碳库动态和稳定性的影响还不清楚,很大程度上限制了预测陆地生态系统碳循环对气候变化的反馈。系统综述国内外大气CO₂浓度升高对植被生产力、植被碳输入和土壤碳库影响的研究进展,旨在揭示土壤碳库物理、化学组成以及周转特征对CO₂浓度升高的响应过程和机理,探讨CO₂升高情境下土壤微生物特征对土壤碳库稳定性的影响和驱动机制,为深入理解全球变化下的土壤碳循环特征提供理论支撑。     

蒙古栎叶片及其土壤碳、氮同位素自然丰度对大气CO2浓度升高的响应

大气CO₂浓度升高对土壤氮素转化过程产生重要影响,研究其变化有助于更好地预测陆地生态系统的固碳潜力.氮同位素自然丰度作为生态系统氮素循环过程的综合指标能够有效地指示CO₂浓度升高对土壤氮素转化过程的影响.本研究采用开顶箱CO₂ 熏蒸法研究连续10年的大气CO₂ 浓度升高对我国东北地区蒙古栎及其土壤和微生物生物量碳、氮同位素自然丰度的影响.结果表明: 大气CO₂浓度升高改变了土壤氮循环过程,增加了土壤微生物和植物叶片δ¹⁵N;促进了富¹³C土壤有机碳分解,中和了贫¹³C植物光合碳输入的效果,导致土壤可溶性有机碳和微生物碳δ¹³C在CO₂升高条件下没有发生显著变化.这些结果表明,CO₂浓度升高很可能促进了土壤有机质矿化过程,并加剧了系统氮限制的状态.     

Glycine uptake in heath plants and soil microbes responds to elevated temperature, CO2 and drought

Temperate terrestrial ecosystems are currently exposed to climatic and air quality changes with increased atmospheric CO₂, increased temperature and prolonged droughts. The responses of natural ecosystems to these changes are focus for research, due to the potential feedbacks to the climate. We here present results from a field experiment in which the effects of these three climate change factors are investigated solely and in all combinations at a temperate heath dominated by heather (Calluna vulgaris) and wavy hair-grass (Deschampsia flexuosa).Climate induced increases in plant production may increase plant root exudation of dissolved organic compounds such as amino acids, and the release of amino acids during decomposition of organic matter. Such free amino acids in soil serve as substrates for soil microorganisms and are also acquired as nutrients directly by plants. We investigated the magnitude of the response to the potential climate change treatments on uptake of organic nitrogen in an in situ pulse labelling experiment with ¹⁵N¹³C₂-labelled glycine (amino acid) injected into the soil.In situ root nitrogen acquisition by grasses responded significantly to the climate change treatments, with larger ¹⁵N uptake in response to warming and elevated CO₂ but not additively when the treatments were combined. Also, a larger grass leaf biomass in the combined T and CO₂ treatment than in individual treatments suggest that responses to combined climate change factors cannot be predicted from the responses to single factors treatments.The soil microbes were superior to plants in the short-term competition for the added glycine, as indicated by an 18 times larger ¹⁵N recovery in the microbial biomass compared to the plant biomass. The soil microbes acquired glycine largely as an intact compound (87%), with no effects of the multi factorial climate change treatment through one year.     

Growth responses of the perennial legume Sesbania sesban to NH4 and NO3 nutrition and effects on root nodulation

Preference for NH₄⁺ or NO₃⁻ nutrition by the perennial legume Sesbania sesban (L.) Merr. was assessed by supplying plants with NH₄⁺ and NO₃⁻ alone or mixed at equal concentrations (0.5 mM) in hydroponic culture. In addition, growth responses of S. sesban to NH₄⁺ and NO₃⁻ nutrition and the effects on root nodulation and nutrient and mineral composition of the plant tissues were evaluated in a hydroponic setup at a range of external concentration of NH₄⁺ and NO₃⁻ (0, 0.1, 0.2, 0.5, 2 and 5 mM). Seedlings of S. sesban grew equally well when supplied with either NH₄⁺ or NO₃⁻ alone or mixed and had high relative growth rates (RGRs) ranging between 0.19 and 0.21 d⁻¹. When larger plants of S. sesban were supplied with NH₄⁺ or NO₃⁻ alone, the RGRs and shoot elongation rates were not affected by the external concentration of inorganic N. At external N concentrations up to 0.5 mM nodulation occurred and contributed to the N nutrition through fixation of gaseous N₂ from the atmosphere. For both NH₄⁺ and NO₃⁻-fed plants the N concentration in the plant tissues, particularly water-extractable NO₃⁻, increased at high supply concentrations, and concentrations of mineral cations generally decreased. It is concluded that S. sesban can grow without an external inorganic N supply by fixing atmospheric N₂ gas via root nodules. Also, S. sesban grows well on both NH₄⁺ and NO₃⁻ as the external N source and the plant can tolerate relatively high concentrations of NH₄⁺. This wide ecological amplitude concerning N nutrition makes S. sesban very useful as a N₂-fixing fallow crop in N deficient areas and also a candidate species for use in constructed wetland systems for the treatment of NH₄⁺ rich waters.     

Photosynthetic responses to elevated CO2and O3 in field-grown potato(Solanum tuberosum)

Potato plants (Solanum tuberosum L. cv. Bintje) were grown to maturity in open-top chambers under three carbon dioxide (CO₂; ambient and 24 h d⁻¹ seasonal mean concentrations of 550 and 680 μmol mol⁻¹) and two ozone levels (O₃; ambient and an 8 h d⁻¹ seasonal mean of 50 nmol mol⁻¹). Chlorophyll content, photosynthetic characteristics, and stomatal responses were determined to test the hypothesis that elevated atmospheric CO₂ may alleviate the damaging influence of O₃ by reducing uptake by the leaves. Elevated O₃ had no detectable effect on photosynthetic characteristics, leaf conductance, or chlorophyll content, but did reduce SPAD values for leaf 15, the youngest leaf examined. Elevated CO₂ also reduced SPAD values for leaf 15, but not for older leaves; destructive analysis confirmed that chlorophyll content was decreased. Leaf conductance was generally reduced by elevated CO₂, and declined with time in the youngest leaves examined, as did assimilation rate (A). A generally increased under elevated CO₂, particularly in the older leaves during the latter stages of the season, thereby increasing instantaneous transpiration efficiency. Exposure to elevated CO₂ and/or O₃ had no detectable effect on dark-adapted fluorescence, although the values decreased with time. Analysis of the relationships between assimilation rate and intercellular CO₂ concentration and photosynthetically active photon flux density showed there was initially little treatment effect on CO₂-saturated assimilation rates for leaf 15. However, the values for plants grown under 550 μmol mol⁻¹ CO₂ were subsequently greater than in the ambient and 680 μmol mol⁻¹ treatments, although the beneficial influence of the former treatment declined sharply towards the end of the season. Light-saturated assimilation was consistently greater under elevated CO₂, but decreased with time in all treatments. The values decreased sharply when leaves grown under elevated CO₂ were measured under ambient CO₂, but increased when leaves grown under ambient CO₂ were examined under elevated CO₂. The results obtained indicate that, although elevated CO₂ initially increased assimilation and growth, these beneficial effects were not necessarily sustained to maturity as a result of photosynthetic acclimation and the induction of earlier senescence.     

CO2浓度升高和施氮条件下小麦根际呼吸对土壤呼吸的贡献

依托FACE技术平台, 采用稳定¹³C同位素技术, 通过将小麦(C₃作物)种植于长期单作玉米(C₄作物)的土壤上, 研究了大气CO₂浓度升高和不同氮肥水平对土壤排放CO₂的δ¹³C值及根际呼吸的影响. 结果表明: 种植小麦后土壤排放CO₂的δ¹³C值随作物生长逐渐降低, CO₂浓度升高200 μmol·mol^-1显著降低了孕穗、抽穗期(施氮量为250 kg·hm^-2, HN)与拔节、孕穗期(施氮量为150 kg·hm^-2, LN)土壤排放CO₂的δ¹³C值, 显著提高了孕穗、抽穗期的根际呼吸比例. 拔节至成熟期, 根际呼吸占土壤呼吸的比例在高CO₂浓度下为24%～48%(HN)和21%～48%(LN), 在正常CO₂浓度下为20%～36% (HN)和19%～32%(LN). 不同CO₂浓度下土壤排放CO₂的δ¹³C值和根际呼吸对氮肥增加的响应不同, CO₂浓度与氮肥用量在拔节期对根际呼吸的交互效应显著.     

氮素添加和CO2浓度升高对白羊草根际和非根际土壤水溶性有机碳、氮的影响

采用盆栽控制试验对黄土丘陵区白羊草在不同CO₂浓度(400和800 μmol·mol^-1)和施氮水平(0、2.5、5.0 g N·m^-2·a^-1)条件下根际和非根际土壤水溶性有机碳(DOC)和水溶性有机氮(DON)的变化特征进行研究.结果表明: CO₂浓度升高对白羊草根际和非根际土壤DOC、水溶性总氮(DTN)、DON、水溶性铵态氮(NH₄⁺-N)、水溶性硝态氮(NO₃^--N)含量均无显著影响.施氮显著提高了根际和非根际土壤DTN、NO₃^--N含量和根际土壤DON含量,显著降低了根际土壤DOC/DON.在各处理条件下,根际土壤DTN、NO₃^--N和DON含量均显著低于非根际土壤,根际土壤DOC/DON显著高于非根际土壤.短期CO₂浓度升高对黄土丘陵区土壤水溶性有机碳、氮含量无显著影响,而氮沉降的增加在一定程度上改善了土壤中水溶性氮素缺乏的状况,但并不足以满足植被对水溶性氮素的需求.     

Laboratory incubations reveal potential responses of soil nitrogen cycling to changes in soil C and N availability in Mojave Desert soils exposed to elevated atmospheric CO2

Elevated atmospheric carbon dioxide (CO₂) has the potential to alter soil carbon (C) and nitrogen (N) cycling in arid ecosystems through changes in net primary productivity. However, an associated feedback exists because any sustained increases in plant productivity will depend upon the continued availability of soil N. We took soils from under the canopies of major shrubs, grasses, and plant interspaces in a Mojave Desert ecosystem exposed to elevated atmospheric CO₂ and incubated them in the laboratory with amendments of labile C and N to determine if elevated CO₂ altered the mechanistic controls of soil C and N on microbial N cycling. Net ammonification increased under shrubs exposed to elevated CO₂, while net nitrification decreased. Elevated CO₂ treatments exhibited greater fluxes of N₂O–N under Lycium spp., but not other microsites. The proportion of microbial/extractable organic N increased under shrubs exposed to elevated CO₂. Heterotrophic N₂‐fixation and C mineralization increased with C addition, while denitrification enzyme activity and N₂O–N fluxes increased when C and N were added in combination. Laboratory results demonstrated the potential for elevated CO₂ to affect soil N cycling under shrubs and supports the hypothesis that energy limited microbes may increase net inorganic N cycling rates as the amount of soil‐available C increases under elevated CO₂. The effect of CO₂ enrichment on N‐cycling processes is mediated by its effect on the plants, particularly shrubs. The potential for elevated atmospheric CO₂ to lead to accumulation of NH₄⁺ under shrubs and the subsequent volatilization of NH₃ may result in greater losses of N from this system, leading to changes in the form and amount of plant‐available inorganic N. This introduces the potential for a negative feedback mechanism that could act to constrain the degree to which plants can increase productivity in the face of elevated atmospheric CO₂.     

Effects of elevated atmospheric CO2 on soil enzyme activities at different nitrogen application treatments

It has been predicted that elevated atmospheric CO₂ will increase enzyme activity as a result of CO₂-induced carbon entering the soil. The objective of this study was to investigate the effects of elevated atmospheric CO₂ on soil enzyme activities under a rice/wheat rotation. This experiment was conducted in Wuxi, Jiangsu, China as part of the China FACE (Free Air Carbon Dioxide Enrichment) Project. Two atmospheric CO₂ concentrations (580±60) and (380±40) μmol·mol^-1) and three N application treatments (low-150, normal-250 and high-350 kg N·hm^-2) were included. Soil samples (0-10 cm) were collected for analysis of β-glucosidase, invertase, urease, acid phosphates and β-glucosaminidase activities. The results revealed that with elevated atmospheric CO₂ β-glucosidase activity significantly decreased (P < 0.05) at low N application rates; had no significant effect with a normal N application rate; and significantly increased (P < 0.05) with a high N application rate. For urease activity, at low and normal N application rates (but not high N application rate), elevated atmospheric CO₂ significantly increased (P < 0.05) it. With acid phosphatase elevated atmospheric CO₂ only had significant higher effects (P < 0.05) at high N application rates. Under different CO₂ concentration, effects of N fertilization are also different. Soil β-glucosidase activity at ambient CO₂ concentration decreased with N fertilization, while it increased at elevated CO₂ concentration. In addition, invertase and acid phosphatase activities at elevated CO₂ concentration, significantly increased (P < 0.05) with N treatments, but there was no effect with the ambient CO₂ concentration. For urease activity, at ambient CO₂ concentration, N fertilization increased it significantly (P < 0.05), whereas at elevated CO₂ concentration it was not significant. Additionally, with β-glucosaminidase activity, there were no significant effects from N application. In general, then, elevated atmospheric CO₂ increased soil enzyme activity, which may be attributed to the following two factors: (1) elevated atmospheric CO₂ led to more plant biomass in the soil, which in turn stimulated soil microbial biomass and activity; and (2) elevated atmospheric CO₂ increased plant photosynthesis, thereby increasing plant-derived soil enzymes.     

Effects of elevated atmospheric CO2 on soil enzyme activities at different nitrogen application treatments

It has been predicted that elevated atmospheric CO₂ will increase enzyme activity as a result of CO₂-induced carbon entering the soil. The objective of this study was to investigate the effects of elevated atmospheric CO₂ on soil enzyme activities under a rice/wheat rotation. This experiment was conducted in Wuxi, Jiangsu, China as part of the China FACE (Free Air Carbon Dioxide Enrichment) Project. Two atmospheric CO₂ concentrations (580±60) and (380±40) μmol·mol^-1) and three N application treatments (low-150, normal-250 and high-350 kg N·hm^-2) were included. Soil samples (0-10 cm) were collected for analysis of β-glucosidase, invertase, urease, acid phosphates and β-glucosaminidase activities. The results revealed that with elevated atmospheric CO₂ β-glucosidase activity significantly decreased (P < 0.05) at low N application rates; had no significant effect with a normal N application rate; and significantly increased (P < 0.05) with a high N application rate. For urease activity, at low and normal N application rates (but not high N application rate), elevated atmospheric CO₂ significantly increased (P < 0.05) it. With acid phosphatase elevated atmospheric CO₂ only had significant higher effects (P < 0.05) at high N application rates. Under different CO₂ concentration, effects of N fertilization are also different. Soil β-glucosidase activity at ambient CO₂ concentration decreased with N fertilization, while it increased at elevated CO₂ concentration. In addition, invertase and acid phosphatase activities at elevated CO₂ concentration, significantly increased (P < 0.05) with N treatments, but there was no effect with the ambient CO₂ concentration. For urease activity, at ambient CO₂ concentration, N fertilization increased it significantly (P < 0.05), whereas at elevated CO₂ concentration it was not significant. Additionally, with β-glucosaminidase activity, there were no significant effects from N application. In general, then, elevated atmospheric CO₂ increased soil enzyme activity, which may be attributed to the following two factors: (1) elevated atmospheric CO₂ led to more plant biomass in the soil, which in turn stimulated soil microbial biomass and activity; and (2) elevated atmospheric CO₂ increased plant photosynthesis, thereby increasing plant-derived soil enzymes.     

Response of coastal phytoplankton to ammonium and nitrate pulses: seasonal variations of nitrogen uptake and regeneration

Seasonal variation in uptake and regeneration of ammonium and nitrate in a coastal lagoon was studied using ¹⁵N incorporation in particulate matter and by measuring changes in particulate nitrogen. Uptake and regeneration rates were two orders of magnitude lower in winter than in summer. Summer uptake values were 2.8 and 2.2 mol N.l^–1.d^–1 for ammonium and nitrate, respectively. Regeneration rates were 2.9 and 2.1 mol N.l^–1.d^–1 for ammonium and nitrate respectively. Regeneration/uptake ratios were often below one, indicating that water column processes were not sufficient to satisfy the phytoplankton nitrogen demand. This implies a role of other sources of nitrogen, such as macrofauna (oysters and epibionts) and sediment. Phytoplankton was well adapted to the seasonal variations in resources, with mixotrophic dinoflagellates dominant in winter, and fast growing diatoms in summer. In winter and spring, ammonium was clearly preferred to nitrate as a nitrogen source, but nitrate was an important nitrogen source in summer because of high nitrification rates. Despite low nutrient levels, the high rates of nitrogen regeneration in summer as well as the simultaneous uptake of nitrate and ammonium allow high phytoplankton growth rates which in turn enable high oyster production.     

Intergenerational above- and belowground responses of spring wheat (Triticum aestivum L.) to elevated CO2

We quantified intergenerational above- and belowground responses of two genotypes of semi-dwarf, hard red, spring wheats (Triticum aestivum L.) to elevated (700 μmol mol⁻¹) CO₂. These plants were progeny of seeds produced from previous generation plants grown at elevated CO₂ under well-watered and high nutrient conditions. Because neither genotype in the first generation exhibited enhanced performance with CO₂ enrichment, our objective in this investigation was to assess if exposure to CO₂ enrichment in subsequent generations resulted in temporal changes in the relative enhancement (elevated/ambient) of above- and belowground plant growth. Relative enhancement occurred in both the second and third generations for both above- and belowground variables. Above- and belowground variables were enhanced by similar relative amounts at elevated CO₂ within a generation at each harvest date. Relative enhancement of measured variables was generally greater in the third than second generation when plants were in the seedling or vegetative stage, but not when plants were reproductive. Additional research is needed to investigate physiological or other limitations of translating above- and belowground responses to CO₂ in vegetative growth stages to reproductive performance. Intergenerational above- and belowground responses of this C₃ annual plant to CO₂ enrichment are not driven by genetic change (selection) that occurred between generations, but rather CO₂-induced changes in seeds that affected seedling responses to CO₂ enrichment. Wir quantifizierten die intergenerationelle ober- und unterirdische Reaktionen von zwei Genotypen mittellangen, hartroten Winterweizen (Triticum aestivum L.) auf erhöhtes CO₂ (700 μmol mol⁻¹). Diese Pflanzen waren Abkömmlinge von Samen, die von Pflanzen der vorherigen Generation produziert wurden, welche ihrerseits bei erhöhtem CO₂ und bei ausreichender Wasserversorgung sowie guten Nährstoffbedingungen kultiviert wurden. Weil keiner der beiden Genotypen in der ersten Generation eine verbesserte Leistung bei CO₂-Anreicherung zeigte, war unser Ziel, in der Untersuchung abzuschätzen, ob die Exposition einer CO₂-Anreicherung in den nachfolgenden Generationen zu temporären Veränderungen in der relativen Förderung (erhöht/umgebend) des ober- und unterirdischen Wachstums führte. Eine relative Steigerung fand in der zweiten und in der dritten Generation sowohl bei den ober- als auch unterirdischen Variablen statt. Bei jedem Erntetermin waren die ober- und unterirdischen Variablen innerhalb einer Generation bei erhöhtem CO₂ mit ähnlichen relativen Anteilen positiv beeinflusst. Die relative Steigerung der gemessenen Variablen war im Allgemeinen bei Pflanzen im Keimlings- oder vegetativen Stadium in der dritten Generation größer als in der zweiten, jedoch nicht bei reproduktiven Pflanzen. Zusätzliche Forschung ist notwendig, um physiologische oder andere Limitierungen zu untersuchen, die ober- und unterirdische Reaktionen von vegetativen Wachstumsstadien auf CO₂ in die reproduktiven Leistung übersetzen. Intergenerationelle, ober- und unterirdische Reaktionen dieser C₃-Pflanze auf CO₂-Anreicherung werden nicht durch genetische Veränderungen (Selektion) im Laufe der Generationen gesteuert, sondern eher durch CO₂-induzierte Veränderungen in den Samen, welche die Reaktion der Keimlinge auf eine CO₂-Anreicherung beeinflussen.     

Involvement of nitrogen and cytokinins in photosynthetic acclimation to elevated CO2 of spring wheat

Acclimation of photosynthetic capacity to elevated CO₂ involves a decrease of the leaf Rubisco content. In the present study, it was hypothesized that nitrogen uptake and partitioning within the leaf and among different aboveground organs affects the down-regulation of Rubisco. Given the interdependence of nitrogen and cytokinin signals at the whole plant level, it is also proposed that cytokinins affect the nitrogen economy of plants under elevated CO₂, and therefore the acclimatory responses. Spring wheat received varying levels of nitrogen and cytokinin in field chambers with ambient (370 μmol mol⁻¹) or elevated (700 μmol mol⁻¹) atmospheric CO₂. Gas exchange, Rubisco, soluble protein and nitrogen contents were determined in the top three leaves in the canopy, together with total nitrogen contents per shoot. Growth in elevated CO₂ induced decreases in photosynthetic capacity only when nitrogen supply was low. However, the leaf contents of Rubisco, soluble protein and total nitrogen on an area basis declined in elevated CO₂ regardless of nitrogen supply. Total nitrogen in the shoot was no lower in elevated than ambient CO₂, but the fraction of this nitrogen located in flag and penultimate leaves was lower in elevated CO₂. Decreased Rubisco: chlorophyll ratios accompanied losses of leaf Rubisco with CO₂ enrichment. Cytokinin applications increased nitrogen content in all leaves and nitrogen allocation to senescing leaves, but decreased Rubisco contents in flag leaves at anthesis and in all leaves 20 days later, together with the amount of Rubisco relative to soluble protein in all leaves at both growth stages. The results suggest that down regulation of Rubisco in leaves at elevated CO₂ is linked with decreased allocation of nitrogen to the younger leaves and that cytokinins cause a fractional decrease of Rubisco and therefore do not alleviate acclimation to elevated CO₂.