Soil DIC uptake and fixation in Pinus taeda seedlings and its C contribution to plant tissues and ectomycorrhizal fungi

Plants can acquire carbon from sources other than atmospheric carbon dioxide (CO(2)), including soil-dissolved inorganic carbon (DIC). Although the net flux of CO(2) is out of the root, soil DIC can be taken up by the root, transported within the plant, and fixed either photosynthetically or anaplerotically by plant tissues. We tested the ability of Pinus taeda L. seedlings exposed to (13)C-labeled soil DIC and two NH(4)(+) availability regimes to take up and fix soil DIC. We also measured the concentration and distribution of the fixed soil DIC within the plant and mycorrhizal tissues, and quantified the contribution of soil DIC to whole-plant carbon (C) gain. Seedlings exposed to labeled DIC were significantly enriched in (13)C compared with seedlings exposed to unlabeled DIC (6.7 versus -31.7 per thousand). Fixed soil DIC was almost evenly distributed between above- and belowground biomass (55 and 45%, respectively), but was unevenly distributed among tissues. Aboveground, stem tissue contained 65% of the fixed soil DIC but represented only 27% of the aboveground biomass, suggesting either corticular photosynthesis or preferential stem allocation. Belowground, soil DIC had the greatest effect (measured as (13)C enrichment) on the C pool of rapidly growing nonmycorrhizal roots. Soil DIC contributed approximately 0.8% to whole-plant C gain, and approximately 1.6% to belowground C gain. We observed a slight but nonsignificant increase in both relative C gain and the contribution of soil DIC to C gain in NH(4)(+)-fertilized seedlings. Increased NH(4)(+) availability significantly altered the distribution of fixed soil DIC among tissue types and increased the amount of fixed soil DIC in ectomycorrhizal roots by 130% compared with unfertilized seedlings. Increased NH(4)(+) availability did not increase fixation of soil DIC in nonmycorrhizal roots, suggesting that NH(4)(+) assimilation may be concentrated in ectomycorrhizal fungal tissues, reflecting greater anaplerotic demands. Soil DIC is likely to contribute only a small amount of C to forest trees, but it may be important in C fixation processes of specific tissues, such as newly formed stems and fine roots, and ectomycorrhizal roots assimilating NH(4)(+).     

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.     

Influence of elevated CO2 and mycorrhizae on nitrogen acquisition: contrasting responses in Pinus taeda and Liquidambar styraciflua

An understanding of root system capacity to acquire nitrogen (N) is critical in assessing the long-term growth impact of rising atmospheric CO2 concentration ([CO2]) on trees and forest ecosystems. We examined the effects of mycorrhizal inoculation and elevated [CO2] on root ammonium (NH4+) and nitrate (NO3-) uptake capacity in sweetgum (Liquidambar styraciflua L.) and loblolly pine (Pinus taeda L.). Mycorrhizal treatments included inoculation of seedlings with the arbuscular mycorrhizal (AM) fungus Glomus intraradices Schenck & Smith in sweetgum and the ectomycorrhizal (EM) fungus Laccaria bicolor (Maire) Orton in loblolly pine. These plants were then equally divided between ambient and elevated [CO2] treatments. After 6 months of treatment, root systems of both species exhibited a greater uptake capacity for NH4+ than for NO3-. In both species, mycorrhizal inoculation significantly increased uptake capacity for NO3-, but not for NH4+. In sweetgum, the mycorrhizal effect on NO3- and NH4+ uptake capacity depended on growth [C02]. Similarly, in loblolly pine, the mycorrhizal effect on NO3- uptake capacity depended on growth [CO2], but the effect on NH4+ uptake capacity did not. Mycorrhizal inoculation significantly enhanced root nitrate reductase activity (NRA) in both species, but elevated [CO2] increased root NRA only in sweetgum. Leaf NRA in sweetgum did not change significantly with mycorrhizal inoculation, but increased in response to [CO2]. Leaf NRA in loblolly pine was unaffected by either treatment. The results indicate that the mycorrhizal effect on specific root N uptake in these species depends on both the form of inorganic N and the mycorrhizal type. However, our data show that in addressing N status of plants under high [CO2], reliable prediction is possible only when information about other root system adjustments (e.g., biomass allocation to fine roots) is simultaneously considered.     

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.     

Changes in above- and belowground biomass in early successional tropical secondary forests after shifting cultivation in Sarawak,Malaysia

Uncertainties in the rate of biomass variation with forest ageing in tropical secondary forests, particularly in belowground components, limit the accuracy of carbon pool estimates in tropical regions. We monitored changes in above- and belowground biomass, leaf area index (LAI), and biomass allocation to the leaf component to determine the variation in carbon accumulation rate with forest age after shifting cultivation in Sarawak, Malaysia. Nine plots in a 4-year-old forest and fourteen plots in a 10-year-old forest were monitored for 5 and 7 years, respectively. Forest and plant part biomass were calculated from an allometric equation obtained from the same forest stands. Both above- and belowground biomass increased rapidly during the initial decade after abandonment. In contrast, a much slower rate of biomass accumulation was observed after the initial decade. LAI also increased by approximately double from the 4-year-old to 10-year-old forest, and then gently increased to the 17-year-old forest. We also found that allocation variation in leaf biomass and nitrogen was closely related to the rate of biomass accumulation as a forest aged. During the first decade after abandonment, a high biomass and nitrogen allocation to the leaf component may have allowed for a high rate of biomass accumulation. However, reduction in those allocations to leaf component after the initial decade may have helped to suppress the biomass accumulation rate in older secondary forests. Roots accounted for 14.0–16.1% of total biomass in the 4–17-year-old abandoned secondary forests. We also verified the model predicted values for belowground biomass by Cairns et al. (1997) and Mokany et al. (2006), although both models overestimated the values throughout our data sets by 45–50% in the 10-year-old forest. The low root:shoot ratio in the secondary forests may have caused this overestimation. Therefore, our results suggest that we should modify the models to estimate belowground biomass considering the low root:shoot ratio in tropical secondary forests.     

Carbon budget of Pinus sylvestris saplings after four years of exposure to elevated atmospheric carbon dioxide concentration

To study the responses of Scots pine (Pinus sylvestris L.), a commercially important tree species in Europe, to future increases in atmospheric CO2 concentration ([CO2]), we grew saplings for 4 years in the ground in open-top chambers in ambient or ambient + 400 micromol mol(-1) CO2, without supplemental addition of nutrients and water. Carbon (C) budgets were developed for trees in both CO2 treatments based on productivity and biomass data obtained from destructive harvests at the end of the third and fourth years of treatment, and simulations of annual gross photosynthesis (P(tot)) and maintenance respiration by the model MAESTRA. Simulated P(tot) was enhanced by elevated [CO2], despite significant down-regulation of photosynthetic capacity. The subsequent increase in C uptake was allocated primarily to tissues with limited longevity (needles and fine roots), which explains why the measured annual increment in woody biomass did not differ between CO2 treatments. Thus, our results suggest that accelerated stem growth only occurs in the first 2 years in the presence of elevated [CO2] and that forest rotations will not be shortened significantly in response to increasing [CO2]. In elevated [CO2], a higher proportion of available C was allocated below ground, resulting in altered biomass distribution patterns. In trees of equal size, measured ratios of fine root/needle biomass and belowground/aboveground biomass were almost twice as large in the elevated [CO2] treatment. Although there are uncertainties in scaling from saplings to mature canopies, the data indicate that, in nutrient-limited Scots pine forests, elevated [CO2] is unlikely to accelerate tree growth significantly, but is likely to increase C inputs to soil.     

Timing and magnitude of C partitioning through a young loblolly pine (Pinus taeda L.) stand using 13C labeling and shade treatments

The dynamics of rapid changes in carbon (C) partitioning within forest ecosystems are not well understood, which limits improvement of mechanistic models of C cycling. Our objective was to inform model processes by describing relationships between C partitioning and accessible environmental or physiological measurements, with a special emphasis on short-term C flux through a forest ecosystem. We exposed eight 7-year-old loblolly pine (Pinus taeda L.) trees to air enriched with (13)CO(2) and then implemented adjacent light shade (LS) and heavy shade (HS) treatments in order to manipulate C uptake and flux. The impacts of shading on photosynthesis, plant water potential, sap flow, basal area growth, root growth and soil CO(2) efflux rate (CER) were assessed for each tree over a 3-week period. The progression of the (13)C label was concurrently tracked from the atmosphere through foliage, phloem, roots and surface soil CO(2) efflux. The HS treatment significantly reduced C uptake, sap flow, stem growth and fine root standing crop, and resulted in greater residual soil water content to 1 m depth. Soil CER was strongly correlated with sap flow on the previous day, but not the current day, with no apparent treatment effect on the relationship. Although there were apparent reductions in new C flux belowground, the HS treatment did not noticeably reduce the magnitude of belowground autotrophic and heterotrophic respiration based on surface soil CER, which was overwhelmingly driven by soil temperature and moisture. The (13)C label was immediately detected in foliage on label day (half-life = 0.5 day), progressed through phloem by Day 2 (half-life = 4.7 days), roots by Days 2-4, and subsequently was evident as respiratory release from soil which peaked between Days 3 and 6. The δ(13)C of soil CO(2) efflux was strongly correlated with phloem δ(13)C on the previous day, or 2 days earlier. While the (13)C label was readily tracked through the ecosystem, the fate of root C through respiratory, mycorrhizal or exudative release pathways was not assessed. These data detail the timing and relative magnitude of C flux through various components of a young pine stand in relation to environmental conditions.     

Above- and belowground biomass and primary productivity of a Larix gmelinii stand near Tura, central Siberia

We assessed above- and belowground biomass and net primary production (NPP) of a mature Larix gmelinii (Rupr.) Rupr. forest (240-280 years old) established on permafrost soils in central Siberia. Specifically, we investigated annual carbon budgets in roots in relation to root system development and availability of soil resources. Total stand biomass estimated by allometry was about 39 Mg per ha. Root biomass (17 Mg per ha) comprised about 43% of total biomass. Coarse root (>/= 5 mm in diameter) biomass was about twice that of fine roots (< 5 mm). The aboveground biomass/root biomass ratio (T/R) of the larch stand was about unity, which is much less than that of other boreal and subalpine conifer forests. The proportion of fine roots in total root biomass (35%) was relatively high compared with other cold-climate evergreen conifer forests. Total NPP, defined as the sum of annual biomass increment of woody parts and needle biomass, was estimated to be 1.8 Mg per ha per year. Allocation of total NPP to needle production was 56%. The proportion of total NPP in belowground production (27%) was less than for evergreen taiga forests. However, belowground NPP was probably under-estimated because root mortality was excluded. We conclude that L. gmelinii trees invested annual carbon gains largely into needle production or roots, or both, at the expense of growth of aboveground woody parts. This carbon allocation pattern, which resulted in the construction of exploitative root networks, appeared to be a positive growth response to the nutrient-poor permafrost soil of central Siberia.     

CDM-ARP火炬松林碳汇监测方法学研究

利用浙江省设立的火炬松样地资料,建立火炬松人工林立地指数模型、生长收获模型、生物量模型等,区分不同立地条件、采用定量方法,并考虑间伐因素,建立了浙江省CDM-ARP火炬松林碳汇监测方法学,对火炬松林分碳密度、碳储量和CO2净吸存量进行监测。监测方法分为项目规划阶段(造林前)预测和林分生长阶段(造林后)监测。选择易测定的碳库作为监测对象,并用常规测树因子来估测较难测定的各器官生物量,用生物量法估算碳汇量。     

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.     

Fine roots branch orders of <Emphasis Type="Italic">Abies faxoniana</Emphasis> respond differentially to warming in a subalpine coniferous forest ecosystem

Root is an important plant organ and has high heterogeneity. Global warming could change root and affect belowground ecological processes. There is little information on how fine roots branch orders responds to global change. This study examined the growth, morphological and physiological responses of fine roots of a subalpine coniferous species to warming. We investigated biomass, average diameter, specific root length (SRL), triphenyltetrazolium chloride (TTC) reducing capacity, carbon (C), total non-structural carbon (TNC) and fractions of the primal five branch order roots of Abies faxoniana in April, August, October and December. The decrease in total fine roots biomass after a growing season was significantly greater under warming treatment compared to control, suggesting that warming could accelerate the carbon input from root to soil, but the increment depended on tree species. Warming did not affect average diameter and SRL. Responses of biomass, TTC reducing capacity, C, TNC and fractions to warming significantly differed with root order and month. Significant warming effects were only observed in C and starch concentration of the first order and also TNC and soluble sugar concentration of the first three orders. The results indicated that the lower order roots (the first three orders) were more sensitive to warming, probably because they had more frequent, intense interactions with soil and low defense capability. Thus, global warming may dramatically alter root functions such as nutrients and water uptake as well as the cycle of C and nutrients at the whole subalpine coniferous forest ecosystem.     

Stand-level allometry in Pinus taeda as affected by irrigation and fertilization

Changing environmental conditions have the potential to alter allometric relationships between plant parts, possibly leading to ecosystem-level feedbacks. We quantified allometric shifts in field-grown loblolly pine (Pinus taeda L.) in response to altered resource availability based on data from multiple harvests to correct for size-related changes in biomass partitioning. A replicated factorial arrangement of irrigation and fertilization treatments was applied for 4 years to an 8-year-old loblolly pine plantation on a well-drained, low fertility site in North Carolina. Destructive and nondestructive growth measurements were used to develop treatment-specific regressions to estimate stand-level biomass for ephemeral and perennial plant parts, both above- and belowground. Stand-level allometric analysis indicated that irrigation increased biomass partitioning to fine roots and decreased partitioning to foliage, relative to other plant parts. Fertilization increased partitioning to perennial tissues (coarse roots, taproots, and branches) and decreased partitioning to ephemeral tissues (foliage and fine roots). Changes in allometry were small (< 6 %) but statistically significant, indicating that biomass partitioning in loblolly pine changes with altered resource availability, but is probably under strong ontogenetic control.     

Free-air CO2 enrichment (FACE) enhances biomass production in a short-rotation poplar plantation

This paper investigates the possible contribution of Short Rotation Cultures (SRC) to carbon sequestration in both current and elevated atmospheric CO2 concentrations ([CO2]). A dense poplar plantation (1 x 1 m) was exposed to a [CO2] of 550 ppm in Central Italy using the free-air CO2 enrichment (FACE) technique. Three species of Populus were examined, namely P. alba L., P. nigra L. and P. x euramericana Dode (Guinier). Aboveground woody biomass of trees exposed to elevated [CO2] for three growing seasons increased by 15 to 27%, depending on species. As a result, light-use efficiency increased. Aboveground biomass allocation was unaffected, and belowground biomass also increased under elevated [CO2] conditions, by 22 to 38%. Populus nigra, with total biomass equal to 62.02 and 72.03 Mg ha-1 in ambient and elevated [CO2], respectively, was the most productive species, although its productivity was stimulated least by atmospheric CO2 enrichment. There was greater depletion of inorganic nitrogen from the soil after three growing seasons in elevated [CO2], but no effect of [CO2] on stem wood density, which differed significantly only among species.     

Effects of elevated carbon dioxide concentration on growth and nitrogen fixation in Alnus glutinosa in a long-term field experiment

Nitrogen-fixing plant species may respond more positively to elevated atmospheric carbon dioxide concentrations ([CO2]) than other species because of their ability to maintain a high internal nutrient supply. A key factor in the growth response of trees to elevated [CO2] is the availability of nitrogen, although how elevated [CO2] influences the rate of N2-fixation of nodulated trees growing under field conditions is unclear. To elucidate this relationship, we measured total biomass, relative growth rate, net assimilation rate (NAR), leaf area and net photosynthetic rate of N2-fixing Alnus glutinosa (L.) Gaertn. (common alder) trees grown for 3 years in open-top chambers in the presence of either ambient or elevated atmospheric [CO2] and two soil N regimes: full nutrient solution or no fertilizer. Nitrogen fixation by Frankia spp. in the root nodules of unfertilized trees was assessed by the acetylene reduction method. We hypothesized that unfertilized trees would show similar positive growth and physiological responses to elevated [CO2] as the fertilized trees. Growth in elevated [CO2] stimulated (relative) net photosynthesis and (absolute) total biomass accumulation. Relative total biomass increased, and leaf nitrogen remained stable, only during the first year of the experiment. Toward the end of the experiment, signs of photosynthetic acclimation occurred, i.e., down-regulation of the photosynthetic apparatus. Relative growth rate was not significantly affected by elevated [CO2] because although NAR was increased, the effect on relative growth rate was negated by a reduction in leaf area ratio. Neither leaf area nor leaf P concentration was affected by growth in elevated [CO2]. Nodule mass increased on roots of unfertilized trees exposed to elevated [CO2] compared with fertilized trees exposed to ambient [CO2]. There was also a biologically significant, although not statistically significant, stimulation of nitrogenase activity in nodules exposed to elevated [CO2]. Root nodules of trees exposed to elevated [CO2] were smaller and more evenly spaced than root nodules of trees exposed to ambient [CO2]. The lack of an interaction between nutrient and [CO2] effects on growth, biomass and photosynthesis indicates that the unfertilized trees maintained similar CO2-induced growth and photosynthetic enhancements as the fertilized trees. This implies that alder trees growing in natural conditions, which are often limited by soil N availability, should nevertheless benefit from increasing atmospheric [CO2].     

Belowground carbon dynamics in loblolly pine (Pinus taeda) immediately following diammonium phosphate fertilization

Forest soils store an immense quantity of labile carbon (C) and a may be a large potential sink for atmospheric C. Forest management practices such as fertilization may enhance overall C storage in soils, yet changes in physiological processes following nutrient amendments have not been widely investigated. We intensively monitored belowground C dynamics for nearly 200 days following diammonium phosphate fertilization of pot-grown loblolly pine (Pinus taeda L.) seedlings in an effort to examine the short-term effects of fertilization on processes involved in soil C sequestration. Soil respiration rates initially increased in fertilized pots relative to controls, followed by a brief reversal in this trend and then a final sustained pattern of elevated rates of soil respiration in the fertilized treatment. Patterns in soil respiration rates over time reflected changes in autotrophic (root) and heterotrophic (microbial) components of soil respiration. Root respiration rates were greater in the fertilized treatment 49 days following fertilization and returned to control rates by the end of the study. In contrast, microbial respiration rates and microbial activity per soil C concentration remained depressed over the same time period. Compared with control seedlings, total root biomass was 27% greater in fertilized seedlings harvested at the end of the study, indicating that the elevated soil respiration rates observed toward the end of the study were a result of increased respiring root biomass. We conclude that fertilization, at least over the short-term, may increase soil C sequestration by increasing belowground biomass production and reducing microbial driven C turnover.     

Carbon and nitrogen allocation in ectomycorrhizal and non-mycorrhizal Pinus sylvestris L. seedlings

We studied carbon and nitrogen allocation in mycorrhizal and non-mycorrhizal Scots pine (Pinus sylvestris L.) seedlings grown in a semi-hydroponic system with nitrogen as the growth limiting factor. Three ectomycorrhizal fungi were compared: one pioneer species (Thelephora terrestris Ehrh.: Fr.) and two late-stage fungi (Suillus bovinus (L.: Fr.) O. Kuntze, and Scleroderma citrinum Pers.). By giving all plants in each treatment the same amount of readily available nitrogen, we ensured that the external mycelium could not increase the total nitrogen content of the plants, thereby guaranteeing that any change in carbon or nitrogen partitioning was a direct effect of the mycorrhizal infection itself. Carbon and nitrogen partitioning were measured at an early and a late stage of mycorrhizal development, and at a low and a high N addition rate. Although mycorrhizal seedlings had a higher net assimilation rate and a higher shoot/root ratio than non-mycorrhizal seedlings, they had a lower rate of shoot growth. The high carbon demand of the mycobionts was consistent with the large biomass of external mycelia and the increased belowground respiration of the mycorrhizal plants. The carbon cost to the host was similar for pioneer and late-stage fungi. Above- and belowground partitioning of nitrogen was also affected by mycorrhizal infection. The external mycelia of Scleroderma citrinum retained 32% of the nitrogen supplied to the plants, thus significantly reducing nitrogen assimilation by the host plants and consequently reducing their growth rate. By contrast, the external mycelia of T. terrestris and Suillus bovinus retained less nitrogen than the mycelia of Scleroderma citrinum, hence we attributed the decreased growth rates of their host plants to a carbon drain rather than a nitrogen deficiency.     

三江平原典型湿地毛果苔草养分含量及累积特征

以三江平原典型湿地毛果苔草群落为研究对象,在2007年5月至2008年9月研究了毛果苔草植物地上和地下部生物量季节动态以及养分（氮、碳、磷）特征。结果表明：毛果苔草地上生物量随着植物的生长符合二次函数模型,地下生物量随着植物的生长符合指数增长模型。毛果苔草地上、地下部分有机碳含量随植物生长均呈显著正的线性相关,相关系数分别是0.983和0.746。地上、地下有机碳累积量随着植物生长的变化趋势与其生物量的变化是一致的,随着植物的生长符合二次函数模型。毛果苔草地上、地下部分总氮含量和积累量随着植物的生长分别符合指数递增模型。毛果苔草磷含量随着植物的生长分别符合二次函数模型,但是磷累积量随着植物的生长分别符合线性增长模型。植物生长季毛果苔草植物体碳/氮的大小与其总氮含量呈反比关系,碳/磷的大小与其总磷含量呈反比关系,研究表明氮含量是毛果苔草生长过程中的限制性养分。     

Discrepancy in fine root turnover estimates between diameter-based and branch-order-based approaches: a case study in two temperate tree species

Fine root turnover plays a key role in carbon(C) budgets and nutrients cycles in forest ecosystems.However,the difference between branch-order-based and diameter-based approaches in estimating fine root turnover is still unclear.We studied root biomass turnover based on multiplying root standing biomass by turnover rate(inverse of median root longevity) in two Chinese temperate tree species,Fraxinus mandshurica Rupr.and Larix gmelinii Rupr.The minirhizotron(MR) technique was used to estimate longevities for first and second order roots,and total roots(R total) apparent on the MR tube surface.The corresponding biomass for each root group was estimated by soil monolith.The difference in biomass turnover between R total and the sum of the first and second order roots was used to represent the discrepancy between diameter-and order-based approaches.First order roots had shorter life spans and higher biomass turnover rates than the second order roots in both species.Biomass turnover estimated by the order-based method for F.mandshurica and L.gmelinii were 155.4 g m-2 a-1 and 158.9 g m-2 a-1,respectively,in comparison with 99.5 g m-2 a-1 and 117.7 g m-2 a-1 estimated by the diameter-based method,indicating that the diameter-based approach underestimated biomass turnover.The most probable reason was that the order-based method enhanced separation of the heterogeneous root population into relatively homogenous root groups with varying turnover rates.We conclude that separating fine root pool into different branch orders can improve the accuracy of estimates for fine root turnover,as well as the understanding of the belowground C allocation and nutrient cycling at ecosystem level.     

Xylem vulnerability to cavitation in Pseudotsuga menziesii and Pinus ponderosa from contrasting habitats

In the Rocky Mountains, ponderosa pine (Pinus ponderosa (ssp.) ponderosa Dougl. ex P. Laws. & C. Laws) often co-occurs with Douglas-fir (Pseudotsuga menziesii var. glauca (Mayr) Franco). Despite previous reports showing higher shoot vulnerability to water-stress-induced cavitation in ponderosa pine, this species extends into drier habitats than Douglas-fir. We examined: (1) whether roots and shoots of ponderosa pine in riparian and slope habitats are more vulnerable to water-stress-induced cavitation than those of Douglas-fir; (2) whether species-specific differences in vulnerability translate into differences in specific conductivity in the field; and (3) whether the ability of ponderosa pine to extend into drier sites is a result of (a) greater plasticity in hydraulic properties or (b) functional or structural adjustments. Roots and shoots of ponderosa pine were significantly more vulnerable to water-stress-induced cavitation (overall mean cavitation pressure, Psi(50%) +/- SE = -3.11 +/- 0.32 MPa for shoots and -0.99 +/- 0.16 MPa for roots) than those of Douglas-fir (Psi(50%) +/- SE = -4.83 +/- 0.40 MPa for shoots and -2.12 +/- 0.35 MPa for roots). However, shoot specific conductivity did not differ between species in the field. For both species, roots were more vulnerable to cavitation than shoots. Overall, changes in vulnerability from riparian to slope habitats were small for both species. Greater declines in stomatal conductance as the summer proceeded, combined with higher allocation to sapwood and greater sapwood water storage, appeared to contribute to the ability of ponderosa pine to thrive in dry habitats despite relatively high vulnerability to water-stress-induced cavitation.     

Coarse and fine root respiration in aspen (Populus tremuloides)

Coarse and fine root respiration rates of aspen (Populus tremuloides Michx.) were measured at 5, 15 and 25 degrees C. Coarse roots ranged from 0.65 to 4.45 cm in diameter, whereas fine roots were less than 5 mm in diameter. To discriminate between maintenance and growth respiration, root respiration rates were measured during aboveground growing periods and dormant periods. An additional measurement of coarse root respiration was made during spring leaf flush, to evaluate the effect of mobilization of resources for leaf expansion on root respiration. Fine roots respired at much higher rates than coarse roots, with a mean rate at 15 degrees C of 1290 micromol CO2 m-3 s-1 during the growing period, and 660 micromol CO2 m-3 s-1 during the dormant period. The temperature response of fine root respiration rate was nonlinear: mean Q10 was 3.90 for measurements made at 5-15 degrees C and 2.19 for measurements made at 15-25 degrees C. Coarse root respiration rates measured at 15 degrees C in late fall (dormant season) were higher (370 micromol CO2 m-3 s-1) than rates from roots collected at leaf flush and early summer (200 micromol CO2 m-3 s-1). The higher respiration rates in late fall, which were accompanied by decreased total nonstructural carbohydrate (TNC) concentrations, suggest that respiration rates in late fall included growth expenditures, reflecting recent radial growth. Neither bud flush nor shoot growth of the trees caused an increase in coarse root respiration or a decrease in TNC concentrations, suggesting a limited role of coarse roots as reserve storage organs for spring shoot growth, and a lack of synchronization between above- and belowground growth. Pooling the data from the coarse and fine roots showed a positive correlation between nitrogen concentration and respiration rate.