Analysis of root fractal characteristics in remote areas of the Taklimakan desert, China

Fractal geometry is a potential new approach to analyze the root architecture, which may offer improved ways to quantify and summarize root system complexity as well as yield ecological and physiological insights into the functional relevance of specific architectural patterns. Fractal analysis is a sensitive measure of root branching intensity and fractal dimension expresses the ＂space filling＂ properties of a structure. The objective of this study was to find out the fractal characteristics of root systems in a remote area of the Taklimakan desert in China. The entire root system of two naturally occurring species were excavated and exposed with shov- els in 2007. The species were Tamarix taklamakanensis and Calligonum roborovskii. A one-factorial ANOVA with species as factor showed statistically a highly significant difference in fractal dimensions, indicating differences in their pattern of root branching. There was no relationship between root diameter and two parameters of fractal root models a and q, representing general characteris- tics of root systems, for either species （a： the ratio of the sum of root cross-sectional areas after a branching to the cross-sectional area before root division; q： the distribution of the cross-sectional areas after branching）. We have found significant linear relation- ships between the diameter after branching and root length and biomass respectively, because of the self-similarity of root branching. Branching rules are the same for roots of all sizes and lengths. Root biomass for the root systems of entire trees can be estimated by measuring the diameter of each root at the base of the trunk or the diameter after branching. We have shown that the diameter of each root at the base of the trunk and the diameter after branching are effective indices that can be measured easily in order to estimate the root lengths, biomass and other parameters of root architecture.     

Contrasting rooting patterns of some arid-zone fruit tree species from Botswana – I. Fine root distribution

To assess the possible degree of root competition from fruit trees which could potentially be used in agroforestry systems, fine root density of fruit trees Strychnos cocculoides Bak., Strychnos spinosa Lam. (Loganiaceae) and Vangueria infausta Burch. (Rubiaceae), as well as from the shrubby species, Grewia flava DC. (Tiliaceae) was investigated. Vangueria infausta had the highest fine root densities in both vertical and horizontal extensions. In Vangueria infausta fine root density decreased with increasing soil depth. For the other species in the 80 cm soil profile investigated, no significant changes in fine root density with soil depth were found. For Strychnos cocculoides almost no fine roots were detected in the upper soil horizon (0–20 cm). Using fine root surface area densities, exploration and exploitation indices were calculated. Vangueria infausta had the highest value of the exploration index compared to the other species. For use in agroforestry systems Vangueria infausta was estimated to be the most competitive of the investigated species, whereas Strychnos cocculoides seems to be the less competitive. Strychnos cocculoides has additionally spatial arrangements of fine roots favourable for agroforestry, slowly increasing with depth and additionally low concentrations in upper soil layers.     

Root architecture in relation to tree-soil-crop interactions and shoot pruning in agroforestry

Desirable root architecture for trees differs between sequential and simultaneous agroforestry systems. In sequential systems extensive tree root development may enhance nutrient capture and transfer to subsequent crops via organic pools. In simultaneous systems tree root development in the crop root zone leads to competition for resources.Fractal branching models provide relationships between proximal root diameter, close to the tree stem, and total root length or surface area. The main assumption is that a root branching proportionality factor is independent of root diameter. This was tested in a survey of 18 multipurpose trees growing on an acid soil in Lampung (Indonesia). The assumption appeared valid for all trees tested, for stems as well as roots. The proportionality factor showed a larger variability in roots than in stems and the effects of this variabilily should be further investigated. A simple index of tree root shallowness is proposed as indicator of tree root competitiveness, based on superficial roots and stem diameter.Pruning trees is a major way to benefit from tree products and at the same time reduce above-ground competition between trees and crops. It may have negative effects, however, on root distribution and enhance below-ground competition. In an experiment with five tree species, a lower height of stem pruning led to a larger number of superficial roots of smaller diameter, but had no effect on shoot:root ratios or the relative importance of the tap root.     

Structural root form and fine root distribution of some woody species evaluated for agroforestry systems

Fine root distribution and structural root characteristics of four-years-old multipurpose woody species with potential for use in agroforestry systems were investigated on an Alfisol in the forest savanna transition zone of south-western Nigeria. Rooting patterns of woody species studied differed considerably. Lonchocarpus sericeus had the lowest percentage (21%) of total fine roots confined to the upper (0 to 30 cm) soil layer, compared to 84% with Tetrapleura tetraptera. Despite the superior tap root system of Enterolobium cyclocarpum and the fine root form of Nauchlea latifolia however, their extensive distribution and very large lateral woody root volume density may pose a major problem for seedbed preparation and tillage operations in simultaneous agroforestry systems. Lonchocarpus sericeus appears to have the most desirable structural and fine root architecture among the species studied for simultaneous agroforestry systems. The total root density below the underlying distal soil layers was linearly related to the sum of square of tap root diameter and the corresponding soil depths. This revised version was published online in June 2006 with corrections to the Cover Date.     

Hydraulic constraints in the functional scaling of trees

I conducted a literature survey to assess the available information on relationships between size--expressed in terms of diameter and dry biomass--and hydraulic efficiency of woody structures at different scales, from stem segments to whole trees. Three data sets were constructed: the first described the relationship between segment diameter and hydraulic conductivity (k(h); kg m s(-1) MPa(-1)) in four species; the second, for the same four species, described the intraspecific trajectories of change in total hydraulic conductance (G; kg s(-1) MPa(-1)) during ontogeny, i.e., from saplings to mature trees, thereby providing a comparison between allometric scaling laws at the scales of segments and whole trees; the third comprised pooled means for nine species that described the interspecific trajectory of change in G with tree size. The scaling coefficients obtained were compared with predictions made with an architectural fractal-like model incorporating tissue-specific hydraulic architecture parameters (West et al. 1999). When data on segment k(h) were examined, the fractal-like model closely predicted the scaling of k(h) with segment diameter in four species. However, the model failed to predict accurately in all species the intraspecific scaling at the branch and whole-tree levels, and consistently overestimated the scaling coefficients. The results suggest that ontogenetic changes in tree size during the life cycle of one tree may result in tradeoffs between optimal hydraulic supply to the existing leaf area and maintenance costs of the supporting xylem tissue. The model of West et al. (1999) may be useful for understanding broad interspecific patterns, but not for understanding more subtle ontogenetic changes.     

Patterns of root respiration rates and morphological traits in 13 tree species in a tropical forest

The root systems of forest trees are composed of different diameters and heterogeneous physiological traits. However, the pattern of root respiration rates from finer and coarser roots across various tropical species remains unknown. To clarify how respiration is related to the morphological traits of roots, we evaluated specific root respiration and its relationships to mean root diameter (D) of various diameter and root tissue density (RTD; root mass per unit root volume; gcm(-3)) and specific root length (SRL; root length per unit root mass; mg(-1)) of the fine roots among and within 14 trees of 13 species from a primary tropical rainforest in the Pasoh Forest Reserve in Peninsular Malaysia. Coarse root (2-269mm) respiration rates increased with decreasing D, resulting in significant relationships between root respiration and diameter across species. A model based on a radial gradient of respiration rates of coarse roots simulated the exponential decrease in respiration with diameter. The respiration rate of fine roots (<2mm) was much higher and more variable than those of larger diameter roots. For fine roots, the mean respiration rates for each species increased with decreasing D. The respiration rates of fine roots declined markedly with increasing RTD and increased with increasing SRL, which explained a significant portion of the variation in the respiration among the 14 trees from 13 species examined. Our results indicate that coarse root respiration in tree species follows a basic relationship with D across species and that most of the variation in fine root respiration among species is explained by D, RTD and SRL. We found that the relationship between root respiration and morphological traits provides a quantitative basis for separating fine roots from coarse roots and that the pattern holds across different species.     

The effect of tree architecture on conduit diameter and frequency from small distal roots to branch tips in Betula pendula, Picea abies and Pinus sylvestris

We studied the effect of tree architecture on xylem anatomy in three Betula pendula Roth., three Picea abies (L.) H. Karst. and three Pinus sylvestris (L.) trees (mean age 35 years). First, the analysis of conduit anatomy in different tree parts showed that conduits tapered and their frequency increased from roots (≥ 2 mm) to stem, from stem to branches and further to leaf petioles in B. pendula. Conduit anatomy in lateral and main roots, as well as lateral and main branches, significantly differed from each other in all the studied species. The increase in conduit diameter and decrease in frequency from the pith to the bark were clear aboveground, but variable patterns were observed belowground. In the leaf petioles of B. pendula, conduit diameter increased and conduit frequency decreased with increasing individual leaf area. Second, the results concerning the scaling of conduit diameter were compared with the predictions of the general vascular scaling model (WBE model) and Murray's law. The scaling parameter values at the tree level corresponded with the predictions of the WBE model in all the studied trees except for one tree of both conifer species. However, the scaling parameter values changed from one tree compartment to another rather than remaining uniform inside a tree, as assumed by the WBE model. The assumptions of the WBE model of a constant conductivity ratio, constant tapering and an unchanged total number of conduits were not fulfilled. When the conductivity ratio and relative tapering were plotted together, the results aboveground corresponded quite well with Murray's law: the conductivity ratio increased when relative tapering decreased. Our results support the theory that trees adjust both their macro- and microstructure to maximize their water transport efficiency, but also to prevent embolism and ensure mechanical safety.     

Fine root dynamics in organic and mineral soil layers of <Emphasis Type="Italic">Cryptomeria japonica</Emphasis> D. Don plantation

Fine root dynamics and root architecture were studied in the organic and mineral soil layers of a Cryptomeria japonica plantation. Fine root biomass (<1 mm) showed seasonal changes whereas fine root biomass (1–2 mm) was unchanged over the study period. Root tips were grouped into size classes based on root tip diameter, including <0.5, 0.5–1, and 1–2 mm. Root tip density (<1 mm) was significantly correlated with fine root biomass (<1 mm). Root tip density and fine root biomass (<1 mm) increased in summer and decreased in winter, and both showed a similar seasonal pattern. Root tip dynamics influenced fine root dynamics. Root architecture as expressed by branching intensity changed with root tip production and mortality. Branching intensity also showed a similar seasonal pattern of root tip density dynamics. Root tips of both <0.5 and 0.5–1 mm were mainly produced in the organic soil layer, while root tips of 0.5–1 mm were mainly produced in the mineral soil layer. Because of the high RT1 root tip production in the organic soil layer, branching intensity was higher in the organic soil than in the mineral soil layer during summer. Root tip dynamics influenced fine root dynamics and the architecture of root systems in both organic and mineral soil layers.     

Assaying the allelopathic effects of <Emphasis Type="Italic">Eucalyptus camaldulensis</Emphasis> in a nursery bed incorporated with leaf litter

Allelopathic effects of Eucalyptus camaldulensis Dehnh. were confirmed in Petri dish and pot experiments in our previous studies. However, the degree to which such effects under controlled experiments exist in more complex ecological settings remains to be tested. Thus, the present study was carried out by incorporating different proportions of ground litter of E. camaldulensis in soil. The growth of three agricultural crops: falen (Vigna unguiculata (L.) Walp.), chickpea (Cicer arietinum L.), and arhor (Cajanus cajan (L.) Millsp.), and two tree species, kala koroi (Albizia procera (Roxb.) Benth.) and ipil ipil (Leucaena leucocephala (Lam.) de Wit) were tested. There were inhibitory effects of leaf litter on germination, shoot and root growth, leaf number, and collar diameter as well as a reduction of nodulation by legume crops (25–80% reduction). The extent of the effects was dependent on the proportion of leaf litter, the species and the type of traits. In contrast to shoot growth, the effect on root growth was more severe. No effect on germination was found with the agriculture crops while the two tree species showed reduced germination. The effect was greater in the presence of higher proportions of leaf litter mixed in soil while in some cases lower proportions stimulated growth. Not all species were suppressed; A. procera, C. cajan, V. unguiculata showed compatible growth while C. arietinum and L. leucocephala were found incompatible. This study provides evidence that E. camaldulensis has allelopathic potential under field conditions and a careful selection of associated crops in agroforestry systems is highly recommended.     

Detection of tree roots and determination of root diameters by ground penetrating radar under optimal conditions

A tree's root system accounts for between 10 and 65% of its total biomass, yet our understanding of the factors that cause this proportion to vary is limited because of the difficulty encountered when studying tree root systems. There is a need to develop new sampling and measuring techniques for tree root systems. Ground penetrating radar (GPR) offers the potential for direct nondestructive measurements of tree root biomass and root distributions to be made. We tested the ability of GPR, with 500 MHz, 800 MHz and 1 GHz antennas, to detect tree roots and determine root size by burying roots in a 32 m3 pit containing damp sand. Within this test bed, tree roots were buried in two configurations: (1) roots of various diameters (1-10 cm) were buried at a single depth (50 cm); and (2) roots of similar diameter (about 5 cm) were buried at various depths (15-155 cm). Radar antennas were drawn along transects perpendicular to the buried roots. Radar profile normalization, filtration and migration were undertaken based on standard algorithms. All antennas produced characteristic reflection hyperbolas on the radar profiles allowing visual identification of most root locations. The 800 MHz antenna resulted in the clearest radar profiles. An unsupervised, maximum-convexity migration algorithm was used to focus information contained in the hyperbolas back to a point. This resulted in a significant gain in clarity with roots appearing as discrete shapes, thereby reducing confusion due to overlapping of hyperbolas when many roots are detected. More importantly, parameters extracted from the resultant waveform through the center of a root correlated well with root diameter for the 500 MHz antenna, but not for the other two antennas. A multiple regression model based on the extracted parameters was calibrated on half of the data (R2 = 0.89) and produced good predictions when tested on the remaining data. Root diameters were predicted with a root mean squared error of 0.6 cm, allowing detection and quantification of roots as small as 1 cm in diameter. An advantage of this processing technique is that it produces results independently of signal strength. These waveform parameters represent a major advance in the processing of GPR profiles for estimating root diameters. We conclude that enhanced data analysis routines combined with improvements in GPR hardware design could make GPR a valuable tool for studying tree root systems.     

Irradiance-induced plasticity in the hydraulic properties of saplings of different temperate broad-leaved forest tree species

We assessed the irradiance-related plasticity of hydraulic architecture in saplings of Betula pendula Roth., a pioneer species; Acer pseudoplatanus L., Fraxinus excelsior L. and Quercus robur L., which are post-pioneer light-requiring species; and Quercus petraea Matt. Liebl. and Fagus sylvatica L. Plants were grown in pots in 36%, 16% and 4% of full sunlight. Hydraulic conductance was measured with a high-pressure flow-meter in entire, in situ root systems and in excised shoots. Leaf-specific whole-plant conductance (LSC) increased with irradiance, due, in part, to an effect of irradiance on plant size. In addition, there was a size-independent effect of irradiance on LSC due, in part, to an increase in root hydraulic conductance paralleled by an increase in root biomass scaled to leaf area. Changes in shoot conductivity also contributed to the size-independent plasticity of LSC. Vulnerability to cavitation measured in current-year twigs was much larger in shade-grown plants. Betula pendula had the highest whole-plant, root and shoot conductances and also the greatest vulnerability to cavitation. The other species were similar in LSC, but showed some variation in root conductance scaled to biomass, with Q. robur, Q. petraea and F. sylvatica having the lowest root conductance and susceptibility to cavitation. All species showed a similar irradiance-related plasticity in LSC.     

植物生态适应性特征及关键种选择研究进展

植物对生境特征的适应是其存活和定居的关键。植被适应性修复研究是退化生态系统修复的前提之一,已经引起科学家们的广泛关注。文中综述了植物生态适应性研究,主要有叶片生理生态特性、解剖结构和根系构型、生态学特征等,但这些研究少有辅助关键树种选择和植物群落结构调控;进一步分析了关键树种选择技术,主要包含与立地条件、生态因子的适应性和种间关系的和谐。为了提高退化生态系统适应性修复水平,未来需要加强以下3个方面的工作:划分树种适应性功能群,综合考虑光、温、肥与水的耦合作用关系,根系构型调控研究。     

Observations on the rooting patterns of some agroforestry trees in an arid region of north-western India

The present study deals with root architecture of 6-year-old trees of 9 indigenous and 3 exotic species growing in arid climate of north-western India. Observations, made on excavated root systems (3 tree replicates of each species) showed large variation in horizontal and vertical spread of roots. In Morus alba, Melia azedarach and Populus deltoides, the roots were confined to 80 cm, while in Prosopis cineraria, Acacia nilotica and Eucalyptus tereticornis, roots penetrated more deeply to 233 cm. The number of total roots ranged from 103 in Acacia catechu to 1932 in Eucalyptus tereticornis, and 62 to 80% of the roots were less than 2 mm in diameter. The primary roots were more horizontal than the secondary roots. The total root biomass varied from 2.2 kg in Acacia catechu to 30.6 kg^–1 tree in Populus deltoides, and top 30 cm soil contained 42 to 78% of the total biomass. The implications of the results are discussed in the context of the ecological niche of the species, and its usefulness in agroforestry systems.     

Nitrogen forms affect root structure and water uptake in the hybrid poplar

The study analyses the effects of two different forms of nitrogen fertilisation (nitrate and ammonium) on root structure and water uptake of two hybrid poplar (Populus maximowiczii x P. balsamifera) clones in a field experiment. Water uptake was studied using sap flow gauges on individual proximal roots and coarse root structure was examined by excavating 18 whole-root systems. Finer roots were scanned and analyzed for architecture. Nitrogen forms did not affect coarse-root system development, but had a significant effect on fine-root development. Nitrate-treated trees presented higher fine:coarse root ratios and higher specific root lengths than control or ammonium treated trees. These allocation differences affected the water uptake capacity of the plants as reflected by the higher sapflow rate in the nitrate treatment. The diameter of proximal roots at the tree base predicted well the total root biomass and length. The diameter of smaller lateral roots also predicted the lateral root mass, length, surface area and the number of tips. The effect of nitrogen fertilisation on the fine root structure translated into an effect on the functioning of the fine roots forming a link between form (architecture) and function (water uptake).     

Relating coarse root respiration to root diameter in clonal Eucalyptus stands in the Republic of the Congo

Root respiration is an important component of the carbon balance of a forest ecosystem. We measured CO2 efflux of excised fine roots and intact coarse roots in 3-, 4- and 13-year-old Eucalyptus stands in the region of Pointe-Noire, Republic of the Congo. A transportable and adaptable closed chamber gas exchange system directly measured CO2 efflux of roots from 0.5 to 32 mm in diameter. Fluxes were corrected for measurement system leaks and normalized to a reference temperature of 30 degrees C. Mean fine root respiration rates at the reference temperature varied between 8.5 and 10.8 micromol CO2 kg(-1) s(-1) depending on the stand. Coarse root respiration was strongly negatively correlated to root diameter. We propose a model based on a radial gradient of respiratory activity within the root to simulate the exponential decrease in respiration with diameter. Although many sources of uncertainty in the measurements remain, as discussed in this paper, these results provide a basis for scaling up organ-level root respiration measurements to the tree and stand levels.     

Tree growth and management in Ugandan agroforestry systems: effects of root pruning on tree growth and crop yield

Tree root pruning is a potential tool for managing belowground competition when trees and crops are grown together in agroforestry systems. We investigated the effects of tree root pruning on shoot growth and root distribution of Alnus acuminata (H.B. & K.), Casuarina equisetifolia L., Grevillea robusta A. Cunn. ex R. Br., Maesopsis eminii Engl. and Markhamia lutea (Benth.) K. Schum. and on yield of adjacent crops in sub-humid Uganda. The trees were 3 years old at the commencement of the study, and most species were competing strongly with crops. Tree roots were pruned 41 months after planting by cutting and back-filling a trench to a depth of 0.3 m, at a distance of 0.3 m from the trees, on one side of the tree row. The trench was reopened and roots recut at 50 and 62 months after planting. We assessed the effects on tree growth and root distribution over a 3 year period, and crop yield after the third root pruning at 62 months. Overall, root pruning had only a slight effect on aboveground tree growth: height growth was unaffected and diameter growth was reduced by only 4%. A substantial amount of root regrowth was observed by 11 months after pruning. Tree species varied in the number and distribution of roots, and C. equisetifolia and M. lutea had considerably more roots per unit of trunk volume than the other species, especially in the surface soil layers. Casuarina equisetifolia and M. eminii were the tree species most competitive with crops and G. robusta and M. lutea the least competitive. Crop yield data provided strong evidence of the redistribution of root activity following root pruning, with competition increasing on the unpruned side of tree rows. Thus, one-sided root pruning will be useful in only a few circumstances.     

Initial seedling morphological characteristics and field performance of two Sudanian savanna species in relation to nursery production period and watering regimes

It has become apparent that some interventions are required to aid the regeneration of woody species in the Sudanian savanna. Direct seeding has been ineffective, thus planting high quality seedlings may be a viable alternative. In this study, we examined the stock quality of two valuable Sudanian savanna species, Acacia macrostachya and Pterocarpus erinaceus. Different nursery production periods were tested as well as the species’ field performance under well-watered and stressed conditions. The results showed that older seedlings (9-month) were morphologically distinct from younger ones (3-month), particularly in the case of P. erinaceus. Eighteen months after planting out, survival and growth of seedlings were not affected by initial seedling size; this was the result of the high root to shoot ratio of seedlings in all age groups at the time of planting. Seedling mortality as high as 30% was observed and attributed to both drought stress and other factors such as herbivory. Regression analyses revealed that initial shoot height was a poor predictor of field performance for both species, but initial root collar diameter accounted for 25% of the variation in diameter of P. erinaceus in the field. We conclude that initial seedling size does not affect survival and growth in the field provided that all sizes of seedling have a high root to shoot ratio at the time of planting. The prediction of field performance could be improved by developing a model that incorporates a wide range of root collar diameter.     

Hydraulic conductance and soil water potential at the soil-root interface of Pinus pinaster seedlings inoculated with different dikaryons of Pisolithus sp

Seedlings of maritime pine (Pinus pinaster Ait.) were inoculated with different dikaryons of Pisolithus sp. from South Africa to determine the influence of extension of the extramatrical phase and diameter of the mycelial strands on water relations parameters including xylem water potential (Psi(x)), soil water potential at the soil-root interface (Psi(s)) and hydraulic conductance (L(p)) during and after a period of water stress. Seedlings inoculated with dikaryons having an extensive extramatrical phase and large diameter mycelial strands showed higher Psi(s) (-2 MPa) during severe water stress than seedlings inoculated with dikaryons producing fine hyphae and sparse extramatrical phases (-3.8 MPa). Seedlings inoculated with strand-forming dikaryons recovered faster from water stress than did non-inoculated seedlings or seedlings inoculated with non-strand-forming dikaryons. Architectural aspects of the extramatrical phase, including the presence of large diameter mycelial strands or fine hyphae, influenced the soil-root contact and the water relations of an inoculated host plant. When water stress was not limiting, the architecture of the extramatrical phase did not have a large effect on Psi(s). It is suggested that the architecture of the extramatrical phase influences the resistance to water flow through the soil-root interface and that large mycelial strands increase the water flow by bridging the gap between the soil and the root. These changes in physiology indicate that dikaryons can improve the survival of Pinus pinaster under dry conditions.