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1.  Variation of Maximum Tree Height and Annual Shoot Growth of Smith Fir at Various Elevations in the Sygera Mountains, Southeastern Tibetan Plateau 
PLoS ONE  2012;7(3):e31725.
Little is known about tree height and height growth (as annual shoot elongation of the apical part of vertical stems) of coniferous trees growing at various altitudes on the Tibetan Plateau, which provides a high-elevation natural platform for assessing tree growth performance in relation to future climate change. We here investigated the variation of maximum tree height and annual height increment of Smith fir (Abies georgei var. smithii) in seven forest plots (30 m×40 m) along two altitudinal transects between 3,800 m and 4,200/4,390 m above sea level (a.s.l.) in the Sygera Mountains, southeastern Tibetan Plateau. Four plots were located on north-facing slopes and three plots on southeast-facing slopes. At each site, annual shoot growth was obtained by measuring the distance between successive terminal bud scars along the main stem of 25 trees that were between 2 and 4 m high. Maximum/mean tree height and mean annual height increment of Smith fir decreased with increasing altitude up to the tree line, indicative of a stress gradient (the dominant temperature gradient) along the altitudinal transect. Above-average mean minimum summer (particularly July) temperatures affected height increment positively, whereas precipitation had no significant effect on shoot growth. The time series of annual height increments of Smith fir can be used for the reconstruction of past climate on the southeastern Tibetan Plateau. In addition, it can be expected that the rising summer temperatures observed in the recent past and anticipated for the future will enhance Smith fir's growth throughout its altitudinal distribution range.
doi:10.1371/journal.pone.0031725
PMCID: PMC3291607  PMID: 22396738
2.  Spatial and seasonal variations in mobile carbohydrates in Pinus cembra in the timberline ecotone of the Central Austrian Alps 
European journal of forest research  2011;130(2):173-179.
To test whether the altitudinal limit of tree growth is determined by carbons shortage or by a limitation in growth we investigated non structural carbohydrates and their components starch and total soluble sugars in Pinus cembra trees along an elevational gradient in the timberline ecotone of the Central Austrian Alps. NSC contents in needles, branches, stems, and coarse roots were measured throughout an entire growing season. At the tissue level NSC contents were not significantly more abundant in treeline trees as compared to trees at lower elevations. Along our 425 m elevational transect from the closed forest to the treeline we failed to find a stable elevational trend in the total NSC pool of entire trees and observed within season increases in the tree’s NSC pool that can be attributed to an altitudinal increase in leaf mass as needles contained the largest NSC fraction of the whole tree NSC pool. Furthermore, whole tree NSC contents were positively correlated with net photosynthetic capacity. Although our observed NSC characteristics do not support the hypothesis that tree life at their upper elevational limit is determined by an insufficient carbon balance we found no consistent confirmation for the sink limitation hypothesis.
doi:10.1007/s10342-010-0419-7
PMCID: PMC3191523  PMID: 22003357
Non structural carbohydrates; seasonal variation; elevational gradient; timberline ecotone; treeline formation; treelife limitation
3.  Seasonal Dynamics of Mobile Carbon Supply in Quercus aquifolioides at the Upper Elevational Limit 
PLoS ONE  2012;7(3):e34213.
Many studies have tried to explain the physiological mechanisms of the alpine treeline phenomenon, but the debate on the alpine treeline formation remains controversial due to opposite results from different studies. The present study explored the carbon-physiology of an alpine shrub species (Quercus aquifolioides) grown at its upper elevational limit compared to lower elevations, to test whether the elevational limit of alpine shrubs (<3 m in height) are determined by carbon limitation or growth limitation. We studied the seasonal variations in non-structural carbohydrate (NSC) and its pool size in Q. aquifolioides grown at 3000 m, 3500 m, and at its elevational limit of 3950 m above sea level (a.s.l.) on Zheduo Mt., SW China. The tissue NSC concentrations along the elevational gradient varied significantly with season, reflecting the season-dependent carbon balance. The NSC levels in tissues were lowest at the beginning of the growing season, indicating that plants used the winter reserve storage for re-growth in the early spring. During the growing season, plants grown at the elevational limit did not show lower NSC concentrations compared to plants at lower elevations, but during the winter season, storage tissues, especially roots, had significantly lower NSC concentrations in plants at the elevational limit compared to lower elevations. The present results suggest the significance of winter reserve in storage tissues, which may determine the winter survival and early-spring re-growth of Q. aquifolioides shrubs at high elevation, leading to the formation of the uppermost distribution limit. This result is consistent with a recent hypothesis for the alpine treeline formation.
doi:10.1371/journal.pone.0034213
PMCID: PMC3316670  PMID: 22479567
4.  Relationships between C3 Plant Foliar Carbon Isotope Composition and Element Contents of Grassland Species at High Altitudes on the Qinghai-Tibet Plateau, China 
PLoS ONE  2013;8(4):e60794.
Relationships of foliar carbon isotope composition (δ13C) with foliar C, N, P, K, Ca, Mg contents and their ratios of 219 C3 species leaf samples, obtained in August in 2004 to 2007 from 82 high altitude grassland sites on the Qinghai-Tibet Plateau China, were examined. This was done with reference to the proposition that foliar δ13C increases with altitude and separately for the life-form groups of graminoids, forbs and shrubs and for the genera Stipa and Kobresia. For all samples, foliar δ13C was negatively related to foliar K, P and ∑K+ Ca+ Mg, and positively correlated to foliar C, C/N and C/P. The significance of these correlations differed for the taxonomic and life-form groups. Lack of a relationship of foliar δ13C with foliar N was inconsistent with the majority of studies that have shown foliar δ13C to be positively related to foliar N due to a decrease of Ci/Ca (the ratio between intercellular and atmospheric concentration of CO2) and explained as a result of greater photosynthetic capacity at higher foliar N concentration. However this inconsistency relates to other high altitude studies that have found that photosynthetic capacity remains constant as foliar N increases. After accounting for the altitudinal relationship with foliar δ13C, of the elements only the K effect was significant and was most strongly expressed for Kobresia. It is concluded that factors critical to plant survival and growth at very high altitudes, such as low atmospheric pressure and low temperatures, may preclude expression of relationships between foliar δ13C and foliar elements that have been observed at lower altitudes.
doi:10.1371/journal.pone.0060794
PMCID: PMC3614917  PMID: 23565275
5.  Girdling Affects Ectomycorrhizal Fungal (EMF) Diversity and Reveals Functional Differences in EMF Community Composition in a Beech Forest▿ † 
The relationships between plant carbon resources, soil carbon and nitrogen content, and ectomycorrhizal fungal (EMF) diversity in a monospecific, old-growth beech (Fagus sylvatica) forest were investigated by manipulating carbon flux by girdling. We hypothesized that disruption of the carbon supply would not affect diversity and EMF species numbers if EM fungi can be supplied by plant internal carbohydrate resources or would result in selective disappearance of EMF taxa because of differences in carbon demand of different fungi. Tree carbohydrate status, root demography, EMF colonization, and EMF taxon abundance were measured repeatedly during 1 year after girdling. Girdling did not affect root colonization but decreased EMF species richness of an estimated 79 to 90 taxa to about 40 taxa. Cenococcum geophilum, Lactarius blennius, and Tomentella lapida were dominant, colonizing about 70% of the root tips, and remained unaffected by girdling. Mainly cryptic EMF species disappeared. Therefore, the Shannon-Wiener index (H′) decreased but evenness was unaffected. H′ was positively correlated with glucose, fructose, and starch concentrations of fine roots and also with the ratio of dissolved organic carbon to dissolved organic nitrogen (DOC/DON), suggesting that both H′ and DOC/DON were governed by changes in belowground carbon allocation. Our results suggest that beech maintains numerous rare EMF species by recent photosynthate. These EM fungi may constitute biological insurance for adaptation to changing environmental conditions. The preservation of taxa previously not known to colonize beech may, thus, form an important reservoir for future forest development.
doi:10.1128/AEM.01703-09
PMCID: PMC2837996  PMID: 20097809
6.  Effects of Soil Compaction and Meloidogyne incognita on Cotton Root Architecture and Plant Growth 
Journal of Nematology  2013;45(2):112-121.
The effects of a soil hardpan and Meloidogyne incognita on cotton root architecture and plant growth were evaluated in microplots in 2010 and 2011. Soil was infested with M. incognita at four different levels with or without a hardpan. The presence of a hardpan resulted in increased plant height, number of main stem nodes, and root fresh weight for cotton seedlings both years. Meloidogyne incognita decreased height and number of nodes for seedlings in 2010. Nematode infestation increased seedling root length and enhanced root magnitude, altitude, and exterior path length in 2010. This was also the case for root length and magnitude in 2011 at lower infestation levels suggesting compensatory growth. A hardpan had no consistent effect on these root parameters but increased root volume in both years. A hardpan hastened crop maturity and increased the number of fruiting branches that were produced, while M. incognita infection delayed crop development and reduced plant height and number of bolls. Both M. incognita infection and a hardpan reduced taproot length and root dry weight below the hardpan in both years. Root topological indices under all the treatments ranged from 1.71 to 1.83 both years indicating that root branching followed a herringbone pattern. The techniques for characterizing root architecture that were used in this study provide a greater understanding of changes that result from disease and soil abiotic parameters affecting root function and crop productivity.
PMCID: PMC3700738  PMID: 23833326
ecology; host-parasite relationship; root-knot nematode; root topology; soil hardpan
7.  Functional traits and root morphology of alpine plants 
Annals of Botany  2011;108(3):537-545.
Background and Aims
Vegetation has long been recognized to protect the soil from erosion. Understanding species differences in root morphology and functional traits is an important step to assess which species and species mixtures may provide erosion control. Furthermore, extending classification of plant functional types towards root traits may be a useful procedure in understanding important root functions.
Methods
In this study, pioneer data on traits of alpine plant species, i.e. plant height and shoot biomass, root depth, horizontal root spreading, root length, diameter, tensile strength, plant age and root biomass, from a disturbed site in the Swiss Alps are presented. The applicability of three classifications of plant functional types (PFTs), i.e. life form, growth form and root type, was examined for above- and below-ground plant traits.
Key Results
Plant traits differed considerably among species even of the same life form, e.g. in the case of total root length by more than two orders of magnitude. Within the same root diameter, species differed significantly in tensile strength: some species (Geum reptans and Luzula spicata) had roots more than twice as strong as those of other species. Species of different life forms provided different root functions (e.g. root depth and horizontal root spreading) that may be important for soil physical processes. All classifications of PFTs were helpful to categorize plant traits; however, the PFTs according to root type explained total root length far better than the other PFTs.
Conclusions
The results of the study illustrate the remarkable differences between root traits of alpine plants, some of which cannot be assessed from simple morphological inspection, e.g. tensile strength. PFT classification based on root traits seems useful to categorize plant traits, even though some patterns are better explained at the individual species level.
doi:10.1093/aob/mcr169
PMCID: PMC3158688  PMID: 21795278
Growth rings; plant diversity; plant functional types; PFTs; ski slope; soil erosion; tensile strength; alpine plants
8.  Evaluation of Radicular Dentin Thickness of Danger Zone in Mandibular First Molars 
Objective:
Better understanding of the furcation anatomy may serve to decrease the risk of root perforation. The purpose of this study was to measure the thickness of root walls in the danger zone in mandibular first molars.
Materials and Methods:
The roots of 53 extracted human mandibular first molars were sectioned in the horizontal plane 4 mm below the orifice of the mesial and distal root canals. For each cut surface buccal, lingual, mesial, and distal thickness of the root wall was measured. Mean values of the thickness at each location were calculated and compared by ANOVA and t-test.
Results:
The results showed that the mean thickness in the distal portion of the mesial root was smaller in comparison to all other portions of the roots (P<0.05) and this difference was statistically significant except for the mesial portion of the distal root (P=0.463). The mean thickness of radicular dentin at the distal aspect of mesial roots was 1.2 millimeter.
Conclusion:
Our study suggests that knowledge of the root dentin thickness in the danger zone is essential for preventing endodontic mishaps leading to failure.
PMCID: PMC3184764  PMID: 21998796
Endodontics; Dentin; Instrumentation; Molar; Mandible
9.  Nitrogen Regulation of Root Branching 
Annals of Botany  2006;97(5):875-881.
• Background Many plant species can modify their root architecture to enable them to forage for heterogeneously distributed nutrients in the soil. The foraging response normally involves increased proliferation of lateral roots within nutrient-rich soil patches, but much remains to be understood about the signalling mechanisms that enable roots to sense variations in the external concentrations of different mineral nutrients and to modify their patterns of growth and development accordingly.
• Scope In this review we consider different aspects of the way in which the nitrogen supply can modify root branching, focusing on Arabidopsis thaliana. Our current understanding of the mechanism of nitrate stimulation of lateral root growth and the role of the ANR1 gene are summarized. In addition, evidence supporting the possible role of auxin in regulating the systemic inhibition of early lateral root development by high rates of nitrate supply is presented. Finally, we examine recent evidence that an amino acid, l-glutamate, can act as an external signal to elicit complex changes in root growth and development.
• Conclusions It is clear that plants have evolved sophisticated pathways for sensing and responding to changes in different components of the external nitrogen supply as well as their own internal nitrogen status. We speculate on the possibility that the effects elicited by external l-glutamate represent a novel form of foraging response that could potentially enhance a plant's ability to compete with its neighbours and micro-organisms for localized sources of organic nitrogen.
doi:10.1093/aob/mcj601
PMCID: PMC2803407  PMID: 16339770
Arabidopsis thaliana; auxin; dissolved organic nitrogen; foraging; glutamate; lateral roots; MADS box transcription factor; nitrate; nitrogen; root architecture; root development; roots; signalling; Thlaspi caerulescens
10.  Genotypic differences in Al resistance and the role of cell-wall pectin in Al exclusion from the root apex in Fagopyrum tataricum 
Annals of Botany  2010;107(3):371-378.
Background and Aims
Aluminium (Al) toxicity is one of the factors limiting crop production on acid soils. However, genotypic differences exist among plant species or cultivars in response to Al toxicity. This study aims to investigate genotypic differences among eight cultivars of tatary buckwheat (Fagopyrum tataricum) for Al resistance and explore the possible mechanisms of Al resistance.
Methods
Al resistance was evaluated based on relative root elongation (root elongation with Al/root elongation without Al). Root apex Al content, pectin content and exudation of root organic acids were determined and compared.
Key Results
Genotypic differences among the eight cultivars were correlated with exclusion of Al from the root apex. However, there was a lack of correlation between Al exclusion and Al-induced oxalate secretion. Interestingly, cell-wall pectin content of the root apex was generally lower in Al-resistant cultivars than in Al-sensitive cultivars. Although we were unable to establish a significant correlation between Al exclusion and pectin content among the eight cultivars, a strong correlation could be established among six cultivars, in which the pectin content in the most Al-resistant cultivar ‘Chuan’ was significantly lower than that in the most Al-sensitive cultivar ‘Liuku2’. Furthermore, root apex cell-wall pectin methylesterase activity (PME) was similar in ‘Chuan’ and ‘Liuku2’ in the absence of Al, but Al treatment resulted in increased PME activity in ‘Liuku2’ compared with ‘Chuan’. Immunolocalization of pectins also showed that the two cultivars had similar amounts of either low-methyl-ester pectins or high-methyl-ester pectins in the absence of Al, but Al treatment resulted in a more significant increase of low-methyl-ester pectins and decrease of high-methyl-ester pectins in ‘Liuku2’.
Conclusions
Cell-wall pectin content may contribute, at least in part, to differential Al resistance among tatary buckwheat cultivars.
doi:10.1093/aob/mcq254
PMCID: PMC3043930  PMID: 21183454
Aluminium resistance; cell wall; exclusion mechanism; Fagopyrum tataricum; pectin; pectin methylesterase; oxalate; toxicity
11.  Fine-root carbon and nitrogen concentration of European beech (Fagus sylvatica L.) in Italy Prealps: possible implications of coppice conversion to high forest 
Fine-root systems represent a very sensitive plant compartment to environmental changes. Gaining further knowledge about their dynamics would improve soil carbon input understanding. This paper investigates C and N concentrations in fine roots in relation to different stand characteristics resulting from conversion of coppiced forests to high forests. In order to evaluate possible interferences due to different vegetative stages of vegetation, fine-root sampling was repeated six times in each stand during the same 2008 growing season. Fine-root sampling was conducted within three different soil depths (0–10; 10–20; and 20–30 cm). Fine-root traits were measured by means of WinRHIZO software which enable us to separate them into three different diameter classes (0–0.5, 0.5–1.0 and 1.0–2.0 mm). The data collected indicate that N concentration was higher in converted stands than in the coppiced stand whereas C concentration was higher in the coppiced stand than in converted stands. Consequently the fine-root C:N ratio was significantly higher in coppiced than in converted stands and showed an inverse relationship with fine-root turnover rate, confirming a significant change of fine-root status after the conversion of a coppice to high forest.
doi:10.3389/fpls.2013.00192
PMCID: PMC3680728  PMID: 23785374
fine-root carbon; fine-root nitrogen; stand characteristics; coppice conversion; Fagus sylvatica L. fine roots; fine roots soil depth; beech fine roots
12.  Relationship between Shoot-rooting and Root-sprouting Abilities and the Carbohydrate and Nitrogen Reserves of Mediterranean Dwarf Shrubs 
Annals of Botany  2007;100(4):865-874.
Background and Aims
This study analysed the differences in nitrogen (N), non-structural carbohydrates (NSC) and biomass allocation to the roots and shoots of 18 species of Mediterranean dwarf shrubs with different shoot-rooting and resprouting abilities. Root N and NSC concentrations of strict root-sprouters and species resprouting from the base of the stems were also compared.
Methods
Soluble sugars (SS), starch and N concentrations were assessed in roots and shoots. The root : shoot ratio of each species was obtained by thorough root excavations. Cross-species analyses were complemented by phylogenetically independent contrasts (PICs).
Key Results
Shoot-rooting species showed a preferential allocation of starch to shoots rather than roots as compared with non-shoot-rooting species. Resprouters displayed greater starch concentrations than non-sprouters in both shoots and roots. Trends were maintained after PICs analyses, but differences became weak when root-sprouters versus non-root-sprouters were compared. Within resprouters, strict root-sprouters showed greater root concentrations and a preferential allocation of starch to the roots than stem-sprouters. No differences were found in the root : shoot ratio of species with different rooting and resprouting abilities.
Conclusions
The shoot-rooting ability of Mediterranean dwarf shrubs seems to depend on the preferential allocation of starch and SS to shoots, though alternative C-sources such as current photosynthates may also be involved. In contrast to plants from other mediterranean areas of the world, the resprouting ability of Mediterranean dwarf shrubs is not related to a preferential allocation of N, NSC and biomass to roots.
doi:10.1093/aob/mcm185
PMCID: PMC2749641  PMID: 17728338
Mediterranean dwarf shrubs; nitrogen; non-structural carbohydrates; sprouting; resprouting; disturbance response; root : shoot ratio; shoot-rooting ability; adventitious roots; allocation pattern; starch; chamaephytes
13.  Do elevations in temperature, CO2, and nutrient availability modify belowground carbon gain and root morphology in artificially defoliated silver birch seedlings? 
Ecology and Evolution  2013;3(9):2783-2794.
Climate warming increases the risk of insect defoliation in boreal forests. Losses in photosynthetically active surfaces cause reduction in net primary productivity and often compromise carbon reserves of trees. The concurrent effects of climate change and removal of foliage on root growth responses and carbohydrate dynamics are poorly understood, especially in tree seedlings. We investigated if exposures to different combinations of elevated temperature, CO2, and nutrient availability modify belowground carbon gain and root morphology in artificially defoliated 1-year-old silver birches (Betula pendula). We quantified nonstructural carbohydrates (insoluble starch as a storage compound; soluble sucrose, fructose, and glucose) singly and in combination in fine roots of plants under winter dormancy. Also the total mass, fine root proportion, water content, and length of roots were defined. We hypothesized that the measured properties are lower in defoliated birch seedlings that grow with ample resources than with scarce resources. On average, fertilization markedly decreased both the proportion and the carbohydrate concentrations of fine roots in all seedlings, whereas the effect of fertilization on root water content and dry mass was the opposite. However, defoliation mitigated the effect of fertilization on the root water content, as well as on the proportion of fine roots and their carbohydrate concentrations by reversing the outcomes. Elevation in temperature decreased and elevation in CO2 increased the absolute contents of total nonstructural carbohydrates, whereas fertilization alleviated both these effects. Also the root length and mass increased by CO2 elevation. This confirms that surplus carbon in birch tissues is used as a substrate for storage compounds and for cell wall synthesis. To conclude, our results indicate that some, but not all elements of climate change alter belowground carbon gain and root morphology in defoliated silver birch seedlings.
doi:10.1002/ece3.665
PMCID: PMC3790529  PMID: 24101972
Betula pendula; climate change; fertilization; fine roots; folivory; plant sugar allocation
14.  Natural Selection and Adaptive Evolution of Leptin in the Ochotona Family Driven by the Cold Environmental Stress 
PLoS ONE  2008;3(1):e1472.
Background
Environmental stress can accelerate the evolutionary rate of specific stress-response proteins and create new functions specialized for different environments, enhancing an organism's fitness to stressful environments. Pikas (order Lagomorpha), endemic, non-hibernating mammals in the modern Holarctic Region, live in cold regions at either high altitudes or high latitudes and have a maximum distribution of species diversification confined to the Qinghai-Tibet Plateau. Variations in energy metabolism are remarkable for them living in cold environments. Leptin, an adipocyte-derived hormone, plays important roles in energy homeostasis.
Methodology/Principal Findings
To examine the extent of leptin variations within the Ochotona family, we cloned the entire coding sequence of pika leptin from 6 species in two regions (Qinghai-Tibet Plateau and Inner Mongolia steppe in China) and the leptin sequences of plateau pikas (O. curzonia) from different altitudes on Qinghai-Tibet Plateau. We carried out both DNA and amino acid sequence analyses in molecular evolution and compared modeled spatial structures. Our results show that positive selection (PS) acts on pika leptin, while nine PS sites located within the functionally significant segment 85-119 of leptin and one unique motif appeared only in pika lineages-the ATP synthase α and β subunit signature site. To reveal the environmental factors affecting sequence evolution of pika leptin, relative rate test was performed in pikas from different altitudes. Stepwise multiple regression shows that temperature is significantly and negatively correlated with the rates of non-synonymous substitution (Ka) and amino acid substitution (Aa), whereas altitude does not significantly affect synonymous substitution (Ks), Ka and Aa.
Conclusions/Significance
Our findings support the viewpoint that adaptive evolution may occur in pika leptin, which may play important roles in pikas' ecological adaptation to extreme environmental stress. We speculate that cold, and probably not hypoxia, may be the primary environmental factor for driving adaptive evolution of pika leptin.
doi:10.1371/journal.pone.0001472
PMCID: PMC2194619  PMID: 18213380
15.  Morphological plasticity of ectomycorrhizal short roots in Betula sp and Picea abies forests across climate and forest succession gradients: its role in changing environments 
Morphological plasticity of ectomycorrhizal (EcM) short roots (known also as first and second order roots with primary development) allows trees to adjust their water and nutrient uptake to local environmental conditions. The morphological traits (MTs) of short-living EcM roots, such as specific root length (SRL) and area, root tip frequency per mass unit (RTF), root tissue density, as well as mean diameter, length, and mass of the root tips, are good indicators of acclimation. We investigated the role of EcM root morphological plasticity across the climate gradient (48–68°N) in Norway spruce (Picea abies (L.) Karst) and (53–66°N) birch (Betula pendula Roth., B. pubescens Ehrh.) forests, as well as in primary and secondary successional birch forests assuming higher plasticity of a respective root trait to reflect higher relevance of that characteristic in acclimation process. We hypothesized that although the morphological plasticity of EcM roots is subject to the abiotic and biotic environmental conditions in the changing climate; the tools to achieve the appropriate morphological acclimation are tree species-specific. Long-term (1994–2010) measurements of EcM roots morphology strongly imply that tree species have different acclimation-indicative root traits in response to changing environments. Birch EcM roots acclimated along latitude by changing mostly SRL [plasticity index (PI) = 0.60], while spruce EcM roots became adjusted by modifying RTF (PI = 0.68). Silver birch as a pioneer species must have a broader tolerance to environmental conditions across various environments; however, the mean PI of all MTs did not differ between early-successional birch and late-successional spruce. The differences between species in SRL, and RTF, diameter, and length decreased southward, toward temperate forests with more favorable growth conditions. EcM root traits reflected root-rhizosphere succession across forest succession stages.
doi:10.3389/fpls.2013.00335
PMCID: PMC3759007  PMID: 24032035
plasticity; ectomycorrhizal roots; morphological characteristics; root acclimation; silver birch; Norway spruce; forest succession; climate gradient
16.  The effects of rising atmospheric carbon dioxide on shoot-root nitrogen and water signaling 
Terrestrial higher plants are composed of roots and shoots, distinct organs that conduct complementary functions in dissimilar environments. For example, roots are responsible for acquiring water and nutrients such as inorganic nitrogen from the soil, yet shoots consume the majority of these resources. The success of such a relationship depends on excellent root–shoot communications. Increased net photosynthesis and decreased shoot nitrogen and water use at elevated CO2 fundamentally alter these source–sink relations. Lower than predicted productivity gains at elevated CO2 under nitrogen or water stress may indicate shoot–root signaling lacks plasticity to respond to rising atmospheric CO2 concentrations. The following presents recent research results on shoot–root nitrogen and water signaling, emphasizing the influence that rising atmospheric carbon dioxide levels are having on these source–sink interactions.
doi:10.3389/fpls.2013.00304
PMCID: PMC3739423  PMID: 23983674
carbon dioxide; nitrogen; nitrate assimilation; water; drought; salinity; chilling
17.  Use of FT-IR Spectra and PCA to the Bulk Characterization of Cell Wall Residues of Fruits and Vegetables Along a Fraction Process 
Food Biophysics  2012;8(1):29-42.
This study focuses on the analysis of polysaccharide residues from the cell walls of fruits and vegetables: tomato, potato, pumpkin, carrot and celery root. An alcohol-insoluble residue was prepared from plant material by extraction using the hot ethyl alcohol method and then cell wall fractions soluble in trans-1,2-diaminocyclohexane-N,N,N′,N′-tetraacetate, sodium carbonate and alkaline solution were sequentially extracted. Infrared spectroscopy combined with Fourier transform (FT-IR) was used to evaluate differences among cell wall residues and among species after each step of sequential extraction of pectins and hemicelluloses. Additionally, pectic substances were identified using an Automated Wet Chemistry Analyser. Principal component analysis (PCA) was applied to FT-IR spectra in two regions: 1,800–1,200 cm−1 and 1,200–800 cm−1 in order to distinguish different components of cell wall polysaccharides. This method also allowed us the possibility of highlighting the most important wavenumbers for each type of polysaccharide: 1,740, 1,610 and 1,240 cm−1 denoting pectins or 1,370 and 1,317 cm−1 denoting hemicelluloses and cellulose, respectively.
doi:10.1007/s11483-012-9279-7
PMCID: PMC3593005  PMID: 23487553
FT-IR spectroscopy; Vegetable and fruit cell walls; Principal component analysis; Pectin; Hemicellulose
18.  Seasonal changes of whole root system conductance by a drought-tolerant grape root system 
Journal of Experimental Botany  2010;62(1):99-109.
The role of root systems in drought tolerance is a subject of very limited information compared with above-ground responses. Adjustments to the ability of roots to supply water relative to shoot transpiration demand is proposed as a major means for woody perennial plants to tolerate drought, and is often expressed as changes in the ratios of leaf to root area (AL:AR). Seasonal root proliferation in a directed manner could increase the water supply function of roots independent of total root area (AR) and represents a mechanism whereby water supply to demand could be increased. To address this issue, seasonal root proliferation, stomatal conductance (gs) and whole root system hydraulic conductance (kr) were investigated for a drought-tolerant grape root system (Vitis berlandieri×V. rupestris cv. 1103P) and a non-drought-tolerant root system (Vitis riparia×V. rupestris cv. 101-14Mgt), upon which had been grafted the same drought-sensitive clone of Vitis vinifera cv. Merlot. Leaf water potentials (ψL) for Merlot grafted onto the 1103P root system (–0.91±0.02 MPa) were +0.15 MPa higher than Merlot on 101-14Mgt (–1.06±0.03 MPa) during spring, but dropped by approximately –0.4 MPa from spring to autumn, and were significantly lower by –0.15 MPa (–1.43±0.02 MPa) than for Merlot on 101-14Mgt (at –1.28±0.02 MPa). Surprisingly, gs of Merlot on the drought-tolerant root system (1103P) was less down-regulated and canopies maintained evaporative fluxes ranging from 35–20 mmol vine−1 s−1 during the diurnal peak from spring to autumn, respectively, three times greater than those measured for Merlot on the drought-sensitive rootstock 101-14Mgt. The drought-tolerant root system grew more roots at depth during the warm summer dry period, and the whole root system conductance (kr) increased from 0.004 to 0.009 kg MPa−1 s−1 during that same time period. The changes in kr could not be explained by xylem anatomy or conductivity changes of individual root segments. Thus, the manner in which drought tolerance was conveyed to the drought-sensitive clone appeared to arise from deep root proliferation during the hottest and driest part of the season, rather than through changes in xylem structure, xylem density or stomatal regulation. This information can be useful to growers on a site-specific basis in selecting rootstocks for grape clonal material (scions) grafted to them.
doi:10.1093/jxb/erq247
PMCID: PMC2993904  PMID: 20851906
Drought tolerance; grape; root hydraulic conductance; root hydraulic conductivity; root water relations; stomatal conductance; Vitis vinifera
19.  No evidence for depletion of carbohydrate pools in Scots pine (Pinus sylvestris L.) under drought stress 
The physiological mechanisms leading to Scots pine (Pinus sylvestris L.) decline in the dry inner Alpine valleys are still unknown. Testing the carbon starvation hypothesis, we analysed the seasonal course of mobile carbohydrate pools (NSC) of Scots pine growing at a xeric and a dry-mesic site within an inner Alpine dry valley (750 m a.s.l., Tyrol, Austria) during the year 2009, which was characterized by exceptional soil dryness. Although, soil moisture content dropped to c. 10% at both sites during the growing season, NSC concentrations were rising in all tissues (branch, stem, root) till end of July, except in needles where maxima were reached around bud break. NSC concentrations were not significantly different in the analysed tissues at the xeric and the dry-mesic site. At the dry-mesic site NSC concentrations in the above ground tree biomass were significantly higher during the period of radial growth. An accumulation of NSC in roots at the end of July indicates a change in carbon allocation after an early cessation in above ground growth, possibly due to elevated below ground carbon demand. In conclusion our results revealed that extensive soil dryness during the growing season did not lead to carbon depletion. However, even though C-reserves were not exhausted, a sequestration of carbohydrate pools during drought periods might lead to deficits in carbon supply that weaken tree vigour and drive tree mortality.
doi:10.1111/j.1438-8677.2011.00467.x
PMCID: PMC3427021  PMID: 21974742
non-structural carbohydrates; Scots pine; drought; dry inner Alpine valley; carbon starvation; tree mortality
20.  Root Exudates from Grafted-Root Watermelon Showed a Certain Contribution in Inhibiting Fusarium oxysporum f. sp. niveum 
PLoS ONE  2013;8(5):e63383.
Grafting watermelon onto bottle gourd rootstock is commonly used method to generate resistance to Fusarium oxysporum f. sp. niveum (FON), but knowledge of the effect of the root exudates of grafted watermelon on this soil-borne pathogen in rhizosphere remains limited. To investigate the root exudate profiles of the own-root bottle gourd, grafted-root watermelon and own-root watermelon, recirculating hydroponic culture system was developed to continuously trap these root exudates. Both conidial germination and growth of FON were significantly decreased in the presence of root exudates from the grafted-root watermelon compared with the own-root watermelon. HPLC analysis revealed that the composition of the root exudates released by the grafted-root watermelon differed not only from the own-root watermelon but also from the bottle gourd rootstock plants. We identified salicylic acid in all 3 root exudates, chlorogenic acid and caffeic acid in root exudates from own-root bottle gourd and grafted-root watermelon but not own-root watermelon, and abundant cinnamic acid only in own-root watermelon root exudates. The chlorogenic and caffeic acid were candidates for potentiating the enhanced resistance of the grafted watermelon to FON, therefore we tested the effects of the two compounds on the conidial germination and growth of FON. Both phenolic acids inhibited FON conidial germination and growth in a dose-dependent manner, and FON was much more susceptible to chlorogenic acid than to caffeic acid. In conclusion, the key factor in attaining the resistance to Fusarium wilt is grafting on the non-host root stock, however, the root exudates profile also showed some contribution in inhibiting FON. These results will help to better clarify the disease resistance mechanisms of grafted-root watermelon based on plant-microbe communication and will guide the improvement of strategies against Fusarium-mediated wilt of watermelon plants.
doi:10.1371/journal.pone.0063383
PMCID: PMC3659071  PMID: 23700421
21.  Modeling a Cortical Auxin Maximum for Nodulation: Different Signatures of Potential Strategies 
Lateral organ formation from plant roots typically requires the de novo creation of a meristem, initiated at the location of a localized auxin maximum. Legume roots can form both root nodules and lateral roots. From the basic principles of auxin transport and metabolism only a few mechanisms can be inferred for increasing the local auxin concentration: increased influx, decreased efflux, and (increased) local production. Using computer simulations we investigate the different spatio-temporal patterns resulting from each of these mechanisms in the context of a root model of a generalized legume. We apply all mechanisms to the same group of preselected cells, dubbed the controlled area. We find that each mechanism leaves its own characteristic signature. Local production by itself can not create a strong auxin maximum. An increase of influx, as is observed in lateral root formation, can result in an auxin maximum that is spatially more confined than the controlled area. A decrease of efflux on the other hand leads to a broad maximum, which is more similar to what is observed for nodule primordia. With our prime interest in nodulation, we further investigate the dynamics following a decrease of efflux. We find that with a homogeneous change in the whole cortex, the first auxin accumulation is observed in the inner cortex. The steady state lateral location of this efflux reduced auxin maximum can be shifted by slight changes in the ratio of central to peripheral efflux carriers. We discuss the implications of this finding in the context of determinate and indeterminate nodules, which originate from different cortical positions. The patterns we have found are robust under disruption of the (artificial) tissue layout. The same patterns are therefore likely to occur in many other contexts.
doi:10.3389/fpls.2012.00096
PMCID: PMC3361061  PMID: 22654886
root nodules; auxin transport manipulation; modeling
22.  Root traits contributing to plant productivity under drought 
Geneticists and breeders are positioned to breed plants with root traits that improve productivity under drought. However, a better understanding of root functional traits and how traits are related to whole plant strategies to increase crop productivity under different drought conditions is needed. Root traits associated with maintaining plant productivity under drought include small fine root diameters, long specific root length, and considerable root length density, especially at depths in soil with available water. In environments with late season water deficits, small xylem diameters in targeted seminal roots save soil water deep in the soil profile for use during crop maturation and result in improved yields. Capacity for deep root growth and large xylem diameters in deep roots may also improve root acquisition of water when ample water at depth is available. Xylem pit anatomy that makes xylem less “leaky” and prone to cavitation warrants further exploration holding promise that such traits may improve plant productivity in water-limited environments without negatively impacting yield under adequate water conditions. Rapid resumption of root growth following soil rewetting may improve plant productivity under episodic drought. Genetic control of many of these traits through breeding appears feasible. Several recent reviews have covered methods for screening root traits but an appreciation for the complexity of root systems (e.g., functional differences between fine and coarse roots) needs to be paired with these methods to successfully identify relevant traits for crop improvement. Screening of root traits at early stages in plant development can proxy traits at mature stages but verification is needed on a case by case basis that traits are linked to increased crop productivity under drought. Examples in lesquerella (Physaria) and rice (Oryza) show approaches to phenotyping of root traits and current understanding of root trait genetics for breeding.
doi:10.3389/fpls.2013.00442
PMCID: PMC3817922  PMID: 24204374
root morphology; root architecture; hydraulic conductance; hydraulic conductivity; QTL; drought tolerance; MAS
23.  Radial Growth of Qilian Juniper on the Northeast Tibetan Plateau and Potential Climate Associations 
PLoS ONE  2013;8(11):e79362.
There is controversy regarding the limiting climatic factor for tree radial growth at the alpine treeline on the northeastern Tibetan Plateau. In this study, we collected 594 increment cores from 331 trees, grouped within four altitude belts spanning the range 3550 to 4020 m.a.s.l. on a single hillside. We have developed four equivalent ring-width chronologies and shown that there are no significant differences in their growth-climate responses during 1956 to 2011 or in their longer-term growth patterns during the period AD 1110–2011. The main climate influence on radial growth is shown to be precipitation variability. Missing ring analysis shows that tree radial growth at the uppermost treeline location is more sensitive to climate variation than that at other elevations, and poor tree radial growth is particularly linked to the occurrence of serious drought events. Hence water limitation, rather than temperature stress, plays the pivotal role in controlling the radial growth of Sabina przewalskii Kom. at the treeline in this region. This finding contradicts any generalisation that tree-ring chronologies from high-elevation treeline environments are mostly indicators of temperature changes.
doi:10.1371/journal.pone.0079362
PMCID: PMC3828375  PMID: 24244488
24.  The forces generated within the musculature of the left ventricular wall 
Heart  2004;90(2):200-207.
Objectives: To test the hypothesis that two populations of myocardial fibres—fibres aligned parallel to the surfaces of the wall and an additional population of fibres that extend obliquely through the wall—when working in concert produce a dualistic, self stabilising arrangement.
Methods: Assessment of tensile forces in the walls of seven porcine hearts by using needle probes. Ventricular diameter was measured with microsonometry and the intracavitary pressure through a fluid filled catheter. Positive inotropism was induced by dopamine, and negative inotropism by thiopental. The preload was raised by volume load and lowered by withdrawal of blood. Afterload was increased by inflation of a balloon in the aortic root. The anatomical orientation of the fibres was established subsequently in histological sections.
Results: The forces in the fibres parallel to the surface decreased 20–35% during systolic shrinkage of the ventricle, during negative inotropism, and during ventricular unloading. They increased 10–30% on positive inotropic stimulation and with augmentation in preload and afterload. The forces in the oblique transmural fibres increased 8–65% during systole, on positive inotropic medication, with an increase in afterload and during ventricular shrinkage, and decreased 36% on negative inotropic medication. There was a delay of up to 147 ms in the drop in activity during relaxation in the oblique transmural fibres.
Conclusion: Although the two populations of myocardial fibres are densely interwoven, it is possible to distinguish their functions with force probes. The delayed drop in force during relaxation in obliquely oriented fibres indicates that they are hindered in their shortening to an extent that parallels any increase in mural thickness. The transmural fibres, therefore, contribute to stiffening of the ventricular wall and hence to confining ventricular compliance.
doi:10.1136/hrt.2003.011650
PMCID: PMC1768069  PMID: 14729798
tangential fibres; unloaded contraction; oblique transmural fibres; auxotonic contraction; ventricular shaping
25.  Plant Neighbour Identity Matters to Belowground Interactions under Controlled Conditions 
PLoS ONE  2011;6(11):e27791.
Background
Root competition is an almost ubiquitous feature of plant communities with profound effects on their structure and composition. Far beyond the traditional view that plants interact mainly through resource depletion (exploitation competition), roots are known to be able to interact with their environment using a large variety of mechanisms that may inhibit or enhance access of other roots to the resource or affect plant growth (contest interactions). However, an extensive analysis on how these contest root interactions may affect species interaction abilities is almost lacking.
Methodology/Principal Findings
In a common garden experiment with ten perennial plant species we forced pairs of plants of the same or different species to overlap their roots and analyzed how belowground contest interactions affected plant performance, biomass allocation patterns, and competitive abilities under abundant resource supply. Our results showed that net interaction outcome ranged from negative to positive, affecting total plant mass and allocation patterns. A species could be a strong competitor against one species, weaker against another one, and even facilitator to a third species. This leads to sets of species where competitive hierarchies may be clear but also to groups where such rankings are not, suggesting that intransitive root interactions may be crucial for species coexistence.
Conclusions/Significance
The outcome of belowground contest interactions is strongly dependent on neighbours' identity. In natural plant communities this conditional outcome may hypothetically help species to interact in non-hierarchical and intransitive networks, which in turn might promote coexistence.
doi:10.1371/journal.pone.0027791
PMCID: PMC3219686  PMID: 22114696

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