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Plant roots and root systems are indispensable for water and nutrient foraging, and are a major evolutionary achievement for plants to cope with dry land conditions. The ability of roots to branch contributes substantially to their capacity to explore the soil for water and nutrients, and led ~400 million years ago to the successful colonization of land by plants, eventually even in arid regions. During this colonization, different forms of root branching evolved, reinforcing step by step the phenotypic plasticity of the root system. Whereas the lycophytes, the most ancient land plants with roots, only branch at the root tip, ferns are able to form roots laterally in a fixed pattern along the main root. Finally, roots of seed plants show the highest phenotypic plasticity, because lateral roots can possibly, dependent on internal and/or external signals, be produced at almost any position along the main root. The competence to form lateral roots in seed plants is based on the presence of internal cell files with stem cell-like features. Despite the dissimilarities between the different clades, a number of genetic modules seem to be co-opted in order to acquire root branching capacity. In this review, starting from the lateral root pathways in seed plants, we review root branching in the different land plant lineages and discuss the hitherto described genetic modules that contribute to their root branching capacity. We try to obtain insight into how land plants have acquired an increasing root branching plasticity during evolution that contributed to the successful colonization of our planet by seed plants.

PREMISE OF THE STUDY Community phylogenetic methods incorporate information on evolutionary relationships into studies of organismal assemblages. We used a community phylogenetic framework to investigate relationships and biogeographic affinities and to calculate phylogenetic signal of endemism and invasiveness for the flora of the pine rocklands-a globally critically imperiled ecosystem with a significant portion of its distribution in South Florida, United States. METHODS We reconstructed phylogenetic relationships of 538 vascular plant taxa, which represent 92.28% of the vascular flora of the pine rocklands. We estimated phylogenetic signal for endemism and invasiveness using phylogenetic generalized linear mixed models. We determined the native range for each species in the data set and calculated the total number of species sourced from each region and all possible combinations of these regions. KEY RESULTS The pine rockland flora includes representatives of all major vascular plant lineages, and most species have native ranges in the New World. There was strong phylogenetic signal for endemism, but not for invasiveness. CONCLUSIONS Community phylogenetics has high potential value for conservation planning, particularly for fragmented and endangered ecosystems like the pine rockland. Strong phylogenetic signal for endemic species in our data set, which also tend to be threatened or endangered, can help to identify species at risk, as well as fragments where those species occur, highlighting conservation priorities. Our results indicate, at least in the pine rockland ecosystem, no phylogenetic signal for invasive species, and thus other information must be used to predict the potential for invasiveness.

Mitochondrial calcium uptake (MICU) plays a vital role in the regulation of mitochondrial calcium homeostasis, and, consequently, influences calcium signaling transduction. Although genes involved in mitochondrial calcium uptake have been well studied in animals, less is known about their ubiquity and function in plants. In this study, we identified 96 MICU genes in land plants. On the basis of phylogenetic analysis of MICU proteins, they were classified into three clades: MICU from eudicots (Clade I), from monocots (Clade II), and from a basal angiosperm, a bryophyte, and a lycophyte (Clade III). Pairwise identity analysis across all MICU proteins showed that they are highly conserved among land plants at the protein level. Conserved motif analysis showed that most MICU proteins contained three EF-hands, and an additional EF-hand motif first identified in the MICU of Arabidopsis thaliana but not mammals was found in all 96 putative MICU proteins. This suggests that a cellular pathway of calcium uptake and signaling that requires three EF-hand motifs is evolutionarily conserved in plants. In addition, we discovered that MICU-defective mutants of Arabidopsis thaliana exhibited longer roots than wild-type under high calcium stress. Concurrently, the mRNA transcription levels of MICU were decreased under high calcium conditions. These results suggest that loss-of-function mutations of MICU may have potential roles in helping plants resist high calcium stress. This study provides clues to the possible role of plant MICU in mitochondrial calcium uptake, as well as useful information to support further studies on MICU function in plants.

Cutin and suberin are hydrophobic lipid biopolyester components of the cell walls of specialized plant tissue and cell-types, where they facilitate adaptation to terrestrial habitats. Many steps in their biosynthetic pathways have been characterized, but the basis of their spatial deposition and precursor trafficking is not well understood. Members of the GDSL lipase/esterase family catalyze cutin polymerization, and candidate proteins have been proposed to mediate interactions between cutin or suberin and other wall components. Comparative genomic studies of charophyte algae and early diverging land plants, combined with knowledge of the biosynthesis, trafficking and assembly mechanisms, suggests an origin for the capacity to secrete waxes, as well as aliphatic and phenolic compounds before the first colonization of true terrestrial habitats.

It is generally accepted that land plants evolved from streptophyte algae. However, there are also many chlorophytes (a sister group of streptophyte algae and land plants) that moved to terrestrial habitats and even resemble mosses. This raises the question of why no land plants evolved from chlorophytes. In order to better understand what enabled streptophyte algae to conquer the land, it is necessary to study the chlorophytes as well. This review will introduce the freshwater filamentous chlorophyte alga Draparnaldia sp. (Chaetophorales, Chlorophyceae) as a model for comparative analyses between these two lineages. It will also focus on current knowledge about the evolution of morphological complexity in chlorophytes versus streptophytes and their respective morphological/behavioural adaptations to semi-terrestrial habitats, and will show why Draparnaldia is needed as a new model system.

Background Sucrose synthase (SUS) is widely considered a key enzyme participating in sucrose metabolism in higher plants and regarded as a biochemical marker for sink strength in crops. However, despite significant progress in characterizing the physiological functions of the SUS gene family, knowledge of the trajectory of evolutionary processes and significance of the family in higher plants remains incomplete. Results In this study, we identified over 100 SUS genes in 19 plant species and reconstructed their phylogenies, presenting a potential framework of SUS gene family evolution in higher plants. Three anciently diverged SUS gene subfamilies (SUS I, II and III) were distinguished based on their phylogenetic relationships and unique intron/exon structures in angiosperms, and they were found to have evolved independently in monocots and dicots. Each subfamily of SUS genes exhibited distinct expression patterns in a wide range of plants, implying that their functional differentiation occurred before the divergence of monocots and dicots. Furthermore, SUS III genes evolved under relaxed purifying selection in dicots and displayed narrowed expression profiles. In addition, for all three subfamilies of SUS genes, the GT-B domain was more conserved than the “regulatory” domain. Conclusions The present study reveals the evolution of the SUS gene family in higher plants and provides new insights into the evolutionary conservation and functional divergence of angiosperm SUS genes.

Aim: Plant trait databases often contain traits that are correlated, but for whom direct (undirected statistical dependency) and indirect (mediated by other traits) connections may be confounded. The confounding of correlation and connection hinders our understanding of plant strategies, and how these vary among growth forms and climate zones. We identified the direct and indirect connections across plant traits relevant to competition, resource acquisition and reproductive strategies using a global database and explored whether connections within and between traits from different tissue types vary across climates and growth forms. Location: Global. Major taxa studied: Plants. Time period: Present. Methods: We used probabilistic graphical models and a database of 10 plant traits (leaf area, specific leaf area, mass- and area-based leaf nitrogen and phosphorous content, leaf life span, plant height, stem specific density and seed mass) with 16,281 records to describe direct and indirect connections across woody and non-woody plants across tropical, temperate, arid, cold and polar regions. Results: Trait networks based on direct connections are sparser than those based on correlations. Land plants had high connectivity across traits within and between tissue types; leaf life span and stem specific density shared direct connections with all other traits. For both growth forms, two groups of traits form modules of more highly connected traits; one related to resource acquisition, the other to plant architecture and reproduction. Woody species had higher trait network modularity in polar compared to temperate and tropical climates, while non-woody species did not show significant differences in modularity across climate regions. Main conclusions: Plant traits are highly connected both within and across tissue types, yet traits segregate into persistent modules of traits. Variation in the modularity of trait networks suggests that trait connectivity is shaped by prevailing environmental conditions and demonstrates that plants of different growth forms use alternative strategies to cope with local conditions. © 2019 John Wiley and Sons Ltd

Mixotrophic plants combine photosynthesis and heterotrophic nutrition. Recent research suggests mechanisms explaining why mixotrophy is so common in terrestrial ecosystems. First, mixotrophy overcomes nutrient limitation and/or seedling establishment constraints. Second, although genetic drift may push mixotrophs to full heterotrophy, the role of photosynthesis in reproduction stabilizes mixotrophy.

International audience; In land plants, the aerial epidermis is essential for growth control, protection and environmental interactions. Epidermal cell fate is specified early during embryogenesis and maintained throughout plant life. Molecular actors of epidermal specification have been characterized, but how epidermal fate is maintained during growth remains unclear. DEFECTIVE KERNEL 1 (DEK1) is required for epidermal cell fate maintenance during both embryonic and postembryonic plant development. The activation of a mechanosensitive Ca2+ channel was recently shown to depend on DEK1, suggesting that the interpretation of mechanical cues could be critical for maintaining epidermal cell fate. Here, we integrate these findings into the epidermal specification network and propose a model explaining why epidermal specification may depend upon the sensing of epidermal tension.

The key structures and functions of land plants are most often studied in flowering plant models. However, the evolution of these traits (character states) is often difficult to infer, because we lack an accurate phylogenetic frame of reference. The potential branching order of the earliest land plants has now been further condensed, narrowing down potential reference frameworks for comparative studies.

Significance The evolution of land plants from algae is an age-old question in biology. The entire terrestrial flora stems from a grade of algae, the streptophyte algae. Recent phylogenomic studies have pinpointed the Zygnematophyceae as the modern-day streptophyte algal lineage that is most closely related to the algal land plant ancestor. Here, we provide insight into the biology of this ancestor that might have aided in its conquest of land. Specifically, we uncover the existence of stress-signaling pathways and the potential for intimate plastid-nucleus communication. Plastids act as environmental sensors in land plants; our data suggest that this feature was present in a common ancestor they shared with streptophyte algae., Streptophytes are unique among photosynthetic eukaryotes in having conquered land. As the ancestors of land plants, streptophyte algae are hypothesized to have possessed exaptations to the environmental stressors encountered during the transition to terrestrial life. Many of these stressors, including high irradiance and drought, are linked to plastid biology. We have investigated global gene expression patterns across all six major streptophyte algal lineages, analyzing a total of around 46,000 genes assembled from a little more than 1.64 billion sequence reads from six organisms under three growth conditions. Our results show that streptophyte algae respond to cold and high light stress via expression of hallmark genes used by land plants (embryophytes) during stress–response signaling and downstream responses. Among the strongest differentially regulated genes were those associated with plastid biology. We observed that among streptophyte algae, those most closely related to land plants, especially Zygnema, invest the largest fraction of their transcriptional budget in plastid-targeted proteins and possess an array of land plant-type plastid-nucleus communication genes. Streptophyte algae more closely related to land plants also appear most similar to land plants in their capacity to respond to plastid stressors. Support for this notion comes from the detection of a canonical abscisic acid receptor of the PYRABACTIN RESISTANCE (PYR/PYL/RCAR) family in Zygnema, the first found outside the land plant lineage. We conclude that a fine-tuned response toward terrestrial plastid stressors was among the exaptations that allowed streptophytes to colonize the terrestrial habitat on a global scale.

International audience; This study introduces a new Bayesian technique for molecular dating that explicitly accommodates for uncertainty in the phylogenetic position of calibrated nodes derived from the analysis of fossil data. The proposed approach thus defines an adequate framework for incorporating expert knowledge and/or prior information about the way fossils were collected in the inference of node ages. Although it belongs to the class of “node-dating” approaches, this method shares interesting properties with “tip-dating” techniques. Yet, it alleviates some of the computational and modeling difficulties that hamper tip-dating approaches. The influence of fossil data on the probabilistic distribution of trees is the crux of the matter considered here. More specifically, among all the phylogenies that a tree model (e.g., the birth–death process) generates, only a fraction of them “agree” with the fossil data. Bayesian inference under the new model requires taking this fraction into account. However, evaluating this quantity is difficult in practice. A generic solution to this issue is presented here. The proposed approach relies on a recent statistical technique, the so-called exchange algorithm, dedicated to drawing samples from “doubly intractable” distributions. A small example illustrates the problem of interest and the impact of uncertainty in the placement of calibration constraints in the phylogeny given fossil data. An analysis of land plant sequences and multiple fossils further highlights the pertinence of the proposed approach.

Biochemical defenses against oomycete infection were already present in early diverging plant lineages, suggesting parallel selection for phenylpropanoid metabolites.

Nitric oxide (NO) is a key signaling molecule in plant physiology. However, its production in photosynthetic organisms remains partially unresolved. The best characterized NO production route involves the reduction of nitrite to NO via different non-enzymatic or enzymatic mechanisms. Nitrate reductases (NRs), the mitochondrial electron transport chain, and the new complex between NR and NOFNiR (nitric oxide-forming nitrite reductase) described in Chlamydomonas reinhardtii are the main enzymatic systems that perform this reductive NO production in plants. Apart from this reductive route, several reports acknowledge the possible existence of an oxidative NO production in an arginine-dependent pathway, similar to the nitric oxide synthase (NOS) activity present in animals. However, no NOS homologs have been found in the genome of embryophytes and, despite an increasing amount of evidence attesting to the existence of NOS-like activity in plants, the involved proteins remain to be identified. Here we review NO production in plants with emphasis on the presentation and discussion of recent data obtained in this field.

One signature of life on Earth is the vegetation red edge (VRE) feature of land plants, a dramatic change of reflectivity at wavelength near 0.7 μm. Potentially habitable planets around M dwarfs are tidally locked, which can limit the distribution of land plants. In this study, we used a biogeochemical model to investigate the distribution of land plants on potentially habitable planets around M dwarfs driven by climate data produced in a general circulation model (GCM). When considering the effects of clouds, the observation time needed for VRE detection on nearby p = 1 exoplanets around nearby M dwarfs is on the order of days using a 25 m2 telescope if a large continent faces Earth during observations. For p = 1.5 exoplanets, the detection time could be similar if land plants developed the capability to endure a dark/cold environment for extended periods of time and the continent configuration favors observations. Our analysis suggests that hypothetical exovegetation VRE features are easier to ...

Tethering complexes mediate the initial, specific contact between donor and acceptor membranes. This review focuses on the modularity and function of multisubunit tethering complexes (MTCs) in higher plants. One emphasis is on molecular interactions of plant MTCs. Here, a number of insights have been gained concerning interactions between different tethering complexes, and between tethers and microtubule-associated proteins. The roles of tethering complexes in abiotic stress responses appear indirect, but in the context of biotic stress responses it has been suggested that some tethers are direct targets of pathogen effectors or virulence factors. In light of the central roles tethering complexes play in plant development, an emerging concept is that tethers may be co-opted for plant adaptive responses.

MicroRNA156 (miR156) and miR529 have high sequence similarity and recognize overlapping sites in the same target genes, SQUAMOSA promoter binding protein-like (SPL or SBP box) genes, making it difficult to accurately distinguish their roles in regulatory networks that affect numerous biological functions. Here, we collected data about miR156 and miR529 family members from representative land plants and performed sequence comparisons, phylogenetic analysis, small RNA sequencing, and parallel analysis of RNA ends (PARE) analysis to dissect their evolutionary and functional differences. Although miR156 and miR529 are highly similar, there are differences in their mismatch-sensitive regions, which are essential for target recognition. In land plants, miR156 precursors are conserved mainly within the hairpin region, whereas miR529 precursors are conserved outside the hairpin region, including both the 5’ and 3’ arms. Phylogenetic analysis showed that MIR156 and MIR529 evolved independently, through divergent evolutionary patterns. The two genes also exhibit different expression patterns, with MIR529 preferentially expressed in reproductive tissues and MIR156 in other tissues. PARE analysis revealed that miR156 and miR529 possess specific targets in addition to common targets in maize, pointing to functional differences between them. Based on our findings, we developed a method for the rapid identification of miR529 and miR156 family members and uncovered the evolutionary divergence of these families, providing insights into their different regulatory roles in plant growth and development.

By a detailed ontogenetic study of Polemonium caeruleum pollen, tracing each stage of development at high TEM resolution, we aim to understand the establishment of the pollen wall and to unravel the mechanisms underlying sporoderm development. The main steps of exine ontogeny in Polemonium caeruleum, observed in the microspore periplasmic space, are spherical units, gradually transforming into columns, then to rod-like units (procolumellae), the appearance of the initial tectum, growth of columellae in height and tectum in thickness and initial sporopollenin accumulation on them, the appearance of the endexine lamellae and of dark-contrasted particles on the tectum, the appearance of a sponge-like layer and of the intine in aperture sites, the appearance of the foot layer on the base of the sponge-like layer and of spinules on the tectum, and massive sporopollenin accumulation. This sequence of developmental events fits well to the sequence of self-assembling micellar mesophases. This gives (together with earlier findings and experimental exine simulations) strong evidence that genome and self-assembly probably share control of exine formation. It is highly probable that self-assembly is an intrinsic instrument of evolution.

There is a public perception that large high-severity wildfires decrease biodiversity and increase fire hazard by homogenizing vegetation composition and increasing the cover of mid-story vegetation. But a growing literature suggests that vegetation responses are nuanced. LiDAR technology provides a promising remote sensing tool to test hypotheses about post-fire vegetation regrowth because vegetation cover can be quantified within different height strata at fine scales over large areas. We assess the usefulness of airborne LiDAR data for measuring post-fire mid-story vegetation regrowth over a range of spatial resolutions (10 × 10 m, 30 × 30 m, 50 × 50 m, 100 × 100 m cell size) and investigate the effect of fire severity on regrowth amount and spatial pattern following a mixed severity wildfire in Warrumbungle National Park, Australia. We predicted that recovery would be more vigorous in areas of high fire severity, because park managers observed dense post-fire regrowth in these areas. Moderate to strong positive associations were observed between LiDAR and field surveys of mid-story vegetation cover between 0.5-3.0 m. Thus our LiDAR survey was an apt representation of on-ground vegetation cover. LiDAR-derived mid-story vegetation cover was 22-40% lower in areas of low and moderate than high fire severity. Linear mixed-effects models showed that fire severity was among the strongest biophysical predictors of mid-story vegetation cover irrespective of spatial resolution. However much of the variance associated with these models was unexplained, presumably because soil seed banks varied at finer scales than our LiDAR maps. Dense patches of mid-story vegetation regrowth were small (median size 0.01 ha) and evenly distributed between areas of low, moderate and high fire severity, demonstrating that high-severity fires do not homogenize vegetation cover. Our results are relevant for ecosystem conservation and fire management because they: indicate that native vegetation are responsive and resilient to high-severity fire, and show the usefulness of remote sensing tools such as LiDAR to monitor post-fire vegetation recovery over large areas in situ.

In land plants, photosystem I (PSI) photoinhibition limits carbon fixation and causes growth defects. In addition, recovery from PSI photoinhibition takes much longer than PSII photoinhibition when the PSI core-complex is degraded by oxidative damage. Accordingly, PSI photoinhibition should be avoided in land plants, and land plants should have evolved mechanisms to prevent PSI photoinhibition. However, such protection mechanisms have not yet been identified, and it remains unclear whether all land plants suffer from PSI photoinhibition in the same way. In the present study, we focused on the susceptibility of PSI to photoinhibition and investigated whether mechanisms of preventing PSI photoinhibition varied among land plant species. To assess the susceptibility of PSI to photoinhibition, we used repetitive short-pulse (rSP) illumination, which specifically induces PSI photoinhibition. Subsequently, we found that land plants possess a wide variety of tolerance mechanisms against PSI photoinhibition. In particular, gymnosperms, ferns and mosses/liverworts exhibited higher tolerance to rSP illumination-induced PSI photoinhibition than angiosperms, and detailed analyses indicated that the tolerance of these groups could be partly attributed to flavodiiron proteins, which protected PSI from photoinhibition by oxidizing the PSI reaction center chlorophyll (P700) as an electron acceptor. Furthermore, we demonstrate, for the first time, that gymnosperms, ferns and mosses/liverworts possess a protection mechanism against photoinhibition of PSI that differs from that of angiosperms.

Growth and productivity in important crop plants is limited by the inefficiencies of the C3 photosynthetic pathway. Introducing CO2-concentrating mechanisms (CCMs) into C3 plants could overcome these limitations and lead to increased yields. Many unicellular microautotrophs, such as cyanobacteria and green algae, possess highly efficient biophysical CCMs that increase CO2 concentrations around the primary carboxylase enzyme, Rubisco, to enhance CO2 assimilation rates. Algal and cyanobacterial CCMs utilize distinct molecular components, but share several functional commonalities. Here we outline the recent progress and current challenges of engineering biophysical CCMs into C3 plants. We review the predicted requirements for a functional biophysical CCM based on current knowledge of cyanobacterial and algal CCMs, the molecular engineering tools and research pipelines required to translate our theoretical knowledge into practice, and the current challenges to achieving these goals.

The evolution of active stomatal closure in response to leaf water deficit, mediated by the hormone abscisic acid (ABA), has been the subject of recent debate. Two different models for the timing of the evolution of this response recur in the literature. A single-step model for stomatal control suggests that stomata evolved active, ABA-mediated control of stomatal aperture, when these structures first appeared, prior to the divergence of bryophyte and vascular plant lineages. In contrast, a gradualistic model for stomatal control proposes that the most basal vascular plant stomata responded passively to changes in leaf water status. This model suggests that active ABA-driven mechanisms for stomatal responses to water status instead evolved after the divergence of seed plants, culminating in the complex, ABA-mediated responses observed in modern angiosperms. Here we review the findings that form the basis for these two models, including recent work that provides critical molecular insights into resolving this intriguing debate, and find strong evidence to support a gradualistic model for stomatal evolution.

Lignin, one of the most abundant biopolymers on Earth, derives from the plant phenolic metabolism. It appeared upon terrestrialization and is thought critical for plant colonization of land. Early diverging land plants do not form lignin, but already have elements of its biosynthetic machinery. Here we delete in a moss the P450 oxygenase that defines the entry point in angiosperm lignin metabolism, and find that its pre-lignin pathway is essential for development. This pathway does not involve biochemical regulation via shikimate coupling, but instead is coupled with ascorbate catabolism, and controls the synthesis of the moss cuticle, which prevents desiccation and organ fusion. These cuticles share common features with lignin, cutin and suberin, and may represent the extant representative of a common ancestor. Our results demonstrate a critical role for the ancestral phenolic metabolism in moss erect growth and cuticle permeability, consistent with importance in plant adaptation to terrestrial conditions., The phenolic polymer lignin is thought to have contributed to adaptation of early land plants to terrestrial environments. Here Renault et al. show that moss, which does not produce lignin, contains an ancestral phenolic metabolism pathway that produces a phenol-enriched cuticle and prevents desiccation.

The divergence of land plants followed by vascular plants has entirely changed the terrestrial ecology. The vascular system is a prerequisite for this evolutionary event, providing upright stature and communication for sink demand-source capacity and facilitating the development of plants and colonization over a wide range of environmental habitats. Various hormonal and non-hormonal regulatory networks have been identified and reviewed as key processes for vascular formation; however, how these factors have evolutionarily emerged and interconnected to trigger the emergence of the vascular system still remains elusive. Here, to understand the intricacy of cross-talks among these factors, we highlight how core hormonal signaling and transcriptional networks are coalesced into the appearance of vascular plants during evolution.

One crucial problem that plants faced during their evolution, particularly during the transition to growth on land, was how to transport water, nutrients, metabolites, and small signaling molecules within a large, multicellular body. As a solution to this problem, land plants developed specific tissues for conducting molecules, called water-conducting cells (WCCs) and food-conducting cells (FCCs). The well-developed WCCs and FCCs in extant plants are the tracheary elements and sieve elements, respectively, which are found in vascular plants. Recent molecular genetic studies revealed that transcriptional networks regulate the differentiation of tracheary and sieve elements, and that the networks governing WCC differentiation are largely conserved among land plant species. In this review, we discuss the molecular evolution of plant conducting cells. By focusing on the evolution of the key transcription factors that regulate vascular cell differentiation, the NAC transcription factor VASCULAR-RELATED NAC-DOMAIN for WCCs and the MYB-coiled-coil (CC)-type transcription factor ALTERED PHLOEM DEVELOPMENT for sieve elements, we describe how land plants evolved molecular systems to produce the specialized cells that function as WCCs and FCCs.

The colonization of land by vascular plants is an extremely important phase in Earth’s life history. This key evolutionary process is thought to have begun during the Middle Cambrian 1 period and culminated in the Silurian/Early Devonian period (interval about 509–393 million years ago (Ma)), and is documented primarily by microfossils (that is, by dispersed spores, phytodebris including fragments of algae, tissues, sporangia and cuticles), tubes and rare megafossils 2 . A newly recognized fossil cooksonioid plant with in situ spores from the Barrandian area, Czech Republic, is of the highest importance because it represents extremely ancient megafossil evidence of land plant diploid generation: sporophytes (~432 Ma). The robust size of this plant places it among the largest known early polysporangiate land plants and it is probable that it attained adequate size for both aeration and effective photosynthetic competence. This would mean not only that sporophytes were photosynthetically autonomous but also that the they might have been able to sustain a relatively gametophyte-independent existence. Ancient fossil Cooksonia plants from the Czech Republic are among the largest known early polysporangiate plants from when vascular plants were colonizing the land. They were of sufficient size to support effective photosynthesis.

The Arabidopsis LSH1 and Oryza G1 (ALOG) protein is a family of plant-specific transcription factors that regulate reproductive growth in angiosperms. Despite their importance in plant development, little research has been conducted on ALOG proteins in basal land plants and the processes involved in their evolution remain largely unknown. Here, we studied the molecular evolution of ALOG family proteins. We found that ALOG proteins are absent in green algae but exist in all land plants analyzed as well as in some Charophycean algae, closest relatives of land plants. Multiple sequence alignments identified the high sequence conservation of ALOG domains in divergent plant lineages. Phylogenetic analyses also identified a distinct clade of ALOG protein member of lycophytes and bryophytes, including two of Marchantia polymorpha LATERAL ORGAN SUPPRESOR (MpLOS1 and MpLOS2) with a long branch length in MpLOS2. Consistent with this, the function of MpLOS1 was replaceable by Phycomitrella patens ALOG proteins, whereas MpLOS2 failed to replace the molecular function of MpLOS1. Moreover, the rice ALOG proteins, OsTAW1 and OsG1, were not able to replace the molecular function of MpLOS1 although we previously found that the function of OsG1 was replaceable by MpLOS1. Altogether, these findings suggest that ALOG proteins emerged before the evolution of land plants and that they exhibit functional conservation and diversification during the evolution of land plants. The finding that MpLOS1 is able to complement rice ALOG mutants but not vice versa also suggest the existence of conserved and the partly divergent functions of ALOG proteins in bryophytes and angiosperms.

The chloroplast genome usually has a quadripartite structure consisting of a large single copy region and a small single copy region separated by two long inverted repeats. It has been known for some time that a single cell may contain at least two structural haplotypes of this structure, which differ in the relative orientation of the single copy regions. However, the methods required to detect and measure the abundance of the structural haplotypes are labour-intensive, and this phenomenon remains understudied. Here we develop a new method, Cp-hap, to detect all possible structural haplotypes of chloroplast genomes of quadripartite structure using long-read sequencing data. We use this method to conduct a systematic analysis and quantification of chloroplast structural haplotypes in 61 land plant species across 19 orders of Angiosperms, Gymnosperms and Pteridophytes. Our results show that there are two chloroplast structural haplotypes which occur with equal frequency in most land plant individuals. Nevertheless, species whose chloroplast genomes lack inverted repeats or have short inverted repeats have just a single structural haplotype. We also show that the relative abundance of the two structural haplotypes remains constant across multiple samples from a single individual plant, suggesting that the process which maintains equal frequency of the two haplotypes operates rapidly, consistent with the hypothesis that flip-flop recombination mediates chloroplast structural heteroplasmy. Our results suggest that previous claims of differences in chloroplast genome structure between species may need to be revisited.

The transition to a terrestrial environment, termed terrestrialization, is generally regarded as a pivotal event in the evolution and diversification of the land plant flora that changed the surface of our planet. Through phylogenomic studies, a group of streptophyte algae, the Zygnematophyceae, have recently been recognized as the likely sister group to land plants (embryophytes). Here, we report genome sequences and analyses of two early diverging Zygnematophyceae (Spirogloea muscicola gen. nov. and Mesotaenium endlicherianum) that share the same subaerial/terrestrial habitat with the earliest-diverging embryophytes, the bryophytes. We provide evidence that genes (i.e., GRAS and PYR/PYL/RCAR) that increase resistance to biotic and abiotic stresses in land plants, in particular desiccation, originated or expanded in the common ancestor of Zygnematophyceae and embryophytes, and were gained by horizontal gene transfer (HGT) from soil bacteria. These two Zygnematophyceae genomes represent a cornerstone for future studies to understand the underlying molecular mechanism and process of plant terrestrialization.

International audience; Genome packaging by nucleosomes is a hallmark of eukaryotes. Histones and the pathways that deposit, remove, and read histone modifications are deeply conserved. Yet, we lack information regarding chromatin landscapes in extant representatives of ancestors of the main groups of eukaryotes, and our knowledge of the evolution of chromatin-related processes is limited. We used the bryophyte Marchantia polymorpha, which diverged from vascular plants circa 400 mya, to obtain a whole chromosome genome assembly and explore the chromatin landscape and three-dimensional genome organization in an early diverging land plant lineage. Based on genomic profiles of ten chromatin marks, we conclude that the relationship between active marks and gene expression is conserved across land plants. In contrast, we observed distinctive features of transposons and other repetitive sequences in Marchantia compared with flowering plants. Silenced transposons and repeats did not accumulate around centromeres. Although a large fraction of constitutive heterochromatin was marked by H3K9 methylation as in flowering plants, a significant proportion of transposons were marked by H3K27me3, which is otherwise dedicated to the transcriptional repression of protein-coding genes in flowering plants. Chromatin compartmentalization analyses of Hi-C data revealed that repressed B compartments were densely decorated with H3K27me3 but not H3K9 or DNA methylation as reported in flowering plants. We conclude that, in early plants, H3K27me3 played an essential role in heterochromatin function, suggesting an ancestral role of this mark in transposon silencing.

The class of conjugating green algae, Zygnematophyceae (Conjugatophyceae), is extremely rich in species and has attracted the interest of phycologists for a long time. It is now widely accepted that this class of charophyte algae holds a key position in the phylogenetic tree of streptophytes, where they represent the closest relatives to all land plants (embryophytes). It is increasingly evident that robust model plants that can be easily cultivated and genetically transformed are necessary to better understand the process of terrestrialization and the related molecular, cellular, and physiological adaptations. Living algae collections play an important role, not only for phylogenomic-based taxonomy but also for screening for suitable model organisms. For this review, we screened six major public algae collections for Zygnematophyceae strains and established a cumulative list comprising 738 different taxa (including species, subspecies, varieties, and forms). From the described biodiversity with 8883 registered taxa (AlgaeBase) the cultured Zygnematophyceae taxa worldwide cover only ~8.3%. We review the past research on this clade of algae and discuss it from the perspective of establishing a model organism. We present data on the life cycle of the genera Micrasterias and Spirogyra, representing the orders Desmidiales and Zygnematales, and outline the current status of genetic transformation of Zygnematophyceae algae and future research perspectives.

Adaptations to environmental stress such as desiccation are needed for land plant transition. Late embryogenesis abundant (LEA) proteins and major intrinsic proteins (MIPs) diversified in ancestors of early land plants., The present review summarizes the effects of desiccation in streptophyte green algae, as numerous experimental studies have been performed over the past decade particularly in the early branching streptophyte Klebsormidium sp. and the late branching Zygnema circumcarinatum. The latter genus gives its name to the Zygenmatophyceae, the sister group to land plants. For both organisms, transcriptomic investigations of desiccation stress are available, and illustrate a high variability in the stress response depending on the conditions and the strains used. However, overall, the responses of both organisms to desiccation stress are very similar to that of land plants. We highlight the evolution of two highly regulated protein families, the late embryogenesis abundant (LEA) proteins and the major intrinsic protein (MIP) family. Chlorophytes and streptophytes encode LEA4 and LEA5, while LEA2 have so far only been found in streptophyte algae, indicating an evolutionary origin in this group. Within the MIP family, a high transcriptomic regulation of a tonoplast intrinsic protein (TIP) has been found for the first time outside the embryophytes in Z. circumcarinatum. The MIP family became more complex on the way to terrestrialization but simplified afterwards. These observations suggest a key role for water transport proteins in desiccation tolerance of streptophytes.

One of the most transformative events in the history of life on earth was the transition of plants from water to land approximately 470 million years ago. Within the Charophyte green algae, the closest living relatives of land plants, body plans have evolved from those that comprise simple unicells to those that are morphologically complex, large and multicellular. The Charophytes developed these broad ranging body plans by exploiting a range of one-dimensional and two-dimensional growth strategies to produce filaments, mats and branches. When plants were confronted with harsh conditions on land, they were required to make significant changes to the way they shaped their body plans. One of the fundamental developmental transitions that occurred was the evolution of three-dimensional growth and the acquisition of apical cells with three or more cutting faces. Plants subsequently developed a range of morphological adaptations (e.g. vasculature, roots, flowers, seeds) that enabled them to colonise progressively drier environments. 3D apical growth also evolved convergently in the brown algae, completely independently of the green lineage. This review summarises the evolving developmental complexities observed in the early divergent Charophytes all the way through to the earliest conquerors of land, and investigates 3D apical growth in the brown algae.

Localized delivery of plasma membrane and cell wall components is an essential process in all plant cells. The vesicle-tethering complex, the exocyst, an ancient eukaryotic hetero-octameric protein cellular module, assists in targeted delivery of exocytosis vesicles to specific plasma membrane domains. Analyses of Arabidopsis and later other land plant genomes led to the surprising prediction of multiple putative EXO70 exocyst subunit paralogues. All land plant EXO70 exocyst subunits (including those of Bryophytes) form three distinct subfamilies-EXO70.1, EXO70.2, and EXO70.3. Interestingly, while the basal well-conserved EXO70.1 subfamily consists of multiexon genes, the remaining two subfamilies contain mostly single exon genes. Published analyses as well as public transcriptomic and proteomic data clearly indicate that most cell types in plants express and also use several different EXO70 isoforms. Here we sum up recent advances in the characterization of the members of the family of plant EXO70 exocyst subunits and present evidence that members of the EXO70.2 subfamily are often recruited to non-canonical functions in plant membrane trafficking pathways. Engagement of the most evolutionarily dynamic EXO70.2 subfamily of EXO70s in biotic interactions and defence correlates well with massive proliferation and conservation of new protein variants in this subfamily.

Since higher plants are unable to move in their environment, interesting and mysterious developmental events (allelopathy, phototropism, apical dominance, nyctinasty, flowering, senescence) are observed in their life cycle. Chemical compounds released from plant organs to the neighboring environment stimulate or suppress the development and/or growth of other plants; this chemical interaction is called "allelopathy." The bending of the organs of a plant toward the light is a well-known phenomenon called "phototropism." The growth of lateral buds of some plants is normally repressed by a strongly growing main shoot apex and is the so-called "apical dominance". Plants open their leaves during the day and close them at night as if sleeping is known as "nyctinasty." Herein, recent studies on isolation and structure elucidation of bioactive compounds involved in their life cycle and determination of the molecular mechanisms for these developmental events are described.

BackgroundIn contrast to the highly labile mitochondrial (mt) genomes of vascular plants, the architecture and composition of mt genomes within the main lineages of bryophytes appear stable and invariant. The available mt genomes of 18 liverwort accessions representing nine genera and five orders are syntenous except forGymnomitrion concinnatumwhose genome is characterized by two rearrangements. Here, we expanded the number of assembled liverwort mt genomes to 47, broadening the sampling to 31 genera and 10 orders spanning much of the phylogenetic breadth of liverworts to further test whether the evolution of the liverwort mitogenome is overall static.ResultsLiverwort mt genomes range in size from 147 Kb in Jungermanniales (clade B) to 185 Kb in Marchantiopsida, mainly due to the size variation of intergenic spacers and number of introns. All newly assembled liverwort mt genomes hold a conserved set of genes, but vary considerably in their intron content. The loss of introns in liverwort mt genomes might be explained by localized retroprocessing events. Liverwort mt genomes are strictly syntenous in genome structure with no structural variant detected in our newly assembled mt genomes. However, by screening the paired-end reads, we do find rare cases of recombination, which means multiple concurrent genome structures may exist in the vegetative tissues of liverworts. Our phylogenetic analyses of the nuclear encoded double stand break repair protein families revealed liverwort-specific subfamilies expansions.ConclusionsThe low repeat recombination level, selection, along with the intensified nuclear surveillance, might together shape the structural evolution of liverwort mt genomes.

The role of understory plants in native forests is critical for ecosystem function,wildlife protection and ecosystem productivity. The interest to estimate biodiversity increased during the last decades at landscape level. The objective was to elaborate a map of potential biodiversity (MPB) of understory species of Nothofagus forest using potential habitat suitability maps (PHS) of 15 plants in Santa Cruz province, Argentina. Additionally, we asked the following questions: (i) Were plant species differentially distributed according to the forest types?, (ii) do forest types represent different plant species assemblagewith specific ecological niche requirements?, and (iii) is it possibleto detect hotspots in the MBP according to the forest types?We used 721 plots database of vascular plants, fromwhere 15 indicator specieswere identified. The assemblage species for different forests (Nothofagus antarctica, N. pumilio and evergreen mixed) were analysed using a detrended correspondence analysis. Also, we explored 41 potential explanatory variables to develop PHS, and combined these maps to obtain one MPB (1?100%). Finally, we analysed the outputs into a GIS through different landscapes alternatives to detect hotspot areas. Marginality and specialization values allowed identifying species assemblage that presented similar variability in the habitat requirements. MPB varied across the landscape, with higher values in the south and lower values near glaciers. MPB had the highest values in N. antarctica forest with N50% cover at landscape level. N. antarctica present more hotspots than N. pumilio forests, mainly in the south, compared to mixed evergreen forests which present few hotspots near glaciers. These results can be used as a tool to design newmanagement and conservation strategies at landscape level. Fil: Rosas, Yamina Micaela. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; Argentina Fil: Peri, Pablo Luis. Instituto Nacional de Tecnología Agropecuaria; Argentina. Universidad Nacional de la Patagonia Austral; Argentina. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina Fil: Lencinas, María Vanessa. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; Argentina Fil: Martínez Pastur, Guillermo José. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Austral de Investigaciones Científicas; Argentina

Background Strigolactones (SLs) are an important class of carotenoid-derived signalling molecule in plants, which function both as exogenous signals in the rhizosphere and as endogenous plant hormones. In flowering plants, SLs are synthesized by a core pathway of four enzymes and are perceived by the DWARF14 (D14) receptor, leading to degradation of SMAX1-LIKE7 (SMXL7) target proteins in a manner dependent on the SCFMAX2 ubiquitin ligase. The evolutionary history of SLs is poorly understood, and it is not clear whether SL synthesis and signalling are present in all land plant lineages, nor when these traits evolved. Results We have utilized recently-generated genomic and transcriptomic sequences from across the land plant clade to resolve the origin of each known component of SL synthesis and signalling. We show that all enzymes in the core SL synthesis pathway originated at or before the base of land plants, consistent with the previously observed distribution of SLs themselves in land plant lineages. We also show that the late-acting enzyme LATERAL BRANCHING OXIDOREDUCTASE (LBO) may be considerably more ancient than previously thought. We perform a detailed phylogenetic analysis of SMXL proteins and show that specific SL target proteins only arose in flowering plants. We also assess diversity and protein structure in the SMXL family, identifying several previously unknown clades. Conclusions Overall, our results suggest that SL synthesis is much more ancient than canonical SL signalling, consistent with the idea that SLs first evolved as rhizosphere signals and were only recruited much later as hormonal signals. Electronic supplementary material The online version of this article (10.1186/s12915-019-0689-6) contains supplementary material, which is available to authorized users.

Repetitive sequences are ubiquitous components of eukaryotic genomes affecting genome size and evolution as well as gene regulation. Among them, short interspersed nuclear elements (SINEs) are non-coding retrotransposons usually shorter than 1000 bp. They contain only few short conserved structural motifs, in particular an internal promoter derived from cellular RNAs and a mostly AT-rich 3' tail, whereas the remaining regions are highly variable. SINEs emerge and vanish during evolution, and often diversify into numerous families and subfamilies that are usually specific for only a limited number of species. In contrast, at the 3' end of multiple plant SINEs we detected the highly conserved 'Angio-domain'. This 37 bp segment defines the Angio-SINE superfamily, which encompasses 24 plant SINE families widely distributed across 13 orders within the plant kingdom. We retrieved 28 433 full-length Angio-SINE copies from genome assemblies of 46 plant species, frequently located in genes. Compensatory mutations in and adjacent to the Angio-domain imply selective restraints maintaining its RNA structure. Angio-SINE families share segmental sequence similarities, indicating a modular evolution with strong Angio-domain preservation. We suggest that the conserved domain contributes to the evolutionary success of Angio-SINEs through either structural interactions between SINE RNA and proteins increasing their transpositional efficiency, or by enhancing their accumulation in genes.

Adaptor protein complexes and the related complexes COPI and TSET function in packaging vesicles for transport among endomembrane compartments in eukaryotic cells. Differences in the complement of these complexes in lineages such as yeast and mammals as well as apicomplexan and kinetoplastid parasites via loss or duplication of subunits appears to reflect specialization in their respective trafficking systems. The model plant Arabidopsis thaliana possesses multiple paralogues for adaptor protein complex subunits, raising questions as to the timing and extent of these duplications in embryophytes (land plants). However, adaptor protein complex evolution in embryophytes is unexplored. Therefore, we analyzed genomes of diverse embryophytes and closely related green algae using extensive homology searches and phylogenetic analysis of 35 complex subunit proteins. The results reveal numerous paralogues, the vast majority of which, approximately 97%, arose from recent duplication events. This suggests that specialization of these protein complexes may occur frequently but independently in embryophytes.

RNA editing alters the identity of nucleotides in an RNA sequence so that the mature transcript differs from the template defined in the genome. This process has been observed in chloroplasts and mitochondria of both seed and early land plants. However, the frequency of RNA editing in plant mitochondria ranges from zero to thousands of editing sites. To date, analyses of RNA editing in mitochondria of early land plants have been conducted on a small number of genes or mitochondrial genomes of a single species. This study provides an overview of the mitogenomic RNA editing potential of the main lineages of these two groups of early land plants by predicting the RNA editing sites of 33 mitochondrial genes of 37 species of liverworts and mosses. For the purpose of the research, we newly assembled seven mitochondrial genomes of liverworts. The total number of liverwort genera with known complete mitogenome sequences has doubled and, as a result, the available complete mitogenome sequences now span almost all orders of liverworts. The RNA editing site predictions revealed that C-to-U RNA editing in liverworts and mosses is group-specific. This is especially evident in the case of liverwort lineages. The average level of C-to-U RNA editing appears to be over three times higher in liverworts than in mosses, while the C-to-U editing frequency of the majority of genes seems to be consistent for each gene across bryophytes.