Effect of wood ash and nitrogen fertilization on soil chemical properties, soil microbial processes, and stand growth in two coniferous stands in Finland

The aim of our study was to investigate long-term effects of wood ash fertilization, given together with nitrogen, on soil chemical properties, soil microbiological processes related to C and N cycling, and tree growth. The study was carried out in a 31-year-old Scots pine stand and in a 45-year-old Norway spruce stand 15 years after application. The treatments were (1) a control with no ash or nutrient addition, (2) wood ash + N (WAN), and (3) a stand-specific fertilization (SSF) formulated on the basis of analyses carried out on needle and soil samples taken from the stand. The SSF treatments included N, Cu and B, and in the spruce stand also P. WAN decreased acidity and increased the extractable Ca, Mg and P concentrations in the organic layer in both stands, but SSF had no effect. The microbial processes reacted more strongly to the treatments in the pine stand, whereas the growth response, although only relatively slight during the third 5-year period after fertilization, was detected only in the spruce stand. WAN increased the NH₄-N concentrations in the organic layer compared to the control and SSF treatments on both sites. In the pine stand, amount of N in microbial biomass and both the C and net N mineralization rates were significantly higher in the WAN treatment than in the SSF treatment. On both sites net nitrification was negligible in all treatments. Soil microbial biomass, microbial respiration and N availability have been used as indices for assessing the biological activity and health of soil, and these parameters either increased or were not affected by the WAN treatment. Hence, with regard to these parameters there are justifiable grounds for applying wood ash.     

Colonisation of spruce roots by two interacting ectomycorrhizal fungi in wood ash amended substrates

Interactions between two ectomycorrhizal fungal species, Piloderma croceum Erikss. and Hjortst. and Piloderma sp. 1 (found to colonise spruce roots and wood ash granules in the field), were investigated in wood ash amended substrates. The comparative ability of these fungi to colonise roots of non-mycorrhizal spruce (Picea abies (L.) Karst.) seedlings was studied in relation to factorial combinations of wood ash and N fertilisation. Non-mycorrhizal spruce seedlings (bait seedlings) were planted together with spruce seedlings colonised by P. croceum or Piloderma sp. 1. The growth substrate was a sand-peat mixture with wood ash or no ash and supplied with two levels of N, so that four substrate combinations were obtained. Piloderma sp. 1 mycelia colonised around 60% of the fine roots of bait seedlings in ash treatments regardless of N level and around 20-26% in treatments without ash. P. croceum only colonised 8% of the root tips in the presence of ash but 56% of the root tips in the low-N treatment without ash. However, in the high-N treatment without ash the colonisation level was reduced to around 30%. Total numbers of root tips per seedling did not vary significantly between the treatments. Possible reasons for the competitive advantage of Piloderma sp. 1 in wood ash fertilised substrate are discussed.     

Ectomycorrhizal fungal biomass in roots and uptake of P from apatite by Pinus sylvestris seedlings growing in forest soil with and without wood ash amendment

Forest soil from an experimental Norway spruce forest with four levels of wood ash addition (0, 1, 3 and 6 tonnes ha^–1) was used to inoculate pine (Pinus sylvestris) seedlings with indigenous ectomycorrhizal (EM) fungi. Uptake of ³²P and ⁸⁶Rb in a root bioassay was used to estimate the demand for P and K by seedlings grown in the different soils. Utilisation of P from apatite was tested in a laboratory system where uptake by the ectomycorrhizal mycelium was separated from uptake by roots. The demand for P and K in the seedlings was similar regardless of the ash treatment. Variation in EM levels, estimated as fungal biomass (ergosterol) in roots, was large in the different soils, but not related to ash addition. Uptake of P from apatite was, on average, 23% of total seedling P and was not related to EM levels. It was concluded that the improved P uptake from apatite by EM fungi found in earlier studies is probably not a general phenomenon among EM fungi. The small effect of ash addition on EM levels and P uptake suggests that addition of granulated wood ash is a forest management treatment that will have only minor influence on ectomycorrhizal symbiosis.     

Influence of elevated ozone and limited nitrogen availability on conifer seedlings in an open-air fumigation system: effects on growth, nutrient content, mycorrhiza, needle ultrastructure, starch and secondary compounds

Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies Karst.) seedlings were exposed to realistically elevated O₃ levels in open‐air experiments over three growing seasons. The total O₃ exposure doses were 1.2 × (1991), 1.5 × (1992) and 1.7 × (1993) ambient levels. During the 1992 and 1993 growing seasons pine and spruce seedlings received two different levels of nitrogen supply. Effects on growth, mycorrhiza formation, needle ultrastructure, primary and secondary compounds were studied. Ozone exposure had only slight effects on biomass production, growth height and nutrient content of studied conifers. Higher nitrogen availability improved growth of the seedlings and resulted in higher concentration of nitrogen in needles. In Scots pine O₃ exposure did not have effects on quantity of total mycorrhizas and short roots, while higher nitrogen availability decreased quantity of mycorrhizas and short roots. In both tree species O₃ exposure induced O₃‐related ultrastructural symptoms, e.g. granulation and dark staining of the chloroplast stroma in the needle mesophyll cells, at both nitrogen availability levels. Ozone exposure and nitrogen availability did not have significant effects on starch concentrations in either tree species. Concentrations of some individual terpenes were higher in O₃‐exposed needles, while concentrations of individual and total resin acids, total phenolics and catechins were not affected by O₃ exposure. Nitrogen availability did not have substantial effects on concentrations of monoterpenes. By contrast, concentrations of some individual and total resin acids were lower in pine needles and higher in spruce needles with higher nitrogen availability, while phenolic concentration in spruce needles decreased at higher nitrogen availability. The results suggest that realistically elevated levels of O₃ in the field can have some negative effects on the mesophyll ultrastructure of conifer needles, but carbon allocation to root and shoot growth and secondary metabolites are not affected substantially.     

Effects of wood ash and nitrogen fertilization on fine root biomass and soil and foliage nutrients in a Norway spruce stand in Finland

We determined the effects of wood ash fertilization, given together with nitrogen (WAN), and nitrogen given together with P, B and Cu (SSF), on soil and foliage nutrients and fine root biomass in a 45-year-old Norway spruce stand in southern Finland. Fine roots were sampled 9 years, and the soil 10 years after ash (3 t/ha) and nitrogen (150 kg/ha) application. Fine root biomass tended to be lower, the necromass higher, and the fine root distribution relatively deeper on the WAN than on the control and SSF plots. The response of fine root biomass to WAN was probably related to changes in soil acidity. pH, base saturation, total and extractable concentrations of Ca, K, Mg and P, and total B, Cd, Mn, Ni and Zn concentrations in the organic layer were significantly higher on the WAN plots than on the SSF and the control plots with no ash and nutrient addition. On the WAN plots, the pH was 1.2 pH-units higher, the exchangeable Ca concentrations fourfold and those of Mg over twofold compared to the control plots. WAN increased the concentrations of K but decreased those of Mn and Ni in the needles compared to the control and SSF treatment. Even though ash and nitrogen fertilisation tended to decrease the fine root biomass, this decrease was not likely to affect tree growth during a 10-year period.     

Frost damage and winter nitrogen uptake by the grass Poa pratensis L.: consequences for vegetative versus reproductive growth

Frost damage can decrease nitrogen uptake by grasses over winter, and it can also decrease biomass production over the following growing season. However, it is not clear to what extent reduced nitrogen uptake over winter decreases grass production, or whether is it merely a symptom of root damage. We examined the growth response of the grass Poa pratensis L. (Kentucky bluegrass) to variation in the timing of freezing and nitrogen availability over winter in London, Ontario, Canada. All tillers were transplanted into untreated soil in early spring, and at peak seed maturation, root, shoot, and reproductive biomass were measured. There was an interaction between freezing and increased winter nitrogen availability, whereby nitrogen addition increased tiller biomass under ambient temperatures, but decreased tiller biomass in combination with a late winter freeze. The nitrogen response of ambient temperature tillers occurred primarily via increased seed production, whereas for frozen tillers seed production was generally absent. Our results support the hypothesis that nitrogen uptake over winter can increase growing season productivity in P. pratensis, but also demonstrate that increased nitrogen availability increases tiller vulnerability to frost. These results have important implications for grass responses to the alteration of soil freezing dynamics with climate change.     

Effect of wood ash application on the biomass distribution and physiological state of Norway spruce seedlings on sandy soils

The effect of wood ash applied as a fertiliser (0.25, 0.5 and 1 kg m^–2) to nutrient poor sandy soil on 4-year-old Norway spruce (Picea abies) growing in small-scale sample plots was studied. Analyses carried out with roots, stems, needles and shoots of different age showed that an increase in the pH level and K and Ca concentrations and a decrease in N and P concentrations in the soil was accompanied by a disbalance of nutrients in tree compartments. Stimulation of pigment synthesis in Norway spruce needles was observed, and no disbalances occurred in the pigment system (Chl a/Chl b, TChl/Car) guaranteeing normal process of photosynthesis. Biomass responses to wood ash application depended on the age of needles and shoots, being only slightly notable in current year organs. Inhibition of height growth of seedlings, but stimulation of root biomass was established.     

Physiological and growth responses of Centaurea maculosa (Asteraceae) to root herbivory under varying levels of interspecific plant competition and soil nitrogen availability

Summary Centaurea maculosa seedlings were grown in pots to study the effects of root herbivory by Agapeta zoegana L. (Lep.: Cochylidae) and Cyphocleonus achates Fahr. (Col.: Curculionidae), grass competition and nitrogen shortage (each present or absent), using a full factorial design. The aims of the study were to analyse the impact of root herbivory on plant growth, resource allocation and physiological processes, and to test if these plant responses to herbivory were influenced by plant competition and nitrogen availability. The two root herbivores differed markedly in their impact on plant growth. While feeding by the moth A. zoegana in the root cortex had no effect on shoot and root mass, feeding by the weevil C. achates in the central vascular tissue greatly reduced shoot mass, but not root mass, leading to a reduced shoot/root ratio. The absence of significant effects of the two herbivores on root biomass, despite considerable consumption, indicates that compensatory root growth occurred. Competition with grass affected plant growth more than herbivory and nutrient status, resulting in reduced shoot and root growth, and number of leaves. Nitrogen shortage did not affect plant growth directly but greatly influenced the compensatory capacity of Centaurea maculosa to root herbivory. Under high nitrogen conditions, shoot biomass of plants infested by the weevil was reduced by 30% compared with uninfested plants. However, under poor nitrogen conditions a 63% reduction was observed compared with corresponding controls. Root herbivory was the most important stress factor affecting plant physiology. Besides a relative increase in biomass allocation to the roots, infested plants also showed a significant increase in nitrogen concentration in the roots and a concomitant reduction in leaf nitrogen concentration, reflecting a redirection of the nitrogen to the stronger sink. The level of fructans was greatly reduced in the roots after herbivore feeding. This is thought to be a consequence of their mobilisation to support compensatory root growth. A preliminary model linking the effects of these root herbivores to the physiological processes of C. maculosa is presented.     

Nutrient and Carbon Additions to the Microbial Soil Community and its Impact on Tree Seedlings in a Boreal Spruce Forest

We have added glucose and nutrients to manipulate soil microbial activity and nutrient availability in a boreal spruce forest to study the performance of birch and spruce seedlings in relation to the soil microbial community. The proportion of aboveground biomass in the seedlings was largest in plots amended with extra nutrients, while ectomycorrhizal (ECM) colonisation was low in these plots. ECM appeared beneficial for growth of both species, but only at low levels of colonisation (<25% ECM colonised root-tips). The soil microbial biomass, as determined by total PLFA, was largest in plots treated with glucose and there was a significant negative relationship between birch seedling size and levels of total PLFA in soil. This could be taken to suggest that poor seedling growth was due to nutrient limitation caused by microbial assimilation. However, the treatment response of the birch seedlings was generally weak, and spruce often showed no response at all to the addition of nutrients and glucose. The most consistent parameter for the variation in plant performance, as well as for the microbial soil community, was the block-effect. This suggests a strong spatial structure in the soil microbial community, and that this structure was robust with respect to our treatments even though they continued over a 3-year period.     

Nitrate reduction and nitrate content in ash trees (Fraxinus excelsior L.): distribution between compartments,site comparison and seasonal variation

Summary Nitrate reductase activity (NRA), nitrate content and biomass components of leaflets, leaf stalks, old stem, current-year stem and roots of ash trees (Fraxinus excelsior L.) growing in their natural habitats were investigated. In addition, NRA, total nitrogen and nitrate concentration were analyzed in the leaves and roots of ash trees from four different field sites. The highest NRA per gram biomass and also per total compartment biomass was found in the leaflets, even though root biomass was much higher than total leaflet biomass. The highest nitrate concentrations were found in the leaf stalks. Correlations between nitrate availability in the soil and NRA in leaves were not significant due to high variability of the actual soil nitrate concentrations. The seasonal variation in foliar NRA, nitrate concentration and total nitrogen concentration is much smaller in F. excelsior than reported for herbaceous species and is mainly caused by changes in the actual soil nitrate availability and by senescence of the leaves.     

Responses in plant, soil inorganic and microbial nutrient pools to experimental fire, ash and biomass addition in a woodland savanna

In order to investigate the effects of savanna fires on nutrient cycling a field experiment was carried out in an open woodland savanna of southwest Ethiopia. This involved manipulations of fire, fuel load and ash fertilisation in a fully factorial design, and recording of responses in plants, soil inorganic and microbial nutrient pools up to 1 year after the disturbances. As plant biomass nitrogen (N) was only 3.5% of that in topsoil the N loss in a single fire event was relatively small. The microbial N pool size in the topsoil was similar to the N pool size in the aboveground part of the plants. Soil microbial biomass carbon increased slightly 12 days after the low severity fire, but the effect was transient and was not accompanied by an increase in microbial N. Instead, the soil inorganic N concentration increased strongly 1 day after the fire, remained higher up to 3 months after the fire and probably caused the 40% higher grass biomass in burned than unburned plots, and the similar sized increase in grass nitrogen, phosphorus and potassium pools in the following rainy season. In contrast, broad-leaved herbs showed less strong increments in biomass and nutrient pool sizes. Fire interacted with fuel load, as burning of plots with double plant biomass led to reduced microbial biomass, plant nutrient pools and herb (but not grass) biomass. Low-severity-fire nutrient losses appear to be moderate and may be replenished from natural sources. However, in areas with frequent fires and high grass biomass (fuel) loads, or with late fires, nutrient losses could be much larger and non-sustainable to the persistence of the woodland savanna ecosystem.     

施氮肥对落叶松和水曲柳人工林土壤动物群落的影响

土壤动物对环境变化反应敏感, 全球变化导致土壤氮(N)有效性增加将影响土壤动物群落结构和功能。本文以落叶松(Larix gmelinii)和水曲柳(Fraxinus mandshurica)人工林为研究对象, 通过施肥处理, 在不同季节和土层取样, 研究土壤N有效性增加对土壤动物的数量、类群数和不同功能团的影响。结果表明: (1) 施肥影响两林分土壤动物密度, 导致当年密度增加, 翌年则下降, 这种先增加后降低的趋势在不同土层中均表现出来; (2) 施肥增加了两林分土壤动物类群数, 其中落叶松林分从34类增加到43类, 水曲柳林分从43类增加到48类; (3) 施肥改变了两林分不同食性土壤动物的密度, 腐食性土壤动物数量降低、植食性数量增加、捕食性数量变化不明显。这些结果说明: 土壤N有效性增加显著影响两林分土壤动物群落结构, 可能改变地下碳分配格局和养分循环过程。     

The significance of Cognettia sphagnetorum(Enchytraeidae) on nitrogen availability and plant growth in wood ash-treated humus soil

A laboratory microcosm experiment was established to study whether the role of Cognettia sphagnetorum (Enchytraeidae) in affecting Scots pine (Pinus sylvestris) seedling growth is influenced by wood ash-amendment, i.e., neutralisation of the raw humus soil. Coniferous forest soil, inoculated with soil microbes and nematodes, was either treated with wood ash or left as ash-free control. Wood ash (corresponding to an amount of 5000 kg ha^–1) was either spread on the soil surface or mixed into the soil. Enchytraeid and pine seedling biomass, abundance of nematodes, and water leachable NH₄ ⁺-N and NO₃ ^–-N were measured 26 and 51 weeks after initiation of the experiment and root length and N concentration of needles were measured 51 weeks after initiation of the experiment. Wood ash when mixed into the soil, reduced the biomass of C. sphagnetorum. Nematodes were unaffected by the treatments. In the ash-free soils C. sphagnetorum had little influence on pine growth, but it did decrease root length and root to shoot ratio. In the absence of enchytraeids wood ash decreased pine biomass production and root length. However, the presence of enchytraeids in the ash-treated soil compensated the ash-induced negative effects on the pine performance. Enchytraeids increased and wood ash decreased water leachable NH₄ ⁺-N in the presence but not in the absence of enchytraeids, while water leachable NO₃ ^–-N was not affected by the treatments. It was concluded that C. sphagnetorum can be important in ensuring nutrient cycling and plant growth in situations when an ecosystem encounters disturbances.     

Short-term regrowth responses of four steppe grassland species to grazing intensity, water and nitrogen in Inner Mongolia

Inner Mongolia steppe grasslands are widely used for livestock farming and the regrowth ability of grassland species is therefore strongly influenced not only by water and nutrient availability but also quite heavily by grazing. However, little is known on how grazing, water and nitrogen interactively affect the dominant C3 species (Leymus chinensis, Stipa grandis, Agropyron cristatum) and the C4 species (Cleistogenes squarrosa). Therefore in the 2007 and 2008 growing seasons, a field experiment was carried out to test the hypothesis that under different grazing intensities the dominant species show different short-term regrowth response to simultaneous variation in the availability of water and nitrogen. Single factor and interaction effects of the addition of water (rainfed vs. simulated wet-year) and nitrogen (0 or 25 kg N ha^?1) were analysed along a gradient of four grazing intensities (ungrazed, lightly, moderately and heavily grazed) after one month of grazing exclusion. Water and nitrogen addition affected short-term regrowth of all species in a similar way whereas species responded differently to grazing. Simulated wet-year water availability consistently resulted in higher standing biomass, relative growth rate and cellulase digestible organic matter yield. Supplementary nitrogen promoted standing biomass and crude protein concentration. The nutritive value of all species’ standing biomass showed a similar increase with more intensive grazing. However, heavy grazing led to a clear shift in the relative biomass of the species, i.e. mainly a promotion of the C4 grass, C. squarrosa. In contrast to our hypothesis, there were no differences among species in their response to water or nitrogen addition, whereas, heavy grazing induced the expected species-specific response. Our results suggest that heavy grazing rather than nitrogen or water determine short-term shifts in species composition of the investigated steppe ecosystem. Furthermore, differences in the species-specific growth response to grazing may increase the proportion of the C4 grass C. squarrosa in steppe communities, whereas higher availability of nitrogen and water may lead to higher forage biomass and nutritive value of all investigated species but in short-term cannot compensate for the grazing induced changes in species composition.     

Long-term addition of fertilizer,labile carbon,and fungicide alters the biomass of plant functional groups in a subarctic-alpine community

In subarctic ecosystems, plant growth is mostly limited by nutrient availability and harsh climate. Investigating how soil nutrient availability controls the plant community composition may therefore help to understand indirect effects of climate change. The study was conducted in a long-term field experiment on a subarctic-alpine fellfield dominated by woody evergreen shrubs, bryophytes, and lichens. To manipulate nutrient availability additions of NPK fertilizer, labile C, and fungicide (benomyl) were done in a fully factorial design, replicated in six blocks. The treatments were run for 10 years, and the aboveground plant biomass was harvested 4 and 16 years after initiating the experiment. In addition, soil inorganic N and P concentration was analyzed the same years. Increased nutrient availability (NPK fertilizer) largely increased the biomass of graminoids and unexpectedly of bryophytes, but not of other vascular plant groups. Also, limitation of soil nutrient availability caused by labile C addition decreased the relative proportion of green shoots in evergreen shrubs, although these were expected to cope better with the nutrient limitation than the opportunistic graminoids, which, by contrast, were unaffected. Reduced fungal biomass due to benomyl addition was accompanied by increased evergreen shrub and clubmoss biomass. Taken together, the effects of treatments were most pronounced 16 years after initiation of the experiment, but despite changes in biomass the overall plant community composition was resistant to environmental changes.     

Growth and reproduction of aphids and levels of free amino acids in Scots pine and Norway spruce in an open-air fumigation with ozone

To estimate the susceptibility of conifer seedlings to aphids under future tropospheric ozone levels, Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings were exposed to ambient and elevated ozone levels in an open-air exposure system. Growth and reproduction of the aphids Schizolachnus pineti and Cinara pinea on Scots pine and Cinara pilicornis on Norway spruce were monitored. Levels of free amino acids in foliage and young shoots were used as indicators of host plant quality. In elevated treatment plots the ozone doses were between 1.2 and 1.7 times the dose in ambient plots in 1990–93. Half of the seedling material in 1992–93 was subjected to nitrogen fertilization treatment to evaluate the effects of increased N deposition. In 1990, population density of S. pineti on pine did not differ between ambient and elevated ozone treatments during growing season, but remained higher in the elevated ozone plot than in the ambient plot at the end of the growing season. This was associated with elevated levels of glutamic acid in foliage. In August 1992, the numbers of S. pineti were consistent between the two ambient ozone plots, but deviated highly between the two ozone-fumigated plots. Glycine concentration in pine foliage was elevated by ozone, but free amino acid concentrations were not related to aphid performance. In 1993, ozone and nitrogen did not significantly affect the relative growth rate (RGR) of S. pineti or C. pinea nymphs on Scots pine, but glutamic acid concentration in foliage was increased by nitrogen fertilization. On Norway spruce, fecundity of C. pilicornis females was higher in elevated ozone treatment, but RGR of nymphs was not affected in 1992. In 1993, RGR of C. pilicornis nymphs was increased by nitrogen fertilization in June, but not affected by ozone. Nitrogen fertilization increased the levels of total free amino acids, aspartic acid, glutamic acid and proline in elongating shoots of Norway spruce, and ozone reduced the concentrations of valine and γ-butyric acid. Our results suggest that availability of nitrogen from soil has a stronger impact on the concentrations of free amino acids in conifer seedlings than ozone. Some episodes of high ozone concentration may increase free amino acids in foliage. Aphid response to ozone was extremely variable, in agreement with previous laboratory experiments. The expected 20–70% increase in ambient concentrations of tropospheric ozone may in some occasions enhance aphid performance on Scots pine and Norway spruce seedlings, but in most cases the ozone effect on the susceptibility of conifer seedlings to sucking insect pests is not important.     

Response of ecosystem carbon exchange to warming and nitrogen addition during two hydrologically contrasting growing seasons in a temperate steppe

A large remaining source of uncertainty in global model predictions of future climate is how ecosystem carbon (C) cycle feedbacks to climate change. We conducted a field manipulative experiment of warming and nitrogen (N) addition in a temperate steppe in northern China during two contrasting hydrological growing seasons in 2006 [wet with total precipitation 11.2% above the long‐term mean (348 mm)] and 2007 (dry with total precipitation 46.7% below the long‐term mean). Irrespective of strong intra‐ and interannual variations in ecosystem C fluxes, responses of ecosystem C fluxes to warming and N addition did not change between the two growing seasons, suggesting independence of warming and N responses of net ecosystem C exchange (NEE) upon hydrological variations in the temperate steppe. Warming had no effect on NEE or its two components, gross ecosystem productivity (GEP) and ecosystem respiration (ER), whereas N addition stimulated GEP but did not affect ER, leading to positive responses of NEE. Similar responses of NEE between the two growing seasons were due to changes in both biotic and abiotic factors and their impacts on ER and GEP. In the wet growing season, NEE was positively correlated with soil moisture and forb biomass. Negative effects of warming‐induced water depletion could be ameliorated by higher forb biomass in the warmed plots. N addition increased forb biomass but did not affect soil moisture, leading to positive effect on NEE. In the dry growing season, NEE showed positive dependence on grass biomass but negative dependence on forb biomass. No changes in NEE in response to warming could result from water limitation on both GEP and ER as well as little responses of either grass or forb biomass. N addition stimulated grass biomass but reduced forb biomass, leading to the increase in NEE. Our findings highlight the importance of changes in abiotic (soil moisture, N availability) and biotic (growth of different plant functional types) in mediating the responses of NEE to climatic warming and N enrichment in the semiarid temperate steppe in northern China.     

夜间增温和施氮对川西亚高山针叶林土壤有效氮和微生物特性的短期影响

开展亚高山针叶林典型林地土壤有效氮和微生物特性对气候变化的响应研究, 对预测未来气候变化背景下亚高山针叶林生态系统C、N的源/汇功能具有重要意义。该文采用红外辐射加热器模拟增温结合外施氮肥的方法, 研究了川西亚高山针叶林下土壤化学特性、有效氮含量以及微生物生物量对夜间增温和施氮的短期响应。结果表明: 在模拟增温试验期间(2009年4月-2010年4月), 空气平均温度和5 cm土壤平均温度分别比对照提高了1.93和4.19 ℃, 增温幅度分别以夏季和冬季最为显著。增温对土壤pH值、有机碳、全氮和微生物生物量无显著影响。增温在试验前期降低了土壤NH₄ ⁺-N含量, 增加了NO₃ ^--N含量, 其影响程度随着增温时间的延长而下降。施氮显著增加了有效氮和微生物生物量氮, 降低了土壤pH值, 使土壤表现出明显的酸化现象。与单独的增温和施氮处理相比, 增温和施氮联合处理对林下土壤的有效氮和微生物特性有显著的交互作用, 显著增加了土壤的有机碳、有效氮及土壤微生物生物量氮含量, 并导致土壤进一步酸化。结果说明, 川西亚高山针叶林的土壤有效氮和微生物特性对土壤氮素状况的变化反应敏感, 而林下土壤有效氮和微生物特性对单独的温度升高表现出一定的适应性, 但更对增温和施氮双因素结合处理反应敏感且表现出不同的响应方式。因此, 该区域在未来全球变化下的氮沉降状况及气候变化的多因素协同效应值得长期深入的探讨。     

Can inconspicuous legumes facilitate alien grass invasions? Partridge peas and fountain grass in Hawai‘i

We investigated the potential role of a nitrogen-fixing legume in facilitating invasion by an alien grass, Pennisetum setaceum (fountain grass) into native Heteropogon contortus grasslands in Hawai'i. Both grasses are C₄ perennials. Many formerly extensive Heteropogon grasslands have been reduced to remnants surrounded by Pennisetum . An inconspicuous annual legume, Chamaecrista nictitans (partridge pea), was commonly associated with Pennisetum stands but was found at higher densities in Pennisetum-Heteropogon mixtures. In the field, we also found higher exchangeable soil nitrogen beneath Chamaecrista-Pennisetum-Heteropogon mixtures than beneath neighboring pure Heteropogon stands. We then used greenhouse experiments to test the hypothesis that increased nitrogen facilitates Pennisetum 's invasion of established Heteropogon . Under low nitrogen conditions, Pennisetum seedlings planted beneath established Heteropogon plants grew very little (<0.3 g biomass after 16 weeks), and their growth was not affected by high or low water treatments. In contrast, under high nitrogen conditions, in both low and high water treatments, Pennisetum seedlings grew rapidly, averaging 60 fold the biomass of seedlings in the low nitrogen treatment and exceeding the biomass of the established Heteropogon plants under which they were planted. We also compared Pennisetum - Heteropogon seedling competition in the presence and absence of live Chamaecrista . The growth of Pennisetum was not directly facilitated by live Chamaecrista in the greenhouse pots, but the Chamaecrista did not increase available soil nitrogen until after it had died. Our results suggest that increases in soil nitrogen, like those associated with an inconspicuous alien legume, can facilitate an alien grass' invasion of a native grassland.     

Effects of soil decomposer invertebrates (protozoa and earthworms) on an above-ground phytophagous insect (cereal aphid) mediated through changes in the host plant

We investigated if the activity of soil invertebrates (protozoa and earthworms) affected the performance of barley and if effects propagated higher up the above-ground food chain into herbivores (cereal aphid, Sitobion avenae ). Barley plants were grown individually in microcosms containing defaunated soil and grass residues. Plants were grown in soil containing: a) no added fauna, b) protozoa, c) earthworms, or d) protozoa and earthworms. After 7 weeks growth at 20°C three adult cereal aphids were added to each plant on separate leaves. The aphids were allowed to grow and reproduce for another 2 weeks before the experiment was destructively sampled. Amounts of mineral N in the soil and leached from the microcosms were significantly reduced by the presence of soil animals. Correspondingly plant biomass and total plant N content were increased significantly by soil animals, protozoa in particular. The different mechanisms responsible for changes in nutrient turnover in presence of protozoa and earthworms are discussed. Aphid performance was strongly influenced by the presence of protozoa, but not by earthworms. In the presence of protozoa the numbers and biomass of adult and juvenile aphids were significantly increased. These effects are likely due to an increased N content in barley plants and consequently increased nitrogen availability to aphids. The results underline that the detritivore and herbivore systems are intimately linked.