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Forestry's fertile crescent: the application of biotechnology to forest trees.

Relative to crop plants, the domestication of forest trees is still in its infancy. For example, the domestication of many crop plants was initiated some 10,000 years ago in the so-called 'Fertile Crescent' of the Middle East. By contrast, the domestication of forest trees for the purposes of producing more fibre began in earnest in the last half century. The application of biotechnology to forest trees offers a great potential to hasten the pace of tree improvement for desirable end uses. This review outlines some of the progress that has been made in the application of biotechnology to forest trees, and considers the prospects for biotechnologically based tree improvement in the future.

Journal Article↗

Phytotoxicity of cyanide to weeping willow trees.

BACKGROUND: Cyanide is found predominantly in industrial effluents generated by metallurgical operations. It is an extremely toxic compound, so that problems and catastrophic accidents have recently occurred all around the globe. The goal of this study was to determine the toxicity of cyanide to a Chinese willow species, and to determine the removal capacity. METHODS: The toxicity of potassium cyanide (KCN) to weeping willow trees (Salix babylonica L.) was tested. The normalized, relative transpiration of the plants was used to determine the phytotoxicity of cyanide. The cyanide removal capacity of weeping willows was also determined. RESULTS AND DISCUSSION: In hydroponic solution, no chlorosis of leaves and only a small reduction in normalized relative transpiration was observed when weeping willows were exposed to low doses of cyanide (< or = 0.93 mg CN/L). Severe signs of toxicity were found for the treatment groups exposed to higher doses of cyanide (> or = 9.3 mg CN/L). Weeping willows grown in sandy soils survived the entire period (216 hours) without any toxic effect when irrigated with low doses of cyanide (3.72 mg CN/L). High doses of cyanide (> or = 18.6 mg CN/L) in irrigation water were fatal for the weeping willows within 216 hours. EC50 values for a 50% inhibition of the transpiration of the trees were estimated to be between 3.27 and 8.23 mg CN/L, depending on the duration of the exposure. CONCLUSIONS: The results obtained for the Chinese willow species Salix babylonica were very similar to those obtained for the European species S. viminalis in earlier studies. Phytotoxic effects were only found at high doses of cyanide. A large proportion of applied cyanide was removed from the contaminated media in the presence of weeping willows. This gives rise to the conclusion that the metabolism of cyanide by weeping willows is possible. RECOMMENDATIONS AND OUTLOOK: Cyanide elimination with trees seems to be a feasible option for cleaning soils and water contaminated with cyanide. A full-scale treatment has been installed in Denmark. For phytoremediation projects in China, weeping willow could be a suitable species. The tree can tolerate and remove cyanide, and it is a native Chinese species. Besides, the tree is of outstanding beauty and is planted as a common park tree in many parts of the world.

Biodegradation, Environmental↗

Bootstrap hypothesis tests for evolutionary trees and other dendrograms.

The bootstrap computer-intensive statistical technique is frequently applied to statistical analyses of phylogenetic trees. The widely used rule that a group is supported significantly if it appears in at least 95% of bootstrap trees is conservative in most situations. This paper describes three ways of using the bootstrap to carry out statistical inference on phylogenies. The first method tests whether there is nonrandom support for a single group or tree. The second method compares the support for two groups or trees. The third method tests whether a single group or tree has better support than the set of all possible alternatives; this may be a replacement for the "95% rule." These tests generally require fewer bootstrap trees to be estimated than do other methods of bootstrapping phylogenies. A simple, sequential statistical method can be used to increase the efficiency further. These methods can be applied to tests of multiple hypotheses about a single phylogeny. Parsimony analyses of 5S rRNA sequences of plants and cluster analyses of randomly amplified polymorphic DNA bands in three pathotypes of the cereal eyespot fungus are used as illustrative examples. The tests can be used to analyze dendrograms in subjects other than taxonomy.

Base Sequence↗

A discrete Fourier analysis for evolutionary trees.

Discrete Fourier transformations have recently been developed to model the evolution of two-state characters (the Cavender/Farris model). We report here the extension of these transformations to provide invertible relationships between a phylogenetic tree T (with three probability parameters of nucleotide substitution on each edge corresponding to Kimura's 3ST model) and the expected frequencies of the nucleotide patterns in the sequences. We refer to these relationships as spectral analysis. In either model with independent and identically distributed site substitutions, spectral analysis allows a global correction for all multiple substitutions (second- and higher-order interactions), independent of any particular tree. From these corrected data we use a least-squares selection procedure, the closest tree algorithm, to infer an evolutionary tree. Other selection criteria such as parsimony or compatibility analysis could also be used; each of these criteria will be statistically consistent for these models. The closest tree algorithm selects a unique best-fit phylogenetic tree together with independent edge length parameters for each edge. The method is illustrated with an analysis of some primate hemoglobin sequences.

Animals↗

Species richness and resource availability: a phylogenetic analysis of insects associated with trees.

The data on the number of species of insects associated with various trees in Britain have been reanalyzed to factor out possible bias from phylogenetic effects. It was found that tree availability (range and abundance) continues to provide a good predictor (r = 0. 852) of insect-species richness, slightly better than straightforward cross-species analyses. Of the two components of tree availability, tree abundance gives a much better prediction than tree range. The species richness on trees of major taxa with similar trophic habits (Lepidoptera and Hymenoptera/Symphyta and the two suborders of the Homoptera-Auchenorrhyncha and Sternorrhyncha) shows positive correlations; there is thus no evidence of competitive exclusion at this taxonomic level.

Journal Article↗

Frequentist properties of Bayesian posterior probabilities of phylogenetic trees under simple and complex substitution models.

What does the posterior probability of a phylogenetic tree mean?This simulation study shows that Bayesian posterior probabilities have the meaning that is typically ascribed to them; the posterior probability of a tree is the probability that the tree is correct, assuming that the model is correct. At the same time, the Bayesian method can be sensitive to model misspecification, and the sensitivity of the Bayesian method appears to be greater than the sensitivity of the nonparametric bootstrap method (using maximum likelihood to estimate trees). Although the estimates of phylogeny obtained by use of the method of maximum likelihood or the Bayesian method are likely to be similar, the assessment of the uncertainty of inferred trees via either bootstrapping (for maximum likelihood estimates) or posterior probabilities (for Bayesian estimates) is not likely to be the same. We suggest that the Bayesian method be implemented with the most complex models of those currently available, as this should reduce the chance that the method will concentrate too much probability on too few trees.

Animals↗

Exploring rate variation among and within sites in a densely sampled tree: species level phylogenetics of north american tiger beetles (genus cicindela).

Species-level phylogenetic studies require fast-evolving nucleotide positions to resolve relationships among close relatives, but these sites may be highly homoplastic and perhaps uninformative or even misleading deeper in the tree. Here we describe a species-level analysis of tiger beetles in the genus Cicindela (Coleoptera: Cicindelidae) for 132 terminal taxa and 1897 nucleotide positions from three regions of mtDNA, comprising 75% coverage of species occurring in North America. Evenly weighted parsimony analysis recovered four major clades representing radiations confined to North and Central America. Relationships near the tips were well supported but signal was contradictory at deeper nodes. Two major categories (3rd positions and all others) can be distinguished in likelihood analysis of character variation, of which only the fast-changing 3rd position characters were affected by saturation. However, their downweighting under a variety of criteria did not improve the tree topology at basal nodes. There was weak conflict between 3rd and non-3rd position characters deep in the tree, but support levels declined towards the root for all categories, even on trees that were reconstructed from 3rd and non-3rd positions separately. Statistical analysis of parsimony-based character transitions along branches showed a largely homogeneous distribution of change along the root-to-tip axis. The comparison of character transitions among the four major portions of the tree revealed deviations from stochastic distribution for the non-3rd positions, but not for 3rd positions. Hence, variability of functionally constrained non-3rd positions differs between clades and may be dependent on the character states at other sites, consistent with the covarion model of molecular evolution. The results suggest that some properties of 3rd positions are less problematic for phylogenetic reconstruction than other categories despite their high total homoplasy. In densely sampled data sets of closely related species, the disadvantages of weighting schemes according to homoplasy levels outweigh the benefits, showing the difficulty of devising meaningful weighting schemes that are applicable universally throughout the tree.

Animals↗

A geometric approach to tree shape statistics.

This article presents a new way to quantify the descriptive ability of tree shape statistics. Where before, tree shape statistics were chosen by their ability to distinguish between macroevolutionary models, the resolution presented in this paper quantifies the ability of a statistic to differentiate between similar and different trees. This is termed the geometric approach to differentiate it from the model-based approach previously explored. A distinct advantage of this perspective is that it allows evaluation of multiple tree shape statistics describing different aspects of tree shape. After developing the methodology, it is applied here to make specific recommendations for a suite of three statistics that may prove useful in applications. The article ends with an application of the statistics to clarify the impact of taxa omission on tree shape.

Classification↗

Multipolar consensus for phylogenetic trees.

Collections of phylogenetic trees are usually summarized using consensus methods. These methods build a single tree, supposed to be representative of the collection. However, in the case of heterogeneous collections of trees, the resulting consensus may be poorly resolved (strict consensus, majority-rule consensus, ...), or may perform arbitrary choices among mutually incompatible clades, or splits (greedy consensus). Here, we propose an alternative method, which we call the multipolar consensus (MPC). Its aim is to display all the splits having a support above a predefined threshold, in a minimum number of consensus trees, or poles. We show that the problem is equivalent to a graph-coloring problem, and propose an implementation of the method. Finally, we apply the MPC to real data sets. Our results indicate that, typically, all the splits down to a weight of 10% can be displayed in no more than 4 trees. In addition, in some cases, biologically relevant secondary signals, which would not have been present in any of the classical consensus trees, are indeed captured by our method, indicating that the MPC provides a convenient exploratory method for phylogenetic analysis. The method was implemented in a package freely available at http://www.lirmm.fr/~cbonnard/MPC.html

Algorithms↗

On the design of the coronary arterial tree: a generalization of Murray's law.

Murray's law has been generalized to provide morphometric relationships among various subtrees as well as between a feeding segment and the subtree it perfuses. The equivalent resistance of each subtree is empirically determined to be proportional to the cube of a subtree's cumulative arterial length (L) and inversely proportional to a subtree's arterial volume (V) raised to a power of approximately 2.6. This relationship, along with a minimization of a cost function, and a linearity assumption between flow and cumulative arterial length, provides a power law relationship between V and L. These results, in conjunction with conservation of energy, yield relationships between the diameter of a segment and the length of its distal subtree. The relationships were tested based on a complete set of anatomical data of the coronary arterial trees using two models. The first model, called the truncated tree model, is an actual reconstruction of the coronary arterial tree down to 500 microm in diameter. The second model, called the symmetric tree model, satisfies all mean anatomical data down to the capillary vessels. Our results show very good agreement between the theoretical formulation and the measured anatomical data, which may provide insight into the design of the coronary arterial tree. Furthermore, the established relationships between the various morphometric parameters of the truncated tree model may provide a basis for assessing the extent of diffuse coronary artery disease.

Arteries↗

Three-dimensional segmentation and skeletonization to build an airway tree data structure for small animals.

Quantitative analysis of intrathoracic airway tree geometry is important for objective evaluation of bronchial tree structure and function. Currently, there is more human data than small animal data on airway morphometry. In this study, we implemented a semi-automatic approach to quantitatively describe airway tree geometry by using high-resolution computed tomography (CT) images to build a tree data structure for small animals such as rats and mice. Silicon lung casts of the excised lungs from a canine and a mouse were used for micro-CT imaging of the airway trees. The programming language IDL was used to implement a 3D region-growing threshold algorithm for segmenting out the airway lung volume from the CT data. Subsequently, a fully-parallel 3D thinning algorithm was implemented in order to complete the skeletonization of the segmented airways. A tree data structure was then created and saved by parsing through the skeletonized volume using the Python programming language. Pertinent information such as the length of all airway segments was stored in the data structure. This approach was shown to be accurate and efficient for up to six generations for the canine lung cast and ten generations for the mouse lung cast.

Algorithms↗

Girdling affects carbohydrate-related gene expression in leaves, bark and roots of alternate-bearing citrus trees.

Effects of girdling on carbohydrate status and carbohydrate-related gene expression in citrus trees were investigated. Alternate-bearing 'Murcott' (a Citrus reticulata hybrid of unknown origin) trees were girdled during autumn (25 Sep. 2001) and examined 10 weeks later. Girdling brought about carbohydrate (soluble sugar and starch) accumulation in leaves and shoot bark above the girdle, in trees during their fruitless, 'off' year. Trees during their heavy fruit load, 'on' year did not accumulate carbohydrates above the girdle due to the high demand for carbohydrates by the developing fruit. Girdling caused a strong decline in soluble sugar and starch concentrations in organs below the girdle (roots), in both 'on' and 'off' trees. Expression of STPH-L and STPH-H (two isoforms of starch phosphorylase), Agps (ADP-glucose pyrophosphorylase, small subunit), AATP (plastidic ADP/ATP transporter), PGM-C (phosphoglucomutase) and CitSuS1 (sucrose synthase), all of which are associated with starch accumulation, was studied. It was found that gene expression is related to starch accumulation in all 'off' tree organs. RNA levels of all the genes examined were high in leaves and bark that accumulated high concentrations of starch, and low in roots with declining starch concentrations. It may be hypothesized that changes in specific sugars signal the up- and down-regulation of genes involved in starch synthesis.

Carbohydrate Metabolism↗

Arbor 3D: an interactive environment for examining phylogenetic and taxonomic trees in multiple dimensions.

UNLABELLED: This paper examines a new technique for the visualization of and the interaction with trees, objects frequently used to convey hierarchical relationships in biological data. Motivated by the quality of 2D tree interaction, we adapt the planar tree-of-life metaphor to a virtual, semi-immersive 3D environment. A 3D environment extends the utility of this metaphor by allowing the user to view an entire data set in a single screen. Interrogation of the tree is implemented using 3D input devices. This real-time interrogation of the tree itself provides a quick means by which to qualitatively analyse the hierarchical data. In this paper, we describe the techniques underlying the implementation of such an environment. We conclude by considering the utility of tree metaphors as a basis for the representation of highly dimensional data sets. AVAILABILITY: Arbor3D (source code, a binary executable for SGI IRIX 6.4, Perl parsers, and sample Newick data files) are available via the Internet (http://xian.tamu.edu/Arbor3D/). Arbor3D can be displayed in "CAVE simulator" mode on an SGI workstation screen, or as an interactive virtual environment on a projection workbench. CONTACT: druths@rice.edu; echen@cs.rice.edu; leland@xian.tamu.edu

Classification↗

PhyloDraw: a phylogenetic tree drawing system.

UNLABELLED: PhyloDraw is a unified viewing tool for phylogenetic trees. PhyloDraw supports various kinds of multi-alignment formats (Dialign2, Clustal-W, Phylip format, NEXUS, MEGA, and pairwise distance matrix) and visualizes various kinds of tree diagrams, e.g. rectangular cladogram, slanted cladogram, phylogram, unrooted tree, and radial tree. By using several control parameters, users can easily and interactively manipulate the shape of phylogenetic trees. This program can export the final tree layout to BMP (bitmap image format) and PostScript. AVAILABILITY: http://pearl.cs.pusan.ac.kr/phylodraw/ CONTACT: jhchoi@pearl.cs.pusan.ac.kr

Computational Biology↗

Single column discrepancy and dynamic max-mini optimizations for quickly finding the most parsimonious evolutionary trees.

MOTIVATION: In the maximum parsimony (MP) method, the tree requiring the minimum number of changes (discrepancy) to explain the given set of DNA or amino acid sequences is chosen to represent their evolutionary relationships. To find the MP tree, the branch-and-bound algorithm is normally used. For a partial phylogenetic-tree (one that has a subset of the organisms) the traditional algorithm assigns a cost equal to the discrepancy of the partial phylogenetic-tree. We propose a single column discrepancy heuristic which increases this cost by predicting a minimum additional discrepancy needed to attach the sequences yet to be added to the partial phylogenetic-tree. A dynamic Max-mini order of sequence addition is also proposed to quickly terminate branch-and-bound search paths that are guaranteed to lead to suboptimal solutions. RESULTS: We studied the running time of 47 problems generated from 17 data sets. The use of single column discrepancy heuristic speeded up the computation to 2.4-fold for static and 18.2-fold for dynamic search order. The improvement appeared to increase exponentially with the number of sequences. The proposed strategies are also likely to be useful in speeding up the MP tree search using heuristic searches that are based on branch-and-bound-like algorithms. CONTACT: s.kumar@asu.edu

Amino Acids↗

Automated ortholog inference from phylogenetic trees and calculation of orthology reliability.

MOTIVATION: Orthologous proteins in different species are likely to have similar biochemical function and biological role. When annotating a newly sequenced genome by sequence homology, the most precise and reliable functional information can thus be derived from orthologs in other species. A standard method of finding orthologs is to compare the sequence tree with the species tree. However, since the topology of phylogenetic tree is not always reliable one might get incorrect assignments. RESULTS: Here we present a novel method that resolves this problem by analyzing a set of bootstrap trees instead of the optimal tree. The frequency of orthology assignments in the bootstrap trees can be interpreted as a support value for the possible orthology of the sequences. Our method is efficient enough to analyze data in the scale of whole genomes. It is implemented in Java and calculates orthology support levels for all pairwise combinations of homologous sequences of two species. The method was tested on simulated datasets and on real data of homologous proteins.

Algorithms↗

A program for drawing evolutionary trees.

From the measures of evolutionary distance between pairs of sequences in a set, it is possible to infer the genetic tree or trees which best fit these known data. DENDRON is a new program, written in FORTRAN 66, which computes an initial tree from the bottom-up, then searches among increasingly divergent trees for a better fit. As a check on the consistency of the measures, the program tests all triplets for the triangle inequality. DENDRON also calculates a single 'top-down' tree, progressing from the trunk to the twigs, for comparison with the 'bottom-up' trees.

Biological Evolution↗

Bit-reproducible parallel phylogenetic tree inference.

MOTIVATION: Phylogenetic trees describe the evolutionary history among biological species based on their genomic data. Maximum likelihood (ML) based phylogenetic inference tools search for the tree and evolutionary model that best explain the observed genomic data. Given the independence of likelihood score calculations between different genomic sites, parallel computation is commonly deployed. This is followed by a parallel summation over the per-site scores to obtain the overall likelihood score of the tree. However, basic arithmetic operations on IEEE 754 floating-point numbers, such as addition and multiplication, inherently introduce rounding errors. Consequently, the order by which floating-point operations are executed affects the exact resulting likelihood value since these operations are not associative. Moreover, parallel reduction algorithms in numerical codes re-associate operations as a function of the core count and cluster network topology, inducing different round-off errors. These low-level deviations can cause heuristic searches to diverge and induce high-level result discrepancies (e.g. yield topologically distinct phylogenies). This effect has also been observed in multiple scientific fields beyond phylogenetics. RESULTS: We observe that varying the degree of parallelism results in diverging phylogenetic tree searches (high-level results) for over 31% out of 10&#xa0;179 empirical datasets. More importantly, 8% of these diverging datasets yield trees that are statistically significantly worse than the best-known ML tree for the dataset (AU-test, P&#x2009;<&#x2009;.05). To alleviate this, we develop a variant of the widely used phylogenetic inference tool RAxML-NG, which does yield bit-reproducible results under varying core-counts, with a slowdown of only 0%-12.7% (median 0.8%) on up to 768 cores. For this, we introduce the ReproRed reduction algorithm, which yields bit-identical results under varying core-counts, by maintaining a fixed operation order that is independent of the communication pattern. ReproRed is thus applicable to all associative reduction operations-in contrast to competitors, which are confined to summation. Our ReproRed reduction algorithm only exchanges the theoretical minimum number of messages, overlaps communication with computation, and utilizes fast base-cases for local reductions. ReproRed is able to all-reduce (via a subsequent broadcast) 4.1&#xd7;106 operands across 48-768 cores in 19.7-48.61&#x2009;&#x3bc;s, thereby exhibiting a slowdown of 13%-93% over a non-reproducible all-reduce algorithm. ReproRed outperforms the state-of-the-art reproducible all-reduction algorithm ReproBLAS (offers summation only) beyond 10&#xa0;000 elements per core. In summary, we re-assess non-reproducibility in parallel phylogenetic inference, present the first bit-reproducible parallel phylogenetic inference tool, as well as introduce a general algorithm and open-source code for conducting reproducible associative parallel reduction operations. AVAILABILITY AND IMPLEMENTATION: ReproRed: https://doi.org/10.5281/zenodo.15004918 (LGPL)-Reproducible RAxML-NG version https://doi.org/10.5281/zenodo.15017407 (GPL).

Phylogeny↗