Global reductions in seafloor biomass in response to climate change

Seafloor organisms are vital for healthy marine ecosystems, contributing to elemental cycling, benthic remineralization, and ultimately sequestration of carbon. Deep‐sea life is primarily reliant on the export flux of particulate organic carbon from the surface ocean for food, but most ocean biogeochemistry models predict global decreases in export flux resulting from 21st century anthropogenically induced warming. Here we show that decadal‐to‐century scale changes in carbon export associated with climate change lead to an estimated 5.2% decrease in future (2091–2100) global open ocean benthic biomass under RCP8.5 (reduction of 5.2 Mt C) compared with contemporary conditions (2006–2015). Our projections use multi‐model mean export flux estimates from eight fully coupled earth system models, which contributed to the Coupled Model Intercomparison Project Phase 5, that have been forced by high and low representative concentration pathways (RCP8.5 and 4.5, respectively). These export flux estimates are used in conjunction with published empirical relationships to predict changes in benthic biomass. The polar oceans and some upwelling areas may experience increases in benthic biomass, but most other regions show decreases, with up to 38% reductions in parts of the northeast Atlantic. Our analysis projects a future ocean with smaller sized infaunal benthos, potentially reducing energy transfer rates though benthic multicellular food webs. More than 80% of potential deep‐water biodiversity hotspots known around the world, including canyons, seamounts, and cold‐water coral reefs, are projected to experience negative changes in biomass. These major reductions in biomass may lead to widespread change in benthic ecosystems and the functions and services they provide.     

水生细菌生物量的估算方法*

细菌的生物量是海洋微生物研究的关键参数之一。目前,细菌生物量估算方法主要根据其体积的大小进行换算,但换算的方式也不尽一致。对细菌体积和细菌碳含量主要的测定方法、体积与生物量之间换算系数、换算模式进行介绍,评述。认为流式细胞术是测定细菌体积较为合适的方法,X-射线微量分析法是测定单个细菌生物量(碳含量)最合适的方法,异速生长模式是目前生物量换算过程较为常用的模式。     

Fertilization effects on fineroot biomass, rhizosphere microbes and respiratory fluxes in hardwood forest soils

Fertilizer-induced reductions in CO(2) flux from soil ((F)CO(2)) in forests have previously been attributed to decreased carbon allocation to roots, and decreased decomposition as a result of nitrogen suppression of fungal activity. Here, we present evidence that decreased microbial respiration in the rhizosphere may also contribute to (F)CO(2) reductions in fertilized forest soils. Fertilization reduced (F)CO(2) by 16-19% in 65-yr-old plantations of northern red oak (Quercus rubra) and sugar maple (Acer saccharum), and in a natural 85-yr-old yellow birch (Betula allegheniensis) stand. In oak plots, fertilization had no effects on fine root biomass but reduced mycorrhizal colonization by 18% and microbial respiration by 43%. In maple plots, fertilization reduced root biomass, mycorrhizal colonization and microbial respiration by 22, 16 and 46%, respectively. In birch plots, fertilization reduced microbial respiration by 36%, but had variable effects on root biomass and mycorrhizal colonization. In plots of all three species, fertilization effects on microbial respiration were greater in rhizosphere than in bulk soil, possibly as a result of decreased rhizosphere carbon flux from these species in fertile soils. Because rhizosphere processes may influence nutrient availability and carbon storage in forest ecosystems, future research is needed to better quantify rhizo-microbial contributions to (F)CO(2).     

Assimilation of toluene carbon along a bacteria-protist food chain determined by 13C-enrichment of biomarker fatty acids

A food chain consisting of toluene, toluene-degrading Pseudomonas sp. PS+ and a bacterivorous flagellated amoebae Vahlkampfia sp. was established in a batch culture. This culture was amended with [U-13C]toluene and served as a model system to elucidate the flux of carbon in the food chain by quantifying bacterial biovolumes and 13C enrichment of phospholipid fatty acid (PLFA) biomarkers of the bacteria and the heterotrophic protists. Major PLFA detected in the batch co-culture included those derived from Pseudomonas sp. PS+ (16:1omega7c and 18:1omega7c) and Vahlkampfia sp. (20:4omega6c and 20:3omega6c). A numerical model including consumption of toluene by the bacteria and predation of the bacteria by the heterotrophic protists was adjusted to the measured toluene carbon, bacterial carbon and delta13C values of bacterial and protist biomass. Using this model, we estimated that 28+/-7% of the consumed toluene carbon was transformed into bacterial biomass, and 12+/-4% of the predated bacterial carbon was incorporated into heterotrophic protist biomass. Our study showed that the 13C enrichment of PLFA biomarkers coupled to biomass determination via biovolume calculations is a suitable method to trace carbon fluxes in protist-inclusive microbial food chains because it does not require the separation of protist cells from bacterial cells and soil particles.     

Models of the rhizosphere

A mathematical model has been developed which is capable of simulating the population dynamics of microbial biomass surrounding a root which is releasing soluble and insoluble carbon compounds into the soil. The model simulates the interconversions of C between different pools within the soil as well as the diffusion and mass flow of soluble carbon. Two main aspects of carbon release were examined: (i) a strategy where exudate was released uniformly over the root surface was compared to the case where exudation was confined to a small region behind the root tip; (ii) the situation in which all the C released was in soluble form was compared to the case of an approximately equal partition between soluble and insoluble forms. Substantial differences between the different simulations were found. It was shown that the maximum concentration and penetration of soluble exudates differed markedly between different simulations and the implications of this for micronutrient acquisition by phytosiderophores and for colonisation of the rhizosphere by root pathogens were discussed. The different simulations also predicted very different biomass distributions in the rhizosphere in both space and time.     

Metabolic Status of Bacteria and Fungi in the Rhizosphere of Ponderosa Pine Seedlings

We determined the quantity and metabolic status of bacteria and fungi in rhizosphere and nonrhizosphere soil from microcosms containing ponderosa pine seedlings. Rhizosphere soil was sampled adjacent to coarse, fine, or young roots. The biovolume and metabolic status of bacterial and fungal cells was determined microscopically and converted to total and active biomass values. Cells were considered active if they possessed the ability to reduce the artificial electron acceptor 2-(4-iodophenyl)-3-(4-nitrophenyl)-5-phenyltetrazolium chloride (INT) to visible intracellular deposits of INT formazan. A colorimetric assay of INT formazan production was also used to assess dehydrogenase activity. INT-active microorganisms made up 44 to 55% of the microbial biomass in the soils studied. The proportion of fungal biomass that exhibited INT-reducing activity (40 to 50%) was higher than previous estimates of the active proportion of soil fungi determined by using fluorescein diacetate. Comparison between soils from different root zones revealed that the highest total and INT-active fungal biomass was adjacent to fine mycorrhizal roots, whereas the highest total and active bacterial biomass was adjacent to the young growing root tips. These observations suggest that fungi are enhanced adjacent to the fine roots compared with the nonrhizosphere soil, whereas bacteria are more responsive than fungi to labile carbon inputs in the young root zone. Colorimetric dehydrogenase assays detected gross differences between bulk and rhizosphere soil activity but were unable to detect more subtle differences due to root types. Determination of total and INT-active biomass has increased our understanding of the role of spatial compartmentalization of bacteria and fungi in rhizosphere carbon flow.     

Species-specific effects of polyploidisation and plant traits of Centaurea maculosa and Senecio inaequidens on rhizosphere microorganisms

Invasive plant species represent a threat to terrestrial ecosystems, but their effects on the soil biota and the mechanisms involved are not yet well understood. Many invasive species have undergone polyploidisation, leading to the coexistence of various cytotypes in the native range, whereas, in most cases, only one cytotype is present in the introduced range. Since genetic variation within a species can modify soil rhizosphere communities, we studied the effects of different cytotypes and ranges (native diploid, native tetraploid and introduced tetraploid) of Centaurea maculosa and Senecio inaequidens on microbial biomass carbon, rhizosphere total DNA content and bacterial communities of a standard soil in relation to plant functional traits. There was no overall significant difference in microbial biomass between cytotypes. The variation of rhizosphere total DNA content and bacterial community structure according to cytotype was species specific. The rhizosphere DNA content of S. inaequidens decreased with polyploidisation in the native range but did not vary for C. maculosa. In contrast, the bacterial community structure of C. maculosa was affected by polyploidisation and its diversity increased, whereas there was no significant change for S. inaequidens. Traits of S. inaequidens were correlated to the rhizosphere biota. Bacterial diversity and total DNA content were positively correlated with resource allocation to belowground growth and late flowering, whereas microbial biomass carbon was negatively correlated to investment in reproduction. There were no correlations between traits of the cytotypes of C. maculosa and corresponding rhizosphere soil biota. This study shows that polyploidisation may affect rhizosphere bacterial community composition, but that effects vary among plant species. Such changes may contribute to the success of invasive polyploid genotypes in the introduced range.     

增温对高寒灌丛根际和非根际土壤微生物生物量碳氮的影响

本文对青藏高原东缘窄叶鲜卑花高寒灌丛生长季根际和非根际土壤微生物生物量碳和氮对增温的响应进行研究.结果表明: 窄叶鲜卑花灌丛生长季初期根际和非根际土壤微生物生物量碳和氮均显著高于生长季中期和末期.在多数时期,增温对根际土壤微生物生物量碳和氮的影响不显著.在非根际土壤中,增温对土壤微生物生物量碳和氮的影响因不同生长季节而不同: 增温使生长季初期土壤微生物生物量碳显著降低,而使土壤微生物生物量氮显著提高;生长季中期增温使土壤微生物生物量碳和氮显著提高;而在生长季末期增温对土壤微生物生物量碳和氮的影响不显著.土壤微生物生物量碳和氮的根际效应也因不同生长季节而不同: 土壤微生物生物量碳和氮在生长季初期表现为负根际效应,而在生长季中期表现为正根际效应;在生长季末期,土壤微生物生物量碳表现为负根际效应,土壤微生物生物量氮则表现为正根际效应.增温在生长季初期使土壤微生物生物量碳和氮的根际效应显著提高,而在生长季中期和末期使土壤微生物生物量碳和氮的根际效应降低.本研究初步阐明了气候变暖背景下高寒灌丛根际和非根际土壤生物学过程变化机理.     

CO2浓度升高对森林土壤微生物呼吸与根（际）呼吸的影响

 根呼吸与微生物呼吸的作用底物不同,二者对高浓度CO₂的响应机理及敏感程度亦不同。在大气CO₂浓度升高的背景下,精确区分根呼吸与微生物呼吸是构建森林生态系统碳循环模型和预测森林生态系统碳源/汇关系所必需的。根（际）呼吸与微生物呼吸对高浓度CO₂的响应呈增加、降低或无明显变化等不同趋势,根（际）呼吸变化主要与根生物量明显相关,细根的作用大于粗根;土壤微生物呼吸变化存在较大的不确定性,微生物量和微生物活性与土壤微生物呼吸相关或不相关。根系统对高浓度CO₂的响应会潜在地影响微生物的代谢底物,进而影响微生物呼吸强度。凡影响土壤总呼吸的生物与非生物因子都会直接或间接地影响根呼吸与土壤微生物呼吸。     

基于模型数据融合的长白山阔叶红松林碳循环模拟

 充分、有效地利用各种陆地生态系统碳观测数据改善陆地生态系统模型, 是当前我国陆地生态系统碳循环研究领域亟待解决的重要问题之一。该研究以2003~2005年长白山阔叶红松林的6组生物计量观测数据和涡度相关技术测定的碳通量数据为基础, 利用马尔可夫链-蒙特卡罗方法对陆地生态系统模型的关键参数(即碳滞留时间)进行了反演, 进而预测了长白山阔叶红松林生态系统碳库、碳通量及其不确定性。反演结果表明, 长白山阔叶红松林叶凋落物和微生物碳的平均滞留时间最短, 为2~6个月; 其次是叶和细根生物量碳, 二者的平均滞留时间为1~2 a; 慢性土壤有机碳的平均滞留时间为8~16 a; 碳在木质生物量和惰性土壤有机质库中的滞留时间最长, 平均滞留时间分别为77~109 a和409~1 879 a。模拟结果显示, 碳库和累积碳通量模拟值的不确定性将随着模拟时间的延长而增大。当气温升高10%和20%时, 长白山阔叶红松林总初级生产力年总量将分别增加6.5%和9.9%, 净生态系统生产力(NEP)年总量的变化取决于土壤温度的变化。若土壤温度保持不变, NEP年总量将分别增加11.4%~21.9%和17.6%~33.1%; 若土壤温度也相应升高10%和20%, NEP年总量的增幅反而下降甚至低于原来的水平。假设气候和植被保持在2003~2005年的状态, 2020年长白山阔叶红松林NEP年总量为(163±12) g C·m^–2·a^–1, 土壤呼吸年总量为(721±14) g C·m^–2·a^–1。马尔可夫链-蒙特卡罗方法是反演模型参数、优化模拟结果和评估模拟结果不确定性的有效方法, 但今后仍需在惰性土壤碳滞留时间的估计、驱动数据和模型结构的不确定性分析、模型数据融合方法方面进行深入研究, 以进一步提高碳循环模拟的准确性。     

Low amounts of herbivory by root-knot nematodes affect microbial community dynamics and carbon allocation in the rhizosphere

Increased carbon translocation to the rhizosphere via 'leakage' induced by low amounts of plant parasitic nematodes can foster microorganisms. The effects of the root-knot nematode Meloidogyne incognita on microbial biomass (C(mic)) and community structure (phospholipid fatty acids) in the rhizosphere of barley were studied. Inoculation densities of 2000, 4000, and 8000 nematodes were well below the threshold level for plant damage. A (13)CO(2) pulse-labelling was performed to assess the distribution of assimilated (13)C in the rhizosphere. Infection with M. incognita increased the carbon concentration in shoots, and enhanced root biomass slightly. The presence of nematodes did not affect microbial biomass, but significantly changed the allocation of the recent photosynthate. Less plant carbon was sequestered by microorganisms with increasing nematode abundance. Microbial community structure was distinctly altered in the early stages of the plant-nematode interactions. Both, bacteria and fungi, showed a positive response with 2000, and a negative one with 4000 and 8000 M. incognita added. The results suggest that low-level root herbivory still imposes a considerable carbon demand, and that proliferation of microorganisms due to increased rhizodeposition may be short-termed. The carbon flow to rhizosphere microbial communities is likely dependent on the specific nematode-plant association and the developmental stage of the nematode in the host.     

Decomposer biomass in the rhizosphere to assess rhizodeposition

Quantification of the organic carbon released from plant roots is a challenge. These compounds of rhizodeposition are quickly transformed into CO₂ and eventually bacterial biomass to be consumed by bacterivores (protozoa and nematodes). Microbes stimulate rhizodeposition several-fold so assays under sterile conditions give an unrealistic value. Quantifying bacterial production from ³H-thymidine incorporation falls short in the rhizosphere and the use of isotopes does not allow clear distinction between labeled CO₂ released from roots or microbes. We reduced rhizodeposition in 3–5 week old barley with a 2 week leaf aphid attack and found that biomass of bacterivores but not bacteria in the rhizosphere correlated with plant–induced respiration activity belowground. This indicated top-down control of the bacteria. Moreover, at increasing density of aphids, bacterivore biomass in the rhizosphere decreased to the level in soil unaffected by roots. This suggests that difference in bacterivore biomass directly reflects variations in rhizodeposition. Rhizodeposition is estimated from plant-induced increases in bacterial and bacterivore biomass, and yield factors, maintenance requirements, and turnover rates from the literature. We use literature values that maximize requirements for organic carbon and still estimate the total organic rhizodeposition to be as little as 4–6% of the plant-induced respiration belowground.     

Estimating diurnal to annual ecosystem parameters by synthesis of a carbon flux model with eddy covariance net ecosystem exchange observations

We performed a synthetic analysis of Harvard Forest net ecosystem exchange of CO₂ (NEE) time series and a simple ecosystem carbon flux model, the simplified Photosynthesis and Evapo‐Transpiration model (SIPNET). SIPNET runs at a half‐daily time step, and has two vegetation carbon pools, a single aggregated soil carbon pool, and a simple soil moisture sub‐model. We used a stochastic Bayesian parameter estimation technique that provided posterior distributions of the model parameters, conditioned on the observed fluxes and the model equations. In this analysis, we estimated the values of all quantities that govern model behavior, including both rate constants and initial conditions for carbon pools. The purpose of this analysis was not to calibrate the model to make predictions about future fluxes but rather to understand how much information about process controls can be derived directly from the NEE observations. A wavelet decomposition enabled us to assess model performance at multiple time scales from diurnal to decadal. The model parameters are most highly constrained by eddy flux data at daily to seasonal time scales, suggesting that this approach is not useful for calculating annual integrals. However, the ability of the model to fit both the diurnal and seasonal variability patterns in the data simultaneously, using the same parameter set, indicates the effectiveness of this parameter estimation method. Our results quantify the extent to which the eddy covariance data contain information about the ecosystem process parameters represented in the model, and suggest several next steps in model development and observations for improved synthesis of models with flux observations.     

Relations Between Bacterial Biomass and Carbon Cycle in Marine Sediments: An Early Diagenetic Model

A new model for early diagenetic processes has been developed through a new formula explicitly accounting for microbial population dynamics. Following a mechanistic approach based on enzymatic reactions, a new model has been proposed for oxic mineralisation and denitrification. It incorporates the dynamics of bacterial metabolism. We find a general formula for inhibition processes of which some other mathematical expressions are particular cases. Moreover a fast numerical algorithm has been developed. It allows us to perform simulations of different diagenetic models in non-steady states. We use this algorithm to compare our model to a classical one (Soetaert et al., 1996). Dynamical evolutions of a perturbation of particulate organic carbon (POC) input are studied for both models. The results are very similar for stationary cases. But with variable inputs, the bacterial biomass dynamics brings about noticeable differences, and these are discussed.     

Bacterial Biovolume and Biomass Estimations

The biomass of bacterial populations in aquatic ecosystems is often estimated by measuring bacterial biovolume and converting this into biomass in terms of carbon. A reliable conversion factor relating the measured bacterial biovolume to bacterial carbon content is essential for this approach. Based on direct measurements of bacterial cell carbon content, cell number, and biovolume, I have derived an average conversion factor of 5.6 × 10⁻¹³ g of C μm⁻³. This conversion factor is 3.4 to 6.6 times higher than most theoretically derived factors currently in use. Both bacterial biomass and bacterial production in aquatic ecosystems may thus have been seriously underestimated.     

The role of carbon flux and biometric observations in constraining a terrestrial ecosystem model: a case study in disturbed forests in East Asia

The process of confining unnecessary freedom is a step toward advanced ecosystem modeling. This study demonstrates the importance of carbon flux and biometric observation in constraining a terrestrial ecosystem model with a simple optimization scheme. At the selected sites from AsiaFlux network, a simultaneous optimization scheme for both carbon flux and biomass was compared with carbon flux-oriented and biomass-oriented optimization schemes using the Biome-BGC model. The optimization scheme oriented to either carbon flux or biomass provided simulation results that were consistent with observations, but with reduced performance in unconstrained variables. The simultaneous optimization scheme yielded results that were consistent with observations for both carbon flux and biomass. By comparing long-term projections simulated by three schemes, it was found that the optimization oriented only to carbon flux has limited performance because misrepresented biomass significantly affected a projection of carbon exchange through heterotrophic respiration. From these experiments, we found that (1) a process model like Biome-BGC is capable of reproducing both carbon flux and biomass within acceptable proximity, (2) constraining biomass is importance not just because it is one of carbon cycle components, but also it significantly affects simulations of carbon flux. Thus, it is important to invest more effort to improve simulation of biomass as well as carbon flux.     

Carbon flow in the rhizosphere: carbon trading at the soil–root interface

The loss of organic and inorganic carbon from roots into soil underpins nearly all the major changes that occur in the rhizosphere. In this review we explore the mechanistic basis of organic carbon and nitrogen flow in the rhizosphere. It is clear that C and N flow in the rhizosphere is extremely complex, being highly plant and environment dependent and varying both spatially and temporally along the root. Consequently, the amount and type of rhizodeposits (e.g. exudates, border cells, mucilage) remains highly context specific. This has severely limited our capacity to quantify and model the amount of rhizodeposition in ecosystem processes such as C sequestration and nutrient acquisition. It is now evident that C and N flow at the soil–root interface is bidirectional with C and N being lost from roots and taken up from the soil simultaneously. Here we present four alternative hypotheses to explain why high and low molecular weight organic compounds are actively cycled in the rhizosphere. These include: (1) indirect, fortuitous root exudate recapture as part of the root’s C and N distribution network, (2) direct re-uptake to enhance the plant’s C efficiency and to reduce rhizosphere microbial growth and pathogen attack, (3) direct uptake to recapture organic nutrients released from soil organic matter, and (4) for inter-root and root–microbial signal exchange. Due to severe flaws in the interpretation of commonly used isotopic labelling techniques, there is still great uncertainty surrounding the importance of these individual fluxes in the rhizosphere. Due to the importance of rhizodeposition in regulating ecosystem functioning, it is critical that future research focuses on resolving the quantitative importance of the different C and N fluxes operating in the rhizosphere and the ways in which these vary spatially and temporally.     

土壤-植物-大气连续体水热、CO2通量估算模型研究进展

土壤-植物-大气连续体(SPAC)水热、CO2通量的准确估算对理解陆地和大气的物质和能量交换过程有着重要意义.重点阐述了基于过程的土壤-植物-大气连续体水热、CO2通量模型,综述了统计模型、综合模型及基于遥感的模型的发展过程.其中水热通量统计模型包括基于温度和湿度以及基于温度和辐射的方法;CO2通量统计模型包括基于气候因子或蒸散因子以及基于光能利用率的方法.水热通量过程模型包括大叶、双源、多源和多层的水热传输物理模型;CO2通量过程模型包括叶片尺度及由大叶、双叶和多层方法扩展到冠层尺度的生理生态模型以及光合-蒸腾耦合模型.综合模型包括生物物理模型、生物化学模型和生物地理模型.统计模型形式简单,资料易得,对大范围的水热通量模拟具有指导意义;过程模型准确的揭示了水热和CO2通量传输的物理和生理过程,是大尺度综合模型的基础.未来生态系统水热、CO2通量估算模型将集成各种技术手段进行多尺度网络观测和大尺度机理模拟.