Above- and belowground biomass and primary productivity of a Larix gmelinii stand near Tura, central Siberia

We assessed above- and belowground biomass and net primary production (NPP) of a mature Larix gmelinii (Rupr.) Rupr. forest (240-280 years old) established on permafrost soils in central Siberia. Specifically, we investigated annual carbon budgets in roots in relation to root system development and availability of soil resources. Total stand biomass estimated by allometry was about 39 Mg per ha. Root biomass (17 Mg per ha) comprised about 43% of total biomass. Coarse root (>/= 5 mm in diameter) biomass was about twice that of fine roots (< 5 mm). The aboveground biomass/root biomass ratio (T/R) of the larch stand was about unity, which is much less than that of other boreal and subalpine conifer forests. The proportion of fine roots in total root biomass (35%) was relatively high compared with other cold-climate evergreen conifer forests. Total NPP, defined as the sum of annual biomass increment of woody parts and needle biomass, was estimated to be 1.8 Mg per ha per year. Allocation of total NPP to needle production was 56%. The proportion of total NPP in belowground production (27%) was less than for evergreen taiga forests. However, belowground NPP was probably under-estimated because root mortality was excluded. We conclude that L. gmelinii trees invested annual carbon gains largely into needle production or roots, or both, at the expense of growth of aboveground woody parts. This carbon allocation pattern, which resulted in the construction of exploitative root networks, appeared to be a positive growth response to the nutrient-poor permafrost soil of central Siberia.     

水曲柳人工林细根季节动态及其影响因素

2002年5-10月,采用连续钻取土芯法对帽儿山实验林场的水曲柳人工林细根(直径＜1 mm)生物量、比根长(SRL)和根长密度(RLD)的季节动态,以及它们与土壤N的有效性、土壤10 cm深处月均温度和含水量的关系进行研究.结果表明:水曲柳细根生物量在春季和秋季分别具有1个明显的高峰,但比根长和根长密度只有1个高峰.在春季和夏季,比根长和根长密度较高,显示细根直径较小,而秋季,这2个参数显著下降,表明细根直径次生增厚或组织密度增加.细根的季节变化与土壤N的有效性、土壤温度和土壤含水量有重要关系.其中细根生物量与土壤铵态氮含量显著相关;硝态氮含量、10 cm深处土壤的温度和土壤含水量与细根的生物量、比根长和根长密度的季节变化正相关,但均不显著(P＞0.05).4种因子的综合作用对水曲柳细根各参数的影响均达到了显著水平.不同季节细根生物量、比根长和根长密度的变化,显示出细根在生长季不同时期具有不同的生理生态功能.     

水曲柳落叶松混交林中细根空间分布

采用根钻取样方法对年生水曲柳落叶松混交林中细根空间分布状况进行了研究。结果表明,水曲柳落叶松地下生物量的空间分配差异显著。在林分水平上,水曲柳的根生物量密度高于落叶松(分别为4442.3和2234.9g/m3)。两树种在相邻区域中分配的细根生物量较高,表明种间根系竞争较弱。落叶松行间的水曲柳细根生物量密度和根长密度均高于水曲柳行间的落叶松细根,表明水曲柳地下部分具有较强能力。根系的空间分布有利于混交林中水曲柳的生长。图1表4参19。     

Definition of fine roots on the basis of the root anatomy,diameter,and branch orders of one-year old Fraxinus mandshurica seedlings

Fine roots are important in root absorption of nutrient and water,and in root turnover.Accurate definition of fine roots is a prerequisite to improved estimation of the physiological and ecological functions of forest ecosystems.Root development and physiological functions are reflections of root anatomical structure.In this study,the anatomical structures of different root orders were analyzed by examining paraffin sections of one-year old Fraxinus mandshurica seedlings.One-year-old F.mandshurica seedlings had over five root orders.The root anatomical structures of all orders showed more differences.First and second order roots consisted of four sections: the epidermis,cortex,pericycle,and vascular bundles.Fourth and fifth order roots were mainly composed of the skin and peripheral vascular bundles(including the xylem and phloem).Third order roots had root epidermal and cortical structures,but the quantity and integrity of the cortical cells were inferior to those of the first and second order roots,and superior to those of the fourth and fifth order roots.All the first and second order roots and some third order roots with discontinuous cork layer(0.4 mm in diameter),but not the fourth and fifth order roots,were the fine roots of one-year old F.mandshurica seedlings.Although they had similar diameters,different portions of root systems had different anatomical structures and therefore,vary in capacity to absorb water and nutrients.Fine roots were accurately defined by root diameter,branch orders,and anatomical structural features of one-year old F.mandshurica seedlings.     

森林细根生产和周转研究

随着近 2 0多年来对细根功能的深入认识和研究方法的发展 ,林分细根生物量、生产和周转及其与环境因子的关系成为森林生态学的研究热点之一 ,开展了大量研究。本文在收集了大量研究报道基础上 ,对森林细根研究结果进行综述。结果如下 :细根 (直径 <2～ 5mm)生物量变化在 46～ 2 80 5g·m- 2 之间 ,大部分在 10 0～ 10 0 0g·m- 2 ;细根生物量分别占地下部分总生物量和林分总生物量的 3%～ 30 %和 0 .5 %～ 10 % ;北方常绿针叶林平均细根生物量最低 (2 16g·m- 2 ) ,热带常绿阔叶林最高 (10 87g·m- 2 )。细根年净生产量 2 0～ 1317g·m- 2 ·a- 1 ,占林分总净初级生产量的3%～ 84% ,大部分在 10 %～ 6 0 % ;从北方森林到温带、亚热带至热带森林 ,细根生产量呈增加趋势 ;针叶林细根生产在总净初级生产中的比例小于阔叶林 (常绿和落叶 )。树木细根生命周期短至数天 ,长达数年。细根年周转率 4.3%～ 2 73.2 % ,阔叶林细根周转率低于针叶林。细根生产和周转是土壤碳和养分的重要来源 ,细根生产向林地输入的生物量占总输入 (细根生产和地上枯落物输入 )的 6 .2 %～ 88.7%。除气候森林类型外 ,森林生态系统细根生物量、分布、生产、周转还因季节、土壤类型、立地条件和生长发育阶段而异。同时 ,还受树木体内碳     

桢楠幼苗对模拟氮沉降的生长响应

为了解桢楠(Phoebe zhennan)幼苗的生长对模拟氮沉降的响应,以1年生桢楠幼苗为材料,设置5个氮水平CK(0)、N1(5.7 g·m-2·a-1)、N2(11.4 g·m-2·a-1)、N3(19.0 g·m-2·a-1)和N4(38.0 g·m-2·a-1),对不同施氮水平下幼苗的株高、地径、生物量和根系指标进行研究。结果表明,不同施氮处理均促进了株高、地径和叶面积的增长,株高增长率、地径增长率和叶面积均以N3处理最高,分别为CK的3.93、1.51和1.33倍;不同施氮处理均提高了幼苗的生物量,各施氮处理下整株鲜重分别较CK提高了22.15%、31.53%、96.45%和45.82%,以N3处理最高;随着施氮量的增加,幼苗的总根长、根直径、根表面积、根体积和根尖数均先升高后降低;隶属函数分析结果显示N3处理的桢楠幼苗质量最佳,说明适度氮沉降能促进幼苗生长,但更高水平的氮沉降可能会产生一定抑制作用。     

林分密度对侧柏人工林细根形态的影响

以徐州林场50年生侧柏人工林为研究对象,采用挖掘法获取土壤根系样品,探究林分密度对侧柏人工林不同根序细根形态的影响。结果表明:细根的直径和根长随着根序上升而显著增大,而比根长则随着根序的上升而显著减小。低林分密度(1 679株/hm2)与中林分密度(2 250株/hm2)相比显著减小了表层土壤1、2级细根的平均直径和平均根长,亚表层土壤3级细根的根长,显著增大了亚表层土壤2级细根的平均比根长;高林分密度(3074株/hm2)比中林分密度显著增大了2级根的平均比根长。与高林分密度相比,低林分密度显著减小了表层土壤1、2级细根的平均直径,增大了亚表层土壤5级细根的平均直径。     

林分年龄对落叶松人工林细根生物量的影响

【目的】随着森林的发育过程,林木个体的生长和生物量分配,以及林分水平的结构和功能均发生了明显的变化。然而,细根生物量与林分年龄的联系,目前仍然了解有限。本研究以黑龙江省帽儿山地区兴安落叶松人工林为研究对象,比较了同一林分在19年和32年生时林分水平(单位面积)和单株水平细根生物量的垂直分布和季节动态,分析了影响细根生物量变化的林分与土壤因子,旨在明确林分年龄对细根生物量的影响和潜在的机制。【方法】在生长季内的5月、7月和9月,采用土钻法获取土壤0~30 cm深度细根并测定生物量,同时测定林分特征和土壤养分和水分含量。【结果】随林龄增加,落叶松人工林单位面积细根生物量显著下降,而单株细根生物量变化不显著;与19年生林分相比,32年生林分土壤表层(0~10 cm)细根生物量占总细根生物量的比例明显下降,土壤亚表层(10~20 cm)和底层(20~30 cm)细根生物量所占比例增加,呈现出细根向深层土壤增生的趋势。土壤表层(0~10cm)单位面积细根生物量随林分年龄的变化趋势与林分密度和胸高断面积、土壤铵态氮浓度变化有关,但是单株细根生物量受林分和土壤因子的影响均不显著。【结论】林分发育过程中,落叶松细根生物量降低,细根的资源吸收策略发生了明显的改变。     

Variations of fine root diameter with root order in Manchurian ash and Dahurian larch plantations

Fine root lifespan and turnover play an important role in carbon allocation and nutrient cycling in forest ecosystems. Fine roots are typically defined as less than 1 or 2 mm in diameter. However, when categorizing roots by this diameter size, the position of an individual root on the complex lateral branching pattern has often been ignored, and our knowledge about relationships between branching order and root function thus remains limited. More recently, studies on root survivals found that longevity was remarkably different in the same branching level due to diameter variations. The objectives of this study were: (1) To examine variations of fine root diameter from the first-to fifth-orders in Fraxinus mandshurica Rupr and Larix gmelinii Rupr roots; and (2) To reveal how the season, soil nutrient, and water availability affect root diameter in different branch order in two species. This study was conducted at Maoershan Forest Research Station (45°21′–45°25′N, 127°30′–127°34′E) owned by Northeast Forestry University in Harbin, northeast China. Both F. mandshurica and L. gmelinii were planted in 1986. In each plantation, fine roots of two species by sampling up to five fine root branch orders three times during the 2003 growing season from two soil depths (i.e., 0–10 and 10–20 cm) were obtained. The results showed that average diameters of fine roots were significantly different among the five branch orders. The first-order had the thinner roots and the fifth order had the thickest roots, the diameter increasing regularly with the ascending branch orders in both species. If the diameter of fine roots was defined as being smaller than 0.5 mm, the first three orders of F. mandshurica roots and the first two orders of L. gmelinii roots would be included in the fine root population. The diameter ranges of the fine roots from first-order to fifth-order were 0.15–0.58, 0.18–0.70, 0.26–1.05, 0.36–1.43, and 0.71–2.96 mm for F. mandshurica, and 0.17–0.76, 0.23–1.02, 0.26–1.10, 0.38–1.77, and 0.84–2.80 mm for L. gmelinii. The average coefficient of variation in first-order roots was less than 10%, second-and third-order was 10–20%, and fourth-and fifth-order was 20–30%. Thus, variation in root diameter also increased with the ascending root order. These results suggest that “fine roots”, which are traditionally defined as an arbitrary diameter class (i.e., <2 mm in diameter) may be too large a size class when compared with the finest roots. The finest roots have much shorter lifespan than larger diameter roots; however, the larger roots are still considered a component of the fine root system. Differences in the lifespan between root diameter and root order affect estimates of root turnover. Therefore, based on this study, it has been concluded that both diameter and branch order should be considered in the estimation of root lifespan and turnover. __________ Translated from Acta Phytoecologica Sinica, 2005, 29(6): 871–877 [译自: 植物生态学报]     

Estimation of the fine root biomass in a Japanese cedar (Cryptomeria japonica) plantation using minirhizotrons

We estimated fine root biomass in a Japanese cedar (Cryptomeria japonica) plantation using a min-irhizotron technique. Since data obtained from minirhizo-trons are limited to the length and diameter of fine roots observed on minirhizotron tubes, data conversion is necessary to determine the fine root biomass per unit soil volume or unit stand area. We first examined the regression between diameter squared and weight per unit length of fine roots in soil core samples, and calculated the fine root biomass on minirhizotron tubes from their length and diameter. Then we determined conversion factors based on the ratio of the fine root biomass in soil core samples to that on minirhizotron tubes. We examined calculation methods, using a single conversion factor for total fine root biomass in the soil for depths of 0–40cm (Cal1), or using four conversion factors for fine roots in the soil at 10-cm intervals (Cal2). Cal1 overestimated fine root biomass in the lower soil or underestimated that in the upper soil, while fine root biomass calculated using Cal2 better matched that in soil core samples. These results suggest that minirhizotron data should be converted separately for different soil depths to better estimate fine root biomass.     

植物细根生产和周转研究进展

细根是植物根系最重要的组成部分,作为衡量植物生产力的重要因素,对森林生态系统生产力具有重要影响。前人研究表明,细根的生产与周转对细根的寿命、分解和生物量估算具有重要意义,并且会影响森林生态系统碳、养分和水循环过程。文中系统阐述了细根生产和周转的研究进展,介绍了细根的3种主要研究方法（根钻法、内生长法和微根管法）,进一步分析细根生产和周转的影响因素,即除了受植物内在因子（细根构型、根序和化学组成）的制约外,细根生产和周转还受到纬度、海拔、气候、土壤条件、土层深度等环境因子及生物因子的影响;探讨了在植物细根研究中存在的问题,并对今后的发展趋势进行了展望,以期为植物细根深入研究和根系生态学学科发展提供参考。     

Fine root biomass,distribution, and production in young pine-hardwood stands

Patterns of fine root biomass, production, and distribution were estimated for pure stands and mixtures of three-year-old loblolly pine (Pinus taeda L.) with red maple (Acer rubrum L.) or black locust (Robinia pseudoacacia L.) on the Virginia Piedmont to determine the role of fine roots in interference between pine and hardwood tree species. Estimates were based on amounts of live and dead fine roots separated from monthly core samples during the third growing season after planting. Live and dead fine root biomass and production varied by species, but mixtures of loblolly pine and black locust generally had greater fine root biomass and fine root production than pure stands or loblolly pine-red maple mixtures. Hardwood species had greater live fine root biomass per tree in mixtures with pine compared to pure stands. Greater live fine root biomass in pine-locust stands may be attributed to differential utilization of the soil volume by fine roots of these species. For all stands, approximately 50% of live five root biomass was located in the upper 10 cm of soil.     

Biomass of fine and small roots in two Japanese black pine stands of different ages

The biomass and the spatial distribution of fine and small roots were studied in two Japanese black pine (Pinus thunbergii Parl.) stands growing on a sandy soil. More biomass of fine and small roots was found in the 17-year-old than in the 40-year-old stand. There were 62 g m⁻² of fine roots and 56 g m⁻² of small roots in the older stand, which represented mean values of 608 g for fine and 552 g for small roots per tree, respectively. In the younger stand, a total of 85 g m⁻² of fine roots and 66 g m⁻² of small roots were determined, representing a mean of 238 g for fine and 186 g for small roots per tree, respectively. Fine and small root biomasses decreased linearly with a soil depth of 0–50 cm in the older stand. In the younger stand, the fine and small roots developed only up to a depth of 30 cm. Horizontal distributions (with regard to distance from a tree) of both root groups were homogeneous. A positive correlation in the amount of biomass of fine and small roots per m² relative to tree size was found. Fine and small root biomasses increased consistently from April to July in both stands. The results also indicated earlier growth activity of the fine roots than small roots at the beginning of the growing season. The seasonal increases in fine and small root biomasses were slightly higher in the younger stand than the older stand.     

Space-time dynamics of fine root biomass of six forests in the Maoershan forest region, northeast China

The Maoershan forestry centre is situated in the Zhangguangcai Mountain of the Changbai mountain range. The main forest types in the Maoershan region are plantation (Pinus sylvestris var. mongolica, Pinus koraiensis and Larix gmelinii) and natural secondary forests (Fraxinus mandshurica, Quercus mongolica and Populus davidiana). Fine roots have enormous surface areas, growing and turning over quickly, which plays an important role in terms of substance cycling and energy flow in the forest ecosystem. This study deals with the dynamics of live, dead, and total fine roots (≤ 5 mm) biomass in the 0–30 cm soil layer using the soil core method. Differences between the six stands in the Maoershan region showed the following results: 1) the fine root biomass in the various stands showed obvious differences. The total fine root biomass of six stands from high to low were F. mandshurica (1,030.0 g/m²) > Q. mongolica (973.4 g/m²) > Pinus koraiensis (780.9 g/m²) > L. gmelinii (718.2 g/m²) > Populus davidiana (709.1 g/m²) > Pinus sylvestris var. mongolica (470.4 g/m²); 2) except for L. gmelinii, the development of live fine root biomass agreed with the trend of total fine root biomass. The maximum biomass of live fine roots in Pinus koraiensis or L. gmelinii stand appeared in May, others in June; in the F. mandshurica stand, the minimum biomass of live fine roots occurred in September, others in July or August; 3) the proportions of dead fine root biomass varied in different stands; 4) the vertical distribution of fine roots was affected by temperature, water, and nutrients; the proportion of fine root biomass was concentrated in the 0–10 cm soil layer. The fine root biomass of six stands in the 0–10 cm soil layer was over 40% of the total fine root biomass; this proportion was 60.3% in F. mandshurica. Space-time dynamics of the various stands had different characteristics. When investigating the substance cycling and energy flows of all forest ecosystems, we should consider the characteristics of different stands in order to improve the precision of our estimates. __________ Translated from Scientia Silvae Sinicae, 2006, 42(6): 13–19 [译自: 林业科学]     

Spatial heterogeneity of fine root biomass of Pinus massoniana forests in the Three Gorges Reservoir Area, China

Environmental heterogeneity is a constant presence in the natural world that significantly affects plant behavior at a variety of levels of complexity. In order to estimate the spatial pattern of fine root biomass in the Three Gorges Reservoir Area, the spatial heterogeneity of fine root biomass in the upper layer of soils (0-10 cm) in three Masson pine (Pinus massoniana) stands in the Three Gorges Reservoir Area, China, was studied in 30 m × 30 m plots with geostatistical analysis. The results indicate that 1) both the live and dead fine root biomass of stand 2 were less than those of other stands, 2) the spatial variation of fine roots in the three stands was caused together by structural and random factors with moderate spatial dependence and 3) the magnitude of spatial heterogeneity of live fine roots ranked as: stand 3 > stand 1 > stand 2, while that of dead fine roots was similar in the three stands. These findings suggested that the range of spatial autocorrelation for fine root biomass varied considerably in the Three Gorges Reservoir Area, while soil properties, such as soil bulk density, organic matter and total nitrogen, may exhibit great effect on the spatial distribution of fine roots. Finally, we express our hope to be able to carry out further research on the quantitative relationship between the spatial heterogeneous patterns of plant and soil properties.     

水曲柳育苗密度对苗木质量的影响

水曲柳(Fraxinus mandshurica Rupr.)是东北地区三大珍贵硬阔叶用材树种之一,也是东北地区的造林树种.垄作时,水曲柳育苗密度决定苗木的产量和质量,试验结果表明,当密度达到50株/m时,是水曲柳的最优育苗密度,其苗高、地径、主根长和侧根数最优,高径比上的比例比较合适,有利于优质壮苗的培育.     

Foliage, fine-root, woody-tissue and stand respiration in Pinus radiata in relation to nitrogen status

We measured respiration of 20-year-old Pinus radiata D. Don trees growing in control (C), irrigated (I), and irrigated + fertilized (IL) stands in the Biology of Forest Growth experimental plantation near Canberra, Australia. Respiration was measured on fully expanded foliage, live branches, boles, and fine and coarse roots to determine the relationship between CO(2) efflux, tissue temperature, and biomass or nitrogen (N) content of individual tissues. Efflux of CO(2) from foliage (dark respiration at night) and fine roots was linearly related to biomass and N content, but N was a better predictor of CO(2) efflux than biomass. Respiration (assumed to be maintenance) per unit N at 15 degrees C and a CO(2) concentration of 400 micro mol mol(-1) was 1.71 micro mol s(-1) mol(-1) N for foliage and 11.2 micro mol s(-1) mol(-1) N for fine roots. Efflux of CO(2) from stems, coarse roots and branches was linearly related to sapwood volume (stems) or total volume (branches + coarse roots) and growth, with rates for maintenance respiration at 15 degrees C ranging from 18 to 104 micro mol m(-3) s(-1). Among woody components, branches in the upper canopy and small diameter coarse roots had the highest respiration rates. Stem maintenance respiration per unit sapwood volume did not differ among treatments. Annual C flux was estimated by summing (1) dry matter production and respiration of aboveground components, (2) annual soil CO(2) efflux minus aboveground litterfall, and (3) the annual increment in coarse root biomass. Annual C flux was 24.4, 25.3 and 34.4 Mg ha(-1) year(-1) for the C, I and IL treatments, respectively. Total belowground C allocation, estimated as the sum of (2) and (3) above, was equal to the sum of root respiration and estimated root production in the IL treatment, whereas in the nutrient-limited C and I treatments, total belowground C allocation was greater than the sum of root respiration and estimated root production, suggesting higher fine root turnover or increased allocation to mycorrhizae and root exudation. Carbon use efficiency, the ratio of net primary production to assimilation, was similar among treatments for aboveground tissues (0.43-0.50). Therefore, the proportion of assimilation used for construction and maintenance respiration on an annual basis was also similar among treatments.     

Individual-based fine root biomass and its functional relationship with leaf for <Emphasis Type="Italic">Pinus tabuliformis</Emphasis> in northern China

Root biomass plays an essential role in carbon sequestration at both individual and ecosystem scales, yet few feasible methods for measuring root biomass of individual trees have been developed. We sampled 11 trees in a pure 20-year-old Pinus tabuliformis plantation to estimate total root, fine root and leaf biomass of individual trees using the nested regression method. The relationships between fine root biomass and surface area as well as leaf biomass and area were examined across tree sizes. Fine roots of P. tabuliformis were estimated to consume 40 % of the net primary production for turnover, and absorbed a total of 1364 g m^?2 year^?1 of major nutrients. A significant linear relationship was determined between roots and leaves in terms of both biomass and surface area. The findings added weight to the concept that the functional balance theory is applicable to even-aged mature trees of P. tabuliformis.     

Impact factors on fine root seasonal dynamics in Fraxinus mandshurica plantations

Fine root turnover plays important roles in carbon allocation and nutrient cycling in forest ecosystems. Seasonal dynamics of fine roots is critical for understanding the processes of fine root turnover. From May to October 2002, soil core method was used for estimating the seasonal pattern of fine root (diameter < 1 mm) parameters (biomass, specific root length (SRL) and root length density (RLD)) in a Manchurian ash (Fraxinus mandshurica) plantation located at the Maoershan Experiment Station, Heilongjiang Province, northeast of China. The relationships of fine root biomass, SRL and RLD with available nitrogen in soil, average soil temperature per month in 10 cm depth and soil moisture content were analyzed. Seasonal variation of fine root biomass was significant (P < 0.05). The peak values of fine root biomass were observed both in spring and in autumn, but SRL and RLD were the highest in spring and lowest in autumn. Specific root length and root length density were higher in spring and summer, which means that fine root diameter was thinner. In autumn, both parameters decreased significantly due to secondary incrassation of fine root diameter or the increase of tissue density. Seasonal dynamics of fine roots was associated with available nitrogen in soil, soil temperature in 10 cm depth and moisture content. Fine root biomass has a significant relationship with available NH₄ ⁺-N in soil. Available NO₃ ⁻-N in soil, soil temperature in 10-cm depth and moisture content have a positive correlation with fine root biomass, SRL and RLD, although these correlations are not significant (P > 0.05). But the compound effects of soil available N, soil temperature and soil moisture content are significant to every root parameter. The variations of these three root parameters in different seasons show different physiological and ecological functions in different growing periods. Translated from Scientia Silvae Sinicae, 2006, 42(9): 7–12 [译自: 林业科学]     

Distribution of root biomass in a low‐shrub pine bog

The average stump and below‐ground biomass of pine was 1464 g/m²; 4% as fine roots (Ø<1 mm), 18% small roots (Ø=1–10 mm), 49% large roots (Ø>10 mm), and 29% stumps), which comprised 35% of the total biomass in the Scots pine stands. The average root length of pine was 728 m/m²: 71 % of this length was fine roots, 29% was small roots, and less than 1 % was large roots. Most of the fine pine roots (80%) were in the 0–10 cm peat layer. The root biomass of the field layer was 548 g/m², which comprised 47 % of the total field layer biomass. Characteristic features of the root systems were: high below‐ground/above‐ground ratios, rather low amounts of root biomass, shallow rooting, and relatively thin roots. Hummocks tended to have less roots in the 0–10 cm layer and more roots in the 10–20 cm layer than the hollows.