Nitrogen Dynamics in Ice Storm-Damaged Forest Ecosystems: Implications for Nitrogen Limitation Theory

Despite the widely recognized importance of disturbance in accelerating the loss of elements from land, there have been few empirical studies of the effects of natural disturbances on nitrogen (N) dynamics in forest ecosystems. We were provided the unusual opportunity for such study, partly because the intensively monitored watersheds at the Hubbard Brook Experimental Forest (HBEF), New Hampshire, experienced severe canopy damage following an ice storm. Here we report the effects of this disturbance on internal N cycling and loss for watershed 1 (W1) and watershed 6 (W6) at the HBEF and patterns of N loss from nine other severely damaged watersheds across the southern White Mountains. This approach allowed us to test one component of N limitation theory, which suggests that N losses accompanying natural disturbances can lead to the maintenance of N limitation in temperate zone forest ecosystems. Prior to the ice storm, fluxes of nitrate (NO₃ ^–) at the base of W1 and W6 were similar and were much lower than N inputs in atmospheric deposition. Following the ice storm, drainage water NO₃ ^– concentrations increased to levels that were seven to ten times greater than predisturbance values. We observed no significant differences in N mineralization, nitrification, or denitrification between damaged and undamaged areas in the HBEF watersheds, however. This result suggests that elevated NO₃ ^- concentrations were not necessarily due to accelerated rates of N cycling by soil microbes but likely resulted from decreased plant uptake of NO₃ ^-. At the regional scale, we observed high variability in the magnitude of NO₃ ^- losses: while six of the surveyed watersheds showed accelerated rates of NO₃ ^– loss, three did not. Moreover, in contrast to the strong linear relationship between NO₃ ^– loss and crown damage within HBEF watersheds [r ²: (W1 = 0.91, W6 = 0.85)], stream water NO₃ ^– concentrations were weakly related to crown damage (r ² = 0.17) across our regional sites. The efflux of NO₃ ^– associated with the ice storm was slightly higher than values reported for soil freezing and insect defoliation episodes, but was approximately two to ten times lower than NO₃ ^– fluxes associated with forest harvesting. Because over one half of the entire years worth of N deposition was lost following the ice storm, we conclude that catastrophic disturbances contribute synergistically to the maintenance of N limitation and widely observed delays of N saturation in northern, temperate zone forest ecosystems. Present address: Department of Ecology and Evolutionary Biology, Princeton University, Guyot Hall, Princeton, New Jersey 08544, USA.     

Carbon Dioxide Exchange Between an Old-growth Forest and the Atmosphere

Eddy-covariance and biometeorological methods show significant net annual carbon uptake in an old-growth Douglas-fir forest in southwestern Washington, USA. These results contrast with previous assumptions that old-growth forest ecosystems are in carbon equilibrium. The basis for differences between conventional biomass-based carbon sequestration estimates and the biometeorologic estimates are discussed. Annual net ecosystem exchange was comparable to younger ecosystems at the same latitude, as quantified in the AmeriFlux program. Net ecosystem carbon uptake was significantly correlated with photosynthetically active radiation and air temperature, as well as soil moisture and precipitation. Optimum ecosystem photosynthesis occurred at relatively cool temperatures (5°–10°C). Understory and soil carbon exchange always represented a source of carbon to the atmosphere, with a strong seasonal cycle in source strength. Understory and soil carbon exchange showed a Q₁₀ temperature dependence and represented a substantial portion of the ecosystem carbon budget. The period of main carbon uptake and the period of soil and ecosystem respiration are out of phase, however, and driven by different climatic boundary conditions. The period of strongest ecosystem carbon uptake coincides with the lowest observed values of soil and ecosystem respiration. Despite the substantial contribution of soil, the overall strength of the photosynthetic sink resulted in the net annual uptake. The net uptake estimates here included two correction methods, one for advection and the other for low levels of turbulence.     

森林土壤氮素的转化与循环

森林土壤氮素转化与循环在森林生态系统功能中占有极其重要的意义。本文综述了森林土壤氮素转化与循环的研究历程和现状 ;介绍了凋落物的归还、施肥、大气沉降、自生固氮、氨化、硝化、反硝化、植物吸收、NH3 的挥发、NO3 -淋溶等土壤氮素输入、转化和输出的途径和过程 ;最后从研究目标、研究方法、研究对象和研究内容 4个方面归纳了森林土壤氮素转化与循环的发展趋势。     

模拟氮沉降对典型阔叶红松林土壤有机碳和养分的影响简

为探讨氮沉降对典型阔叶红松（Pinus koraiensis）林的影响,从2008年6月~2010年8月进行了人工模拟氮沉降实验,实验分为对照、低N、中N、高N4个处理,每个处理3个重复。所施氮肥为CO(NH₂)₂,以溶液的形式喷施,4个处理浓度分别为0、30、60、120 kg·hm^-2·a^-1。在氮沉降进行1年后,采集各处理0～20、20～40和40～60 cm的土壤样品,测定其土壤有机C、全N、碱解N和速效P、速效K。结果表明：相同处理下,有机C和全N含量随土层的加深均逐渐减少。总体上低、中N处理显著增加了土壤有机C、碱解N和速效K含量,中、高N处理显著降低了土壤速效P含量（P<0.05）,而对全N含量影响不显著（P>0.05）。土壤有机C与土壤全N、碱解N、速效P、速效K之间存在极显著正相关关系（P<0.001）。有机C和土壤养分对氮沉降的响应说明氮沉降在短期内可能影响阔叶红松林土壤碳库积累和土壤肥力水平。     

Element Fluxes and Landscape Position in a Northern Hardwood Forest Watershed Ecosystem

Chemical changes along headwater streams at the Hubbard Brook Experimental Forest in New Hampshire suggest that important differences exist in biogeochemical cycles along an altitudinal gradient within small watershed ecosystems. Using data collected during the period 1982–92, we have constructed element budgets [Ca, Mg, K, Na, Si, Al, dissolved organic carbon (DOC), S, and N] for three subcatchments within watershed 6, a forested watershed last logged around 1917–20. The biogeochemistry of the high-elevation spruce-fir–white birch subcatchment was dominated by processes involving naturally occuring organic compounds. Stream water and soil solutions in this zone had elevated concentrations of organic acidity, DOC, and organically bound monomeric aluminum (Al_o), relative to lower-elevation sites. The middle-elevation subcatchment, dominated by hardwood vegetation, had the greatest net production of inorganic-monomeric aluminum (Al_i), and exhibited net immobilization of DOC and Al_o. The low-elevation subcatchment, also characterized by deciduous vegetation, had the highest rates of net production of base cations (Ca²⁺, Mg²⁺, K⁺, Na⁺) among the subcatchments. Living biomass of trees declined slightly in the spruce-fir–white birch subcatchment during the study period, remained constant in the middle-elevation zone, and increased by 5% in the low-elevation subcatchment. Coupling the corresponding changes in biomass nutrient pools with the geochemical patterns, we observed up to 15-fold differences in the net production of Ca, Mg, K, Na, and Si in soils of the three subcatchments within this 13.2-ha watershed. Release of Ca, Na, and dissolved Si in the highest-elevation subcatchment could be explained by the congruent dissolution of 185 mol ha⁻¹ y⁻¹ of plagioclase feldspar. The rate of plagioclase weathering, based on the net output of Na, increased downslope to 189 and 435 mol ha⁻¹ y⁻¹ in the middle-elevation and low-elevation subcatchments, respectively. However, the dissolution of feldspar in the hardwood subcatchments could account for only 26%–37% of the observed net Ca output. The loss of Ca from soil exchange sites and organic matter is the most likely source of the unexplained net export. Furthermore, this depletion appears to be occurring most rapidly in the lower half of watershed 6. The small watersheds at the Hubbard Brook Experimental Forest occupy a soil catena in which soil depth and soil-water contact time increase downslope. By influencing hydrologic flowpaths and acid neutralization processes, these factors exert an important influence on biogeochemical fluxes within small watersheds, but their influence on forest vigor is less clear. Our results illustrate the sensitivity of watershed-level studies to spatial scale. However, it appears that much of the variation in element fluxes occurs in the first 10–20 ha of drainage area. Received 13 August 1998; accepted 7 September 1999.     

Effects of Soil Texture on Belowground Carbon and Nutrient Storage in a Lowland Amazonian Forest Ecosystem

Soil texture plays a key role in belowground C storage in forest ecosystems and strongly influences nutrient availability and retention, particularly in highly weathered soils. We used field data and the Century ecosystem model to explore the role of soil texture in belowground C storage, nutrient pool sizes, and N fluxes in highly weathered soils in an Amazonian forest ecosystem. Our field results showed that sandy soils stored approximately 113 Mg C ha^-1 to a 1-m depth versus 101 Mg C ha^-1 in clay soils. Coarse root C represented a large and significant ecosystem C pool, amounting to 62% and 48% of the surface soil C pool on sands and clays, respectively, and 34% and 22% of the soil C pool on sands and clays to 1-m depth. The quantity of labile soil P, the soil C:N ratio, and live and dead fine root biomass in the 0–10-cm soil depth decreased along a gradient from sands to clays, whereas the opposite trend was observed for total P, mineral N, potential N mineralization, and denitrification enzyme activity. The Century model was able to predict the observed trends in surface soil C and N in loams and sands but underestimated C and N pools in the sands by approximately 45%. The model predicted that total belowground C (0–20 cm depth) in sands would be approximately half that of the clays, in contrast to the 89% we measured. This discrepancy is likely to be due to an underestimation of the role of belowground C allocation with low litter quality in sands, as well as an overestimation of the role of physical C protection by clays in this ecosystem. Changes in P and water availability had little effect on model outputs, whereas adding N greatly increased soil organic matter pools and productivity, illustrating the need for further integration of model structure and tropical forest biogeochemical cycling. Received 3 March 1999; accepted 27 August 1999.     

Seasonal and annual variation in nitrogen mineralization and nitrification along an elevational gradient in New Mexico

Patterns and amounts of nitrogen loss from disturbed ecosystems vary widely. The mineralization of organic nitrogen to ammonium and then nitrification to nitrate are important processes regulating nitrogen cycling rates and nitrogen losses. Nitrification is a significant process because of the production of the nitrate anion which is easily leached or denitrified. Most studies of these processes do not evaluate their seasonal and yearly variations. This study demonstrates that marked seasonal and yearly variations can occur in these processes in different ecosystems and suggests that nitrogen loss or other system properties correlated with one arbitrarily selected collection can be misleading. Spruce-fir and ponderosa pine ecosystems demonstrated little actual orpotential nitrification. Aspen and mixed conifer ecosystems demonstrated distinct seasonal patterns with increased rates of mineralization and nitrification during spring and summer months and a precipitous decline in both rates coincident with autumn foliage litterfall.The relative availability of soil nitrogen along with the amount of nitrogen circulating annually in litterfall prior to disturbance are useful predictors of the potential for nitrate production and loss following disturbance. However, other controls, including regulation by organic compounds, appear important in determining seasonal and annual variation in actual nitrification rates.     

Nitrogen Oxide Fluxes and Nitrogen Cycling during Postagricultural Succession and Forest Fertilization in the Humid Tropics

The effects of changes in tropical land use on soil emissions of nitrous oxide (N₂O) and nitric oxide (NO) are not well understood. We examined emissions of N₂O and NO and their relationships to land use and forest composition, litterfall, soil nitrogen (N) pools and turnover, soil moisture, and patterns of carbon (C) cycling in a lower montane, subtropical wet region of Puerto Rico. Fluxes of N₂O and NO were measured monthly for over 1 year in old (more than 60 years old) pastures, early- and mid-successional forests previously in pasture, and late-successional forests not known to have been in pasture within the tabonuco (Dacryodes excelsa) forest zone. Additional, though less frequent, measures were also made in an experimentally fertilized tabonuco forest. N₂O fluxes exceeded NO fluxes at all sites, reflecting the consistently wet environment. The fertilized forest had the highest N oxide emissions (22.0 kg N · ha⁻¹· y⁻¹). Among the unfertilized sites, the expected pattern of increasing emissions with stand age did not occur in all cases. The mid-successional forest most dominated by leguminous trees had the highest emissions (9.0 kg N · ha⁻¹· y⁻¹), whereas the mid-successional forest lacking legumes had the lowest emissions (0.09 kg N · ha⁻¹· y⁻¹). N oxide fluxes from late-successional forests were higher than fluxes from pastures. Annual N oxide fluxes correlated positively to leaf litter N, net nitrification, potential nitrification, soil nitrate, and net N mineralization and negatively to leaf litter C:N ratio. Soil ammonium was not related to N oxide emissions. Forests with lower fluxes of N oxides had higher rates of C mineralization than sites with higher N oxide emissions. We conclude that (a) N oxide fluxes were substantial where the availability of inorganic N exceeded the requirements of competing biota; (b) species composition resulting from historical land use or varying successional dynamics played an important role in determining N availability; and (c) the established ecosystem models that predict N oxide loss from positive relationships with soil ammonium may need to be modified. Received 22 February 2000; accepted 6 September 2000.     

Forest nitrogen sinks in large eastern U.S. watersheds: estimates from forest inventory and an ecosystem model

The eastern U.S. receives elevated rates of Ndeposition compared to preindustrial times, yetrelatively little of this N is exported indrainage waters. Net uptake of N into forestbiomass and soils could account for asubstantial portion of the difference between Ndeposition and solution exports. We quantifiedforest N sinks in biomass accumulation andharvest export for 16 large river basins in theeastern U.S. with two separate approaches: (1)using growth data from the USDA ForestService's Forest Inventory and Analysis (FIA)program, and (2) using a model of forestnitrogen cycling (PnET-CN) linked to FIAinformation on forest age-class structure. Themodel was also used to quantify N sinks in soiland dead wood, and nitrate losses below therooting zone. Both methods agreed that netgrowth rates were highest in the relativelyyoung forests on the Schuylkill watershed, andlowest in the cool forests of northern Maine. Across the 16 watersheds, wood export removedan average of 2.7 kg N ha^–1 yr^–1(range: 1–5 kg N ha^–1 yr^–1), andstanding stocks increased by 4.0 kg N ha^–1yr^–1 (–3 to 8 kg N ha^–1 yr^–1). Together, these sinks for N in woody biomassamounted to a mean of 6.7 kg N ha^–1yr^–1 (2–9 kg N ha^–1 yr^–1), or73% (15–115%) of atmospheric N deposition. Modeled rates of net N sinks in dead wood andsoil were small; soils were only a significantnet sink for N during simulations ofreforestation of degraded agricultural sites. Predicted losses of nitrate depended on thecombined effects of N deposition, and bothshort- and long-term effects of disturbance. Linking the model with forest inventoryinformation on age-class structure provided auseful step toward incorporating realisticpatterns of forest disturbance status acrossthe landscape.     

Biomass,Carbon, and Nitrogen Pools in Mexican Tropical Dry Forest Landscapes

Tropical dry forest is the most widely distributed land-cover type in the tropics. As the rate of land-use/land-cover change from forest to pasture or agriculture accelerates worldwide, it is becoming increasingly important to quantify the ecosystem biomass and carbon (C) and nitrogen (N) pools of both intact forests and converted sites. In the central coastal region of México, we sampled total aboveground biomass (TAGB), and the N and C pools of two floodplain forests, three upland dry forests, and four pastures converted from dry forest. We also sampled belowground biomass and soil C and N pools in two sites of each land-cover type. The TAGB of floodplain forests was as high as 416 Mg ha^–1, whereas the TAGB of the dry forest ranged from 94 to 126 Mg ha^–1. The TAGB of pastures derived from dry forest ranged from 20 to 34 Mg ha^–1. Dead wood (standing and downed combined) comprised 27%–29% of the TABG of dry forest but only about 10% in floodplain forest. Root biomass averaged 32.0 Mg ha^–1 in floodplain forest, 17.1 Mg ha^–1 in dry forest, and 5.8 Mg ha^–1 in pasture. Although total root biomass was similar between sites within land-cover types, root distribution varied by depth and by size class. The highest proportion of root biomass occurred in the top 20 cm of soil in all sites. Total aboveground and root C pools, respectively, were 12 and 2.2 Mg ha^–1 in pasture and reached 180 and 12.9 Mg ha^–1 in floodplain forest. Total aboveground and root pools, respectively, were 149 and 47 kg ha^–1 in pasture and reached 2623 and 264 kg ha^–1 in floodplain forest. Soil organic C pools were greater in pastures than in dry forest, but soil N pools were similar when calculated for the same soil depths. Total ecosystem C pools were 306. The Mg ha^–1 in floodplain forest, 141 Mg ha^–1 in dry forest, and 124 Mg ha^–1 in pasture. Soil C comprised 37%–90% of the total ecosystem C, whereas soil N comprised 85%–98% of the total. The N pools lack of a consistent decrease in soil pools caused by land-use change suggests that C and N losses result from the burning of aboveground biomass. We estimate that in México, dry forest landscapes store approximately 2.3 Pg C, which is about equal to the C stored by the evergreen forests of that country (approximately 2.4 Pg C). Potential C emissions to the atmosphere from the burning of biomass in the dry tropical landscapes of México may amount to 708 Tg C, as compared with 569 Tg C from evergreen forests.     

Fifteen Years of Vegetation and Soil Development after Brackish-Water Marsh Creation

Aboveground biomass, macro‐organic matter (MOM), and wetland soil characteristics were measured periodically between 1983 and 1998 in a created brackish‐water marsh and a nearby natural marsh along the Pamlico River estuary, North Carolina to evaluate the development of wetland vegetation and soil dependent functions after marsh creation. Development of aboveground biomass and MOM was dependent on elevation and frequency of tidal inundation. Aboveground biomass of Spartina alterniflora, which occupied low elevations along tidal creeks and was inundated frequently, developed to levels similar to the natural marsh (750 to 1,300 g/m²) within three years after creation. Spartina cynosuroides, which dominated interior areas of the marsh and was flooded less frequently, required 9 years to consistently achieve aboveground biomass equivalent to the natural marsh (600 to 1,560 g/m²). Aboveground biomass of Spartina patens, which was planted at the highest elevations along the terrestrial margin and seldom flooded, never consistently developed aboveground biomass comparable with the natural marsh during the 15 years after marsh creation. MOM (0 to 10 cm) generally developed at the same rate as aboveground biomass. Between 1988 and 1998, soil bulk density decreased and porosity and organic C and N pools increased in the created marsh. Like vegetation, wetland soil development proceeded faster in response to increased inundation, especially in the streamside zone dominated by S. alterniflora. We estimated that in the streamside and interior zones, an additional 30 years (nitrogen) to 90 years (organic C, porosity) are needed for the upper 30 cm of created marsh soil to become equivalent to the natural marsh. Wetland soil characteristics of the S. patens community along upland fringe will take longer to develop, more than 200 years. Development of the benthic invertebrate‐based food web, which depends on organic matter enrichment of the upper 5 to 10 cm of soil, is expected to take less time. Wetland soil characteristics and functions of created irregularly flooded brackish marshes require longer to develop compared with regularly flooded salt marshes because reduced tidal inundation slows wetland vegetation and soil development. The hydrologic regime (regularly vs. irregularly flooded) of the “target” wetland should be considered when setting realistic expectations for success criteria of created and restored wetlands.     

Interactions between Litter Lignin and Nitrogenitter Lignin and Soil Nitrogen Availability during Leaf Litter Decomposition in a Hawaiian Montane Forest

Previous work in a young Hawaiian forest has shown that nitrogen (N) limits aboveground net primary production (ANPP) more strongly than it does decomposition, despite low soil N availability. In this study, I determined whether (a) poor litter C quality (that is, high litter lignin) poses an overriding constraint on decomposition, preventing decomposers from responding to added N, or (b) high N levels inhibit lignin degradation, lessening the effects of added N on decomposition overall. I obtained leaf litter from one species, Metrosideros polymorpha, which dominates a range of sites in the Hawaiian Islands and whose litter lignin concentration declines with decreasing precipitation. Litter from three dry sites had lignin concentrations of 12% or less, whereas litter from two wet sites, including the study site, had lignin concentrations of more than 18%. This litter was deployed 2.5 years in a common site in control plots (receiving no added nutrients) and in N-fertilized plots. Nitrogen fertilization stimulated decomposition of the low-lignin litter types more than that of the high-lignin litter types. However, in contrast to results from temperate forests, N did not inhibit lignin decomposition. Rather, lignin decay increased with added N, suggesting that the small effect of N on decomposition at this site results from limitation of decomposition by poor C quality rather than from N inhibition of lignin decay. Even though ANPP is limited by N, decomposers are strongly limited by C quality. My results suggest that anthropogenic N deposition may increase leaf litter decomposition more in ecosystems characterized by low-lignin litter than in those characterized by high-lignin litter. Received 26 October 1999; accepted 2 June 2000.     

Relationship between leaf characteristics, tree sizes and species distribution along a slope in a warm temperate forest

The relationships between some leaf characteristics, tree size and species distribution were investigated for evergreen tree species along a slope in a warm temperate forest in Japan. Tree species were classified into three groups based on their dominance on the slope: ridge species that were aggregated in an uppersite, valley species that were aggregated in a lowersite, and uniform species that were distributed almost uniformly. The ridge species had a more positive leaf carbon isotope ratio than the valley species, which suggests that the ridge species have larger water use efficiency than the valley species. This may give some advantage to the ridge species over the valley species in the uppersite where water availability would be limited. However, the ridge species had smaller leaf nitrogen content on a mass basis and larger leaf mass per area than the valley species, which suggests that the ridge species had a smaller mass-based leaf photosynthetic capacity than the valley species. This may be disadvantageous to ridge species in the lowersite, because smaller leaf photosynthetic capacities cause lower leaf carbon gain and thus lower growth than the valley species. These differences in leaf characteristics between the ridge and the valley species were affected by microenvironments, and were also partly affected by the difference in species specific responses to microenvironments on the slope.     

Linking Foliar Chemistry to Forest Floor Solid and Solution Phase Organic C and N in Picea abies [L.] Karst Stands in Northern Bohemia

Dissolved organic carbon and nitrogen (DOC and DON) produced in the forest floor are important for ecosystem functions such as microbial metabolism, pedogenesis and pollutant transport. Past work has shown that both DOC and DON production are related to litterfall and standing stocks of C and N in the forest floor. This study, conducted in spring, 2003, investigated variation in forest floor water extractable DOC (WEDOC) and DON (WEDON) and forest floor C and N as a function of lignin, cellulose and N contained in live canopy foliage across eight Picea abies [L.] Karst stands in northern Bohemia. Based on Near Infrared Spectroscopy (NIR) analysis of foliar materials, lignin:N and cellulose:N content of the youngest needles (those produced in 2002) were positively and significantly related to WEDOC (R² = 0.82–0.97; P<0.01) and to forest floor C:N ratio (R = 0.72–0.78; P<0.01). Foliar N was strongly and negatively related to WEDOC and C:N ratio (R = −0.91 and 0.72; P<0.05) among our study sites. WEDON was positively correlated to foliar lignin:N (R = 0.48; P<0.05; n=40). Forest floor C pools were not positively correlated with foliar lignin and cellulose and forest floor N pools were not positively correlated with foliar N. Instead, a significant negative correlation was found between forest floor N pools and foliar cellulose (R=−0.41; P<0.05), and between forest floor C pools and foliar N (R = −0.44; P<0.05). From a remote sensing standpoint, our results are important because canopy reflectance properties are primarily influenced by the most recent foliage, and it was the chemistry of the most recently produced needles that showed a stronger relationship with forest floor WEDOC and C:N ratio suggesting forest floor production of WEDOC can be calculated regionally with remote sensing.     

Biological and Geochemical Sinks for Phosphorus in Soil from a Wet Tropical Forest

In many tropical and volcanic soils, phosphorus (P) availability is strongly influenced by geochemical sorption, which binds P to soil minerals. The aim of this study was to determine whether biological demand or soil sorption strength was the primary control over phosphate availability and retention in a wet tropical soil with high sorption capacity and low P availability. We added ³²PO₄ to soil from the upper two horizons and assessed the ability of soil microbes to immobilize the added phosphate in the presence of strong sorption. We added phosphate at two concentrations, one representing background turnover that adds low concentrations of P to the soil solution, and the other representing nutrient pulses that can add fairly high fluxes of P to the soil solution. Sorption and microbial immobilization were rapid for both concentrations, consuming most added P within 30 min. Thus, little P remained in the soil solution or extractable pools, which are considered more available to plants. Although soil sorption strength was almost identical for the two horizons, immobilization of tracer P was approximately three times greater in the upper horizon, where most microbial activity was located. This result suggests that microbial demand controlled how P was partitioned into biological versus geochemical sinks. Further evidence for microbial control is suggested by the movement of tracer P from the sorbed pool into the microbial pool when demand was stimulated by the addition of carbon (C). We also explored how increased nitrogen (N) and P availability changed P dynamics in this nutrient poor soil. In contrast to the unfertilized soil, long-term N and P fertilization substantially reduced biological control over inorganic P. P fertilization saturated the soils, overwhelming biological P demand, whereas N fertilization appeared to increase available P through reduced P sorption. Where biological demand for P is high and P becomes available in the soil solution, microbes may play an important role in controlling P partitioning into biological versus geochemical sinks even in soils that have high sorption capacity.     

Implications of pre- and post-fertilizing changes in growth and nitrogen pools following multiple applications of nitrogen fertilizer to a Pinus radiata stand over 12 years

Around the world large tracts of forest, previously available for production, have been reserved for nature conservation. This means that wood supply must be met from a reduced land base, including land of low productivity. In addition there are likely to be increasing demands on the use of managed forests for sequestering C as one means of reducing the build up of atmospheric CO2. One way for the forest industry to meet the demands of increased production would be through the use of fertilizers. Substantially increased growth from fertilizer N application has been measured in many cases while, in the northern hemisphere, atmospheric N deposition has been associated with increased growth in some forests.The possibility of using fertilizer N to increase growth, and the effect on the forest and soil, was studied in a research trial area in north-east Tasmania, Australia. Nitrogen was applied for 12 years to a 16 year old P. radiata stand in a low rainfall zone. Growth and foliar nutrient concentrations were measured to age 34 years, to determine change after attainment of steady state growth at age 25 years, and following cessation of fertilizing at age 29 years. Biomass sampling was carried out at ages 25 years, 29 years, and 34 years.Growth at the steady peak rate achieved in the fertilizer plots, of 31 m3 ha-1 periodic annual increment, was accompanied by changes in the N nutrient pools. Surface applied N built up in the surface litter layer while this layer was increasing, from 15 t ha-1 to nearly 50 t ha-1, and fertilizer was being applied between ages 25 and 29 years. Decline of total N in the soil, between ages 25 and 29 years, indicated continued uptake from that source even though fertilizer was being applied. Subsequent increases in soil N, through leaching from the litter layer, followed cessation of fertilizing.Following cessation of annual applications of N fertilizer, growth rates declined by less than 15% and remained substantially (>150%) above the control growth rates for 4 years. Over a 3 year period foliar N concentrations declined steadily, to the same as control plots, although mass of needles was not reduced 5 years after cessation of fertilizer applications and remained significantly above that of the control plots.Without fertilizers, N for new foliage must be supplied from internal recycling and from mineralization of litter and soil organic matter. There were net losses of N from the foliage and the wood, estimated at around 12 kg N ha-1 yr-1, for the fertilized trees for the 5 years following cessation of fertilizing. This indicated retention and recycling within the tree of a considerable proportion of the 104 kg N ha-1 in the foliage, at age 29 years. As rates of mineralization of soil N were likely to be low, this retention of N within the tree was important in maintaining growth rates.In the forest ecosystems investigated, with low initial content of soil organic matter and N, fertilizer N produced improved tree health and substantially increased growth, thus providing the opportunity to manage this forest for increased wood production or C sequestration. Although there was a substantial build up of N and C in the litter on the fertilized treatment, incorporation of N and C into the mineral soil was slow. Over 12 years 1.34 t N ha-1 had been applied, comparable to 40 years of atmospheric deposition at 30 kg N ha-1. There were no signs of detrimental effects from this application.     

The Fate and Retention of Organic and Inorganic <Superscript>15</Superscript>N-Nitrogen in an Old-Growth Forest Soil in Western Oregon

Forests in the American Pacific Northwest receive very little nitrogen (N) through atmospheric deposition; therefore, they can provide insights into how the N cycle functioned in other regions before heavy atmospheric deposition of inorganic N began. Our objectives were to determine (a) if the fate of organic N differed from the fate of inorganic N, (b) the effect that polyphenols have on the fate of organic N, and (c) the effect of season of addition on the fate of N inputs. We traced N added to in situ soil cores as ammonium, organic N, tannin-complexed organic N, and the N₂-fixing lichen Lobaria oregana. Total ¹⁵N recovery was between 74% and 109% for all N additions. Total ¹⁵N recovery did not vary significantly from the first sampling date to the last date. The litter/organic horizon, as a bulk pool, was the largest N retention pool for all forms of N addition. Within the litter/organic horizon, the chloroform-extractable microbial biomass initially accounted for nearly all of the added N from the ammonium additions. On a different time scale, microbial biomass also played a noteworthy role in the retention of N from organic N, tannin-complexed organic N, and Lobaria. Complexing organic matter with tannin appeared to slow N cycling, but it did not significantly change the ultimate distribution of added organic N. Season of N addition had little effect on the retention of added N; however, where differences did occur, spring additions had lower recoveries than autumn additions.     

水稻受精前后胚囊内钙调素分布的变化：免疫金电镜观察

用胶体金免疫电镜技术观察了水稻 (Oryzasativasubsp .japonica)受精前后胚囊内钙调素的分布变化。授粉后 ,卵细胞、助细胞和中央细胞内的钙调素较授粉前均有所增加。中央细胞内钙调素的增加要比卵细胞中约早 2h ,退化助细胞与宿存助细胞之间的钙调素含量无明显差异。授粉到受精期间 ,钙调素的主要分布形式由分散的单颗粒转变为聚集颗粒 ,受精完成后再变为分散的单颗粒形式。胚囊壁及珠心细胞的细胞壁和胞间隙中也观察到钙调素的分布和数量变化。初步讨论了胞内和胞外钙调素在水稻受精与合子形成中的作用。     

Spatial and Temporal Variation of Roots, Arbuscular Mycorrhizal Fungi, and Plant and Soil Nutrients in a Mature Pinot Noir (Vitis vinifera L.) Vineyard in Oregon, USA

Soil and crop management practices may influence biomass growth and yields of cotton (Gossypium hirsutum L.) and sorghum (Sorghum bicolorL.) and sequester significant amount of atmospheric CO₂in plant biomass and underlying soil, thereby helping to mitigate the undesirable effects of global warming. This study examined the effects of three tillage practices [no-till (NT), strip till (ST), and chisel till (CT)], four cover crops [legume (hairy vetch) (Vicia villosa roth), nonlegume (rye) (Secale cerealeL), hairy vetch/rye mixture, and winter weeds orno covercrop], and three N fertilization rates (0, 60–65, and 120–130 kg N ha ^–1) on the amount of C sequestered in cotton lint (lint + seed), sorghum grain, their stalks (stems + leaves) and roots, and underlying soil from 2000 to 2002 in central Georgia, USA. A field experiment was conducted on a Dothan sandy loam (fine-loamy, kaolinitic, thermic, Plinthic Kandiudults). In 2000, C accumulation in cotton lint was greater in NT with rye or vetch/rye mixture but in stalks, it was greater in ST with vetch or vetch/rye mixture than in CT with or without cover crops. Similarly, C accumulation in lint was greater in NT with 60 kg N ha ^–1 but in stalks, it was greater in ST with 60 and 120 kg N ha ^–1 than in CT with 0 kg N ha ^–1. In 2001, C accumulation in sorghum grains and stalks was greater in vetch and vetch/rye mixture with or without N rate than in rye without N rate. In 2002, C accumulation in cotton lint was greater in CT with or without N rate but in stalks, it was greater in ST with 60 and 120 kg N ha ^–1 than in NT with or without N rate. Total C accumulation in the above- and belowground biomass in cotton ranged from 1.7 to 5.6 Mg ha ^–1 and in sorghum ranged from 3.4 to 7.2 Mg ha ^–1. Carbon accumulation in cotton and sorghum roots ranged from 1 to 14% of the total C accumulation in above- and belowground biomass. In NT, soil organic C at 0–10 cm depth was greater in vetch with 0 kg N ha ^–1 or in vetch/rye with 120–130 kg N ha ^–1 than in weeds with 0 and 60 kg N ha ^–1 but at 10–30 cm, it was greater in rye with 120–130 kg N ha ^–1 than in weeds with or without rate. In ST, soil organic C at 0–10 cm was greater in rye with 120–130 kg N ha ^–1 than in rye, vetch, vetch/rye and weeds with 0 and 60 kg N ha ^–1. Soil organic C at 0–10 and 10–30 cm was also greater in NT and ST than in CT. Since 5 to 24% of C accumulation in lint and grain were harvested, C sequestered in cotton and sorghum stalks and roots can be significant in the terrestrial ecosystem and can significantly increase C storage in the soil if these residues are left after lint or grain harvest, thereby helping to mitigate the effects of global warming. Conservation tillage, such as ST, with hairy vetch/rye mixture cover crops and 60–65 kg N ha ^–1 can sustain C accumulation in cotton lint and sorghum grain and increase C storage in the surface soil due to increased C input from crop residues and their reduced incorporation into the soil compared with conventional tillage, such as CT, with no cover crop and N fertilization, thereby maintaining crop yields, improving soil quality, and reducing erosion.     

Net Primary Production and Canopy Nitrogen in a Temperate Forest Landscape: An Analysis Using Imaging Spectroscopy, Modeling and Field Data

Understanding spatial patterns of net primary production (NPP) is central to the study of terrestrial ecosystems, but efforts are frequently hampered by a lack of spatial information regarding factors such as nitrogen availability and site history. Here, we examined the degree to which canopy nitrogen can serve as an indicator of patterns of NPP at the Bartlett Experimental Forest in New Hampshire by linking canopy nitrogen estimates from two high spectral resolution remote sensing instruments with field measurements and an ecosystem model. Predicted NPP across the study area ranged from less than 700 g m⁻² year⁻¹ to greater than 1300 g m⁻² year⁻¹ with a mean of 951 g m⁻² year⁻¹. Spatial patterns corresponded with elevation, species composition and historical forest management, all of which were reflected in patterns of canopy nitrogen. The relationship between production and elevation was nonlinear, with an increase from low- to mid-elevation deciduous stands, followed by a decline in upper-elevation areas dominated by evergreens. This pattern was also evident in field measurements and mirrored an elevational trend in foliar N concentrations. The increase in production from low-to mid-elevation deciduous stands runs counter to the generally accepted pattern for the northeastern U.S. region, and suggests an importance of moisture limitations in lower-elevation forests. Field measurements of foliar N, wood production and leaf litterfall were also used to evaluate sources of error in model estimates and to determine how predictions are affected by different methods of acquiring foliar N input data. The accuracy of predictions generated from remotely sensed foliar N approached that of predictions driven by field-measured foliar N. Predictions based on the more common approach of using aggregated foliar N for individual cover types showed reasonable agreement in terms of the overall mean, but were in poor agreement on a plot-by-plot basis. Collectively, these results suggest that variation in foliar N exerts an important control on landscape-level spatial patterns and can serve as an integrator of other underlying factors that influence forest growth rates.