Soil ecological interactions: comparisons between tropical and subalpine forests

Soil fauna can influence soil processes through interactions with the microbial community. Due to the complexity of the functional roles of fauna and their effects on microbes, little consensus has been reached on the extent to which soil fauna can regulate microbial activities. We quantified soil microbial biomass and maximum growth rates in control and fauna-excluded treatments in dry and wet tropical forests and north- and south-facing subalpine forests to test whether soil fauna effects on microbes are different in tropical and subalpine forests. Exclusion of fauna was established by physically removing the soil macrofauna and applying naphthalene. The effect of naphthalene application on the biomass of microbes that mineralize salicylate was quantified using the substrate induced growth response method. We found that: (1) the exclusion of soil fauna resulted in a higher total microbial biomass and lower maximum growth rate in the subalpine forests, (2) soil fauna exclusion did not affect the microbial biomass and growth rate in the tropical forests, and (3) the microbial biomass of salicylate mineralizers was significantly enhanced in the fauna-exclusion treatment in the tropical wet and the south-facing subalpine forests. We conclude that non-target effects of naphthalene on the microbial community alone cannot explain the large differences in total microbial biomass found between control and fauna-excluded treatments in the subalpine forests. Soil fauna have relatively larger effects on the microbial activities in the subalpine forests than in tropical dry and wet forests.     

Influence of benthic macrofauna on microbial production of methylmercury in sediments on the New England continental shelf

Microbial methylation processes in sediment are an important source of toxic monomethylmercury (MMHg) to aquatic ecosystems. Although bioturbation activities (feeding, digging of galleries, excavations, bioirrigation) by benthic fauna are known to affect many biogeochemical processes, their influence on benthic MMHg production is poorly understood. We investigated the effect of benthic fauna on the microbial production of MMHg in sediments on the continental shelf of the northwest Atlantic Ocean in September 2009. Replicate cores of sieved (control) and unaltered sediment containing native macrofauna were incubated to examine the influence of benthic macrofauna on net MMHg production, potential gross rates of Hg methylation, sediment reworking, dissolved oxygen and organic carbon concentrations, and microbial metabolic activities. The presence of macrofauna stimulated aerobic microbial respiration and net MMHg production, but had no observed effect on short-term gross rates of Hg methylation. This suggests that bioturbation may promote net MMHg production by inhibiting demethylating microorganisms, although overall community metabolism was increased. Results from this work emphasize the need to enhance our knowledge and understanding of the interactions among benthic fauna, microorganisms, and geochemistry in affecting MMHg production.     

Climate and litter quality differently modulate the effects of soil fauna on litter decomposition across biomes

Climate and litter quality have been identified as major drivers of litter decomposition at large spatial scales. However, the role played by soil fauna remains largely unknown, despite its importance for litter fragmentation and microbial activity. We synthesised litterbag studies to quantify the effect sizes of soil fauna on litter decomposition rates at the global and biome scales, and to assess how climate, litter quality and soil fauna interact to determine such rates. Soil fauna consistently enhanced litter decomposition at both global and biome scales (average increment ~ 27%). However, climate and litter quality differently modulated the effects of soil fauna on decomposition rates between biomes, from climate‐driven biomes to those where climate effects were mediated by changes in litter quality. Our results advocate for the inclusion of biome‐specific soil fauna effects on litter decomposition as a mean to reduce the unexplained variation in large‐scale decomposition models.     

土壤动物与N素循环及对N沉降的响应

以主要的生态过程之一——N循环为对象,论述了土壤动物不仅对凋落物的分解有重要影响,而且在N素矿化和植物对N的吸收过程中也起着重要作用。同时,日益严重的全球变化问题之一——N沉降对土壤动物的多样性及其在生态系统中的功能构成了极大的威胁。另还对土壤动物与N循环研究的方法、土壤动物在N循环过程中的作用机制、热带地区的需求及N沉降下土壤动物的响应作了探讨,并提出,开展大尺度的专类研究及长期定位研究成为下一步研究的需要。     

Exogenous carbon turnover within the soil food web strengthens soil carbon sequestration through microbial necromass accumulation

Exogenous carbon turnover within soil food web is important in determining the trade-offs between soil organic carbon (SOC) storage and carbon emission. However, it remains largely unknown how soil food web influences carbon sequestration through mediating the dual roles of microbes as decomposers and contributors, hindering our ability to develop policies for soil carbon management. Here, we conducted a ¹³C-labeled straw experiment to demonstrate how soil food web regulated the residing microbes to influence the soil carbon transformation and stabilization process after 11 years of no-tillage. Our work demonstrated that soil fauna, as a “temporary storage container,” indirectly influenced the SOC transformation processes and mediated the SOC sequestration through feeding on soil microbes. Soil biota communities acted as both drivers of and contributors to SOC cycling, with 32.0% of exogenous carbon being stabilizing in the form of microbial necromass as “new” carbon. Additionally, the proportion of mineral-associated organic carbon and particulate organic carbon showed that the “renewal effect” driven by the soil food web promoted the SOC to be more stable. Our study clearly illustrated that soil food web regulated the turnover of exogenous carbon inputs by and mediated soil carbon sequestration through microbial necromass accumulation.     

半干旱草原大中型土壤动物在畜粪分解中的作用

土壤动物是陆地生态系统的重要组分, 在有机质分解过程中具有重要作用。目前有关土壤动物在生态系统分解中的作用研究主要聚焦于植物凋落物的分解, 而对动物粪便分解的研究稀少。本研究在内蒙古典型草原设置了马粪和牛粪分解原位实验, 使用不同孔径的金属隔离网排除不同体型大小的土壤动物, 通过测定大中型土壤动物对畜粪分解过程中质量损失、碳氮含量和微生物呼吸以及土壤养分动态变化的影响, 解析其在分解中的作用。设置5个处理, 即CK, 仅土壤, 无粪; T0, 粪添加+0.425 mm隔离网(排除了粪居型和掘洞型粪金龟和中型土壤动物); T1, 粪添加+1 mm隔离网(排除了粪居型和掘洞型粪金龟); T2, 粪添加+2 mm隔离网(排除了掘洞型粪金龟); T3, 仅粪添加(不排除土壤动物)。结果表明: (1)在畜粪分解60天内, 土壤动物对畜粪的干质量损失没有显著的促进作用(P > 0.05); 相反, 在畜粪分解360天, 不隔离土壤动物处理(T3)显著地提高了牛粪干质量损失(P < 0.05), 而降低了马粪干质量损失(P < 0.05)。(2)在畜粪分解的60天内, 畜粪中碳和氮含量下降速度在有土壤动物存在的情况下(T3)快于隔离土壤动物(T0和T1)。(3)两种畜粪添加增加了土壤微生物的呼吸, 且这种增加趋势在实验的第15天和第30天在土壤动物存在时(T3)最明显。(4)与对照(CK)相比, 马粪添加处理提高了土壤速效氮、有机碳的含量和土壤含水量, 且这种增加趋势在排除掘洞型粪金龟(T2)和不排除土壤动物(T3)条件下表现更显著(P < 0.05), 而牛粪添加处理没有明显改变这些指标(P > 0.05)。研究表明, 分解初期粪金龟的取食和活动会改变畜粪的理化性质, 进而影响分解后期土壤生物在畜粪分解中的作用。     

Invertebrate communities and turnover in wetland ponds affected by drought

1. Changes to the macroinvertebrate fauna found in small ponds on a freshwater marsh (Aberlady Bay, Scotland) in 1986, 1987 and 1992 were used to assess the impact of the 1988–92 drought on taxon turnover in pond communities. 2. Permanent ponds accumulated taxa over the study period. Ponds that were wet throughout 1986–87 but dried in 1992 lost some of the many taxa associated with permanent water but acquired a reduced fauna typical of temporary ponds. Ponds that were temporary in 1986–87 were dry during most of 1992 and lost almost all aquatic taxa. 3. Faunal turnover was considerable even in permanent ponds. Extinction rates were high for taxa typical of permanent or temporary ponds. Colonization rates were poor for the taxa from permanent water, but high for the taxa from temporary ponds. 4. The importance of colonization and extinction rates as main predictors of the distribution of species between the ponds was looked at using metapopulation incidence functions, where observed incidence of a taxon can be used to predict colonization and extinction rates. Predicted rates were then compared with observed rates. Incidence functions gave reasonable predictions of observed colonization rates but were poor predictors of extinction, even for taxa that appeared likely to be true metapopulations. 5. For the pond fauna, including fugitive species adapted to temporary ponds, whilst colonization may well depend on environmental stochasricity (how long a pond holds water), subsequent survival depends on other, demographic, processes (e.g. finding a mate, predation) rather than the pond drying out.     

Glycine mineralization in situ closely correlates with soil carbon availability across six North American forest ecosystems

Free amino acids (FAA) constitute a significant fraction of dissolved organic nitrogen (N) in forest soils and play an important role in the N cycle of these ecosystems. However, comparatively little attention has been given to their role as labile carbon (C) substrates that might influence the metabolic status of resident microbial populations. We hypothesized that the residence time of simple C substrates, such as FAA, are mechanistically linked to the turnover of endogenous soil C pools. We tested this hypothesis across a latitudinal gradient of forested ecosystems that differ sharply with regard to climate, overstory taxon, and edaphic properties. Using a combined laboratory and field approach, we compared the turnover of isotopically labeled glycine in situ to the turnover of mineralizable soil C (C_min) at each site. The turnover of glycine was rapid (residence times <2 h) regardless of soil type. However, across all ecosystems glycine turnover rates were strongly correlated with indices of soil organic matter quality. For example, C:N ratios for the upper soil horizons explained ~80% of the variability observed in glycine turnover, and there was a strong positive correlation between in situ glycine-C turnover and C_min measured in the laboratory. The turnover of glycine in situ was better explained by changes in soil C availability than cross-ecosystem variation in soil temperature or concentrations of dissolved inorganic N and FAA-N. This suggests the consumption of these low-molecular-weight substrates by soil microorganisms may be governed as much by the overall decomposability of soil C as by N limitation to microbial growth.     

Linking soil food web structure to above‐ and belowground ecosystem processes: a meta‐analysis

Soil fauna can be an important regulator of community parameters and ecosystem processes, but there have been few quantitative syntheses of the role of soil fauna in terrestrial soil communities and ecosystems. Here, we conducted a meta‐analysis to investigate the impacts of invertebrate soil micro‐ and mesofauna (grazers and predators) on plant productivity and microbial biomass. Overall our results indicate that an increase in the biomass of soil fauna increased aboveground plant productivity across ecosystems by 35% and decreased microbial biomass by 8%. In addition, we found no evidence for trophic cascades in terrestrial soil food webs, but the bacterivorous component of soil fauna influenced plant productivity and microbial biomass more than did the fungivorous component. Furthermore, changes in the biomass of soil fauna differentially affected plant productivity among plant functional groups: a higher biomass of soil fauna increased aboveground productivity by 70% in coniferous systems. However, in ecosystems dominated by legumes, a functional group with lower inorganic nitrogen requirements, there was no response of aboveground productivity to increases in the biomass of soil fauna. In sum, the results of this meta‐analysis indicate that soil fauna help to regulate ecosystem production, especially in nutrient‐limited ecosystems.     

Contrasting patterns of soil N-cycling in model ecosystems of Fennoscandian boreal forests

The low plant productivity of boreal forests in general has been attributed to low soil N supply and low temperatures. Exceptionally high productivity occurs in toe-slope positions, and has been ascribed to influx of N from surrounding areas and higher rates of soil N turnover in situ. Despite large apparent natural variations in forest productivity, rates of gross soil N mineralization and gross nitrification have never been compared in Fennoscandian boreal forests of contrasting productivity. We report contrasting patterns of soil N turnover in three model ecosystems, representing the range in soil C-to-N ratios (19–41) in Fennoscandian boreal forests and differences in forest productivity by a factor close to 3. Gross N mineralization was seven times higher when soil, microbial, and plant C-to-N ratios were the lowest compared to the highest. This process, nitrification and potential denitrification correlated with inorganic, total and microbial biomass N, but not microbial C. There was a constant ratio between soil and microbial C-to-N ratio of 3.7±0.2, across wide ratios of soil C-to-N and fungi-to-bacteria. Soil N-cycling should be controlled by the supplies of C and N to the microbes. In accordance with plant allocation theory, we discuss the possibility that the high fungal biomass at high soil C-to-N ratio reflects a particularly high supply of plant photosynthates, substrates of high-quality C, to mycorrhizal fungi. Methods to study soil N turnover and N retention should be developed to take into account the impact of mycorrhizal fungi on soil N-cycling.     

四川盆地亚热带常绿阔叶林不同物候期凋落物分解与土壤动物群落结构的关系

为了解植物生长不同物候时期凋落物分解过程中土壤动物群落结构动态及其与凋落物分解的关系,以四川盆地亚热带常绿阔叶林典型人工林树种马尾松和柳杉,次生林树种香樟和麻栎凋落物为研究对象,采用凋落物分解袋试验研究,凋落物分解过程中土壤动物的群落特征。4种凋落物分解袋共获得土壤动物8047只,其中,柳杉(2341只)>香樟(2105只)>马尾松(2046只)>麻栎(1555只)。其中,秋末落叶期、萌动期和展叶期,马尾松凋落物袋中主要以捕食性土壤动物为优势类群,而后以菌食性土壤动物为主;香樟凋落物袋中除秋末落叶期和叶衰期以菌食性土壤动物为主要优势类群外,其他各时期均以捕食性土壤动物为主要优势类群;柳杉凋落物分解各时期均以菌食性土壤动物为主要优势类群;麻栎凋落物分解在前3个时期以菌食性为主,而后以植食性土壤动物为主要优势类群。相关分析表明,在秋末落叶期和萌动期土壤动物的个体密度主要和氮、磷含量及其格局密切相关,叶衰期主要和难分解组分木质素显著相关。除在秋末落叶期土壤动物对凋落物分解的贡献率与土壤动物的个体密度显著相关外,其余主要物候关键时期均与土壤动物的类群密度及其食性显著相关。     

Elevated temperature increases the accumulation of microbial necromass nitrogen in soil via increasing microbial turnover

Microbial‐derived nitrogen (N) is now recognized as an important source of soil organic N. However, the mechanisms that govern the production of microbial necromass N, its turnover, and stabilization in soil remain poorly understood. To assess the effects of elevated temperature on bacterial and fungal necromass N production, turnover, and stabilization, we incubated ¹⁵N‐labeled bacterial and fungal necromass under optimum moisture conditions at 10°C, 15°C, and 25°C. We developed a new ¹⁵N tracing model to calculate the production and mineralization rates of necromass N. Our results showed that bacterial and fungal necromass N had similar mineralization rates, despite their contrasting chemistry. Most bacterial and fungal necromass ¹⁵N was recovered in the mineral‐associated organic matter fraction through microbial anabolism, suggesting that mineral association plays an important role in stabilizing necromass N in soil, independently of necromass chemistry. Elevated temperature significantly increased the accumulation of necromass N in soil, due to the relatively higher microbial turnover and production of necromass N with increasing temperature than the increases in microbial necromass N mineralization. In conclusion, we found elevated temperature may increase the contribution of microbial necromass N to mineral‐stabilized soil organic N.     

土壤动物多样性及其生态功能

土壤无脊椎动物生物量通常小于土壤生物总生物量的10%,但它们种类丰富,取食行为及生活史策略多种多样,且土壤动物之间,土壤动物与微生物之间存在着复杂的相互作用关系。土壤动物的生态功能主要通过取食作用(trophic effect)和非取食作用(non-trophic effect)来实现。原生动物数量大、周转快,故原生动物本身的代谢活动(即取食作用)对碳氮矿化的贡献可以接近甚至超过细菌的贡献;然而大多数中小型土壤动物的本身代谢过程对碳氮矿化的贡献远低于土壤微生物,但它们可以通过取食作用来调节微生物进而影响碳氮的矿化。大型节肢动物中的蜘蛛和地表甲虫等捕食者经常活跃于地表,它们常常会通过级联效应对土壤生态系统产生重要的影响。蚯蚓、白蚁等大型土壤动物除可以通过取食作用以外,还可以通过非取食作用调控土壤微生物,进而显著影响土壤碳氮过程。土壤动物取食行为的多样性和复杂的非营养关系的存在造就了多维度的土壤食物网,给土壤动物的生态功能研究带来了巨大的挑战。介绍了土壤动物的多样性及主要的生态功能,并对研究的热点和前沿问题进行了探讨,以期引起关于土壤动物多样性及其生态功能的深入思考。     

Contrasting above‐ and belowground organic matter decomposition and carbon and nitrogen dynamics in response to warming in High Arctic tundra

Tundra regions are projected to warm rapidly during the coming decades. The tundra biome holds the largest terrestrial carbon pool, largely contained in frozen permafrost soils. With warming, these permafrost soils may thaw and become available for microbial decomposition, potentially providing a positive feedback to global warming. Warming may directly stimulate microbial metabolism but may also indirectly stimulate organic matter turnover through increased plant productivity by soil priming from root exudates and accelerated litter turnover rates. Here, we assess the impacts of experimental warming on turnover rates of leaf litter, active layer soil and thawed permafrost sediment in two high‐arctic tundra heath sites in NE‐Greenland, either dominated by evergreen or deciduous shrubs. We incubated shrub leaf litter on the surface of control and warmed plots for 1 and 2 years. Active layer soil was collected from the plots to assess the effects of 8 years of field warming on soil carbon stocks. Finally, we incubated open cores filled with newly thawed permafrost soil for 2 years in the active layer of the same plots. After field incubation, we measured basal respiration rates of recovered thawed permafrost cores in the lab. Warming significantly reduced litter mass loss by 26% after 1 year incubation, but differences in litter mass loss among treatments disappeared after 2 years incubation. Warming also reduced litter nitrogen mineralization and decreased the litter carbon to nitrogen ratio. Active layer soil carbon stocks were reduced 15% by warming, while soil dissolved nitrogen was reduced by half in warmed plots. Warming had a positive legacy effect on carbon turnover rates in thawed permafrost cores, with 10% higher respiration rates measured in cores from warmed plots. These results demonstrate that warming may have contrasting effects on above‐ and belowground tundra carbon turnover, possibly governed by microbial resource availability.     

颗粒有机质的来源、测定及其影响因素

土壤活性有机质及其组分作为土壤质量的重要指标在土壤化学、物理和生物性质方面起着重要作用。颗粒有机质能够有效地反映有机质的特性,与微生物生长、营养供给及C、N的生物学调节密切相关。作为活性有机质的一个量度指标,颗粒有机质越来越受到人们的重视。本文综述了土壤颗粒有机质的来源及其在土壤有机质转化过程中的作用,对其测定方法作了系统的描述,阐明了土壤理化性质、农业措施(施肥与耕作)及土地利用类型对土壤颗粒有机质在土壤形成及维持其稳定性方面的影响。     

Empirical evidence that soil carbon formation from plant inputs is positively related to microbial growth

Plant-carbon inputs to soils in the form of dissolved sugars, organic acids and amino acids fuel much of heterotrophic microbial activity belowground. Initial residence times of these compounds in the soil solution are on the order of hours, with microbial uptake a primary removal mechanism. Through microbial biosynthesis, the dissolved compounds become dominant precursors for formation of stable soil organic carbon. How the chemical class (e.g. sugar) of a dissolved compound influences stabilization in field soils is unknown and predictions from our understanding of microbial metabolism, turnover and identity are contradictory. We show that soil carbon formation, from chronic amendments of dissolved compounds to fertilized and unfertilized grasslands, is 2.4-times greater from a sugar than an amino acid. Formation rates are negatively correlated with respiration rates of the compounds, and positively correlated with their recovery in microbial biomass. These relationships suggest that the efficiency of microbial growth on a compound is positively related to formation rates of soil organic carbon. Fertilization does not alter these findings, but together nitrogen and phosphorus additions reduce soil carbon formation. Our results highlight the need to consider both nutrient enrichment and global-change induced shifts in the form of dissolved root inputs to soils to predict future soil carbon stocks and hence phenomena such as climate warming and food security to which these stock sizes are intimately tied.     

川西亚高山森林土壤动物对杨树和箭竹凋落物分解过程中微生物群落的影响

为了解凋落物分解过程中碎屑食物链土壤动物与微生物的相互联系,本研究以川西亚高山森林杨树和箭竹凋落叶为对象,通过原位控制试验,于2016年4月至2018年4月采用磷脂脂肪酸(PLFAs)生物标记法,研究了土壤动物对两种凋落叶分解过程中微生物丰度、群落结构和多样性的影响.结果表明: 土壤动物的参与显著影响两个树种凋落叶分解过程中微生物PLFAs含量,降低了分解前240天的PLFAs含量,增加了分解360~480 d的PLFAs含量;土壤动物的参与降低了杨树分解过程中的真菌/细菌比值,增加了革兰氏阳性菌(G⁺)/革兰氏阴性菌(G^-)比值,对箭竹分解过程中真菌/细菌和G⁺/G^-比值的影响作用相反;两个树种凋落叶的微生物多样性和均匀性在分解的120 d和480 d维持着较高水平,在分解的360 d和720 d急剧降低,土壤动物的参与显著影响杨树凋落叶的微生物多样性和均匀性,但对箭竹影响不显著;土壤动物对凋落叶微生物PLFAs变化的影响随分解持续时间和树种变化存在差异.亚高山森林凋落物分解过程中土壤动物与微生物群落的相互作用随季节和树种变化具有变异性.     

Enhanced root exudation induces microbial feedbacks to N cycling in a pine forest under long-term CO2 fumigation

The degree to which rising atmospheric CO(2) will be offset by carbon (C) sequestration in forests depends in part on the capacity of trees and soil microbes to make physiological adjustments that can alleviate resource limitation. Here, we show for the first time that mature trees exposed to CO(2) enrichment increase the release of soluble C from roots to soil, and that such increases are coupled to the accelerated turnover of nitrogen (N) pools in the rhizosphere. Over the course of 3 years, we measured in situ rates of root exudation from 420 intact loblolly pine (Pinus taeda L.) roots. Trees fumigated with elevated CO(2) (200 p.p.m.v. over background) increased exudation rates (μg C cm(-1) root h(-1) ) by 55% during the primary growing season, leading to a 50% annual increase in dissolved organic inputs to fumigated forest soils. These increases in root-derived C were positively correlated with microbial release of extracellular enzymes involved in breakdown of organic N (R(2) = 0.66; P = 0.006) in the rhizosphere, indicating that exudation stimulated microbial activity and accelerated the rate of soil organic matter (SOM) turnover. In support of this conclusion, trees exposed to both elevated CO(2) and N fertilization did not increase exudation rates and had reduced enzyme activities in the rhizosphere. Collectively, our results provide field-based empirical support suggesting that sustained growth responses of forests to elevated CO(2) in low fertility soils are maintained by enhanced rates of microbial activity and N cycling fuelled by inputs of root-derived C. To the extent that increases in exudation also stimulate SOM decomposition, such changes may prevent soil C accumulation in forest ecosystems.     

Increased microbial growth,biomass, and turnover drive soil organic carbon accumulation at higher plant diversity

Species‐rich plant communities have been shown to be more productive and to exhibit increased long‐term soil organic carbon (SOC) storage. Soil microorganisms are central to the conversion of plant organic matter into SOC, yet the relationship between plant diversity, soil microbial growth, turnover as well as carbon use efficiency (CUE) and SOC accumulation is unknown. As heterotrophic soil microbes are primarily carbon limited, it is important to understand how they respond to increased plant‐derived carbon inputs at higher plant species richness (PSR). We used the long‐term grassland biodiversity experiment in Jena, Germany, to examine how microbial physiology responds to changes in plant diversity and how this affects SOC content. The Jena Experiment considers different numbers of species (1–60), functional groups (1–4) as well as functional identity (small herbs, tall herbs, grasses, and legumes). We found that PSR accelerated microbial growth and turnover and increased microbial biomass and necromass. PSR also accelerated microbial respiration, but this effect was less strong than for microbial growth. In contrast, PSR did not affect microbial CUE or biomass‐specific respiration. Structural equation models revealed that PSR had direct positive effects on root biomass, and thereby on microbial growth and microbial biomass carbon. Finally, PSR increased SOC content via its positive influence on microbial biomass carbon. We suggest that PSR favors faster rates of microbial growth and turnover, likely due to greater plant productivity, resulting in higher amounts of microbial biomass and necromass that translate into the observed increase in SOC. We thus identify the microbial mechanism linking species‐rich plant communities to a carbon cycle process of importance to Earth's climate system.