Biological effects of water-soluble soil phenol and soil humic extracts on plant systems

The aim of this research was to extract, with a sequential method, HS free from water-soluble phenols (HS-WP) and water-soluble phenols free from humic substances (WP-HS), and to evaluate and to compare the biological effects of these two extracts to humic substances (HS) and water-soluble phenols (WP) traditionally extracted. In each extract we determined the concentration of low molecular weight organic acids, soluble carbohydrates, fatty acids, phenolic acids and total proteins. We tested the biological activity of each soil extract and of the single identified compounds on different plant organs (Pinus laricio callus, Daucus carota cells, and Pinus laricio, Pinus halepensis, Lens culinaris and Cichorium intybus seeds). The results showed that eliminating from HS the WP fraction, and from WP the HS fraction we obtained extracts chemically different from HS as such, and WP as such. HS and HS-WP increased callus and cell growth and also root elongation of the different species used; in contrast, WP and WP-HS had negative effects inhibiting callus and cell growth and seed germination percentage of coniferous, herbaceous and leguminous species. The negative effects can be ascribed to the presence of phenolic acids in the extracts while the positive biological activity can be attributed to the presence of tartaric acid, and fatty acids. In conclusion, this study helps to discriminate the effects of humic substances against phenolic compounds extracted from soils explaining the different and in some way contradictory biological behaviour of these two main fractions of SOM.     

Growth and elemental content of several sagebrush-steppe species in unburned and post-wildfire soil and plant effects on soil attributes

Fire is the principal means of stand renewal in big sagebrush-steppe communities of western North America. Plant growth following fire may be influenced by heat-induced changes in the nutrient status of the soil. Moreover, post-wildfire pioneer plant species may alter soil properties, and thereby, impact subsequent plant recruitment. Our study compared the growth and elemental content of big sagebrush (Artemisia tridentata), squirreltail (Elymus elymoides), cheatgrass (Bromus tectorum), and Indian ricegrass (Achnatherum hymenoides), grown under greenhouse conditions in post-wildfire and similar unburned soil. We also examined soil attributes following plant growth. Cheatgrass and squirreltail, grown in post-wildfire soil, had significantly (p≤0.05) greater aboveground mass than plants grown in unburned soil. As compared with unburned soil, post-wildfire soil engendered the following significant (p≤0.05) differences in leaf elemental content: 1) big sagebrush had higher levels of P and lower levels of Mn; 2) squirreltail accumulated more P and N; and 3) all grass species had higher SiO₂ content. Following harvest of plants, post-wildfire soil generally contained significantly (p≤0.05) more KCl-extractable ortho-P, NH _inf4 ⁺ , and SO ₄ ⁻ , than unburned soil. Plant growth in both burned and unburned soils fostered a significant (p≤0.05) increase in the bicarbonate-extractable pool of P as compared with unplanted controls. Soil Kjeldahl-N was significantly (p≤0.05) greater after plant growth in burned treatments as compared with the control. This study demonstrates that post-wildfire soil can have a stimulatory effect on plant growth for some species. Squirreltail deserves consideration as a post-wildfire revegetation species. Furthermore, pioneer plant growth following wildfires can attenuate soil properties and therefore influence plant succession.     

Competition alters plant species response to nickel and zinc

Phytoextraction can be a cost-efficient method for the remediation of contaminated soils. Using species mixtures instead of monocultures might improve this procedure. In a species mixture, an effect of heavy metals on the species' performance can be modified by the presence of a co-occuring species. We hypothesised that (a) a co-occuring species can change the effect of heavy metals on a target species, and (b) heavy metal application may modifiy the competitive behaviour between the plants. We investigated these mechanisms in a greenhouse experiment using three species to serve as a model system (Carex flava, Centaurea angustifolia and Salix caprea). The species were established in pots of monocultures and mixtures, which were exposed to increasing concentrations of Ni and Zn, ranging from 0 to 2,500 mg/kg. Increased heavy metal application reduced the species' relative growth rate (RGR); the RGR reduction being generally correlated with Ni and Zn concentrations in plant tissue. S. caprea was an exception in that it showed considerable Zn uptake but only moderate growth reduction. In two out of six cases, competitors significantly modified the influence of heavy metals on a target species. The interaction can be explained by an increased uptake of Zn by one species (in this case S. caprea) that reduced the negative heavy metal effect on a target species (C. flava). In two further cases, increasing heavy metal application also altered competitive effects between the species. The mechanisms demonstrated in this experiment could be of relevance for the phytoextraction of heavy metals. The total uptake of metals might be maximised in specific mixtures, making phytoextraction more efficient.     

Resource heterogeneity, soil fertility, and species diversity: effects of clonal species on plant communities

Spatial heterogeneity in soil resources is widely thought to promote plant species coexistence, and this mechanism figures prominently in resource-ratio models of competition. However, most experimental studies have found that nutrient enhancements depress diversity regardless of whether nutrients are uniformly or heterogeneously applied. This mismatch between theory and empirical pattern is potentially due to an interaction between plant size and the scale of resource heterogeneity. Clonal plants that spread vegetatively via rhizomes or stolons can grow large and may integrate across resource patches, thus reducing the positive effect of small-scale resource heterogeneity on plant species richness. Many rhizomatous clonal species respond strongly to increased soil fertility, and they have been hypothesized to drive the descending arm of the hump-shaped productivity-diversity relationship in grasslands. We tested whether clonals reduce species richness in a grassland community by manipulating nutrient heterogeneity, soil fertility, and the presence of rhizomatous clonal species in a 6-year field experiment. We found strong and consistent negative effects of clonals on species richness. These effects were greatest at high fertility and when soil resources were applied at a scale at which rhizomatous clonals could integrate across resource patches. Thus, we find support for the hypothesis that plant size and resource heterogeneity interact to determine species diversity.     

Techniques for boron determination and their application to the analysis of plant and soil samples

This paper reviews techniques for determining B concentration and isotopic ratio and their application to soil and plant samples. Boron concentration has been determined utilising spectrophotometry, potentiometry, chromatography, flame atomic emission and absorption spectrometry, inductively coupled plasma (ICP) optical emission (OES) and mass spectrometry (MS), and neutron activation analysis using neutron radiography and prompt- activation analysis. Isotopic ratios of B have been measured by ICP–MS, thermal ionisation mass spectrometry (TIMS) and secondary ion mass spectrometry (SIMS). For isotopic measurements, TIMS and SIMS are more sensitive and provide higher degrees of accuracy and resolution than ICP–MS, however, extensive sample preparation and purification, and time-consuming measurements limit their usefulness for routine analyses.While the spectrophotometric technique using a colorimetric reaction of B with azomethine-H has been the most extensively applied B determination method for soil and plant samples, colorimetric methods, in general, suffer from numerous interferences and have poor sensitivity and precision. The prompt- method can determine B concentration in intact samples which enables this method to be especially useful for some applications in agriculture. Research involving B behaviour in plant and soil environments would benefit from this technology. In recent years, the use of ICP–OES and ICP–MS for B determination in plant and soil samples has grown tremendously. The application of ICP–OES brought a significant improvement in B analysis because of its simplicity, sensitivity and multielement detection capability. However, besides matrix interferences, the two most sensitive emission lines for B suffer strong spectral interference from Fe. The ICP–OES is not adequately sensitive for some nutritional work involving low B concentrations and B translocation studies using the isotope tracer technique.Plasma is one of the most effective analyte ionisers and MS is the most sensitive ion detector. Coupling of plasma with MS resulted in the development of plasma source MS technology (ICP–MS) which has outperformed all previous analytical methods for trace element determination. Boron determination by ICP–MS suffers no spectroscopic interferences, and is considered the most practical and convenient technique for B isotope determination. The ability of ICP–MS to measure isotopic ratios as well as B concentration enables: (1) B concentration determination by the isotope dilution method, (2) verification of B concentration by isotope fingerprinting in routine analysis and (3) determination of total B concentration as well as B isotope ratio in the same run for biological tracer studies. Therefore, ICP–MS is the method of choice among the present-day technologies for determining B concentration and a convenient method for B isotope determination. In recent years, new generations of plasma-source MS instruments have been developed using alternative plasma generation methods and high-resolution mass spectrometers. These instruments are expected to bring further improvements in accuracy, sensitivity and precision of B determination.     

Evaluation of three herbaceous index plant species for bioavailability of soil cadmium, chromium, nickel and vanadium

Uncultivated plants growing on disturbed sites may be useful for assessing the bioavailability of some metals in soils, and thus the potential for metal mobilization up the terrestrial food chain, an important element in ecological risk assessment. A planted chicory cultivar (Cichorium intybus L. var. foliosum Hegi.) and the uncultivated plants horseweed (Canada fleabane) (Erigeron canadensis L.) and dogfennel (Eupatorium capillifolium (Lam.) Small) were evaluated for their ability to act as index plant species for soil Cd, Cr, Ni, and V at two field sites where these metals had been applied five yr previously to two highly weathered sandy Ultisols. Soil Cd was available to all analyzed plant tissues of all three plant species at both sites, particularly on the sandier Blanton soil. Chicory was an effective index plant for Cd on the finer textured Orangeburg soil but functioned as an indicator plant (toxicity symptoms were observed) on the sandier Blanton soil. Horseweed and dogfennel were effective index plants for Cd in both contaminated soils. Soil Cr, Ni, and V were less bioavailable than soil Cd and plant metal uptake was more sensitive to residual soil Cr, Ni, and V than was soil extraction with double acid. Horseweed and chicory may have potential as index plants for soil Cr. Chicory may have potential as a Ni index plant. Chicory and dogfennel may have potential as V index plants.     

Reciprocal effects of litter from exotic and congeneric native plant species via soil nutrients

Invasive exotic plant species are often expected to benefit exclusively from legacy effects of their litter inputs on soil processes and nutrient availability. However, there are relatively few experimental tests determining how litter of exotic plants affects their own growth conditions compared to congeneric native plant species. Here, we test how the legacy of litter from three exotic plant species affects their own performance in comparison to their congeneric natives that co-occur in the invaded habitat. We also analyzed litter effects on soil processes. In all three comparisons, soil with litter from exotic plant species had the highest respiration rates. In two out of the three exotic-native species comparisons, soil with litter from exotic plant species had higher inorganic nitrogen concentrations than their native congener, which was likely due to higher initial litter quality of the exotics. When litter from an exotic plant species had a positive effect on itself, it also had a positive effect on its native congener. We conclude that exotic plant species develop a legacy effect in soil from the invaded range through their litter inputs. This litter legacy effect results in altered soil processes that can promote both the exotic plant species and their native congener.     

Root, mycorrhiza and earthworm interactions: their effects on soil structuring processes, plant and soil nutrient concentration and plant biomass

Earthworms, arbuscular mycorrhiza fungi (AMF) and roots are important components of the belowground part of terrestrial ecosystem. However, their interacting effects on soil properties and plant growth are still poorly understood. A compartmental experimental design was used in a climate chamber in order to investigate, without phosphorus (P) addition, the single and combined effects of earthworms (Allolobophora chlorotica), AMF (Glomus intraradices) and roots (Allium porrum) on soil structure, nutrient concentration and plant growth. In our experimental conditions, plant roots improved soil structure stability (at the level of macroaggregates) whereas earthworms decreased it. AMF had no effect on soil structure stability but increased P transfer from the soil to the plant and significantly increased plant biomass. Earthworms had no direct influence on P uptake or plant biomass, and the N/P ratio measured in the shoots indicated that P was limiting. Interactions between AMF and earthworms were also observed on total C and N content in the soil and on total root biomass. Their effects varied temporally and between the different soil compartments (bulk soil, rhizosphere and drilosphere). After comparison with other similar studies, we suggest that effects of earthworms and AMF on plant production may depend on the limiting factors in the soil, mainly N or P. Our experiment highlights the importance of measuring physical and chemical soil parameters when studying soil organism interactions and their influence on plant performance.     

The plant beneficial effects of Metarhizium species correlate with their association with roots

Metarhizium species have recently been found to be plant rhizosphere associates as well as insect pathogens. Because of their abundance, rhizospheric Metarhizium could have enormous environmental impact, with co-evolutionary implications. Here, we tested the hypothesis that some Metarhizium spp. are multifactorial plant growth promoters. In two consecutive years, corn seeds were treated with entomopathogenic Metarhizium spp. and field tested at the Beltsville Facility in Maryland. Seed treatments included application of green fluorescent protein (GFP)-tagged strains of Metarhizium brunneum, Metarhizium anisopliae, Metarhizium robertsii, and M. robertsii gene disruption mutants that were either avirulent (Δmcl1), unable to adhere to plant roots (Δmad2), or poorly utilized root exudates (Δmrt). Relative to seeds treated with heat-killed conidia, M. brunneum, M. anisopliae, and M. robertsii significantly increased leaf collar formation (by 15, 14, and 13 %), stalk length (by 16, 10, and 10 %), average ear biomass (by 61, 56, and 36 %), and average stalk and foliage biomass (by 46, 36, and 33 %). Their major impact on corn yield was during early vegetative growth by allowing the plants to establish earlier and thereby potentially outpacing ambient biotic and abiotic stressors. Δmcl1 colonized roots and promoted plant growth to a similar extent as the parent wild type, showing that Metarhizium populations are plant growth promoters irrespective of their role as insect pathogens. In contrast, rhizospheric populations and growth promotion by Δmrt were significantly reduced, and Δmad2 failed to colonize roots or impact plant growth, suggesting that colonization of the root is a prerequisite for most, if not all, of the beneficial effects of Metarhizium.     

Grazing-induced changes in plant species composition affect plant and soil properties of grassland mesocosms

Grazing-induced floristic changes in plant communities may accelerate or reduce plant and soil processes through changes in litter quality. Here, we intended to elucidate if the joint action of live and senescing plant tissue of palatable and non-palatable species differentially influences soil processes and properties. We conducted a 1-year experiment with mesocosms from a subhumid grassland. Mesocosms were monocultures of palatable or non-palatable species and a multispecific control. Palatable species included a legume and annual and perennial grasses, whereas non-palatable species included a perennial grass and annual and perennial forbs. Palatable monocultures showed greater soil mineral nitrogen, soil bacterial diversity, and lower soil pH than non-palatable monocultures. These differences were not accounted for by differences in plant biomass. The multispecific control treatment only exhibited greater shoot biomass than the monocultures, and lower root biomass than the palatable monocultures. Our results suggest that the whole (live + dead plant tissue) had a specific imprint on soil system even when variation was not very apparent in terms of plant biomass, and that this effect was associated with plant palatability to domestic large herbivores.     

Quantifying effects of legal and non-legal designations of alien plant species on their control and profile

Invasive alien species (IAS) pose serious threats to native ecosystems worldwide. In some regions, laws and/or lists related to alien species have been made, but their effects on actual measures against alien species have been little studied. In Japan the IAS act, which came into force in 2005, lists many harmful species under the heading “Designated IAS”. The importation, domestication, sale or release of these species is tightly regulated. In addition, other species are named as alerted alien species (AASs) on the “alien species alert list” to raise public awareness. In this paper we examined the situation in Japan as case study that is exposed to threats from alien species under such laws in recent year to evaluate the effects of that for public interesting. We conducted a survey of the effects of the legal designation and other classification of alien plant species, examining a number of aspects, including research publications, eradications and public awareness. In our analysis, the degree of harmfulness of IASs was considered on the basis of weed risk assessment (WRA). The results show that legal classification of IASs has led to increases in research focused on the designated species, eradication efforts and public awareness. Similar results were also apparent for some AASs, but there was a great deal of variability among the many sub-categories of AASs. This may be partly because AAS sub-categories are based on different premises, and partly because the identification of AASs is not necessarily linked to legal controls that affect the behavior of individuals.     

Interaction of nutrients and four antifungal antibiotics in their effects on Pythium species in vitro and in soil

Summary Griseofulvin, streptomycin, aureomycin, and actidione, with and without nutrients, were tested againstPythium ultimum Trow (two strains),P. irregulare Buis., andP. rostratum Butler. Streptomycin was tolerated in the highest concentration, and actidione and aureomycin in the lowest.P. rostratum, a saprophytic species, was more resistant to the antibiotics than the pathogenic species, which were inhibited at low concentrations. The two strains ofP. ultimum behaved differently, the oogonial strain being more tolerant than the sporangial strain. Yeast extract was usually superior to sucrose or asparagine in counteracting the toxicity of antibiotics.Germination of sporangia ofPythium irregulare was studiedin vitro and in a natural soil in the presence of antibiotic and antibiotic-nutrient solutions. Very meager germination of sporangia occurred in soil amended with antibiotics. Actidione severely restricted germination in soil. Nutrients nullified the toxicity to varying degrees. Usually higher germination of sporangia occurredin vitro than in soil. Sub-lethal concentrations of antibiotics induced morphological abnormalities in germ tubes. Aureomycin caused a dendroid type of branching; actidione caused distortion, vacuolation, and vesiculation; griseofulvin caused lysis and septation; and streptomycin caused distortion.Germ tubes were lysed in a natural soil and in soils amended with antibiotics, but nutrients, in general, prevented early lysis at low concentrations of antibiotics. No resistant bodies were formed prior to the initiation of lysis of germ tubes.     

Interactive effects between plant functional types and soil factors on tundra species diversity and community composition

Plant communities are coupled with abiotic factors, as species diversity and community composition both respond to and influence climate and soil characteristics. Interactions between vegetation and abiotic factors depend on plant functional types (PFT) as different growth forms will have differential responses to and effects on site characteristics. However, despite the importance of different PFT for community assembly and ecosystem functioning, research has mainly focused on vascular plants. Here, we established a set of observational plots in two contrasting habitats in northeastern Siberia in order to assess the relationship between species diversity and community composition with soil variables, as well as the relationship between vegetation cover and species diversity for two PFT (nonvascular and vascular). We found that nonvascular species diversity decreased with soil acidity and moisture and, to a lesser extent, with soil temperature and active layer thickness. In contrast, no such correlation was found for vascular species diversity. Differences in community composition were found mainly along soil acidity and moisture gradients. However, the proportion of variation in composition explained by the measured soil variables was much lower for nonvascular than for vascular species when considering the PFT separately. We also found different relationships between vegetation cover and species diversity according the PFT and habitat. In support of niche differentiation theory, species diversity and community composition were related to edaphic factors. The distinct relationships found for nonvascular and vascular species suggest the importance of considering multiple PFT when assessing species diversity and composition and their interaction with edaphic factors. Synthesis: Identifying vegetation responses to edaphic factors is a first step toward a better understanding of vegetation–soil feedbacks under climate change. Our results suggest that incorporating differential responses of PFT is important for predicting vegetation shifts, primary productivity, and in turn, ecosystem functioning in a changing climate.     

The role of arbuscular mycorrhizal fungi and glomalin in soil aggregation: comparing effects of five plant species

Soil aggregation and soil structure are fundamental properties of natural and managed ecosystems. However, most of our knowledge on the role of plant species in soil aggregation is derived from work in agroecosystems or with agriculturally important plants. Here we examined the effects of five plant species on soil aggregate water stability. The five species (three grasses, one forb, and a legume) were from the same natural grassland, and were grown in monoculture plots in the field. Our first goal was to test if productivity-related or species-specific factors would prevail in determining soil aggregation. We also tested what the relative importance of the soil protein glomalin (produced by arbuscular mycorrhizal fungi, AMF) in soil aggregation is, compared to other factors, including AMF hyphal and root length and percent plant cover. We found significant differences in soil aggregate water stability (1–2 mm size class) for the five plant species examined, and corresponding differences in plant cover, root weight and length, AMF soil hyphal length, and glomalin concentrations. A structural equation modeling approach (path analysis) was used to distinguish direct from indirect effects of factors on soil aggregation based on covariance structures. Root length, soil glomalin, and percent cover contributed equally strong paths to water-stable aggregation. The direct effect of glomalin was much stronger than the direct effect of AMF hyphae themselves, suggesting that this protein is involved in a very important hypha-mediated mechanism of soil aggregate stabilization, at least for the 1–2-mm size class of aggregates.     

Punching above their weight: low-biomass non-native plant species alter soil properties during primary succession

Non‐native invasive plants can greatly alter community and ecosystem properties, but efforts to predict which invasive species have the greatest impacts on these properties have been generally unsuccessful. An hypothesis that has considerable promise for predicting the effects of invasive non‐native plant species is the mass ratio hypothesis (i.e. that dominant species exert the strongest effects). We tested this hypothesis using data from a four year removal experiment in which the presence of two dominant shrub species (one native and the other not), and subordinate plant species, were manipulated in factorial combinations over four years in a primary successional floodplain system. We measured the effects of these manipulations on the plant community, soil nutrient status and soil biota in different trophic levels of the soil food web. Our experiment showed that after four years, low‐biomass non‐native plant species exerted disproportionate belowground effects relative to their contribution to total biomass in the plant community, most notably by increasing soil C, soil microbial biomass, altering soil microbial community structure and increasing the abundance of microbial‐feeding and predatory nematodes. Low‐biomass, non‐native plant species had distinct life history strategies and foliar traits (higher foliar N concentrations and higher leaf area per unit mass) compared with the two dominant shrub species (97% of total plant mass). Our results have several implications for understanding species’ effects in communities and on soil properties. First, high‐biomass species do not necessarily exert the largest impacts on community or soil properties. Second, low‐biomass, inconspicuous non‐native species can influence community composition and have important trophic consequences belowground through effects on soil nutrient status or resource availability to soil biota. Our finding that low‐biomass non‐native species influence belowground community structure and soil properties more profoundly than dominant species demonstrates that the mass ratio hypothesis does not accurately predict the relative effects of different coexisting species on community‐ and ecosystem‐level properties.     

镍污染对土壤微生物的生态效应

镍是高等植物和某些微生物必需的微量营养元素之一,在它们的生命活动中起着重要作用;但浓度较高时,也是一种极毒元素。大量的研究表明,镍污染土壤中微生物的生长、代谢、群落结构和种群多样性会受到不同程度的影响;微生物在长期受重金属威迫的环境中形成其适应性。利用微生物形成的这种适应机制,采用微生物技术治理重金属污染的土壤是可能的。本文还对镍污染土壤的微生物评价指标体系、土壤环境容量、微生物技术开发和综合治理技术开发等的进一步研究作了展望。     

森林植物多样性、树种重要值与土壤理化性质对球囊霉素相关土壤蛋白的影响

球囊霉素相关土壤蛋白(glomalin-related soil protein, GRSP)在土壤物理结构调节和土壤碳库稳定性中发挥着重要作用,但植物多样性和优势种如何影响GRSP还缺乏系统性研究。本研究依托东北林业大学哈尔滨实验林场的72块样地, 对1 m深土壤剖面分5层采样, 测定土壤易提取球囊霉素(easily extractable GRSP, EEG)、总提取球囊霉素(total GRSP, TG)及土壤理化性质, 并同时计算植物多样性指数及优势种重要值(importance value, IV), 进一步通过相关分析和冗余排序分析判断影响GRSP的主要因素与贡献。结果表明: (1)在整个土壤剖面上均表现为TG和EEG与土壤有机碳(SOC)正相关, 在部分土层深度与全氮(total nitrogen, TN)和含水量(moisture content, MC)正相关, 而与电导率(electrical conductivity, EC)和pH值负相关。(2)部分土层TG和EEG与黑皮油松(Pinus tabuliformis var. mukdensis)、樟子松(P. sylvestris var. mongolica)、胡桃楸(Juglans mandshurica)、黄檗(Phellodendron amurense)、榆树(Ulmus pumila)优势种重要值显著相关, 表现为黑皮油松重要值越高, 而黄檗、榆树重要值越小, 越有利于EEG的积累, 并且伴随EEG-C/SOC (EEG中C占SOC比例)增加、EEG/TG增大; 群落中胡桃楸、黄檗、榆树更有利于TG积累, 黑皮油松、落叶松(Larix gmelinii)、樟子松不利于TG的积累。(3)植物Simpson指数、Shannon-Wiener指数、物种丰富度与EEG、TG、EEG/TG无显著相关性, 而与EEG-C/SOC、EEG-N/TN (EEG中N占TN的比例)、TG-C/SOC (TG中C占SOC比例)、TG-N/TN (TG中N占TN的比例)显著负相关; 土壤EEG/TG和EEG-N/TN与植物均匀度指数显著正相关, 在1 m土壤不同土层趋势类似。(4)方差分解分析表明: 生物因子对GRSP变化的解释率是20.2%, 土壤理化因子解释率为7.8%, 而生物因子中植物优势种重要值的解释率最大(16.4%), 而植物物种多样性指数解释率仅为0.4%。冗余排序发现常绿针叶树种(黑皮油松和樟子松)越多且阔叶树种越少时, GRSP含量和GRSP对土壤碳氮的贡献越高(P < 0.01), 其机制可能与树种菌根类型有关: 外生菌根树种重要值与TG显著负相关, 丛枝菌根树种重要值与TG显著正相关。本研究解析了植物物种多样性对GRSP含量的重要影响, 并强调未来土壤管理和评估可以通过调整优势物种而不是树种多样性来促进GRSP积累。