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PubMed2000

TrExML: a maximum-likelihood approach for extensive tree-space exploration.

MOTIVATION: Maximum-likelihood analysis of nucleotide and amino acid sequences is a powerful approach for inferring phylogenetic relationships and for comparing evolutionary hypotheses. Because it is a computationally demanding and time-consuming process, most algorithms explore only a minute portion of tree-space, with the emphasis on finding the most likely tree while ignoring the less likely, but not significantly worse, trees. However, when such trees exist, it is equally important to identify them to give due consideration to the phylogenetic uncertainty. Consequently, it is necessary to change the focus of these algorithms such that near optimal trees are also identified. RESULTS: This paper presents the Advanced Stepwise Addition Algorithm for exploring tree-space and two algorithms for generating all binary trees on a set of sequences. The Advanced Stepwise Addition Algorithm has been implemented in TrExML, a phylogenetic program for maximum-likelihood analysis of nucleotide sequences. TrExML is shown to be more effective at finding near optimal trees than a similar program, fastDNAml, implying that TrExML offers a better approach to account for phylogenetic uncertainty than has previously been possible. A program, TreeGen, is also described; it generates binary trees on a set of sequences allowing for extensive exploration of tree-space using other programs. AVAILABILITY: TreeGen, TrExML, and the sequence data used to test the programs are available from the following two WWW sites: http://whitetail.bemidji.msus. edu/trexml/and http://jcsmr.anu.edu.au/dmm/humgen.+ ++html.

Algorithms↗

PubMed2003

Pair hidden Markov models on tree structures.

MOTIVATION: Computationally identifying non-coding RNA regions on the genome has much scope for investigation and is essentially harder than gene-finding problems for protein-coding regions. Since comparative sequence analysis is effective for non-coding RNA detection, efficient computational methods are expected for structural alignments of RNA sequences. On the other hand, Hidden Markov Models (HMMs) have played important roles for modeling and analysing biological sequences. Especially, the concept of Pair HMMs (PHMMs) have been examined extensively as mathematical models for alignments and gene finding. RESULTS: We propose the pair HMMs on tree structures (PHMMTSs), which is an extension of PHMMs defined on alignments of trees and provides a unifying framework and an automata-theoretic model for alignments of trees, structural alignments and pair stochastic context-free grammars. By structural alignment, we mean a pairwise alignment to align an unfolded RNA sequence into an RNA sequence of known secondary structure. First, we extend the notion of PHMMs defined on alignments of 'linear' sequences to pair stochastic tree automata, called PHMMTSs, defined on alignments of 'trees'. The PHMMTSs provide various types of alignments of trees such as affine-gap alignments of trees and an automata-theoretic model for alignment of trees. Second, based on the observation that a secondary structure of RNA can be represented by a tree, we apply PHMMTSs to the problem of structural alignments of RNAs. We modify PHMMTSs so that it takes as input a pair of a 'linear' sequence and a 'tree' representing a secondary structure of RNA to produce a structural alignment. Further, the PHMMTSs with input of a pair of two linear sequences is mathematically equal to the pair stochastic context-free grammars. We demonstrate some computational experiments to show the effectiveness of our method for structural alignments, and discuss a complexity issue of PHMMTSs.

Algorithms↗

PubMed2003

Parametric alignment of ordered trees.

MOTIVATION: Computing the similarity between two ordered trees has applications in RNA secondary structure comparison, genetics and chemical structure analysis. Alignment of tree is one of the proposed measures. Similar to pair-wise sequence comparison, there is often disagreement about how to weight matches, mismatches, indels and gaps when we compare two trees. For sequence comparison, the parametric sequence alignment tools have been developed. The users are allowed to see explicitly and completely the effect of parameter choices on the optimal sequence alignments. A similar tool for aligning two ordered trees is required in practice. RESULTS: We develop a parametric tool for aligning two ordered trees that allow users to see the effect of parameter choices on the optimal alignment of trees. Our contributions include: (1) develop a parametric tool for aligning two ordered trees; (2) design an efficient algorithm for aligning two ordered trees with gap penalties that runs in O(n(2)deg(2)) time, where n is the number of nodes in the trees and deg is the degree of the trees; and (3) reduce the space of the algorithm from O(n(2)deg(2)) to O(n log n. deg(2)). AVAILABILITY: The software is available at http://www.cs.cityu.edu.hk/~lwang/software/ParaTree

Algorithms↗

PubMed2006

Salinity tolerance of 'Valencia' orange trees on rootstocks with contrasting salt tolerance is not improved by moderate shade.

The effects of shading in combination with salinity treatments were studied in citrus trees on two rootstocks with contrasting salt tolerance to determine if shading could reduce the negative effects of salinity stress. Well-nourished 2-year-old 'Valencia' orange trees grafted on Cleopatra mandarin (Cleo, relatively salt tolerant) or Carrizo citrange (Carr, relatively salt sensitive), were grown either under a 50% shade cloth or left unshaded in full sunlight. Half the trees received no salinity treatment and half were salinized with 50 mM Cl- during two 9 week salinity periods in the spring and autumn interrupted by an 11 week rainy period. The shade treatment reduced midday leaf temperature and leaf-to-air vapour pressure deficit regardless of salinity treatments. In non-salinized trees, shade increased midday CO2 assimilation rate (A(CO2)) and stomatal conductance, but had no effect on leaf transpiration (E(lf)). Shade also increased leaf chlorophyll and photosynthetic water use efficiency (A(CO2)/E(lf)) in leaves on both rootstocks and increased total plant dry weight in Cleo. The salinity treatment reduced leaf growth and leaf gas exchange parameters. Shade decreased Cl- concentrations in leaves of salinized Carr trees, but had no effect on leaf or root Cl- of trees on Cleo. There were no significant differences in leaf gas exchange parameters of shaded and unshaded salinized plants but the growth reduction from salinity stress was actually greater for shaded than for unshaded trees. Shaded trees on both rootstocks had higher leaf Na+ than unshaded trees after the first salinity period, and this shade-induced elevated leaf Na+ persisted after the second salinity period in trees on Carr. Thus, shading did not alleviate the negative effects of salinity on growth and Na+ accumulation.

Carbon Dioxide↗

PubMed2004

The MinMax Squeeze: guaranteeing a minimal tree for population data.

We report that for population data, where sequences are very similar to one another, it is often possible to use a two-pronged (MinMax Squeeze) approach to prove that a tree is the shortest possible under the parsimony criterion. Such population data can be in a range where parsimony is a maximum likelihood estimator. This is in sharp contrast to the case with species data, where sequences are much further apart and the problem of guaranteeing an optimal phylogenetic tree is known to be computationally prohibitive for realistic numbers of species, irrespective of whether likelihood or parsimony is the optimality criterion. The Squeeze uses both an upper bound (the length of the shortest tree known) and a lower bound derived from partitions of the columns (the length of the shortest tree possible). If the two bounds meet, the shortest known tree is thus proven to be a shortest possible tree. The implementation is first tested on simulated data sets and then applied to 53 complete human mitochondrial genomes. The shortest possible trees for those data have several significant improvements from the published tree. Namely, a pair of Australian lineages comes deeper in the tree (in agreement with archaeological data), and the non-African part of the tree shows greater agreement with the geographical distribution of lineages.

Computational Biology↗

PubMed2006

Is there a star tree paradox?

Concerns have been raised that posterior probabilities on phylogenetic trees can be unreliable when the true tree is unresolved or has very short internal branches, because existing methods for Bayesian phylogenetic analysis do not explicitly evaluate unresolved trees. Two recent papers have proposed that evaluating only resolved trees results in a "star tree paradox": when the true tree is unresolved or close to it, posterior probabilities were predicted to become increasingly unpredictable as sequence length grows, resulting in inflated confidence in one resolved tree or another and an increasing risk of false-positive inferences. Here we show that this is not the case; existing Bayesian methods do not lead to an inflation of statistical confidence, provided the evolutionary model is correct and uninformative priors are assumed. Posterior probabilities do not become increasingly unpredictable with increasing sequence length, and they exhibit conservative type I error rates, leading to a low rate of false-positive inferences. With infinite data, posterior probabilities give equal support for all resolved trees, and the rate of false inferences falls to zero. We conclude that there is no star tree paradox caused by not sampling unresolved trees.

Bayes Theorem↗

PubMed1986

An estimate of the photosynthetic production of individual trees in a Chamaecyparis obtusa plantation.

A Weibull function was used to model the vertical distribution of leaf area of individual trees in a 25-year-old Chamaecyparis obtusa (Siebold & Zucc.) Endl. plantation. The parameter representing the shape of the leaf distribution was independent of tree size. A scale parameter tended to decrease with tree size suggesting a critical minimum height for retention of foliage by trees. On the basis of leaf distribution, the photosynthetic production of individual trees was estimated from the canopy photosynthetic production, which was determined from a model of canopy photosynthesis. The data indicated that the photosynthesis of a tree was proportional to the corresponding tree weight to the power of 1.84. Furthermore, the photosynthetic production varied as the 3/2nd power of total leaf area of the tree. Thus, it was concluded that the photosynthetic production per unit of leaf area, that is, the mean photosynthetic activity of a tree, is proportional to the stem girth at clear length, or the square root of the leaf area of the tree.

Journal Article↗

PubMed1994

Effects of soil water deficit on gas exchange characteristics and water relations of orchard lychee (Litchi chinensis Sonn.) trees.

Eight-year-old lychee (Litchi chinensis Sonn.) trees, cv. 'Bengal,' growing in krasnozem soil were subjected to soil water deficit from one month before flowering until harvest by covering the ground with polyethylene sheeting and withholding irrigation. The ratio of daytime stomatal conductance of unirrigated to irrigated trees decreased 20% during the three months of increasing water deficit. Predawn leaf water potentials of irrigated trees averaged about -0.3 MPa throughout the period, whereas they declined progressively to -0.9 MPa in unirrigated trees. Minimum daytime leaf water potential in the unirrigated trees decreased from -1.0 to -1.1 MPa at the beginning of the drought period to -2.2 to -2.4 MPa after three months, and calculated whole-plant conductance did not change with decreasing availability of water. The calculated soil-root water potential declined to less than -1.0 MPa in unirrigated trees. Capacitance effects on the relationship between leaf water potential and transpiration were significant only at low transpiration rates. Although unirrigated trees reduced soil water content at 0-30 cm depths to an equivalent water potential of -1.0 MPa, fruit shedding was significantly less than in irrigated trees. Water deficit had no effect on the fresh weight of pericarp, but caused increased seed size and decreased fresh weight of flesh, resulting in fruit from unirrigated trees being 16% lower in total fresh weight per fruit than fruit from irrigated trees.

Journal Article↗

PubMedDate not supplied

Sap flow in Bornean heath and dipterocarp forest trees during wet and dry periods.

Daytime xylem sap flow was measured by the compensation heat pulse velocity technique in dipterocarp forest trees growing on sandy loam (haplic acrisol) and in heath forest trees growing on bleached sand (albic arenosol) in Brunei during wet and dry periods. In dipterocarp forest trees, daily sap flow was stable during the dry period but declined on rainy days presumably as a result of reduced transpiration. At the heath site, daily sap flow of the small trees was stable during the dry period, whereas it declined in the large trees as the dry season progressed. In some trees at both sites, the hydroactive area of xylem was less during the dry period than during the wet period. Daily sap flow was linearly related to projected crown area, and a common regression fitted trees of different species from both forest types. The dry/wet period ratio of sap flow (about 0.7) was statistically equivalent for heath and dipterocarp forest trees, whereas it had been expected that trees on the freely draining arenosol would suffer greater water stress during drought. For well-watered trees >/= 0.2 m dbh (diameter at 1.3 m above ground) in both the dipterocarp and heath forests, stand water use was estimated to be 40 mm month(-1).

Journal Article↗

PubMedDate not supplied

Adaptive growth of tree root systems in response to wind action and site conditions.

Soil-root plate dimensions and structural root architecture were examined on 46-year-old Sitka spruce (Picea sitchensis (Bong.) Carr.) trees that had been mechanically uprooted. Rooting depth was restricted by a water table, and root system morphology had adapted to resist the wind movement associated with shallow rooting. The spread of the root system and the ratio of root mass to shoot mass (root/shoot ratio) were both negatively related to soil-root plate depth. Root systems had more structural root mass on the leeward side than the windward side of the tree relative to the prevailing wind direction. Cross sections of structural roots were obtained at distances of 0.5, 0.75, 1.0, and 1.25 m from the tree center. Buttressed parts of roots had greater lateral and vertical secondary thickening above rather than below the biological center. This uneven growth, which produced a shape similar in cross section to a T-beam, was greater on the leeward side of the tree, and was greatest at 0.5 m from the tree center of shallow rooted trees. Further from the tree, particularly on the windward side, many roots developed eccentric cross-sectional shapes comparable to I-beams, which would efficiently resist vertical flexing. Roots became more ovoid in shape with increasing distance from the tree, especially on deep rooted trees where lateral roots tapered rapidly to a small diameter. We conclude that these forms of adaptive growth in response to wind movement improve the rigidity of the soil-root plate and counteract the increasing vulnerability to windthrow as the tree grows.

Journal Article↗

PubMed1990

Seasonal changes in amino acids, protein and total nitrogen in needles of fertilized Scots pine trees.

Seasonal changes in amino acids, protein and total nitrogen in needles of 30-year-old, fertilized Scots pine (Pinus sylvestris L.) trees growing in Northern Sweden were investigated over two years in field experiments. The studied plots had been fertilized annually for 17 years with (i) a high level of N, (ii) a medium level of N, or (iii) a medium level of N, P and K. Trees growing on unfertilized plots served as controls. In control trees, glutamine, glutamic acid, gamma-aminobutyric acid, aspartic acid and proline represented 50-70% of the total free amino acids determined. Arginine was present only in low concentrations in control trees throughout the year, but it was usually the most abundant amino acid in fertilized trees. Glutamine concentrations were high during the spring and summer in both years of study, whereas proline concentrations were high in the spring but otherwise low throughout the year. In the first year of study, glutamic acid concentrations were high during the spring and summer, whereas gamma-aminobutyric acid was present in high concentrations during the winter months. This pattern was less pronounced in the second year of investigation. The concentrations of most amino acids, except glutamic acid, increased in response to fertilization. Nitrogen fertilization increased the foliar concentration of arginine from < 1 micromol g(dw) (-1) in control trees to a maximum of 110 micromol g(dw) (-1). Trees fertilized with nitrogen, phosphorus and potassium had significantly lower arginine concentrations than trees fertilized with the same amount of nitrogen only. Protein concentrations were similar in all fertilized trees but higher than those in control trees. For all treatments, protein concentrations were high in winter and at a minimum in early spring. In summer, the protein concentration remained almost constant except for a temporary decrease which coincided with the expansion of new shoots. Apart from arginine, the amino acid composition of proteins was similar in all treatments.

Journal Article↗

PubMed1990

Characteristics of high- and low-vigor lodgepole pine trees in old-growth stands.

Lodgepole pine (Pinus contorta var. latifolia Engelm.) trees in an old stand differed widely in appearance. Some appeared healthy, with deep crowns, high leaf areas, nearly conical-shaped crown tops, and branches with live needles extending well back from the tip. Others with similar-sized boles appeared to be in a state of decline, having much shallower crowns, lower leaf areas, crowns that were flattened at the top, and "tufted" branches with needles only near the ends of the branches. Ten trees, five each in these two apparent vigor classes, were selected from a one-hectare area of a mature stand. Trees in the two groups were found to have similar height, diameter at breast height, age, and total volume. As expected from the selection criteria, high-vigor trees had much deeper crowns (P = 0.01) and higher leaf areas (P = 0.10). The volume of heartwood in the bole was higher in the low-vigor trees (P = 0.08), and the volume of the sapwood was higher in the high-vigor trees (P = 0.13). Sapwood and heartwood ages were similar in both classes. This suggests that, in earlier years, low-vigor trees reached a larger volume, represented by the present heartwood, than did the high-vigor trees (i.e., the current vigor rankings are the reverse of what they had been). The ratio of sapwood volume to total leaf area was not significantly higher in the low-vigor trees (P = 0.24), although this ratio varied tenfold among the data. Results suggest that a physiological basis for assessing vigor involving leaf area, sapwood, heartwood, age, and incremental growth data would identify a continuum of response from high- to low-vigor trees rather than discrete groups based on visual appearance.

Journal Article↗

PubMed2000

Whole-genome trees based on the occurrence of folds and orthologs: implications for comparing genomes on different levels.

We built whole-genome trees based on the presence or absence of particular molecular features, either orthologs or folds, in the genomes of a number of recently sequenced microorganisms. To put these genomic trees into perspective, we compared them to the traditional ribosomal phylogeny and also to trees based on the sequence similarity of individual orthologous proteins. We found that our genomic trees based on the overall occurrence of orthologs did not agree well with the traditional tree. This discrepancy, however, vanished when one restricted the tree to proteins involved in transcription and translation, not including problematic proteins involved in metabolism. Protein folds unite superficially unrelated sequence families and represent a most fundamental molecular unit described by genomes. We found that our genomic occurrence tree based on folds agreed fairly well with the traditional ribosomal phylogeny. Surprisingly, despite this overall agreement, certain classes of folds, particularly all-beta ones, had a somewhat different phylogenetic distribution. We also compared our occurrence trees to whole-genome clusters based on the composition of amino acids and di-nucleotides. Finally, we analyzed some technical aspects of genomic trees-e.g., comparing parsimony versus distance-based approaches and examining the effects of increasing numbers of organisms. Additional information (e.g. clickable trees) is available from http://bioinfo.mbb.yale.edu/genome/trees.

Computational Biology↗

PubMed2000

Identification of natural rubber and characterization of rubber biosynthetic activity in fig tree.

Natural rubber was extracted from the fig tree (Ficus carica) cultivated in Korea as part of a survey of rubber producing plants. Fourier transform infrared and (13)C nuclear magnetic resonance analysis of samples prepared by successive extraction with acetone and benzene confirmed that the benzene-soluble residues are natural rubber, cis-1,4-polyisoprene. The rubber content in the latex of fig tree was about 4%, whereas the rubber content in the bark, leaf, and fruit was 0.3%, 0.1%, and 0.1%, respectively. Gel-permeation chromatography revealed that the molecular size of the natural rubber from fig tree is about 190 kD. Similar to rubber tree (Hevea brasiliensis) and guayule (Parthenium argentatum Gray), rubber biosynthesis in fig tree is tightly associated with rubber particles. The rubber transferase in rubber particles exhibited a higher affinity for farnesyl pyrophosphate than for isopentenyl pyrophosphate, with apparent K(m) values of 2.8 and 228 microM, respectively. Examination of latex serum from fig tree by sodium dodecyl sulfate-polyacrylamide gel electrophoresis revealed major proteins of 25 and 48 kD in size, and several proteins with molecular mass below 20 and above 100 kD. Partial N-terminal amino acid sequencing and immunochemical analyses revealed that the 25- and 48-kD proteins were novel and not related to any other suggested rubber transferases. The effect of EDTA and Mg(2+) ion on in vitro rubber biosynthesis in fig tree and rubber tree suggested that divalent metal ion present in the latex serum is an important factor in determining the different rubber biosynthetic activities in fig tree and rubber tree.

Cations, Divalent↗

PubMed2005

Genome trees and the nature of genome evolution.

Genome trees are a means to capture the overwhelming amount of phylogenetic information that is present in genomes. Different formalisms have been introduced to reconstruct genome trees on the basis of various aspects of the genome. On the basis of these aspects, we separate genome trees into five classes: (a) alignment-free trees based on statistic properties of the genome, (b) gene content trees based on the presence and absence of genes, (c) trees based on chromosomal gene order, (d) trees based on average sequence similarity, and (e) phylogenomics-based genome trees. Despite their recent development, genome tree methods have already had some impact on the phylogenetic classification of bacterial species. However, their main impact so far has been on our understanding of the nature of genome evolution and the role of horizontal gene transfer therein. An ideal genome tree method should be capable of using all gene families, including those containing paralogs, in a phylogenomics framework capitalizing on existing methods in conventional phylogenetic reconstruction. We expect such sophisticated methods to help us resolve the branching order between the main bacterial phyla.

Evolution, Molecular↗

PubMed2006

Uptake, removal, accumulation, and phytotoxicity of phenol in willow trees (Salix viminalis).

Phenol is an intermediate in the metabolism of plants, but it also is a high-volume production compound frequently found in the environment. The relation between removal, uptake, accumulation, and toxicity of phenol in willow trees (Salix viminalis) was determined. Willows were exposed to phenol levels up to 1000 mg/L in hydroponic solution and sand, respectively. The change on water transpiration of the trees was used to determine toxic effects. In both hydroponic solution and sand, only small inhibition of transpiration was detected up to phenol concentrations of 250 mg/L. For concentrations of 500 mg/L or greater, transpiration fell to 50% or less, and the trees wilted. The trees exposed to 1000 mg/L wilted and eventually died. For initial nominal concentrations of 1000 mg/L, a significantly higher amount of phenol remained at the end of the experiment compared to the amount remaining with all other concentrations. Phenol loss was approximately 90% for the trees that survived, except for trees exposed to the lowest concentration (10 mg/L). The loss of phenol was only 15 to 18% in experiments performed without trees. The results demonstrate a clear relation between removal, accumulation, and toxic effects of phenol in trees. Phenol levels of less than 250 mg/L could be degraded by "in planta" metabolism; therefore, no significant buildup of phenol nor any toxic effects in plants occurred. With higher levels, phenol was still degraded quickly, but an accumulation in trees accompanied by severe toxic effects occurred. Phytoremediation of phenol would be best with concentrations in water or soil solution of less than 250 mg/L, at which the degradation of phenol by willows or associated bacteria is rapid and efficient and the toxic effects on trees are negligible.

Animals↗

PubMed2006

Predicting impaired glucose metabolism in women with polycystic ovary syndrome by decision tree modelling.

AIMS/HYPOTHESIS: Polycystic ovary syndrome (PCOS) is a risk factor of type 2 diabetes. Screening for impaired glucose metabolism (IGM) with an OGTT has been recommended, but this is relatively time-consuming and inconvenient. Thus, a strategy that could minimise the need for an OGTT would be beneficial. MATERIALS AND METHODS: Consecutive PCOS patients (n=118) with fasting glucose <6.1 mmol/l were included in the study. Parameters derived from medical history, clinical examination and fasting blood samples were assessed by decision tree modelling for their ability to discriminate women with IGM (2-h OGTT value >/=7.8 mmol/l) from those with NGT. RESULTS: According to the OGTT results, 93 PCOS women had NGT and 25 had IGM. The best decision tree consisted of HOMA-IR, the proinsulin:insulin ratio, proinsulin, 17-OH progesterone and the ratio of luteinising hormone:follicle-stimulating hormone. This tree identified 69 women with NGT. The remaining 49 women included all women with IGM (100% sensitivity, 74% specificity to detect IGM). Pruning this tree to three levels still identified 53 women with NGT (100% sensitivity, 57% specificity to detect IGM). Restricting the data matrix used for tree modelling to medical history and clinical parameters produced a tree using BMI, waist circumference and WHR. Pruning this tree to two levels separated 27 women with NGT (100% sensitivity, 29% specificity to detect IGM). The validity of both trees was tested by a leave-10%-out cross-validation. CONCLUSIONS/INTERPRETATION: Decision trees are useful tools for separating PCOS women with NGT from those with IGM. They can be used for stratifying the metabolic screening of PCOS women, whereby the number of OGTTs can be markedly reduced.

Adult↗

PubMed2005

Association between tree-ring and needle delta13C and leaf gas exchange in Pinus halepensis under semi-arid conditions.

Associations between delta13C values and leaf gas exchanges and tree-ring or needle growth, used in ecophysiological compositions, can be complex depending on the relative timing of CO2 uptake and subsequent redistribution and allocation of carbon to needle and stem components. For palaeoenvironmental and dendroecological studies it is often interpreted in terms of a simple model of delta13C fractionation in C3 plants. However, in spite of potential complicating factors, few studies have actually examined these relationships in mature trees over inter- and intra-annual time-scales. Here, we present results from a 4 years study that investigated the links between variations in leaf gas-exchange properties, growth, and dated delta13C values along the needles and across tree rings of Aleppo pine trees growing in a semi-arid region under natural conditions or with supplemental summer irrigation. Sub-sections of tissue across annual rings and along needles, for which time of formation was resolved from growth rate analyses, showed rapid growth and delta13C responses to changing environmental conditions. Seasonal cycles of growth and delta13C (up to approximately 4 per thousand) significantly correlated (P < 0.01) with photosynthetically active radiation, vapour pressure deficit, air temperature, and soil water content. The irrigation significantly increased leaf net assimilation, stomatal conductance and needle and tree-ring growth rate, and markedly decreased needle and tree-ring delta13C values and its sensitivity to environmental parameters. The delta13C estimates derived from gas-exchange parameters, and weighted by assimilation, compared closely with seasonal and inter-annual delta13C values of needle- and tree-ring tissue. Higher stomatal conductances of the irrigated trees (0.22 vs. 0.08 mol m(-2) s(-1) on average) corresponded with approximately 2.0 per thousand lower average delta13C values, both measured and derived. Derived and measured delta13C values also indicated that needle growth, which occurs throughout the stressful summer was supported by carbon from concurrent, low rate assimilation. For Aleppo pine under semi-arid and irrigated conditions, the delta13C of tree-ring and needle material proved, in general, to be a reasonable indicator of integrated leaf gas-exchange properties.

Carbon Dioxide↗