Chronic nitrogen additions at the Harvard Forest (USA): the first 15 years of a nitrogen saturation experiment

The delivery of reactive forms of nitrogen to the environment through the sum of agricultural and industrial activities now exceeds that from natural processes. Potential negative effects on forests were first proposed in 1985, and in the ensuing two decades, the process of N saturation has become a well-established and generally understood phenomenon, with a few remaining, significant unknowns.One goal of this special section in Forest Ecology and Management is to report in detail on results from the first 15 years of chronic nitrogen additions to two contrasting forest types at the Harvard Forest in Petersham, MA, USA, with special reference to these two central questions. As similar projects elsewhere come to an end, the Harvard Forest experiment remains as one of the few on-going, long-term N saturation experiments. Longevity has enhanced the value of the chronic N experiment, and lead to a series of collaborative studies on plant, soil and microbial responses. Another goal of this special issue is to bring together and present the findings resulting from a diverse set of measurements enabled by the presence of this long-term experiment. A total of 11 papers are presented, in addition to this brief introduction.     

Response of soil microbial biomass and community composition to chronic nitrogen additions at Harvard forest

Soil microbial communities may respond to anthropogenic increases in ecosystem nitrogen (N) availability, and the microbial response may ultimately feed back on ecosystem carbon and N dynamics. We examined the long-term effects of chronic N additions on soil microbes by measuring soil microbial biomass, composition and substrate utilization patterns in pine and hardwood forests at the Harvard Forest Chronic N Amendment Study. Functional and structural genes for important N cycling processes were studied using DNA community profiles. In the O horizon soil of both stands, N additions decreased microbial biomass C as determined by chloroform fumigation-extraction. Utilization of N-containing substrates was lower in N-treated pine soils than in the controls, suggesting that N additions reduced potential microbial activity in the pine stand. Counts of fungi and bacteria as determined by direct microscopy and culture techniques did not show a clear response to N additions. Nitrogen additions, however, strongly influenced microbial community DNA profiles. The ammonia monooxygenase gene (amoA) generally was found in high N-treated soils, but not in control soils. The nifH gene for N₂-fixation was generally found in all soils, but was more difficult to amplify in the pine N-treated soil than the controls, suggesting that the population of N₂-fixers was altered by N additions. The 16S rDNA gene for Nitrobacter was found in all samples, but distinct differences among DNA profiles were observed in the pine B horizon in the control, low N, and high N-treated plots. Our findings indicate that chronic N additions decreased chloroform microbial carbon and altered microbial community profiles. These changes in microbial community structure may be an important component of the response of terrestrial ecosystems to human-accelerated N supply.     

Effect of six years of nitrogen additions on soil chemistry in a subtropical Pleioblastus amarus forest,Southwest China

Soil chemistry influences plant health and carbon storage in forest ecosystems. Increasing nitrogen(N) deposition has potential effect on soil chemistry. We studied N deposition effects on soil chemistry in subtropical Pleioblastus amarus bamboo forest ecosystems. An experiment with four N treatment levels(0, 50, 150,and300 kg N ha~(-1)a~(-1),applied monthly, expressed as CK,LN,MN, HN,respectively) in three replicates. After6 years of N additions, soil base cations, acid-forming cations, exchangeable acidity(EA), organic carbon fractions and nitrogen components were measured in all four seasons. The mean soil pH values in CK,LN,MN and HN were 4.71, 4.62, 4.71, and 4.40, respectively, with a significant difference between CK and HN. Nitrogen additions significantly increased soil exchangeable Al3+,EA, and Al/Ca,and exchangeable Al3+ in HN increased by 70%compared to CK. Soil base cations(Ca~(2+), Mg~(2+), K+, and Na+) did not respond to N additions. Nitrogen treatments significantly increased soil NO3--N but had little effect on soil total nitrogen, particulate organic nitrogen, or NH4~+-N. Nitrogen additions did not affect soil total organic carbon, extractable dissolved organic carbon,incorporated organic carbon, or particulate organic carbon.This study suggests that increasing N deposition could increase soil NO3--N, reduce soil pH, and increase mobilization of Al~(3+). These changes induced by N deposition can impede root grow and function, further may influence soil carbon storage and nutrient cycles in the future.     

Acid deposition effects on forest composition and growth on the Monongahela National Forest,West Virginia

The northern and central Appalachian forests are subject to high levels of atmospheric acid deposition (AD), which has been shown in some forests to negatively impact forest growth as well as predispose the forest system to damage from secondary stresses. The purpose of this study was to evaluate the possible contribution of AD to changes in composition and productivity of the Monongahela National Forest, and to evaluate soil-based indicators of acidification that might be useful for detecting AD-related forest changes. Soils adjacent to 30 Forest Inventory and Analysis (FIA) sites were sampled and analyzed for a suite of acidity indicators. These indicators were correlated with the periodic mean annual volume increment (PMAVI) of the forest stands on FIA plots for the 10-yr period 1989–2000. PMAVI ranged from −9.5 to 11.8 m³ ha⁻¹ yr⁻¹, with lower-than-expected growth (<3 m³ ha⁻¹ yr⁻¹) on two-thirds of the sites. In the surface horizon, effective base saturation, Ca²⁺ concentration, base saturation, K⁺ concentration, Ca/Al molar ratio, and Mg/Al molar ratio, were positively correlated with PMAVI and Fe concentration was negatively correlated with PMAVI (p ≤ 0.1). In the subsurface horizon pH_(w) and effective base saturation were positively correlated and Al³⁻ concentration and K⁺ concentration were negatively correlated with PMAVI. We hypothesized that NO₃-N/NH₄-N ratio would also be correlated with PMAVI, but it was not. Correlations between soil chemical indicators and PMAVI suggest that AD may contribute, in part, to the lower-than-expected forest growth on the Monongahela National Forest.     

Physiological responses of radiata pine roots to soil strength and soil water deficit

We investigated physiological responses of radiata pine (Pinus radiata D. Don) roots to soil strength and soil water deficit by measuring the osmotic potential (Psi(pi)) and yield turgor (Y) in the elongation zone of root segments of seedlings growing (i) in polyethylene glycol 4000-containing rooting solution of different water potentials (Psi(s)) and (ii) in soil of different soil strengths (Q) at the same soil matric potential (Psi(m)). Root elongation rate (Deltal/Deltat) decreased progressively with decreasing Psi(s) and was associated with decreased Psi(pi) and decreased turgor pressure (P). Osmotic adjustment occurred at Psi(s) < -0.2 MPa. Over a range in Psi(s) of -0.01 to -1.0 MPa, Psi(pi) fell 0.3 MPa whereas P fell 0.7 MPa. Mean Psi in the solution experiment was 0.37 MPa and did not differ significantly with Psi(s) (P = 0.10). Root elongation rate decreased exponentially as Q increased from 0 to 3.0 MPa, and was associated with an increase in P of 0.11 MPa as a consequence of Psi(pi) decreasing by the same amount. Mean Y in the soil experiment was 0.49 MPa and did not change significantly with Q (P = 0.87).     

The responses of carbon-and nitrogen-acquiring enzymes to nitrogen and phosphorus additions in two plantations in southern China

In tropical plantations,nutrients such as nitrogen(N) or phosphorus(P) are often applied as management practices.However,the effects of such nutrient additions on topsoil C-and N-acquiring enzymes activities are unclear.In this study,the impacts of fertilization on β-1,4-glucosidase(BG),β-D-cellobiosidase(CBH),β-1,4-xylosidase(BX),β-1,4-N-acetyl-glucosaminidase(NAG),and leucine amino peptidase(LAP) enzymes activities from topsoil and litter layer of two tropical plantations(Acacia auriculiformis and Eucalyptus urophylla) were measured.The results showed that N addition had neutral impact on topsoil enzymes,while significantly elevating the activities of BG,CBH,BX,and NAG in the litter layer.P fertilization had no impacts except for an elevation of NAG in litter sample.There was no interactions found between N and P additions on these enzyme activities.The clearer impacts of N over P fertilization were unexpected because that the study site receives a high rate of atmospheric N deposition,and has low soil P availability.The impact of P fertilization on hydrolytic enzyme activities may be less important compared with that of N.     

Growth, nutrition, photosynthesis and transpiration responses of longleaf pine seedlings to light, water and nitrogen

Early growth and physiology of longleaf pine (Pinus palustris Mill.) seedlings were studied in response to light, water and nitrogen under greenhouse conditions. The experiment was conducted with 1-year-old seedlings grown in 11.3 l pots. The experimental design was a split-plot factorial with two levels (low and high) of each of the factors, replicated in three blocks. The four factorial combinations of water and nitrogen were randomly applied to 15 pots (sub-plots) in each of the light treatment (main plot). Data were collected on survival, root collar diameter (RCD), and height on a monthly basis. Biomass (shoot, root and needle), leaf area index, specific needle area, and needle nutrient (N, P, K, Ca, and Mg) concentrations were determined following final harvest after 16 months. Physiological data (net photosynthesis and transpiration) were collected monthly from March to July during the second growing season.

Height and RCD were significantly influenced by nitrogen and water and by the interaction between them with no apparent effect of light. Seedlings grew 93% taller in the high nitrogen and well watered (HNWW) treatment compared to the low nitrogen and water stressed (LNWS) treatment. Similarly, a significant increase (78%) in RCD was observed for seedlings in the HNWW treatment over the LNWS treatment. Light, along with water and nitrogen, played an important role in seedling biomass growth, especially when water was not limiting. Biomass partitioning (as measured by root:shoot ratio) was affected only by nitrogen and water. Nutrient stress had a greater influence on carbon allocation (69% increase in root:shoot ratio) than water stress (19% increase). Net photosynthesis (P_net) was significantly higher for seedlings in the high resource than in the low resource treatments with significant light×water and nitrogen×water interactions. Transpiration rate was higher (75%) under the WW treatment compared to the WS treatment. Longleaf pine seedlings grown under the LNWW treatment had the lowest foliar nitrogen (0.71%) whereas seedlings in the HNWS treatment had the highest (1.46%). Increasing the availability of light (through larger canopy openings or controlling midstory density) and soil nitrogen (through fertilization) may not result in greater P_net and improved seedling growth unless soil water is not limiting.     

The effect of soil moisture on the response by fungi and bacteria to nitrogen additions for N_2O production

In addition to bacteria,the contribution of fungi to nitrous oxide(N_2 O) production has been recognized but the responses of these two broad and unrelated groups of microorganisms to global environmental changes,atmospheric nitrogen(N) deposition,and precipitation in terms of N_2 O production are unclear.We studied how these two microbial-mediated N_2 O production pathways responded to soil moisture conditions and to N addition in an N-limited temperate forest.Soils from a long-term N addition experiment in Changbai Mountain,northeastern China were incubated.Varied concentrations of cycloheximide and streptomycin,both inhibitors of fungal and bacterial activity,were used to determine the contributions of both to N~2 O production in 66%,98% and 130% water-filled pore spaces(WFPS).The results showed that N_2 O production decreased significantly with increasing cycloheximide concentration whereas streptomycin was only inhibiting N_2 O emissions at 98% and 130% WFPS.The bacterial pathway of N_2 O production in N-addition(N_(add)) soil was significantly more dominant than that in untreated(N_(amb)) soil.The difference in the fungal pathway of N_2 O production between the soil with nitrogen addition and the untreated soil was not significant.Net N_2 O emissions increased with increasing soil moisture,especially at 130% WFPS,a completely flooded condition.Bacteria dominated carbon dioxide(CO_2) and N_2 O emissions in Nadd soil and at 130% WFPS regardless of N status,while fungi dominated CO_2 and N_2 O emissions in soil without N addition at 66% and 98% WFPS.The results suggest that flooded soil is an important source of N_2 O emissions and that bacteria might be better adapted to compete in fertile soils under anoxic conditions.     

Forest products,markets and socio-economics: ten years of pan-European research co-operation at EFI

An insight of the EFI Programme on Forest Products, Markets and Socio-economics should consider that Forest Economics research is generally done by small groups of scientists dispersed around Universities and Research Institutes. EFI has provided the European forest economists a new arena—laboratory, an encounter place, where they can meet, find new opportunities and challenges through co-operation and joint research work. The three main themes of research considered by the Programme have been: (i) market analysis; (ii) forest related industries and regional development; (iii) business management and accounting. An extensive list of publications demonstrate the outstanding quantity of research work undertaken during the last decade. All in all, it has to be acknowledged also that the quality of the research work, undertaken according to the tradition of forest economists, open to both scientific innovation and attention to the real world. The ability to attract human and financial resources in forestry economics research must also be acknowledged. A shortcoming is the poor coverage of certain regions of Europe. However, it should be mentioned that the large spectrum of nationalities involved in the most recent years as well as the launch of Regional Project Centres are now paying major attention to regional issues. Relationships with EFI members have been essential to the Programme: when research has been done by networking with members, an added value can always be seen and the light of the ‘diamond’ can be fully appreciated. Otherwise, it has remained shadowed by regional narrow perspectives missing the main scope of EFI.     

Effects of freeze-thaw on soil nitrogen and phosphorus availability at the Keerqin Sandy Lands, China

A laboratory simulated freeze-thaw was conducted to determine the effects of freeze-thaw on soil nutrient availability in temperate semi-arid regions. Soil samples were collected from sandy soils （0-20 cm） of three typical ecosystems （grassland, Mongolian pine plantation and poplar plantation） in southeastern Keerqin Sandy Lands of China and subjected to freeze-thaw treatment （-12℃ for 10 days, then r 20℃ for 10 days） or incubated at constant temperature （20℃ for 20 days）. Concentrations of the soil NO3^--N, NH4^＋-N, NaHCO3 extractable inorganic P （LPi） and microbial biomass P （MBP） were determined on three occasions： at the start of the incubation, immediate post-thawing and at the 10th day post-thawing. The results showed that soil net nitrification and N mineralization rates at three sites were negatively affected by freeze-thaw treatment, and decreased by 50%-85% as compared to the control, of which the greatest decline occurred in the soil collected from poplar plantation. In contrast, the concentration of soil NH4^＋-N, NaHCO3 extractable inorganic P （LPi） and microbial biomass P were insignificantly influenced by freeze-thaw except that LPi and NH4^＋-N showed a slight increase immediate post-thawing. The effects of freeze-thaw on soil N transformation were related to soil biological processes and the relatively constant available P was ascribed to severe soil aridity.     

Elevation dependent sensitivity of northern hardwoods to Ca addition at Hubbard Brook Experimental Forest,NH, USA

Acidic deposition has caused a depletion of calcium (Ca) in the northeastern forest soils. Wollastonite (Ca silicate) was added to watershed 1 (WS1) at the Hubbard Brook Experimental Forest (HBEF) in 1999 to evaluate its effects on various functions of the HBEF ecosystem. The effects of Ca addition on foliar soluble (extractable in 5% HClO₄) ions, chlorophyll, polyamines, and amino acids were studied in three hardwood species, namely sugar maple, yellow birch, and American beech. We further analyzed these effects in relation to elevation at Ca-supplemented WS1 and reference WS3 watersheds. Foliar soluble Ca increased significantly in all species at mid and high elevations at Ca-supplemented WS1. This was accompanied by increases in soluble P, chlorophyll, and two amino acids, glutamate and glycine. A decrease in known metabolic indicators of physiological stress (i.e., the amino acids, arginine and γ-aminobutyric acid (GABA), and the diamine, putrescine) was also observed. In general, these changes were species-specific and occurred in an elevation dependent manner. Despite an observed increase in Ca at high elevation for all three species, only sugar maple exhibited a decrease in foliar putrescine at this elevation indicating possible remediation from Ca deficiency. At higher elevations of the reference WS3 site, foliar concentrations of Ca and Mg, as well as Ca:Mn ratios were lower, whereas Al, putrescine, spermidine, and GABA were generally higher. Comparison of metabolic data from these three species reinforces the earlier findings that sugar maple is the most sensitive and American beech the least sensitive species to soil Ca limitation. Furthermore, there was an increase in sensitivity with an increase in elevation.     

Effect of six years of nitrogen additions on soil chemistry in a subtropical <Emphasis Type="Italic">Pleioblastus amarus</Emphasis> forest,Southwest China

Soil chemistry influences plant health and carbon storage in forest ecosystems. Increasing nitrogen (N) deposition has potential effect on soil chemistry. We studied N deposition effects on soil chemistry in subtropical Pleioblastus amarus bamboo forest ecosystems. An experiment with four N treatment levels (0, 50, 150, and 300 kg N ha^?1 a^?1, applied monthly, expressed as CK, LN, MN, HN, respectively) in three replicates. After 6 years of N additions, soil base cations, acid-forming cations, exchangeable acidity (EA), organic carbon fractions and nitrogen components were measured in all four seasons. The mean soil pH values in CK, LN, MN and HN were 4.71, 4.62, 4.71, and 4.40, respectively, with a significant difference between CK and HN. Nitrogen additions significantly increased soil exchangeable Al³⁺, EA, and Al/Ca, and exchangeable Al³⁺ in HN increased by 70% compared to CK. Soil base cations (Ca²⁺, Mg²⁺, K⁺, and Na⁺) did not respond to N additions. Nitrogen treatments significantly increased soil NO₃^?–N but had little effect on soil total nitrogen, particulate organic nitrogen, or NH₄⁺–N. Nitrogen additions did not affect soil total organic carbon, extractable dissolved organic carbon, incorporated organic carbon, or particulate organic carbon. This study suggests that increasing N deposition could increase soil NO₃^?–N, reduce soil pH, and increase mobilization of Al³⁺. These changes induced by N deposition can impede root grow and function, further may influence soil carbon storage and nutrient cycles in the future.     

Respiratory responses of Scots pine stems to 5 years of exposure to elevated CO2 concentration and temperature

Stem respiration in 20-year-old Scots pine (Pinus sylvestris L.) trees was examined following 5 years of exposure to ambient conditions (CON), elevated atmospheric carbon dioxide concentration ([CO2]) (ambient + 350 micromol mol(-1), (EC)), elevated temperature (ambient + 2-6 degrees C, (ET)) or a combination of elevated [CO2] and elevated temperature (ECT). Stem respiration varied seasonally regardless of the treatment and displayed a similar trend to temperature, with maximum rates occurring around Day 190 in summer and minimum rates in winter. Respiration normalized to 15 degrees C (R15) was higher in the growing season than in the non-growing season, whereas the temperature coefficient (Q10) was lower in the growing season. Annually averaged R15 was 0.36, 0.43, 0.40 and 0.44 micromol m(-2) s(-1) under CON, EC, ET and ECT conditions, respectively, whereas the corresponding values for total stem respiration were 6.55, 7.69, 7.50 and 7.90 mol m(-2) year(-1). The EC, ET and ECT treatments increased R15 by 18, 11 and 22%, respectively, relative to CON, and increased the modeled annual total stem respiration by 18, 15 and 21%. The increase in modeled annual stem respiration under EC and ECT conditions was caused mainly by higher maintenance respiration (22 and 25%, respectively, whereas the increase in growth respiration was 9 and 12%). Growth respiration was unaltered by ET. The treatments did not significantly affect the respiratory response to stem temperature; the mean Q10 value was 2.04, 2.10, 1.99 and 2.12 in the CON, EC, ET and ECT treatments, respectively. It is suggested that the increase in stem respiration was partly a result of the increased growth rate. We conclude that elevated [CO2] increased the maintenance component of respiration more than the growth component.     

热带林业实验中心森林资源评价分析与发展研究

通过对热带林业实验中心森林资源状况的评价与分析可知,热林中心林地与林木资源丰富,林地生产力较高,但存在林种结构、树种结构不合理以及林地利用不合理的状况。根据森林资源现实状况,采用系统工程的决策分析方法,对热林中心2006—2020年的森林资源发展策略进行了分析与研究。     

森林的价值、效益与森林经营之关系概述

文章阐述了森林价值、效益的概念,相互关系,各类不同森林的价值、效益概述及森林价值最大化之森林经营措施等内容.     

Autumnal changes of sulfur fractions and the ratio of organic sulfur to total nitrogen in leaves and adjacent bark of eastern cottonwood, white basswood and actinorhizal black alder

Autumnal changes in organic-S, sulfate-S, total-S and the ratios of organic-S to total-N and sulfate-S to organic-S were followed in leaves and adjacent bark of actinorhizal (Frankia-nodulated) black alder (Alnus glutinosa (L.) Gaertn.) and eastern cottonwood (Populus deltoides Bartr. ex Marsh.) trees growing on a minespoil site high in extractable soil sulfate, and in black alder and white basswood (Tilia heterophylla Venten.) trees growing on a prairie-derived soil in Illinois. Organic-S concentrations decreased significantly (P < 0.05) during autumn only in foliage of trees growing on the prairie-derived soil where losses of leaf organic-S were 65% for black alder and 100% for white basswood. Leaf sulfate concentrations were relatively stable throughout autumn in white basswood growing on prairie-derived soil and in black alder at both sites. Sulfate-S concentrations in leaves were significantly (P < 0.05) higher in trees at the minespoil site than in trees growing in the prairie-derived soil (5.1 mg g(-1) for the minespoil site and 1.2 mg g(-1) for the prairie-derived soil), and in the non-actinorhizal species during late summer. During the autumn, the ratio of organic-S to total-N doubled in leaves of eastern cottonwood at the minespoil site, but in black alder and white basswood growing on the prarie-derived soil, it decreased by 60 and 74%, respectively. Organic-S concentrations in bark increased more during autumn in species unable to fix atmospheric N(2), than in black alder. The results suggest that patterns of autumnal translocation of leaf S can be site-dependent and that leaf S and leaf N are, at least in part, translocated independently in the fall. Black alder and eastern cottonwood seemed to incorporate sulfate-S readily into organic substances in leaves when grown in soils with a high sulfate content.     

A meta-analysis of the effects of nitrogen additions on base cations: Implications for plants, soils, and streams

The dominant base cations (BC; i.e., Ca²⁺, Mg²⁺, K⁺, and Na⁺) are important in buffering soil and water acidity in both terrestrial and aquatic ecosystems. Ca²⁺, Mg²⁺, and K⁺ are also important in many plant physiological functions. Because BC availability is affected by changes in the nitrogen (N) cycle, we conducted a meta-analysis of previously published data to determine if N fertilization alters the availability of BC in terrestrial and stream ecosystems across biomes. We include data from 107 independent studies published in 62 different articles, taking a holistic perspective on BC by examining their responses to added N in plant foliage, bulk soil, soil solution, and stream water. Our results suggest N fertilization may accelerate BC loss from terrestrial ecosystems over time periods less than five years. We found that N additions resulted in an overall 24% decrease in the availability of exchangeable Ca²⁺, Mg²⁺, and K⁺ in the bulk soil of boreal forest, temperate forest, and grassland biomes. Collectively, responses of BC in boreal forest, temperate forest, tropical forest, and grassland biomes increased following N fertilization by about 71% in soil solution and 48% in stream waters. Additionally, BC responses in foliage decreased in boreal forest and temperate forest biomes following N additions over time periods less than five years, but there were no significant changes over longer time periods. Despite large short-term shifts in BC responses following N additions, we did not find evidence of widespread negative impacts on ecosystems over time periods greater than five years. This analysis suggests effects of N addition on the availability of exchangeable BC may diminish over time. Although the effects on BC can be substantial over periods less than five years, there is little available evidence that N fertilization has had large-scale detrimental effects on the availability of BC needed for plant growth within terrestrial or aquatic ecosystems.     

Responses of secondary chemicals in sugar maple (Acer saccharum) seedlings to UV-B, springtime warming and nitrogen additions

Anticipated effects of climate change involve complex interactions in the field. To assess the effects of springtime warming, ambient ultraviolet-B radiation (UV-B) and nitrogen fertilization on the foliar chemistry and herbivore activity of native sugar maple (Acer saccharum Marsh.) seedlings, we carried out a field experiment for 2 years at two sugar maple forests growing on soils of contrasting acidity. At the Oliver site, soils are derived from a strongly calcareous till, whereas the naturally acidic soils and base-poor soils of the Haliburton site are derived from the largely granitic Precambrian Shield. At both sites, removal of ambient UV-B led to increases in chlorogenic acid and some flavonoids and reduced herbivore activity. At Haliburton, ammonium nitrate fertilization led to further increases in foliar manganese (Mn), whereas at Oliver there were no such changes. Nitrogen additions led to decreases in the concentrations of some flavonoids at both sites, but seedlings at Oliver had significantly higher concentrations of flavonoids and chlorogenic acid than seedlings at Haliburton. We suggest that this could be associated with increased mobilization of Mn due to increased soil acidity, which interferes with the role of calcium (Ca) in the phenolic biosynthetic pathway. It appears that the composition of the forest soil governs the response of seedlings when they are exposed to abiotic stressors.     

Variable responses of the depth of tree nitrogen uptake to pruning and competition

Trees in cropped fields may improve nitrogen (N) use efficiency by intercepting leached N, but crop yield will be reduced if the trees compete strongly with crops for N. Ideal trees for intercropping will take up N from deeper soil layers not accessed by the crop species. Spatiotemporal aspects of tree nitrogen capture niches were investigated within a hedgerow intercropping system by placing 15N at three depths and monitoring 15N uptake by trees pruned either 25 or 4 days before application of 15N. Trees with contrasting rooting patterns (Gliricidia sepium L. and Peltophorum dasyrrachis (Miq.) Kurz) were grown in mixed hedgerows and intercropped with maize (Zea mays L.). Neither species showed significant N uptake during the 5-14 days after pruning, even though some shoot regrowth occurred during this time. Mean topsoil (0-5 cm depth) root length density of G. sepium was 520% greater than that of P. dasyrrachis, whereas total root length (0-100 cm depth) of G. sepium was only 450% greater. On average, G. sepium recovered 15 times as much 15N as P. dasyrrachis, following application of 15N at 5 cm depth, but the two species recovered a similar amount following application at 80 cm depth, suggesting that P. dasyrrachis had better niche complementarity with shallow rooting crops. However, both species showed strong plasticity in vertical N uptake pattern in response to competition from establishing maize plants. The species differed in their response: N uptake activity by G. sepium shifted down the soil profile in response to increasing competition from a growing maize crop (uptake from 80 cm depth changed from 4 to 9% of uptake from 5 cm depth), whereas N uptake by P. dasyrrachis became relatively shallow (uptake from 80 cm depth changed from 305 to 25% of uptake from 5 cm depth). Niche avoidance and increased competitiveness within the topsoil represent alternative responses to competition. The response displayed may be related to soil fertility in the species' natural habitats.     

Photosynthetic responses of forest understory tree species to long-term exposure to elevated carbon dioxide concentration at the Duke Forest FACE experiment

We examined the photosynthetic responses of four species of saplings growing in the understory of the Duke Forest FACE experiment during the seventh year of exposure to elevated CO2 concentration ([CO2]). Saplings of these same species were measured in the first year of the Duke Forest FACE experiment and at that time showed only seasonal fluctuations in acclimation of photosynthesis to elevated [CO2]. Based on observations from the Duke Forest FACE experiment, we hypothesized that after seven years of exposure to elevated [CO2] significant photosynthetic down-regulation would be observed in these tree species. To test our hypothesis, photosynthetic CO2-response and light-response curves, along with chlorophyll fluorescence, chlorophyll concentration and foliar N were measured twice during the summer of 2003. Exposure to elevated [CO2] continued to increase photosynthesis in all species measured after seven years of treatment with the greatest photosynthetic increase observed near saturating irradiances. In all species, elevated [CO2] increased electron transport efficiency but did not significantly alter carboxylation efficiency. Quantum yield estimated by light curves, chlorophyll concentration, and foliar nitrogen concentrations were unaffected by elevated [CO2]. Contrary to our hypothesis, there is little evidence of progressive N limitation of leaf-level processes in these understory tree species after seven years of exposure to elevated [CO2] in the Duke Forest FACE experiment.