Correlates of diversification in the plant clade Dipsacales: geographic movement and evolutionary innovations

We explore patterns of diversification in the plant clades Adoxaceae and Valerianaceae (within Dipsacales), evaluating correlations between biogeographic change (i.e., movements into new areas), morphological change (e.g., the origin of putative key innovations associated with vegetative and reproductive characters), and shifts in rates of diversification. Our findings indicate that rates of diversification in these plants tend to be less tightly correlated with the evolution of morphological innovations but instead exhibit a pronounced correlation with movement into new geographic areas, particularly the dispersal of lineages into new mountainous regions. The interdependence among apparent novelties (arising from their nested phylogenetic distribution) and the correlation between morphological and biogeographic change suggests a complex history of diversification in Dipsacales. Overall, these findings highlight the importance of incorporating biogeographic history in studies of diversification rates and in the study of geographic gradients in species richness. Furthermore, these results argue against a simple deterministic relationship between dispersal and diversification: like other factors that may influence the probability of speciation and/or extinction, the impact of dispersal on diversification rates depends on being in the right place at the right time.     

Mitochondrial sequence data and Dipsacales phylogeny: mixed models, partitioned Bayesian analyses, and model selection

Phylogenetic analyses of chloroplast DNA sequences, morphology, and combined data have provided consistent support for many of the major branches within the angiosperm clade Dipsacales. Here we use sequences from three mitochondrial loci to test the existing broad scale phylogeny and in an attempt to resolve several relationships that have remained uncertain. Parsimony, maximum likelihood, and Bayesian analyses of a combined mitochondrial data set recover trees broadly consistent with previous studies, although resolution and support are lower than in the largest chloroplast analyses. Combining chloroplast and mitochondrial data results in a generally well-resolved and very strongly supported topology but the previously recognized problem areas remain. To investigate why these relationships have been difficult to resolve we conducted a series of experiments using different data partitions and heterogeneous substitution models. Usually more complex modeling schemes are favored regardless of the partitions recognized but model choice had little effect on topology or support values. In contrast there are consistent but weakly supported differences in the topologies recovered from coding and non-coding matrices. These conflicts directly correspond to relationships that were poorly resolved in analyses of the full combined chloroplast-mitochondrial data set. We suggest incongruent signal has contributed to our inability to confidently resolve these problem areas.     

Fundamental links between genes and elements: evolutionary implications of ecological stoichiometry

Organisms require elements to live and reproduce. We already know that availability of certain elements [e.g. phosphorus (P)] is highly variable spatiotemporally. In addition, there is variability in demand for various elements ontogenetically, as well as phylogenetically. Nonetheless, we know little about the underlying causes for such variation. In this study, we surveyed the literature to identify genes involved in the homeostasis of one biogenic element, P. Evidence from the literature suggests that variation in the environmental supply of P affects expression of highly conserved genes (e.g. the phosphate transporter system). In addition, we found evidence for genetic variation in the acquisition, assimilation, and allocation of P. Such effects of P supply should impinge on fitness, and drive evolutionary change. Further understanding of the inter- and intraspecific fitness consequences to imbalances in the availability of P in relation to other biogenic elements may be useful to disentangle primary mechanisms driving diversifications. We argue that studying the interactions between stoichiometric constraints and underlying genes is relevant to the majority of organisms, and conclude by furnishing information on designing and interpreting experiments using this approach.     

Atavisms: medical, genetic, and evolutionary implications

Traits expected to be lost in the evolutionary history of a species occasionally reappear apparently out of the blue. Such traits as extra nipples or tails in humans, hind limbs in whales, teeth in birds, or wings in wingless stick insects remind us that certain genetic information is not completely lost, but can be reactivated. Atavisms seem to violate one of the central evolutionary principles, known as Dollo's law, that "an organism is unable to return, even partially, to a previous stage already realized in the ranks of its ancestors." Although it is still not clear what triggers and controls the reactivation of dormant traits, atavisms are a challenge to evolutionary biologists and geneticists. This article presents some of the more striking examples of atavisms, discusses some of the currently controversial issues like human quadrupedalism, and reviews the progress made in explaining some of the mechanisms that can lead to atavistic features.     

Expression of Dlx genes during the development of the zebrafish pharyngeal dentition: evolutionary implications

In order to investigate similarities and differences in genetic control of development among teeth within and between species, we determined the expression pattern of all eight Dlx genes of the zebrafish during development of the pharyngeal dentition and compared these data with that reported for mouse molar tooth development. We found that (i) dlx1a and dlx6a are not expressed in teeth, in contrast to their murine orthologs, Dlx1 and Dlx6; (ii) the expression of the six other zebrafish Dlx genes overlaps in time and space, particularly during early morphogenesis; (iii) teeth in different locations and generations within the zebrafish dentition differ in the number of genes expressed; (iv) expression similarities and differences between zebrafish Dlx genes do not clearly follow phylogenetic and linkage relationships; and (v) similarities and differences exist in the expression of zebrafish and mouse Dlx orthologs. Taken together, these results indicate that the Dlx gene family, despite having been involved in vertebrate tooth development for over 400 million years, has undergone extensive diversification of expression of individual genes both within and between dentitions. The latter type of difference may reflect the highly specialized dentition of the mouse relative to that of the zebrafish, and/or genome duplication in the zebrafish lineage facilitating a redistribution of Dlx gene function during odontogenesis.     

The heat shock genes in theDrosophila montium subgroup: Chromosomal localization and evolutionary implications

The hsp70, hsp83, hsrω, and the small heat shock protein genes were mapped on the polytene chromosomes of six species, representative of the geographical distribution of the Drosophila montium subgroup of the melanogaster species group. In addition, based on hybridization conditions, the putative locus of the hsp68 gene is given. In contrast to the situation in the melanogaster subgroup species, the hsp70 locus is single in the montium species. The hsp83, hsrωand the small hsp loci are also single in the montium genomes studied here, a common feature of all Drosophila species. Among the hsp genes studied, the small hsp genes and the hsrω-homologous sequences exhibit a higher degree of divergence between the melanogaster and the montium subgroups. Our results support the idea that the montium subgroup species has a genome organization closer to that of the common ancestor compared with the melanogaster subgroup species. Received: 25 July 1995; in revised form: 31 January 1996 / Accepted: 1 April 1996     

Digest: Fossils,evolutionary models,and diatoms*

Adaptation to new environments by organisms has been and is of much interest in scientific research. Does the diatom lineage Stephanodiscus niagarae/yellowstonensis follow models compatible with adaptation to a new environment? Voje (2019) shows that in this lineage, none of the current models tested—decelerated evolution, Ornstein-Uhlenbeck, and random walk—fully explained how multiple traits—valve diameter, the number of costae, and spines per valve—evolved. More accurate models are needed to understand the evolutionary history of these diatoms.     

"Bioplutonism" and the evolutionary implications of beneficial genes from another biosphere

Could exogenous genes from another biosphere have aided the evolution of life on Earth's surface over the last half-billion years? That possibility was considered by Thomas Gold in 1992, when he hypothesized that a “deep hot biosphere” (DHB) resides independently well below its cooler surface counterpart. And he suggested that “… in the long term … there may occasionally be beneficial exchanges of genetic material between microbial life at depth and the surface life.” Thus, the question: what evidence is there to support Gold's notion that exogenous genes from the DHB – let us call them “bioplutons” – ever bestowed benefits on the evolution of surface life? In pursuit of this question I drafted a null hypothesis: “Nothing beyond our own biosphere, as we know it today, renders any kind of genetic benefits to biological evolution.” After objectively analyzing the evidence and arguments pro and con I failed to reject the null hypothesis, given what we know today, especially the fact that no genetic imprint from the DHB has been identified in eukaryotic genomes. But my conclusion is regarded as tentative, because the fundamentals of Gold's argument, collectively referred to herein as “bioplutonism,” might be confirmed eventually with successful probes into the DHB, and with the sampling of its alleged genetic material.     

Structure and expression of three Emx genes in the dogfish Scyliorhinus canicula: functional and evolutionary implications

We report the characterization of three Emx genes in a chondrichthyan, the dogfish Scyliorhinus canicula. Comparisons of these genes with their osteichthyan counterparts indicate that the gnathostome Emx genes belong to three distinct orthology classes, each containing one of the dogfish genes and either the tetrapod Emx1 genes (Emx1 class), the osteichthyan Emx2 genes (Emx2 class) or the zebrafish Emx1 gene (Emx3 class). While the three classes could be retrieved from the pufferfish genome data, no indication of an Emx3-related gene in tetrapods could be found in the databases, suggesting that this class may have been lost in this taxon. Expression pattern comparisons of the three dogfish Emx genes and their osteichthyan counterparts indicate that not only telencephalic, but also diencephalic Emx expression territories are highly conserved among gnathostomes. In particular, all gnathostomes share an early, dynamic phase of Emx expression, spanning presumptive dorsal diencephalic territories, which involves Emx3 in the dogfish, but another orthology class, Emx2, in tetrapods. In addition, the dogfish Emx2 gene shows a highly specific expression domain in the cephalic paraxial mesoderm from the end of gastrulation and throughout neurulation, which suggests a role in the segmentation of the cephalic mesoderm.     

The RNA polymerase III-dependent family of genes in hemiascomycetes: comparative RNomics, decoding strategies, transcription and evolutionary implications

We present the first comprehensive analysis of RNA polymerase III (Pol III) transcribed genes in ten yeast genomes. This set includes all tRNA genes (tDNA) and genes coding for SNR6 (U6), SNR52, SCR1 and RPR1 RNA in the nine hemiascomycetes Saccharomyces cerevisiae, Saccharomyces castellii, Candida glabrata, Kluyveromyces waltii, Kluyveromyces lactis, Eremothecium gossypii, Debaryomyces hansenii, Candida albicans, Yarrowia lipolytica and the archiascomycete Schizosaccharomyces pombe. We systematically analysed sequence specificities of tRNA genes, polymorphism, variability of introns, gene redundancy and gene clustering. Analysis of decoding strategies showed that yeasts close to S.cerevisiae use bacterial decoding rules to read the Leu CUN and Arg CGN codons, in contrast to all other known Eukaryotes. In D.hansenii and C.albicans, we identified a novel tDNA-Leu (AAG), reading the Leu CUU/CUC/CUA codons with an unusual G at position 32. A systematic 'p-distance tree' using the 60 variable positions of the tRNA molecule revealed that most tDNAs cluster into amino acid-specific sub-trees, suggesting that, within hemiascomycetes, orthologous tDNAs are more closely related than paralogs. We finally determined the bipartite A- and B-box sequences recognized by TFIIIC. These minimal sequences are nearly conserved throughout hemiascomycetes and were satisfactorily retrieved at appropriate locations in other Pol III genes.     

Diversity of secondary endosymbiont-derived actin-coding genes in cryptomonads and their evolutionary implications

In the secondary endosymbiotic organisms of cryptomonads, the symbiont actin genes have been found together with the host one. To examine whether they are commonly conserved and where they are encoded, host and symbiont actin genes from Pyrenomonas helgolandii were isolated, and their specific and homologous regions were digoxigenin (DIG) labeled separately. Using these probes, Southern hybridization was performed on 13 species of cryptomonads. They were divided into three groups: (1) both host and symbiont actin gene signals were detected, (2) only the host actin gene signal was detected, and (3) host and unknown actin signals were detected. The phylogenetic analysis of these actin gene sequences indicated that the evolutionary rates of the symbiont actin genes were accelerated more than those of the hosts. The unknown actin signals were recognized as the highly diverged symbiont actin genes. One of the diverged symbiont actin sequences from Guillardia theta is presumed to be as a pseudogene or to its precursor. Southern hybridizations based on the samples divided by pulsed-field gel electrophoresis showed that all actin genes were encoded by the host nuclei. These results possibly represent the evolutionary fate of the symbiont actin gene in cryptomonads, which was firstly transferred from the symbiont nucleus or nucleomorph, to the host nucleus and became a pseudogene and then finally disappeared there.     

The legumin gene family: structure and evolutionary implications of Vicia faba B-type genes and pseudogenes

We have characterized several Vicia faba genes encoding methionine residue-free group B subunits of the 11S or legumin storage proteins. The respective gene subfamily consists of 10 to 15 members, six of them having been studied by DNA sequence analysis. Four functional genes (LeB2, LeB4, LeB6, LeB7) are highly homologous in their coding region and 0.3 kb of their 3 flanking sequences. On the other hand, two pseudogenes (LeB1, LeB5) have accumulated a large number of mutations including an identical 0.7 kb internal deletion; they are both flanked by a repetitive element. Analysis of sequence changes show that transitions are nearly double as frequent as transversions. CpG is the most infrequent dinucleotide whereas TpA is significantly underrepresented in exon sequences. End points of deletions are correlated with short direct repeats and preferentially found in the two introns. Our studies indicate that the Vicia faba legumin B gene subfamily contains a group of expressed, highly homologous genes as well as more diverged pseudogenes.     

Pollen viability and longevity: Practical, ecological and evolutionary implications

The present article reviews the various definitions and terminology of pollen viability and longevity as well as the various tests of its assessment. We compare the advantages and the disadvantages of each method and suggest some practical implications as revealed by the extensive data. We recognize eight main hypotheses concerning the ecology and the evolution of pollen longevity and critically evaluated them according to the literature. The hypotheses are grouped as follows: (1) Desiccation risk-carbohydrate content; (2) Pollen packaging; (3) Pollen competitive ability; (4) Pollinator activity-stigma receptivity duration; (5) Self-pollination chance; (6) Pollen exposure schedule; (7) Pollen travel distance, and (8) Pollen removal chance.     

Structural differences between the repertoires of mouse and human germline genes and their evolutionary implications

 Although human and mouse antibodies are similar when one considers their diversification strategies, they differ in the extent to which kappa and lambda light chains are present in their respective variable light chain repertoires. While the Igk-V germline genes are preponderant in mice (95% or more), they comprise only 60% in humans. This may account for differences in the structural repertoire encoded in the Igk-V germline genes of these species. However, this subject has not been properly investigated, partially because a systematic structural characterization of the mouse Igk-V germline genes has not been undertaken. In the present study we compiled all available information on mouse Igk-V germline genes to characterize their structural repertoire. As expected, comparison with the structural repertoire of human Igk-V germline genes indicates differences. The most interesting is that the mouse Igk-V germline gene repertoire is more diverse in structural terms than its human counterpart: the mouse encodes seven canonical structure classes (combination of canonical structures in L1 and L3). In contrast, the human encodes only four. Analysis of the evolutionary relationships of human and mouse Igk-V germline genes led us to propose that the difference reflects a strategy of mice to compensate for the small lambda chain contribution to the repertoire of their variable light chains. Received: 1 June 1997 / Revised: 6 October 1997     

全基因组复制事件的绝对定年揭示莲座蕨属植物的迟滞演化

全基因组复制在维管植物的物种形成过程中普遍存在, 被认为是物种适应极端环境的重要机制之一。确定全基因组复制事件的发生时间对理解生物的适应性演化具有重要意义。然而, 在维管植物, 特别是蕨类植物中, 全基因组复制事件的发生时间及其演化意义仍知之甚少。本研究以蕨类植物重要基部类群——福建莲座蕨(Angiopteris fokiensis)为例, 基于不同采样点(广东、广西、上海)的3个转录组学数据, 利用同义替换率(Ks)和绝对定年的方法分析全基因组复制事件的发生时间和物种单位时间内的分子演化速率, 并对事件发生后保留下的基因进行基因功能注释和富集分析。结果表明, 福建莲座蕨在159‒165 Mya发生了一次全基因组复制事件, 该复制事件优先保留的基因主要与营养代谢、信号传导、适应调节和组织结构生长相关。另外, 福建莲座蕨的分子演化速率为1.66 × 10^‒9 (同义替换/位点/年), 是除裸子植物外, 陆生植物中已知演化速率最缓慢的类群。综合以上研究结果, 我们推测福建莲座蕨全基因组复制的发生可能与裸子植物繁盛、核心被子植物集中兴起或托阿尔阶灭绝事件有关。而复制后显著保留基因可能促进了莲座蕨属(Angiopteris)植物的遗传和形态创新, 从而帮助其快速适应环境的剧烈变化。进一步对该类群植物演化速率缓慢的原因进行讨论, 推测莲座蕨属缓慢的演化速率可能与其本身世代周期长、基因组较大及其生长环境稳定有关。本研究通过分析福建莲座蕨的全基因组复制历史和复制基因的保留模式, 推测全基因组复制事件对促进演化速率较慢的植物适应极端环境变化具有重要意义, 可为理解其他陆生植物的适应性演化提供更多启发。     

Energy,genes and evolution: introduction to an evolutionary synthesis

Life is the harnessing of chemical energy in such a way that the energy-harnessing device makes a copy of itself. No energy, no evolution. The ‘modern synthesis’ of the past century explained evolution in terms of genes, but this is only part of the story. While the mechanisms of natural selection are correct, and increasingly well understood, they do little to explain the actual trajectories taken by life on Earth. From a cosmic perspective—what is the probability of life elsewhere in the Universe, and what are its probable traits?—a gene-based view of evolution says almost nothing. Irresistible geological and environmental changes affected eukaryotes and prokaryotes in very different ways, ones that do not relate to specific genes or niches. Questions such as the early emergence of life, the morphological and genomic constraints on prokaryotes, the singular origin of eukaryotes, and the unique and perplexing traits shared by all eukaryotes but not found in any prokaryote, are instead illuminated by bioenergetics. If nothing in biology makes sense except in the light of evolution, nothing in evolution makes sense except in the light of energetics. This Special Issue of Philosophical Transactions examines the interplay between energy transduction and genome function in the major transitions of evolution, with implications ranging from planetary habitability to human health. We hope that these papers will contribute to a new evolutionary synthesis of energetics and genetics.     

Mathematical models, explanation, laws, and evolutionary biology

It is commonly agreed in the literature on laws of nature that there are at least two necessary conditions for lawhood--that a law must have empirical content and that it must be universal. The main reason offered for the requirement that laws be empirical is as follows: a priori statements are consistent with any imaginable set of observations, so they cannot be informative about the world and therefore they cannot provide explanations. However, we care about laws because we think that laws provide explanations and allow us to make predictions. Thus, if one of the functions of laws is to provide explanations and a priori propositions cannot fulfill this function, they cannot properly be viewed as laws. In this paper, I will aim to show that this argument for the claim that laws must be empirical does not work.