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.     

Foliar nitrogen and phosphorus accumulation responses after fertilization: an example from nutrient-limited Hawaiian forests

How plants respond to long-term nutrient enrichment can provide insights into physiological and evolutionary constraints in various ecosystems. The present study examined foliar concentrations after fertilization—to determine if nutrient accumulation responses of the most abundant species in a plant community reflect differences in N and P uptake and storage. Using a chronosequence in the Hawaiian Islands that differs in N and P availability, it was shown that after fertilization, plants increase foliar P to a much greater degree than foliar N, as indicated by response ratios. In addition, foliar P responses after fertilization were more variable and largely driving the observed changes in N:P values. Across species, both inorganic and organic P increased but neither form of N increased significantly. This pattern of P accumulation was consistent across 13 species of varying life forms and occurred at both the N-limited and P-limited site, although its magnitude was larger at the P-limited site. Foliar P accumulation after nutrient enrichment may indicate nutrient storage and may have evolved to be a general strategy to deal with uncertainties in P availability. Storage of P complicates interpretations of N:P values and the determination of nutrient limitation.     

氮磷营养限制影响三角褐指藻光合无机碳利用和碳酸酐酶活性

以海洋硅藻三角褐指藻为实验材料, 研究了不同氮磷比培养对其光合无机碳利用和碳酸酐酶活性的影响, 结果显示三角褐指藻生长速率在N:P=16:1时最大, 高于或低于16:1时明显下降, 表明其最适生长受到氮磷的限制。氮限制(N:P=4:1或1:1)导致叶绿素a含量分别下降30.1% 和47.6%, 磷限制(N:P=64:1或256:1)下降39.1%和52.4%, 但氮或磷限制对叶绿素c含量并没有明显影响。不同营养水平培养对光饱和光合速率具有明显的影响, 与营养充足培养相比, 在严重氮磷限制(N:P=1:1或256:1)培养下光饱和光合速率分别下降39.7%和48.0%, 光合效率与暗呼吸速率也明显下降。在氮磷限制培养下藻细胞pH补偿点明显下降; K0.5CO2值在磷限制下降低30%, 表明磷限制有助于提高细胞对CO2的亲和力, 但氮限制并没有明显影响。在氮磷限制培养的细胞反应液中Fe (CN)63-浓度下降速率较慢, 表明在氮磷限制环境中生长的细胞质膜氧化还原能力明显低于营养充足条件下生长的细胞。氮磷限制也导致胞内、外碳酸酐酶活性明显下降, 其中在氮限制下胞外碳酸酐酶活性分别下降50%和37.5%, 在磷限制下下降22.3%和42.1%。严重的氮(N:P=1:1)或磷(N:P=256:1)限制导致胞内碳酸酐酶活性下降36.5%和42.9%。研究结果表明, 三角褐指藻细胞在氮磷营养限制的环境中, 可以通过调节叶绿素含量、无机碳的利用方式和碳酸酐酶的活性以维持适度的生长。       

Nitrogen limitation of growth and nutrient dynamics in a disturbed mangrove forest,Indian River Lagoon,Florida

The objectives of this study were to determine effects of nutrient enrichment on plant growth, nutrient dynamics, and photosynthesis in a disturbed mangrove forest in an abandoned mosquito impoundment in Florida. Impounding altered the hydrology and soil chemistry of the site. In 1997, we established a factorial experiment along a tree-height gradient with three zones, i.e., fringe, transition, dwarf, and three fertilizer treatment levels, i.e., nitrogen (N), phosphorus (P), control, in Mosquito Impoundment 23 on the eastern side of Indian River. Transects traversed the forest perpendicular to the shoreline, from a Rhizophora mangle-dominated fringe through an Avicennia germinans stand of intermediate height, and into a scrub or dwarf stand of A. germinans in the hinterland. Growth rates increased significantly in response to N fertilization. Our growth data indicated that this site is N-limited along the tree-height gradient. After 2 years of N addition, dwarf trees resembled vigorously growing saplings. Addition of N also affected internal dynamics of N and P and caused increases in rates of photosynthesis. These findings contrast with results for a R. mangle-dominated forest in Belize where the fringe is N-limited, but the dwarf zone is P-limited and the transition zone is co-limited by N and P. This study demonstrated that patterns of nutrient limitation in mangrove ecosystems are complex, that not all processes respond similarly to the same nutrient, and that similar habitats are not limited by the same nutrient when different mangrove forests are compared.     

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.     

Growth, photosynthesis, respiration, and intracellular free amino acid profiles in the unicellular alga Cyanidium caldarium. Effect of nutrient limitation and resupply

Growth, intrácellular free amino acid pools and photosynthetic and respiratory activities in nutrient sufficient cells and in N- K- and P-limited cells of Cyanidium caldarium (Tilden) Geitler, and responses to nutrient resupply were investigated. Addition of ammonium to N-limited cells and of phosphate to P-limited cells resulted in a stimulation of dark respiration and in a decrease in photosynthetic oxygen evolution. Addition of K to K-limited cells had no effect on rates of photosynthesis and respiration. Nutrient limited cells and sufficient cells exhibited different free amino acid profiles. Upon resupply of ammonium to N-limited cells levels of glutamine, citrulline, arginine, alanine, and serine increased. Also the levels of δ-aminolevulinic acid (δ-ALA) and putrescine increased notably. On adding phosphate to P-limited cells the level of glutamate decreased significantly whereas the level of alanine increased and the concentrations of other amino acids remained unaffected. On adding potassium to K-limited cells there was an increase in glutamate and citrulline concentrations, and a decrease in putrescine concentration, whereas concentrations of arginine and alanine remained at the very high levels observed already before addition. Resuspension of N- and K-limited cells in a complete growth only after 25-30h. In P-limited cells resumption of growth in complete medium occurred progressively and reached the maximum rate 30h later. P-, K- and N- limited cells resuspended into sufficient media showed different rates of ammonium and phosphate assimilation. The pattern of recovery from nutrient limitation is discussed according to the cellular role fulfilled by the nutrient which was growth rate-limiting.     

Potential ecosystem-level effects of genetic variation among populations of Metrosideros polymorpha from a soil fertility gradient in Hawaii

This study assessed intrinsic differences in tissue quality and growth rate among populations of Metrosideros polymorpha native to sites with a range of soil fertilities. We collected seedlings from three Hawaiian mesic forests that were either phosphorus-limited, nitrogen-limited, or relatively fertile. These individuals were grown in a common garden under a factorial high/low, N/P fertilization regime for 1.5 years and then harvested to determine genetic divergence; aboveground growth rate; and lignin, N, and P concentrations in leaves and roots. Allozyme analyses indicated that the three groups had genetically diverged to some degree (genetic distance = 0.036-0.053 among populations). Relative growth rate did not differ significantly among the populations. Senescent leaves from the fertile-site population had the highest N concentrations (due to low N resorption) and had lower lignin concentrations than plants from the N-limited site. Across treatments, P concentrations in senescent leaves were highest in plants from the fertile and P-limited site. Root tissue quality did not generally differ significantly among populations. Since decomposition rate of senescent leaves in this system is related positively to N concentration and negatively to lignin concentration, senescent leaves from the fertile-site population may have a genetic tendency toward faster decay than the others. The intrinsic qualities of the three populations may provide positive feedbacks on nutrient cycling at each site-nutrient availability may be raised to some degree at the fertile site, and reduced at the N- or P-limited sites. Our results suggest that even a small degree of genetic differentiation among groups can influence traits related to nutrient cycling.     

Production and Resource Use Efficiencies in N- and P-Limited Tropical Forests: A Comparison of Responses to Long-term Fertilization

At two sites at the extreme ends of a soil development chronosequence in Hawaii, we investigated whether forest responses to fertilization on young soils were similar to those on highly weathered soils and whether the initial responses were maintained after 6–11 years of fertilization. Aboveground net primary production (ANPP) was increased by nitrogen (N) application at the 300-year-old site and phosphorus (P) application at the 4.1-million-year-old site, thus confirming earlier results and their designations as N- and P-limited forests. Along with ANPP, application of the limiting element consistently increased leaf area index (LAI), radiation conversion efficiency (RCE), and foliar and litter nutrient concentrations. Fertilization did not consistently alter N or P retranslocation from senescent leaves at either site, but a comparison with other sites on the chronosequence and with a common-garden study suggests that there is a genetic basis for low foliar and litter nutrients and higher retranslocation at infertile sites vs more fertile sites. N limitation appears to be expressed as limitation to carbon gain, with long leaf lifespans and high leaf mass per area. P limitation results in high P-use efficiency and disproportionally large increases in P uptake after fertilization; a comparison with other studies indicates large investments in acquiring and storing P. Although the general responses of ANPP, LAI, and RCE were similar for the two sites, other aspects of nutrient use differ in relation to the physiological and biogeochemical roles of the two elements. Received 2 June 2000; Accepted 4 April 2001.     

Fertilization with nitrogen and phosphorus increases abundance of non-native species in Hawaiian montane forests

Weexamined the effects of fertilization on the diversity, abundance, and cover ofthe understory plant community of two montane wet forests in Hawaii. One siteoccupies a young substrate, where aboveground tree growth is limited bynitrogen(N), while the other site is on an older substrate, where aboveground treegrowth is limited by phosphorus (P). Both sites contained an on-going,long-termfactorial fertilization experiment in which plots were fertilized semi-annuallywith N, P, or N and P in combination. In each fertilization treatment, wemeasured density of species 0.5 m tall and percent cover ofspecies <0.5 m tall. Fertilization with N reducedspeciesrichness at the young, N-limited site, but none of the nutrient additionsaltered species richness at the older, P-limited site. Species diversity andevenness were not affected by fertilization at either site. At the site withlowN availability, plots fertilized with NP had higher densities of the non-nativeginger Hedychium gardnerianum, and at the site with lowP-availability, densities of the exotic shrub Rubusargutuswere higher in P- and NP-fertilized plots. Other effects included declines inmoss cover with fertilization at both sites, and reduced abundance of nativeseedlings in response to N and NP addition at the N-limited site. Continuedlong-term fertilization could lead to greater dominance of non-native speciesbyencouraging their growth at the expense of native species, which may sufferdecreased recruitment as fertilization and increased abundance of thenon-nativespecies may reduce suitable substrates for seedling establishment.     

Soil Fertility and the Impact of Exotic Invasion on Microbial Communities in Hawaiian Forests

Exotic plant invasions into Hawaiian montane forests have altered many important nutrient cycling processes and pools. Across different ecosystems, researchers are uncovering the mechanisms involved in how invasive plants impact the soil microbial community-the primary mediator of soil nutrient cycling. We examined whether the invasive plant, Hedychium gardnerianum, altered microbial community composition in forests dominated by a native tree, Metrosideros polymorpha, under varying soil nutrient limitations and soil fertility properties within forest plots of the Hawaii long-term substrate age gradient (LSAG). Microbial community lipid analysis revealed that when nutrient limitation (as determined by aboveground net primary production [ANPP]) and soil fertility were taken into account, plant species differentially altered soil microbial community composition. Microbial community characteristics differed under invasive and native plants primarily when N or P was added to the older, highly weathered, P-limited soils. Long-term fertilization with N or P at the P-limited site led to a significant increase in the relative abundance of the saprophytic fungal indicator (18:2 omega 6c,9c) under the invasive plant. In the younger, N-limited soils, plant species played a minor role in influencing soil microbial community composition. We found that the general rhizosphere microbial community structure was determined more by soil fertility than by plant species. This study indicates that although the aggressive invasion of a nutrient-demanding, rapidly decomposable, and invasive plant into Hawaiian forests had large impacts on soil microbial decomposers, relatively little impact occurred on the overall soil microbial community structure. Instead, soil nutrient conditions were more important determinants of the overall microbial community structure within Hawaii's montane forests.     

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.     

Nutrient limitation of phytoplankton communities in Subarctic lakes and ponds in Wapusk National Park, Canada

We determined the limiting nutrient of phytoplankton in 21 lakes and ponds in Wapusk National Park, Canada, using nutrient enrichment bioassays to assess the response of natural phytoplankton communities to nitrogen and phosphorus additions. The goal was to determine whether these Subarctic lakes and ponds were nutrient (N or P) limited, and to improve the ability to predict future impacts of increased nutrient loading associated with climate change. We found that 38% of lakes were not limited by nitrogen or phosphorus, 26% were co-limited by N and P, 26% were P-limited and 13% were N-limited. TN/TP, DIN/TP and NO₃ ⁻/TP ratios from each lake were compared to the Redfield ratio to predict the limiting nutrient; however, these predictors only agreed with 29% of the bioassay results, suggesting that nutrient ratios do not provide a true measure of nutrient limitation within this region. The N-limited lakes had significantly different phytoplankton community composition with more chrysophytes and Anabaena sp. compared to all other lakes. N and P limitation of phytoplankton communities within Wapusk National Park lakes and ponds suggests that increased phytoplankton biomass may result in response to increased nutrient loading associated with environmental change.     

Nutrient limitation in rainforests and cloud forests along a 3,000-m elevation gradient in the Peruvian Andes

We report results from a large-scale nutrient fertilization experiment along a “megadiverse” (154 unique species were included in the study) 3,000-m elevation transect in the Peruvian Andes and adjacent lowland Amazonia. Our objectives were to test if nitrogen (N) and phosphorus (P) limitation shift along this elevation gradient, and to determine how an alleviation of nutrient limitation would manifest in ecosystem changes. Tree height decreased with increasing elevation, but leaf area index (LAI) and diameter at breast height (DBH) did not vary with elevation. Leaf N:P decreased with increasing elevation (from 24 at 200 m to 11 at 3,000 m), suggesting increased N limitation and decreased P limitation with increasing elevation. After 4 years of fertilization (N, P, N + P), plots at the lowland site (200 m) fertilized with N + P showed greater relative growth rates in DBH than did the control plots; no significant differences were evident at the 1,000 m site, and plots fertilized with N at the highest elevation sites (1,500, 3,000 m) showed greater relative growth rates in DBH than did the control plots, again suggesting increased N constraint with elevation. Across elevations in general N fertilization led to an increase in microbial respiration, while P and N + P addition led to an increase in root respiration and corresponding decrease in hyphal respiration. There was no significant canopy response (LAI, leaf nutrients) to fertilization, suggesting that photosynthetic capacity was not N or P limited in these ecosystems. In sum, our study significantly advances ecological understanding of nutrient cycling and ecosystem response in a region where our collective knowledge and data are sparse: we demonstrate N limitation in high elevation tropical montane forests, N and P co-limitation in lowland Amazonia, and a nutrient limitation response manifested not in canopy changes, but rather in stem and belowground changes.     

PHOSPHATE UPTAKE UNDER NITRATE LIMITATION BY SCENEDESMUS SP. AND ITS ECOLOGICAL IMPLICATIONS1

Under nitrogen limitation the phosphate content of Scenedesmus sp. shows little variation regardless of growth rate and the N/P atomic ratio of the medium. P uptake therefore can be calculated as the product of P content and N-dependent growth rate. The maximum rate of P uptake in N limitation is lower by a factor of about 8 than the rate in P limitation. As reported earlier, P uptake by this alga under P limitation is described by the kinetics resembling non-competitive enzyme inhibition, with one or several intracellular P fractions as inhibitors. These fractions include surplus P (water extractable) and inorganic polyphosphate fractions A (acid soluble) and B, C, and D (acid insoluble). In N limitation, the ratios of fractions A, B, C, and D are quite different from the ratios of P limitation at comparable growth rates. The concentrations of polyphosphate fraction A in N-limited cells are much, higher than the levels in P-limited cells, and this fraction becomes more predominant at low growth rates in N limitation. This fraction, if introduced as the inhibitor into the noncompetitive scheme, explains the uptake kinetics in both N- and P-limited cells and the low maximum uptake rate in N limitation. This finding may have two significant ecological implications: (1) A nutrient imbalance which brings about changes in the internal, level or the metabolism, of fraction A would affect P uptake. (2) Nitrogen sufficiency would cause a competitive advantage in P uptake. This advantage would be shared by N₂ fixers and algae with low optimum N/P ratios. In Scenedesmus sp. P limitation switches to N limitation and vice versa when the cell N/P atomic ratio is about 30.     

INTERACTING EFFECTS OF LIGHT AND NUTRIENT LIMITATION ON THE GROWTH RATE OF SYNECHOCOCCUS LINEARIS (CYANOPHYCEAE)1

The blue-green alga Synechococcus linearis (Naeg.) Kom. was grown in P- and N-limited chemostats over a range of potentially limiting irradiances in order to determine the combined effects of light and nutrient limitation on some aspects of the composition and metabolism of this alga. Over a narrow range of low irradiances, simultaneous limitation of growth rate by light and either N or P was shown. This simultaneous limitation of growth rate by a nutrient and a physical factor can be explained by the ability of an increased supply of one to compensate in part for a decreased supply of the other. At all irradiances, the internal concentration of the limiting nutrient increased with increasing dilution rate, and the results could be fitted to the Droop relationship. With decreasing irradiance, the internal concentration of the limiting nutrient increased. There appeared to be little or no effect of light on the minimum internal concentration of P but that of N increased with decreasing light. Both chlorophyll a and biliprotein per unit particulate C increased with increasing dilution rate and decreasing irradiance. The critical N/P ratio increased with decreasing light as the N requirement of N-limited cells increased faster than did the P requirement of P-limited cells. The composition of exponentially growing cells in complete medium varied much less with light. Neither dilution rate nor irradiance during growth had a great effect on saturated rates of P or N uptake or alkaline phosphatase activity. Calculated assimilation ratios increased with light and dilution rate. The role of the flexibility of nutrient composition in adaptation to adverse conditions and the implications of the results for the use of physiological indicators of nutrient status are discussed.     

Nitrogen vs. phosphorus limitation across an ecotonal gradient in a mangrove forest

Mangrove forests are characterized by distinctive tree-height gradientsthat reflect complex spatial, within-stand differences in environmentalfactors,including nutrient dynamics, salinity, and tidal inundation, across narrowgradients. To determine patterns of nutrient limitation and the effects ofnutrient availability on plant growth and within-stand nutrient dynamics, weused a factorial experiment with three nutrient treatment levels (control, N,P)and three zones along a tree-height gradient (fringe, transition, dwarf) onoffshore islands in Belize. Transects were laid out perpendicular to theshoreline across a mangrove forest from a fringe stand along the seaward edge,through a stand of intermediate height, into a dwarf stand in the interior ofthe island. At three sites, three trees were fertilized per zone for 2yr. Although there was spatial variability in response, growth byR. mangle was generally nitrogen (N) -limited in thefringe zone;phosphorus (P) -limited in the dwarf zone; and, N- and/or P-limited in thetransition zone. Phosphorus-resorption efficiency decreased in all three zones,and N-resorption efficiency increased in the dwarf zone in response to Penrichment. The addition of N had no effect on either P or N resorptionefficiencies. Belowground decomposition was increased by P enrichment in allzones, whereas N enrichment had no effect. This study demonstrated thatessential nutrients are not uniformly distributed within mangrove ecosystems;that soil fertility can switch from conditions of N to P limitation acrossnarrow ecotonal gradients; and, that not all ecological processes respondsimilarly to, or are limited by, the same nutrient.     

Nitrogen supply effects on productivity and potential leaf litter decay of Carex species from peatlands differing in nutrient limitation

We investigated the effect of increased N-supply on productivity and potential litter decay rates of Carex species, which are the dominant vascular plant species in peatlands in the Netherlands. We hypothesized that: (1) under conditions of N-limited plant growth, increased N-supply will lead to increased productivity but will not affect C:N ratios of plant litter and potential decay rates of that litter; and (2) under conditions of P-limited plant growth, increased N-supply will not affect productivity but it will lead to lower C:N ratios in plant litter and thereby to a higher potential decay rate of that litter. These hypotheses were tested by fertilization experiments (addition of 10 g N m^-2 year^-1) in peatlands in which plant growth was N-limited and P-limited, respectively. We investigated the effects of fertilization on net C-fixation by plant biomass, N uptake, leaf litter chemistry and potential leaf litter decay. In a P-limited peatland, dominated by Carex lasiocarpa, there was no significant increase of net C-fixation by plant biomass upon enhanced N-supply, although N-uptake had increased significantly compared with the unfertilized control. Due to the N-fertilization the C:N ratio in the plant biomass decreased significantly. Similarly, the C:N ratio of leaf litter produced at the end of the experiment showed a significant decrease upon enhanced N-supply. The potential decay rate of that litter, measured as CO₂-evolution from the litter under aerobic conditions, was significantly increase upon enhanced N-supply. In a N-limited peatland, dominated by C. acutiformis, the net C-fixation by plant biomass increased with increasing N-supply, whereas the increase in N-uptake was not significant. The C:N ratio of both living plant material and of dead leaves did not change in response to N-fertilization. The potential decay rate of the leaf litter was not affected by N-supply. The results agree with our hypotheses. This implies that atmospheric N-deposition may affect the CO₂-sink function of peatlands, but the effect is dependent on the nature of nutrient limitation. In peatlands where plant growth is N-limited, increased N-supply leads to an increase in the net accumulation of C. Under conditions of P-limited plant growth, however, the net C-accumulation will decrease, because productivity is not further increased, whereas the amount of C lost through decomposition of dead organic matter is increased. As plant growth in most terrestrial ecosystems is N-limited, increased N-supply will in most peatlands lead to an increase of net C-accumulation.     

A model analysis of N and P limitation on carbon accumulation in Amazonian secondary forest after alternate land-use abandonment

Productivity and carbon (C) storage in many mature tropical forests are considered phosphorus (P) limited because of advanced soil weathering. However, disturbance can shift limitation away from P and toward nitrogen (N) because of disproportionately large N losses associated with its mobility relative to P in ecosystems. This shift was illustrated by model analyses in which large disturbances including timber extraction and slash-burn were simulated in a P-limited tropical forest. Re-accumulation of ecosystem C during secondary forest growth was initially N-limited, but long term limitation reverted to P. Mechanisms controlling shifts between N and P limitation included: (1) N volatility during slash combustion produced ash that increased soil solution P more than N, (2) a wide N:P ratio in residual fuel and belowground necromass relative to soil organic matter (SOM) N:P produced a simultaneous P sink and N source during decomposition, (3) a supplemental (to aerosol deposition) external N source via biological N fixation. Redistribution of N and P from low C:nutrient SOM to high C:nutrient vegetation was the most important factor contributing to the resilience of ecosystem C accumulation during secondary growth. Resilience was diminished when multiple harvest and re-growth cycles depleted SOM. Phosphorus losses in particular resulted in long-term reductions of C storage capacity because of slow re-supply rates via deposition and the absence of other external sources. Sensitivity analyses limiting the depth of microbially active SOM in soil profiles further illustrated the importance of elements stored in SOM to ecosystem resilience, pointing to a need for better knowledge on the functioning of deeply buried SOM.     

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.