Studies of Physcomitrella patens reveal that ethylene‐mediated submergence responses arose relatively early in land‐plant evolution

Colonization of the land by multicellular green plants was a fundamental step in the evolution of life on earth. Land plants evolved from fresh‐water aquatic algae, and the transition to a terrestrial environment required the acquisition of developmental plasticity appropriate to the conditions of water availability, ranging from drought to flood. Here we show that extant bryophytes exhibit submergence‐induced developmental plasticity, suggesting that submergence responses evolved relatively early in the evolution of land plants. We also show that a major component of the bryophyte submergence response is controlled by the phytohormone ethylene, using a perception mechanism that has subsequently been conserved throughout the evolution of land plants. Thus a plant environmental response mechanism with major ecological and agricultural importance probably had its origins in the very earliest stages of the colonization of the land.     

Cereal mycorrhiza: an ancient symbiosis in modern agriculture

The majority of terrestrial plants live in association with symbiotic fungi that facilitate mineral nutrient uptake. The oldest and most prevalent of these associations are the arbuscular mycorrhizal (AM) symbioses that first evolved approximately 400 million years ago, coinciding with the appearance of the first land plants. Crop domestication, in comparison, is a relatively recent event, beginning approximately 10000 years ago. How has the dramatic change from wild to cultivated ecosystems impacted AM associations, and do these ancient symbioses potentially have a role in modern agriculture? Here, we review recent advances in AM research and the use of breeding approaches to generate new crop varieties that enhance the agronomic potential of AM associations.     

Origin of land plants: Do conjugating green algae hold the key?

Background  

The terrestrial habitat was colonized by the ancestors of modern land plants about 500 to 470 million years ago. Today it is widely accepted that land plants (embryophytes) evolved from streptophyte algae, also referred to as charophycean algae. The streptophyte algae are a paraphyletic group of green algae, ranging from unicellular flagellates to morphologically complex forms such as the stoneworts (Charales). For a better understanding of the evolution of land plants, it is of prime importance to identify the streptophyte algae that are the sister-group to the embryophytes. The Charales, the Coleochaetales or more recently the Zygnematales have been considered to be the sister group of the embryophytes However, despite many years of phylogenetic studies, this question has not been resolved and remains controversial.     

INDEPENDENT EVOLUTION OF TERRESTRIALITY IN ATLANTIC TRUNCATELLID GASTROPODS

Phylogenetic analysis of truncatellid gastropods using comparative anatomy and ribosomal RNA sequences shows that terrestrial truncatellids likely evolved three times independently in the Caribbean. The terrestrial subfamily Geomelaniinae, characterized in part by pallial fertilization and uniquely derived features of radula and protoconch, occurs in the Greater Antilles and Cayman Islands. Truncatellinae, with renopericardial fertilization, has several widespread amphibious species and two terrestrial species restricted to Trinidad and Barbados. The species in Barbados may be the most recent animal species to evolve full terrestriality; Barbados emerged above sea level only about one million years ago. By the mid-Cenozoic, truncatellids had traits enabling them to colonize land in appropriate tectonic settings. Parallel trends in character evolution occurred in the terrestrial lineages. In older terrestrial radiations, transitional character states would likely be lost, potentially allowing parallelism to confound phylogenetic analysis of morphological characters.     

Origin and evolution of the grazing guild in new world terrestrial mammals

Although vertebrate herbivory has existed on land for about 300 million years, the grazingadaptation, principally developed in mammals, did not appear until the middle Cenozoic about 30 million years ago. Paleontological evidence indicates that grazing mammals diversified at the time of the spread of grasslands. Recently revised fossil calibrations reveal that the grazing mammal guild originated during the early Miocene in South America about 10-15 million years earlier than it did during the late Miocene in the northern hemisphere. Carbon isotopic analyses of extinct grazers' teeth reveal that this guild originated predominantly in C(3) terrestrial ecosystems. The present-day distribution of C(3) and C(4) grasslands evolved on the global ecological landscape since the late Miocene, after about 7 million years ago.     

Perception mechanism of gravistimuli in gravity resistance responses of plants.

Gravity resistance is a response that enables plants to develop against the gravitational force. Hypergravity conditions produced by centrifugation have been used to analyze the mechanisms of gravity resistance responses. Under hypergravity conditions, plants construct short and thick shoots and increase cell wall rigidity for resisting the gravitational force. Hypergravity caused a decrease in the percentage of cells with transverse microtubules, and an increase in that with longitudinal microtubules. Such a prompt reorientation of cortical microtubules is involved in the changes in morphology of shoots by gravity. Hypergravity increased cell wall rigidity by increasing the molecular mass of xyloglucans via suppression of xyloglucan breakdown as well as by the thickening of cell walls. Blocker reagents of mechanoreceptors nullified the above-mentioned changes induced by hypergravity. Gravity resistance responses were brought about normally in mutants deprived of gravitropism. This result indicates that the graviperception mechanism in gravity resistance is independent of that in gravitropism. Gravity resistance responses were brought about independently of the direction of gravistimuli, but the responses disappeared in the presence of blockers of mechanoreceptors. Thus, in gravity responses, plants may perceive the gravitational force independently of the direction of stimuli by mechanoreceptors on the plasma membrane, and may utilize the signal to construct a tough body.     

Three-dimensional growth: a developmental innovation that facilitated plant terrestrialization

Journal of Plant Research - 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...     

Biotic interactions of marine algae

Marine algae encompass lineages that diverged about one billion years ago. Recent results suggest that they feature natural immunity traits that are conserved, as well as others that appear to be phylum- or environment-specific. In particular, marine plants resemble terrestrial plants and animals in their basic mechanisms for pathogen recognition and signaling, suggesting that these essential cell functions arose in the sea. Specific traits are based on the synthesis of unique secondary defense metabolites, often making use of the variety of halides found in the sea.     

A horizontal gene transfer at the origin of phenylpropanoid metabolism: a key adaptation of plants to land

Background  

The pioneering ancestor of land plants that conquered terrestrial habitats around 500 million years ago had to face dramatic stresses including UV radiation, desiccation, and microbial attack. This drove a number of adaptations, among which the emergence of the phenylpropanoid pathway was crucial, leading to essential compounds such as flavonoids and lignin. However, the origin of this specific land plant secondary metabolism has not been clarified.     

Molecular identification and phylogeny of arbuscular mycorrhizal fungi

The fossil record and molecular data show that the evolutionary history of arbuscular mycorrhizal fungi (Glomales) goes back at least to the Ordovician (460 million years ago), coinciding with the colonization of the terrestrial environment by the first land plants. At that time, the land flora only consisted of plants on the bryophytic level. Ribosomal DNA sequences indicate that the diversity within the Glomales on the family and genus level is much higher than previously expected from morphology-based taxonomy. Two deeply divergent lineages were found and described in two new genera, Archaeospora and Paraglomus, each in its own family. Based on a fast-growing number of available DNA sequences, several systems for molecular identification of the Glomales within roots have been designed and tested in the past few years. These detection methods have opened up entirely new perspectives for studying the ecology of arbuscular mycorrhiza.     

Plant evolution: TALES of development

TALE homeodomain proteins regulate development in many eukaryotes. Now, Lee et al. (2008) report that two TALE homeodomain proteins control zygote development of the unicellular green alga Chlamydomonas. This implicates TALE gene loss and diversification in the evolution of new diploid body plans that appeared when land plants evolved from algal ancestors over 450 million years ago.     

Woody plant and gravity]

In this review, we attempted to summarize the effect of gravity on growth of woody plants, broad leaved trees, on earth. It is well known that in tilted broad leaved trees, tension wood formed in the secondary xylem causes negative gravitropism. Gibberellin has been shown to induce tension wood in weeping branch, causing its upright growth. Recent study has shown that seedling of Japanese cherry tree grown on three dimensional clinostat, a device that simulates microgravity, grew at random angles, and that the formation of secondary xylem, as supporting tissue for upright growth, decreased. In the decreased xylem formation, the inhibition of the differentiation and development of fiber cell was clearly observed. These results suggest that in attitude control and morphogenesis of stem in woody plant, secondary xylem formation seriously relates to gravity on earth. In woody plant, the mechanism of gravity perception and the following signal transduction have not yet been elucidated, although the recent study reported the possibility that endodermal starch sheath cells and plant hormones may play some role in the mechanism. Space experiment for woody plant is expected to study these problem.     

Intron phylogeny: a new hypothesis

The three major classes of intron are clearly of unequal antiquity. Structured (often self-splicing and sometimes mobile) introns are the most ancient, probably dating (at least for group I) from the ancestral (eubacterial) cell 3500 million years ago, and were originally restricted to tRNA. Protein-spliced introns (usually in tRNA) probably evolved from them by a radical change in splicing mechanism in the common ancestor of eukaryotes and archaebacteria, perhaps only about 1700 million years ago. Spliceosomal introns probably evolved from group-II-like self-splicing introns after the origin of the nucleus between 1700 and 1000 million years ago, and were probably mostly inserted into previously unsplit protein-coding genes after the origin of mitochondria 1000 million years ago.     

The response to gravity is correlated with the number of statoliths in Chara rhizoids.

In contrast to higher plants, Chara rhizoids have single membrane-bound compartments that appear to function as statoliths. Rhizoids were generated by germinating zygotes of Chara in either soil water (SW) medium or artificial pond water (APW) medium. Differential-interference-contrast microscopy demonstrated that rhizoids form SW-grown plants typically contain 50 to 60 statoliths per cell, whereas rhizoids from APW-grown plants contain 5 to 10 statoliths per cell. Rhizoids from SW are more responsive to gravity than rhizoids from APW because (a) SW rhizoids were oriented to gravity during vertical growth, whereas APW rhizoids were relatively disoriented, and (b) curvature of SW rhizoids was 3 to 4 times greater throughout the time course of curvature. The growth rate of APW rhizoids was significantly greater than that of SW-grown rhizoids. This latter result suggests that APW rhizoids are not limited in their ability for gravitropic curvature by growth and that these rhizoids are impaired in the early stages of gravitropism (i.e. gravity perception). Plants grown in APW appeared to be healthy because of their growth rate and the vigorous cytoplasmic streaming observed in the rhizoids. This study is comparable to earlier studies of gravitropism in starch-deficient mutants of higher plants and provides support for the role of statoliths in gravity perception.     

Azospirillum genomes reveal transition of bacteria from aquatic to terrestrial environments

Fossil records indicate that life appeared in marine environments ～3.5 billion years ago (Gyr) and transitioned to terrestrial ecosystems nearly 2.5 Gyr. Sequence analysis suggests that "hydrobacteria" and "terrabacteria" might have diverged as early as 3 Gyr. Bacteria of the genus Azospirillum are associated with roots of terrestrial plants; however, virtually all their close relatives are aquatic. We obtained genome sequences of two Azospirillum species and analyzed their gene origins. While most Azospirillum house-keeping genes have orthologs in its close aquatic relatives, this lineage has obtained nearly half of its genome from terrestrial organisms. The majority of genes encoding functions critical for association with plants are among horizontally transferred genes. Our results show that transition of some aquatic bacteria to terrestrial habitats occurred much later than the suggested initial divergence of hydro- and terrabacterial clades. The birth of the genus Azospirillum approximately coincided with the emergence of vascular plants on land.     

Resistance of plants to gravitational force

Developing resistance to gravitational force is a critical response for terrestrial plants to survive under 1 × g conditions. We have termed this reaction “gravity resistance” and have analyzed its nature and mechanisms using hypergravity conditions produced by centrifugation and microgravity conditions in space. Our results indicate that plants develop a short and thick body and increase cell wall rigidity to resist gravitational force. The modification of body shape is brought about by the rapid reorientation of cortical microtubules that is caused by the action of microtubule-associated proteins in response to the magnitude of the gravitational force. The modification of cell wall rigidity is regulated by changes in cell wall metabolism that are caused by alterations in the levels of cell wall enzymes and in the pH of apoplastic fluid (cell wall fluid). Mechanoreceptors on the plasma membrane may be involved in the perception of the gravitational force. In this review, we discuss methods for altering gravitational conditions and describe the nature and mechanisms of gravity resistance in plants.     

Is water immersion useful for analyzing gravity resistance responses in terrestrial plants?

Water immersion has been used as a simulator of microgravity for analyzing gravity responses in semiaquatic plants such as rice. To examine whether or not water immersion for a short experimental period is a useful microgravity simulator even in terrestrial plants, we analyzed effects of water immersion on the cell wall rigidity and metabolisms of its constituents in azuki bean epicotyls. The cell wall rigidity of epicotyls grown underwater was significantly lower than that in the control. Water immersion also caused a decrease in molecular mass of xyloglucans as well as the thinning of the cell wall. Such changes in the mechanical and chemical properties of the cell wall underwater were similar to those observed in microgravity conditions in space. These results suggest that water immersion for a short period is a useful system for analyzing gravity resistance responses even in terrestrial plants.     

Oldest Pathology in a Tetrapod Bone Illuminates the Origin of Terrestrial Vertebrates

The origin of terrestrial tetrapods was a key event in vertebrate evolution, yet how and when it occurred remains obscure, due to scarce fossil evidence. Here, we show that the study of palaeopathologies, such as broken and healed bones, can help elucidate poorly understood behavioural transitions such as this. Using high-resolution finite element analysis, we demonstrate that the oldest known broken tetrapod bone, a radius of the primitive stem tetrapod Ossinodus pueri from the mid-Viséan (333 million years ago) of Australia, fractured under a high-force, impact-type loading scenario. The nature of the fracture suggests that it most plausibly occurred during a fall on land. Augmenting this are new osteological observations, including a preferred directionality to the trabecular architecture of cancellous bone. Together, these results suggest that Ossinodus, one of the first large (>2m length) tetrapods, spent a significant proportion of its life on land. Our findings have important implications for understanding the temporal, biogeographical and physiological contexts under which terrestriality in vertebrates evolved. They push the date for the origin of terrestrial tetrapods further back into the Carboniferous by at least two million years. Moreover, they raise the possibility that terrestriality in vertebrates first evolved in large tetrapods in Gondwana rather than in small European forms, warranting a re-evaluation of this important evolutionary event.     

Molecular evidence for the common origin of snap-traps among carnivorous plants

The snap-trap leaves of the aquatic waterwheel plant (Aldrovanda) resemble those of Venus' flytrap (Dionaea), its distribution and habit are reminiscent of bladderworts (Utricularia), but it shares many reproductive characters with sundews (Drosera). Moreover, Aldrovanda has never been included in molecular phylogenetic studies, so it has been unclear whether snap-traps evolved only once or more than once among angiosperms. Using sequences from nuclear 18S and plastid rbcL, atpB, and matK genes, we show that Aldrovanda is sister to Dionaea, and this pair is sister to Drosera. Our results indicate that snap-traps are derived from flypaper-traps and have a common ancestry among flowering plants, despite the fact that this mechanism is used by both a terrestrial species and an aquatic one. Genetic and fossil evidence for the close relationship between these unique and threatened organisms indicate that carnivory evolved from a common ancestor within this caryophyllid clade at least 65 million years ago.     

Regulatory mechanism controlling stomatal behavior conserved across 400 million years of land plant evolution

Stomatal pores evolved more than 410 million years ago [1,?2]?and allowed vascular plants to regulate transpirational water loss during the uptake of CO(2) for photosynthesis [3]. Here, we show that stomata on the sporophytes of the moss Physcomitrella patens [2] respond to environmental signals in a similar way to those of flowering plants [4] and that a homolog of a key signaling component in the vascular plant drought hormone abscisic acid (ABA) response [5] is?involved in stomatal control in mosses. Cross-species complementation experiments reveal that the stomatal ABA response of a flowering plant (Arabidopsis thaliana) mutant, lacking the ABA-regulatory protein kinase OPEN STOMATA 1 (OST1) [6], is rescued by substitution with the moss P.?patens homolog, PpOST1-1, which evolved more than 400 million years earlier. We further demonstrate through the targeted knockout of the PpOST1-1 gene in P.?patens that its role in guard cell closure is conserved, with stomata of mutant mosses exhibiting a significantly attenuated ABA response. Our analyses indicate that core regulatory components involved in guard cell ABA signaling of flowering plants are operational in mosses and likely originated in the last common ancestor of these lineages more than 400 million years ago [7], prior to the evolution of ferns [8, 9].