林木细根寿命及其影响因子研究进展

 细根周转要消耗大量的C,它影响森林生态系统C分配格局与过程和养分循环,对生态系统生产力具有重要意义。细根的周转取决于细根的寿命,细根寿命越短,周转越快,根系对C的消耗也越多。大量研究表明,细根的寿命与地上部分C向根系供应的多少有密切关系,同时也与细根直径大小、土壤中N和水分的有效性、土壤温度以及根际周围的土壤动物和微生物的活动有关。本文综述了国外近年来在该领域里的研究进展,特别是对控制细根寿命的机理和主要影响因子进行了评述,目的是引起国内研究者的关注,促进我国根系生态学的研究与发展。     

微根管法和同位素法在细根寿命研究中的应用及比较

细根的生产和周转在陆地生态系统的碳和养分循环中起着重要作用,并且对全球环境变化具有一定的指示意义。细根寿命是估计细根周转的关键,其长短决定了养分和碳消耗与循环的速度。由于采用的研究方法不同,导致所得细根寿命估计值存在较大差异,目前最新的同位素和微根管2种方法之间寿命估计值差异可达10倍以上。本文对这2种研究方法的原理和优点进行了阐述,并从细根定义、细根寿命理论分布假设、细根取样误差等方面对导致这2种方法研究结果存在差异的原因进行分析,以期有助于今后根系研究的发展。     

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。     

遮荫对水曲柳幼苗细根衰老的影响

细根周转对森林生态系统碳地下分配和养分循环具有重要影响,而衰老是细根周转过程中最重要的阶段。根据“源-汇”理论,细根衰老受碳向细根分配的影响。为此,该研究通过控制水曲柳(Fraxinus mandshurica)苗木向根系的碳分配的遮荫处理试验,采用树木生理分析技术,重点研究了在光合产物供应停止情况下水曲柳幼苗根系的生理变化(即根尖、1级根到3级根的细胞活力、可溶性糖含量、可溶性蛋白含量和膜透性的变化)。目的是从生理水平上证实:1)碳分配对细根衰老产生怎样的影响;2)细根衰老的顺序是否与分支顺序相反。实验结果表明,遮荫处理使细根活力、可溶性糖含量、可溶性蛋白含量显著降低和膜透性增大,导致细根出现明显衰老。从根系顶端向基部随着根序增加,细胞活力、可溶性糖含量和可溶性蛋白含量提高,膜透性降低,从生理水平上表明细根衰老具有逐渐变化的顺序性,并且这种顺序性与根发育的顺序性相反。     

国外对树木细根的研究动态

森林生态系统的研究已经深入到了生态系统的各个层次,但无论是在哪一层次上的研究,其重点往往是树木的地上部分,而对地下部分——根系的研究相对较为薄弱。主要原因是对根系的生态学作用认识不足以及根系研究的困难。近来,随着对根系特别是细根在养分循环及能流中的重要作用的认识,细根的研究已成为森林生态系统研究的一个热点。根系中粗根(直径>5mm)虽然占总生物量的20%左     

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。     

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。     

增温对林木细根寿命影响的研究进展

细根寿命作为林木生长的重要参数，不仅对林木吸收水分和养分能力具有重要作用，而且影响林木对光合产物的地下分配以及森林生态系统碳循环过程。气候变暖对细根寿命的影响已成为全球变化背景下关注的热点问题，细根寿命对全球变暖的响应将影响到生态系统的碳平衡。本文就林木细根寿命对增温的响应特征及其响应机制的研究进展进行综述，以期揭示增温对林木细根寿命的影响。目前已开展的大部分研究认为，增温会通过改变根际土壤环境、细根形态以及林木物候等指标来影响细根寿命。但细根生长和死亡受诸多因素的影响，导致增温对细根寿命影响的研究结果因研究区域自然环境、增温方式和研究对象等因素存在分歧。因此，综合分析气候变暖背景下林木细根寿命的变化对研究地下生态学过程至关重要。今后还应加强以下研究：1)结合多种方式进行地下、地上同时增温，并探索更准确和有效的无损观测方法；2)结合多种观测手段研究增温对细根寿命的影响；3)开展增温对不同树种细根寿命的研究，深入理解不同林木细根寿命对增温的响应机理；4)多角度综合分析增温对细根寿命的影响特征，探究多种因素综合作用对细根寿命的影响机理；5)增温与其他环境因子对细根寿命交互作用；6)增温后根...     

瓦屋山常绿阔叶次生林建群种扁刺栲的细根分布及其碳氮特征

四川瓦屋山国家森林公园是我国西部的中亚热带湿性常绿阔叶林的典型代表,具有四川现存的较为完好的扁刺栲(Castanopsis platyacantha)-华木荷(Schima sinensis)群系,该研究利用土钻法探讨了该群系内主要建群种扁刺栲标准木的细根分布及其碳氮特征。结果表明:(1)扁刺栲细根总生物量为173.62 g·m~(-2),其中活根生物量为135.29 g·m~(-2)。(2)随着土层深度的增加,扁刺栲细根生物量、根长密度、根系表面积和比根长呈下降趋势,0~30 cm土层所占比例分别为67.23%、69.53%、69.48%和57.20%;根长密度、根系表面积和比根长均随细根直径的增加而显著下降,直径小于1 mm的根系所占比例分别为58.84%、52.59%和51.36%。(3)扁刺栲细根生物量、根长和表面积消弱系数β均随根系直径的增加而增加。(4)根系C含量在第Ⅰ土层中随细根直径的增加而显著增加,在其他土层则无显著差异;直径小于2 mm的根系C含量在第Ⅰ土层中显著低于其他土层,大于2 mm的根系C含量在各土层间的差异不大。(5)根系N含量随根系直径和土层深度的增加而减少,C/N值则与之相反。该研究结果在一定程度上反映了该次生林地下细根的垂直分布及养分特征,为揭示该生态系统地下生态过程及今后在该生态系统研究环境变化对地下生态过程的影响提供了基础数据。     

模拟氮沉降对杉木幼苗细根的生理生态影响

细根对氮沉降的生理生态响应将显著影响森林生态系统的生产力和碳吸存。为了揭示氮沉降对杉木细根的生理生态影响,对一年生杉木(Cunninghamia lanceolata)幼苗进行了模拟氮沉降试验,并测定施氮1年后杉木幼苗细根生物量、细根形态学特征(比根长、比表面积)、元素化学计量学指标(C、N、P、C/N、C/P、N/P)、细根代谢特征(细根比呼吸速率、非结构性碳水化合物)。结果表明:(1)杉木细根生物量随氮添加水平的升高而显著降低,尤其是0—1 mm细根生物量;细根比根长和比表面积随氮添加水平升高而显著增大。(2)氮添加后杉木细根C含量、C/N、C/P显著降低,高氮添加导致1—2 mm细根N含量和N/P显著升高,而低氮添加导致1—2 mm细根P含量显著升高、N/P显著降低,而0—1 mm细根的N、P含量则保持相对稳定。(3)氮添加后杉木细根比呼吸速率无显著变化,细根可溶性糖含量随氮添加增加而显著增加,而淀粉含量和NSC显著降低。综合以上结果表明:氮添加后用于细根形态构建的碳分配减少,这可能会减少土壤中有机碳的保留,0—1 mm细根的形态更易发生变化,但是其内部N、P养分含量相对更稳定以维持生理活动,细根NSC对氮添加的响应表明施氮可能导致细根受光合产物的限制。     

Seasonal dynamics of fine root biomass,root length density,specific root length,and soil resource availability in a Larix gmelinii plantation

Fine root tumover is a major pathway for carbon and nutrient cycling in terrestrial ecosystems and is most likely sensitive to many global change factors.Despite the importance of fine root turnover in plant C allocation and nutrient cycling dynamics and the tremendous research efforts in the past,our understanding of it remains limited.This is because the dynamics processes associated with soil resources availability are still poorly understood.Soil moisture,temperature,and available nitrogen are the most important soil characteristics that impact fine root growth and mortality at both the individual root branch and at the ecosystem level.In temperate forest ecosystems,seasonal changes of soil resource availability will alter the pattern of carbon allocation to belowground.Therefore,fine root biomass,root length density(RLD)and specific root length(SRL)vary during the growing season.Studying seasonal changes of fine root biomass,RLD,and SRL associated with soil resource availability will help us understand the mechanistic controls of carbon to fine root longevity and turnover.The objective of this study was to understand whether seasonal variations of fine root biomass,RLD and SRL were associated with soil resource availability,such as moisture,temperature,and nitrogen,and to understand how these soil components impact fine root dynamics in Larix gmelinii plantation.We used a soil coring method to obtain fine root samples(≤2 mm in diameter)every month from Mav to October in 2002 from a 17-year-old L.gmelinii plantation in Maoershan Experiment Station,Northeast Forestry University,China.Seventy-two soil cores(inside diameter 60 mm;depth intervals:0-10 cm,10-20 cm,20-30 cm)were sampled randomly from three replicates 25 m×30 m plots to estimate fine root biomass(live and dead),and calculate RLD and SRL.Soil moisture,temperature,and nitrogen(ammonia and nitrates)at three depth intervals were also analyzed in these plots.Results showed that the average standing fine root biomass(live (32.2 g.m-2.a-1)in the middle(10-20 cm)and deep layer (20-30cm),respectively.Live and dead fine root biomass was the highest from May to July and in September,but lower in August and October.The live fine root biomass decreased and dead biomass increased during the growing soil layer.RLD and SRL in May were the highestthe other months,and RLD was the lowest in Septemberdynamics of fine root biomass,RLD,and SRL showed a close relationship with changes in soil moisture,temperature,and nitrogen availability.To a lesser extent,the temperature could be determined by regression analysis.Fine roots in the upper soil layer have a function of absorbing moisture and nutrients,while the main function of deeper soil may be moisture uptake rather than nutrient acquisition.Therefore,carbon allocation to roots in the upper soil layer and deeper soil layer was different.Multiple regression analysis showed that variation in soil resource availability could explain 71-73% of the seasonal variation of RLD and SRL and 58% of the variation in fine root biomass.These results suggested a greater metabolic activity of fine roots living in soil with higher resource availability,which resulted in an increased allocation of carbohydrate to these roots,but a lower allocation of carbohydrate to those in soil with lower resource availability.     

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。     

落叶松和水曲柳人工林细根生长、死亡和周转

 细根周转是陆地生态系统碳分配格局与过程的核心环节,而细根周转估计的关键是了解细根的生长和死亡动态。该研究以18年生落叶松(Larix gmelinii)和水曲柳(Fraxi nus mandshurica)人工林为对象,采用微根管（Minirhizotron）技术对两树种0～40 cm深度的细根生长和死亡动态进行了为期1年的观测,研究了两树种细根在不同土层深度的生长与死亡动态、细根周转以及与土壤有效氮含量、土壤温度、大气温度和降水的关系。结果表明：1） 落叶松平均细根生长（Root length density production, RLDP）0.0045 mm•cm^－2•d^－1）明显低于水曲柳RLDP（0.0077 mm•cm^－2•d^－1）。两个树种细根平均RLDP在表层（0～10 cm）最大,而底层（30～40 cm）最小 ,两树种平均细根死亡（Root length density mortality, RLDM）也表现同样规律 。水曲柳春季生长的细根占41.7%,夏季占39.7%,而落叶松细根生长分别是24.0%和51.2%,水曲柳细根死亡主要发生在春季（34.3%） 和夏季（34.0%）,而落叶松细根死亡主要发生在夏季和秋季（分别占28.5%和32.3%）,两 树种细根生长与死亡在冬季均较小;2）落叶松细根年生长量（0.94 mm•cm^－2•a^－1）和年死亡量（0.72 mm•cm^－2•a^－1）明显低于水曲柳（1.52和1.21 mm•cm^－2•a^－1）,两树种细根表层年生长量和年死亡量均最高,底层最低。落叶松细根年周转为3.1次•a^－1（按年生长量计算）和2.4次•a^－1（按年死亡量计算）,相比较,水曲柳细根年周转分别为2.7次•a^－1和2.2次•a^－1;3）土壤有效氮含量、土壤温度、大气温度和降水综合作用影响细根生长和死亡动态,可以解释细根生长80%的变异和细根死亡95%以上的变异。     

Herbivore-induced changes in plant carbon allocation: assessment of below-ground C fluxes using carbon-14

Effects of above-ground herbivory on short-term plant carbon allocation were studied using maize (Zea mays) and a generalist lubber grasshopper (Romalea guttata). We hypothesized that above-ground herbivory stimulates current net carbon assimilate allocation to below-ground components, such as roots, root exudation and root and soil respiration. Maize plants 24 days old were grazed (c. 25–50% leaf area removed) by caging grasshoppers around individual plants and 18 h later pulse-labelled with¹⁴CO₂. During the next 8 h,¹⁴C assimilates were traced to shoots, roots, root plus soil respiration, root exudates, rhizosphere soil, and bulk soil using carbon-14 techniques. Significant positive relationships were observed between herbivory and carbon allocated to roots, root exudates, and root and soil respiration, and a significant negative relationship between herbivory and carbon allocated to shoots. No relationship was observed between herbivory and¹⁴C recovered from soil. While herbivory increased root and soil respiration, the peak time for¹⁴CO₂ evolved as respiration was not altered, thereby suggesting that herbivory only increases the magnitude of respiration, not patterns of translocation through time. Although there was a trend for lower photosynthetic rates of grazed plants than photosynthetic rates of ungrazed plants, no significant differences were observed among grazed and ungrazed plants. We conclude that above-ground herbivory can increase plant carbon fluxes below ground (roots, root exudates, and rhizosphere respiration), thus increasing resources (e.g., root exudates) available to soil organisms, especially microbial populations.     

森林生态系统细根周转规律及影响因素

根系周转是陆地生态系统碳循环的关键过程, 对研究土壤碳库变化及全球气候变化均具有重要意义。然而由于根系周转率的测量计算方法较多, 不同方法得出的结果差异较大, 且目前对全球区域尺度上森林生态系统根系周转的研究还不够充分, 使得全球森林生态系统根系周转变化规律仍不清楚。该研究通过收集文献数据并统一周转率计算方法, 对全球5种森林类型的细根周转空间格局进行整合, 同时结合土壤理化性质和气候数据, 得出影响森林生态系统细根周转的因子。结果表明, 不同森林类型细根周转率存在显著差异, 且随着纬度的升高逐渐降低; 森林生态系统细根周转率与年平均温度和年平均降水量呈正相关; 森林生态系统细根周转率与土壤有机碳含量呈正相关但与土壤pH值呈负相关。该研究为揭示森林生态系统细根周转规律及机制提供了科学依据。     

Allometric constraints on,and trade‐offs in,belowground carbon allocation and their control of soil respiration across global forest ecosystems

To fully understand how soil respiration is partitioned among its component fluxes and responds to climate, it is essential to relate it to belowground carbon allocation, the ultimate carbon source for soil respiration. This remains one of the largest gaps in knowledge of terrestrial carbon cycling. Here, we synthesize data on gross and net primary production and their components, and soil respiration and its components, from a global forest database, to determine mechanisms governing belowground carbon allocation and their relationship with soil respiration partitioning and soil respiration responses to climatic factors across global forest ecosystems. Our results revealed that there are three independent mechanisms controlling belowground carbon allocation and which influence soil respiration and its partitioning: an allometric constraint; a fine‐root production vs. root respiration trade‐off; and an above‐ vs. belowground trade‐off in plant carbon. Global patterns in soil respiration and its partitioning are constrained primarily by the allometric allocation, which explains some of the previously ambiguous results reported in the literature. Responses of soil respiration and its components to mean annual temperature, precipitation, and nitrogen deposition can be mediated by changes in belowground carbon allocation. Soil respiration responds to mean annual temperature overwhelmingly through an increasing belowground carbon input as a result of extending total day length of growing season, but not by temperature‐driven acceleration of soil carbon decomposition, which argues against the possibility of a strong positive feedback between global warming and soil carbon loss. Different nitrogen loads can trigger distinct belowground carbon allocation mechanisms, which are responsible for different responses of soil respiration to nitrogen addition that have been observed. These results provide new insights into belowground carbon allocation, partitioning of soil respiration, and its responses to climate in forest ecosystems and are, therefore, valuable for terrestrial carbon simulations and projections.     

落叶松人工林细根动态与土壤资源有效性关系研究

树木细根在森林生态系统C和养分循环中具有重要的作用。由于温带土壤资源有效性具有明显的季节变化, 导致细根生物量、根长密度 (Rootlengthdensity, RLD) 和比根长 (Specificrootlength, SRL) 的季节性变化。以 17年生落叶松 (Larixgmelini) 人工林为研究对象, 采用根钻法从 5月到 10月连续取样, 研究了不同土层细根 (直径≤ 2mm) 生物量、RLD和SRL的季节动态, 以及这些根系指标动态与土壤水分、温度和N有效性的关系。结果表明 :1) 落叶松细根年平均生物量 (活根 +死根 ) 为 189.1g·m-2 ·a-1, 其中 5 0 %分布在表层 (0~ 10cm), 33%分布在亚表层 (11~ 2 0cm), 17%分布在底层 (2 1~ 30cm) 。活根和死根生物量在 5~ 7月以及 9月较高, 8月和 10月较低。从春季 (5月 ) 到秋季 (10月 ), 随着活细根生物量的减少, 死细根生物量增加 ;2 ) 土壤表层 (0~ 10cm) 具有较高的RLD和SRL, 而底层 (2 1~ 30cm) 最低。春季 (5月 ) 总RLD和SRL最高, 分别为 10 6 2 1.4 5m·m-3 和 14.83m·g-1, 到秋季 (9月 ) 树木生长结束后达到最低值, 分别为 2 198.2 0m·m-3 和 3.77m·g-1;3) 细根生物量、RLD和SRL与土壤水分、温度和有效N存在不同程度的相关性。从单因子分析来看, 土壤水分和有效N对细根的影响明显大于温度, 对活根的影响大于死根。由于土壤资源有效性的季节变化, 使得C的地下分配格局发生改变。各土层细根与有效性资源之间的相关性反映了细根功能季节性差异。细根 (生物量、RLD和SRL) 的季节动态 (5 8%~ 73%的变异 ) 主要由土壤资源有效性的季节变化引起。