亚热带次级森林演替过程中模拟氮磷沉降对土壤微生物生物量及土壤养分的影响

氮（N）沉降深刻影响着森林生态系统的生物多样性、生产力和稳定性。亚热带地区森林土壤磷（P）的有效性较低,N沉降将更突显P的限制作用。N、P输入对亚热带次级森林土壤的影响是否依赖于森林演替阶段知之甚少。选取两种不同演替年龄阶段（年轻林：<40 a;老年林：>85 a）的亚热带常绿阔叶林,设置模拟N和/或P沉降（10 g m^-2 a^-1）4个处理（Ctrl、N、P、NP）,连续处理4.5年后采集表层、次表层和下底层（0-15、15-30、30-60 cm）土壤样品,综合分析了土壤微生物生物量碳（MBC）氮（MBN）和多种土壤养分含量。结果表明,MBC、MBN及土壤养分含量均随土壤深度增加而降低。N添加对两种演替阶段森林土壤中MBC和MBN均无显著影响。施P相关处理（P和NP）对年轻林表层土壤MBC和MBN无显著影响,但显著增加了老年林表层土壤MBC和MBN（P<0.05）,表明老年林可能比年轻林更易受P限制。N添加显著增加了两种演替森林表层土壤可溶性有机氮（DON）、氨态氮（NH₄⁺-N）和硝态氮（NO₃^--N）的含量（P<0.05）;P相关处理（P和NP）显著增加两种演替阶段表层和次表层土壤速效磷（AP）以及表层土壤全磷（TP）的含量（P<0.05）。土壤MBC和MBN与土壤中各养分指标（可溶性有机碳DOC、DON、NH₄⁺-N、NO₃^--N、AP、全碳TC、全氮TN和TP）呈显著正相关关系,土壤TC、TN和DOC是影响土壤微生物生物量的主要因子。研究可为评估和揭示未来全球环境变化背景下不同演替林龄亚热带森林的土肥潜力及土壤质量的演变提供一定的科学理论依据。     

荒漠草原区弃耕地植被演替过程中植物群落生物量及土壤养分变化

采用空间变化代替时间变化的方法,以荒漠草原区不同年限(1、4、9、12和20年)弃耕地为对象,研究弃耕演替过程中植物群落生物量与土壤养分的变化特征.结果表明： 随弃耕年限的增加,弃耕地植物群落地上生物量呈先减少后增加的趋势,0～60 cm土层的土壤全氮、全磷和有机碳含量及碳密度均呈先增大后减小再增大的趋势,4年和20年弃耕地的土壤全氮、全磷含量达到峰值.弃耕演替过程中土壤全氮和有机碳含量对植物群落生物量的影响大于土壤容重和土壤全磷.     

Carbon and nitrogen cycling during old-field succession: Constraints on plant and microbial biomass

Soil C and N dynamics were studied in a sequence of old fields of increasing age to determine how these biogeochemical cycles change during secondary succession. In addition, three different late-successional forests were studied to represent possible "steady state" conditions. Surface soil samples collected from the fields and forests were analyzed for total C, H₂O-soluble C, total N, potential net N mineralization, potential net nitrification, and microbial biomass. Above-and belowground plant biomass was estimated within each of the old field sites.Temporal changes in soil organic C, total N and total plant biomass were best described by a gamma function [y =at ^b e ^ctd +f] whereas a simple exponential model [y =a(l – e^–bt ) + c] provided the best fit to changes in H₂O-soluble C, C:N ratio, microbial C, and microbial N. Potential N mineralization and nitrification linearly increased with field age; however, rates were variable among the fields. Microbial biomass was highly correlated to soil C and N pools and well correlated to the standing crop of plant biomass. In turn, plant biomass was highly correlated to pools and rates of N cycling.Patterns of C and N cycling within the old field sites were different from those in a northern hardwood forest and a xeric oak forest; however, nutrient dynamics within an oak savanna were similar to those found in a 60-yr old field. Results suggest that patterns in C and N cycling within the old-field chronosequence were predictable and highly correlated to the accrual of plant and microbial biomass.     

长白山次生林演替过程中土壤动物群落的变化

通过对长白山白河局不同林龄的次生林土壤动物的组成进行比较。探讨了土壤动物群落随森林采伐更新演替的变化以及演替阶段(增长期、过渡期和稳定期)．研究表明,不同类群土壤动物的演变趋势是腐食类群数量相对较多,其变化趋势接近植被的变化规律,均是经过上升、下降,最后达到稳定;植食类群演替前期在整个群落中所占比例较高,随着演替年代的增加,比例呈下降的趋势;捕食类群经过100年左右的演替,数量明显增加;从昆虫类群组成来看,林龄较短者,未稳定的类群数较多,林龄较长者,未稳定的类群较少．     

Plant strategies and secondary succession on Brittany heathlands after severe fire

Abstract. Plant succession on heathlands subjected to major fire disturbance and humus burn in 1976 was studied over twelve years following burning. Life history strategies of principal heathland species are described with reference to concepts outlined by Grime (1979) and Whittaker & Goodman (1979). Heathlands and closely related communities are characterised by dominance of speciestolerant of physical stress (‘S’ strategists) whereas species which colonise disturbed sites are closer to rude-rals (‘R’ strategists). After severe burning three main successional patterns were identified. They depend on water and nutrient availability relative to temporal population dynamics. Recovery of heathland is often retarded due to monospecific dominance, e.g. of Polytrichum commune, Molinia caerulea and Betula pubescens. These patterns of secondary succession illustrate the inhibition model advanced by Connell & Slatyer (1977).     

N limitation increases along a temperate forest succession: evidences from leaf stoichiometry and nutrient resorption

温带森林演替加剧了氮限制：来自叶片化学计量和养分重吸收的证据 森林生产力和碳汇功能在很大程度上取决于土壤氮和磷的有效性。然而,迄今为止,养分限制随森林演替的时间变化仍存在争议。叶片化学计量和养分重吸收是预测植物生长养分限制的重要指标。基于此,本研究测定了温带森林4个演替阶段所有木本植物叶片和凋落叶中氮和磷的含量,并分析了演替过程中非生物因子和生物因子如何影响叶片化学计量和养分重吸收。研究结果表明,在个体尺度上,叶片氮磷含量在演替末期显著增加,而叶片氮磷比无显著变化;氮的重吸收效率随演替显著增加,然而磷的重吸收效率先增加后减少;氮重吸收效率与磷重吸收效率的比值仅在演替末期显著增加。此外,植物氮素循环对土壤养分的响应比磷素循环更弱。在群落尺度上,叶片氮磷含量随森林演替呈现先降低后升高的趋势,主要受香农-维纳多样性指数和物种丰富度的影响;叶片氮磷比随演替而显著变化,主要由胸径的群落加权平均值决定;氮的重吸收效率增加,主要受物种丰富度和胸径的影响,而磷的重吸收效率相对稳定。因此,氮重吸收效率与磷重吸收效率的比值显著增加,表明随着温带森林演替,氮限制加剧。这些结果可能反映了较高生物多样性群落中物种间对有限资源的激烈竞争,强调了生物因子在驱动森林生态系统养分循环中的重要性,为中国温带和北方森林可持续经营的施肥管理提供了参考。     

The role of remnant forest trees in tropical secondary succession

This paper emphasizes the contribution of remnant trees to the establishment of woody species during succession on abandoned fields and pastures in the Mexican rain forest area, Los Tuxtlas, Veracruz. Remnant trees, original large forest trees left in the clearings by traditional farmers will become natural perching sites for both passing and resident birds. Frugivorous birds drop or regurgitate seeds and fruits which fall under the canopies of remnant trees during their stay, thus contributing to an accumulation of species, which make these remnant trees into ‘regeneration nuclei’. The species transported into these sites belong chiefly to older stages of successional development and reach these otherwise isolated areas, counteracting the depauperization of tropical land, brought about by both intensive and extensive clearing. In a study of seven remnant trees, 29 woody species and two climbers were found, 86% of which are bird dispersed. The total number of species per tree varied from 6 to 15 and was higher under remnant trees with fruits attractive to birds. Floristic variations of the understorey as detected by detrended correspondence analysis was correlated with the relative amount of shade-tolerant primary and late secondary trees versus light dependent pioneers and early successional trees.     

崇明东滩湿地植被演替过程中生物量与氮含量的时空变化

研究了长江口东滩潮滩植物群落各演替阶段优势种的生物量与氮含量的季节变化以及演替过程中的空间变化格局。结果表明,潮滩植被生物量在生长季内随时间呈增长趋势,氮含量则在植被生长初期最大,生长末期最小。在演替过程中,植被总体生物量变化趋势为:演替中期>演替后期>演替早期。植物单位氮含量的变化是:地下部分氮含量随演替呈降低趋势,地上部分氮含量从演替中期开始随演替呈增长趋势。每1 hm2湿地植被氮含量平均值变化趋势为:演替晚期>演替中期>演替早期。根据计算,崇明东滩鸟类自然保护区内芦苇地上部分蓄积的氮含量为227.5 t,互花米草地上部分蓄积的氮含量45.9 t,海三棱草体内蓄积的氮含量为68.2 t。     

营养盐加富和鱼类添加对浮游植物群落演替和多样性的影响

浮游植物是水体生态系统敞水区最重要的初级生产者,其组成与多样性反映了群落的结构类型和存在状态。通过围隔实验,模拟水库春季发生的营养盐加富和鱼类放养的干扰,分析在这两种干扰下的浮游植物群落演替过程中优势种和稀有种的变化,并通过以丰度与生物量为变量的香农和辛普森多样性指数的计算,分析浮游植物群落演替过程中的多样性变化特征。结果表明,营养盐加富干扰下的浮游植物群落的优势种变化和演替更为明显,营养盐加富与鱼类添加对浮游植物群落多样性变化的影响符合中度干扰理论。在优势种优势度变化较大的浮游植物群落演替过程中,多样性指数与浮游植物生物量有较高的负相关性。在浮游植物群落演替过程中,香农和辛普森多样性指数的变化趋势基本一致,采用丰度与生物量为变量的两种多样性指数的计算结果对实验系统中浮游植物群落多样性的分析结果没有明显的影响。     

Characterisation of the type and extent of nutrient limitation in grassland vegetation using a bioassay with intact sods

An important methodological problem in plant ecology concerns the way in which the type and extent of nutrient limitation in terrestrial communities should be assessed. Conclusions on nutrient limitation have been founded mainly on soil extractions, fertiliser trials and tissue nutrient concentrations. In order to avoid some of the problems associated with these methods, we employed a special technique using intact sods which rooted both in the intact soil and in a nutrient solution, from which N, P and K were omitted stepwise. The method was applied to hay-field communities which differed in their history of fertiliser application. Four fields were compared which were not fertilised for 2, 6, 19 or 45 years, while hay making continued. This was done to restore former species-rich grassland communities. We tested the hypothesis that the increase in species diversity in these grasslands was attended by an increase in the number of limiting nutrients.We observed clear shifts in the type and extent of nutrient limitation. Fields which were recently fertilised were characterised by nitrogen and potassium limitation, while phosphorus limitation increased in importance towards the later stages of succession. In the last field (45 years unfertilised) N, P and K equally limited production at the community level. These conclusions differed from those drawn from a fertiliser trial in these same four fields, which failed to detect phosphorus limitation.It is concluded that the use of this method provides a valuable extra source of information while studying relationships between nutrient limitation and species diversity in grassland communities.     

Changes in the seasonal rhythm of two forest communities during secondary succession

Changes in the seasonal rhythm of two plant phytocoenoses in a submountain beech forest during secondary succession were studied. Investigations were done on four monitoring plots with different stand density over the period of four successive years. The rhythm of the associations Dentario bulbiferae-Fagetum and Carici pilosae-Fagetum reflects the course of succession processes running six years after the human impact (cutting) in the ecosystem. Results of the phenological observations of the understorey species with the focus on the changes in flowering and colour spectrum allowed to make the comparisons between both associations in connection with different phyto-climatic conditions and in dependence on time. The most conspicuous changes in the seasonal rhythm and structure of the examined associations were found in conditions of the former clear-cut, currently in succession phase. A clear decrease (56%) in number of taxons with the dominance > 1% in one association towards the end of the 4-year study period was detected here. Simultaneously, a decrease in the number of flowering species was observed, while the relative rate of species being in the vegetative stage increased considerably (from 6 to 67%) over the growing season. The course of flowering of both of the associations missed discernible trends and peaks as well as colour spectra were partially changed during four monitored successive years on the formerly unstocked area.     

Towards comparable assessment of the soil nutrient status across scales—Review and development of nutrient metrics

Nutrient availability influences virtually every aspect of an ecosystem, and is a critical modifier of ecosystem responses to global change. Although this crucial role of nutrient availability in regulating ecosystem structure and functioning has been widely acknowledged, nutrients are still often neglected in observational and experimental synthesis studies due to difficulties in comparing the nutrient status across sites. In the current study, we explain different nutrient‐related concepts and discuss the potential of soil‐, plant‐ and remote sensing‐based metrics to compare the nutrient status across space. Based on our review and additional analyses on a dataset of European, managed temperate and boreal forests (ICP [International Co‐operative Programme on Assessment and Monitoring of Air Pollution Effects on Forests] Forests dataset), we conclude that the use of plant‐ and remote sensing‐based metrics that rely on tissue stoichiometry is limited due to their strong dependence on species identity. The potential use of other plant‐based metrics such as Ellenberg indicator values and plant‐functional traits is also discussed. We conclude from our analyses and review that soil‐based metrics have the highest potential for successful intersite comparison of the nutrient status. As an example, we used and adjusted a soil‐based metric, previously developed for conifer forests across Sweden, against the same ICP Forests data. We suggest that this adjusted and further adaptable metric, which included the organic carbon concentration in the upper 20 cm of the soil (including the organic fermentation‐humus [FH] layer), the C:N ratio and of the FH layer, can be used as a complementary tool along with other indicators of nutrient availability, to compare the background nutrient status across temperate and boreal forests dominated by spruce, pine or beech. Future collection and provision of harmonized soil data from observational and experimental sites is crucial for further testing and adjusting the metric.     

缙云山森林群落次生演替中土壤理化性质的动态变化

选择典型、有代表性的不同演替阶段群落,进行了植被调查和土壤分析.结果表明,土壤理化性质在演替方向和土壤剖面上表现出较强的规律.土壤有机质随植被从低级向高级演进逐渐积累,分别是19.5(X₁)、37.0(X²)、50.1(X₃)和71.6 g·kg^-1(X₄);土壤全N、碱解氮和速效钾等也呈上升趋势;土壤pH和盐基饱和度降低,阳离子交换量增加.在土壤剖面上,有机质、全N等指标表现出A＞B＞C层的趋势.灰色关联度分析表明,随着演替的进行,土壤肥力提高,物种丰富度和郁闭度也相应增加.不同群落土壤理化性质在不同季节有显著差异,但这种差异并不影响植物与土壤在大时间尺度下的演变方向.土壤理化性质的动态变化与植物演替相适应.     

云南松林的根系生物量及其分布规律的研究

利用平均标准木机械布点法测定了云南省永仁林业局云南松不同龄组林分的根系生物量及其沿土壤剖面深度的分布规律.结果表明,林分根系总生物量随林龄而增加,幼龄林(15～17年)的根系生物量为8.50 t·hm^-2,中龄林(30～32年)为11.70 t·hm^-2,成熟林为(＞62年)18.91 t·hm^-2.在不同龄组林分中,粗根(＞10mm)生物量差异最大(1.5～12.3 t·hm^-2),而中根(5～10 mm)(1.4～1.6 t·hm^-2)及小根(＜5 mm)(5.3～6.2 t·hm^-2)的生物量差异最小.根系生物量沿土壤深度迅速减少,约93%的根系生物量集中分布在0～30 cm土层中,深土层(30～115 cm)的根系生物量仅占7%左右.     

Changes in the microbial community structure during thermophilic composting of manure as detected by the quinone profile method

Quinone profiles and physico-chemical properties were measured to characterize the microbial community structure during a 14-day thermophilic composting of cattle manure mixed with rice straw as a bulking agent. The change in total quinone content (TQ) and the divergence of quinone (DQ) indicated that the microbial biomass reached a peak followed by a decrease, whereas the microbial community diversity increased continuously during the composting process. The high mole fraction of ubiquinones in the raw materials, and at the beginning of the composting period suggested that fungi and/or Proteobacteria were present. The predominance of MK-7 from days 3 to 7 suggested that Bacillus spp. were the main microbial species. An increase in partially saturated and long-chain menaquinones during the latter composting period indicated that the proliferation of various species of Actinobacteria was occurring. The microbial community structure, as expressed by TQ and DQ, corresponded well to physico-chemical properties such as the C/N ratio, pH, O₂ consumption and compost mass reduction.     

黄土高原次生林演替过程土壤氮矿化特征及其影响因素

土壤氮矿化速率限制着森林土壤有效氮素供给能力,但目前关于黄土高原次生林演替过程土壤氮矿化特征的认识还不清楚。选择黄土高原黄龙山次生林区草本阶段、灌木阶段、先锋乔木群落阶段、混交森林群落阶段和顶级森林群落阶段等5个演替阶段的9种群落类型,测定了其林下土壤氮矿化速率,并分析了土壤理化性质、凋落物养分特征和土壤酶活性及其对氮矿化的影响。结果表明:随着演替进展,土壤有机碳和氮不断积累,土壤硝态氮和铵态氮含量逐渐增加,凋落物养分和土壤酶活性也发生了显著变化。土壤硝化速率从草本群落阶段到顶级森林群落阶段显著增加了71.73%;土壤氨化速率一直为负值且持续负向增加;土壤净矿化速率从草本群落阶段的(0.16±0.06) mg kg^-1 d^-1逐渐增加到混交森林阶段的(0.31±0.08)mg kg^-1 d^-1,但从混交林群落到顶级森林群落阶段有所下降。土壤理化性质(路径系数-0.530)和酶活性(路径系数-0.268)对氮矿化速率有显著的直接作用;凋落物养分对氮矿化速率的影响来自于其直接作用(路径系数-0.283)和通过调控土壤理化性质、酶活性产生的间接影响(路径系数-0.315)。结果有助于认识黄土高原植被演替过程中土壤氮循环过程,对森林可持续经营也有一定的意义。     

Ecosystem properties and microbial community changes in primary succession on a glacier forefront

We studied microbial community composition in a primary successional chronosequence on the forefront of Lyman Glacier, Washington, United States. We sampled microbial communities in soil from nonvegetated areas and under the canopies of mycorrhizal and nonmycorrhizal plants from 20- to 80-year-old zones along the successional gradient. Three independent measures of microbial biomass were used: substrate-induced respiration (SIR), phospholipid fatty acid (PLFA) analysis, and direct microscopic counts. All methods indicated that biomass increased over successional time in the nonvegetated soil. PLFA analysis indicated that the microbial biomass was greater under the plant canopies than in the nonvegetated soils; the microbial community composition was clearly different between these two types of soils. Over the successional gradient, the microbial community shifted from bacterial-dominated to fungal-dominated. Microbial respiration increased while specific activity (respiration per unit biomass) decreased in nonvegetated soils over the successional gradient. We proposed and evaluated new parameters for estimating the C use efficiency of the soil microbial community: “Max” indicates the maximal respiration rate and “Acc” the total C released from the sample after a standard amount of substrate is added. These, as well as the corresponding specific activities (calculated as Max and Acc per unit biomass), decreased sharply over the successional gradient. Our study suggests that during the early stages of succession the microbial community cannot incorporate all the added substrate into its biomass, but rapidly increases its respiration. The later-stage microbial community cannot reach as high a rate of respiration per unit biomass but remains in an “energy-saving state,” accumulating C to its biomass. Received: 4 June 1998 / Accepted: 11 January 1999     

黑石顶自然保护区森林次生演替过程中群落主要种的种间协变分析

采用2×2列联表,通过Pearson积矩相关系数和Speaman秩相关系数对黑石顶森林次生演替过程中各阶段群落优势种的种间协变进行了定量分析。结果表明,在演替初期的裸地恢复阶段,显著种间协变的种对较多,有14对(正协变9对负协变5对),在针阔混交林阶段,显著种间协变的种对较少,仅有5种。全为正协变,表现为主要种间对群落环境的相同要求,并形成了主要种间的竞争格局,在阳生性常绿阔叶林阶段。显著种间协变的种对最少,只有4对,其中3对为负协变,这是主要种之间对群落环境相同要求的激烈竞争结果,在中生性常绿阔叶林阶段,显著种间协变的种对最多有20对,19对为正协变,主要表现为群落上层种和中下层种间形成了大量的正协变,但群落上层主要种间无显著的种间协变。     

热带人工林土壤动物群落的次生演替和发展过程探讨

在广东小良地区的桉林迹地上,对不同林龄的阔叶混交林土壤动物的组成进行比较,探讨群落随林龄的增加而变化的过程。以多样性为基础,可将这里土壤动物群落的演替和发展分成3个阶段,即增长期、过渡期和稳定期。在林龄20多年后,群落发展可进入最后阶段。不同类群的演变程序:(1)湿生动物个体数逐渐增加,直至平稳;(2)昆虫中不稳定类群逐渐减少;(3)白蚁种群在增长期前段迅速增加,随后即在较低水平上受到控制;(4)蚯蚓在增长期后段才出现,并迅速成为优势类群。过渡期及其以前的蚯蚓种类主要为寒(虫宪)蚓属(Ocnerodrilus),稳定后为环毛蚓属(Pheretima)。     

The role of environmental, root, and microbial biomass characteristics in soil respiration in temperate secondary forests of Northeast China

For secondary forests, the major forest resources in China (accounting for more than 50% of the national total), soil respiration (R _S) and the relationship between R _S and various biotic/abiotic factors are poorly understood. The objectives of the present study were to examine seasonal variations in soil respiration during the growing season, and to explore the factors affecting the variation in soil respiration rates for three forest types (Mongolian oak, Manchurian walnut and mixed forests) of temperate secondary forest in Northeast China. The results showed that (1) the maximum total R _S rate occurred in July, following a bell-shaped curve with season, (2) for all forest types, the total R _S was significantly influenced by soil temperature (P < 0.01), and did not significantly correlate with soil moisture, (3) compared with fine root biomass, coarse root biomass was more closely related with the root respiration in mixed forest (R ² = 0.711, P = 0.017) and in Manchurian walnut forest (R ² = 0.768, P = 0.010), and (4) microbial biomass carbon (MBC) and nitrogen were significantly correlated with heterotrophic R _S in Mongolian oak forest (R ² = 0.664, P = 0.026; R ² = 0.784, P = 0.008, respectively) and in mixed forest (R ² = 0.918, P = 0.001; R ² = 0.967, P = 0.001, respectively). We can conclude that in temperate secondary forests: (1) the R _S rate and the relationships between R _S and abiotic/biotic factors change greatly with forest types, and (2) R _S is strongly influenced by soil temperature, MBC, microbial biomass nitrogen and coarse root biomass in temperate secondary forests.