Influence of nutrients,disturbances and site conditions on carbon stocks along a boreal forest transect in central Canada

The interacting influence of disturbances and nutrient dynamics on aboveground biomass, forest floor, and mineral soil C stocks was assessed as part of the Boreal Forest Transect Case Study in central Canada. This transect covers a range of forested biomes–-from transitional grasslands (aspen parkland) in the south, through boreal forests, and into the forested subarctic woodland in the north. The dominant forest vegetation species are aspen, jack pine and spruce. Disturbances influence biomass C stocks in boreal forests by determining its age-class structure, altering nutrient dynamics, and changing the total nutrient reserves of the stand. Nitrogen is generally the limiting nutrient in these systems, and N availability determines biomass C stocks by affecting the forest dynamics (growth rates and site carrying capacity) throughout the life cycle of a forest stand. At a given site, total and available soil N are determined both by biotic factors (such as vegetation type and associated detritus pools) and abiotic factors (such as N deposition, soil texture, and drainage). Increasing clay content, lower temperatures and reduced aeration are expected to lead to reduced N mineralization and, ultimately, lower N availability and reduced forest productivity. Forest floor and mineral soil C stocks vary with changing balances between complex sets of organic carbon inputs and outputs. The changes in forest floor and mineral soil C pools at a given site, however, are strongly related to the historical changes in biomass at that site. Changes in N availability alter the processes regulating both inputs and outputs of carbon to soil stocks. N availability in turn is shaped by past disturbance history, litter fall rate, site characteristics and climatic factors. Thus, understanding the life-cycle dynamics of C and N as determined by age-class structure (disturbances) is essential for quantifying past changes in forest level C stocks and for projecting their future change.     

蚂蚁筑巢对高檐蒲桃热带森林群落土壤呼吸的影响

蚂蚁筑巢能够改变热带森林土壤微生物与土壤理化性质的状况,从而对土壤呼吸时间动态产生重要影响。本研究以西双版纳高檐蒲桃热带森林群落为研究对象,采用Li-6400-09便携式土壤呼吸测定仪对蚂蚁筑巢地与非筑巢地土壤呼吸进行测定。研究结果表明:(1)高檐蒲桃群落土壤呼吸呈明显的单峰型季节变化趋势,且土壤呼吸速率蚂蚁筑巢地(4.96μmol CO_2m~(-2)s~(-1))高于非筑巢地(4.42μmol CO_2m~(-2)s~(-1))。(2)土壤温度和土壤水分显著影响土壤呼吸的时间动态(P0.01);蚂蚁筑巢显著改变巢内温度与水分(P0.05),进而影响土壤呼吸动态。土壤温度对土壤呼吸动态的贡献:蚁巢(83.8%—91.8%)大于非巢地(81.2%—83.1%),但由于筑巢地土壤湿度低于非巢地,土壤水分对土壤呼吸动态的贡献率表现为蚁巢低于非筑巢地。(3)蚂蚁筑巢显著增加土壤微生物生物量(P0.05),从而对土壤呼吸速率产生极显著的影响(P0.01)。蚂蚁筑巢引起微生物生物量碳的增加能够解释76.9%—71.1%的土壤呼吸变化。(4)蚂蚁筑巢引起土壤理化性质变化对土壤呼吸产生一定的影响。土壤容重与土壤呼吸速率呈显著负相关;土壤呼吸速率与土壤微生物量碳、有机质、易氧化有机碳、全氮、硝氮和铵氮显著正相关(P0.05或P0.01)。因此,蚂蚁筑巢显著改变土壤微生物(如微生物生物量碳)、土壤物理性质(如土壤温度与水分)、土壤化学性质(如碳和氮养分),进而对热带森林土壤呼吸产生重要影响。     

Interactions among nitrogen-transforming bacteria and nitrogen-fixingPisum sativum L. in laboratory sand columns

Plants of barley (Hordeum vulgare), ryegrass (Lolium perenne), pea (Pisum sativum) or turnip (Brassica campestris rapifera) were grown in pots of unfertilised soil for 10 weeks together with unplanted control pots. A wide range of soil microbiological parameters was measured on bulk soil samples 2, 4, 7 and 10 weeks after seedlings were transplanted. There was no effect of planting or differential effect of plant species upon respiration rate, microbial biomass N, or biomass of microbial predators, but these parameters all varied significantly over time. Respiration, biomass N and nematode biomass all increased, whilst protozoan biomass decreased. Microbial biomass C showed no significant temporal changes or effect of planting. There was evidence for differential plant effects on potential nitrification and denitrification. Nitrification rates were depressed, compared with the fallow, in all treatments except the pea, at some time in the experiment. Conversely denitrification rates were enhanced in all treatments, except the grass, at specific times. Denitrification rates were greater in the pea treatment than the fallow on all occasions. These results demonstrate that plants do not necessarily influence the gross microbiology of the soil, but may affect physiologically distinct sub-components of the microbial biomass.     

Nitrogen addition reduces soil respiration in a mature tropical forest in southern China

Response of soil respiration (CO₂ emission) to simulated nitrogen (N) deposition in a mature tropical forest in southern China was studied from October 2005 to September 2006. The objective was to test the hypothesis that N addition would reduce soil respiration in N saturated tropical forests. Static chamber and gas chromatography techniques were used to quantify the soil respiration, following four‐levels of N treatments (Control, no N addition; Low‐N, 5 g N m^?2 yr^?1; Medium‐N, 10 g N m^?2 yr^?1; and High‐N, 15 g N m^?2 yr^?1 experimental inputs), which had been applied for 26 months before and continued throughout the respiration measurement period. Results showed that soil respiration exhibited a strong seasonal pattern, with the highest rates found in the warm and wet growing season (April–September) and the lowest rates in the dry dormant season (December–February). Soil respiration rates showed a significant positive exponential relationship with soil temperature, whereas soil moisture only affect soil respiration at dry conditions in the dormant season. Annual accumulative soil respiration was 601±30 g CO₂‐C m^?2 yr^?1 in the Controls. Annual mean soil respiration rate in the Control, Low‐N and Medium‐N treatments (69±3, 72±3 and 63±1 mg CO₂‐C m^?2 h^?1, respectively) did not differ significantly, whereas it was 14% lower in the High‐N treatment (58±3 mg CO₂‐C m^?2 h^?1) compared with the Control treatment, also the temperature sensitivity of respiration, Q₁₀ was reduced from 2.6 in the Control with 2.2 in the High‐N treatment. The decrease in soil respiration occurred in the warm and wet growing season and were correlated with a decrease in soil microbial activities and in fine root biomass in the N‐treated plots. Our results suggest that response of soil respiration to atmospheric N deposition in tropical forests is a decline, but it may vary depending on the rate of N deposition.     

Earthworm Invasion,Fine-root Distributions,and Soil Respiration in North Temperate Forests

The efflux of carbon from soils is a critical link between terrestrial ecosystems and the atmosphere. Current concerns about rising atmospheric carbon dioxide (CO₂) concentrations highlight the need to better understand the dynamics of total soil respiration (TSR, sum of root and heterotroph respiration) in changing environments. We investigated the effects of exotic earthworm invasion on TSR, fine-root distributions, and aboveground litterfall flux in two sugar maple-dominated forests in two locations in New York State, USA. The Arnot Forest in central New York was harvested in the late 19th century and has no history of cultivation. Tompkins Farm in eastern New York regenerated following abandonment from cultivation approximately 75 years ago. Arnot had 20% higher total soil CO₂ efflux (880 g C m^–2year^–1) than Tompkins (715 g C m^–2year^–1). The presence of earthworms had no influence on TSR at either location. However, fine-root (< 1 mm diameter) biomass in earthworm plots (350 g/m²) was significantly lower than in worm-free reference plots (440 g/m²) at Arnot. Fine-root nitrogen (N) concentrations were not influenced by earthworms, and total fine-root N content was significantly reduced in the presence of earthworms at Arnot. Our results indicate that the presence of exotic earthworms is not presently affecting net C emission from soil in these forests. They also suggest a change in root function in earthworm plots that is not associated with higher fine-root N concentration, but that increases efficiency of nutrient uptake and also may enhance the belowground supply of C for heterotroph metabolism.     

Linking aphid honeydew, throughfall, and forest floor solution chemistry of Norway spruce

In a seminatural manipulation experiment with artificial irrigation we followed throughfall and forest floor solution chemistry collected underneath aphid infested and uninfested Norway spruce. Solutions underneath infested trees showed significantly higher concentrations of dissolved organic carbon (DOC) but lower concentrations of dissolved organic nitrogen (DON), NO₃-N, and NH₄-N in throughfall solutions and of NH₄-N in forest floor solutions. Average concentrations were 40.5% (DON), 27.5% (NO₃-N), and 46.2% lower (NH₄-N) underneath infested trees in throughfall solutions, and 19.5% (DON), 9.4% (NO₃-N), and 42.0% (NH₄-N) lower in forest floor solutions. Differences in throughfall were more pronounced than in forest floor leachates. It is likely that honeydew is fuelling the metabolism of micro-organisms and thus critically affects above and below ground nutrient cycles. We emphasize the importance of linking the biology of herbivores and micro-organisms with geochemical processes.     

川西亚高山不同森林生态系统碳氮储量及其分配格局

森林采伐和恢复是影响森林碳氮储量的重要因素。以川西亚高山岷江冷杉原始林、粗枝云杉阔叶林、天然次生林和粗枝云杉人工林为研究对象,采用样地调查和生物量实测的方法,研究了不同森林生态系统各组分碳、氮储量及其分配特征。结果表明岷江冷杉原始林、粗枝云杉阔叶林、天然次生林和粗枝云杉人工林生态系统碳储量分别为611.18、252.31、363.07 tC/hm~2和239.06 tC/hm~2;氮储量分别为16.44、12.11、15.48 tN/hm~2和8.92 tN/hm~2。恢复林分与原始林碳储量在土壤—植被的分配格局发生了变化,而氮储量未发生变化。岷江冷杉原始林以植被碳储量为主,恢复林分以土壤为主,氮储量均以土壤为主。乔木层碳储量分别占生态系统总储量的56.65%、17.63%、13.57%和22.05%,土壤层(0—80 cm)分别占32.03%、69.87%、76.20%和72.12%;土壤层氮储量占生态系统总储量的76.80%—92.58%。植物残体碳氮储量分别占生态系统总储量的4.40%—9.83%和2.94%—7.08%,林下植被所占比例最小。空间格局上,岷江冷杉原始林植被部分具有较高的碳储量,应进行保护。3种恢复林分具有较高的碳汇潜力,且地上/地下碳储量较低,表明其碳汇潜力尤其表现在地上部分。天然次生林利于土壤有机碳的积累,而人工林乔木层碳储量较高。     

Mineral nitrogen and microbial dynamics in the forest floor of clearcut or partially harvested successional boreal forest stands

The effects of clearcut and partial harvesting of early-seral trembling aspen plots were compared to conventional clearcut harvesting in mid-seral mixedwood and late-seral conifer plots. Twice a year, for three consecutive years, we assessed mineral N and microbial dynamics in the forest floor of these plots to test three hypotheses related to the higher litter quality of aspen leaves and to the sustained inputs of available C on partially harvested plots: (1) the post-clearcutting mineral N flush and the net [(NO₃^–): (NO₃^– + NH₄⁺)] production ratio (RNI) are higher in aspen plots than in black spruce plots, with intermediate values occurring in mixedwood plots; (2) net N mineralization rates in aspen plots are higher in spring than in autumn; and (3) compared to clearcutting, partial harvesting reduces potential ammonification and nitrification rates. Initial NH₄⁺ and NO₃^– concentrations respectively ranged between 1.7–4.4 and 0.2–1.5 g N kg^–1 N_total, net ammonification and nitrification rates (30 d incubations) respectively ranged between 5.3–17.8 and 0.1–27.6 g N kg^–1 N_total, basal respiration ranged between 20.9–38.9 mg CO₂-C kg^–1 h^–1, and microbial biomass ranged between 6.1–8.7 g C_mic kg^–1. Although clearcutting increased NO₃^– concentrations in aspen plots, the balance of our results did not support our first hypothesis, because NH₄⁺ concentrations increased in conifer plots only, potential ammonification was unaffected by clearcutting, potential nitrification increased in mixedwood plots only, and RNI increased in all plots. In each seral stage, basal respiration, microbial biomass, and metabolic quotient either increased or were unaffected by clearcutting, suggesting that increases in RNI after disturbance were not related to lower microbial immobilisation of NO₃^– due to lower available C. Forest floors in mid-seral mixedwood plots exhibited a distinct combination of mineral N and microbial properties, suggesting that the functional richness of the forest is enhanced not only by the number of species, but also by the diversity of assemblages that are present. Results supported our second hypothesis and showed, furthermore, that net N mineralization in conifer stands is greater in autumn than in spring. Partial harvesting in aspen stands resulted in lower potential mineralization of N and lower RNI, compared to clearcutting. Further lysimetry studies are needed to confirm whether partial harvesting mitigates NO₃^– leaching following disturbance.     

林床清理对落叶松(Larix gmelinii)人工林土壤呼吸和物理性质的影响

林下可再生生物质资源的利用是当今森林资源利用的热点,通过林床清理可以获得廉价可再生生物质资源,但其对林分土壤碳收支的影响尚不清楚。运用红外气体分析法（IRGA法）连续两年观测了林床清理对落叶松（Larix gmelinii）人工林土壤呼吸及物理性质的影响,并估算了林床清理生物质资源利用对落叶松人工林碳收支的影响。结果表明：林床清理能够降低落叶松人工林的土壤呼吸,2a的平均值由2．20μmol&#183;m^-2s^-1降低到1．18μmol&#183;m^-2s^-1,平均降低幅度1．02μmol&#183;m^-2s^-1,年呼吸总量由41．2μmol&#183;m^-2a^-1降至22．4μmol&#183;m^-2a^-1,而且,使土壤呼吸Q10值从2．33降低到2．22,R0值从0．61μmol&#183;m^-2s^-1降至0．36μmol&#183;m^-2s^-1;林床清理能够使林床土壤温度冬季低于对照,而夏季则有相反趋势,清理使得林床土壤湿度变化幅度加大,而且秋季和春季较对照低,而夏季偏高;林床清理使得表层土壤容重要比对照未清理样地高53％（P〈0．05）,土壤非毛管孔隙度比未处理样地低49．5％（P〈0．001）,毛管孔隙度较对照未清理降低约15％（P〈0．001）。林床清理导致林下生物质资源所储藏的碳非呼吸性释放约175．0mol&#183;m^-2,当考虑到林床清理导致的土壤呼吸的降低作用时,所测定的2a内土壤净碳支出由175．0mol&#183;m^-2降低至137．4mol&#183;m^-2。林床清理措施增加生物质资源利用和其所导致的土壤呼吸释放减少,能够减少非冉生资源利用导致的碳释放压力。但仍然需要注意到林床清理使得土壤物理结构发生改变,可能不利于落叶松的生长和落叶松林生态系统的稳定。     

秦岭火地塘林区油松林土壤呼吸时空变异

土壤呼吸是陆地生态系统碳循环的关键生态过程,土壤呼吸的时空变异及其影响因子已成为生态学研究的主要内容之一。采用红外线开路气室法和便携式微气象站,连续测定了秦岭火地塘林区天然次生油松林地不同部位土壤呼吸速率和不同土层深度土壤温度和土壤体积含水率,结果表明:(1)植物生长季,试验地上部与中部、中部与下部,土壤呼吸日均值间存在显著差异。植物休眠季,全坡面土壤呼吸日均值差异不显著。同一观测部位植物生长季与休眠季,土壤呼吸日均值差异显著。观测期内全样地土壤呼吸日均值为(38.64±6.43)gm-2d-1;(2)同一地形部位不同观测月中和不同地形部位同一观测时间,土壤呼吸月均值大多存在显著差异,植物生长季和休眠季,全样地土壤呼吸均值分别为(46.98±2.21)gm-2d-1和(35.94±1.01)gm-2d-1,全样地土壤呼吸月均值为(1.18±0.20)kgm-2月-1,休眠季土壤日均呼吸约为整个观测季的43.34%;(3)当土壤温度9.0℃时,土壤温度与土壤呼吸速率间均存在显著的指数关系。回归模型的决定系数均大于0.87,均方差根不超过0.21,模型有效性系数不小于0.85,残差系数的绝对值不超过0.007。(4)植物生长季0-5cm和5-10cm土层及植物休眠季0-5cm土层,土壤呼吸日累积值均值与相应土层深度土壤体积含水率均值间存在三次函数关系,回归模型的决定系数分别为0.456,0.513和0.143;植物休眠季5-10cm土层,土壤呼吸日累积值均值与土壤体积含水率均值间存在幂函数关系,回归模型的决定系数为0.650。     

Reduction of forest floor respiration by fertilization on both carbon dioxide-enriched and reference 17-year-old loblolly pine stands

Elevated atmospheric carbon dioxide (CO₂^e) increases soil respiration rates in forest, grassland, agricultural and wetland systems as a result of increased growth, root biomass and enhanced biological activity of soil microorganisms. Less is known about how forest floor fluxes respond to the combined effects of elevated CO₂ and nutrient amendments; until now no experiments have been in place with large forest trees to allow even preliminary investigations. We investigated changes in forest floor respiration (S_ff) in a Pinus taeda L. plantation fumigated with CO₂ by using free‐air CO₂ enrichment (FACE) technology and given nutrient amendments. The prototype FACE apparatus (FACEp; 707 m²) was constructed in 1993, 10 years after planting, on a moderate fertility site in Duke Forest, North Carolina, USA, enriching the stand to 55 Pa (CO₂^e). A nearby ambient CO₂ (CO₂^a) plot (117 m²) was designated at the inception of the study as a reference (Ref). Both FACEp and Ref plot were divided in half and urea fertilizer was applied to one half at an annual rate of 11.2 g N m^?2 in the spring of 1998, 1999 and 2000. Forest floor respiration was monitored continuously for 220 days – March through November 2000 – by using two Automated Carbon Efflux Systems. Thirty locations (491 cm² each) were sampled in both FACEp and Ref, about half in each fertility treatment. Forest floor respiration was strongly correlated with soil temperature at 5 cm. Rates of S_ff were greater in CO₂^e relative to CO₂^a (an enhancement of ～178 g C m^?2) during the measurement period. Application of fertilizer resulted in a statistically significant depression of respiration rates in both the CO₂^a and CO₂^e plots (a reduction of ～186 g C m^?2). The results suggest that closed canopy forests on moderate fertility sites cycle back to the atmosphere more assimilated carbon (C) than similar forests on sites of high fertility. We recognize the limitations of this non‐replicated study, but its clear results offer strong testable hypotheses for future research in this important area.     

不同林龄落叶松人工林土壤微生物生物量碳氮的季节变化

为从土壤微生物生物量角度分析不同林龄落叶松人工林的土壤肥力状况,对辽宁东部山区两种林龄(9年生,幼龄林;43年生,成熟林)落叶松人工林不同土层(腐殖质层和矿化层)微生物生物量碳、氮季节变化进行了监测,并分析了微生物生物量碳氮的季节变化与土壤养分及水分的关系.结果表明:两种林龄落叶松腐殖质层微生物生物量碳、氮含量均高于矿化层;在腐殖质层,幼龄林微生物生物量碳、氮含量高于成熟林.方差分析表明,在春、秋季节,同一土层两林龄土壤微生物生物量碳、氮含量之间差异达到显著水平(P<0.01).在观测的3个季节内,幼龄林腐殖质层的微生物生物量碳基本无变化,而成熟林的微生物生物量碳在秋季达到最高;两种林龄落叶松微生物生物量氮均在夏季达到最高.在矿化层,两种林龄落叶松微生物生物量碳、氮均在秋季达到最大.相关分析发现,微生物生物量碳、氮之间以及土壤微生物生物量碳、氮与土壤有机碳、全氮呈显著正相关,而与土壤水分无相关性;另外,落叶松人工林内的灌木种类和数量以及季节性温度变化对土壤微生物生物量碳氮也有影响.上述结果表明,研究区域土壤微生物生物量碳、氮的季节波动与土壤养分状况密切相关,幼龄林土壤养分状况优于成熟林.     

固氮树种在混交林中的作用研究Ⅱ.固氮树木叶部N、P养分动态特征

研究了23种固氮与非固氮树种叶部N、P含量的季节变化、输出率以及叶片衰老过程中N、P的输出过程.结果表明,供试树种叶部N、P含量均存在着明显的季节变化,平均变化率为21～23%,最大变化率出现在叶片衰老期.在衰老期内不同树种表现出不同的N、P动态特征.固氮树种表现出低输出率,落叶中N含量是非固氮树种的1.6～3.7倍.固氮树种生长季叶部N含量在2.5%左右.     

Experimental assessment of salamanders as predators of detrital food webs: effects on invertebrates, decomposition and the carbon cycle

Declines in amphibian populations and the loss of some species has spurred research into the causes of declines. Little research has been done on what the loss of amphibian species may mean to ecosystem function. I describe a field experiment where the number of Plethodon cinereus (the red-backed salamander) was manipulated in enclosures buried in the forest floor. I determined the composition of the forest floor invertebrate community and the rate of decomposition of leaf litter in enclosures with and without salamanders over 4.5 months. Salamander predation in enclosures significantly reduced invertebrate numbers compared to control enclosures. Salamander predation also indirectly reduced rate of decomposition by between 11 and 17%. This was probably due to salamanders consuming a significant number of leaf fragmenters (e.g. Coleoptera and Diptera larvae). Predation on leaf litter fragmenters may alter carbon dynamics in forested ecosystems.     

采伐和火烧对森林氮动态的影响

森林是重要的陆地生态系统类型,它通过特有的养分循环机制维持其结构和功能.其中氮素对林木生长和发育十分重要,而且常是森林生产力的限制因素.另一方面,森林氮动态又常受到人类活动干扰的影响.根据国内外研究结果综述了采伐和火烧对森林氮动态的影响.结果表明采伐后环境因素的变化将影响森林N动态,其中最为关注的是采伐后一系列因素引起的N损失,如:N淋溶增加、伴随生物量的N迁移以及因径流或侵蚀增加造成的枯枝落叶层和土壤层N流失.这些N损失又将影响更新林分的生长和生产力.此外,采伐后N吸收速率一般下降,但随着植被快速生长N吸收速率将不断增加.采伐后氨化和硝化过程增强,但因短期内同化作用较弱,生态系统中大部分N将发生损失.火烧对森林N动态的短期影响主要包括:第一,火烧时N直接挥发损失;第二,火烧后N有效性增加,这主要由灰分沉积、根和微生物死亡及有机质N矿化增强等综合造成.随着时间延长,N有效性逐渐降低,这可能与火烧引起的有机质损失、植物N吸收增加、淋溶或侵蚀损失有关.然而,目前关于火烧造成的长期生态影响,如火烧后地上植被恢复与地下生物地球化学过程变化有何联系仍不太清楚.未来研究应着重于探讨氮素对森林采伐和火烧作出的短期响应将如何长期影响森林的结构和功能.此外,建议在实施营林方案时需考虑采伐和火烧对生态系统氮的影响.