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Phylogenetic trees: visualizing, customizing and detecting incongruence.

SUMMARY: TreeIllustrator is a user-friendly application to visualize and customize phylogenetic trees. It has a broad range of functions and capabilities, such as dragging of nodes, different tree shapes, zooming and searching capabilities, and support for large trees. It acts as a solution that integrates the specificity of visualizing phylogenetic trees and the customization options of a drawing program. It also contains a simple and effective method that compares a custom tree with the Tree of Life, by detecting incongruence.

Chromosome Mapping↗

Improving the efficiency of SPR moves in phylogenetic tree search methods based on maximum likelihood.

MOTIVATION: Maximum likelihood (ML) methods have become very popular for constructing phylogenetic trees from sequence data. However, despite noticeable recent progress, with large and difficult datasets (e.g. multiple genes with conflicting signals) current ML programs still require huge computing time and can become trapped in bad local optima of the likelihood function. When this occurs, the resulting trees may still show some of the defects (e.g. long branch attraction) of starting trees obtained using fast distance or parsimony programs. METHODS: Subtree pruning and regrafting (SPR) topological rearrangements are usually sufficient to intensively search the tree space. Here, we propose two new methods to make SPR moves more efficient. The first method uses a fast distance-based approach to detect the least promising candidate SPR moves, which are then simply discarded. The second method locally estimates the change in likelihood for any remaining potential SPRs, as opposed to globally evaluating the entire tree for each possible move. These two methods are implemented in a new algorithm with a sophisticated filtering strategy, which efficiently selects potential SPRs and concentrates most of the likelihood computation on the promising moves. RESULTS: Experiments with real datasets comprising 35-250 taxa show that, while indeed greatly reducing the amount of computation, our approach provides likelihood values at least as good as those of the best-known ML methods so far and is very robust to poor starting trees. Furthermore, combining our new SPR algorithm with local moves such as PHYML's nearest neighbor interchanges, the time needed to find good solutions can sometimes be reduced even more.

Algorithms↗

Rates of population increase, abundance, and life stage distribution of Rhynchites cribripennis (Coleoptera: Attelabidae) on trees and in the soil in an olive grove.

The seasonal population abundance of Rhynchites cribripennis (Desbrochers) adults on olive trees was studied by collecting samples from an olive grove on the island of Zakynthos, Greece, from April 1994 to the end of July 1995. Moreover, the population abundance of larvae, pupae, and adults of R. cribripennis was recorded in soil samples from two soil depths (0-4 and 4-8 cm) from October 1994 to October 1995. Results showed that adult populations increased considerably on trees during May and June and peaked on 16 June (19.9 adults per twig) and 8 July (7.7 adults per twig) in 1994 and 1995, respectively. In 1994, no significant differences were found in the number of adults sampled from the different tree quadrants (northwest [NW], northeast [NE], southwest [SW], and southeast [SE]). However, in 1995, adult numbers in the NW quadrant of trees were significantly higher than those in the SE or SW quadrants. In soil samples, larvae were recorded throughout the sampling period with the highest numbers occurring in December, 2.4 larvae per soil sample, whereas pupae were found in lowest numbers in October and November. Adults were present in the soil samples from December to May, but the highest numbers were recorded in December, with a peak of one adult per sample. The number of adults was significantly higher, and that of larvae numerically higher, in the upper compared with lower soil layer, whereas pupae were found with similar numbers in both soil layers. Results of these studies suggest that this weevil exhibits a prolonged larval diapause and a 2-yr life cycle. The ecological implications of this behavior are discussed. Moreover, a prediction of the highest adult population on trees was estimated by taking into account the rate of increase of adult numbers in the early period of adult occurrence on trees.

Animals↗

Evaluation of biological and biorational control tactics for suppression of Nantucket pine tip moth damage in Virginia pine Christmas trees.

A study was performed to evaluate the potential of the egg parasitoid Trichogramma exiguum Pinto and Platner (Hymenoptera: Trichogrammatidae), the newly registered insect growth regulator (IGR) tebufenozide (Confirm), and a modified spray technique (top whorl only pesticide application) for suppression of Nantucket pine tip moth, Rhyacionia frustrana (Comstock) (Lepidoptera: Tortricidae), damage in Virginia pine, Pinus virginiana Mill., Christmas trees. Augmentative releases of T. exiguum failed to increase parasitism levels in release plots compared with controls, and significant reduction in tip moth damage did not occur. High predation levels on released T. exiguum may have contributed to the failure of parasitoid augmentations. Whole-tree and top whorl tebufenozide treatments provided significantly greater damage control than corresponding applications of acephate (Orthene), a commonly used pesticide in Christmas trees. Damage to trees receiving whole-tree chemical applications did not differ significantly from trees receiving top whorl treatments for most measurements of damage. Top whorl chemical treatments resulted in a 67% reduction in time required for application and a 70% reduction in pesticide used.

Animals↗

Theoretical foundation of the balanced minimum evolution method of phylogenetic inference and its relationship to weighted least-squares tree fitting.

Due to its speed, the distance approach remains the best hope for building phylogenies on very large sets of taxa. Recently (R. Desper and O. Gascuel, J. Comp. Biol. 9:687-705, 2002), we introduced a new "balanced" minimum evolution (BME) principle, based on a branch length estimation scheme of Y. Pauplin (J. Mol. Evol. 51:41-47, 2000). Initial simulations suggested that FASTME, our program implementing the BME principle, was more accurate than or equivalent to all other distance methods we tested, with running time significantly faster than Neighbor-Joining (NJ). This article further explores the properties of the BME principle, and it explains and illustrates its impressive topological accuracy. We prove that the BME principle is a special case of the weighted least-squares approach, with biologically meaningful variances of the distance estimates. We show that the BME principle is statistically consistent. We demonstrate that FASTME only produces trees with positive branch lengths, a feature that separates this approach from NJ (and related methods) that may produce trees with branches with biologically meaningless negative lengths. Finally, we consider a large simulated data set, with 5,000 100-taxon trees generated by the Aldous beta-splitting distribution encompassing a range of distributions from Yule-Harding to uniform, and using a covarion-like model of sequence evolution. FASTME produces trees faster than NJ, and much faster than WEIGHBOR and the weighted least-squares implementation of PAUP*. Moreover, FASTME trees are consistently more accurate at all settings, ranging from Yule-Harding to uniform distributions, and all ranges of maximum pairwise divergence and departure from molecular clock. Interestingly, the covarion parameter has little effect on the tree quality for any of the algorithms. FASTME is freely available on the web.

Algorithms↗

Bayesian estimation of concordance among gene trees.

Multigene sequence data have great potential for elucidating important and interesting evolutionary processes, but statistical methods for extracting information from such data remain limited. Although various biological processes may cause different genes to have different genealogical histories (and hence different tree topologies), we also may expect that the number of distinct topologies among a set of genes is relatively small compared with the number of possible topologies. Therefore evidence about the tree topology for one gene should influence our inferences of the tree topology on a different gene, but to what extent? In this paper, we present a new approach for modeling and estimating concordance among a set of gene trees given aligned molecular sequence data. Our approach introduces a one-parameter probability distribution to describe the prior distribution of concordance among gene trees. We describe a novel 2-stage Markov chain Monte Carlo (MCMC) method that first obtains independent Bayesian posterior probability distributions for individual genes using standard methods. These posterior distributions are then used as input for a second MCMC procedure that estimates a posterior distribution of gene-to-tree maps (GTMs). The posterior distribution of GTMs can then be summarized to provide revised posterior probability distributions for each gene (taking account of concordance) and to allow estimation of the proportion of the sampled genes for which any given clade is true (the sample-wide concordance factor). Further, under the assumption that the sampled genes are drawn randomly from a genome of known size, we show how one can obtain an estimate, with credibility intervals, on the proportion of the entire genome for which a clade is true (the genome-wide concordance factor). We demonstrate the method on a set of 106 genes from 8 yeast species.

Algorithms↗

Four new mitochondrial genomes and the increased stability of evolutionary trees of mammals from improved taxon sampling.

We have sequenced four new mitochondrial genomes to improve the stability of the tree for placental mammals; they are two insectivores (a gymnure, Echinosorex gymnurus and Formosan shrew Soriculus fumidus); a Formosan lesser horseshoe bat (Rhinolophus monoceros); and the New Zealand fur seal (Arctocephalus forsteri). A revision to the hedgehog sequence (Erinaceus europaeus) is also reported. All five are from the Laurasiatheria grouping of eutherian mammals. On this new data set there is a strong tendency for the hedgehog and its relative, the gymnure, to join with the other Laurasiatherian insectivores (mole and shrews). To quantify the stability of trees from this data we define, based on nuclear sequences, a major four-way split in Laurasiatherians. This ([Xenarthra, Afrotheria], [Laurasiatheria, Supraprimates]) split is also found from mitochondrial genomes using either protein-coding or RNA (rRNA and tRNA) data sets. The high similarity of the mitochondrial and nuclear-derived trees allows a quantitative estimate of the stability of trees from independent data sets, as detected from a triplet Markov analysis. There are significant changes in the mutational processes within placental mammals that are ignored by current tree programs. On the basis of our quantitative results, we expect the evolutionary tree for mammals to be resolved quickly, and this will allow other problems to be solved.

Animals↗

Methods for computing the standard errors of branching points in an evolutionary tree and their application to molecular data from humans and apes.

Statistical methods for computing the standard errors of the branching points of an evolutionary tree are developed. These methods are for the unweighted pair-group method-determined (UPGMA) trees reconstructed from molecular data such as amino acid sequences, nucleotide sequences, restriction-sites data, and electrophoretic distances. They were applied to data for the human, chimpanzee, gorilla, orangutan, and gibbon species. Among the four different sets of data used, DNA sequences for an 895-nucleotide segment of mitochondrial DNA (Brown et al. 1982) gave the most reliable tree, whereas electrophoretic data (Bruce and Ayala 1979) gave the least reliable one. The DNA sequence data suggested that the chimpanzee is the closest and that the gorilla is the next closest to the human species. The orangutan and gibbon are more distantly related to man than is the gorilla. This topology of the tree is in agreement with that for the tree obtained from chromosomal studies and DNA-hybridization experiments. However, the difference between the branching point for the human and the chimpanzee species and that for the gorilla species and the human-chimpanzee group is not statistically significant. In addition to this analysis, various factors that affect the accuracy of an estimated tree are discussed.

Amino Acid Sequence↗

Seasonal patterns of radial root growth and starch dynamics in plantation-grown Sitka spruce trees of different ages.

Seasonal patterns of radial root growth within 1 m of tree stems were examined in Scottish plantations of Sitka spruce trees aged 9, 15 and 20 years. Results were compared with parallel measurements of shoot extension, radial growth of stems and amounts of starch stored in tissues external to root wood. Youngest trees produced the largest annual increments in root cross-sectional area and numbers of new cells along radial files of tracheids. Irrespective of tree age, new cells were present in roots before bud burst and the onset of radial growth occurred progressively later with increasing distances from the stems. At ages 15 and 20, both stem cross-sectional area and radial root growth up to 0.5 m from the stem base had a minor peak of activity preceding and a major peak following shoot elongation. Further than 0.5 m from the stem, root growth was frequently restricted to the period following shoot extension. Starch storage in the roots reached a maximum in April and May, which was greatest for 9-year-old trees and least for 20-year-old trees. At all ages, radial root growth in early spring occurred concurrently with increased starch storage. Later in the season starch reserves declined rapidly during the period of shoot elongation and root growth occurred whilst reserves were low. At all ages for positions on the root at the base of the stem and 0.25 m from it, starch depletion, at its maximum rate during June, accounted for less than the measured increment of root wood growth at that point. This indicates a substantial translocation of substrates to these zones during growth. At the same time, the reduction in starch concentrations at more distal points from the stem far exceeded that required for local root thickening.

Journal Article↗

Patterns in aboveground carbon allocation and tree architecture that favor stem growth in young Scots pine from high latitudes.

The production and allocation of aboveground biomass and the characteristics of tree architecture were examined in eight-year-old Scots pine (Pinus sylvestris L.). Considerable among-tree variation existed in tree architecture, total aboveground dry mass production, and dry mass partitioning among tree parts. A linear relationship existed between needle and branch mass. Stem mass was directly proportional to tree height, which in turn was directly proportional to the allocation ratio between stem mass and total needle + branch mass production. The architectural characteristics that were related to a high proportional allocation to stem and high stemwood production were a large mean shoot volume, large mean number of branches per whorl, long needle retention and a high crown length/crown width ratio. Individual trees were found that combined high stemwood production with both high harvest index and high stemwood specific gravity.

Journal Article↗

Water use efficiency of short-rotation Salix viminalis at leaf, tree and stand scales.

Fluxes of water and carbon dioxide from short-rotation Salix viminalis L. were measured at leaf, tree and stand scales using porometer, chamber and micrometeorological techniques, respectively. At all three scales, instantaneous water use efficiency exhibited a similar dependence on vapor pressure deficit (deltae) with values of 21, 13 and 10 g CO(2) per kg H(2)O for deltae of 0.5, 1.0 and 1.5 kPa, respectively. The fluxes of CO(2) and H(2)O plotted against incident light above the stand differed in magnitude according to scale, with highest and lowest values being obtained for leaf- and stand-level measurements, respectively. At the tree level, the fluxes of CO(2) and H(2)O reflected the social position of the sample trees with the dominant tree having higher values than the subdominant tree. Average long-term water use efficiency, estimated from stand-level measurements, was 6.3 g dry biomass per kg of transpired water. This value is high compared with values for other tree species and may be associated with a high foliar nitrogen concentration. We conclude that water availability will be a critical factor in short-rotation willow forestry despite the relatively high water use efficiency of the species.

Journal Article↗

Storage of foliar-absorbed nitrogen and remobilization for spring growth in young nectarine (Prunus persica var. nectarina) trees.

The effectiveness of spraying foliage with urea to provide nitrogen (N) to augment the seasonal internal cycling of N in young nectarine trees (Prunus persica (L.) Batsch var. nectarina (Ait. f. Maxim.), cv. Stark Red Gold) was studied. One-year-old trees were grown with contrasting N supplies during the summer and foliage was sprayed with a 2% urea solution labeled with (15)N just before leaf senescence started. After leaf abscission had finished, the trees were repotted in sand and given no further N. Remobilization of both labeled and unlabeled N for leaf growth the following spring was quantified. Leaves absorbed between 58 and 69% of the (15)N intercepted by the canopy irrespective of tree N status. During leaf senescence, the majority of (15)N was withdrawn from the leaves into the shoot and roots. Remobilization of (15)N the following spring was also unaffected by tree N status. About 38-46% of (15)N in the trees was recovered in the new growth. More unlabeled N (derived from root uptake) was remobilized for leaf growth in the spring than was withdrawn from leaves during canopy senescence the previous autumn. Therefore, soil-applied N augmented N storage pools directly, and contributed more to N remobilization the following spring than did foliar-absorbed (15)N.

Journal Article↗

Estimating stem and root-anchorage flexibility in trees.

This paper describes a nondestructive method for distinguishing root flexibility from stem flexibility in living trees. It is used here for Sitka spruce (Picea sitchensis (Bong.) Carrière.), but is applicable to any species where the main stem is normally straight and near-vertical. Well-known engineering equations permit the calculation of deflected shape for a vertical cantilever with arbitrary distribution of mass and bending stiffness, when infjected to a lateral force. The equations are used to calculate stem deflections of four Sitka spruce trees for which the stem and branch mass distribution and stem taper have been measured. Free parameters in the mathematical model are a nominal value of Young's Modulus E (assumed uniform and isotropic over the cross section and height of the tree stem) and a root-anchorage stiffness k. The former allows the stem to curve, whereas the latter represents the flexibility of the roots and allows the stem to tilt elastically at ground level. For each of the four trees, the calculated deflection curve is compared with actual deflections measured when the living tree is pulled by a rope at a specified point. By adjusting both E and k, iteratively, a best fit solution is obtained. This provides a simple and effective way to determine both stem stiffness and root hinge stiffness from a single experiment on a living tree.

Journal Article↗

Radial variation in sap flow in five laurel forest tree species in Tenerife, Canary Islands.

Variations in radial patterns of xylem water content and sap flow rate were measured in five laurel forest tree species (Laurus azorica (Seub.) Franco, Persea indica (L.) Spreng., Myrica faya Ait., Erica arborea L. and Ilex perado Ait. ssp. platyphylla (Webb & Berth.) Tutin) growing in an experimental plot at Agua García, Tenerife, Canary Islands. Measurements were performed around midday during warm and sunny days by the heat field deformation method. In all species, water content was almost constant (around 35% by volume) over the whole xylem cross-sectional area. There were no differences in wood color over the whole cross-sectional area of the stem in most species with the exception of E. arborea, whose wood became darker in the inner layers. Radial patterns of sap flow were highly variable and did not show clear relationships with tree diameter or species. Sap flow occurred over the whole xylem cross-sectional area in some species, whereas it was limited to the outer xylem layers in others. Sap flow rate was either similar along the xylem radius or exhibited a peak in the outer part of the xylem area. Low sap flow rates with little variation in radial pattern were typical for shaded suppressed trees, whereas dominant trees exhibited high sap flow rates with a peak in the radial pattern. Stem damage resulted in a significant decrease in sap flow rate in the outer xylem layers. The outer xylem is more important for whole tree water supply than the inner xylem because of its larger size. We conclude that measurement of radial flow pattern provides a reliable method of integrating sap flow from individual measuring points to the whole tree.

Journal Article↗

Tree water use and rainfall partitioning in a mature poplar-pasture system.

Traditionally, poplars (Populus) have been planted to control erosion on New Zealand's hill-slopes, because of their capacity to dry out and bind together the soil, by reducing effective rainfall and increasing evapotranspiration and soil strength. However, the effect of widely spaced poplars on the partitioning of soil water and rainfall has not been reported. This study determined rainfall partitioning for 18 mid-spring days in a mature P. deltoides (Bart. ex Marsh, Clone I78)-pasture association (37 stems per hectare, unevenly spaced at 16.4 +/- 0.4 m) and compared it with a traditional open pasture system in grazed areas of a hill environment. Tree transpiration was measured by the heat pulse technique. A time-driven mathematical model was used to set a zero offset, adjust anomalous values and describe simultaneous sap velocity time courses of trees. The model showed that daylight sap flow velocities can be represented with a nonlinear Beta function (R(2) > 0.98), and differences in the parameters representing the initiation, duration and conformation of the sap velocity can be tested statistically to discern tree transpiration differences during the day. Evapotranspiration was greater for the poplar-pasture association than for the open pasture (2.7-3.0 versus 2.2 mm day(-1)). The tree canopy alone contributed 0.92 mm day(-1) as transpiration and 1.37 mm day(-1) as interception, whereas evapotranspiration of the pasture understory was only 0.4-0.6 mm day(-1). Despite the higher water use of the poplar-pasture association, soil water in the 0-300 mm soil stratum was higher than, or similar to, that of the open pasture. Tree shading decreased evapotranspiration and pasture accumulation under the trees.

Journal Article↗

Distribution of leaf mass per unit area and leaf nitrogen concentration determine partitioning of leaf nitrogen within tree canopies.

Distribution of leaf nitrogen with respect to leaf mass per unit area (M(a)), nitrogen per unit mass (N(m)) and nitrogen per unit area (N(a)) within peach (Prunus persica L.) tree canopies was studied in two field experiments. In one experiment, leaf light exposure and M(a) were measured on leaves from different canopy positions of peach trees subjected to five nitrogen (N) fertilization treatments. Leaf light exposure and M(a) were linearly related and the relationship was independent of N fertilization. In a subsequent experiment, N fertilizer was applied to previously unfertilized trees in midsummer, after shoot growth had terminated. Application of N fertilizer did not affect mean canopy M(a). Fertilization increased N(m) of all leaves throughout the canopy compared with non-fertilized trees. No significant relationship between N(m) and M(a) was found in either fertilized or control trees. There was a linear relationship between N(a) and M(a) and the slope of the relationship was increased by N fertilizer application. We conclude that distribution of N(a) in peach tree canopies is primarily a function of M(a) partitioning with light and N(m), which is related to soil N availability.

Journal Article↗

Radial variation in sap velocity as a function of stem diameter and sapwood thickness in yellow-poplar trees.

Canopy transpiration and forest water use are frequently estimated as the product of sap velocity and cross-sectional sapwood area. Few studies, however, have considered whether radial variation in sap velocity and the proportion of sapwood active in water transport are significant sources of uncertainty in the extrapolation process. Therefore, radial profiles of sap velocity were examined as a function of stem diameter and sapwood thickness for yellow-poplar (Liriodendron tulipifera L.) trees growing on two adjacent watersheds in eastern Tennessee. The compensation heat pulse velocity technique was used to quantify sap velocity at four equal-area depths in 20 trees that ranged in stem diameter from 15 to 69 cm, and in sapwood thickness from 2.1 to 14.8 cm. Sap velocity was highly dependent on the depth of probe insertion into the sapwood. Rates of sap velocity were greatest for probes located in the two outer sapwood annuli (P1 and P2) and lowest for probes in closest proximity to the heartwood (P3 and P4). Relative sap velocities averaged 0.98 at P1, 0.66 at P2, 0.41 at P3 and 0.35 at P4. Tree-specific sap velocities measured at each of the four probe positions, divided by the maximum sap velocity measured (usually at P1 or P2), indicated that the fraction of sapwood functional in water transport (f(S)) varied between 0.49 and 0.96. There was no relationship between f(S) and sapwood thickness, or between f(S) and stem diameter. The fraction of functional sapwood averaged 0.66 +/- 0.13 for trees on which radial profiles were determined. No significant depth-related differences were observed for sapwood density, which averaged 469 kg m(-3) across all four probe positions. There was, however, a significant decline in sapwood water content between the two outer probe positions (1.04 versus 0.89 kg kg(-1)). This difference was not sufficient to account for the observed radial variation in sap velocity. A Monte-Carlo analysis indicated that the standard error in estimated mean f(S) declined rapidly with increasing sample size. At n = 10, the coefficient of variation in mean f(S) was 7% and at n = 15 it was slightly less than 5%. These observations indicate that radial variation in sap velocity is an important, albeit often overlooked, source of uncertainty in the scaling process. Failure to recognize that not all sapwood is functional in water transport will introduce systematic bias into estimates of both tree and stand water use. Future studies should devise sampling strategies for assessing radial variation in sap velocity and such strategies should be used to identify the magnitude of this variation in a range of non-, diffuse- and ring-porous trees.

Journal Article↗

Nitrogen availability, local light regime and leaf rank effects on the amount and sources of N allocated within the foliage of young walnut (Juglans nigra x regia) trees.

Early season leaf growth depends largely on nitrogen (N) provided by remobilization from storage, and many studies have tested the effect of N availability to roots on the amount of N provided for new leaf development by remobilization. Although it is well known that the light regime experienced by a leaf influences the amount of N per unit leaf area (LA), the effect of the local light regime on the amount of N derived either directly from root uptake or from remobilization for early season leaf growth has never been tested at an intra- canopy scale. The objective of this study was to quantify the relative importance of (1) N availability to roots, (2) local light regime experienced by the foliage (at the shoot scale) and (3) leaf rank along the shoot, on the total amount of N allocated to leaves and on the proportions of N provided by remobilization and root uptake. To quantify the importance of N uptake and remobilization as sources of leaf N, potted hybrid walnut trees (Juglans nigra L. x regia L.) were grown outdoors in sand and fed with a labeled ((15)N) nutrient solution. By removing the apical bud, the trees were manipulated to produce only two shoots. The experimental design had two factors: (1) high (HN; 8 mol N m(-3)) and low (LN; 2 mol N m(-3)) N availability; and (2) high (HL; 90% of incident photosynthetically active photon flux (PPF)) and low (LL; 10% of incident PPF) light. Total leaf N per tree was unaffected by either N availability or irradiance. The HN treatment increased the amount of leaf N derived from root uptake at the whole-tree scale (typically around 8 and 2% in the HN and LN treatments, respectively). Nitrogen allocation within foliage of individual trees was controlled by the local light regime, which strongly affected individual leaf characteristics as leaf mass per unit LA and area- based amount of leaf (N(a)). Decreasing the light availability to a branch decreased the amount of N allocated to it, benefiting the less shaded branches. In contrast, shading of the lower branch did not affect the fraction of total leaf N remobilized for either the lower, shaded branch or the upper, unshaded branch. The relevance of these findings for tree growth modeling is discussed.

Biological Transport↗