Competitive interactions between Nardus stricta L. and Calluna vulgaris (L.) Hull: the effect of fertilizer and defoliation on above- and below-ground performance

1 The role of nutrient supply and defoliation on the competitive interactions between pot-grown Calluna vulgaris and Nardus stricta plants was investigated.WP leading adjustment
2 Young plants were grown alone and together in pots under a combination of fertilizer and defoliation treatments. After 18 months, parameters reflecting both above- and below-ground performance were measured, namely: total above-ground biomass, shoot nitrogen and phosphorus content, root length and the extent of mycorrhizal infection of the roots.
3 In the pots that received fertilizer, the shoot nutrient content and above-ground biomass of Nardus plants increased to a greater extent than those of Calluna plants; this effect was more marked for Nardus plants growing with Calluna plants than for those growing with other Nardus plants. In contrast , Calluna plants growing in competition with Nardus failed to respond to the addition of nutrients. However, in unfertilized pots, Calluna gained more above-ground biomass during the experimental period than Nardus.
4 Calluna had greater root length than Nardus , but Nardus had a higher proportion of its root length infected by mycorrhizal fungi. In both plants, the addition of fertilizer reduced the mycorrhizal infection and increased the root length. Nardus root length was decreased when grown in competition with Calluna only in pots where no nutrients were added. Defoliation decreased the extent of mycorrhizal infection in Calluna roots but not in those of Nardus; defoliation decreased the shoot nutrient content in Calluna plants, but not in Nardus plants.
5 These results suggest that the competitive balance between Nardus and Calluna may be altered by the addition of nutrients, and by defoliation, which may have serious implications for the future dominance of Calluna in heathland ecosystems, particularly those where nutrient inputs are increasing significantly or where grazing pressures are high.     

异质养分环境中一年生分蘖草本黍根系的生长特征

为揭示黍（Panicum miliaceum L.)根系对异质养分环境的生长反应,作研究了黍根系从起始斑块向目标斑块水平生长时,时始斑块和目标斑块养分水平根生长的影响,就低养分起始珏块而言,粗根生物量,粗根长度,粗根表面积和细极长度在高养分目标斑块中的分配比例均小于其在低养分目标斑块中的分配比例,而细根长度及其密度,细根表面积指及其密度的变化恰好相反,就高养分起始斑块而言,高养分目标斑块的细根长度,细根长度密度,细根表面积指数和细根表面积密均不于低养分目标斑块,而粗根对目标斑块中养分状的反应不明显。当黍根系从桢的起始斑块进入不同的目标斑块后,目标斑块的养分状况对细根生物量及其分配无影响,而显影响细根长度和表现积,这指示细根是通过长度和表面积可塑性而不是生物量变化响应目标斑块中的养分差异。     

Root Growth of the Annual Tillering Grass Panicum miliaceum in Heterogeneous Nutrient Environments (in English)

To study growth responses of the roots of Panicum miliaceum L. to heterogeneous supply of nutrients. The authors analyzed the effects of the nutrient levels in both original patches (O) and destination patches (D) on the root growth of P. miliaceum when its roots were allowed to extend from original patch into destination patch. When the nutrient levels in the original patches were low, coarse root biomass ratio (coarse root biomass in the D/total coarse root biomass), coarse root length ratio (coarse root length in the D/total coarse root length), coarse root surface area ratio (coarse root surface area in the D/total coarse root surface area) and fine root length ratio (fine root length in the D/total fine root length) were greater in the destination patches with lower nutrient levels than in the destination patches with higher nutrient levels, while fine root length, fine root length density, fine root surface index, and fine root surface area density were smaller in the former than in the latter. When the nutrient levels in the original patches were high, fine root length, fine root length density, fine root surface area index and fine root surface density were greater in the destination patches with lower nutrient levels than in the destination patches with higher nutrient levels, coarse roots did not respond to the nutrient levels in the destination patches significantly. When the roots extended from the original patches with the same nutrient level into the destination patches with contrasting nutrient levels, fine root biomass and its percentage allocation did not respond to the nutrient levels in the destination patches significantly, whereas both root length and root surface area did. This indicates that the fine roots of P. miliaceum responded to difference in nutrient supply by plasticity in their length and surface area, rather than in their root biomass.     

两种入侵植物与三种本地植物根系特征的比较研究

以菊科2种入侵植物飞机草(Chromolaena odorata(L.)R.M.King&H.Rob)和紫茎泽兰(Eupatorium adenophorum Speng)以及生活型相似的3种本地植物异叶泽兰(Eupatorium heterophyllum DC.)、佩兰(Eupatorium fortune Turcz.)和白头婆(Eupatorium japonicum Thunb.)为研究对象,以单种为对照,分析5种植物的株高、生物量以及细根形态和结构特征对混种(竞争)的响应。结果显示,在单种情况下,2种入侵植物的株高和地上生物量比3种本地植物高。2种入侵植物的株高和地上生物量在单种和混种下无显著差异,但在混种下3种本地植物的株高、地上生物量、根生物量、细根长度、细根表面积和体积与单种相比显著降低。在混种情况下,2种入侵植物的根组织密度显著降低,但比根长和比根面积仅在紫茎泽兰中显著增加。5种植物在单种和混种下的株高与细根长度、根生物量呈显著正相关,地上生物量与细根体积、表面积呈显著正相关。主成分分析结果表明入侵植物位于株高和地上生物量更高的一端。说明在与本地植物竞争的过程中,飞机草和紫茎泽兰能通过调整根系碳投资策略实现更强的竞争力。     

Development of fine and coarse roots of Thuja occidentalis `Brabant' in non-irrigated and drip irrigated field plots

Aboveground dry mass, total root dry mass and root length density of the fine roots of Thuja occidentalis `Brabant' were determined under non- and drip-irrigated field conditions. Two-dimensional diffusion parameters for dynamic root growth were estimated based on dry mass production of the fine roots and the concept of the convective-diffusion model of cylindrical root growth and proliferation. Drip irrigation increased above-ground dry mass and the shoot:root ratio compared with no irrigation. Dry mass of the coarse roots increased as well due to drip irrigation. No effect on total or fine root dry mass was found. Drip irrigation increased root length densities in the top 0.1 m but not significantly. However, drip irrigation decreased root proliferation in depth by 27%, whereas proliferation in the horizontal direction was not altered. Measured root length densities were overestimated by 6–21% by the model (0.68 < R ² < 0.92).     

色季拉山急尖长苞冷杉叶片及细根性状随海拔的变异特征

叶片和细根是植物对环境变化响应的主要功能器官,在气候变化趋势下,研究其随环境梯度的变化,对理解植物对环境的适应策略具有重要意义。本文是在色季拉山急尖长苞冷杉分布区,按海拔梯度(3800、3900、4000、4100、4200、4300、4400m)选择7个典型样地,在样地内对其叶片基本特征、叶绿素含量、比叶重和细根特征(0—60cm)等参数进行了测量。结果表明:(1)随海拔梯度升高,急尖长苞冷杉叶片叶面积减小、叶片厚度增加、叶绿素含量逐渐降低、比叶重显著增大。3900m处急尖长苞冷杉的叶片形态特征表现最好,叶片长度、宽度和面积均显著优于其他海拔,海拔4200m叶片厚度达到最大值,叶片面积、叶绿素含量随海拔升高呈下降趋势,但在4200m处出现第二峰值。(2)随海拔梯度增加,细根各性状与海拔表现出非线性关系,其中细根生物量和细根体积在3900m和4200m处出现两次峰值,3900m处细根生物量达到最大值,4200m处细根体积达到最大值,4400m处细根生物量和细根体积均显著少于其他海拔。细根根长密度在海拔3900、4200、4300m较高,三者之间差异不显著,但显著高于其余海拔,4400m海拔细根根长密度最小。细根表面积在3900m海拔处达到最大值,显著高于其他海拔,4200和4300m次之,3800、4000和4400m海拔下细根表面积相对较小。4400m处细根比根长达到最大值。各海拔上细根均主要分布在0—20cm土层。随土层厚度增加,各海拔细根生物量和根系体积在0—60cm土层范围内均逐渐减小;细根根长密度、表面积在20—40cm和40—60cm显著提高;同一海拔细根比根长随土层深度增加呈明显的增加趋势。各海拔40—60cm土层细根比根长显著高于20—40和0—20cm土层。(3)综合叶片及细根特征,海拔3900m为急尖长苞冷杉的最适宜生长区域;随海拔升高,环境因子逐渐恶劣,环境胁迫加剧,急尖长苞冷杉最终形成在4400m处为其分布上限的海拔梯度格局。     

杉木成熟林细根形态与功能特征的海拔梯度变异特点

为探究植物对环境变化的适应策略,在安徽省金寨县天马国家自然保护区,以不同海拔高度(750、850、1000、1150 m)杉木(Cunninghamia lanceolata)成熟林为对象,采用土钻法获取土壤细根样品,分别测定了不同海拔不同土层(0—10 cm、10—20 cm、20—30 cm)土壤细根生物量、形态特征参数和碳氮含量。结果表明:(1)随海拔梯度增加,0—30 cm土层细根生物量、根长密度、比根长、表面积密度、体积密度均呈先减少后增加趋势,在海拔750 m生物量最大,其余指标在海拔1150 m最大;随土层深度增加,同一海拔细根生物量、根长密度、表面积密度、体积密度均呈减少趋势。(2)随海拔梯度增加,0—30 cm土层细根C和N含量呈先增加后减少趋势,C/N比呈先减少后增加再减少趋势;随土层深度增加,同一海拔细根C含量呈先减少后增加趋势,N含量呈降低趋势,C/N比呈上升趋势。(3)细根N含量与生物量、根长密度和体积密度显著正相关,C/N比与生物量、根长密度、表面积密度和体积密度极显著负相关。(4)土壤水分对细根生物量及其形态指标影响显著。     

杨树人工林细根数量和形态特征的季节动态及代际差异

采集欧美杨107Ⅰ代和Ⅱ代人工林细根样品,分析杨树不同根序细根数量特征(根长度、表面积和生物量)和形态特征(比根长、根长密度、根组织密度)对季节波动的响应及其代际差异.结果表明： 杨树各根序细根数量特征(根长度、表面积和生物量)均呈明显的季节变化,且具有明显的根序差异性.低级根序细根数量特征季节差异显著,细根生物量在生长季显著增加而生长季后显著下降.高级根序细根比根长季节波动显著,而根长密度和根组织密度等形态特征波动较小.连作导致人工林杨树1～2级细根长度、生物量、比根长和根长密度在生长季显著增大.1级细根数量特征与土壤温湿度呈显著正相关,与土壤有机质和速效氮含量呈显著负相关;而2级细根数量特征仅与土壤养分显著相关.杨树人工林细根特征的季节动态及代际差异体现了杨树对细根的碳投入变化,因连作引发的土壤养分匮乏可能引发植株对根系的碳投入增加,这种碳分配格局与人工林地上部分生产力形成密切相关.     

A cyclical but asynchronous pattern of fine root and woody biomass production in a hardwood forest of southern Quebec and its relationships with annual variation of temperature and nutrient availability

In contrast to the well-documented seasonal variation in growth of below- and above-ground components of trees, the annual variation in below- and aboveground production is not well understood. In this study, we report on the monitoring of an unmanaged hardwood forest ecosystem in a small watershed of southern Quebec between 1993 and 1999. Below- and above-ground biomass production, leaf and soil solution chemistry, and air temperature were measured at different regular intervals and are reported on an annual basis. The objective of the study was to describe the annual dynamics of carbon partitioning between below- and above-ground tree components and to gain a better understanding of the soil and climatic factors that govern it. Fine root production peaked one year earlier than woody biomass production and years with high production of fine roots had low woody biomass production. All models that included May temperature in the calculation of the predicting/independent variables were significant predictors of total tree biomass production (r > 0.87). Fine root production was associated negatively with the previous year average growing season temperature (r < -0.84). Soil solution NO₃ ^–, NH₄ ⁺ and NO₃ ^– + NH₄ ⁺ concentrations were positively correlated with fine root production (r > 0.72) and negatively correlated with woody biomass production (r < -0.84). Leaf N and P concentrations were negatively correlated (r = -0.99 and r = -0.98, respectively) with fine root production for the period of 1994–1998. Our results suggest that a cool growing season, and in particular a cool month of October, is likely to result in low fine root production and nutrient uptake the following year and, therefore, to increase soil N availability and decrease leaf N. This initial response is thought to be the first step of a feedback loop involving plant N nutrition, soil N availability, fine root growth and aboveground biomass production that led to a cyclical (3–4 years) but asynchronous production of fine roots and aboveground biomass production.     

Fibrous root turnover and growth in sugar beet (Beta vulgaris var. saccharifera) as affected by nitrogen shortage

The root system of plants is subject to fast cycles of renewal and decay within the growing season. In water and/or nutrient stress conditions, this turnover may become strategic for plant survival and productivity, but knowledge about its mechanisms is still insufficient. In order to investigate the effects of nitrogen fertilization on growth and turnover of sugar beet roots, an experiment was carried out over two growing seasons in northern Italy with two levels of N supply (0, 100 kg ha^–1). Biomass production and partitioning were followed during growth, and fibrous root dynamics were inspected by means of computer-aided procedures applied to minirhizotron images.In conditions of N shortage, lower yields (storage roots) were associated with greater allocation of biomass to tap roots (final tap-root/shoot ratio = 5.6 vs. 4.1) and shallower distribution of fibrous root length density. The maximum depth of roots was not affected by N, but unfertilized plants reached it more slowly.The ratio of cumulative root dead length to produced length at the end of the growing period (TDL _max/TPL _max) was used as the most suitable approach for assessing overall root turnover. This ratio was greater in controls (0.73 vs. 0.50), which showed lower root longevity (–34% life-span on average), indicating that a greater proportion of root growth was renewed by unfertilized plants over the season.     

郭岩山不同海拔丝栗栲细根功能性状及其与土壤因子的关系

为揭示丝栗栲(Castanopsis fargesii)细根功能性状对环境变化的适应机制,对郭岩山500、700、900 m海拔处丝栗栲细根功能性状及其与土壤因子的关系进行研究。结果表明,丝栗栲细根生物量与细根根长密度、表面积密度、组织密度及体积密度呈正相关,细根根长密度、体积密度、表面积密度和比根长4个性状间均呈极显著正相关关系,且均与细根组织密度呈显著负相关。根际土含水量、C和N含量与细根比根长、根长密度、体积密度、表面积密度均存在显著正相关关系,而土壤容重与细根组织密度呈正相关。海拔700 m的细根生物量、根长密度、表面积密度及体积密度显著大于海拔500和900 m的。500和900 m海拔的根长密度、表面积密度与土壤深度呈负相关,而500 m海拔细根的组织密度与土壤深度呈正相关。因此,郭岩山丝栗栲通过改变细根功能性状来适应海拔和土壤的变化。     

Fine‐root exploitation strategies differ in tropical old growth and logged‐over forests in Ghana

Understanding the changes in root exploitation strategies during post‐logging recovery is important for predicting forest productivity and carbon dynamics in tropical forests. We sampled fine (diameter < 2 mm) roots using the soil core method to quantify fine‐root biomass and architectural and morphological traits to determine root exploitation strategies in an old growth forest and in a 54‐yr‐old logged‐over forest influenced by similar parent material and climate. Seven root traits were considered: four associated with resource exploitation potential or an ‘extensive’ strategy (fine‐root biomass, length, surface area, and volume), and three traits which reflect exploitation efficiency or an ‘intensive’ strategy (specific root area, specific root length, and root tissue density). We found that total fine‐root biomass, length, surface area, volume, and fine‐root tissue density were higher in the logged‐over forest, whereas the old growth forest had higher total specific root length and specific root surface area than the logged‐over forest. The results suggest different root exploitation strategies between the forests. Plants in the old growth forest invest root biomass more efficiently to maximize soil volume explored, whereas plants in the logged‐over forest increase the spatial distribution of roots resulting in the expansion of the rhizosphere.     

水曲柳苗木地下竞争与地上竞争的定量研究

对3种密度进行栽培试验,利用通径分析的方法,研究了水曲柳地下竞争和地上竞争的关系及对总竞争的影响。结果表明,水曲柳苗木的地下部分生物量、地上部分生物量和总生物量与营养空间有密切关系。随着苗木空间距离增加,由生物量计算的竞争指数下降。在同一密度条件下,地下竞争指数明显大于地上竞争指数。由于地下生长与地上生长的相互作用,各竞争指数之间具有明显的相关性。但是地下竞争和地上竞争对总竞争的影响是不同的,通径分析可以定量的区分地下竞争和地上竞争的相对大小。地下竞争对总竞争的直接作用范围在0．5543～0．7426之间,明显大于地上竞争对总竞争的直接作用(0．2851～0．5282)。随着距离的增加,单株苗木的生长空间加大,地上部分的竞争作用增加,地下部分的竞争程度减弱。但是,地下根系的竞争在水曲柳苗木总的竞争中占有重要地位。     

弓杠岭不同海拔云杉细根生物量及形态特征

海拔变化是多环境因子的梯度效应,细根作为植物吸收水分与养分的重要器官,其性状特征在指示植物的生长和分布等方面意义重大.该研究以弓杠岭2500～3300 m海拔地的云杉(Picea asperata)细根为研究对象,采用根序分级法对云杉1～5级根序的生物量及细根形态(平均直径、比根长、根长密度、比表面积)进行测定,以明确...     

Size and morphology of root systems of perennial grasses from contrasting habitats as affected by nitrogen supply

This paper discusses interspecific differences and phenotypic responses to nitrogen supply in various root parameters of five perennial grasses from contrasting habitats. The following root parameters were studied: root:shoot ratio, specific root length, specific root area, mean root diameter, frequency of fine roots, and the length and density of root hairs. Significant between-species variation was found in all of these features. Species from fertile sites had higher root:shoot ratios at high nitrogen supply than species from infertile habitats. All species growing at low nitrogen supply showed a significant increase in root:shoot ratio. Specific root length, specific root area, mean root diameter and frequency of fine roots were not affected significantly by nitrogen supply. Species from infertile sites responded to low nitrogen supply by a significant increase in root hair length and root hair density.     

Dry matter partitioning and root length/leaf area ratios in herbaceous perennial plants with diverse altitudinal distribution

Summary Partitioning patterns in 22 exclusively low and 27 exclusively high altitude perennial herbaceous species were examined in order to test the hypothesis that plants of high altitudes allocate more dry matter to below-ground parts and in particular to storage organs, than typical low altitude plants. Our results raise some doubts about the general validity of this hypothesis. The mean fractions of total dry matter allocated to green leaves (22±2% s.e. at low and 24±2% at high altitude) and special storage organs (28±4% at both altitudes) do not differ significantly among sites. The mean relative portions of total dry matter allocated to above-ground plant parts amount to 57±3% at low and 42±3% at high elevation (P=0.002) and differ less than often assumed. The greater below-ground fraction at high altitude results from reduced stem and proportionally increased fine root compartments. At high altitude specific root length is increased by 50% and mean individual rooting density is tripled. Fine root length per unit leaf area is 4.5 times greater (P<0.001). However, interspecific variation in all these quantities is considerable and species with quite contrasting partitioning patterns coexist at both elevations. This suggests that the success of perennial herbaceous plants at high elevations does not necessarily depend on a large below ground biomass fraction. The increased fine root length at high altitude may substitute for reduced mycorrhizal infection. Figure 1 provides a graphical summary.     

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.     

Root and leaf functional trait relations in Poaceae species: implications of differing resource-acquisition strategies

Aims Root systems play an essential role in grassland functioning in both acquisition and storage of resources. Nevertheless, root functional traits have not received as much attention as those measured on above-ground organs, and little is known about their relations. Our objectives were to test whether morphological and root system traits allowed identification of grass species' functional strategies and to determine whether a relation exists between above- and below-ground traits.Methods Functional traits of root tissues (specific root length, diameter, tissue density and nitrogen concentration), whole root systems (root mass, root length density, root mass percentage below a depth of 20cm and fine root %) and two major leaf traits (specific leaf area and leaf dry matter content) were determined under field conditions and their relations were analysed in eleven perennial temperate Poaceae species.Important findings Canonical correspondence analysis along Axis 1 revealed a gradient of species, from those with deep, dense and coarse root systems with a large root mass to those with shallow root systems, thin roots and high specific root length; this suggests strong correlations among root traits. Correlations between specific root length and specific leaf area reveal two groups of species, which probably indicates different drought-tolerance capacities. Root trait syndromes enable ranking grasses along a gradient from conservative-strategy species (from stressful habitats), which display a deep and coarse root system, to acquisitive species (from rich and moist meadows), which display a shallow and thin root system. Although both types display similar above-ground strategies, drought-tolerant species have lower specific root lengths than drought-sensitive species, revealing more conservative root strategies.     

The effect of neighbors on the root system of the desert annualEremobium aegyptiacum

Intra-specific competition with a primary focus on root competition between plants living in an area with low resource levels, was studied using the natural monotypic population of a desert annual plantEremobium aegyptiacum (Cruciferae). We tested the effect of neighboring distance on shoot and root biomass, and such root parameters as root length, diameter of root neck, number of first order lateral roots and number of lateral roots per unit of main root length. Our results indicate a strong negative relationship between neighboring plant density and such plant parameters as shoot and root biomass, and root neck diameter. The number of first order lateral roots and the number of lateral roots per unit of main root length were negatively related to the distance between neighboring plants. Contrary to predictions, there was no influence of competition on node distribution: neither root overlap nor root avoidance was observed in pairs of adjacent plants.