Prediction of tree species composition in fine root mixed samples using near-infrared reflectance spectroscopy

Fine roots ( ≤ 2 mm diameter) are of great value when investigating belowground interactions among different plant species and soil nutrient cycling in forest ecosystems. However, fine root separation and species identification are labor-intensive and time-consuming processes. This study aimed to evaluate the aptitude of near-infrared reflectance spectroscopy (NIRS) in predicting tree species composition in fine root mixed samples. The coniferous species Cunninghamia lanceolata and Pinus massoniana, the deciduous species Alniphyllum fortunei and Liquidambar formosana, and the evergreen broadleaved species Cyclobalanopsis glauca represent the five subtropical tree species selected for this investigation. To obtain near-infrared reflectance spectral data, 20 samples taken in the field and 70 artificially mixed samples of the five species were produced after root samples were oven-dried and ground. Calibration was performed with partial least squares regression and leave-one-out cross-validation. Root mass proportions of the mixed samples showed good predictive capacity for C. lanceolata, P. massoniana, and C. glauca with low root mean square error of prediction ( < 6.82%) and high determination coefficients (R²>0.944). Predictions for A. fortunei and L. formosana were acceptable with R²>0.819. NIRS shows potential in predicting tree species composition with suitable accuracy.     

Quantifying species composition in root mixtures using two methods: near-infrared reflectance spectroscopy and plant wax markers

Understanding of plant interactions is greatly limited by our ability to identify and quantify roots belonging to different species. We proposed and compared two methods for estimating the root biomass proportion of each species in artificial mixtures: near-infrared reflectance spectroscopy (NIRS) and plant wax markers. Two sets of artificial root mixtures composed of two or three herbaceous species were prepared. The proportion of root material of each species in mixtures was estimated from NIRS spectral data (i) and the concentration patterns of n-alkanes (ii), n-alcohols (iii), and n-alkanes +n-alcohols combined (iv). For each data set, calibration equations were developed using multivariate statistical models. The botanical composition of root mixtures was predicted well for all the species considered. The accuracy varied slightly among methods: alkanes < alcohols = alkanes + alcohols < NIRS. Correlation coefficients between predicted and actual root proportions ranged from 0.89 to 0.99 for alkanes + alcohols predictions and from 0.97 to 0.99 for NIRS predictions. These two methods provide promising potential for understanding allocation patterns and competitive interactions.     

Are fine roots of both shrubs and perennial grasses able to occupy the upper soil layer? A case study in the arid Patagonian Monte with non-seasonal precipitation

We tested whether both shrubs and grasses are able to develop similar active fine-root systems in the upper soil layer of the arid Patagonian Monte ecosystem with non-seasonal precipitation. We selected in the field shrub patches consisting of one isolated modal plant of the dominant shrub Larrea divaricata Cav., grass patches formed by one or more bunches of the dominant grass Stipa tenuis Phil. (15 cm diameter), and mixed patches consisting of one individual of L. divaricata with bunches of S. tenuis under its canopy. We assessed the biomass, regrowth, and activity of fine roots (diameter <1.4 mm) of each species in the upper soil (50 cm depth) of each patch type at 3-month intervals. We also measured the N concentration in fine roots to estimate the relative contribution of each species to fine-root biomass of mixed patches. We injected Li⁺ in the soil as a chemical tracer to detect fine-root activity of each species in the upper soil. Fine-root biomass was higher in mixed patches than in grass patches while fine-root biomass in shrub patches did not differ from the two former. We did not find differences in fine-root regrowth among patch types. Li⁺ injection provided evidence of active fine roots of both species in the upper soil when it was wet. N concentration in fine roots suggested the prevalence of fine roots of L. divaricata in the upper soil of mixed patches. Our results support evidence of the ability of fine roots of both the shrub and the grass species to occupy the upper soil. These findings did not support the two-layer model (H Walter, Ecology of tropical and subtropical vegetation, Oliver and Boyd, Edinburgh, 1971) and provide evidence of this model would be less applicable to arid ecosystems with non-seasonal precipitation. Further, our results highlighted some issues deserving more research such as the outcome of belowground competition between neighboring plants of both contrasting life forms, the eventual limited fine-root carrying capacity of the upper soil, and differences in fine-root lifespan between species of both contrasting life form.     

The effect of tree species diversity on fine-root production in a young temperate forest

The phenomenon of overyielding in species-diverse plant communities is mainly attributed to complementary resource use. Vertical niche differentiation belowground might be one potential mechanism for such complementarity. However, most studies that have analysed the diversity/productivity relationship and belowground niche differentiation have done so for fully occupied sites, not very young tree communities that are in the process of occupying belowground space. Here we used a 5–6 year old forest diversity experiment to analyse how fine-root (<2 mm) production in ingrowth cores (0–30 cm) was influenced by tree species identity, as well as the species diversity and richness of tree neighbourhoods. Fine-root production during the first growing season after the installation of ingrowth cores increased slightly with tree species diversity, and four-species combinations produced on average 94.8% more fine-root biomass than monocultures. During the second growing season, fine-root mortality increased with tree species diversity, indicating an increased fine-root turnover in species-rich communities. The initial overyielding was attributable to the response to mixing by the dominant species, Pseudotsuga menziesii and Picea abies, which produced more fine roots in mixtures than could be expected from monocultures. In species-rich neighbourhoods, P. abies allocated more fine roots to the upper soil layer (0–15 cm), whereas P. menziesii produced more fine roots in the deeper layer (15–30 cm) than in species-poor neighbourhoods. Our results indicate that, although there may be no lasting overyielding in the fine-root production of species-diverse tree communities, increasing species diversity can lead to substantial changes in the production, vertical distribution, and turnover of fine roots of individual species.     

Fine-root morphological and growth traits in a Turkey-oak stand in relation to seasonal changes in soil moisture in the Southern Apennines, Italy

We investigated the effects of seasonal changes in soil moisture on the morphological and growth traits of fine roots (<2?mm in diameter) in a mature Turkey-oak stand (Quercus cerris L.) in the Southern Apennines of Italy. Root samples (diameter:?<0.5, 0.5?C1.0, 1.0?C1.5, and 1.5?C2.0?mm) were collected with the Auger method. Mean annual fine-root mass and length on site was 443?g?m^?2 (oak fine roots 321?g?m^?2; other species 122?g?m^?2) and 3.18?km?m^?2 (oak fine roots 1.14?km?m^?2; other species 2.04?km?m^?2), respectively. Mean specific root length was 8.3?m?g^?1. All fine-root traits displayed a complex pattern that was significantly related to season. In the four diameter classes, both fine-root biomass and length peaked in summer when soil water content was the lowest and air temperature the highest of the season. Moreover, both fine-root biomass and length were inversely related with soil moisture (p?<?0.001). The finest roots (<0.5?mm in diameter) constituted an important fraction of total fine-root length (79?%), but only 21?% of biomass. Only in this root class, consequent to change in mean diameter, specific root length peaked when soil water content was lowest showing an inverse relationship (p?<?0.001). Furthermore, fine-root production and turnover decreased with increasing root diameter. These results suggest that changes in root length per unit mass, and pulses in root growth to exploit transient periods of low soil water content may enable trees to increase nutrient and water uptake under seasonal drought conditions.     

The age of fine-root carbon in three forests of the eastern United States measured by radiocarbon

Using a new approach involving one-time measurements of radiocarbon (¹⁴C) in fine (<2 mm diameter) root tissues we have directly measured the mean age of fine-root carbon. We find that the carbon making up the standing stock of fine roots in deciduous and coniferous forests of the eastern United States has a mean age of 3-18 years for live fine roots, 10-18 years for dead fine roots, and 3-18 years for mixed live+dead fine roots. These ¹⁴C-derived mean ages represent the time C was stored in the plant before being allocated for root growth, plus the average lifespan (for live roots), plus the average time for the root to decompose (for dead roots and mixtures). Comparison of the ¹⁴C content of roots known to have grown within 1 year with the ¹⁴C of atmospheric CO₂ for the same period shows that root tissues are derived from recently fixed carbon, and the storage time prior to allocation is <2 years and likely <1 year. Fine-root mean ages tend to increase with depth in the soil. Live roots in the organic horizons are made of C fixed 3-8 years ago compared with 11-18 years in the mineral B horizons. The mean age of C in roots increases with root diameter and also is related to branching order. Our results differ dramatically from previous estimates of fine-root mean ages made using mass balance approaches and root-viewing cameras, which generally report life spans (mean ages for live roots) of a few months to 1-2 years. Each method for estimating fine-root dynamics, including this new radiocarbon method, has biases. Root-viewing approaches tend to emphasize more rapidly cycling roots, while radiocarbon ages tend to reflect those components that persist longest in the soil. Our ¹⁴C-derived estimates of long mean ages can be reconciled with faster estimates only if fine-root populations have varying rates of root mortality and decomposition. Our results indicate that a standard definition of fine roots, as those with diameters of <2 mm, is inadequate to determine the most dynamic portion of the root population. Recognition of the variability in fine-root dynamics is necessary to obtain better estimates of belowground C inputs.     

Effect of species, root branching order and season on the root traits of 13 perennial grass species

Aims

Inter-specific comparisons of plant traits may vary depending on intra-specific variation. Here we examine the impact of root branching order and season on key functional root traits for grass species. We also compare root traits among co-existing grass species as a step towards defining root trait syndromes.

Methods

Monocultures of 13 grass species, grown under field conditions and subjected to intensive management, were used to record root trait values for coarse roots (1st order, >0.3?mm), fine roots (2nd and 3rd orders, <0.2?mm) and mixed root samples over three growing seasons.

Results

Branching order and species had a significant effect on root trait values, whereas season showed a marginal effect. The diameter of coarse roots was more variable than that of fine roots and, as expected, coarse roots had higher tissue density and lower specific root length values than fine roots. Principal component analysis run on eight root traits provided evidence for two trait syndromes related to resource acquisition and conservation strategies across grass species.

Conclusions

Our data show that root branching order is the main determinant of root trait variation among species. This highlights the necessity to include the proportion of fine vs coarse roots when measuring traits of mixed root samples.     

Rooting strategy of naturally regenerated beech in Silver birch and Scots pine woodlands

This study investigated the belowground development and strategy of late-successional European beech (Fagus sylvatica L.) in ageing natural Scots pine (Pinus sylvestris L.) and Silver birch (Betula pendula Roth.) woodlands in a French volcanic mid-elevation area. For this purpose root biomass, root profile and fine-root architecture of competitor trees were examined in 53 mixed pine–beech and 42 birch–beech woodlands along a stand maturation gradient, using the root auger technique (0–75-cm). The total beech fine-root biomass highly correlated with aerial dimensions such as stem height and girth, whereas it moderately correlated with its age, thus indicating the effects of competition. Basic stand biometric data such as stand density and basal area had no significant effect on beech root biomass. Conversely, competition indices taking into account the vertical dimensions of competitor trees were efficient, probably due to redundancy with beech height. At similar age and height, beeches under birch had a greater belowground development than beeches under pine. Each species exhibited specific rooting pattern and plasticity of fine-root architecture along the gradients of stand maturation and competition. Beech had a heart-shaped rooting habit in both mixings, which strongly increased along stand maturation. Its fine-root system adopted a foraging strategy to respond to increasing stand competition. The Scots pine fine-root system was plate-like and showed a low morphological plasticity, thus presumably a conservative strategy. Silver birch exhibited a high biomass and a foraging capacity in the topsoil but a loose root system in the subsoil. The coexistence of pine and beech roots in the upper soil presumably leads to a high belowground competition. Beech root system becomes predominant throughout the soil profile and it adopts an efficient foraging strategy, but at the expense of its belowground development. Conversely, the niche partitioning strategy between beech and birch may explain why beech develops strongly belowground in spite of the fact that birch has a dense rooting and a competitive fine-root architecture. As a consequence, beech mid-term regeneration and development may be facilitated under birch as compared with pine.     

岷江上游4个栽培树种细根功能性状垂直分布的差异性

细根(直径≤2 mm)功能性状及垂直分布格局是反映植物对土壤资源吸收策略和影响森林地下生态过程的关键。本研究以岷江上游4个人工林树种连香树(Cercidiphyllum japonicum)、白桦(Betula platyphylla)、华山松(Pinus armandii)和油松(P.tabuliformis)为对象,调查不同海拔树木细根功能性状及其在不同土层间的垂直分布格局,并分析细根功能性状分布与构型之间的相关关系。结果表明:阔叶树种比针叶树种有更大的根长密度、生物量、比根长和比表面积,而直径反之;4个树种细根集中在0~20 cm土层,根长密度和生物量在较高海拔地段均显著大于较低海拔,且均随土壤深度增加而减少,但比根长、比表面积和直径无显著的海拔差异,随土层加深也无明显的垂直变化规律;针阔树种间的细根构型差异显著,但不受海拔差异的影响,阔叶树的细根分支强度与一级根数量显著大于针叶树种;一级根数和根尖密度与比根长以及分根比与根长密度和生物量均呈显著正相关,而分叉与几个细根功能参数均呈负相关;随着土层深度增加,细根总生长量明显减少,但细根资源利用效率和策略不变;5个细根功能性状的垂直分布格局差异明显,且受树种特性影响显著,细根分支构型对其垂直分布及资源获取策略具有显著影响。     

On the significance of belowground overyielding in temperate mixed forests: separating species identity and species diversity effects

Complementary soil exploration by the root systems of coexisting tree species has been hypothesised to result in a higher root biomass of mixed forests than of monocultures but the existing evidence for a belowground diversity effect in forests is scarce and not conclusive. In a species‐rich temperate broad‐leaved forest, we analysed the fine root biomass (roots ≤ 2 mm) and necromass in 100 plots differing in tree species diversity (one to three species) and species composition (all possible combinations of five species of the genera Acer, Carpinus, Fagus, Fraxinus and Tilia) which allowed us to separate possible species diversity and species identity effects on fine root biomass. We found no evidence of a positive diversity effect on standing fine root biomass and thus of overyielding in terms of root biomass. Root necromass decreased with increasing species diversity at marginal significance. Various lines of evidence indicate significant species identity effects on fine root biomass (10–20% higher fine root biomass in plots with presence of maple and beech than in plots with hornbeam; 100% higher fine root biomass in monospecific beech and ash plots than in hornbeam plots; differences significant). Ash fine roots tended to be over‐represented in the 2‐ and 3‐species mixed plots compared to monospecific ash plots pointing at apparent belowground competitive superiority of Fraxinus in this mixed forest. Our results indicate that belowground overyielding and spatial complementarity of root systems may be the exception rather than the rule in temperate mixed forests.     

Root Growth and Recovery in Temperate Broad-Leaved Forest Stands Differing in Tree Species Diversity

In contrast to studies on aboveground processes, the effect of species diversity on belowground productivity and fine-root regrowth after disturbance is still poorly studied in forests. In 12 old-growth broad-leaved forest stands, we tested the hypotheses that (i) the productivity and recovery rate (regrowth per standing biomass) of the fine-root system (root diameter < 2 mm) increase with increasing tree species diversity, and that (ii) the seasonality of fine-root biomass and necromass is more pronounced in pure than in tree species-rich stands as a consequence of non-synchronous root biomass peaks of the different species. We investigated stands with 1, 3, and 5 dominant tree species growing under similar soil and climate conditions for changes in fine-root biomass and necromass during a 12-month period and estimated fine-root productivity with two independent approaches (ingrowth cores, sequential coring). According to the analysis of 360 ingrowth cores, fine-root growth into the root-free soil increased with tree species diversity from 72 g m⁻² y⁻¹ in the monospecific plots to 166 g m⁻² y⁻¹ in the 5-species plots, indicating an enhanced recovery rate of the root system after soil disturbance with increasing species diversity (0.26, 0.34, and 0.51 y⁻¹ in 1-, 3-, and 5-species plots, respectively). Fine-root productivity as approximated by the sequential coring data also indicated a roughly threefold increase from the monospecific to the 5-species stand. We found no indication of a more pronounced seasonality of fine-root mass in species-poor as compared to species-rich stands. We conclude that species identification on the fine root level, as conducted here, may open new perspectives on tree species effects on root system dynamics. Our study produced first evidence in support of the hypothesis that the fine-root systems of more diverse forest stands are more productive and recover more rapidly after soil disturbance than that of species-poor forests.     

Further evidence for slow decomposition of very fine roots using two methods: litterbags and intact cores

Background and aims

Root decomposition studies have rarely considered the heterogeneity within a fine-root system. Here, we investigated fine root (< 0.5 and 0.5–2 mm in diameter) decomposition and accompanying nutrient dynamics of two temperate tree species—Betula costata Trautv and Pinus koraiensis Sieb. et Zucc.

Methods

Both litterbag and intact-core techniques were used to examine decomposition dynamic and nutrient release of the two size class roots over a 498-day period. Moreover, we examined differences between the two approaches.

Results

The very fine roots (< 0.5 mm) with an initially lower C:N ratio, decomposed more slowly than 0.5–2 mm roots of both tree species. The differences in mass loss between size classes were smaller when using the intact-core technique compared with litterbag technique. In contrast to root biomass loss, net N release was much higher in the fine roots (< 0.5 mm). All fine roots initially released N (0–75 days), but immobilized N to varying extent in the following days (75–498 days) during decomposition.

Conclusions

Our results suggest that the slow decomposition rate of very fine roots (< 0.5 mm) may be determined by their high concentration of acid-unhydrolyzable structural components. Additionally, the heterogeneity within a bulk fine-root system could lead to differences in their contribution to soil in terms of carbon and nitrogen dynamics.     

长白山原始阔叶红松（Pinus koraiensis）林及其次生林细根生物量与垂直分布特征

以我国东北长白山自然保护区内同一海拔梯度的原始阔叶红松林及其次生林——白桦山杨成熟林和白桦山杨幼林为对象,采用土钻取样法对不同演替阶段细根生物量的变化、细根垂直分布规律及其影响因子进行系统地研究。研究结果表明,在原始阔叶红松林的正向演替过程中,林地细根的总生物量逐渐增加,其中主要乔木细根的生物量逐渐增加,而灌木和草本细根的生物量则逐渐降低。在演替过程中,细根的垂直分布逐渐加深。在长白山地区,3块林地中细根生物量的组成分布受林分植被组成的影响;细根的垂直分布与土壤容重、水分含量以及不同土层中C、N含量存在一定的相关性,但与土壤温度则不存在相关关系。     

基于土芯法的亚热带常绿阔叶林细根空间变异与取样数量估计

树木细根具有高度空间异质性,确定合理的细根取样策略是林木细根研究的前提。通过在福建省三明米槠天然常绿阔叶林内随机钻取96个土芯,分析细根生物量和形态特征的空间变异特征,并估计各指标所需的取样数量。结果表明:(1)随着径级增加,细根各指标变异系数增大,相应的取样数量增加;(2)随着土壤深度增加,单位面积细根生物量变异程度和相应的取样数量均增加。在置信水平为95%、精度为80%的条件下,直径为0-1 mm和1-2 mm的细根,分别采集16和42个样品可以满足测定单位面积细根生物量,采集17和31个样品可以满足测定单位面积细根长度,采集25和33个样品可以满足测定单位面积细根表面积。Shapiro-Wilk检验表明,除表层土壤0-1 mm细根单位面积生物量符合正态分布外,其余细根生物量和形态指标数据均不符合正态分布。研究结果为亚热带常绿阔叶林细根的合理取样提供了科学依据。     

Contrasting fine-root production,survival and soil CO2 efflux in pine and poplar plantations

Tree root activity, including fine-root production, turnover and metabolic activity are significant components of forest productivity and nutrient cycling. Differences in root activity among forest types are not well known. A 3-year study was undertaken in red pine (Pinus resinosa Ait.) and hybrid poplar (Populus tristis X P. balsamifera cv `Tristis no. 1') plantations to compare belowground root dynamics. We measured fine-root production, mortality and standing crop, as well as soil CO₂ efflux. Pine fine-root production was only 2.9% of that of poplar during three years; 85 pine roots were observed in minirhizotron tubes compared with 4088 poplar roots. Live-root density oscillated seasonally for both species with late winter minimum and autumn maximum. Poplar reached constant maximum live-root length within the first growing season, but pine continued to increase observed fine-root length for three growing seasons. Within the first 100 days following initial appearance, 22% of the pine roots disappeared and 38% of the poplar roots disappeared. Median fine-root longevity of pine was 291 days compared with 149 days for poplar roots. Fine-root longevity increased with depth in the soil, and was greater for roots with initial diameter >0.5 mm. The probability of poplar root death from late February to May was more than three times that in any other season, regardless of root age. Despite the greater poplar root production and live-root length, fine-root biomass and soil CO₂ efflux was greater in pine. Greater metabolic activity in the pine stand may be due to greater fine-root biomass or greater heterotrophic respiration. This revised version was published online in June 2006 with corrections to the Cover Date.     

Root biomass,root distribution and the fine-root growth dynamics of Quercus coccifera L. in the garrigue of southern France

Quercus coccifera L., the characteristic scrub oak of the garrigue, covers more than 100,000 ha in southern France alone. Precipitation in this area averages 900 mm/year and summer rains are not rare. A total belowground biomass of 7.2 kg/m², including rhizomes and lignotubers, was harvested. Roots were concentrated in the uppermost 50 cm of the soil. It was hypothesized that low winter temperatures inhibit active fine-root growth. This hypothesis was tested by means of fine-root extractions of soil samples from 0–50 cm depth from November 1987 to June 1988. Although the fine-root analysis could not be extended into late summer and fall, the data supported the hypothesis. Ratios of live/dead fine roots reached their minimum at 0.2–0.3 from December to April. They increased to 1.0–1.2 during late spring and early summer. Initiation of fine-root growth in early April was synchronous with bud break. Starch contents of roots, rhizomes, and lignotubers fluctuated from 4.3% in January to 8.3% in April. The starch stored in belowground organs of Q. coccifera in a closed canopy stand amounted to about 500 g/m² in April. This amount declined to 400 g with bud burst and fine-root growth initiation.     

The dependence of root system properties on root system biomass of 10 North American grassland species

Dependence of the properties of root systems on the size of the root system may alter conclusions about differences in plant growth in different environments and among species. To determine whether important root system properties changed as root systems aged and accumulated biomass, we measured three important properties of fine roots (tissue density, diameter, and C:N) and three biomass ratios (root:shoot, fine:coarse, and shallow:deep) of monocultures of 10 North American grassland species five times during their second and third years of growth. With increasing belowground biomass, root tissue density increased and diameter decreased. This may reflect cortical loss associated with the aging of roots. For non-legumes, fine root C:N decreased with increasing root biomass, associated with decreases in soil solution NO^{3 –} concentrations. No changes in fine root C:N were detected with increasing belowground biomass for the two legumes we studied. Among all 10 species, there were generally no changes in the relative amounts of biomass in coarse and fine roots, root:shoot, or the depth placement of fine roots in the soil profile as belowground biomass increased. Though further research is needed to separate the influence of root system size, age of the roots, and changes in nutrient availability, these factors will need to be considered when comparing root functional traits among species and treatments.     

Fine-root vitality in a Norway spruce stand subjected to various nutrient supplies

The vitality of fine roots in a Norway spruce stand subjected to application of ammonium sulphate (NS), wood ash (A) and nitrogen-free fertilizer (V) respectively, was investigated using i) vitality classification of fine roots based on morphological characteristics and ii) the triphenyl tetrazolium chloride (TTC) method of estimating dehydrogenase activity.Although the NS-treated areas showed a 30% increase in above-ground production in response to the NS-application, the vitality of the fine-root system in the humus layer appeared to be in a state of deterioration, as indicated by a decrease in fine-root biomass, an increased amount of dead fine (0–1 mm) and small (1–2 mm) roots, a decreased specific root length (SRL = fine root length/fine root dry weight) and an increased dehydrogenase activity. The impact of the the A and V treatments was reflected in a decrease in fine-root biomass and an increase in SRL. The results make it clear that in order to study the vitality of forest trees, both fine-root studies and studies of above-ground tree parts are necessary.     

Anatomical characteristics of individual roots within the fine-root architecture of Chamaecyparis obtusa (Sieb. & Zucc.) in organic and mineral soil layers

Nutrient availability and temporal variation of physical stress are usually higher in organic soil layers than in mineral soils. Individual roots within the fine-root system adjust anatomical, morphological, and turnover characteristics to soil conditions, for example nutrient availability and physical stresses. We investigated anatomical traits, including cork formation and passage and protoxylem cell numbers, in cross-sections of individual fine roots of the conifer Chamaecyparis obtusa (Siebold & Zucc.) growing under different soil conditions. The fine-root systems in different soil layers were compared by sampling ingrowth cores buried for 1 year and filled with organic and mineral soil substrates. The number of exodermal passage cells was lower in roots from organic soils than in those from mineral soils, suggesting that apical roots tend to be more stress-tolerant in the organic layer than in mineral soils. In contrast, both root tip and specific root tip density were higher in roots from organic soils than in those from mineral soil layers. The proportion of roots with two strands of protoxylem (diarch) was greater in organic (90%) than in mineral (25%) soils. Thus, although the absorptivity of individual apical roots decreases in organic layers, the absorptivity of the entire fine-root system of C. obtusa may be increased as a result of the increase in apical root density and the proportion of ephemeral roots. We found that the fine-root system had simultaneous plasticity in density, anatomy, and architecture in response to complex soil conditions.     

Is there coordination of leaf and fine root traits at local scales? A test in temperate forest swamps

Examining the coordination of leaf and fine root traits not only aids a better understanding of plant ecological strategies from a whole‐plant perspective, but also helps improve the prediction of belowground properties from aboveground traits. The relationships between leaf and fine root traits have been extensively explored at global and regional scales, but remain unclear at local scales. Here, we measured six pairs of analogous leaf and fine root traits related to resource economy and organ size for coexisting dominant and subordinate vascular plants at three successional stages of temperate forest swamps in Lingfeng National Nature Reserve in the Greater Hinggan Mountains, NE China. Leaf and fine root traits related to resource acquisition (e.g., specific leaf area [SLA], leaf N, leaf P, root water content, and root P) decreased with succession. Overall, we found strong linear relationships between leaf dry matter content (LDMC) and root water content, and between leaf and root C, N, and P concentrations, but only weak correlations were observed between leaf area and root diameter, and between SLA and specific root length (SRL). The strong relationships between LDMC and root water content and between leaf and root C, N, and P held at the early and late stages, but disappeared at the middle stage. Besides, C and P of leaves were significantly correlated with those of roots for woody plants, while strong linkages existed between LDMC and root water content and between leaf N and root N for herbaceous species. These results provided evidence for the existence of strong coordination between leaf and root traits at the local scale. Meanwhile, the leaf–root trait relationships could be modulated by successional stage and growth form, indicating the complexity of coordination of aboveground and belowground traits at the local scale.