Adaptive walks through fitness landscapes for early vascular land plants

Hypothetical adaptive walks (i. e., morphological transformation series gaining increasing relative fitness) were simulated through a computer-generated domain for early vascular land plant morphologies to examine the relationship between the dynamics of adaptive walks and the topologies of fitness landscapes. A total of 15 hypothetical adaptive walks were simulated, assuming that relative fitness was based on performing one or more of four biological tasks: maximizing light interception, mechanical stability, and reproductive success, and minimizing total surface area. Morphologies occupying fitness peaks typically were similar to some early vascular land plant remains. The most stringent task (the minimization of total surface area) resulted in a few, comparatively small Y-shaped morphologies. Based on the 15 walks, the number of fitness peaks increased and their heights decreased as the number of tasks simultaneously performed increased. These results (which are consistent with prior computer-simulated walks treating light interception, mechanical stability, and reproductive success) suggest that the biological requirement to conserve water reduced the number of phenotypic options available to the earliest land plants, and that, once this adaptive hurtle was overcome, the simultaneous performance of two or more tasks, increased the number of phenotypic options with equivalent relative fitnesses that could be rapidly reached due to the comparatively small fitness differential between derived and ancestral morphologies.     

Evolutionary genetics of plant adaptation

Plants provide unique opportunities to study the mechanistic basis and evolutionary processes of adaptation to diverse environmental conditions. Complementary laboratory and field experiments are important for testing hypotheses reflecting long-term ecological and evolutionary history. For example, these approaches can infer whether local adaptation results from genetic tradeoffs (antagonistic pleiotropy), where native alleles are best adapted to local conditions, or if local adaptation is caused by conditional neutrality at many loci, where alleles show fitness differences in one environment, but not in a contrasting environment. Ecological genetics in natural populations of perennial or outcrossing plants can also differ substantially from model systems. In this review of the evolutionary genetics of plant adaptation, we emphasize the importance of field studies for understanding the evolutionary dynamics of model and nonmodel systems, highlight a key life history trait (flowering time) and discuss emerging conservation issues.     

Evolutionary pathways to self-fertilization in a tristylous plant species

Evolutionary transitions from outcrossing to selfing occur commonly in heterostylous genera. The morphological polymorphisms that characterize heterostyly provide opportunities for different pathways for selfing to evolve. Here, we investigate the origins and pathways by which selfing has evolved in tristylous Eichhornia paniculata by providing new evidence based on morphology, DNA sequences and genetic analysis. The primary pathway from outcrossing to selfing involves the stochastic loss of the short-styled morph (S-morph) from trimorphic populations, followed by the spread of selfing variants of the mid-styled morph (M-morph). However, the discovery of selfing variants of the long-styled morph (L-morph) in Central America indicates a secondary pathway and distinct origin for selfing. Comparisons of multi-locus nucleotide sequences from 27 populations sampled from throughout the geographical range suggest multiple transitions to selfing. Genetic analysis of selfing variants of the L- and M-morphs demonstrates recessive control of the loss of herkogamy, although the number of factors appears to differ between the forms. Early stages in the establishment of selfing involve developmental instability in the formation of flowers capable of autonomous self-pollination. The relatively simple genetic control of herkogamy reduction and frequent colonizing episodes may often create demographic conditions favouring transitions to selfing in E. paniculata .     

The Devonian land plant Protosalvinia

Gray, Jane&Boucot, ArthurJ. 197901 15: The Devonian land plant Protosalvinia. Lethaia . Vol. 12, pp. 57–63. Oslo. ISSN 0024–1164.
Protosalvinia (=Foerstia ) shows a number of morphological features which suggest that it was a land plant, possibly an emergent aquatic. Certain kinds of biochemical data, as presently interpreted, also support the morphological evidence. The weight of evidence supports neither a fucoid nor even an algal attribution for this enigmatic plant. Protosalvinia should be treated as the representative of a separate group of land plants, as previously advocated. Tracheid-like tubes with internal spiral thickenings appear on the basis of direct comparison to be morphologically identical to those of nematophytalean origin. They may have functioned in a capacity similar to the tracheids of vascular plants. The Nematophytales are non-marine plants of uncertain taxonomic position.     

Evolutionary history of a mosquito endosymbiont revealed through mitochondrial hitchhiking

Due to cytoplasmic inheritance, spread of maternally inherited Wolbachia symbionts can result in reduction of mitochondrial variation in populations. We examined sequence diversity of the mitochondrial NADH dehydrogenase subunit 4 (ND4) gene in Wolbachia-infected (South Africa (SA), California and Thailand) and uninfected (SA) Culex pipiens complex populations. In total, we identified 12 haplotypes (A-L). In infected populations, 99% of individuals had haplotype K. In the uninfected SA population, 11 haplotypes were present, including K. Nuclear allozyme diversity was similar between infected and uninfected SA populations. Analysis of nuclear DNA sequences suggested that haplotype K presence in uninfected SA Cx. pipiens was probably due to a shared ancestral polymorphism rather than hybrid introgression. These data indicate that Wolbachia spread has resulted in drastic reduction of mitochondrial variability in widely separated Cx. pipiens complex populations. In contrast, the uninfected SA population is probably a cryptic species where Wolbachia introgression has been prevented by reproductive isolation, maintaining ancestral levels of mitochondrial diversity. Molecular clock analyses suggest that the Wolbachia sweep occurred within the last 47000 years. The effect of Wolbachia on mitochondrial dynamics can provide insight on the potential for Wolbachia to spread transgenes into mosquito populations to control vector-borne diseases.     

Evolutionary origins of a novel host plant detoxification gene in butterflies

Chemical interactions between plants and their insect herbivoresprovide an excellent opportunity to study the evolution of speciesinteractions on a molecular level. Here, we investigate themolecular evolutionary events that gave rise to a novel detoxifyingenzyme (nitrile-specifier protein [NSP]) in the butterfly familyPieridae, previously identified as a coevolutionary key innovation.By generating and sequencing expressed sequence tags, genomiclibraries, and screening databases we found NSP to be a memberof an insect-specific gene family, which we characterized andnamed the NSP-like gene family. Members consist of variabletandem repeats, are gut expressed, and are found across Insectaevolving in a dynamic, ongoing birth–death process. Inthe Lepidoptera, multiple copies of single-domain major allergengenes are present and originate via tandem duplications. Multipledomain genes are found solely within the brassicaceous-feedingPieridae butterflies, one of them being NSP and another calledmajor allergen (MA). Analyses suggest that NSP and its paralogMA have a unique single-domain evolutionary origin, being formedby intragenic domain duplication followed by tandem whole-geneduplication. Duplicates subsequently experienced a period ofrelaxed constraint followed by an increase in constraint, perhapsafter neofunctionalization. NSP and its ortholog MA are stillexperiencing high rates of change, reflecting a dynamic evolutionconsistent with the known role of NSP in plant–insectinteractions. Our results provide direct evidence to the hypothesisthat gene duplication is one of the driving forces for speciationand adaptation, showing that both within- and whole-gene tandemduplications are a powerful force underlying evolutionary adaptation.     

Charophyte algae and land plant origins

The charophyte algae are six distinct groups of mostly freshwater green algae that are related to modern land plants. Charophyte algae exhibit diverse morphologies and reproductive strategies, from unicells to branching erect forms, and from swimming asexual spores to sex involving eggs and sperm, respectively. The green algae known as stoneworts (Charales) are suggested to be the extant sister group to all land plants, although the phylogeny is not conclusive. Here we review recent molecular phylogenetic work on the charophyte algae and its implications for our understanding of the origins of land plants and of characters in their aquatic ancestors that might have played a role in the explosive diversification of plants on land.     

The evolution of land plant cilia

Eukaryotic cilia/flagella are ancient organelles with motility and sensory functions. Cilia display significant ultrastructural conservation where present across the eukaryotic phylogeny; however, diversity in ciliary biology exists and the ability to produce cilia has been lost independently on a number of occasions. Land plants provide an excellent system for the investigation of cilia evolution and loss across a broad phylogeny, because early divergent land plant lineages produce cilia, whereas most seed plants do not. This review highlights the differences in cilia form and function across land plants and discusses how recent advances in genomics are providing novel insights into the evolutionary trajectory of ciliary proteins. We propose a renewed effort to adopt ciliated land plants as models to investigate the mechanisms underpinning complex ciliary processes, such as number control, the coordination of basal body placement and the regulation of beat patterns.     

Evolutionary responses of a dominant plant along a successional gradient in a salt-marsh system

Plant Ecology - The ecological responses of plant populations along a successional gradient have been intensively examined; however, the evolutionary responses received much less attention. Here, I...     

Evolutionary genomics of LysM genes in land plants

Background  

The ubiquitous LysM motif recognizes peptidoglycan, chitooligosaccharides (chitin) and, presumably, other structurally-related oligosaccharides. LysM-containing proteins were first shown to be involved in bacterial cell wall degradation and, more recently, were implicated in perceiving chitin (one of the established pathogen-associated molecular patterns) and lipo-chitin (nodulation factors) in flowering plants. However, the majority of LysM genes in plants remain functionally uncharacterized and the evolutionary history of complex LysM genes remains elusive.     

Photosynthesis during desiccation in an intertidal alga and a land plant

This study was undertaken to determine how photosynthesis tolerates desiccation in an intertidal alga Fucus vesiculosus L. and a terrestrial sunflower Helianthus annuus L. Photosynthetic O2 evolution generally was inhibited at low water potentials (psiw) but more in sunflower leaves than in Fucus fronds at the same psiw. As psiw decreased, less carbon accumulated in an organic carbon store in Fucus. The inhibition of photosynthesis appeared to be mostly biochemical because it could not be prevented by supplying additional CO2 or by supplying CO2 from the internal organic carbon store. The inhibition of photosynthesis and carbon storage occurred after turgor disappeared and thus when solute concentrations were increasing in the cells. Solute concentrations were much higher in Fucus than in sunflower. After desiccation to the air-dry state (psiw below - 10 MPa), photosynthesis could not recover in sunflower but it recovered rapidly when Fucus was exposed to seawater. The lack of recovery in sunflower was associated with inability to recover turgor probably because of breaks in cell membranes. The ability to recover in Fucus was gradually lost during 1.5 d of desiccation at 45% relative humidity. At lower humidities, recovery was lost sooner as small amounts of water were removed. We conclude that photosynthesis tolerated desiccation more in Fucus than in sunflower because of differences in the molecular environment around the photosynthetic enzymes. Important aspects of this environment were features that prevented membrane breakage but promoted the retention of small amounts of water that were critical for viability.     

A chromium-tolerant plant growing in Cr-contaminated land

In this paper, a kind of Typhaceae plant species, Typha angustifolia L, with a high tolerance to Cr is described. Experiments were carried out to examine its ability to tolerant Cr and its physiological response. The results showed that there was no difference in growth, plant height, and biomass response to external Cr (VI) between the plants exposed to 100 microM Cr (VI) and control (0 microM), while increasing Cr levels to 200-800 microM induced a significant decrease in plant height and biomass, but no significant injury was detected, even for the plants exposed to 800 microM Cr. Chromium induced significant increases in superoxide dismutase (SOD) and peroxidase (POD) activities. Meanwhile, a significantly positive correlation was found between Cr and Mn or Cu in leaves and roots, respectively. The Cr tolerance of the plant appeared to be associated with the enhancement of SOD and POD activities and the improvement in uptake and translocation of the essential microelements.     

Microfossils and evidence of land plant evolution

Resolving the question of (1) land plant and (2) vascular land plant evolution is a complex problem. It requires a continued, multidisciplinary effort. We are all grateful to Dianne Edwards and her colleagues (Edwards et al. 1979) for pursuing one such line of research with careful attention to morphology and stratigraphy. Their efforts, however, are confined to one small part of the globe, and they perforce involve only one type of evidence effective with regard to questions of vascular plant evolution once the land estate has been reached. They do not deal with the basic question of vascular and non-vascular land plant origins. The so-called Přídolí benchmark that remained the sine qua nun for vascular plant evolution when Dianne Edwards began her studies has now been irrevocably breached. What other supposedly sacrosanct benchmarks will also crumble as the result of future discoveries, now that vascular plants have been recovered from lower Ludlovian strata? Nevertheless, pre-Devonian plant megafossils are too rare to yield definitive results with regard to the origin of land plants prior to the vascular estate, even though they could in principle (as emphasized by Edwards et al. 1979) provide the least ambiguous evidence. We must look elsewhere than to megafossils for evidence suggesting links in time between green algal ancestors, major groups of living green land Plants, and the many extinct groups of enigmatic higher land plants in the crucial Early Paleozoic time interval.     

Evolutionary conservation of plant gibberellin signalling pathway components

Background:  

Gibberellins (GA) are plant hormones that can regulate germination, elongation growth, and sex determination. They ubiquitously occur in seed plants. The discovery of gibberellin receptors, together with advances in understanding the function of key components of GA signalling in Arabidopsis and rice, reveal a fairly short GA signal transduction route. The pathway essentially consists of GID1 gibberellin receptors that interact with F-box proteins, which in turn regulate degradation of downstream DELLA proteins, suppressors of GA-controlled responses.     

Polyploidy-associated genome modifications during land plant evolution

The occurrence of polyploidy in land plant evolution has led to an acceleration of genome modifications relative to other crown eukaryotes and is correlated with key innovations in plant evolution. Extensive genome resources provide for relating genomic changes to the origins of novel morphological and physiological features of plants. Ancestral gene contents for key nodes of the plant family tree are inferred. Pervasive polyploidy in angiosperms appears likely to be the major factor generating novel angiosperm genes and expanding some gene families. However, most gene families lose most duplicated copies in a quasi-neutral process, and a few families are actively selected for single-copy status. One of the great challenges of evolutionary genomics is to link genome modifications to speciation, diversification and the morphological and/or physiological innovations that collectively compose biodiversity. Rapid accumulation of genomic data and its ongoing investigation may greatly improve the resolution at which evolutionary approaches can contribute to the identification of specific genes responsible for particular innovations. The resulting, more ‘particulate’ understanding of plant evolution, may elevate to a new level fundamental knowledge of botanical diversity, including economically important traits in the crop plants that sustain humanity.     

Parka decipiens and land plant spore evolution

The morphology and ultrastructure of the spores of the enigmatic Lower Devonian plant Parka decipiens are briefly described and previous interpretations of the plant are discussed. It is suggested that the alete spores of Parka could have been haploid or diploid. The possibility that some early plant spores were diploid may be viewed as supporting the interpolation theory for the origin of alternation of generations in land plants. The spores of Parka may lie (in morphological and developmental terms) somewhere between the oospores produced by algae and the triradiate haploid spores of the bryophytes.     

Adaptive walks toward a moving optimum

We investigate how the dynamics and outcomes of adaptation by natural selection are affected by environmental stability by simulating adaptive walks in response to an environmental change of fixed magnitude but variable speed. Here we consider monomorphic lineages that adapt by the sequential fixation of beneficial mutations. This is modeled by selecting short RNA sequences for folding stability and secondary structure conservation at increasing temperatures. Using short RNA sequences allows us to describe adaptive outcomes in terms of genotype (sequence) and phenotype (secondary structure) and to follow the dynamics of fitness increase. We find that slower rates of environmental change affect the dynamics of adaptive walks by reducing the fitness effect of fixed beneficial mutations, as well as by increasing the range of time in which the substitutions of largest effect are likely to occur. In addition, adaptation to slower rates of environmental change results in fitter endpoints with fewer possible end phenotypes relative to lineages that adapt to a sudden change. This suggests that care should be taken when experiments using sudden environmental changes are used to make predictions about adaptive responses to gradual change.