Leaf litter production and decomposition in a poplar short-rotation coppice exposed to free air CO2 enrichment (POPFACE)

The capacity of forest ecosystems to sequester C in the soil relies on the net balance between litter production above, as well as, below ground, and decomposition processes. Nitrogen mineralization and its availability for plant growth and microbial activity often control the speed of both processes. Litter production, decomposition and N mineralization are strongly interdependent. Thus, their responses to global environmental changes (i.e. elevated CO₂, climate, N deposition, etc.) cannot be fully understood if they are studied in isolation. In the present experiment, we investigated litter fall, litter decomposition and N dynamics in decomposing litter of three Populus spp., in the second and third growing season of a short rotation coppice under FACE. Elevated CO₂ did not affect annual litter production but slightly retarded litter fall in the third growing season. In all species, elevated CO₂ lowered N concentration, resulting in a reduction of N input to the soil via litter fall, but did not affect lignin concentrations. Litter decomposition was studied in bags incubated in situ both in control and FACE plots. Litter lost between 15% and 18% of the original mass during the eight months of field incubation. On average, litter produced under elevated CO₂ attained higher residual mass than control litter. On the other end, when litter was incubated in FACE plots it exhibited higher decay rates. These responses were strongly species‐specific. All litter increased their N content during decomposition, indicating immobilization of N from external sources. Independent of the initial quality, litter incubated on FACE soils immobilized less N, possibly as a result of lower N availability in the soil. Indeed, our results refer to a short‐term decomposition experiment. However, according to a longer‐term model extrapolation of our results, we anticipate that in Mediterranean climate, under elevated atmospheric CO₂, soil organic C pool of forest ecosystems may initially display faster turnover, but soil N availability will eventually limit the process.     

冬季土壤呼吸：不可忽视的地气CO2交换过程

 冬季土壤呼吸是生态系统释放CO₂的极为重要的组成部分,并显著地影响着碳收支。然而,过去绝大多数工作集中在生长季节土壤呼吸的测定,对年土壤呼吸量的估算大多基于冬季土壤呼吸为零的假设。目前为数不多的研究集中在极地苔原和亚高山,其它植被类型的研究只有零星报道。极地苔原和森林冬季土壤呼吸速率分别为0.002～1.359和0.22～0.67 μmol C．m^-2·s^-1;土壤呼吸的CO2释放量分别为0.55～26.37和22.4～152.0 g C·m^－2,是地气CO₂交换过程中不可忽视的环节。雪是土壤呼吸过程的重要调节者,积雪厚度和覆盖时间的长短均会影响土壤呼吸的强弱;水分的可获取性是重要的限制因素;对于维持活跃的土壤呼吸有一个关键的土壤温度临界值（-7～-5 ℃）,低于这个值会因自由水的缺乏而抑制异养微生物的呼吸。如果存在绝缘的积雪层,可溶性碳底物在自由水存在的情况下可控制异养微生物的活力。该文对冬季土壤呼吸的重要性、研究方法、土壤呼吸强度及其影响机制等进行了综述,并讨论了冬季土壤呼吸研究中存在的问题及未来研究方向。     

Elevated CO2 and O3 Alter Soil Nitrogen Transformations beneath Trembling Aspen, Paper Birch, and Sugar Maple

Nitrogen cycling in northern temperate forest ecosystems could change under increasing atmospheric CO₂ and tropospheric O₃ as a result of quantitative and qualitative changes in plant litter production. At the Aspen Free Air CO₂–O₃ Enrichment (FACE) experiment, we previously found that greater substrate inputs to soil under elevated CO₂ did not alter gross N transformation rates in the first 3 years of the experiment. We hypothesized that greater litter production under elevated CO₂ would eventually cause greater gross N transformation rates and that CO₂ effects would be nullified by elevated O₃. Following our original study, we continued measurement of gross N transformation rates for an additional four years. From 1999 to 2003, gross N mineralization doubled, N immobilization increased 4-fold, but changes in microbial biomass N and soil total N were not detected. We observed year-to-year variation in N transformation rates, which peaked during a period of foliar insect damage. Elevated CO₂ caused equivalent increases in gross rates of N mineralization (+34%) and NH ₄ ⁺ immobilization (+36%). These results indicate greater rates of N turnover under elevated CO₂, but do not indicate a negative feedback between elevated CO₂ and soil N availability. Elevated O₃ decreased gross N mineralization (−16%) and had no effect on NH ₄ ⁺ immobilization, indicating reduced N availability under elevated O₃. The effects of CO₂ and O₃ on N mineralization rates were mainly related to changes in litter production, whereas effects on N immobilization were likely influenced by changes in litter chemistry and production. Our findings also indicate that concomitant increases in atmospheric CO₂ and O₃ could lead to a negative feedback on N availability.     

Forest Productivity and Efficiency of Resource Use Across a Chronosequence of Tropical Montane Soils

We tested the hypothesis that plants adjust to nutrient availability by altering carbon allocation patterns and nutrient-use efficiency (NUE = net primary production [NPP] per unit nutrient uptake), but are constrained by a trade-off between NUE and light-use efficiency () = NPP per unit intercepted light). NPP, NUE and ) were measured in montane Metrosideros polymorpha forest across a 4.1 x 10⁶ yr space for time substitution chronosequence in which available soil N and P pools change with site age. Although the range of N and P availability across sites was broad, there was little difference in NPP between sites, and in contrast to theories of carbon allocation relative to limiting resources, we found no consistent relationships in production allocation to leaves, fine roots or wood. However, canopy nutrient pools and fluxes were correlated with the mass of fine roots per unit soil volume and there was a weak but positive correlation of NPP with LAI. Patterns of ) and NUE across the soil developmental sequence were opposite to each other. ) increased as nutrient availability and nutrient turnover increased, while NUE decreased in response to the same influences but reached its highest values where either N or P availability and turnover of both N and P were low. A negative correlation between ) and NUE supports the hypothesis that a trade-off exists between ) and leaf characteristics affecting NUE.     

Soil–Nitrogen Cycling in a Pine Forest Exposed to 5 Years of Elevated Carbon Dioxide

Empirical and modeling studies have shown that the magnitude and duration of the primary production response to elevated carbon dioxide (CO₂) can be constrained by limiting supplies of soil nitrogen (N). We have studied the response of a southern US pine forest to elevated CO₂ for 5 years (1997–2001). Net primary production has increased significantly under elevated CO₂. We hypothesized that the increase in carbon (C) fluxes to the microbial community under elevated CO₂ would increase the rate of N immobilization over mineralization. We tested this hypothesis by quantifying the pool sizes and fluxes of inorganic and organic N in the forest floor and top 30 cm of mineral soil during the first 5 years of CO₂ fumigation. We observed no statistically significant change in the gross or net rate of inorganic N mineralization and immobilization in any soil horizon under elevated CO₂. Similarly, elevated CO₂ had no statistically significant effect on the concentration or flux of organic N, including amino acids. Microbial biomass N was not significantly different between CO₂ treatments. Thus, we reject our hypothesis that elevated CO₂ increases the rate of N immobilization. The quantity and chemistry of the litter inputs to the forest floor and mineral soil horizons can explain the limited range of microbially mediated soil–N cycling responses observed in this ecosystem. Nevertheless a comparative analysis of ecosystem development at this site and other loblolly pine forests suggests that rapid stand development and C sequestration under elevated CO₂ may be possible only in the early stages of stand development, prior to the onset of acute N limitation.     

Elevated CO2 influences nutrient availability in young beech-spruce communities on two soil types

The objectives of this study were to investigate how different soil types and elevated N deposition (0.7 vs 7 g N m^-2a^-1) influence the effects of elevated CO₂ (370 vs 570 µmol CO₂ mol^-1) on soil nutrients and net accumulation of N, P, K, S, Ca, Mg, Fe, Mn, and Zn in spruce (Picea abies) and beech (Fagus sylvatica). Model ecosystems were established in large open-top chambers on two different forest soils: a nutrient-poor acidic loam and a nutrient-rich calcareous sand. The response of net nutrient accumulation to elevated atmospheric CO₂ depended upon soil type (interaction soil 2 CO₂, P<0.05 for N, P, K, S, Ca, Mg, Zn) and differed between spruce and beech. On the acidic loam, CO₂ enrichment suppressed net accumulation of all nutrients in beech (P<0.05 for P, S, Zn), but stimulated it for spruce (P<0.05 for Fe, Zn) On the nutrient-rich calcareous sand, increased atmospheric CO₂ enhanced nutrient accumulation in both species significantly. Increasing the N deposition did not influence the CO₂ effects on net nutrient accumulation with either soil. Under elevated atmospheric CO₂, the accumulation of N declined relative to other nutrients, as indicated by decreasing ratios of N to other nutrients in tree biomass (all ratios: P<0.001, except the N to S ratio). In both the soil and soil solution, elevated CO₂ did not influence concentrations of base cations and available P. Under CO₂ enrichment, concentrations of exchangeable NH₄⁺ decreased by 22% in the acidic loam and increased by 50% in the calcareous sand (soil 2 CO₂, P<0.001). NO₃^- concentrations decreased by 10-70% at elevated CO₂ in both soils (P<0.01).     

Combined effects of atmospheric CO<Subscript>2</Subscript> and N availability on the belowground carbon and nitrogen dynamics of aspen mesocosms

It is uncertain whether elevated atmospheric CO₂ will increase C storage in terrestrial ecosystems without concomitant increases in plant access to N. Elevated CO₂ may alter microbial activities that regulate soil N availability by changing the amount or composition of organic substrates produced by roots. Our objective was to determine the potential for elevated CO₂ to change N availability in an experimental plant-soil system by affecting the acquisition of root-derived C by soil microbes. We grew Populus tremuloides (trembling aspen) cuttings for 2 years under two levels of atmospheric CO₂ (36.7 and 71.5 Pa) and at two levels of soil N (210 and 970 μg N g^–1). Ambient and twice-ambient CO₂ concentrations were applied using open-top chambers, and soil N availability was manipulated by mixing soils differing in organic N content. From June to October of the second growing season, we measured midday rates of soil respiration. In August, we pulse-labeled plants with ¹⁴CO₂ and measured soil ¹⁴CO₂ respiration and the ¹⁴C contents of plants, soils, and microorganisms after a 6-day chase period. In conjunction with the August radio-labeling and again in October, we used ¹⁵N pool dilution techniques to measure in situ rates of gross N mineralization, N immobilization by microbes, and plant N uptake. At both levels of soil N availability, elevated CO₂ significantly increased whole-plant and root biomass, and marginally increased whole-plant N capital. Significant increases in soil respiration were closely linked to increases in root biomass under elevated CO₂. CO₂ enrichment had no significant effect on the allometric distribution of biomass or ¹⁴C among plant components, total ¹⁴C allocation belowground, or cumulative (6-day) ¹⁴CO₂ soil respiration. Elevated CO₂ significantly increased microbial ¹⁴C contents, indicating greater availability of microbial substrates derived from roots. The near doubling of microbial ¹⁴C contents at elevated CO₂ was a relatively small quantitative change in the belowground C cycle of our experimental system, but represents an ecologically significant effect on the dynamics of microbial growth. Rates of plant N uptake during both 6-day periods in August and October were significantly greater at elevated CO₂, and were closely related to fine-root biomass. Gross N mineralization was not affected by elevated CO₂. Despite significantly greater rates of N immobilization under elevated CO₂, standing pools of microbial N were not affected by elevated CO₂, suggesting that N was cycling through microbes more rapidly. Our results contained elements of both positive and negative feedback hypotheses, and may be most relevant to young, aggrading ecosystems, where soil resources are not yet fully exploited by plant roots. If the turnover of microbial N increases, higher rates of N immobilization may not decrease N availability to plants under elevated CO₂. Received: 12 February 1999 / Accepted: 2 March 2000     

Elevated carbon dioxide and ozone alter productivity and ecosystem carbon content in northern temperate forests

Three young northern temperate forest communities in the north‐central United States were exposed to factorial combinations of elevated carbon dioxide (CO₂) and tropospheric ozone (O₃) for 11 years. Here, we report results from an extensive sampling of plant biomass and soil conducted at the conclusion of the experiment that enabled us to estimate ecosystem carbon (C) content and cumulative net primary productivity (NPP). Elevated CO₂ enhanced ecosystem C content by 11%, whereas elevated O₃ decreased ecosystem C content by 9%. There was little variation in treatment effects on C content across communities and no meaningful interactions between CO₂ and O₃. Treatment effects on ecosystem C content resulted primarily from changes in the near‐surface mineral soil and tree C, particularly differences in woody tissues. Excluding the mineral soil, cumulative NPP was a strong predictor of ecosystem C content (r² = 0.96). Elevated CO₂ enhanced cumulative NPP by 39%, a consequence of a 28% increase in canopy nitrogen (N) content (g N m^?2) and a 28% increase in N productivity (NPP/canopy N). In contrast, elevated O₃ lowered NPP by 10% because of a 21% decrease in canopy N, but did not impact N productivity. Consequently, as the marginal impact of canopy N on NPP (?NPP/?N) decreased through time with further canopy development, the O₃ effect on NPP dissipated. Within the mineral soil, there was less C in the top 0.1 m of soil under elevated O₃ and less soil C from 0.1 to 0.2 m in depth under elevated CO₂. Overall, these results suggest that elevated CO₂ may create a sustained increase in NPP, whereas the long‐term effect of elevated O₃ on NPP will be smaller than expected. However, changes in soil C are not well‐understood and limit our ability to predict changes in ecosystem C content.     

Mechanical properties of spruce and beech wood grown in elevated CO2

Biomechanical responses of stems of 6- to 7-year-old spruce [Picea abies (L.) Karst.] and beech (Fagus sylvatica L) trees were studied after 4 years of growth in elevated atmospheric CO₂ in combination with a nitrogen treatment and on two different soil types. At the end of the treatment, stems were harvested and tested in fresh and air-dried status. Bending characteristics of juvenile wood (modulus of elasticity, termed rigidity) were determined by bending tests. Fracture characteristics (termed toughness) were determined by stroke-pendulum tests. From the base disk of each stem densitometric data were obtained. In spruce, wood produced under elevated CO₂ was tougher on both soil types; enhanced N deposition made wood less rigid only on acidic soils. In contrast, beech wood samples showed no significant reaction to CO₂ but were significantly tougher under high nitrogen depositions on acidic soil. Effects on wood density of both CO₂ and N treatments were not significant, but wood density was higher on acidic soil and so were rigidity and toughness (soil effect). Different genotypes of spruce and beech reacted significantly differently to the treatments. Some genotypes reacted strongly to CO₂ or N, whereas others did not react or showed interactions between CO₂ and N. This underlines the importance of genetic diversity in tree communities.     

Assessment of soil nitrogen and phosphorous availability under elevated CO2 and N-fertilization in a short rotation poplar plantation

Photosynthetic stimulation by elevated [CO₂] is largely regulated by nitrogen and phosphorus availability in the soil. During a 6 year Free Air CO₂ Enrichment (FACE) experiment with poplar trees in two short rotations, inorganic forms of soil nitrogen, extractable phosphorus, microbial and total nitrogen were assessed. Moreover, in situ and potential nitrogen mineralization, as well as enzymatic activities, were determined as measures of nutrient cycling. The aim of this study was to evaluate the effects of elevated [CO₂] and fertilization on: (1) N mineralization and immobilization processes; (2) soil nutrient availability; and (3) soil enzyme activity, as an indication of microbial and plant nutrient acquisition activity. Independent of any treatment, total soil N increased by 23% in the plantation after 6 years due to afforestation. Nitrification was the main process influencing inorganic N availability in soil, while ammonification being null or even negative. Ammonium was mostly affected by microbial immobilization and positively related to total N and microbial biomass N. Elevated [CO₂] negatively influenced nitrification under unfertilised treatment by 44% and consequently nitrate availability by 30% on average. Microbial N immobilization was stimulated by [CO₂] enrichment and probably enhanced the transformation of large amounts of N into organic forms less accessible to plants. The significant enhancement of enzyme activities under elevated [CO₂] reflected an increase in nutrient acquisition activity in the soil, as well as an increase of fungal population. Nitrogen fertilization did not influence N availability and cycling, but acted as a negative feed-back on phosphorus availability under elevated CO₂.     

Net mineralization of N at deeper soil depths as a potential mechanism for sustained forest production under elevated [CO2]

Elevated atmospheric carbon dioxide concentrations [CO₂] is projected to increase forest production, which could increase ecosystem carbon (C) storage. This study contributes to our broad goal of understanding the causes and consequences of increased fine‐root production and mortality under elevated [CO₂] by examining potential gross nitrogen (N) cycling rates throughout the soil profile. Our study was conducted in a CO₂‐enriched sweetgum (Liquidambar styraciflua L.) plantation in Oak Ridge, TN, USA. We used ¹⁵N isotope pool dilution methodology to measure potential gross N cycling rates in laboratory incubations of soil from four depth increments to 60 cm. Our objectives were twofold: (1) to determine whether N is available for root acquisition in deeper soil and (2) to determine whether elevated [CO₂], which has increased inputs of labile C resulting from greater fine‐root mortality at depth, has altered N cycling rates. Although gross N fluxes declined with soil depth, we found that N is potentially available for roots to access, especially below 15 cm depth where rates of microbial consumption of mineral N were reduced relative to production. Overall, up to 60% of potential gross N mineralization and 100% of potential net N mineralization occurred below 15 cm depth at this site. This finding was supported by in situ measurements from ion‐exchange resins, where total inorganic N availability at 55 cm depth was equal to or greater than N availability at 15 cm depth. While it is likely that trees grown under elevated [CO₂] are accessing a larger pool of inorganic N by mining deeper soil, we found no effect of elevated [CO₂] on potential gross or net N cycling rates. Thus, increased root exploration of the soil volume under elevated [CO₂] may be more important than changes in potential gross N cycling rates in sustaining forest responses to rising atmospheric CO₂.     

Effects of grassland management on plant nitrogen use efficiency (NUE): Evidence from a long-term experiment

Grassland management intensification can greatly influence nitrogen (N) dynamics between aboveground and belowground compartments mainly because of the large amount of available N forms, which are repeatedly added to soils. A better understanding of how chronic fertilisation might affect N use efficiency (NUE) in plants can contribute to reducing N losses from soils and improve the sustainability of managed grasslands. Here we address how NUE might be affected by (1) the addition of key nutrients (e.g. N, P, K, Mg) in different combinations, (2) grazing by rabbits, and (3) liming (i.e. CaCO₃ applications) in a 22-year-old permanent grassland experiment established in Berkshire, UK, in 1991. We first calculate seven different NUE indexes, which are known to respond either to changes in soil N availability (i.e. endogenous N inputs from soil N mineralization processes) or to exogenous N inputs (i.e. synthetic N fertiliser). We found that plant NUE calculated as plant biomass produced per unit of N acquired significantly decreased under the chronic addition of multiple nutrients (NPKMg) and was even lower under N-only applications. Most NUE indexes significantly decreased under grazing but greatly increased under liming applications. We found evidence that NUE indexes, which accounted for endogenous N sources decreased at increased rates of soil N mineralization. Finally, we found no significant relationships between any of the NUE indexes and estimates of soil N losses (Mg N ha⁻¹) or N retention in soils (i.e. units of soil N retained per unit of N added) calculated from changes in net soil N budget over 22 years. Our study carried out on relatively acidic sandy soils suggests how liming applications in combination with low levels of multi-nutrient additions (NPKMg) can significantly improve plant biomass production per unit of N added thus contributing to enhance the sustainability of managed grassland ecosystems.     

Soil carbon sequestration in a pine forest after 9 years of atmospheric CO2 enrichment

The impact of anthropogenic CO₂ emissions on climate change may be mitigated in part by C sequestration in terrestrial ecosystems as rising atmospheric CO₂ concentrations stimulate primary productivity and ecosystem C storage. Carbon will be sequestered in forest soils if organic matter inputs to soil profiles increase without a matching increase in decomposition or leaching losses from the soil profile, or if the rate of decomposition decreases because of increased production of resistant humic substances or greater physical protection of organic matter in soil aggregates. To examine the response of a forest ecosystem to elevated atmospheric CO₂ concentrations, the Duke Forest Free‐Air CO₂ Enrichment (FACE) experiment in North Carolina, USA, has maintained atmospheric CO₂ concentrations 200 μL L^?1 above ambient in an aggrading loblolly pine (Pinus taeda) plantation over a 9‐year period (1996–2005). During the first 6 years of the experiment, forest‐floor C and N pools increased linearly under both elevated and ambient CO₂ conditions, with significantly greater accumulations under the elevated CO₂ treatment. Between the sixth and ninth year, forest‐floor organic matter accumulation stabilized and C and N pools appeared to reach their respective steady states. An additional C sink of ～30 g C m^?2 yr^?1 was sequestered in the forest floor of the elevated CO₂ treatment plots relative to the control plots maintained at ambient CO₂ owing to increased litterfall and root turnover during the first 9 years of the study. Because we did not detect any significant elevated CO₂ effects on the rate of decomposition or on the chemical composition of forest‐floor organic matter, this additional C sink was likely related to enhanced litterfall C inputs. We also failed to detect any statistically significant treatment effects on the C and N pools of surface and deep mineral soil horizons. However, a significant widening of the C : N ratio of soil organic matter (SOM) in the upper mineral soil under both elevated and ambient CO₂ suggests that N is being transferred from soil to plants in this aggrading forest. A significant treatment × time interaction indicates that N is being transferred at a higher rate under elevated CO₂ (P=0.037), suggesting that enhanced rates of SOM decomposition are increasing mineralization and uptake to provide the extra N required to support the observed increase in primary productivity under elevated CO₂.     

易分解有机碳对不同恢复年限森林土壤激发效应的影响

土壤有机碳库作为陆地生态系统最大的碳库,其微小的改变都将引起大气CO_2浓度的急剧改变。易分解有机碳的输入可以通过正/负激发效应加快/减缓土壤有机碳(SOC)的矿化,并最终影响土壤碳平衡。以长汀县不同恢复年限森林(裸地、5年、15年、30年马尾松林以及天然林)土壤为研究对象,通过室内培养向土壤中添加~(13)C标记葡萄糖研究易分解有机碳输入对不同恢复阶段森林土壤激发效应的影响。研究结果表明,易分解有机碳输入引起的土壤激发效应的方向和强度因不同恢复阶段而异。易分解有机碳输入的初期对各恢复阶段森林土壤均产生正的激发效应,然而随着时间的推移,15年、30年马尾松林以及天然林相继出现负的激发效应。从整个培养期(59 d)来看,易分解有机碳的输入促进了裸地与5年生马尾松林土壤有机碳的矿化,有机碳的矿化量分别提高了131%±27%与25%±5%;但是减缓了15年生马尾松林土壤有机碳的矿化,使其矿化量减少了10%±1%;然而,易分解有机碳输入对30年生马尾松林及天然林土壤有机碳的矿化则无明显影响。土壤累积激发碳量与葡萄糖添加前后土壤氮素的改变百分比呈显著正相关关系(R~2=0.44,P0.05),表明易分解有机碳输入诱导的土壤激发效应受土壤氮素可利用性的调控,土壤微生物需要通过分解原有土壤有机碳释放的氮素来满足自身的需求。     

Influence of High CO2Partial Pressure on Nitrogen Use Efficiency of the C4Grasses Panicum coloratum and Cenchrus ciliaris

Australia's tropical grasslands are dominated by C₄grasses,characterized by their unique biochemistry and anatomy. Twonaturalized C₄grasses (Panicum coloratum and Cenchrus ciliaris)were used to investigate whether high CO₂partial pressure [p(CO₂)] influences photosynthetic nitrogen use efficiency andplant nitrogen use efficiency (PNUE and NUE respectively). Plantswere grown for 30 d with four levels of N at p(CO₂) of 38 or86 Pa. PNUE was calculated from leaf CO₂assimilation rates (A)and leaf N concentrations, and NUE from total leaf N contentand plant dry mass. At each p(CO₂), PNUE and NUE were greaterfor C. ciliaris than for P. coloratum due to higher A and drymass combined with lower leaf N concentrations. Elevatedp (CO₂)increased PNUE of C. ciliaris only. This effect was due to lowerleaf N concentrations (area basis). At high p(CO₂), NUE of C.ciliaris was also greater. This resulted from a 1.6-fold stimulationof dry mass by high p(CO₂). Although dry mass of P. coloratumwas increased 1.2-fold by elevated p(CO₂), its NUE was unaffected.Leaf transpiration rates were halved at elevated p(CO₂), andwe suggest that this factor plays a major role in the growthresponse of C₄grasses to high p(CO₂). Copyright 2001 Annalsof Botany Company Panicum coloratum, Cenchrus ciliaris, nitrogen use efficiency, elevated CO₂, leaf N concentration, growth, photosynthesis     

Net primary production,nitrogen cycling,biomass allocation,and resource use efficiency along a topographical soil water and nitrogen gradient in a semi-arid forest near an arid boundary

Aims

The objective of this study was to investigate how plants maintain productivity under a limited supply of water and N along the topographical soil water and N gradients in semi-arid forests.

Methods

We investigated forest structure and productivity, N cycling, and water and N use by plants at three different slope positions in a forested area near an arid boundary on a loess plateau in China.

Results

Net primary production (NPP) and aboveground N uptake decreased as soil water and/or N availability decreased on upper slopes; however, NPP and aboveground N uptake were only slightly lower than those of more humid forest ecosystems. Water use efficiency (WUE), N use efficiency (NUE), and fine root biomass increased as soil water and/or N supply decreased with altitude. High NUE was linked to higher N mean residence time, caused by higher N resorption efficiency rather than increasing N productivity.

Conclusions

Our results suggest that NPP and N uptake can be maintained by increasing WUE and NUE and increasing fine root biomass in water and N co-limited semi-arid forest ecosystems near arid boundaries. Such changes in resource use and acquisition strategy can affect production and N cycling via plant-soil feedback systems.

     

Sustained effects of atmospheric [CO2] and nitrogen availability on forest soil CO2 efflux

Soil CO₂ efflux (F_soil) is the largest source of carbon from forests and reflects primary productivity as well as how carbon is allocated within forest ecosystems. Through early stages of stand development, both elevated [CO₂] and availability of soil nitrogen (N; sum of mineralization, deposition, and fixation) have been shown to increase gross primary productivity, but the long‐term effects of these factors on F_soil are less clear. Expanding on previous studies at the Duke Free‐Air CO₂ Enrichment (FACE) site, we quantified the effects of elevated [CO₂] and N fertilization on F_soil using daily measurements from automated chambers over 10 years. Consistent with previous results, compared to ambient unfertilized plots, annual F_soil increased under elevated [CO₂] (ca. 17%) and decreased with N (ca. 21%). N fertilization under elevated [CO₂] reduced F_soil to values similar to untreated plots. Over the study period, base respiration rates increased with leaf productivity, but declined after productivity saturated. Despite treatment‐induced differences in aboveground biomass, soil temperature and water content were similar among treatments. Interannually, low soil water content decreased annual F_soil from potential values – estimated based on temperature alone assuming nonlimiting soil water content – by ca. 0.7% per 1.0% reduction in relative extractable water. This effect was only slightly ameliorated by elevated [CO₂]. Variability in soil N availability among plots accounted for the spatial variability in F_soil, showing a decrease of ca. 114 g C m^?2 yr^?1 per 1 g m^?2 increase in soil N availability, with consistently higher F_soil in elevated [CO₂] plots ca. 127 g C per 100 ppm [CO₂] over the +200 ppm enrichment. Altogether, reflecting increased belowground carbon partitioning in response to greater plant nutritional needs, the effects of elevated [CO₂] and N fertilization on F_soil in this stand are sustained beyond the early stages of stand development and through stabilization of annual foliage production.     

Nitrogen response efficiency increased monotonically with decreasing soil resource availability: a case study from a semiarid grassland in northern China

The concept of nutrient use efficiency is central to understanding ecosystem functioning because it is the step in which plants can influence the return of nutrients to the soil pool and the quality of the litter. Theory suggests that nutrient efficiency increases unimodally with declining soil resources, but this has not been tested empirically for N and water in grassland ecosystems, where plant growth in these ecosystems is generally thought to be limited by soil N and moisture. In this paper, we tested the N uptake and the N use efficiency (NUE) of two Stipa species (S. grandis and S. krylovii) from 20 sites in the Inner Mongolia grassland by measuring the N content of net primary productivity (NPP). NUE is defined as the total net primary production per unit N absorbed. We further distinguished NUE from N response efficiency (NRE; production per unit N available). We found that NPP increased with soil N and water availability. Efficiency of whole-plant N use, uptake, and response increased monotonically with decreasing soil N and water, being higher on infertile (dry) habitats than on fertile (wet) habitats. We further considered NUE as the product of the N productivity (NP the rate of biomass increase per unit N in the plant) and the mean residence time (MRT; the ratio between the average N pool and the annual N uptake or loss). The NP and NUE of S. grandis growing usually in dry and N-poor habitats exceeded those of S. krylovii abundant in wet and N-rich habitats. NUE differed among sites, and was often affected by the evolutionary trade-off between NP and MRT, where plants and communities had adapted in a way to maximize either NP or MRT, but not both concurrently. Soil N availability and moisture influenced the community-level N uptake efficiency and ultimately the NRE, though the response to N was dependent on the plant community examined. These results show that soil N and water had exerted a great impact on the N efficiency in Stipa species. The intraspecific differences in N efficiency within both Stipa species along soil resource availability gradient may explain the differences in plant productivity on various soils, which will be conducive to our general understanding of the N cycling and vegetation dynamics in northern Chinese grasslands.     

Four-year growth dynamics of beech-spruce model ecosystems under CO2 enrichment on two different forest soils

To elucidate how atmospheric CO₂ enrichment, enhanced nutrient supply and soil quality interact to affect regrowth of temperate forests, young Fagus sylvatica and Picea abies trees were grown together in large model ecosystems. Identical communities were established on a nutrient-poor acidic and on a more fertile calcareous soil and tree growth, leaf area index, fine root density and soil respiration monitored over four complete growing seasons. Biomass responses to CO₂ enrichment and enhanced N supply at the end of the experiment reflected compound interest effects of growth stimulation during the first two to three seasons rather than persistent stimulation over the whole duration of the experiment. Whereas biomass of Picea was enhanced in elevated CO₂ on both soils, Fagus responded negatively to CO₂ on acidic but positively on calcareous soil. Biomass of both species profited from enhanced N supply on the poor acidic soil only. Leaf area index on both soils was greater in high N supply as a consequence of a stimulation early in the experiment, but was unaffected by CO₂ enrichment. Fine root density on acidic soil was increased in high N supply, but this did not stimulate soil respiration rate. In contrast, elevated CO₂ stimulated both fine root density and soil CO₂ efflux on calcareous soil, especially towards the end of the experiment. Our experiment suggests that future species dominance in beech-spruce forests is likely to change in response to CO₂ enrichment, but this response is subject to complex interactions with environmental factors other than CO₂, particularly soil type.     

Long‐term dynamics of mycorrhizal root tips in a loblolly pine forest grown with free‐air CO2 enrichment and soil N fertilization for 6 years

Large‐scale, long‐term FACE (Free‐Air CO₂ enrichment) experiments indicate that increases in atmospheric CO₂ concentrations will influence forest C cycling in unpredictable ways. It has been recently suggested that responses of mycorrhizal fungi could determine whether forest net primary productivity (NPP) is increased by elevated CO₂ over long time periods and if forests soils will function as sources or sinks of C in the future. We studied the dynamic responses of ectomycorrhizae to N fertilization and atmospheric CO₂ enrichment at the Duke FACE experiment using minirhizotrons over a 6 year period (2005–2010). Stimulation of mycorrhizal production by elevated CO₂ was observed during only 1 (2007) of 6 years. This increased the standing crop of mycorrhizal tips during 2007 and 2008; during 2008, significantly higher mortality returned standing crop to ambient levels for the remainder of the experiment. It is therefore unlikely that increased production of mycorrhizal tips can explain the lack of progressive nitrogen limitations and associated increases in N uptake observed in CO₂‐enriched plots at this site. Fertilization generally decreased tree reliance on mycorrhizae as tip production declined with the addition of nitrogen as has been shown in many other studies. Annual NPP of mycorrhizal tips was greatest during years with warm January temperatures and during years with cool spring temperatures. A 2 °C increase in average late spring temperatures (May and June) decreased annual production of mycorrhizal root tip length by 50%. This has important implications for ecosystem function in a warmer world in addition to potential for forest soils to sequester atmospheric C.