Correlation Between Ka/Ks and Ks is Related to Substitution Model and Evolutionary Lineage

In 2005, Wyckoff and coworkers described a surprisingly strong correlation between Ka/Ks and Ks in several data sets using the LPB93 algorithm. This finding indicated the possibility of a paradigm shift in the way selection strength can be measured using the Ka/Ks ratio. We carried out a calculation of Ka and Ks using six different algorithms on three cross-species orthologous data sets and found a highly variable correlation among the algorithms and lineages. Algorithms based on the GY-HKY substitution model exhibit a weaker positive correlation or a stronger negative correlation than those based on the K2P and JC69 substitution model. Even if one algorithm shows a positive correlation between Ka/Ks and Ks in a warm-blooded lineage, it may show no correlation in a cold-blooded lineage. This algorithm-related and evolutionary lineage-related correlation indicates the need for great caution in drawing conclusions when using only one Ka and Ks algorithm in a genomewide analysis of selection strength. Our results indicated that currently used algorithms for Ka and Ks calculations are flawed and need improvements.     

KaKs_Calculator： Calculating Ka and Ks Through Model Selection and Model Averaging

KaKs_Calculator is a software package that calculates nonsynonymous （Ka） and synonymous （Ks） substitution rates through model selection and model averaging. Since existing methods for this estimation adopt their specific mutation （substitution） models that consider different evolutionary features, leading to diverse estimates, KaKs_Calculator implements a set of candidate models in a maximum likelihood framework and adopts the Akaike information criterion to measure fitness between models and data, aiming to include as many features as needed for accurately capturing evolutionary information in protein-coding sequences. In addition, several existing methods for calculating Ka and Ks are also incorporated into this software. KaKs_Calculator, including source codes, compiled executables, and documentation, is freely available for academic use at http：//evolution.genomics.org.cn/software.htm.     

Average allozyme heterozygosity in vertebrates correlates with Ka/Ks measured in the human-mouse lineage

It is well established that different allozyme proteins vary in heterozygosity in averages made over large numbers of species. For example, the enzyme 6-phosphogluconate dehydrogenase has a much higher average heterozygosity than glutamate dehydrogenase. Allozyme data alone provide insufficient power to determine the evolutionary cause of such a difference. Many studies have now been carried out on the DNA sequences coding for allozymes. These have identified diverse selective and nonselective causes of polymorphisms at individual loci. However the studies are mainly in a small number of model species; thus, it is difficult to identify from these DNA studies specific causes of global average heterozygosity differences among allozyme proteins. Here we demonstrate that estimates of average heterozygosity for 37 allozyme proteins in vertebrates correlate positively with Ka and Ka/Ks but not with Ks, measured in the human-mouse lineage. The values of Ka/Ks are less than 0.25, and Ka/Ks is negatively correlated with subunit number (quaternary structure), a measure of structural constraint. Proteins with lower levels of constraint have higher values of both Ka/Ks and heterozygosity. These results better support the hypothesis that differences in average allozyme diversity between proteins are more closely related to differences in the level of purifying selection than to differences in the underlying mutation rate or level of positive selection.     

Computing Ka and Ks with a consideration of unequal transitional substitutions

Background  

Approximate methods for estimating nonsynonymous and synonymous substitution rates (Ka and Ks) among protein-coding sequences have adopted different mutation (substitution) models. In the past two decades, several methods have been proposed but they have not considered unequal transitional substitutions (between the two purines, A and G, or the two pyrimidines, T and C) that become apparent when sequences data to be compared are vast and significantly diverged.     

基于Ks分布的被子植物演化的时间尺度研究

物种演化时间估算是生命演化研究的重要部分。近年来,许多研究发现由于不同基因和不同物种的进化速率差异显著,因此需要新的方法对进化事件发生时间进行重新估计。为了对被子植物演化时间的重新估计,该研究基于共享多倍化事件或共享分歧事件应该有共同同义突变率(Ks)峰值的理念,建立了基于基因组数据的进化速率矫正模型。结果表明:(1)对获取Ks分布三种常见方式进行比较分析,明确了通过提取共线性区块上Ks值的中位数的方式最优。(2)模拟了Ks值随时间累积系数(v)变化过程下的Ks分布,当假设v服从正态分布时,Ks分布出现了长尾现象。(3)将矫正方法应用到被子植物中,发现不同谱系的被子植物具有同步的辐射进化和适应性进化现象。并且,被子植物的进化速率虽然差异显著,但不同分支间的进化速率仍具有部分一致性,如木兰类植物进化速率最慢,真双子叶植物次之,单子叶植物进化速率最快。最终得到了相对可靠的物种同义突变率演化时间轴,为植物研究提供了系统发育和演化的支撑。     

The nonsynonymous/synonymous substitution rate ratio versus the radical/conservative replacement rate ratio in the evolution of mammalian genes

There are 2 ways to infer selection pressures in the evolution of protein-coding genes, the nonsynonymous and synonymous substitution rate ratio (K(A)/K(S)) and the radical and conservative amino acid replacement rate ratio (K(R)/K(C)). Because the K(R)/K(C) ratio depends on the definition of radical and conservative changes in the classification of amino acids, we develop an amino acid classification that maximizes the correlation between K(A)/K(S) and K(R)/K(C). An analysis of 3,375 orthologous gene groups among 5 mammalian species shows that our classification gives a significantly higher correlation coefficient between the 2 ratios than those of existing classifications. However, there are many orthologous gene groups with a low K(A)/K(S) but a high K(R)/K(C) ratio. Examining the functions of these genes, we found an overrepresentation of functional categories related to development. To determine if the overrepresentation is stage specific, we examined the expression patterns of these genes at different developmental stages of the mouse. Interestingly, these genes are highly expressed in the early middle stage of development (blastocyst to amnion). It is commonly thought that developmental genes tend to be conservative in evolution, but some molecular changes in developmental stages should have contributed to morphological divergence in adult mammals. Therefore, we propose that the relaxed pressures indicated by the K(R)/K(C) ratio but not by K(A)/K(S) in the early middle stage of development may be important for the morphological divergence of mammals at the adult stage, whereas purifying selection detected by K(A)/K(S) occurs in the early middle developmental stage.     

The D/H ratio and the evolution of water in the terrestrial planets

The presence of liquid water at the surface of the Earth has played a major role in the biological evolution of the Earth. None of the other terrestrial planets — Mercury, Venus and Mars — has liquid water at its surface. However, it has been suggested, since the early seventies, from both geological and atmospheric arguments that, although Venus and Mars are presently devoid of liquid water, their surfaces could have been partially or completely covered by water at some time of their evolution. There are many possible diagnostics of the long-term evolution of the planets, either from the present characteristics of their surfaces or from their present atmospheric compositions. Among them, the present value of the D/H ratio is of particular interest, although its significance in terms of long term evolution has been challenged by some authors. Recent progress has been made in this field. We now have evidence for higher D/H ratios on Mars and Venus than on Earth, with an enrichment factor of the order of 5 on Mars, and about 100 on Venus. Any scenario for the evolution of these planets must take this into account. The most recent models on the evolution of Mars and Venus are reviewed in light of these new measurements.Presented at the Session Water in the Solar System and Its Role in Exobiology during the 26th General Assembly of the European Geophysical Society, 22–26 April 1991 in Wiesbaden, Germany.     

The basic depression ratio of the host: the evolution of host resistance to microparasites

The basic reproduction ratio R0 occupies a central position in the theory of host pathogen interactions. However, this quantity stresses the role of the pathogen. This paper proposes an additional, more host-centred char acterization using the basic depression ratio D0. This quantity is the number of host individuals per infected by which the infected host population is depressed below its uninfected level. This paper shows that a baseline criterion for the evolution of host resistance to microparasites is that resistance evolves to minimize D0. This parallels the result for pathogen virulence where R0 is maximized. The tension between these two criteria is noted. The framework established allows a discussion of trade-offs between aspects of the pathogen-free host biology and the host pathogen interaction. For certain linear and convex trade-offs it is shown that the strain with the lowest transmission parameter beta wins (despite the fact that it has the lowest intrinsic birth rate a). For corresponding concave trade-offs, either the strain with minimum beta and a or the strain with maximum beta and a wins. Finally the connection with the techniques of adaptive dynamics is made. Evolutionary singular points are shown to occur at extrema of D0. The evolutionary attainment of the results is discussed.     

The evolution of signal form: effects of learned versus inherited recognition

Organisms can learn by individual experience to recognize relevant stimuli in the environment or they can genetically inherit this ability from their parents. Here, we ask how these two modes of acquisition affect signal evolution, focusing in particular on the exaggeration and cost of signals. We argue first, that faster learning by individual receivers cannot be a driving force for the evolution of exaggerated and costly signals unless signal senders are related or the same receiver and sender meet repeatedly. We argue instead that biases in receivers' recognition mechanisms can promote the evolution of costly exaggeration in signals. We provide support for this hypothesis by simulating coevolution between senders and receivers, using artificial neural networks as a model of receivers' recognition mechanisms. We analyse the joint effects of receiver biases, signal cost and mode of acquisition, investigating the circumstances under which learned recognition gives rise to more exaggerated signals than inherited recognition. We conclude the paper by discussing the relevance of our results to a number of biological scenarios.     

Sex ratio polymorphism: The impact of mutation and drift on evolution

This paper addresses the question, which sex ratio will evolve in a population that is subject to mutation and drift. The problem is analyzed using a simulation model as well as analytical methods. A detailed simulation model for the evolution of a population's allele distribution shows that for the sex ratio game a wide spectrum of different population states may evolve from on the one hand a monomorphic state with one predominant allele and with all other alleles suppressed by the forces of selection, to on the other hand a polymorphism determined by recurrent mutations. Which of these states will evolve depends on the population size, the mating system and the rate of mutations. For the sex ratio game the evolutionary stable strategy (ESS), as defined by evolutionary game theory, can only predict the population sex ratio but not the underlying stable population state. A comparison of different approaches to the problem shows that false predictions of the stable population states might result from two simplifying assumptions that are fairly common in evolutionary biology: a) it is assumed that mutations are rare events and there is never more than one mutant gene present in a population at any one time; b) a deterministic relationship is assumed between the fitness assigned to an individual's strategy and the individual's contribution to the gene pool of future generations.     

The human olfactory subgenome: from sequence to structure and evolution

Olfactory receptors (ORs) constitute the largest multigene family in multicellular organisms. Their evolutionary proliferation has been driven by the need to provide recognition capacity for millions of potential odorants with arbitrary chemical configurations. Human genome sequencing has provided a highly informative picture of the "olfactory subgenome", the repertoire of OR genes. We describe here an analysis of 224 human OR genes, a much larger number than hitherto systematically analyzed. These are derived by literature survey, data mining at 14 genomic clusters, and by an OR-targeted experimental sequencing strategy. The presented set contains at least 53% pseudogenes and is minimally divided into 11 gene families. One of these (no. 7) has undergone a particularly extensive expansion in primates. The analysis of this collection leads to insight into the origin of OR genes, suggesting a graded expansion through mammalian evolution. It also allows us to delineate a structural map of the respective proteins. A sequence database and analysis package is provided (http://bioinformatics.weizmann.ac.il/HORDE), which will be useful for analyzing human OR sequences genome-wide.     

CS-PSeq-Gen: simulating the evolution of protein sequence under constraints

CS-PSeq-Gen is a program derived from PSeq-Gen, designed to perform simulations of the evolution of protein sequences under the constraints of a reconstructed phylogeny. It also provides a basis for the investigation of the correlated evolution of sites. AVAILABILITY: http://condor.urbb.jussieu.fr/CS-PSeq-Gen.html     

The complete amino acid sequence of the mature form of rat sepiapterin reductase

The partial amino acid sequence of rat sepiapterin reductase was determined using peptides generated by cleavage of the S-carboxyamidomethylated protein with Achromobacter protease I, cyanogen bromide, chymotrypsin or BNPS-skatole. The protein began with N-acetyl methionyl residue at the N-terminus and ended with isoleucyl residue at the C-terminus. The present results essentially coincided with the amino acid sequence predicted from the nucleotide sequence of the cDNA recently reported by Citron et al. (Proc. Natl. Acad. Sci. USA 87, 6436-6440 (1990)), clarified the processing event during the biosynthesis and provided the complete amino acid sequence of the mature form of the enzyme.     

The development and evolution of hydrozoan polyp and colony form

Hydrozoans represent an extremely diverse group of mostly colonial forms. Despite this tremendous diversity, many of the morphological differences between hydrozoan species can be attributed to simple changes in the relative position of regions/structures along the axes of the polyp and the stolon or hydrocaulus from which polyps bud. Many genes have been implicated in the specification of positional information along the axis of the polyp. Knowledge from these studies in Hydra, and from comparative studies in Hydractinia polyp polymorphs, suggests that evolutionary changes in the regulation of axial patterning genes may be a prominent mechanism underlying hydrozoan evolution. Despite the paucity of interspecies comparative expression information, hypotheses can be formulated about the role of developmental regulatory genes in hydrozoan evolution from information available from Hydra.