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1.  Deep Phenotyping of Coarse Root Architecture in R. pseudoacacia Reveals That Tree Root System Plasticity Is Confined within Its Architectural Model 
PLoS ONE  2013;8(12):e83548.
This study aims at assessing the influence of slope angle and multi-directional flexing and their interaction on the root architecture of Robinia pseudoacacia seedlings, with a particular focus on architectural model and trait plasticity. 36 trees were grown from seed in containers inclined at 0° (control) or 45° (slope) in a glasshouse. The shoots of half the plants were gently flexed for 5 minutes a day. After 6 months, root systems were excavated and digitized in 3D, and biomass measured. Over 100 root architectural traits were determined. Both slope and flexing increased significantly plant size. Non-flexed trees on 45° slopes developed shallow roots which were largely aligned perpendicular to the slope. Compared to the controls, flexed trees on 0° slopes possessed a shorter and thicker taproot held in place by regularly distributed long and thin lateral roots. Flexed trees on the 45° slope also developed a thick vertically aligned taproot, with more volume allocated to upslope surface lateral roots, due to the greater soil volume uphill. We show that there is an inherent root system architectural model, but that a certain number of traits are highly plastic. This plasticity will permit root architectural design to be modified depending on external mechanical signals perceived by young trees.
doi:10.1371/journal.pone.0083548
PMCID: PMC3873950  PMID: 24386227
2.  Linking carbon supply to root cell-wall chemistry and mechanics at high altitudes in Abies georgei 
Annals of Botany  2010;107(2):311-320.
Background and Aims
The mobile carbon supply to different compartments of a tree is affected by climate, but its impact on cell-wall chemistry and mechanics remains unknown. To understand better the variability in root growth and biomechanics in mountain forests subjected to substrate mass movement, we investigated root chemical and mechanical properties of mature Abies georgei var. smithii (Smith fir) growing at different elevations on the Tibet–Qinghai Plateau.
Methods
Thin and fine roots (0·1–4·0 mm in diameter) were sampled at three different elevations (3480, 3900 and 4330 m, the last corresponding to the treeline). Tensile resistance of roots of different diameter classes was measured along with holocellulose and non-structural carbon (NSC) content.
Key Results
The mean force necessary to break roots in tension decreased significantly with increasing altitude and was attributed to a decrease in holocellulose content. Holocellulose was significantly lower in roots at the treeline (29·5 ± 1·3 %) compared with those at 3480 m (39·1 ± 1·0 %). Roots also differed significantly in NSC, with 35·6 ± 4·1 mg g−1 dry mass of mean total soluble sugars in roots at 3480 m and 18·8 ± 2·1 mg g−1 dry mass in roots at the treeline.
Conclusions
Root mechanical resistance, holocellulose and NSC content all decreased with increasing altitude. Holocellulose is made up principally of cellulose, the biosynthesis of which depends largely on NSC supply. Plants synthesize cellulose when conditions are optimal and NSC is not limiting. Thus, cellulose synthesis in the thin and fine roots measured in our study is probably not a priority in mature trees growing at very high altitudes, where climatic factors will be limiting for growth. Root NSC stocks at the treeline may be depleted through over-demand for carbon supply due to increased fine root production or winter root growth.
doi:10.1093/aob/mcq237
PMCID: PMC3025735  PMID: 21186240
Abies georgei; biomechanics; tensile resistance; cellulose content; non-structural carbon; root strength; Tibet–Qinghai; slope stability
3.  (Not) Keeping the stem straight: a proteomic analysis of maritime pine seedlings undergoing phototropism and gravitropism 
BMC Plant Biology  2010;10:217.
Background
Plants are subjected to continuous stimuli from the environment and have evolved an ability to respond through various growth and development processes. Phototropism and gravitropism responses enable the plant to reorient with regard to light and gravity.
Results
We quantified the speed of maritime pine seedlings to reorient with regard to light and gravity over 22 days. Seedlings were inclined at 15, 30 and 45 degrees with vertical plants as controls. A lateral light source illuminated the plants and stem movement over time was recorded. Depending on the initial angle of stem lean, the apical response to the lateral light source differed. In control and 15° inclined plants, the apex turned directly towards the light source after only 2 h. In plants inclined at 30° and 45°, the apex first reoriented in the vertical plane after 2 h, then turned towards the light source after 24 h. Two-dimensional gel electrophoresis coupled with mass spectrometry was then used to describe the molecular response of stem bending involved in photo- and gravi-tropism after 22 hr and 8 days of treatment. A total of 486 spots were quantitatively analyzed using image analysis software. Significant changes were determined in the protein accumulation of 68 protein spots. Early response gravitropic associated proteins were identified, which are known to function in energy related and primary metabolism. A group of thirty eight proteins were found to be involved in primary metabolism and energy related metabolic pathways. Degradation of Rubisco was implicated in some protein shifts.
Conclusions
Our study demonstrates a rapid gravitropic response in apices of maritime pine seedlings inclined >30°. Little or no response was observed at the stem bases of the same plants. The primary gravitropic response is concomitant with a modification of the proteome, consisting of an over accumulation of energy and metabolism associated proteins, which may allow the stem to reorient rapidly after bending.
doi:10.1186/1471-2229-10-217
PMCID: PMC3017815  PMID: 20925929
4.  Plant Growth Modelling and Applications: The Increasing Importance of Plant Architecture in Growth Models 
Annals of Botany  2008;101(8):1053-1063.
Background
Modelling plant growth allows us to test hypotheses and carry out virtual experiments concerning plant growth processes that could otherwise take years in field conditions. The visualization of growth simulations allows us to see directly and vividly the outcome of a given model and provides us with an instructive tool useful for agronomists and foresters, as well as for teaching. Functional–structural (FS) plant growth models are nowadays particularly important for integrating biological processes with environmental conditions in 3-D virtual plants, and provide the basis for more advanced research in plant sciences.
Scope
In this viewpoint paper, we ask the following questions. Are we modelling the correct processes that drive plant growth, and is growth driven mostly by sink or source activity? In current models, is the importance of soil resources (nutrients, water, temperature and their interaction with meristematic activity) considered adequately? Do classic models account for architectural adjustment as well as integrating the fundamental principles of development? Whilst answering these questions with the available data in the literature, we put forward the opinion that plant architecture and sink activity must be pushed to the centre of plant growth models. In natural conditions, sinks will more often drive growth than source activity, because sink activity is often controlled by finite soil resources or developmental constraints.
PMA06
This viewpoint paper also serves as an introduction to this Special Issue devoted to plant growth modelling, which includes new research covering areas stretching from cell growth to biomechanics. All papers were presented at the Second International Symposium on Plant Growth Modeling, Simulation, Visualization and Applications (PMA06), held in Beijing, China, from 13–17 November, 2006. Although a large number of papers are devoted to FS models of agricultural and forest crop species, physiological and genetic processes have recently been included and point the way to a new direction in plant modelling research.
doi:10.1093/aob/mcn050
PMCID: PMC2710283  PMID: 18387970
Biomechanics; carbon allocation; functional–structural plant models; meristem; nitrogen; phenotypic plasticity; root architecture; simulation; sink; source; PMA06
5.  Understanding the Impact of Root Morphology on Overturning Mechanisms: A Modelling Approach 
Annals of Botany  2007;101(8):1267-1280.
Background and Aims
The Finite Element Method (FEM) has been used in recent years to simulate overturning processes in trees. This study aimed at using FEM to determine the role of individual roots in tree anchorage with regard to different rooting patterns, and to estimate stress distribution in the soil and roots during overturning.
Methods
The FEM was used to carry out 2-D simulations of tree uprooting in saturated soft clay and loamy sand-like soil. The anchorage model consisted of a root system embedded in a soil block. Two root patterns were used and individual roots removed to determine their contribution to anchorage.
Key Results
In clay-like soil the size of the root–soil plate formed during overturning was defined by the longest roots. Consequently, all other roots localized within this plate had no influence on anchorage strength. In sand-like soil, removing individual root elements altered anchorage resistance. This result was due to a modification of the shape and size of the root–soil plate, as well as the location of the rotation axis. The tap root and deeper roots had more influence on overturning resistance in sand-like soil compared with clay-like soil. Mechanical stresses were higher in the most superficial roots and also in leeward roots in sand-like soil. The relative difference in stresses between the upper and lower sides of lateral roots was sensitive to root insertion angle. Assuming that root eccentricity is a response to mechanical stresses, these results explain why eccentricity differs depending on root architecture.
Conclusions
A simple 2-D Finite Element model was developed to better understand the mechanisms involved during tree overturning. It has been shown how root system morphology and soil mechanical properties can modify the shape of the root plate slip surface as well as the position of the rotation axis, which are major components of tree anchorage.
doi:10.1093/aob/mcm245
PMCID: PMC2710277  PMID: 17942593
Acclimative growth; anchorage; biomechanics; tree uprooting; rotation axis; root architecture; root eccentricity; secondary growth; von Mises stresses
6.  Using Three-dimensional Plant Root Architecture in Models of Shallow-slope Stability 
Annals of Botany  2007;101(8):1281-1293.
Background
The contribution of vegetation to shallow-slope stability is of major importance in landslide-prone regions. However, existing slope stability models use only limited plant root architectural parameters. This study aims to provide a chain of tools useful for determining the contribution of tree roots to soil reinforcement.
Methods
Three-dimensional digitizing in situ was used to obtain accurate root system architecture data for mature Quercus alba in two forest stands. These data were used as input to tools developed, which analyse the spatial position of roots, topology and geometry. The contribution of roots to soil reinforcement was determined by calculating additional soil cohesion using the limit equilibrium model, and the factor of safety (FOS) using an existing slope stability model, Slip4Ex.
Key Results
Existing models may incorrectly estimate the additional soil cohesion provided by roots, as the spatial position of roots crossing the potential slip surface is usually not taken into account. However, most soil reinforcement by roots occurs close to the tree stem and is negligible at a distance >1·0 m from the tree, and therefore global values of FOS for a slope do not take into account local slippage along the slope.
Conclusions
Within a forest stand on a landslide-prone slope, soil fixation by roots can be minimal between uniform rows of trees, leading to local soil slippage. Therefore, staggered rows of trees would improve overall slope stability, as trees would arrest the downward movement of soil. The chain of tools consisting of both software (free for non-commercial use) and functions available from the first author will enable a more accurate description and use of root architectural parameters in standard slope stability analyses.
doi:10.1093/aob/mcm199
PMCID: PMC2710273  PMID: 17766845
Landslide; root area ratio; slope stability; 3D digitizing; Quercus alba; soil cohesion; soil internal friction angle

Results 1-6 (6)