Belowground carbon allocation in unfertilized and fertilized red pine plantations in northern Wisconsin

We estimated carbon allocation to belowground processes in unfertilized and fertilized red pine (Pinus resinosa Ait.) plantations in northern Wisconsin to determine how soil fertility affects belowground allocation patterns. We used soil CO(2) efflux and litterfall measurements to estimate total belowground carbon allocation (root production and root respiration) by the carbon balance method, established root-free trenched plots to examine treatment effects on microbial respiration, estimated fine root production by sequential coring, and developed allometric equations to estimate coarse root production. Fine root production ranged from 150 to 284 g m(-2) year(-1) and was significantly lower for fertilized plots than for unfertilized plots. Coarse root production ranged from 60 to 90 g m(-2) year(-1) and was significantly lower for fertilized plots than for unfertilized plots. Annual soil CO(2) fluxes ranged from 331 to 541 g C m(-2) year(-1) and were significantly lower for fertilized plots than for unfertilized plots. Annual foliage litterfall ranged from 110 to 187 g C m(-2) year(-1) and was significantly greater for fertilized plots than for unfertilized plots. Total belowground carbon allocation ranged from 188 to 395 g C m(-2) year(-1) and was significantly lower for fertilized than for unfertilized plots. Annual soil CO(2) flux was lower for trenched plots than for untrenched plots but did not differ between fertilized and unfertilized trenched plots. Collectively, these independent estimates suggest that fertilization decreased the relative allocation of carbon belowground.     

Effects of carbon and nitrogen amendment on soil carbon and nitrogen mineralization in volcanic immature soil in southern Kyushu,Japan

Soil N mineralization is affected by microbial biomass and respiration, which are limited by available C and N. To examine the relationship between C and N for soil microbial dynamics and N dynamics, we conducted long-term laboratory incubation (150 days) after C and N amendment and measured changes in C and N mineralization, microbial biomass C, and dissolved C and N throughout the incubation period. The study soil was volcanic immature soil from the southern part of Japan, which contains lower C and N compared with other Japanese forest soils. Despite this, the area is covered by well-developed natural and plantation forests. Carbon amendment resulted in an increase in both microbial biomass and respiration, and net N mineralization decreased, probably due to increasing microbial immobilization. In contrast, N amendment resulted in a decrease in microbial respiration and an increase in net N mineralization, possibly due to decreased immobilization by microbes. Amendment of both C and N simultaneously did not affect microbial biomass and respiration, although net N mineralization was slightly increased. The results suggested that inhibitory effect on microbial respiration by N amendment should be reduced if carbon availability is higher. Thus, soil available C may limit microbial biomass and respiration in this volcanic immature soil. Even in immature soil where C and N substrate is low, soil C, such as plant root exudates and materials from above- and belowground dead organisms, might help to maintain microbial activity and N mineralization in this study site.     

Short-term soil respiration and nitrogen immobilization response to nitrogen applications in control and nitrogen-enriched temperate forests

Forest stands at the Harvard Forest, Petersham, MA, receiving experimentally elevated N inputs have shown greatly increased N leaching loss yet still retain over 70% of the added N in soils, presumably in organic form. Whether microbial or abiotic mechanisms are responsible for the high N retention is not well understood. We monitored soil respiration and extractable NH₄-N and NO₃-N following monthly applications of NH₄NO₃ to a hardwood forest and a pine plantation during the fifth year of chronic fertilizer applications (15 g N as NH₄NO₃ m⁻² per year). We hypothesized that individual N applications would increase short-term soil respiration (within 1 month) in previously unamended and N-limited soil, but that little or no increase would occur following N applications to chronically N-amended soils, assumed to be carbon-limited to some degree after 5 years of N additions. Short-term soil respiration did not increase after N additions in either the chronically amended or previously untreated soils except for one instance in the latter. However, extractable N levels in both previously unamended plots returned to pre-application levels within 2 weeks of the N addition. This rapid disappearance of the applied N suggests microbial immobilization, but in all but one instance there was no accompanying CO₂ efflux increase indicating increased microbial biomass growth. A model of N immobilization through microbial biomass production, driven by the observed apparent net N immobilization, predicted soil CO₂ efflux 4–17 times greater than measured rates. Microbial biomass production does not appear to be the mechanism by which the fertilizer N immobilization occurred, according to our assumptions about microbial C:N ratios and carbon use efficiency. Hardwood stand average soil respiration rates over the study period were significantly higher in the previously unamended plot than in the control, and the control and chronically N-treated plot respiration rates were similar. Soil respiration rates for all pine stand treatments were similar. These results are insufficient to support our hypotheses concerning carbon versus nitrogen limitation in these soils. Our results, along with evidence from other studies, suggest that abiotic mechanisms play a role in the high retention of long-term N additions in these soils.     

Forest soil carbon cycle under elevated CO2 - a case of increased throughput?

Forest soils account for a large part of the stable carbon poolheld in terrestrial ecosystems. Future levels of atmosphericCO₂ are likely to increase C input into the soils through increasedabove- and below-ground production of forests. This increasedinput will result in greater sequestration of C only if theadditional C enters stable pools. In this review, we comparecurrent observations from four large-scale Free Air FACE Enrichment(FACE) experiments on forest ecosystems (EuroFACE, Aspen-FACE,Duke FACE and ORNL-FACE) and consider their predictive powerfor long-term C sequestration. At all sites, FACE increasedfine root biomass, and in most cases higher fine root turnoverresulted in higher C input into soil via root necromass. However,at all sites, soil CO₂ efflux also increased in excess of theincreased root necromass inputs. A mass balance calculationsuggests that a large part of the stimulation of soil CO₂ effluxmay be due to increased root respiration. Given the durationof these experiments compared with the life cycle of a forestand the complexity of processes involved, it is not yet possibleto predict whether elevated CO₂ will result in increased C storagein forest soil.     

Thinning reduces soil carbon dioxide but not methane flux from southwestern USA ponderosa pine forests

Forest soils are important components of the global carbon cycle because they both store and release carbon. Carbon dioxide is released from soil to the atmosphere as a result of plant root and microbial respiration. Additionally, soils in dry forests are often sinks of methane from the atmosphere. Both carbon dioxide and methane are greenhouse gases whose increasing concentration in the atmosphere contributes to climate warming. Thinning treatments are being implemented in ponderosa pine forests across the southwestern United States to restore historic forest structure and reduce the risk of severe wildfire. This study addresses how thinning alters fluxes of carbon dioxide and methane in ponderosa pine forest soils within one year of management and examines mechanisms of change. Carbon dioxide and methane fluxes, soil temperature, soil water content, forest floor mass, root mass, understory plant biomass, and soil microbial biomass carbon were measured before and after the implementation of a thinning and in an unthinned forest. Carbon dioxide efflux from soil decreased as a result of thinning in two of three summer months. Average summer carbon dioxide efflux declined by an average of 34 mg C m⁻² hr⁻¹ in the first year after thinning. Methane oxidation did not change in response to thinning. Thinning had no significant short-term effect on total forest floor mass, total root biomass, or microbial biomass carbon in the mineral soil. Understory plant biomass increased after thinning. Thinning increased carbon available for decomposition by killing tree roots, but our results suggest that thinning reduced carbon dioxide emissions from the soil because the reduction in belowground autotrophic respiration was larger than the stimulation of heterotrophic respiration. Methane oxidation was probably not affected by thinning because thinning did not alter the forest floor mass enough to affect methane diffusion from the atmosphere into the soil.     

Effects of elevated CO2 concentrations on soil microbial respiration and root/rhizosphere respiration in forest soils

The two main components of soil respiration, i.e., root/rhizosphere and microbial respiration, respond differently to elevated atmospheric CO₂ concentrations both in mechanism and sensitivity because they have different substrates derived from plant and soil organic matter, respectively. To model the carbon cycle and predict the carbon source/sink of forest ecosystems, we must first understand the relative contributions of root/rhizosphere and microbial respiration to total soil respiration under elevated CO₂ concentrations. Root/rhizosphere and soil microbial respiration have been shown to increase, decrease and remain unchanged under elevated CO₂ concentrations. A significantly positive relationship between root biomass and root/rhizosphere respiration has been found. Fine roots respond more strongly to elevated CO₂ concentrations than coarse roots. Evidence suggests that soil microbial respiration is highly variable and uncertain under elevated CO₂ concentrations. Microbial biomass and activity are related or unrelated to rates of microbial respiration. Because substrate availability drives microbial metabolism in soils, it is likely that much of the variability in microbial respiration results from differences in the response of root growth to elevated CO₂ concentrations and subsequent changes in substrate production. Biotic and abiotic factors affecting soil respiration were found to affect both root/rhizosphere and microbial respiration. __________ Translated from Journal of Plant Ecology, 2007, 31(3): 386–393 [译自: 植物生态学报]     

Foliage, fine-root, woody-tissue and stand respiration in Pinus radiata in relation to nitrogen status

We measured respiration of 20-year-old Pinus radiata D. Don trees growing in control (C), irrigated (I), and irrigated + fertilized (IL) stands in the Biology of Forest Growth experimental plantation near Canberra, Australia. Respiration was measured on fully expanded foliage, live branches, boles, and fine and coarse roots to determine the relationship between CO(2) efflux, tissue temperature, and biomass or nitrogen (N) content of individual tissues. Efflux of CO(2) from foliage (dark respiration at night) and fine roots was linearly related to biomass and N content, but N was a better predictor of CO(2) efflux than biomass. Respiration (assumed to be maintenance) per unit N at 15 degrees C and a CO(2) concentration of 400 micro mol mol(-1) was 1.71 micro mol s(-1) mol(-1) N for foliage and 11.2 micro mol s(-1) mol(-1) N for fine roots. Efflux of CO(2) from stems, coarse roots and branches was linearly related to sapwood volume (stems) or total volume (branches + coarse roots) and growth, with rates for maintenance respiration at 15 degrees C ranging from 18 to 104 micro mol m(-3) s(-1). Among woody components, branches in the upper canopy and small diameter coarse roots had the highest respiration rates. Stem maintenance respiration per unit sapwood volume did not differ among treatments. Annual C flux was estimated by summing (1) dry matter production and respiration of aboveground components, (2) annual soil CO(2) efflux minus aboveground litterfall, and (3) the annual increment in coarse root biomass. Annual C flux was 24.4, 25.3 and 34.4 Mg ha(-1) year(-1) for the C, I and IL treatments, respectively. Total belowground C allocation, estimated as the sum of (2) and (3) above, was equal to the sum of root respiration and estimated root production in the IL treatment, whereas in the nutrient-limited C and I treatments, total belowground C allocation was greater than the sum of root respiration and estimated root production, suggesting higher fine root turnover or increased allocation to mycorrhizae and root exudation. Carbon use efficiency, the ratio of net primary production to assimilation, was similar among treatments for aboveground tissues (0.43-0.50). Therefore, the proportion of assimilation used for construction and maintenance respiration on an annual basis was also similar among treatments.     

Seasonal variation of soil respiration rates in a secondary forest and agroforestry systems

Agroforestry systems are widely practiced in tropical forests to recover degraded and deforested areas and also to balance the global carbon budget. However, our understanding of difference in soil respiration rates between agroforestry and natural forest systems is very limited. This study compared the seasonal variations in soil respiration rates in relation to fine root biomass, microbial biomass, and soil organic carbon between a secondary forest and two agroforestry systems dominated by Gmelina arborea and Dipterocarps in the Philippines during the dry and the wet seasons. The secondary forest had significantly higher (p < 0.05) soil respiration rate, fine root biomass and soil organic matter than the agroforestry systems in the dry season. However, in the wet season, soil respiration and soil organic matter in the G. arborea dominated agroforestry system were as high as in the secondary forest. Whereas soil respiration was generally higher in the wet than in the dry season, there were no differences in fine root biomass, microbial biomass and soil organic matter between the two seasons. Soil respiration rate correlated positively and significantly with fine root biomass, microbial biomass, and soil organic C in all three sites. The results of this study indicate, to some degree, that different land use management practices have different effects on fine root biomass, microbial biomass and soil organic C which may affect soil respiration as well. Therefore, when introducing agroforestry system, a proper choice of species and management techniques which are similar to natural forest is recommended.     

中亚热带天然林改造成人工林后土壤呼吸的变化特征

【目的】研究中亚热带常绿阔叶林(天然林)改造成人工林后土壤碳排放量的变化及主要影响因子,为评估森林类型转换对土壤碳排放的影响提供科学依据。【方法】在福建农林大学西芹教学林场的常绿阔叶林及由其改造而来的38年生闽楠人工林与35年生杉木人工林中分别设置4块20 m×20 m样地,利用Li-8100土壤碳通量观测系统于2014年9月—2016年9月进行定点观测,并同期观测土壤温度、含水量、有机碳含量(SOC)、微生物生物量碳含量(MBC)、可溶性有机碳含量(DOC)、0～20 cm土层细根生物量和年凋落物量及凋落物碳氮比(C/N)。【结果】常绿阔叶林改造成闽楠(38年后)和杉木人工林(35年后),年均土壤碳排放通量由16. 22显著降为12. 71和4. 83 tC·hm^-2a^-1,分别减少21. 60%和70. 20%;各林分类型的土壤呼吸温度敏感性Q¹⁰值表现为常绿阔叶林(1. 97)<闽楠人工林(2. 03)<杉木人工林(2. 91),转换为杉木人工林后,Q¹⁰值显著升高(P<0. 05);土壤温度能分别解释常绿阔叶林、闽楠人工林与杉木人工林土壤呼吸速率变化的89. 70%、88. 50%和87. 90%,土壤呼吸速率和土壤含水量相关不显著(P>0. 05);土壤呼吸速率和SOC、MBC、DOC、年凋落物量及0～20 cm土层细根生物量均极显著正相关(P<0. 01);土壤呼吸温度敏感性指数Q¹⁰值和凋落物C/N极显著正相关(P<0. 01),而与年均土壤呼吸速率及MBC极显著负相关(P<0. 01);进一步分析发现土壤MBC和SOC含量是影响土壤呼吸速率的2个最重要因子,而凋落物C/N在影响土壤呼吸温度敏感性中的贡献最大。【结论】中亚热带地区常绿阔叶林改造成闽楠(38年)或杉木(35年)人工林后,土壤碳排放通量显著降低。林分类型转换后树种组成和林分结构发生改变,凋落物数量、质量及细根生物量显著降低,土壤SOC和MBC含量显著下降可共同导致土壤呼吸通量的下降。土壤温度是3种林分类型土壤呼吸季节变化的主导因素,而土壤总有机碳库和土壤微生物量碳库的差异是不同林分之间土壤呼吸差异的主导因素,凋落物C/N对土壤呼吸的Q¹⁰影响最大。为提高模型预测森林类型转换影响土壤碳排放的精度,应综合考虑土壤有机碳库、易变性有机碳库及底物质量的变化。     

Belowground carbon pools and processes in different age stands of Douglas-fir

Forest floor material and soil organic matter may act as both a source and a sink in global CO2 cycles. Thus, the ecosystem processes controlling these pools are central to understanding the transfers of carbon (C) between the atmosphere and terrestrial systems. To examine these ecosystem processes, the effect of stand age on temporal carbon source-sink relationships was examined in 20-year-old, 40-year-old and old-growth stands of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) in the Cascade Mountains of south-central Washington State. Belowground C and nitrogen (N) storage and soil respiration were measured. In addition, nylon mesh bags containing homogenized soils from each site were buried at the respective sites to quantify root ingrowth and potential C sequestration and loss. The sites supporting the 20- and 40-year-old stands had soil C stores reflecting the C contributions from logging residue, coarse woody debris and stumps left after harvest. Because the N-fixer red alder (Alnus rubra Bong.) comprised 33% of the 40-year-old stand, this site had significantly greater concentrations and pools of N in the forest floor than sites without red alder. This N-rich site had consistently lower soil CO2 efflux rates during the growing season than the sites supporting the 20-year-old and old-growth stands. Estimated annual soil C efflux was 1367, 883 and 1194 g m-2 for the sites supporting the 20-, 40- and old-growth stands, respectively. These values are higher than previously reported values. Root ingrowth was significantly less in the 40-year-old stand than in the 20-year-old stand, and both young stands showed markedly less fine root growth than the old-growth stand. At the sites supporting the young stands, C and N were lost from the soil bags, whereas there was an increase in C and N in the soil bags at the site supporting the old-growth stand. The fine root growth and soil respiration data support the hypothesis that belowground C allocation decreases with increasing fertility. Quantification of the source-sink relationship of soil C at the three stands based on litterfall, relative root ingrowth and soil respiration measurements was compromised because of significant CO2 flux from decaying organic matter in the young stands.     

Kalmia removal increases nutrient supply and growth of black spruce seedlings: An effect fertilizer cannot emulate

In Canada's eastern boreal forest, the stagnant growth of black spruce (Picea mariana (Mill.) BSP) seedlings is often observed in the presence of ericaceous shrubs such as Kalmia angustifolia L. Many mechanisms, including allelopathic interference, reduced soil N mineralization, soil enzymes inhibition, and direct resource competition have been proposed to explain poor spruce growth in the presence of Kalmia. However, the relative importance of direct competition versus indirect interference remains unclear. Our objective was thus to adequately isolate the “Kalmia effect” from other growth-limiting factors and to determine if removal of Kalmia also resulted in fundamental changes in the biochemical properties of the forest floor. By sampling plots established in 2000, we evaluated how Kalmia eradication and spot fertilization influenced soil nutrient availability, N mineralization rates, microbial basal respiration and biomass, as well as planted black spruce seedling growth, dimensions, and foliar nutrient concentrations 6 years later. We measured higher extractable-P, mineralizable-N, seedling dimensions and growth rates, as well as lower extractable-K, total-K, basal respiration and microbial biomass, in plots without Kalmia than in those where Kalmia had been maintained from 2000 to 2006. Our results thus confirmed that Kalmia eradication over 6 years not only improved the growth and nutrition of black spruce seedlings, but also resulted in fundamental changes in the biochemical properties of the forest floor. We demonstrated that along with direct competition for resources, Kalmia interferes indirectly with black spruce by modifying nutrient cycling and energy fluxes in soil. Higher indices of available C in plots with Kalmia corroborates that Kalmia tannins or rhizodeposition may reduce N mineralization by stimulating microbial immobilization, a relation that however needs to be confirmed with longer term laboratory incubations. Our results indicated that although it had a positive influence on seedling growth, the fertilization effect was confined to the first few years following treatment application, and failed to influence soil processes as did Kalmia eradication. Further monitoring will indicate if the increased litterfall in fertilized plots will eventually initiate a second wave of fertilizer-induced changes to soil processes, as observed in other ecosystems.     

Soil and microbial respiration in a loblolly pine plantation in response to seven years of irrigation and fertilization

Because soil CO₂ efflux or soil respiration (R_S) is the major component of forest carbon fluxes, the effects of forest management on R_S and microbial biomass carbon (C), microbial respiration (R_H), microbial activity and fine root biomass were studied over two years in a loblolly pine (Pinus taeda L.) plantation located near Aiken, SC. Stands were six-years-old at the beginning of the study and were subjected to irrigation (no irrigation versus irrigation) and fertilization (no fertilization versus fertilization) treatments since planting. Soil respiration ranged from 2 to 6 μmol m⁻² s⁻¹ and was strongly and linearly related to soil temperature. Soil moisture and C inputs to the soil (coarse woody debris and litter mass) which may influence R_H were significantly but only weakly related to R_S. No interaction effects between irrigation and fertilization were observed for R_S and microbial variables. Irrigation increased R_S, fine root mass and microbial biomass C. In contrast, fertilization increased R_H, microbial biomass C and microbial activity but reduced fine root biomass and had no influence on R_S. Predicted annual soil C efflux ranged from 8.8 to 10.7 Mg C ha⁻¹ year⁻¹ and was lower than net primary productivity (NPP) in all stands except the non-fertilized treatment. The influence of forest management on R_S was small or insignificant relative to biomass accumulation suggesting that NPP controls the transition between a carbon source and sink in rapidly growing pine systems.     

Carbon and nitrogen cycling immediately following bark beetle outbreaks in southwestern ponderosa pine forests

Bark beetle infestation is a well-known cause of historical low-level disturbance in southwestern ponderosa pine forests, but recent fire exclusion and increased tree densities have enabled large-scale bark beetle outbreaks with unknown consequences for ecosystem function. Uninfested and beetle-infested plots (n = 10 pairs of plots on two aspects) of ponderosa pine were compared over one growing season in the Sierra Ancha Experimental Forest, AZ to determine whether infestation was correlated with differences in carbon (C) and nitrogen (N) pools and fluxes in aboveground biomass and soils. Infested plots had at least 80% of the overstory ponderosa pine trees attacked by bark beetles within 2 years of our measurements. Both uninfested and infested plots stored ∼9 kg C m⁻² in aboveground tree biomass, but infested plots held 60% of this aboveground tree biomass in dead trees, compared to 5% in uninfested plots. We hypothesized that decreased belowground C allocation following beetle-induced tree mortality would alter soil respiration rates, but this hypothesis was not supported; throughout the growing season, soil respiration in infested plots was similar to uninfested plots. In contrast, several results supported the hypothesis that premature needlefall from infested trees provided a pulse of low C:N needlefall that altered soil N cycling. The C:N mass ratio of pine needlefall in infested plots (∼45) was lower than uninfested plots (∼95) throughout the growing season. Mineral soils from infested plots had greater laboratory net nitrification rates and field resin bag ammonium accumulation than uninfested plots. As bark beetle outbreaks become increasingly prevalent in western landscapes, longer-term biogeochemical studies on interactions with other disturbances (e.g. fire, harvesting, etc.) will be required to predict changes in ecosystem structure and function.     

Changes in aboveground and belowground properties during secondary natural succession of a cool-temperate forest in Japan

Forest development in temperate regions is considered to be a global carbon sink. Many studies have examined forest development after harvesting or fire from aboveground (e.g., biomass) or belowground (e.g., soil nutrient) perspectives. However, few studies have explored forest development from both perspectives simultaneously in cool-temperate forests in Japan. In this study, we examined changes over 105 years in both aboveground and belowground components during secondary natural succession. The aboveground biomass increased for 50 years and reached a plateau in a 105-year-old stand. The N mineralization rate increased during succession for 50 years, but showed a decline in the 105-year-old stand due to the decrease in the nitrification rate in late succession. The percent nitrification (i.e., relative contribution of nitrification to N mineralization) decreased significantly with increasing forest stand age. The N mineralization rates had significant relationships with N concentrations of the dominant tree foliage and litter fall and with the amount of litter fall N. Meanwhile, other belowground properties (i.e., soil pH, phenol concentration, soil microbial respiration, and litter mass loss) did not show any significant relationship with forest stand age. This may be because the soil at the study sites was heterogeneous and consisted of Cambisols and Andosols, the latter of which originally has high organic matter content, and thus may have buffered the effect of the aboveground development. These results indicate that belowground N dynamics are more closely associated with aboveground development than other belowground properties in these forests.     

Influences of canopy photosynthesis and summer rain pulses on root dynamics and soil respiration in a young ponderosa pine forest

Our first objective was to link the seasonality of fine root dynamics with soil respiration in a ponderosa pine (Pinus ponderosa P. & C. Lawson) plantation located in the Sierra Nevada of California. The second objective was to examine how canopy photosynthesis influences fine root initiation, growth and mortality in this ecosystem. We compared CO2 flux measurements with aboveground and belowground root dynamics. Initiation of fine root growth coincided with tree stem thickening and shoot elongation, preceding new needle growth. In the spring, root, shoot and stem growth occurred simultaneously with the increase in canopy photosynthesis. Compared with the other tree components, initial growth rate of fine roots was the highest and their growing period was the shortest. Both above and belowground components completed 90% of their growth by the end of July and the growing season lasted approximately 80 days. The period for optimal growth is short at the study site because of low soil temperatures during winter and low soil water content during summer. High photosynthetic rates were observed following unusual late-summer rains, but tree growth did not resume. The autotrophic contribution to soil respiration was 49% over the whole season, with daily contributions ranging between 18 and 87%. Increases in soil and ecosystem respiration were observed during spring growth; however, the largest variation in soil respiration occurred during summer rain events when no growth was observed. Both the magnitude and persistence of the soil respiration pulses were positively correlated with the amount of rain. These pulses accounted for 16.5% of soil respiration between Days 130 and 329.     

Soil respiration and microbial biomass in a pecan — cotton alley cropping system in Southern USA

Little information is available on soil respiration and microbial biomass in soils under agroforestry systems. We measured soil respiration rate and microbial biomass under two age classes (young and old) of a pecan (Carya illinoinensis) — cotton (Gossypium hirsutum) alley cropping system, two age classes of pecan orchards, and a cotton monoculture on a well-drained, Redbay sandy loam (a fine-loamy, siliceous, thermic Rhodic Paleudult) in southern USA. Soil respiration was quantified monthly during the growing season from May to November 2001 using the soda-lime technique and was corrected based on infrared gas analyzer (IRGA) measurements. The overall soil respiration rates ranged from 177 to 776 mg CO₂ m^–2 h^–1. During the growing season, soil respiration was higher in the old alley cropping system than in the young alley cropping system, the old pecan orchard, the young pecan orchard, and the monoculture. Microbial biomass C was higher in the old alley cropping system (375 mg C kg^–1) and in the old pecan orchard (376 mg C kg^–1) compared to the young alley cropping system (118 mg C kg^–1), young pecan orchard (88 mg C kg^–1), and the cotton monoculture (163 mg C kg^–1). Soil respiration was correlated positively with soil temperature, microbial biomass, organic matter, and fine root biomass. The effect of alley cropping on soil properties during the brief history of alley cropping was not significant except in the old systems, where there was a trend of increasing soil respiration with short-term alley cropping. Over time, different land use and management practices influenced soil properties such as soil temperature, moisture, microbial biomass, organic matter, and fine root biomass, which in turn affected the magnitude of soil respiration. Our results suggest that trees in agroforestry systems have the potential to enhance soil fertility and sustainability of farmlands by improving soil microbial activity and accreting residual soil carbon.This revised version was published online in November 2005 with corrections to the Cover Date.     

Fine root respiration in the mangrove Rhizophora mangle over variation in forest stature and nutrient availability

Root respiration uses a significant proportion of photosynthetically fixed carbon (C) and is a globally important source of C liberated from soils. Mangroves, which are an important and productive forest resource in many tropical and subtropical countries, sustain a high ratio of root to shoot biomass which may indicate that root respiration is a particularly important component in mangrove forest carbon budgets. Mangroves are often exposed to nutrient pollution from coastal waters. Here we assessed the magnitude of fine root respiration in mangrove forests in Belize and investigated how root respiration is influenced by nutrient additions. Respiration rates of excised fine roots of the mangrove, Rhizophora mangle L., were low (4.01 +/- 0.16 nmol CO(2) g(-1) s(-1)) compared to those measured in temperate tree species at similar temperatures. In an experiment where trees where fertilized with nitrogen (N) or phosphorus (P) in low productivity dwarf forests (1-2 m height) and more productive, taller (4- 7 m height) seaward fringing forests, respiration of fine roots did not vary consistently with fertilization treatments or with forest stature. Fine roots of taller fringe trees had higher concentrations of both N and P compared to dwarf trees. Fertilization with P enhanced fine root P concentrations in both dwarf and fringe trees, but reduced root N concentrations compared to controls. Fertilization with N had no effect on root N or P concentrations. Unlike photosynthetic C gain and growth, which is strongly limited by P availability in dwarf forests at this site, fine root respiration (expressed on a mass basis) was variable, but showed no significant enhancements with nutrient additions. Variation in fine root production and standing biomass are, therefore, likely to be more important factors determining C efflux from mangrove sediments than variations in fine root respiration per unit mass.     

Autumn Fertilization in the Nursery Affects Growth of Picea abies Container Seedlings After Transplanting

Second-year Norway spruce seedlings [Picea abies (L.) Karst.] grown in containers were divided into three fertilization levels in August [5, 15 and 25 mg nitrogen (N) seedling^?1]. The resulting foliar concentrations of N were 11.0, 13.1 and 15.8 g kg^?1, respectively. Seedling height (mean 26.0 cm) did not differ among treatments. The next spring, the seedlings were tested in two experiments. (1) The seedlings were transplanted into pots containing sandy soil in the greenhouse, where they were fertilized with either pure water or nutrient solution (22 mg N l^?1). (2) The outplanting performance of the autumn-fertilized seedlings was tested on a sandy field. In the greenhouse experiment, the autumn fertilization level affected height growth and root egress of the seedlings significantly, but less than fertilization with a nutrient solution after planting. In the field experiment, during the first season after transplanting shoot growth of the seedlings increased with the level of autumn fertilization. After the second and third seasons, the seedling stem volume was highest with the highest level of autumn fertilization. These results suggest that, by improving the preplanting nutrient status of seedlings, the growth of shoot, stem diameter and root biomass can be enhanced after planting, especially on nutrient-poor soils. However, heavier autumn fertilization than that used here may yield a greater and more persistent increment in growth.     

Nutrient availability alters belowground respiration of ozone-exposed ponderosa pine

Exposure to ozone (O(3)) and changes in soil fertility influence both the metabolism of plant roots and their interaction with rhizosphere organisms. Because one indication of altered root metabolism is a change in belowground respiratory activity, we used specially designed measurement chambers to assess the effects of O(3) and nutrient availability on belowground respiratory activity of potted three-year-old ponderosa pine (Pinus ponderosa Dougl. ex Laws.). Seedlings were exposed to a factorial combination of three O(3) treatments and three fertilization treatments in open-top O(3) exposure chambers. Ozone exposure decreased and high nutrient supply increased total plant dry weight, but root/shoot ratios were not affected. In general, exposure to O(3) increased rates of belowground O(2) uptake and CO(2) release and the respiratory quotient (RQ, CO(2)/O(2)), although seasonal differences were detected. In October, following the second season of O(3) exposure, rates of belowground O(2) uptake and CO(2) release and RQ were increased in trees in the high-O(3) exposure treatment by 22, 73 and 32%, respectively, over values in control trees in charcoal-filtered air. Increasing nutrient supply resulted in decreasing rates of belowground O(2) uptake and CO(2) release but it had little effect on RQ. In the high-nutrient supply treatment, rates of belowground O(2) uptake and CO(2) release were decreased by 38 and 39%, respectively, compared with rates in the low-nutrient supply treatment. At the end of the second growing season, the high-nutrient supply treatment had decreased lateral root total nonstructural carbohydrates by 22% compared with the low-nutrient supply treatment. Nutrient availability altered the belowground respiratory response to O(3), such that the response to O(3) was greatest in the low-nutrient supply treatment. Significant O(3) effects on belowground respiratory activity were apparent before any reduction in total plant growth was found, suggesting that roots and rhizosphere organisms may be early indicators of physiological dysfunction in stressed seedlings.     

Forest thinning and soil respiration in a ponderosa pine plantation in the Sierra Nevada

Soil respiration is controlled by soil temperature, soil water, fine roots, microbial activity, and soil physical and chemical properties. Forest thinning changes soil temperature, soil water content, and root density and activity, and thus changes soil respiration. We measured soil respiration monthly and soil temperature and volumetric soil water continuously in a young ponderosa pine (Pinus ponderosa Dougl. ex P. Laws. & C. Laws.) plantation in the Sierra Nevada Mountains in California from June 1998 to May 2000 (before a thinning that removed 30% of the biomass), and from May to December 2001 (after thinning). Thinning increased the spatial homogeneity of soil temperature and respiration. We conducted a multivariate analysis with two independent variables of soil temperature and water and a categorical variable representing the thinning event to simulate soil respiration and assess the effect of thinning. Thinning did not change the sensitivity of soil respiration to temperature or to water, but decreased total soil respiration by 13% at a given temperature and water content. This decrease in soil respiration was likely associated with the decrease in root density after thinning. With a model driven by continuous soil temperature and water time series, we estimated that total soil respiration was 948, 949 and 831 g C m(-2) year(-1) in the years 1999, 2000 and 2001, respectively. Although thinning reduced soil respiration at a given temperature and water content, because of natural climate variability and the thinning effect on soil temperature and water, actual cumulative soil respiration showed no clear trend following thinning. We conclude that the effect of forest thinning on soil respiration is the combined result of a decrease in root respiration, an increase in soil organic matter, and changes in soil temperature and water due to both thinning and interannual climate variability.