Application of Picea wilsonii roots to determine erosion rates in eastern Qilian Mountains, Northwest China

As a consequence of global change and human activities, processes of soil erosion are expected to increase in forested areas, resulting in exposed roots. Dendrogeomorphic research was conducted by analyzing exposed roots of Picea wilsonii subjected to continuous denudation along a main road in Tulugou National Forest Reserve, eastern Qilian Mountains, to reconstruct the local soil erosion dynamics. We determined the start of the exposure by examining the shifts in the ring-growth patterns from concentric to eccentric and by analyzing the detailed changes of wood anatomical features of exposed roots. We also find that the width of growth ring, the percentage of latewood and the average cell size of earlywood tracheids are all remarkable signs for soil lowering. According to the analysis of ANOVA (Fisher’s least significant difference method), the reduction of the cell size of earlywood tracheids is verified to be the key indicator for dating the first year of exposure and occurs prior to the other two indicators. Using 40 roots from 23 trees spread along the road, it has been found that erosion rates vary between 3.3 and 13.5 mm/year with an average value about 5.3 ± 2.1 mm/year. The intensity and occurrence of soil erosion may be influenced by the increase of human activities.     

Exposed roots as indicators of geomorphic processes: A case-study from Polylepis mountain woodlands of Central Argentina

Soil erosion is a serious problem of land degradation in many parts of the world, and particularly in mountain rangelands. To understand this process it is necessary to develop methods to assess soil erosion rate in a quick, economic and accurate manner. Based on the analysis of exposed Polylepis australis roots, we tested a dendrogeomorphological method for determining soil loss rate in rills and gullies. Few studies considered non-coniferous tree rings in soil erosion analysis and we used, for the first time, an experimental procedure of root exposure and provided a comparison with roots exposed by gully erosion. Our main results showed that as a consequence of soil erosion, exposed roots changed from root-like to a more stem-like wood anatomical structure. The percentage of vessel area per tree-ring area decreases by an average of 22% to 43% during the first and second year after exposure, respectively. Moreover, and during the same time interval, the mean vessel area decreased 32% and 65%, and the number of vessels increased 7% and 48%, respectively. Scars formed at the upper side of the exposed roots are coincident with changes in wood anatomy, and both evidences may be applied to reconstruct an erosion process. This study confirms that the wood anatomy analysis of partially exposed roots can be used to determine the year in which roots are exposed and provides a useful tool to monitor soil erosion rates with a high accuracy.     

Root site occupancy modelling of young New Zealand native plants: implications for soil reinforcement

Plants are widely used in soil conservation to control and prevent erosion on hillslopes and on riverbanks. Previous research has shown the mechanical root reinforcement on soil stability can be considerable. However, land and forest managers still require information and simple tools to enable them to determine how and when a species becomes effective in terms of soil stabilisation. This paper uses root length data from a trial of young New Zealand trees and shrubs to develop a simple model to account for the spatial occupancy of a planting site by roots, and by implication their potential strength contribution to soil reinforcement. It is developed by calculating root surface area in contact with the soil to obtain an effective radius of the root spread about the stem. The approach generates a set of coefficients that are unique to a species for a given site which can then be used in the generalised model to predict root site occupancy, which is taken as a proxy for when soil reinforcement is attained. This information can then be used to assess effectiveness of different species mixes in planting plans.     

Comparative analysis of annual rings of perennial forbs in the Loess Plateau,China

Although recent studies have demonstrated that annual growth rings are present among perennial forbs species at high northern latitudes, little is known about whether there are demarcated growth rings of perennial forbs in the Loess Plateau of China where plant growth is strongly limited by dry climate conditions and severe soil erosion. In this study, we collected the main roots of 11 perennial forbs species along the precipitation gradient in the Loess Plateau, and analyzed the growth rings in the secondary root xylem. We found that ten species showed distinct annual growth rings, and the anatomical patterns, including vessel size and density, varied considerably among different families. Our results suggest, for forbs species in the Loess Plateau, that vessel diameter in the root xylem was strongly correlated with growth rate of the forb’s roots. Ring widths of the forbs showed a significant declining trend, reflecting the deteriorating signal of growth condition with age. In comparison to other families, forb species of Fabaceae usually have the evidently larger vessels that link directly to higher hydraulic capacity and growth rate. In terms of annual ring width patterns, this study provides an applicable approach to detecting effects of limited climatic conditions and life history strategies on herbaceous vegetation in the Loess Plateau.     

Are radial changes in vascular anatomy mechanically induced or an ageing process? Evidence from observations on buttressed tree root systems

To investigate the effect of changes in mechanical loading conditions on radial anatomical patterns, we here compare the trunk with the roots in two locations of three species of buttressed trees. The proximal part of the buttress roots is highly mechanically loaded throughout juvenile growth whereas the distal part of the buttresses is though to be mechanically unimportant at formation but become progressively more mechanically loaded during growth. We measured the frequency and diameter of the vessels and the vessel area fraction, and from this calculated the specific conductivity of tissue samples of the core-, intermediate- and outer wood. As in previous studies there was an increase in vessel size, vessel area fraction and specific conductivity from the pith to the bark in the trunk. In the proximal part of the buttress roots, however, there was no increase in vessel size and conductivity from core wood and out in agreement with the high mechanical loading found here throughout growth. There was instead a decrease in vessel size, vessel area fraction and specific conductivity from core- to outer wood in the distal part of the buttress roots in accordance with the increase in mechanical loading. Hence, it appears that the radial anatomical patterns are not a passive function of cambial ageing but may be modified in response to local mechanical loading.     

Wood anatomy and vessel characteristics of spiny monkey orange (Strychnos spinosa) in Benin (West Africa)

The response of plant species to varying climate conditions in tropical Africa remains poorly understood but can be assessed using wood anatomical traits. These traits play an important role for the adaptive capacity of a species to environmental stress, since environmental conditions can modify the proportion, size, and morphology of wood anatomical elements. This study reports quantitative data on vessel characteristics of the diffuse porous angiosperm Spiny monkey orange (Strychnos spinosa Lam.) in Benin in tropical West Africa. The vessel-related anatomical traits varied with high amplitude (coefficient of variation CV ˃ 25%) between different sites located in different climate zones. The variability of the traits is higher within one climatic zone than between climatic zones, and even more pronounced within trees. Consequently, the climatic zones have less influence on the studied features than local site conditions. However, the study showed that S. spinosa individuals that have numerous vessels also have a high lumen fraction and total ring area. On the other hand, individuals presenting a high vessel density also display vessels of smaller size. The correlation between vessel number and total ring area on the one hand, and between vessel size and lumen fraction on the other hand are highly significant and positive. In Benin, S. spinosa wood anatomical traits are likely linked to local site factors rather than to regional climatic factors.     

Angiosperm wood structure: Global patterns in vessel anatomy and their relation to wood density and potential conductivity

Woody stems comprise a large biological carbon fraction and determine water transport between roots and leaves; their structure and function can influence both carbon and hydrological cycles. While angiosperm wood anatomy and density determine hydraulic conductivity and mechanical strength, little is known about interrelations across many species. We compiled a global data set comprising two anatomical traits for 3005 woody angiosperms: mean vessel lumen area (ā) and number per unit area (N). From these, we calculated vessel lumen fraction (F = āN) and size to number ratio (S = ā/N), a new vessel composition index. We examined the extent to which F and S influenced potential sapwood specific stem conductivity (K(S)) and wood density (D; dry mass/fresh volume). F and S varied essentially independently across angiosperms. Variation in K(S) was driven primarily by S, and variation in D was virtually unrelated to F and S. Tissue density outside vessel lumens (D(N)) must predominantly influence D. High S should confer faster K(S) but incur greater freeze-thaw embolism risk. F should also affect K(S), and both F and D(N) should influence mechanical strength, capacitance, and construction costs. Improved theory and quantification are needed to better understand ecological costs and benefits of these three distinct dimensions.     

Changes in vessel anatomy in response to mechanical loading in six species of tropical trees

It is well known that trees adapt their supportive tissues to changes in loading conditions, yet little is known about how the vascular anatomy is modified in this process. We investigated this by comparing more and less mechanically loaded sections in six species of tropical trees with two different rooting morphologies. We measured the strain, vessel size, frequency and area fraction and from this calculated the specific conductivity, then measured the conductivity, modulus of elasticity and yield stress. The smallest vessels and the lowest vessel frequency were found in the parts of the trees subjected to the greatest stresses or strains. The specific conductivity varied up to two orders of magnitude between mechanically loaded and mechanically unimportant parts of the root system. A trade-off between conductivity and stiffness or strength was revealed, which suggests that anatomical alterations occur in response to mechanical strain. By contrast, between-tree comparisons showed that average anatomical features for the whole tree seemed more closely related to their ecological strategy.     

Axial Water Flux Dynamics in Small Diameter Roots of a Fast Growing Tropical Tree

Field measurements of water flux in small diameter roots are important to the study of whole plant water transport systems. Miniature sap flow gauges were used to capture high resolution water flux patterns in small roots of 2 – 5 mm diameter and simultaneously in the canopy branches of a Eucalyptus saligna tree growing in Hawaii. The axial transport flux rates were then correlated with anatomical measurements to describe the internal hydraulic capacity of the tree. The daily patterns of water flux showed a strong coupling between the canopy and root systems and both systems were tightly synchronized with rapid fluctuations in photosynthetic photon flux density, vapour pressure deficit, and wind speed. When flow rates were normalized by the total vessel lumen area, branches had daily totals equivalent to the surface roots. Daily flows of water through surface roots were consistently 30% greater than through deep roots. Results of an experiment where a portion of the canopy was removed showed the decrease in water flux for all roots was in nearly direct proportion to the decrease in leaf area. The root anatomical measurements suggested a high capacity axial root water transport system with roots containing a smaller number of vessels per unit of sapwood area than branches but with vessel diameters twice that of the branches. However, relative conductivity values of roots and branches were similar and comparable to some of the highest values reported. Overall, the results suggested a highly efficient axial water transport system that would help to maintain a favorable plant water status for maximal stomatal opening. This revised version was published online in July 2006 with corrections to the Cover Date.     

黄河三角洲滨海盐碱地11个造林树种细根解剖性状对土壤条件的响应

细根作为植物与土壤连接的重要部位,能够反映植物对生存环境的适应性。以黄河三角洲滨海盐碱地不同立地条件下11个造林树种为对象,基于细根分支等级划分1-4级根序并进行解剖特征测定,分析细根解剖性状对滨海盐碱地不同土壤条件的响应规律。结果表明：（1）不同根序的细根直径存在显著差异,细根直径随根序升高呈增大趋势,而同根序的细根直径在不同树种间表现出显著的种间差异（P < 0.05）。1-2级细根皮层厚度、3-4级细根导管密度在树种间的差异均达显著水平（P < 0.05）。（2）在较为严重盐渍化土壤条件下（立地1）,细根皮层厚度较其他立地显著增大,但细根导管密度较小;在轻度盐碱立地条件下（立地3）,细根导管密度较大;较为严重的盐碱立地具有更为发达的细根直径及维管柱直径。（3）树种1-2级细根解剖结构与土壤环境关系最为密切,其中1级根直径与土壤pH值显著正相关（P < 0.05）,与土壤硝态氮含量呈显著负相关（P < 0.05）。对土壤理化性质与细根解剖性状的冗余分析表明,前两个轴的特征值达0.640和0.196,土壤速效养分含量与轴一（RDA1）呈正相关,低级根解剖性状则与轴二（RDA2）呈显著负相关。低级根解剖结构以及土壤的pH值能解释较多树种的差异性,其中低级根直径与皮层厚度对盐碱环境表现出较强的响应。     

Duration and extension of anatomical changes in wood structure after cambial injury

Cambial injury has been reported to alter wood structure in broad-leaved trees. However, the duration and extension of associated anatomical changes have rarely been analysed thoroughly. A total of 18 young European ash (Fraxinus excelsior L.) trees injured on the stem by a spring flood were sampled with the aim of comparing earlywood vessels and rays formed prior to and after the scarring event. Anatomical and hydraulic parameters were measured in five successive rings over one-quarter of the stem circumference. The results demonstrate that mechanical damage induces a decrease in vessel lumen size (up to 77%) and an increase in vessel number (up to 475%) and ray number (up to 115%). The presence of more earlywood vessels and rays was observed over at least three years after stem scarring. By contrast, abnormally narrow earlywood vessels mainly developed in the first ring formed after the event, increasing the thickness-to-span ratio of vessels by 94% and reducing both xylem relative conductivity and the index for xylem vulnerability to cavitation by 54% and 32%, respectively. These vessels accumulated in radial groups in a 30° sector immediately adjacent to the wound, raising the vessel grouping index by 28%. The wound-induced anatomical changes in wood structure express the functional need of trees to improve xylem hydraulic safety and mechanical strength at the expense of water transport. Xylem hydraulic efficiency was restored in one year, while xylem mechanical reinforcement and resistance to cavitation and decay lasted over several years.     

Strain distribution during anchorage failure of Pinus pinaster Ait. at different ages and tree growth response to wind-induced root movement

Winching tests were carried out on 5- 13- and 17-year-old tap rooted Maritime pine (Pinus pinaster Ait.) in order to determine how the mode of anchorage failure changes throughout the life of a tree. As trees were pulled sideways, strain along the lateral roots was recorded using strain gauges attached to a strain indicator. Measurements of strain in the root system, taken during winching, provide information about root movement when loaded by wind. The mode of mechanical failure appeared to depend on tree age. The youngest trees bent over completely during winching, but did not break due to the plasticity of their trunks. The 13-year-old trees either broke at the base of the tree (due to the presence of grafting scar tissue) or at the base of the tap-root. The oldest trees broke at the base of the tap-root and sounds of roots breaking were also heard. Strain was twice as great in the trunk compared to the roots in the 5- and 13-year-old trees and was three times greater in the compression roots of 17-year–old trees compared to that in the trunk. In one 17-year-old tree, strain was found to increase at a distance of 35 cm in tension roots before decreasing again. Although the mode of failure changed with tree size, anchorage strength increased proportionally with the third power of trunk diameter, therefore another reason why failure differs with tree age must exist. In order to determine if different types of wood were being laid down in the lateral roots in response to wind loading, maturation strains, indicating the existence of mechanical stress in developing wood cells, were measured at different points along the roots. A high correlation was found between maturation strain and strain measured during winching, in roots that lay in the wind direction only. Therefore, trees appear to be able to respond to external loading stress, even at a local level within a root. This revised version was published online in June 2006 with corrections to the Cover Date.     

Assessing root death and root system dynamics in a study of grape canopy pruning

Defining root death in studies of root dynamics is problematic because cell death occurs gradually and the resulting effects on root function are not well understood. In this study, metabolic activity of grape roots of different ages was assessed by excised root respiration and tetrazolium chloride reduction. We investigated changes in metabolic activity and patterns of cell death occurring with root age and changes in root pigmentation. Tetrazolium chloride reduction of roots of different ages was strongly correlated to respiration ( R ² = 0.786). As roots aged, respiration and tetrazolium chloride reduction declined similarly, with minimum metabolic activity reached at six weeks. Tetrazolium chloride reduction indicated that the onset of root browning corresponded to a 77% reduction in metabolic activity ( P < 0.001). Anatomical examination of roots at each pigmentation stage showed that even though some cells in brown roots were still alive, these roots were functionally dead. The effect of using different definitions of root death in relation to root survivorship was determined in a study of 'Concord' grapes with two pruning treatments, using three criteria for root death: browning, blackening or shriveling, and disappearance. There was no effect of vine pruning on root life span when life span was defined as the time from first appearance to the onset of browning. However, if death was judged as the point when roots either became black or shriveled or disappeared, vine pruning decreased root life span by 34% and 40%, respectively ( P < 0.001), and also increased the decay constant for root decomposition by about 45% ( P < 0.001). We conclude that the discrepancy among determinations of root life span assessed with different definitions of death might be partly caused by the latter evaluations of root life span incorporating a portion of root decomposition in definitions of root death.     

半干旱矿区侵蚀营力对黑沙蒿根系生长特性的影响及其自修复

以半干旱矿区典型分布的黑沙蒿为研究对象,通过野外原位不离体试验,模拟侵蚀拉拔破坏对黑沙蒿根系生长特性的影响,并分析其受损自修复能力,旨在揭示植物根系在脆弱生态区中抵御外力侵蚀的生存策略.结果 表明,拉拔破坏形成机械损伤后,根系生长速率和活性均明显降低,持续拉拔对二者的抑制作用显著大于瞬时拉拔,重度损伤产生的负反馈显著大...     

Effects of experimental stem burial on radial growth and wood anatomy of pedunculate oak

In dendrogeomorphology, abrupt changes in wood anatomy are frequently used to date the exact year of burial and exposure events. However, few studies have addressed the precision and underlying mechanisms of these changes. In a field experiment, performed in a drift-sand area in the Netherlands, we buried the stems of mature pedunculate oak trees (Quercus robur L.) up to a height of 50 cm and analysed the responses in ring width and earlywood-vessel characteristics, while monitoring the course of temperature above and below the soil surface.After 3 years of stem burial, we found no significant differences in ring width and earlywood-vessel characteristics between control and buried trees both above and below the burial level. Burial however strongly reduced temperature amplitude and the occurrence of sub-zero temperatures around the buried stems. All buried trees formed epitropic roots that grew upward into the new sediment layer, but no adventitious roots were formed on the buried stems. Irrespective of the burial treatments, we found that the mean ring width was largest at the original stem base and lowest at breast height. In contrast, vessel sizes were significantly larger at breast height compared with the stem base. Differences in vessel density barely differed between years and heights.In our field experiment on mature pedunculate oak trees, the burial of stems by 50 cm of drift sand did not induce any local growth suppression or detectable changes in wood anatomy. As wood-anatomical changes in response to burial have previously been reported for trees that had formed adventitious roots, we stress the role of adventitious-root formation as a possible trigger behind the local changes in wood anatomy, reflecting a functional change of a buried stem towards a root. Based on our field experiment, it seems unlikely that years of shallow or moderate burial events (≤50 cm) can be reconstructed using the wood structure of buried stems. As epitropic roots develop quickly after burial, dating such roots may potentially yield better estimates of burial events. Further research on the relation between adventitious root and changes in stem anatomy is needed to ascertain the precision of dating sand-burial events using tree rings.     

树木不同着生位置1级根的形态、解剖结构和氮含量

树木根系中1级根在养分和水分吸收方面发挥着重要作用。研究1级根的形态结构与功能的联系,对了解1级根的生理功能和寿命,以及森林生态系统碳和养分的循环具有重要的理论意义。但是,1级根在根系统中,因着生的位置不同,可能表现出不同的生理生态功能。该研究以胡桃楸(Juglans mandshurica)、黄波罗(Phellodendron amurense)和水曲柳(Fraxinus mandshurica)人工林根系的1级根为研究对象,研究了不同着生位置的1级根的形态、解剖结构和组织化学特征。将1级根按着生位置的不同分成3类:Ar类根为2级根上的1级根;Br类根为2级–4级根的根尖;Cr类根为3级–5级根上的1级根。结果表明:不同着生位置的1级根,形态、解剖结构和组织化学方面都具有高度的异质性。3类1级根中,Ar类根数量多、根长较短、直径细,N含量高,皮层比例高、维根比低,主要由二原型原生木质部的根组成;Br类和Cr类根数量较少、单根较长、直径粗,N含量低,皮层比例低、维根比高,主要由多原型原生木质部的根组成。研究结果对了解不同着生位置的1级根的吸收功能和寿命具有重要的理论意义。     

Tracheid anatomical responses to climate in a forest-steppe in Southern Siberia

Tree-rings are precious natural archives to assess ecosystem variability over time. Xylem anatomy in woody tissue is a promising source of information in tree-rings since it is closely linked to tree hydraulics and carbon fixation. However, despite the rising interest for cell anatomy in dendrochronology, still little is known about the interpretation of the variability of cell anatomical responses observed across different environments and species.Here we analyze cell anatomical responses to increasing summer drought on 18 trees from 3 conifer species (Picea obovata, Pinus sylvestris, and Larix sibirica) growing in the transition zone between forest and steppe in the Republic of Khakassia (Russia). Analyses include the comparison of tracheid size distributions along climatic gradients and contrasting micro-topography from 1986 to 2008.Results indicate an overall decrease of earlywood tracheid lumen and cell wall thickness to high temperature and drought regardless of species and site conditions. In particular an increase of one degree Celsius during the summer caused up to 5% reduction of earlywood cell lumen and wall thickness. These anatomical shifts suggest that a downscaling of hydraulic efficiency is not paralleled by increased hydraulic safety, presumably due to carbon limitation.Based on the results of this case study, we suggest that increasing drought stress might hamper the formation of a functional xylem structure, thus being a possible trigger for a miss-acclimation causing long-term decline and higher exposure to hydraulic failures. Despite the promising study approach, more studies including more data (trees, years) and broader climatic gradients would be needed to further improve our mechanistic understanding.     

栓皮栎根系解剖结构、水力特性及碳、氮含量研究

为阐明栓皮栎根系随径级的变化规律,探究其细根的合理划分标准。以1年生栓皮栎幼苗为研究对象,将其根系分为1、1～2、2～3、3～4 mm四个径级,分别制作石蜡切片观察解剖结构,比较木质部水力特性,测定碳氮含量及其比值,并采用主成分法对根系进行分类。结果表明:(1)随着径级增加,栓皮栎根系周皮、韧皮部和形成层组织厚度增加而占径比降低,木质部直径及其占径比均增加。(2)直径2 mm以上的栓皮栎根系木质部平均最大和最小导管直径、根比导水率和栓塞脆弱性指数增加显著; 而导管密度显著下降,导管面积与木质部面积之比变化不显著。(3)直径2 mm以上栓皮栎根系碳含量表现出显著增加,随着径级增加,根系氮含量下降、碳氮比升高。(4)主成分分析表明,13项根系结构和元素含量指标降维后,前2个主分量方差贡献率达62%,PCA双序轴显示栓皮栎根系可划分为2 mm以下的吸收根群和2 mm以上的运输根群。综上认为,以2 mm作为栓皮栎细根划分的标准兼顾了形态和功能的特点,更具有准确性。     

Application of growth rings and scars in exposed roots of Schizolobium parahyba as a tool for dating geomorphic processes in the State of São Paulo,Brazil

Water erosion is an important degradation process, which results in loss of soil, reduction in agricultural productivity, and causes severe environmental impact. Dendrogeomorphology has methods in which the structure of the wood of the stem and roots of tree and shrub species affected by sediment deposition or by root exposure is analysed, to establish the chronology of erosive events. The objective of the present work was to describe the modifications and scar formation in Schizolobium parahyba growth rings, attributed to the effect of roots exposure for determining the first year of exposure. The study area presents erosion features, such as gullies, with the consequent exposure of tree roots. The growth rings and the scars formed by the flow of water and soil particles were analysed in cross-sections of exposed roots to date the erosion processes. This paper demonstrates the potential of S. parahyba for dendrogeomorphological studies, validating dendrogeomorphology as a research tool in tropical climate. Scars used for erosion dating in cross-sections have been proven as good indicators of geomorphic processes. The relevance of this work is to become the first attempt in tropical regions to date erosion processes using dendrogeomorphological techniques on exposed roots.     

Ephemeral root modules in Fraxinus mandshurica

Historically, ephemeral roots have been equated with 'fine roots' (i.e. all roots of less than an arbitrary diameter, such as 2 mm), but evidence shows that 'fine roots' in woody species are complex branching systems with both rapid-cycling and slow-cycling components. A precise definition of ephemeral roots is therefore needed. Using a branch-order classification, a rhizotron method and sequential sampling of a root cohort, we tested the hypothesis that ephemeral root modules exist within the branching Fraxinus mandshurica (Manchurian ash) root system as distal nonwoody lateral branches, which show anatomical, nutritional and physiological patterns distinct from their woody mother roots. Our results showed that in F. mandshurica, distal nonwoody root branch orders die rapidly as intact lateral branches (or modules). These nonwoody branch orders exhibited highly synchronous changes in tissue nitrogen concentrations and respiration, dominated root turnover, nutrient flux and root respiration, and never underwent secondary development. The ephemeral root modules proposed here may provide a functional basis for differentiating and sampling short-lived absorptive roots in woody plants, and represent a conceptual leap over the traditional coarse-fine root dichotomies based on arbitrary size classes.