Developing and applying heterogeneous phylogenetic models with XRate

Modeling sequence evolution on phylogenetic trees is a useful technique in computational biology. Especially powerful are models which take account of the heterogeneous nature of sequence evolution according to the "grammar" of the encoded gene features. However, beyond a modest level of model complexity, manual coding of models becomes prohibitively labor-intensive. We demonstrate, via a set of case studies, the new built-in model-prototyping capabilities of XRate (macros and Scheme extensions). These features allow rapid implementation of phylogenetic models which would have previously been far more labor-intensive. XRate 's new capabilities for lineage-specific models, ancestral sequence reconstruction, and improved annotation output are also discussed. XRate 's flexible model-specification capabilities and computational efficiency make it well-suited to developing and prototyping phylogenetic grammar models. XRate is available as part of the DART software package: http://biowiki.org/DART.     

Fels-Rand: an Xlisp-Stat program for the comparative analysis of data under phylogenetic uncertainty

MOTIVATION: Currently available programs for the comparative analysis of phylogenetic data do not perform optimally when the phylogeny is not completely specified (i.e. the phylogeny contains polytomies). Recent literature suggests that a better way to analyse the data would be to create random trees from the known phylogeny that are fully-resolved but consistent with the known tree. A computer program is presented, Fels-Rand, that performs such analyses. A randomisation procedure is used to generate trees that are fully resolved but whose structure is consistent with the original tree. Statistics are then calculated on a large number of these randomly-generated trees. Fels-Rand uses the object-oriented features of Xlisp-Stat to manipulate internal tree representations. Xlisp-Stat's dynamic graphing features are used to provide heuristic tools to aid in analysis, particularly outlier analysis. The usefulness of Xlisp-Stat as a system for phylogenetic computation is discussed. AVAILABILITY: Available from the author or at http://www.uq.edu.au/~ansblomb/Fels-Rand.sit.hqx. Xlisp-Stat is available from http://stat.umn.edu/~luke/xls/xlsinfo/xlsinfo.html. CONTACT: s.blomberg@abdn.ac.uk     

What Can Molecular and Morphological Markers Tell Us About Plant Hybridization?

The study of hybrids and their evolutionary significance is often based on a number of tacit assumptions regarding character expression in hybrids. This article examines morphological, chemical, and molecular character expression in hybrids to determine whether traditionally recognized properties of hybrid plants, such as hybrid intermediacy and character coherence, are actually supported by empirical evidence, and also examines the impact of hybrids on phylogenetic analyses. We show that hybrids are a mosaic of both parental and intermediate morphological characters rather than just intermediate ones, and that a large proportion of first (64%) and later generation hybrids (89%) exhibit extreme or novel characters. Chemical character expression in hybrids is more predictable, with predominantly additive or complementary expression for both first generation hybrids (68%) and hybrid taxa (54%). Likewise, the genetic basis, and thus the expression of molecular characters, is well-worked out and predictable, although non-Mendelian inheritance has been reported in a few instances for molecular markers as well.

There is even less empirical support for the concept of character coherence than hybrid intermediacy. Although morphological character coherence has been reported in both artificial and natural hybrid populations, it appears to be the exception rather than the rule. This idiosyncratic relationship among morphological characters is shown to be the norm for the relationship of morphological to molecular characters, as well as for the relationship among molecular characters.

Hybrids are shown to have little impact on the topology of nonhybrid taxa in phylogenetic trees. However, the expression of primitive vs. derived character states, the placement of hybrids in cladograms, the number of equally parsimonious trees produced, and the effects of hybrids on phylogenetic topology do not appear to be predictable. Thus, cladistic identification of hybrid taxa is difficult and may not be possible based on phenotypic data, regardless of the analytical tools used.     

PhyloWidget: web-based visualizations for the tree of life

PhyloWidget is a web-based tool for the visualization and manipulation of phylogenetic tree data. It can be accessed online or downloaded as a standalone application. A simple URL-based API allows databases to easily link to and customize PhyloWidget for interactively viewing medium- to large-sized trees. AVAILABILITY: PhyloWidget is available for online use or download at http://www.phylowidget.org/.     

Evolutionary HMMs: a Bayesian approach to multiple alignment

MOTIVATION: We review proposed syntheses of probabilistic sequence alignment, profiling and phylogeny. We develop a multiple alignment algorithm for Bayesian inference in the links model proposed by Thorne et al. (1991, J. Mol. Evol., 33, 114-124). The algorithm, described in detail in Section 3, samples from and/or maximizes the posterior distribution over multiple alignments for any number of DNA or protein sequences, conditioned on a phylogenetic tree. The individual sampling and maximization steps of the algorithm require no more computational resources than pairwise alignment. METHODS: We present a software implementation (Handel) of our algorithm and report test results on (i) simulated data sets and (ii) the structurally informed protein alignments of BAliBASE (Thompson et al., 1999, Nucleic Acids Res., 27, 2682-2690). RESULTS: We find that the mean sum-of-pairs score (a measure of residue-pair correspondence) for the BAliBASE alignments is only 13% lower for Handelthan for CLUSTALW(Thompson et al., 1994, Nucleic Acids Res., 22, 4673-4680), despite the relative simplicity of the links model (CLUSTALW uses affine gap scores and increased penalties for indels in hydrophobic regions). With reference to these benchmarks, we discuss potential improvements to the links model and implications for Bayesian multiple alignment and phylogenetic profiling. AVAILABILITY: The source code to Handelis freely distributed on the Internet at http://www.biowiki.org/Handel under the terms of the GNU Public License (GPL, 2000, http://www.fsf.org./copyleft/gpl.html).     

Effects of dominant tree species on insectivorous birds breeding in Mediterranean oak woodlands

Capsule Differences in Cork Oak Quercus suber and Holm Oak Quercus rotundifolia dominance had little influence on bird communities though bark-gleaners showed a foraging preference for Cork Oak.

Aims Examine the use of Cork and Holm Oak trees by insectivorous birds in Mediterranean oak woodlands.

Methods Point-counts were used to compare species abundance among Cork Oak-dominated, Holm Oak-dominated and mixed woodlands. Focal foraging observations were used to evaluate the use of Cork and Holm Oaks in the three habitats and to relate tree characteristics with the foraging time of foliage- and bark-gleaners.

Results Bird densities in the three habitats were not different for most foliage- and bark-gleaners. Tree preference index values and foraging time per tree showed no significant differences between tree species and foraging guilds, however bark-gleaners had positive index values for Cork Oak in the three habitats. The foraging time of foliage- and bark-gleaners on both tree species showed a positive relationship with characteristics associated with arthropod abundance.

Conclusion Cork and Holm Oak trees are equally preferred by foliage-gleaners but bark-gleaners moderately preferred Cork Oak. Characteristics regarding morphology, phenology and physiological condition of trees can be used to predict habitat quality for insectivorous forest birds in Mediterranean oak woodlands.     

Habitat use and tree selection of a declining Afro-Palaearctic migrant at sub-Saharan staging and wintering sites

Capsule: Wood Warblers Phylloscopus sibilatrix showed significant selection for tree species and woodland characteristics at staging and wintering sites in sub-Saharan Africa.

Aims: To investigate home range size, habitat and tree species selection of Wood Warblers at a staging site in Burkina Faso (Koubri) and a wintering site in Ghana (Pepease).

Methods: Comparing habitat recorded at locations of radio-tagged birds and at control points, we investigated whether there was habitat and tree species selection. We also compared home range size of individual birds between the two sites.

Results: Home range size did not differ between the two sites. There was significant selection for tree species at both Koubri and Pepease: Anogeissus leiocarpus and Albizia zygia, respectively. At Koubri, there was significant avoidance of the most common tree species (Azadirachta indica, Mangifera indica (both non-native), Vitellaria paradoxa and Acacia spp.). In addition, there was a preference for taller trees and greater tree density at both sites. However, the probability of a point being used declined with increasing number of taller (>14?m) trees.

Conclusion: Fine-scale selection of woodland habitats suggests that Wood Warblers are likely to suffer the consequences of ongoing land-use change in their West African wintering grounds.     

TREE-PUZZLE: maximum likelihood phylogenetic analysis using quartets and parallel computing

SUMMARY: TREE-PUZZLE is a program package for quartet-based maximum-likelihood phylogenetic analysis (formerly PUZZLE, Strimmer and von Haeseler, Mol. Biol. Evol., 13, 964-969, 1996) that provides methods for reconstruction, comparison, and testing of trees and models on DNA as well as protein sequences. To reduce waiting time for larger datasets the tree reconstruction part of the software has been parallelized using message passing that runs on clusters of workstations as well as parallel computers. AVAILABILITY: http://www.tree-puzzle.de. The program is written in ANSI C. TREE-PUZZLE can be run on UNIX, Windows and Mac systems, including Mac OS X. To run the parallel version of PUZZLE, a Message Passing Interface (MPI) library has to be installed on the system. Free MPI implementations are available on the Web (cf. http://www.lam-mpi.org/mpi/implementations/).     

Diverse scaling relationships of tree height and diameter in five tree species

Background: Allometric scaling relationships are important in ecology and forestry. The metabolic scaling theory (MST) predicts a universal invariant scaling relationship for tree growth, relating height and diameter to each other.

Aims: Data on five tree species (Pinus taeda L., Pinus virginiana Mill., Liquidambar styraciflua L., Liriodendron tulipifera L., and Pinus palustris Mill.) were used to test the predictions from MST on the scaling of height–diameter and also diameter growth.

Methods: Data on tree height and diameter for five tree species from both natural forests and plantations were collected to study two types of scaling relationships: tree height–diameter and stem diameter growth. A reduced major axis of regression analysis model type II was used to determine scaling exponents from each species under different environmental conditions.

Results: No universal invariant scaling exponent was found in height–diameter and diameter growth for the five species. The scaling varied across natural forests, plantations and scales (e.g., time and number of measured trees). However, in some situations the scaling exponents failed to show significant difference with the predicted values (e.g., 2/3 or 1).

Conclusions: Diverse scaling exponents were observed for the five tree species with the scaling relationships varying with environmental settings.     

Species notions that combine phylogenetic trees and phenotypic partitions

A recent paper (Manceau and Lambert in bioRxiv, 2017. https://doi.org/10.1101/075580) developed a novel approach for describing two well-defined notions of ‘species’ based on a phylogenetic tree and a phenotypic partition. In this paper, we explore some further combinatorial properties of this approach and describe an extension that allows an arbitrary number of phenotypic partitions to be combined with a phylogenetic tree for these two species notions.

     

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     

Forests for Energy and the Role of Planted Trees

Woodfuels currently supply 7 percent of worldwide energy use and are much more important in developing than developed countries. Worldwide fuelwood use may be slowly declining, while charcoal production is increasing. In developing countries, woodfuel comes from natural forests, woodlands, shrublands, plantations and trees outside forests; the last is often the most important source. In developed countries, woodfuels are largely from industrial roundwood processing where up to 50 percent of logs may be used for energy. In urban areas, of both developing and developed countries, woodfuels also arises from wood processing and construction and other urban residues including tree trimming. Currently, apart from some nonindustrial plantations and the small areas of energy plantations, woodfuel largely results from by-products and residues.

Forest plantation may currently provide 15 to 20 percent of all fuelwood and this contribution will rise due to continued tree planting plus the rapidly increasing industrial plantation harvest. However, forest and woodfuel-use data are of uneven quality, making estimates tenuous.

There is potential to increase the amount of woodfuels in developed countries but its use will depend on fossil fuel price increases, and on ecological and social issues. In Nordic countries, forest residue use has grown, being underpinned by carbon taxes and research and development. Site nutrient conservation is assisted by leaving foliage on site and by returning the ash after burning. Leaving bark on site may be important for eucalypts.

In developing countries participatory programmes are critical for encouraging tree planting. Farmers seldom plant for fuelwood alone. Large-scale fuelwood plantations programs have not always been successful. Agroforestry practices are often practiced as they provide multiple benefits, including the provision of woodfuel. Silviculture should be readily adoptable, with special attention being given to species selection, high-quality planting stock, establishment methods and protection from animals.

Because of woodfuel's low relative value, energy is seldom a management objective of industrial plantations. However, often silviculture could be altered to increase bioenergy output, particularly with those sawlog regimes based on low stockings and non-extraction thinnings, and through more intensive silviculture. As a supplement to current silvicultural decision-making tools, energy analysis of silvicultural options would assist in efficient allocation of fossil fuels.     

T-REX: reconstructing and visualizing phylogenetic trees and reticulation networks.

T-REX (tree and reticulogram reconstruction) is an application to reconstruct phylogenetic trees and reticulation networks from distance matrices. The application includes a number of tree fitting methods like NJ, UNJ or ADDTREE which have been very popular in phylogenetic analysis. At the same time, the software comprises several new methods of phylogenetic analysis such as: tree reconstruction using weights, tree inference from incomplete distance matrices or modeling a reticulation network for a collection of objects or species. T-REX also allows the user to visualize obtained tree or network structures using Hierarchical, Radial or Axial types of tree drawing and manipulate them interactively. AVAILABILITY: T-REX is a freeware package available online at: http://www.fas.umontreal.ca/biol/casgrain/en/labo/t-rex     

Special Care Is Needed in Applying Phylogenetic Comparative Methods to Gene Trees with Speciation and Duplication Nodes

How gene function evolves is a central question of evolutionary biology. It can be investigated by comparing functional genomics results between species and between genes. Most comparative studies of functional genomics have used pairwise comparisons. Yet it has been shown that this can provide biased results, as genes, like species, are phylogenetically related. Phylogenetic comparative methods should be used to correct for this, but they depend on strong assumptions, including unbiased tree estimates relative to the hypothesis being tested. Such methods have recently been used to test the “ortholog conjecture,” the hypothesis that functional evolution is faster in paralogs than in orthologs. Although pairwise comparisons of tissue specificity (τ) provided support for the ortholog conjecture, phylogenetic independent contrasts did not. Our reanalysis on the same gene trees identified problems with the time calibration of duplication nodes. We find that the gene trees used suffer from important biases, due to the inclusion of trees with no duplication nodes, to the relative age of speciations and duplications, to systematic differences in branch lengths, and to non-Brownian motion of tissue specificity on many trees. We find that incorrect implementation of phylogenetic method in empirical gene trees with duplications can be problematic. Controlling for biases allows successful use of phylogenetic methods to study the evolution of gene function and provides some support for the ortholog conjecture using three different phylogenetic approaches.     

PhyloDraw: a phylogenetic tree drawing system

SUMMARY: 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     

MetaPhOrs: orthology and paralogy predictions from multiple phylogenetic evidence using a consistency-based confidence score

Reliable prediction of orthology is central to comparative genomics. Approaches based on phylogenetic analyses closely resemble the original definition of orthology and paralogy and are known to be highly accurate. However, the large computational cost associated to these analyses is a limiting factor that often prevents its use at genomic scales. Recently, several projects have addressed the reconstruction of large collections of high-quality phylogenetic trees from which orthology and paralogy relationships can be inferred. This provides us with the opportunity to infer the evolutionary relationships of genes from multiple, independent, phylogenetic trees. Using such strategy, we combine phylogenetic information derived from different databases, to predict orthology and paralogy relationships for 4.1 million proteins in 829 fully sequenced genomes. We show that the number of independent sources from which a prediction is made, as well as the level of consistency across predictions, can be used as reliable confidence scores. A webserver has been developed to easily access these data (http://orthology.phylomedb.org), which provides users with a global repository of phylogeny-based orthology and paralogy predictions.     

Emerging single-cell technologies for functional proteomics in oncology

Introduction: Cellular heterogeneity has challenged current cancer therapeutics and hindered the discovery and development of cancer drugs. The heterogeneity in functional proteome is of particular interest because many cancer drugs are developed to target signaling proteins. The complex nature of tumor systems calls for more advanced multiplexed single-cell tools to address the heterogeneity issue.

Area covered: Over the past five years, there are a few single-cell functional proteomics tools introduced with unprecedented multiplexity and performance that are transforming the oncology field. Those tools are generally categorized as cytometry-based tools and microfluidics-based tools, and we discuss the representatives in both categories.

Expert commentary: The single-cell tools have provided an avenue to understand the multifaceted differences of cancer cells, the complex signaling networks, and the relationship of intercellular interaction and tumor architecture. We also provide an outlook of single-cell tools in five years and the challenges to address before a greater impact on oncology can be made.     

phangorn: phylogenetic analysis in R

SUMMARY: phangorn is a package for phylogenetic reconstruction and analysis in the R language. Previously it was only possible to estimate phylogenetic trees with distance methods in R. phangorn, now offers the possibility of reconstructing phylogenies with distance based methods, maximum parsimony or maximum likelihood (ML) and performing Hadamard conjugation. Extending the general ML framework, this package provides the possibility of estimating mixture and partition models. Furthermore, phangorn offers several functions for comparing trees, phylogenetic models or splits, simulating character data and performing congruence analyses. AVAILABILITY: phangorn can be obtained through the CRAN homepage http://cran.r-project.org/web/packages/phangorn/index.html. phangorn is licensed under GPL 2.     

Climate and fragmentation affect forest structure at the southern border of Amazonia

Background: The remaining forests in the extensive contact zone between southern Amazonia (seasonal rain forest) and the Cerrado (savanna) biomes are at risk due to intense land-use and climate change.

Aims: To explore the vulnerability of these transitional forests to changes in land use and climate, we evaluated the effects of fragmentation and climatic variables on forest structure.

Methods: We measured the diameter and height of 14,185 trees with diameter ≥10 cm at 24 forest plots distributed over an area of 25,000 km². For each plot, we obtained data on contemporary fragmentation and climatic variables.

Results: Forest structure variables (height, diameter, height:diameter allometry, biomass) varied significantly both within and among plots. The height, H:D and biomass of trees were positively correlated with annual precipitation and fragment area.

Conclusions: The association between forest structure and precipitation indicates that these forests plots are likely to be vulnerable to dry season intensification anticipated for the southern edge of the Amazon. Additionally, the reduction in the fragment area may contribute to reductions in forest biomass and tree height, and consequently ecosystem carbon stocks. Given the likely susceptibility of these forests, urgent conservation action is needed to prevent further habitat degradation.