Changing leaf litter feedbacks on plant production across contrasting sub-arctic peatland species and growth forms

Plant species and growth forms differ widely in litter chemistry, which affects decay and may have important consequences for plant growth via e.g. the release of nutrients and growth-inhibitory compounds. We investigated the overall short-term (9.5 months) and medium-term (21.5 months) feedback effects of leaf litter quality and quantity on plant production, and tested whether growth forms can be used to generalise differences among litter species. Leaf litter effects of 21 sub-arctic vascular peatland species on Poa alpina test plants changed clearly with time. Across all growth forms, litter initially reduced plant biomass compared with untreated plants, particularly litters with a high decomposition rate or low initial lignin/P ratio. In the second year, however, litter effects were neutral or positive, and related to initial litter N concentration (positive), C/N, polyphenol/N and polyphenol/P ratios (all negative), but not to decomposability. Differences in effect size among several litter species were large, while differences in response to increasing litter quantities were not significant or of similar magnitude to differences in response to three contrasting litter species. Growth forms did not differ in initial litter effects, but second-year plant production showed a trend (P < 0.10) for differences in response to litters of different growth forms: evergreen shrubs < graminoids or deciduous shrubs < forbs. While long-persisting negative litter effects were predominant across all growth forms, our data indicate that even within nutrient-constrained ecosystems such as northern peatlands, vascular plant species, and possibly growth forms, differ in litter feedbacks to plant growth. Differences in the composition of undisturbed plant communities or species shifts induced by external disturbance, such as climate change, may therefore feedback strongly to plant biomass production and probably nutrient cycling rates in northern peatlands. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Nutrient Addition Differentially Affects Ecological Processes of <Emphasis Type="Italic">Avicennia germinans</Emphasis> in Nitrogen versus Phosphorus Limited Mangrove Ecosystems

Nutrient over-enrichment is a major threat to marine environments, but system-specific attributes of coastal ecosystems may result in differences in their sensitivity and susceptibility to eutrophication. We used fertilization experiments in nitrogen (N)- and phosphorus (P)-limited mangrove forests to test the hypothesis that alleviating different kinds of nutrient limitation may have different effects on ecosystem structure and function in natural systems. We compared a broad range of ecological processes to determine if these systems have different thresholds where shifts might occur in nutrient limitation. Growth responses indicated N limitation in Avicennia germinans (black mangrove) forests in the Indian River Lagoon (IRL), Florida, and P limitation at Twin Cays, Belize. When nutrient deficiency was relieved, A. germinans grew out of its stunted form by increasing wood relative to leaf biomass and shoot length relative to lateral growth. At the P-limited site, P enrichment (+P) increased specific leaf area, N resorption, and P uptake, but had no effect on P resorption. At the N-limited site, +N increased both N and P resorption, but did not alter biomass allocation. Herbivory was greater at the P-limited site and was unaffected by +P, whereas +N led to increased herbivory at the N-limited site. The responses to nutrient enrichment depended on the ecological process and limiting nutrient and suggested that N- versus P-limited mangroves do have different thresholds. +P had a greater effect on more ecological processes at Twin Cays than did +N at the IRL, which indicated that the P-limited site was more sensitive to nutrient loading. Because of this sensitivity, eutrophication is more likely to cause a shift in nutrient limitation at P-limited Twin Cays than N-limited IRL.     

Differences in plant function in phosphorus- and nitrogen-limited mangrove ecosystems

Mangrove ecosystems can be either nitrogen (N) or phosphorus (P) limited and are therefore vulnerable to nutrient pollution. Nutrient enrichment with either N or P may have differing effects on ecosystems because of underlying differences in plant physiological responses to these nutrients in either N- or P-limited settings. Using a common mangrove species, Avicennia germinans, in sites where growth was either N or P limited, we investigated differing physiological responses to N and P limitation and fertilization. We tested the hypothesis that water uptake and transport, and hydraulic architecture, were the main processes limiting productivity at the P-limited site, but that this was not the case at the N-limited site. We found that plants at the P-deficient site had lower leaf water potential, stomatal conductance and photosynthetic carbon-assimilation rates, and less conductive xylem, than those at the N-limited site. These differences were greatly reduced with P fertilization at the P-limited site. By contrast, fertilization with N at the N-limited site had little effect on either photosynthetic or hydraulic traits. We conclude that growth in N- and P-limited sites differentially affect the hydraulic pathways of mangroves. Plants experiencing P limitation appear to be water deficient and undergo more pronounced changes in structure and function with relief of nutrient deficiency than those in N-limited ecosystems.     

Polyphenols in litter from tropical montane forests across a wide range in soil fertility

In nutrient-poor ecosystems high polyphenol concentrations in plant litter have been proposed to influence soil nutrient availability in benefit of the plants. We addressed the question whether litter polyphenol concentrations vary across a soil chronosequence of almost identical geology, climate and plant species composition, but of a wide range in nitrogen (N) and phosphorus (P) availability in the Hawaiian Islands. Concentrations of total phenolics (TPh) and proanthocyanidins (PA) in leaf litter of the dominant tree species Metrosideros polymorpha were higher at the oldest, P-limited site compared to the youngest, N-limited site, with intermediate values at the two relatively fertile sites co-limited by N and P. Polyphenol concentrations in fine root litter differed considerably from those observed in leaf litter and varied differently across the soil age gradient. Long-term fertilization did not significantly alter polyphenol concentrations in Metrosideros litter at either site. Moreover, green leaves and leaf litter of Metrosideros showed similar relative differences among sites when compared between natural populations and plants from the same populations but grown in a common garden. These results suggest that polyphenol concentrations inherently vary among populations of the dominant tree species in Hawaiian montane forests possibly indicating an adaptation to ecosystem properties such as substrate age related differences in soil fertility. The combined above- and below-ground input rate of TPh ranged from 62.4 to 170.8 g/m²/yr and was significantly higher at the P-limited than at the N-limited site. Root-derived polyphenols contributed a much higher absolute and relative amount of phenolic input at the N-limited than at the P-limited site. The differences in amount, quality, and pathways of input might suggest specific interactions with soil processes and nutrient cycling among the Hawaiian rainforests studied here.     

The effect of nitrogen nutrition on growth and biomass partitioning of annual plants originating from habitats of different nitrogen availability

Summary The hypothesis was tested that faster growth of nitrophilic plants at high nitrogen (N) nutrition is counterbalanced by faster growth of non-nitrophilic plants at low N-nutrition. Ten annual plant species were used which originated from habitats of different N-availability. The species' preference for N was quantified by the N-number of Ellenberg (1979), a relative measure of nitrophily. The plants were cultivated in a growth cabinet at five levels of ammonium-nitrate supply. At low N-supply, the relative growth rate (RGR) was independent of nitrophily. At high N-supply, RGR tended to be higher in nitrophilic than in non-nitrophilic species. However, the response of RGR to N-supply was strongly and positively correlated with the nitrophily of species. Increasing N-supply enhanced partitioning to leaf weight per total biomass (LWR) and increased plant leaf area per total biomass (LAR). Specific leaf weight (SLW) and LWR were both higher in non-nitrophilic than in nitrophilic species at all levels of N-nutrition. NAR (growth per leaf area or net assimilation rate) increased with nitrophily only under conditions of high N-supply. RGR correlated positively with LAR, irrespective of N-nutrition. Under conditions of high N-supply RGR correlated with SLW negatively and with NAR positively.     

Effects of elevated [CO2] on forest growth and carbon storage: a modelling analysis of the consequences of changes in litter quality/quantity and root exudation

Many researchers have proposed that the stimulus of plant growth under elevated [CO₂] observed in short-term experiments will be moderated in the longer term by a reduction in soil nitrogen (N) availability linked to decreased litter quality and/or increased litter production. However, these negative feedbacks may be offset to some extent by a stimulus in N fixation linked to increased root exudation. The aim of this modelling study is to examine how changes in litter quality/quantity and root exudation –- if they occur –- will affect the CO₂ responses of net primary productivity and ecosystem carbon (C) storage on different timescales. We apply a model of C and N cycling in forest ecosystems (G’DAY) to stands of Norway spruce (Picea abies, L. Cast) growing at a N-limited experimental site at Flakaliden, Sweden, and draw the following conclusions: (1) in the absence of changes in litter quality and root exudation, the short-term CO₂ stimulus of litter quantity leads to only a minimal CO₂ stimulus of productivity or C storage in the medium term (≈ 20 years) and long term (≈ 200 years), because of constraints on soil N availability; (2) increasing plant nitrogen use efficiency (via a decrease in the N:C ratio of new litter) makes little impact on these results; (3) a significant CO₂ response in the medium term requires a substantial decrease in the N:C ratio of older litter, when it is approaching stabilisation as soil organic matter, although the long-term CO₂ response remains small; and (4) an increase in N fixation leads to a small effect on productivity in the short term, but a very large effect on both productivity and C storage in the long term. These results suggest that soil N constraints on the long-term CO₂-fertilisation effect can be overcome to a significant extent only by increases in N acquisition, although only modest increases may be required. This revised version was published online in June 2006 with corrections to the Cover Date.     

What are the effects of nitrogen deficiency on growth components of lettuce?

Relationships between nitrogen (N) content and growth are routinely measured in plants. This study determined the effects of N on the separate morphological and physiological components of plant growth, to assess how N-limited growth is effected through these components. Lettuce ( Lactuca sativa ) plants were grown hydroponically under contrasting N-supply regimes, with the external N supply either maintained continuously throughout the period of study, or withdrawn for up to 14 d. Richards' growth functions, selected using an objective curve-fitting technique, accounted for 99.0 and 99.1% of the variation in plant dry weight for control and N-limited plants respectively. Sublinear relationships occurred between N and relative growth rates under restricted N-supply conditions, consistent with previous observations. There were effects of treatment on morphological and physiological components of growth. Leaf weight ratio increased over time in control plants and decreased in N- limited plants. Shoot:root ratio followed a similar pattern. On a whole-plant basis, assimilation of carbon decreased in N-limited plants, a response paralleled by differences in stomatal conductance between treatments. Changes in C assimilation, expressed as a function of stomatal conductance to water vapour, suggest that the effects of N limitation on growth did not result directly from a lack of photosynthetic enzymes. Relationships between plant N content and components of growth will depend on the availability of different N pools for remobilization and use within the plant.     

Foliar and Litter Nutrients, Nutrient Resorption, and Decomposition in Hawaiian Me t rosideros polymorpha

The native tree Metrosideros polymorpha dominates Hawaiian forests across a very wide range of soil fertility, including both sites where forest production is limited by nitrogen (N) and others where it is limited by phosphorus (P). Five long-term fertilization experiments have further broadened the range of nutrient availabilities experienced by Metrosideros. Adding P to P-limited sites increased foliar P concentrations threefold and litter P concentrations up to 10-fold; lignin concentrations decreased, and the decomposability of leaf litter increased from 32%–35% to 36%–46% mass loss in the first year. Adding N to N-limited sites increased leaf and litter N concentrations by only 15%–20%, with little or no effect on the decomposability of tissue. Received 22 January 1998; accepted 4 May 1998.     

Belowground positive and negative feedbacks on CO2 growth enhancement

In this paper we present a conceptual model of integrated plant-soil interactions which illustrates the importance of identifying the primary belowground feedbacks, both positive and negative, which can simultaneously affect plant growth responses to elevated CO₂. The primary negative feedbacks share the common feature of reducing the amount of nutrients available to plants. These negative feedbacks include increased litter C/N ratios, and therefore reduced mineralization rates, increased immobilization of available nutrients by a larger soil microbial pool, and increased storage of nutrients in plant biomass and detritus due to increases in net primary productivity (NPP). Most of the primary positive feedbacks share the common feature of being plant mediated feedbacks, the only exception being Zak et al.'s hypothesis that increased microbial biomass will be accompanied by increased mineralization rates. Plant nutrient uptake may be increased through alterations in root architecture, physiology, or mycorrhizal symbioses. Further, the increased C/N ratios of plant tissue mean that a given level of NPP can be achieved with a smaller supply of nitrogen.Identification of the net plant-soil feedbacks to enhanced productivity with elevated CO₂ are a critical first step for any ecosystem. It is necessary, however, that we first identify how universally applicable the results are from one study of one ecosystem before ecosystem models incorporate this information. The effect of elevated CO₂ on plant growth (including NPP, tissue quality, root architecture, mycorrhizal symbioses) can vary greatly for different species and environmental conditions. Therefore it is reasonable to expect that different ecosystems will show different patterns of interacting positive and negative feedbacks within the plant-soil system. This inter-ecosystem variability in the potential for long-term growth responses to rising CO₂ levels implies that we need to parameterize mechanistic models of the impact of elevated CO₂ on ecosystem productivity using a detailed understanding of each ecosystem of interest.     

Nitrogen supply differentially affects litter decomposition rates and nitrogen dynamics of sub-arctic bog species

High-latitude peatlands are important soil carbon sinks. In these ecosystems, the mineralization of carbon and nitrogen are constrained by low temperatures and low nutrient concentrations in plant litter and soil organic matter. Global warming is predicted to increase soil N availability for plants at high-latitude sites. We applied N fertilizer as an experimental analogue for this increase. In a three-year field experiment we studied N fertilization effects on leaf litter decomposition and N dynamics of the four dominant plant species (comprising >75% of total aboveground biomass) in a sub-arctic bog in northern Sweden. The species were Empetrum nigrum (evergreen shrub), Eriophorum vaginatum (graminoid), Betula nana (deciduous shrub) and Rubus chamaemorus (perennial forb). In the controls, litter mass loss rates increased in the order: Empetrum < Eriophorum < Betula < Rubus. Increased N availability had variable, species-specific effects: litter mass loss rates (expressed per unit litter mass) increased in Empetrum, did not change in Eriophorum and Betula and decreased in Rubus. In the leaf litter from the controls, we measured no or only slight net N mineralization even after three years. In the N-fertilized treatments we found strong net N immobilization, especially in Eriophorum and Betula. This suggests that, probably owing to substantial chemical and/or microbial immobilization, additional N supply does not increase the rate of N cycling for at least the first three years.     

Dynamics and Nutritional Ecology of a Nanoflagellate Preying Upon Bacteria

Ingestion and growth rates of the nanoflagellate predator Ochromonas danica feeding on the bacterium Pseudomonas fluorescens were quantified in laboratory cultures. Bacterial prey were grown under four nutritional conditions with respect to macronutrient elements: C-limited, N-limited, P-limited, and balanced. Ingestion and growth rates were saturating functions of prey abundance when preying upon nutritionally balanced, C-limited, and P-limited bacteria but were unimodal functions of abundance when preying on N-limited bacteria. At saturating prey concentrations, the ingestion rate of C-limited prey was about twice that of prey in other nutritional states, while at subsaturating prey concentrations, the ingestion rates of both C- and N-limited prey were higher than those of prey in other nutritional states. Over all prey concentrations, growth was most rapid on balanced and C-limited prey and generally lowest for P-limited prey. Due to the unimodal response of growth rate to abundance of N-limited prey, growth rate on N-limited prey approached that obtained on balanced and C-limited prey when prey were available at intermediate abundances. The accumulation of recycled N increased with the growth rate of O. danica. Recycling of N was highest when O. danica was feeding upon P-limited prey. The accumulation of recycled P increased with growth rate for balanced and N-limited prey, but not for P-limited prey, which consistently had low accumulation of recycled P. The low growth rate and negligible recycling of P for O. danica preying on P-limited prey is consistent with the theory of ecological stoichiometry and resembles results found for crustacean zooplankton, especially in the genus Daphnia. Potentially, the major predators of bacterioplankton and a major predator of phytoplankton play analogous roles in the trophic dynamics and biogeochemistry of aquatic ecosystems.     

Litter N:P ratios indicate whether N or P limits the decomposability of graminoid leaf litter

The N:P ratio of leaf litter may determine if decomposability is N-limited (litter with low N:P ratio) or P-limited (litter with high N:P ratio). To test this hypothesis and to determine the threshold between N and P limitation, we studied relationships between litter N and P concentrations, litter mass loss and effects of fertilisation on litter mass loss in laboratory experiments. Leaf litter of 11 graminoid species was collected in Swiss and Dutch wetlands, yielding 84 litter samples with a broad range of N and P concentrations (3.2–15.1 mg N g⁻¹, 0.04–1.93 mg P g⁻¹) and with N:P mass ratios ranging from 5 to 100. On nutrient-free sand, dry mass loss after five or ten weeks (5.5–53% of initial mass) correlated positively with the N and P concentrations of the litter. Within species, mass loss correlated mainly with N for litter with low N:P ratio, and with P for litter with high N:P ratio, in agreement with our hypothesis. Among species, however, these relationships did not exist, and decomposition rather correlated with the specific leaf area. When the litter was incubated on fertilised sand, 35 out of 50 litter samples decomposed faster than on nutrient-free sand. Decomposition was generally accelerated by P fertilisation (i.e. P-limited) when the N:P ratio of the litter was above 25 and the P concentration below 0.22 mg g⁻¹, supporting our hypothesis. N-limited decomposition was not significantly related to the litter N:P ratio but occurred rarely for litter with N:P ratio greater than 25, and only for litter with N concentration below 11.3 mg g⁻¹. We conclude that the N:P ratio of leaf litter indicates whether its decomposability is more likely to be N- or P-limited. The critical N:P ratio (threshold between N and P limitation) appeared to be 25 for graminoid leaf litter.     

喀斯特区不同退化程度植被群落植物-凋落物-土壤-微生物生态化学计量特征

为摸清喀斯特植被退化对群落各组分C、N、P生态化学计量特征及内稳态特征的影响，为喀斯特退化生态系统植被恢复与重建提供科学依据，以桂西北喀斯特地区5种退化程度植被群落为研究对象，测定了不同退化程度植被群落植物叶片、凋落物、土壤和微生物生物量的C、N、P含量，分析其化学计量比特征、相互关系及植物内稳性特征。结果表明：(1)随着退化程度加剧，叶片C、N、P含量、N∶P和凋落物N∶P、微生物量C显著下降，而叶片C∶N、C∶P则显著增加，且植物叶片N∶P<14;随退化程度加剧，凋落物N、P含量、土壤C、N、P含量、微生物量N、P呈先略有增后显著降低的趋势，且不同退化程度群落土壤N∶P和微生物量C∶N无显著差异。(2)叶片N、P含量与土壤N、P含量，叶片C∶P与土壤C∶N、C∶P、N∶P,叶片N∶P与凋落物N、N∶P,叶片C、N、P含量与微生物量C呈显著或极显著正相关关系；叶片C∶N与土壤C、N,叶片C∶P与土壤N、P,叶片N∶P与土壤P呈显著或极显著负相关关系。(3)喀斯特地区植物叶片N、P元素的内稳性指数(H)平均值分别为2.74和2.31,属于弱稳态型，叶片N∶P的H值为5.14,为稳...     

Nutrient limitation in species-rich lowland fens

Abstract. Nitrogen, phosphorus and potassium were supplied to some Belgian fens of varying nutrient status and productivity. Plant growth in the lowest productive fen with a species-rich Caricion davallianae vegetation was strongly P-limited. N was ineffective when applied alone, but increased the effect of P-addition when applied together. Summer biomass and plant nutrient concentrations were monitored for four years, and showed partial recovery of nutrient limitation. In a more productive fen dominated by Carex lasiocarpa and in a fen meadow, nutrient limitation was less strong. N limited growth in the productive fen, and N and K were co-limiting in the fen meadow. The P-concentration in the productive fen vegetation showed a marked increase after P-fertilization, but it did not result in higher standing crop. The significance of P-limitation for the conservation of species rich low productive fens is discussed. P-limitation may be an essential feature in the conservation of low productive rich fens: because it is less mobile in the landscape than N and/or because it is an intrinsic property of this vegetation type. Plant nutrient concentrations and N:P-ratios may be used as an indication for the presence and type of nutrient limitation in the vegetation. We found N:P-ratios of 23 to 31 for a P-limited site and 8 to 15 in N-limited sites. This was in agreement with critical values from the literature: N:P > ca. 20 for P-limitation and N:P < 14 for N-limitation. Thus, this technique appears valid in the vegetation types that were studied here.     

Effects of nutrient addition on vegetation and carbon cycling in an ombrotrophic bog

We measured net ecosystem CO₂ exchange (NEE), plant biomass and growth, species composition, peat microclimate, and litter decomposition in a fertilization experiment at Mer Bleue Bog, Ottawa, Ontario. The bog is located in the zone with the highest atmospheric nitrogen deposition for Canada, estimated at 0.8–1.2 g N m⁻² yr⁻¹ (wet deposition as NH₄ and NO₃). To establish the effect of nutrient addition on this ecosystem, we fertilized the bog with six treatments involving the application of 1.6–6 g N m⁻² yr⁻¹ (as NH₄NO₃), with and without P and K, in triplicate 3 m × 3 m plots. The initial 5–6 years have shown a loss of first Sphagnum, then Polytrichum mosses, and an increase in vascular plant biomass and leaf area index. Analyses of NEE, measured in situ with climate‐controlled chambers, indicate that contrary to expectations, the treatments with the highest levels of nutrient addition showed lower rates of maximum NEE and gross photosynthesis, but little change in ecosystem respiration after 5 years. Although shrub biomass and leaf area increased in the high nutrient plots, loss of moss photosynthesis owing to nutrient toxicity, increased vascular plant shading and greater litter accumulation contributed to the lower levels of CO₂ uptake. Our study highlights the importance of long‐term experiments as we did not observe lower NEE until the fifth year of the experiment. However, this may be a transient response as the treatment plots continue to change. Higher levels of nutrients may cause changes in plant composition and productivity and decrease the ability of peatlands to sequester CO₂ from the atmosphere.     

Decomposability of C3 and C4 grass litter sampled under different concentrations of atmospheric carbon dioxide at a natural CO2 spring

Elevated concentrations of atmospheric CO₂ can influence the relative proportions, biomass and chemical composition of plant species in an ecosystem and, thereby, the input of litter nutrients to soil. Plant growth under elevated CO₂ appears to have no consistent effect on rates of litter decomposition; decomposition can, however, differ in C3 and C4 plant material from the same CO₂ environment. We here describe the decomposability of leaf litter of two grass species – the C3 Holcus lanatus L. (Yorkshire fog) and C4 Pennisetum clandestinum Hochst. (kikuyu) - from an unfertilized, ungrazed grassland at a cold CO₂ spring in Northland, New Zealand. Decomposability was measured by net CO₂–C production from litter incubated for 56 days at 25 °C in a gley soil from the site; net mineral-N production from litter was also determined. Both litter and soils were sampled under `low' and `high' concentrations of atmospheric CO₂. Decomposition of H. lanatus litter was greater than that of P. clandestinum litter throughout the 56-day incubation. Decomposition tended to be greater in `high-CO<sub>2</sub>' than in `low-CO₂' H. lanatus litter, but lower in `high-CO<sub>2</sub>' than `low-CO₂' P. clandestinum litter; differences were, however, non-significant after 28 days. Overall, litter decomposition was greater in the `low-CO<sub>2</sub>' than `high-CO₂' soil. Differences in decomposition rates were related negatively to litter N concentrations and positively to C:N ratios, but were not predictable from lignin:total N ratios. Net mineral-N production from litter decomposition did not differ significantly in `high-CO<sub>2</sub>' and `low-CO₂' samples incubated in `low-CO<sub>2</sub>' soil; in `high-CO₂' soil some net immobilization was observed. Overall, results indicate the likely complexity of litter decomposition in the field but, nevertheless, strongly suggest that rates of decomposition will not necessarily decline in a `high-CO₂' environment.     

Litter species traits, but not richness, contribute to carbon and nitrogen dynamics in an alpine meadow on the Tibetan Plateau

Aims

Litter, as afterlife of plants, plays an important role in driving belowground decomposition processes. Here we tested effects of litter species identity and diversity on carbon (C) and nitrogen (N) dynamics during litter decomposition in N-limited alpine meadow soil from the Qinghai–Tibet Plateau.

Methods

We incubated litters of four meadow species, a sedge (“S”, Kobresia humilis), a grass (“G”, Elymus nutans), a herb (“H”, Saussurea superba), and a legume (“L”, Oxytropis falcata), in monoculture and in mixture with meadow soil. CO₂ release was measured 21 times during the incubation, and soil available N and microbial biomass C and N were measured before and after the experiment.

Results

The organic C decay rate did not differ much among soils amended with monocultures or mixtures of litter, except in the H, S, L, and S+H treatments, which had much higher decay rates. Potential decomposable C pools were lowest in the control, highest in the L treatment, and intermediate in the S treatment. Mineralized N was completely immobilized by soil microbes in all treatments except the control, S+L, and S+G+L treatments. Litter mixtures had both additive and non-additive effects on CO₂-C emission (mainly antagonistic effects), net N mineralization (mainly synergistic), and microbial biomass C and N (both). Overall, these parameters were not significantly correlated with litter species richness. Similarly, microbial C or N was not significantly correlated with litter N content or C/N. However, cumulative CO₂-C emission and net N mineralization were positively correlated with litter N content and negatively correlated with litter C/N.

Conclusions

Litter N content and C/N rather than litter species richness drove the release of CO₂-C and net available N in this ecosystem. The antagonistic effects of litter mixtures contributed to a modest release of CO₂-C, but their synergistic effects enhanced net available N. We suggest that in alpine meadow communities, balancing species with high and low N contents will benefit soil carbon sequestration and plant competition for available N with soil microbes.     

The role of vegetation and litter in the nitrogen dynamics of riparian buffer zones in Europe

Plant uptake and denitrification are considered to be the most important processes responsible for N retention and mitigation in riparian buffers. In many riparian buffers, however, nutrients taken up by plants remain in the system only temporarily and may be gradually released by mineralization later. Still, plants increase the residence time of nutrients considerably by reducing their mobility. We investigated the importance of plant N uptake and N immobilization in litter for N retention in riparian buffers. Nitrogen uptake in vegetation and N dynamics in litter were measured over a two-year period in a range of forested and herbaceous riparian buffers along a climatic gradient in Europe, receiving different loadings of N-enriched groundwater. Plant production, nitrogen uptake, and N retention were significantly higher in the forested buffer sites compared to the herbaceous buffer sites. However, in herbaceous buffers, periodic harvesting of herbaceous biomass contributed considerably to the N retention. No relationship between lateral N loading and plant productivity or N uptake was observed; this indicated that plant growth was not N-limited. In the winter period, decaying leaf litter had a small but significant role in N retention in a majority of the riparian ecosystems studied. Moreover, no responses to the climatic gradient were found. Generally, we can state that annual N retention in the vegetation and litter compartment is substantial, making up 13–99% of the total N mitigation.     

Growth rate and biomass partitioning of wildtype and low-gibberellin tomato (Solanum lycopersicum) plants growing at a high and low nitrogen supply

The growth-promoting effects of gibberellins (GAs) on plants are well documented, but a complete growth analysis at the whole plant level on plants with an altered GA biosynthesis has never been reported. In the present work, the relative growth rate (RGR), biomass partitioning and morphological parameters of wildtype (Wt) tomato ( Solanum lycopersicum L. cv. Moneymaker) plants were compared with those of isogenic ( gib ) mutants with a reduced biosynthesis of gibberellins. GA deficiency reduced RGR and specific leaf area (SLA, leaf area per unit leaf mass) and increased the net assimilation rate (NAR, the rate of biomass increment per unit leaf area). Despite the free access to nitrogen in the rooting medium, the low-GA mutants had a much higher root mass ratio (RMR, the root mass per unit plant biomass) than the Wt, suggesting that the mutants were disturbed in their growth response to nitrate supply. The experiment was repeated at a low exponential nitrate supply, which forced all plants to grow at the same low RGR. The persistence of the differences in RMR at low N-supply indicated that the high RMR of the mutants was a direct effect of low GA, which was independent of nitrate supply. Because the low N-supply increased the RMRs of all genotypes to the same extent, the response of RMR to N-supply does not seem to depend on GA. Although many of the traits of the slow growing GA mutants were very similar to those of inherently slow growing plant species from unproductive habitats, gibberellins are unlikely to be a main determinant of a plant's potential RGR.     

Responses of wetland graminoids to the relative supply of nitrogen and phosphorus

The biomass production of wetland vegetation can be limited by nitrogen or phosphorus. Some species are most abundant in N-limited vegetation, and others in P-limited vegetation, possibly because growth-related traits of these species respond differently to N versus P supply. Two growth experiments were carried out to examine how various morphological and physiological traits respond to the relative supply of N and P, and whether species from sites with contrasting nutrient availability respond differently. In experiment 1, four Carex species were grown in nutrient solutions at five N:P supply ratios (1.7, 5, 15, 45, 135) combined with two levels of supply (geometric means of N and P supply). In experiment 2, two Carex and two grass species were grown in sand at the same .ve N:P supply ratios combined with three levels of supply and two light intensities (45% or 5% daylight). After 12-13 weeks of growth, plant biomass, allocation, leaf area, tissue nutrient concentrations and rates and nutrient uptake depended signi.cantly on the N:P supply ratio, but the type and strength of the responses differed among these traits. The P concentration and the N:P ratio of shoots and roots as well as the rates of N and P uptake were mainly determined by the N:P supply ratio; they showed little or no dependence on the supply level and relatively small interspeci.c variation. By contrast, the N concentration, root mass ratio, leaf dry matter content and speci.c leaf area were only weakly related to the N:P supply ratio; they mainly depended on plant species and light, and partly on overall nutrient supply. Plant biomass was determined by all factors together. Within a level of light and nutrient supply, biomass was generally maximal (i.e. co-limited by N and P) at a N:P supply ratio of 15 or 45. All species responded in a similar way to the N:P supply ratio. In particular, the grass species Phalaris arundinacea and Molinia caerulea showed no differences in response that could clearly explain why P. arundinacea tends to invade P-rich (N-limited) sites, and M. caerulea P-limited sites. This may be due to the short duration of the experiments, which investigated growth and nutrient acquisition but not nutrient con­servation.