植物对有机氮源的利用及其在自然生态系统中的意义

近来大量实验研究表明,许多植物能够在不经矿化的情况下直接吸收、利用环境介质中的生物有机氮,尤其氨基酸类。而且,有些植物利用氨基酸的效率可以与矿质氮源(NH4 、NO3)相当或更高。自然界植物赖以生存的土壤生境中同时存在多种有机氮和矿质氮养分,这是导致植物(至少部分植物)进化产生利用各种不同氮源能力的环境驱动力。土壤中的游离氨基酸尽管含量不高,但其周转快、通量大,理论上可远大于植物的氮需求。尽管植物在与土壤微生物的有机氮源竞争中处于根本性劣势,但土壤中氨基酸的巨大潜在通量和植物相对于微生物的生命周期仍可使植物在长期竞争中获取数量可观的氮。基于植物根对氨基酸的吸收能力、土壤中游离氨基酸库的大小和通量、植物与土壤微生物对氨基酸氮源的竞争以及有关的原位实验结果,近来许多研究者都认为植物有机氮营养在多种生态系统中是重要或潜在重要的。尤其是在一些极地、高山、亚高山、北方针叶林或泰加林生态系统中,由于低温等因素限制有机氮矿化,土壤氨基酸浓度常超过矿质氮(NH4 、NO3-)浓度,氨基酸可能代表着植物的一个主要氮源。认识到现实生态系统中植物对有机氮源利用的重要性意味着传统的矿质营养观念的更新,这将在很大程度上改变人们对某些重要生态过程的理解,并导致对若干生态学中心问题的再认识。研究以森林生态系统为例,阐述了我国开展该领域研究的科学意义和基本框架。     

Amino acid uptake: a widespread ability among boreal forest plants

Amino acids constitute a potentially important source of nitrogen for plants in boreal forest ecosystems. Accordingly, it may be suggested that distinct plant species differing abilities to take up amino acids constitutes an important factor in determining plant ecosystem composition. Using GC-MS and isotopically labelled amino acids, we measured the simultaneous uptake of 15 different amino acids by 31 common boreal forest plant species. The results from this study show that all plant species tested, representing a wide variety of plant types, have the ability to take up amino acids from an incubation solution. Furthermore, uptake rates were unrelated to mycorrhizal associations as well as habitat soil amino acid concentrations and plant nitrogen availability dependence as measured by Ellenberg nitrogen indicator values. These results suggest that mycorrhiza is of minor importance for discrete plant amino acid uptake rates and further points out the potential importance of amino acids to plant nitrogen nutrition in boreal forest ecosystems.     

Soil amino-acid availability across a temperate-forest fertility gradient

Despite increasing recognition that free amino acids can be an important source of N for plant uptake, we have a poor understanding of environmental variation in the availability of amino-acid N in soils outside of arctic, alpine and boreal regions. I investigated patterns of amino-acid availability along a temperate forest fertility gradient ranging from low mineral N availability, oak-dominated forests to high mineral N availability, maple-basswood forests (5 sites). I measured standing pools of free amino acids, soluble peptides, ammonium and nitrate, rates of amino acid production (native proteolysis activity) and rates of consumption of a ¹⁵N-labeled leucine tracer. Standing pools of amino acid N decreased consistently along the fertility gradient from the low fertility black oak/white oak system to the high fertility sugar maple/basswood system, with a 25-fold difference in pool sizes between the poorest and richest sites. Standing pools of soluble peptides varied little among sites, instead, the relationship between free amino acids and peptides changed markedly across the gradient. At low fertility sites free amino acids were positively correlated with soluble peptides, whereas free amino acid pools were universally low at high fertility sites, regardless of peptide pools. Assays for native proteolysis activity indicated that amino acid production did not vary significantly among sites. Recovery of leucine tracer in inorganic (NH₄ ⁺ and NO₃ ⁻) pools and in residual soil organic matter both increased with increasing soil fertility; however, total consumption of the added amino-acid tracer did not vary among sites. Results from this study demonstrate that free amino acids can make an important contribution to potentially plant-available N pools in temperate forest soils, particularly at low fertility sites.     

Ungulate and topographic control of nitrogen: phosphorus stoichiometry in a temperate grassland; soils, plants and mineralization rates

Although the link between the nitrogen (N): phosphorus (P) stoichiometry of biota and availability has received considerable attention in aquatic systems, there has been relatively little effort to compare the elemental composition of biota and supply in terrestrial habitats. In this study, I explored the effects of a prominent topo-edaphic gradient, from dry hilltop to wet slope-base, and native ungulates on N and P of soils, plants, and rates of in situ net mineralization in grasslands of Yellowstone National Park. Nitrogen and P measurements were made May–September, 2000, in paired, grazed and 38–42 year fenced, ungrazed grassland at five topographically variable sites. Similar to findings from other grassland ecosystems, several site factors associated with organic activity, including soil moisture, C, and plant biomass, covaried with soil N concentration and/or net N mineralization. Soil P concentration and net P mineralization, however, were unrelated to those factors. Instead, net P mineralization was negatively related to soil pH, which is known to control the form of inorganic P and its availability, and soil P was uncorrelated with any soil or plant variable measured in the study. Because of being influenced by different soil properties, N and P net mineralization were unrelated among grasslands. Furthermore, supply and plant N:P ratios were uncorrelated in this grassland system. Based on critical N:P ratios reflecting nutritional limitation of plants, Yellowstone grassland vegetation ranged from being N limited to N-P co-limited. Grazers increased N-P co-limitation by enhancing plant N concentrations and the soil pH gradient across grassland sites regulated plant nutritional limitation by affecting plant-available P. These findings showed how ungulates and a landscape factor, i.e. soil pH, determined plant nutrient status among YNP grasslands differently by influencing plant N concentration versus plant P concentration, respectively.     

高等植物对有机氮吸收与利用研究进展

主要综述植物氨基酸营养生理生化和分子生物学研究的最新进展。长期以来,人们一直认为植物只能吸收无机态N,有机N必须矿化为无机N后才能被植物吸收利用,而近年来越来越多实验证明植物能吸收有机N,特别是氨基酸,其吸收能力因植物种类而异,生长在有机N丰富的北极,高山和亚高山生态环境中的植物甚至嗜好氨基酸,因此,不应过分夸大有机N矿化的重要性,迄今一些植物细胞质膜上的氨基酸转运子基因已被描述并加以克隆。     

喀斯特不同植被恢复阶段土壤有机酸季节变化与有效氮磷的关系

植物分泌有机酸在提高土壤养分有效性方面起到重要作用。为了解喀斯特地区不同植被恢复阶段土壤有机酸含量季节性变化与氮磷有效性的关系,在灌木林和原生林各选择3种优势植物,测定雨季和旱季两个季节根际土和非根际土的有机酸含量、碳氮磷含量和比值、有效性氮磷含量及微生物生物量碳。结果表明:原生林植物根际土的草酸含量高于灌木林,而苹果酸和乙酸含量则低于灌木林;根际土草酸含量均高于非根际土; 2个植被根际土和非根际土的草酸含量在雨季高于旱季,而苹果酸和乙酸含量则低于旱季;土壤草酸含量与有机碳、全氮、全磷和N∶P值呈显著正相关,与C∶N呈显著负相关;土壤有效氮和有效磷与草酸和微生物生物量碳呈显著正相关。上述结果表明,植物分泌有机酸的季节性变化与土壤养分状态和自身养分需求相关,而有机酸耦合微生物对养分有效性的提高具有积极的作用。因此,根际土的有机酸季节变化可能是喀斯特生态系统中植物适应土壤养分限制的一种重要机制。     

氨基酸添加对亚热带森林红壤氮素转化的影响

为探究氨基酸氮形态对亚热带土壤氮素含量及转化的影响,选择建瓯市万木林保护区的山地红壤为对象,采用室内培养实验法,通过设计60%和90%WHC两种土壤含水量并添加不同性质氨基酸,测定了土壤中铵态氮、硝态氮、可溶性有机氮的含量和氧化亚氮的释放量,分析了可溶性有机碳、土壤p H值的大小变化及其与氮素的相互关系。结果表明:与对照处理相比,氨基酸添加显著增加了土壤NH_4~+-N含量并使土壤p H值升高,且在一定程度上解除了高含水量(90%WHC)对NH_4~+-N产生的抑制,其中甲硫氨基酸的效果最为明显。酸性、碱性、中性氨基酸对土壤NO_3~--N含量和N_2O释放影响不显著,但甲硫氨基酸可显著抑制土壤硝化从而导致NH_4~+-N的积累,并在培养前期抑制土壤N_2O产生而在培养后期促进N_2O释放,总体上促进N_2O释放。60%WHC的氨基酸添加处理较90%WHC条件下降低土壤可溶性有机氮的幅度更大。氨基酸对土壤氮素转化的影响与带电性关系较小,而可能与其分解产物密切相关。可见,不同性质氨基酸处理对森林土壤氮素含量及转化存在不同程度的影响,且甲硫氨基酸对土壤氮素转化的影响机理值得深入研究。     

Ectomycorrhizal Fungal Protein Degradation Ability Predicted by Soil Organic Nitrogen Availability

In temperate and boreal forest ecosystems, nitrogen (N) limitation of tree metabolism is alleviated by ectomycorrhizal (ECM) fungi. As forest soils age, the primary source of N in soil switches from inorganic (NH₄⁺ and NO₃⁻) to organic (mostly proteins). It has been hypothesized that ECM fungi adapt to the most common N source in their environment, which implies that fungi growing in older forests would have greater protein degradation abilities. Moreover, recent results for a model ECM fungal species suggest that organic N uptake requires a glucose supply. To test the generality of these hypotheses, we screened 55 strains of 13 Suillus species with different ecological preferences for their in vitro protein degradation abilities. Suillus species preferentially occurring in mature forests, where soil contains more organic matter, had significantly higher protease activity than those from young forests with low-organic-matter soils or species indifferent to forest age. Within species, the protease activities of ecotypes from soils with high or low soil organic N content did not differ significantly, suggesting resource partitioning between mineral and organic soil layers. The secreted protease mixtures were strongly dominated by aspartic peptidases. Glucose addition had variable effects on secreted protease activity; in some species, it triggered activity, but in others, activity was repressed at high concentrations. Collectively, our results indicate that protease activity, a key ectomycorrhizal functional trait, is positively related to environmental N source availability but is also influenced by additional factors, such as carbon availability.     

Fire effects on ecosystem nitrogen cycling in a Californian bishop pine forest

Fire can cause severe nitrogen (N) losses from grassland, chaparral, and temperate and boreal forest ecosystems. Paradoxically, soil ammonium levels are markedly increased by fire, resulting in high rates of primary production in re-establishing plant communities. In a manipulative experiment, we examined the influence of wild-fire ash residues on soil, microbial and plant N pools in a recently burned Californian bishop pine (Pinus muricata D. Don) forest. Ash stimulated post-fire primary production and ecosystem N retention through direct N inputs from ash to soils, as well as indirect ash effects on soil N availability to plants. These results suggest that redistribution of surface ash after fire by wind or water may cause substantial heterogeneity in soil N availability to plants, and could be an important mechanism contributing to vegetation patchiness in fire-prone ecosystems. In addition, we investigated the impact of fire on ecosystem N cycling by comparing ¹⁵N natural abundance values from recently burned and nearby unburned P. muricata forest communities. At the burned site, ¹⁵N natural abundance in recolonising species was similar to that in bulk soil organic matter. By contrast, there was a marked ¹⁵N depletion in the same species relative to the total soil N pool at the unburned site. These results suggest that plant uptake of nitrate (which tends to be strongly depleted in ¹⁵N because of fractionation during nitrification) is low in recently burned forest communities but could be an important component of eco- system N cycling in mature conifer stands. Received: 29 June 1999 / Accepted: 24 October 1999     

落叶松幼苗碳素和氮素的获取与分配对供氮水平的响应

落叶松(Larix gmelinii)是中国东北林区最重要的工业用材树种,而且在北温带森林中具有重要的生态学意义。落叶松的种植区域内气温低、冬季长,氮素矿化速度低,供氮不足常常成为落叶松生长的限制因素。为揭示落叶松生长与氮素营养的关系,采用沙培法设置了1、4、8和16 mmol·L^-1 4个供氮水平,研究了不同供氮条件下落叶松一年生幼苗对碳和氮的获取与分配的规律。结果显示,落叶松幼苗的生物量、全株氮浓度、氮含量、比氮吸收速率均随供氮水平的增加而升高,叶重比(LWR)、茎重比(SWR)及叶氮比(LNR)、茎氮比(SNR)亦随供氮水平的增加而增加,而根重比(RWR)和根氮比(RNR)则随供氮水平的增加而降低。当供氮水平从1 mmol·L^-1增加至8 mmol·L^-1时,落叶松幼苗相对生长速率呈线性增加,而全株氮生产力几乎未受供氮水平的影响;当供氮水平从8 mmol·L^-1增加至16 mmol·L^-1时,全株相对生长速率不再增加,全株氮生产力则显著下降。与全株氮生产力的变化不同,落叶松幼苗的叶氮生产力与供氮水平呈负相关。     

氮沉降对森林植物的影响

综述了氮沉降对森林植物的影响。氮沉降对森林植物的影响主要表现在以下6个方面：(1)在一定量范围内的氮沉降有利于植物的光合作用,但过量后则会引起植物的光合速率下降;(2)当植物生长受氮限制时,在一定程度上的氮沉降增加植物生产力,但当氮过量后,氮沉降则使植物的生产力下降;(3)过量的氮沉降导致植物体各种营养元素含量的比例失衡;(4)氮沉降会改变植物的形态结构,集中表现为根／冠比减小;(5)氮沉降会增加植物对天然胁迫如干旱、病虫害和风的敏感性,减少其抵御能力;(6)氮沉降会改变植物组成和降低森林植物的多样性。     

Soil acidification and adaptations of plants and microorganisms in Bornean tropical forests

In tropical forest ecosystems, a paradoxical relationship is commonly observed between massive biomass production and low soil fertility (low pH). The loss and deficiency of soil phosphorus (P) and bases generally constrain biomass production; however, high productivity on nutrient-deficient soils of Bornean tropical forests is hypothesized to be maintained by plant and microorganism adaptation to an acidic soil environment. Proton budgets in the plant–soil system indicated that plants and microorganisms promote acidification to acquire bases, even in highly acidic tropical soils. The nitric and organic acids they produce contribute to the mobilization of basic cations and their uptake by plants. In response to soil P deficiency and the recalcitrance of lignin-rich organic matter, specific trees and fungi can release organic acids and enzymes for nutrient acquisition. Organic acids exuded by roots and rhizosphere microorganisms can promote the solubilization of P bonded to aluminum and iron oxides and its uptake by plants from P-poor soils. Lignin degradation, a rate-limiting step in organic matter decomposition, is specifically enhanced in acidic organic layers by lignin peroxidase, produced by white-rot fungi, which may solubilize recalcitrant lignin and release soluble aromatic substances into the soil solution. This dissolved organic matter functions in the transport of nitrogen, P, and basic cations in acidic soils without increasing leaching loss. In Bornean tropical forests, soil acidification is promoted by plants and microorganisms as a nutrient acquisition strategy, while plant roots and fungi can develop rhizosphere and enzymatic processes that promote tolerance of low pH.     

Differential uptakes of different forms of soil nitrogen among major tree species in subalpine coniferous forests of western Sichuan,China

Aims Although acquisition of soil organic nitrogen (N)(mainly amino acids) by plants is a widespread ecological phenomenon in many terrestrial ecosystems, the rate of organic N uptake and their contributions to plant nutrient supply are poorly understood. Our objective was to determine the relative contributions of inorganic N (NO₃^–-N and NH₄⁺-N) and organic N (amino acids) to plant N uptake in a high-frigid forest ecosystem.Methods The differences in the uptake rate of three different forms of N (NO₃^–-N, NH₄⁺-N and glycine) were quantified by exposing seedlings of two dominant tree species (Picea asperata and Betula albo-sinensis) in subalpine coniferous forests of western Sichuan, China, to trace quantities of K¹⁵NO₃,¹⁵NH₄Cl and (U-¹³C₂/¹⁵N) glycine.Important findings Both ¹³C and ¹⁵N were significantly enriched in fine roots 2 h after tracer application, indicating the occurrence of glycine uptake in P. asperata and B. albo-sinensis seedlings. The seedlings of two tree species had a significant preference for NO₃^–-N compared with glycine and NH₄⁺-N, and the uptake rate of NO₃^–-N was 5 to 10 times greater than that of glycine and NH₄⁺-N. The roots of seedlings in the two species took up glycine more rapidly than NH₄⁺-N, implying that soil organic N (i.e., amino acids) could be an important N source for the two species in subalpine coniferous forests. The results of this study are of great theoretical significance for understanding N utilization strategies and nutrient regulation processes in plants of the high-frigid forest ecosystems.     

A novel method to measure the effect of temperature on diffusion of plant-available nitrogen in soil

Background and aims

Soil temperature influences nitrogen (N) diffusion in soil but until now, such effects have been difficult to quantify. This study aimed at estimating the effect of temperature on the diffusive fluxes of plant-available N forms in two contrasting soils.

Methods

Using a novel technique based on micro-dialysis, we established relationships between diffusive fluxes and temperature in aqueous solutions and in soil samples.

Results

Averaged for all compounds, the decreases of diffusive fluxes from the soil to the microdialysis sampler were 3.8 and 4.7% per degree Celsius in an agricultural and a boreal forest soil, respectively. The temperature-related shift of diffusive flux was, however, significantly dependent on molecular weight of the N compound. In accordance with established functions for temperature effects on diffusive fluxes, the non-linearity of this relationship resulted in a greater temperature response for small N compounds compared to larger compounds.

Conclusions

Our results show that, all other factors being equal, the relative contribution of smaller N compounds to the diffusive flux of total plant-available N increases with increasing soil temperatures.     

Relationships Among pH,Minerals, and Carbon in Soils from Tundra to Boreal Forest Across Alaska

Tundra and boreal forests in northern high latitudes contain significant amounts of carbon (C) in the soil, indicating the importance of clarifying controls on soil C dynamics in the region and their feedback effects on climate systems. In northern Alaska, variations in soil C processes are closely associated with variations in soil acidity within ecosystems; however, the reason for this association remains unclear. In this study, we demonstrate that it results from weathering and subsequent changes in soil geochemical characteristics, including minerals and adsorptive organic C. We sampled soils from 12 sites in Alaska along a 600-km transect from the Arctic Ocean to interior Alaska, spanning the biomes of tundra, tundra–boreal forest ecotone, and boreal forest. Mineral soil analyses revealed that soils with low pH have fewer base cations, more aluminum/iron minerals, and lower base saturation, indicating that weathering is a major function of these geochemical characteristics in the broad area over northern Alaska. Adsorbed organic C in soil presented strong correlations with Al and Fe minerals, soil pH, and soil total C and represented approximately 30–55% of total soil C, suggesting that soil C accumulation in the Alaskan ecosystems is strongly controlled by weathering-related changes in geochemical characteristics. An adsorption test supported these observations and illustrated a greater capacity for acidic soil to adsorb organic C. These findings demonstrate that variations in weathering-associated characteristics have a strong influence on the regional variation in C dynamics and biogeochemical consequences in the Alaskan ecosystems.     

Amino acid uptake in deciduous and coniferous taiga ecosystems

We measured in situ uptake of amino acids and ammonium across deciduous and coniferous taiga forest ecosystems in interior Alaska to examine the idea that late successional (coniferous) forests rely more heavily on dissolved organic nitrogen (DON), than do early successional (deciduous) ecosystems. We traced ¹⁵N-NH₄⁺ and ¹³C-¹⁵N-amino acids from the soil solution into plant roots and soil pools over a 24 h period in stands of early successional willow and late successional black spruce. Late successional soils have much higher concentrations of amino acid in soil solution and a greater ratio of DON to dissolved inorganic N (DIN) (ammonium plus nitrate) than do early successional soils. Moreover, late successional coniferous forests exhibit higher rates of soil proteolytic activity, but lower rates of inorganic N turnover. Differences in ammonium and amino acid uptake by early successional willow stands were insignificant. By contrast, the in situ uptake of amino acid by late successional black spruce forests were approximately 4-fold greater than ammonium uptake. The relative difference in uptake of ammonium and amino acids in these forests was approximately proportional to the relative difference of these N forms in the soil solution. Thus, we suggest that differences in uptake of different N forms across succession in these boreal forests largely reflect edaphic variation in available soil N (composition), rather than any apparent physiological specialization to absorb particular forms of N. These finding are relevant to our understanding of how taiga ecosystems may respond to increases in temperature, fire frequency, N deposition, and other potential consequences of global change.     

森林生态系统碳循环对全球氮沉降的响应

森林土壤和植被储存着全球陆地生态系统大约46%的碳,在全球碳平衡中起着非常重要的作用。过去几十年来,森林生态系统的碳循环和碳吸存受到了全球氮沉降的深刻影响,因为氮沉降改变了陆地生态系统的生产力和生物量积累。以欧洲和北美温带森林区域开展的研究为基础,综述了氮沉降对植物光合作用、土壤呼吸、土壤DOM及林木生长的影响特征和机理,探讨了森林生态系统碳动态对氮沉降响应的不确定性因素。热带森林C、N循环与大部分温带森林不同,人为输入的氮对热带生态系统过程的影响也可能不同,因此指出了在热带地区开展碳氮循环耦合研究的必要性和紧迫性。     

菌根真菌影响森林生态系统碳循环研究进展

森林生态系统在全球碳(C)储量中占据极为重要的地位。菌根真菌广泛存在于森林生态系统中,在森林生态系统C循环过程中发挥重要的作用。阐述了不同菌根类型真菌在森林生态系统C循环过程中的功能,对比了温带/北方森林与热带/亚热带森林中菌根真菌介导的C循环研究方面新近取得的研究结果。发现温带和北方森林的外生菌根(EcM)植物对地上生物量C的贡献相对较小,然而是地下C储量的主要贡献者;以丛枝菌根(AM)共生为主的热带/亚热带森林地表生物量占比较高,表明AM植被对热带/亚热带森林地上生物量C的贡献相对较大。我们还就全球变化背景下,菌根真菌及其介导的森林生态系统C汇功能,以及不同菌根类型树种影响C循环的机制等进行了总结。菌根真菌通过影响凋落物分解、土壤有机质形成及地下根系生物量,进而影响整个森林生态系统的C循环功能。菌根介导的森林C循环过程很大程度上取决于(优势)树木的菌根类型和森林土壤中菌根真菌的群落结构。最后指出了当前研究存在的主要问题以及未来研究展望。本文旨在明确菌根真菌在森林生态系统C循环转化过程中的重要生态功能,有助于准确地评估森林生态系统C汇现状,为应对全球变化等提供重要的依据。     

Ammonium assimilation and amino acid metabolism in conifers

Conifers are the most important group of gymnosperms, which include tree species of great ecological and economic importance that dominate large ecosystems and play an essential role in global carbon fixation. Nitrogen (N) economy has a special importance in these woody plants that are able to cope with seasonal periods of growth and development over a large number of years. As N availability in the forest soil is extremely low, efficient mechanisms are required for the assimilation, storage, mobilization, and recycling of inorganic and organic forms of N. The cyclic interconversion of arginine and the amides glutamine and asparagine plays a central role in the N metabolism of conifers and the regulation of these pathways is of major relevance to the N economy of the plant. In this paper, details of recent progress in our understanding of the metabolism of arginine and the other major amino acids glutamine, glutamate, aspartate, and asparagine in pine, a conifer model tree, are presented and discussed.     

陆地生态系统碳密度格局研究概述

 准确了解陆地生态系统中碳密度的时空格局及其影响因子和作用机制,对于估算和预测不同类型生态系统中的植被和土壤的碳存储能力、判定碳汇、制定缓解全球变化的合理政策措施,具有重要意义。该文综述了现有研究中发现的世界陆地生态系统碳密度空间分布的地带性规律及中国陆地生态系统碳密度格局的独特特点。在全球尺度上,植被碳密度分布与植物生物量格局基本一致,除北方森林外其余大部分随纬度升高而减小;土壤碳密度则随纬度升高而增大。陆地生态系统中北方森林和热带森林的总体碳密度最高,不同的是,前者的碳主要集中在土壤中,而后者则集中在植被中。但在区域尺度上,由于气候、地形及人类活动影响,这种规律性可能会发生变化甚至不起作用。水热条件、土壤养分、生物多样性、气候和大气CO2浓度的变化以及土地利用与覆盖变化等是碳密度空间格局形成和发生变化的驱动因子。在某一特定区域,它们通过直接或间接提高植被净初级生产力,抑制呼吸和分解作用来增加陆地生态系统碳密度。综合分析特定时空条件下各因子对碳存储量的影响是解释碳密度分布现状,预测碳密度格局变化的关键,但目前的研究对各项驱动因子的作用机制、影响强度及多个因子间的相互作用仍不是很清楚,需要加强该方面的研究力度。碳密度研究中的数据获取、机理分析和过程模拟等方面仍存在很大的不确定性,因此有必要建立规范统一的碳密度测量估算系统和更为精准有效的估算模型,进行多尺度、多精度水平的综合研究。