应用微根管法测定细根指标方法评述

树木细根(直径<2mm)在森林生态系统能量流动和物质循环中起着重要的作用。原有的细根生产周转研究中常采用的土钻法、内生长法、挖掘法、根室法和土柱法等,均不能直接观察到细根的动态变化。微根管法是一种非破坏性、可定点直接观察和研究植物根系的方法,为研究细根的生长、衰老、死亡、分解和再生长的过程提供了有效的工具,尤其适用于细根周转、寿命和分解等方面的研究。但该技术不能直接测定单位面积的细根生物量、细根化学组成及细根周转对土壤碳和养分循环的影响,需要与土钻法结合。本文就运用微根管法对细根生物量、生产、周转和寿命等指标的研究方法进行了评述。     

帽儿山温带落叶阔叶林细根生物量、生产力和周转率

细根在森林生态系统能量流动与物质循环中占有重要地位,但其生物量、生产和周转测定尚存在很大的不确定性,而且局域尺度空间变异机制尚不清楚。本研究分析了帽儿山温带天然次生林活细根生物量和死细根生物量在0～100 cm剖面的垂直分布与0～20 cm细根的季节动态、生产力和周转率,对比了采用连续根钻法(包括决策矩阵法和极差法)和内生长袋(直径3和5 cm)估测细根生产力和细根周转率,并探讨了可能影响细根的林分因子。结果表明: 76.8%的活细根生物量和62.9%的死细根生物量均集中在0～20 cm土层,随着深度增加,二者均呈指数形式减少。活细根生物量和死细根生物量的季节变化不显著,可能与冬季几乎无降雪而夏季降雨异常多有关。2种直径内生长袋估计的细根生产力无显著差异;对数转换后决策矩阵、极差法和内生长法估计的细根生产力和细根周转率差异显著。随着土壤养分增加,活细根生物量和死细根生物量比值显著增加,死细根生物量显著减少,但活细根生物量、细根生产力和细根周转率均无显著变化;细根周转率与前一年地上木质生物量增长量呈显著正相关,但与当年地上木质生物量增长量无显著相关关系。     

杉木林年龄序列地下碳分配变化

  森林地下碳分配在森林碳平衡和碳吸存中具有重要作用, 而揭示人工林生长过程中地下碳分配变化对于人工林碳汇估算和碳汇管理等有重要意义。通过采用年龄序列方法研究了杉木(Cunninghamia lanceolata)林生长过程中地下碳分配变化特点。年龄序列为福建省南平7 a生(幼龄林)、16 a生(中龄林)、21 a生(近熟林)、41 a生(成熟林)和88 a生(老龄林)的杉木林。细根净生产力测定采用连续土芯法, 根系呼吸测定采用壕沟法, 生物量增量测定采用异速生长方程, 地上年凋落物量采用凋落物收集框测定。结果表明: 杉木林细根净生产力在中龄林前没有显著差异, 维持在较高水平; 但此后则显著下降。细根净生产力/地上凋落物量比值随林龄增加而显著下降。老龄林的根系呼吸显著低于其它林龄林分, 根系呼吸与细根生物量间呈显著线性相关。中龄林和近成熟林的地下碳分配(Total belouground carbon allocation, TBCA)显著高于幼龄林和成熟林, 而老龄林的则最低。中龄林、近成熟林和成熟林的地上部分净生产力/TBCA比值显著高于幼龄林和老龄林, 而杉木林的根系碳利用效率(RCUE)则呈现出随林龄增加而降低的趋势。     

功能划分方法在树木细根生物量研究中的应用:进展与评述

树木细根具有高度的结构和功能的异质性。近20年来,人们逐渐认识到采用某一直径阈值来定义细根并开展研究的方法(即直径法)不能准确地描述相关的生理学和生态学过程。随着越来越多的研究证实根系功能与其分支等级密切相关,根序划分的方法(即根序法)在根系研究中得到更多的应用。但是,采用根序法测定相关功能属性需耗费大量的人力和时间。最近,有研究者在结合直径法和根序法优点的基础上提出了功能划分的方法。该方法将传统的细根区分为吸收根和运输根,在充分考虑根系结构和功能联系的同时,又能兼顾研究工作的效率和结果间的可比性,特别适用于根系生物量与周转的研究。采用功能划分方法来研究根系生物量(包括其他功能属性)是一个较新的观点,研究结果的规律性和存在的主要问题仍有待总结。该文作者通过查阅近年来的相关研究报告发现吸收根与运输根生物量在树种间存在较大的差异,目前对吸收根生物量在全球尺度上的变异格局并不清楚,吸收根与运输根对细根生物量周转的相对贡献有待探究,运输根在界定上存在着很大的挑战性。该文最后讨论了在根系研究中应用功能划分方法的优势与不足,并提出了建议。     

科尔沁沙地赤松和樟子松根系生物量分配与构型特征

沙地赤松(Pinus densiflora)在科尔沁沙地南缘区已有50年的引种历史,但其生长表现和根系生物量分配与构型特征还很少被报道。本研究以同龄(40 a)樟子松为对照,在生长指标测定基础上,采用分层分段全挖法采集根系,对沙地赤松不同径级根生物量分配规律进行研究,测定根长、连接数量、平均连接长度等指标,同时计算分形维数和分形丰度。结果表明:与樟子松相比,沙地赤松具有较大生长量和生物量,其根生物量显著高于樟子松,是其1.96倍,细根(直径≤0.2 cm)生物量更显著高于樟子松,是其4.76倍;沙地赤松根系生物量占总生物量的29.0%,细根生物量占总根系生物量的1.1%,细根长度占根总长度的44.3%;樟子松根系生物量占总生物量的25.6%,细根生物量占总根系生物量的0.4%,细根长度占根总长度的28.8%;从根系垂直分布看,沙地赤松地下0~180 cm均有细根分布,且40~180 cm范围内细根生物量占总细根生物量的65.2%;樟子松几乎全部细根分布于0~100 cm范围内,此范围细根生物量占总细根生物量的99.2%,且0~40 cm土层细根生物量占63.4%;虽然两树种根系平均连接长度没有显著差异,但沙地赤松细根及部分中根(0.2~2.0 cm)连接数量显著高于樟子松;沙地赤松根系分形维数为1.548±0.251,是樟子松(1.293±0.190)的1.2倍,并且分形丰度是樟子松的1.3倍;与樟子松相比,沙地赤松根系具有较强的吸收能力,能够利用较大范围的深层水分和养分,根系分支多,拓扑结构更加复杂。     

广西不同林龄杉木、马尾松人工林根系生物量及碳储量特征

为了解不同林龄杉木、马尾松人工林地地下根系生物量及碳储量特征,以广西杉木、马尾松主产区5个不同林龄阶段(幼龄林、中龄林、近熟林、成熟林、过熟林)的人工林为研究对象,采用全根挖掘法和土钻法获取标准木根系生物量、灌草根系生物量和林分细根生物量,并测定其碳含量,分析其不同林龄阶段地下根系生物量和碳储量分配特征。结果表明:杉木、马尾松林地下根系总生物量分别在9.06—31.40Mg/hm~2和7.91—53.40Mg/hm~2之间,各林龄阶段根系总生物量总体上呈现随林龄增加而增加的趋势,杉木林细根生物量随林龄的增加呈现出先减后增的趋势,马尾松呈现出逐渐减小的趋势;林分各层次根系碳含量表现为乔木灌木草本、细根;杉木、马尾松地下根系碳储量变化趋势与生物量变化趋势相同,杉木、马尾松林不同林龄阶段各层次根系和土壤细根总碳储量分别在7.56—21.97Mg/hm~2和8.86—29.95Mg/hm~2之间;地下根系碳储量总体上以乔木根系占优势,且随林龄的增大其比例呈增加的趋势。     

长白山阔叶红松林细根周转的研究

系统研究了长白山阔叶红松林细根生物量、生产力、年周转率及其在净生产力分配中的作用,生物量调查结果表明,阔叶红松林活细根的生物量为5049kg.ha~(-1),死细根生物量的平均值为1883kg.ha~(-1),细根的年周转率为0.96,年生产量为4860kg.ha~(-1),约占总净初级生产力的19.40％,年死亡量为2343kg.ha~(-1),相当于阔叶红松林枯枝落叶年凋落量的60％,由此提出了森林凋落物应包括枯枝落叶和根系凋落物的论点。     

实验增温对陆地生态系统根系生物量的影响

根系是植物吸收土壤水分和养分的重要器官, 驱动着多个生态系统过程, 该研究揭示了实验增温对根系生物量的影响及机制, 可为气候变暖背景下土壤碳动态和生态系统过程的变化提供理论依据。该研究从已发表的151篇国内外研究论文中收集到611组数据, 通过整合分析(meta-analysis)方法研究了实验增温对根系生物量(根系总生物量、粗根生物量、细根生物量、根冠比)的影响, 并探讨了增温幅度、增温年限、增温方式的影响, 以及根系生物量对增温的响应与本底环境条件(生态系统类型、年平均气温、年降水量、干旱指数)的关系。结果表明: (1)模拟增温使细根生物量显著增加8.87%, 而对根系总生物量、粗根生物量、根冠比没有显著影响; (2)中等强度增温(1-2 ℃)使得细根生物量和根冠比分别提高14.57%和23.63%; 中短期增温实验(<5年)对细根生物量具有促进影响, 而长期增温实验(≥5年)使细根生物量有降低的趋势; 开顶箱增温和红外辐射增温分别使细根生物量显著提高了17.50%和12.16%, 而电缆加热增温使细根生物量和粗根生物量显著降低了23.44%和43.23%; (3)不同生态系统类型对于增温响应不一致, 模拟增温使苔原生态系统细根生物量显著提高了21.03%, 细根生物量对增温的响应与本底年平均气温、年降水量、干旱指数均呈显著负相关关系。     

模拟氮沉降对中亚热带杉木幼树根系生物量的影响

植物根系是全球陆地生态系统碳储量的重要组成部分,在全球生态系统碳循环中起着重要作用,日益加剧的氮沉降会影响根系生物量在空间和不同径级的分配,进而影响森林生态系统的生产力和土壤养分循环。以杉木幼树为研究对象,通过野外氮沉降模拟实验,研究氮沉降四年后对不同土层、不同径级根系生物量的影响。结果发现：（1）低氮和高氮处理总细根生物量较对照均无显著差异（P > 0.05）,高氮处理粗根生物量及总根系生物量较对照分别增加45%和40%（P < 0.05）;（2）与对照相比,施氮处理显著增加20-40 cm与40-60 cm土层细根和粗根生物量,且在低氮处理下,20-40 cm土层细根、粗根在总土层细根与粗根生物量的占比显著提高。（3）与对照相比,高氮处理显著增加了2-5 mm、5-10 mm及10-20 mm径级的根系生物量,低氮处理显著增加2-5 mm、5-10 mm径级根系生物量,且显著降低20-50 mm径级根系生物量。综上所述表明：氮沉降后杉木幼树通过增加较粗径级根系来增加对养分及水分的输送,同时通过增加深层根系生物量及其比例的策略来维持杉木幼树的快速生长;而根系生物量的增加,在一定程度上会增加根系碳源的输入,影响土壤碳循环过程。     

高寒草甸植被细根生产和周转的比较研究

植物根系是陆地生态系统重要的碳汇和养分库,细根周转过程是陆地生态系统地下部分碳氮循环的核心环节,在陆地生态系统如何响应全球变化中起着关键作用。在全球变化敏感地区之一的青藏高原,对该地区的主要植被类型矮嵩草草甸同时采用根钻法、内生长袋法和微根管法3种观测方法研究细根生产和周转速率,并探讨了极差法、积分法、矩阵法和Kaplan-Meier法等数据处理方法对计算值的影响。研究结果显示:在估算细根净初级生产力时,根钻法宜采用积分法,内生长袋法宜选用矩阵法;由此进一步以最大细根生物量为基础,根钻法和内生长袋法估测的细根年周转速率分别为0.36 a-1和0.52 a-1,内生长袋法的估算结果是根钻法的1.44倍。对于微根管法,将其观测得到的细根长度转换为单位面积的生物量值后,采用积分法计算出细根周转速率为0.84 a-1,远高于传统方法的估算结果;若采用Kaplan-Meier生存分析方法,则计算出的细根周转速率更高达3.41 a-1。     

Root sampling methods - applications and limitations of the minirhizotron technique

Applications and limitations of the minirhizotron technique (non-destructive) in relation to two frequently used destructive methods (soil coreing and ingrowth cores) is discussed. Sequential coreing provides data on standing crop but it is difficult to obtain data on root biomass production. Ingrowth cores can provide a quick estimate of relative fine-root growth when root growth is rapid. One limitation of the ingrowth core is that no information on the time of ingrowth and mortality is obtained.The minirhizotron method, in contrast to the destructive methods permits simultaneous calculation of fine-root length production and mortality and turnover. The same fine-root segment in the same soil space can be monitored for its life time, and stored in a database for processing. The methodological difficulties of separating excavated fine roots into living and dead vitality classes are avoided, since it is possible to judge directly the successive ageing of individual roots from the images. It is concluded that the minirhizotron technique is capable of quantifying root dynamics (root-length production, mortality and longevity) and fine-root decomposition. Additionally, by combining soil core data (biomass, root length and nutrient content) and minirhizotron data (length production and mortality), biomass production and nutrient input into the soil via root mortality and decomposition can be estimated.     

Measuring Fine Root Turnover in Forest Ecosystems

Development of direct and indirect methods for measuring root turnover and the status of knowledge on fine root turnover in forest ecosystems are discussed. While soil and ingrowth cores give estimates of standing root biomass and relative growth, respectively, minirhizotrons provide estimates of median root longevity (turnover time) i.e., the time by which 50% of the roots are dead. Advanced minirhizotron and carbon tracer studies combined with demographic statistical methods and new models hold the promise of improving our fundamental understanding of the factors controlling root turnover. Using minirhizotron data, fine root turnover (y⁻¹) can be estimated in two ways: as the ratio of annual root length production to average live root length observed and as the inverse of median root longevity. Fine root production and mortality can be estimated by combining data from minirhizotrons and soil cores, provided that these data are based on roots of the same diameter class (e.g., < 1 mm in diameter) and changes in the same time steps. Fluxes of carbon and nutrients via fine root mortality can then be estimated by multiplying the amount of carbon and nutrients in fine root biomass by fine root turnover. It is suggested that the minirhizotron method is suitable for estimating median fine root longevity. In comparison to the minirhizotron method, the radio carbon technique favor larger fine roots that are less dynamics. We need to reconcile and improve both methods to develop a more complete understanding of root turnover.     

微根管在细根研究中的应用

细根（直径≤2 mm）的周转在植物生态系统碳分配过程中具有重要意义.已往细根周转研究主要采用根钻法、分室模型法和内生长法等.这些方法由于不能直接观测到细根生长动态,导致细根周转估计不准确.微根管法是一种非破坏性野外观察细根动态的方法.本文从微根管的发展、功能、安装步骤、图像采集、参数计算、影响观测因素和存在问题等方面逐一进行介绍,并通过水曲柳和落叶松微根管细根观测实例介绍在细根周转过程研究中的应用. 结果表明,微根管可以比较精确地估计出细根长度、单位面积上根长密度、单位体积上根长密度、细根生长量、细根死亡量和细根周转等.微根管是一个观察细根生长、衰老、死亡和分解过程的有效工具.微根管观测精度主要取决于微根管安装的质量和数量、微根管取样间隔期和取样数量、微根管图像分析技术等.此外,土壤质地、石砾多少、微根管材料选择、减少光系统对根系的干扰等也是影响微根管测定精度的因素.如何提高微根管测定精度将成为今后微根管在细根研究中的主要问题.     

Estimating Root Production: Comparison of 11 Methods in Shortgrass Steppe and Review of Biases

Estimating root production has been difficult due to multiple potential biases associated with both old and new methods. This shortgrass steppe site is the only place we are aware of that can compare most methods including sequential coring, ingrowth cores, and ingrowth donuts, ¹⁴C pulse-isotope dilution, ¹⁴C pulse-isotope turnover, rhizotron windows, and minirhizotron, and indirect methods including nitrogen budget, carbon flux, simulation carbon flow model, and regression model. We used the studies at this site, other comparisons, a summary of potential directional biases, and different ways of calculating estimates in a logical, comparative approach of evaluating methods. Much of the literature for root production is based on sequential biomass coring, a method resulting in erroneous estimates. Root ingrowth estimates of production are generally conservative compared to minirhizotron and isotope turnover methods. The size of the ingrowth area may be the most important determinant of the underestimation. Estimates based on pulse-isotope dilution are also erroneous due to non-uniform labeling of tissues. Uniform labeling is not an assumption of the pulse-isotope turnover method, and this method has the least severe potential biases. Root production estimates from pulse-isotope turnover were lower than those using minirhizotron when the most common method of calculation was used. This agrees with literature concerning bomb ¹⁴C continuous-isotope labeling comparisons with minirhizotron, although some potential biases between isotope methods are different. However, good agreement between pulse-isotope turnover and minirhizotron were obtained when minirhizotron estimates were calculated from regression of decomposition versus production to equilibrium and when pulse-isotope turnover estimates were calculated from two-phase life-span regressions. This minirhizotron method bypasses biases associated with the artificial surface similar to root-cohort methods that may be practical only in mesic systems, and takes into account both short- and long-lived roots and corrects for soil-isotope contamination that the continuous-isotope labeling bomb ¹⁴C method is not able to account for. Comparisons of these direct methods are also made with four indirect methods.     

Belowground drought response of European beech: fine root biomass and carbon partitioning in 14 mature stands across a precipitation gradient

How tree root systems will respond to increased drought stress, as predicted for parts of Central Europe, is not well understood. According to the optimal partitioning theory, plants should enhance root growth relative to aboveground growth in order to reduce water limitations. We tested this prediction in a transect study with 14 mature forest stands of European beech (Fagus sylvatica L.) by analysing the response of the fine root system to a large decrease in annual precipitation (970–520 mm yr⁻¹). In 3 years with contrasting precipitation regimes, we investigated leaf area and leaf biomass, fine root biomass and necromass (organic layer and mineral soil to 40 cm) and fine root productivity (ingrowth core approach), and analysed the dependence on precipitation, temperature, soil nutrient availability and stand structure. In contrast to the optimal partitioning theory, fine root biomass decreased by about a third from stands with >950 mm yr⁻¹ to those with <550 mm yr⁻¹, while leaf biomass remained constant, resulting in a significant decrease, and not an increase, in the fine root/leaf biomass ratio towards drier sites. Average fine root diameter decreased towards the drier stands, thereby partly compensating for the loss in root biomass and surface area. Both δ¹³C‐signature of fine root mass and the ingrowth core data indicated a higher fine root turnover in the drier stands. Principal components analyses (PCA) and regression analyses revealed a positive influence of precipitation on the profile total of fine root biomass in the 14 stands and a negative one of temperature and plant‐available soil phosphorus. We hypothesize that summer droughts lead to increased fine root mortality, thereby reducing root biomass, but they also stimulate compensatory fine root production in the drier stands. We conclude that the optimal partitioning theory fails to explain the observed decrease in the fine root/leaf biomass ratio, but is supported by the data if carbon allocation to roots is considered, which would account for enhanced root turnover in drier environments.     

Fine root dynamics in lodgepole pine and white spruce stands along productivity gradients in reclaimed oil sands sites

Open‐pit mining activities in the oil sands region of Alberta, Canada, create disturbed lands that, by law, must be reclaimed to a land capability equivalent to that existed before the disturbance. Re‐establishment of forest cover will be affected by the production and turnover rate of fine roots. However, the relationship between fine root dynamics and tree growth has not been studied in reclaimed oil sands sites. Fine root properties (root length density, mean surface area, total root biomass, and rates of root production, turnover, and decomposition) were assessed from May to October 2011 and 2012 using sequential coring and ingrowth core methods in lodgepole pine (Pinus contorta Dougl.) and white spruce (Picea glauca (Moench.) Voss) stands. The pine and spruce stands were planted on peat mineral soil mix placed over tailings sand and overburden substrates, respectively, in reclaimed oil sands sites in Alberta. We selected stands that form a productivity gradient (low, medium, and high productivities) of each tree species based on differences in tree height and diameter at breast height (DBH) increments. In lodgepole pine stands, fine root length density and fine root production, and turnover rates were in the order of high > medium > low productivity sites and were positively correlated with tree height and DBH and negatively correlated with soil salinity (P < 0.05). In white spruce stands, fine root surface area was the only parameter that increased along the productivity gradient and was negatively correlated with soil compaction. In conclusion, fine root dynamics along the stand productivity gradients were closely linked to stand productivity and were affected by limiting soil properties related to the specific substrate used for reconstructing the reclaimed soil. Understanding the impact of soil properties on fine root dynamics and overall stand productivity will help improve land reclamation outcomes.     

植物地下生物量研究进展

在生物量的研究过程中,地下部分生物量十分重要却又多被忽视,本文首先比较了当地地下生物量研究中较为常用的4种研究方法：挖土块法、钻土芯法、内生长土芯法和微根区管法,然后总结了地下生物量在空间和时间上变化规律的研究成果,最后又讨论了环境因素诸如水分、温度和开垦、放牧对地下生物量的影响以及地下生物量的周转。