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50 I. 50 II. 52 III. 53 IV. 66 V. 71 VI. 72 VII. 74 75 References 75 SUMMARY: Cryptospores, recovered from Ordovician through Devonian rocks, differ from trilete spores in possessing distinctive configurations (i.e. hilate monads, dyads, and permanent tetrads). Their affinities are contentious, but knowledge of their relationships is essential to understanding the nature of the earliest land flora. This review brings together evidence about the source plants, mostly obtained from spores extracted from minute, fragmented, yet exceptionally anatomically preserved fossils. We coin the term ‘cryptophytes’ for plants that produced the cryptospores and show them to have been simple terrestrial organisms of short stature (i.e. millimetres high). Two lineages are currently recognized. Partitatheca shows a combination of characters (e.g. spo‐rophyte bifurcation, stomata, and dyads) unknown in plants today. Lenticulatheca encompasses discoidal sporangia containing monads formed from dyads with ultrastructure closer to that of higher plants, as exemplified by Cooksonia. Other emerging groupings are less well characterized, and their precise affinities to living clades remain unclear. Some may be stem group embryophytes or tracheophytes. Others are more closely related to the bryophytes, but they are not bryophytes as defined by extant representatives. Cryptophytes encompass a pool of diversity from which modern bryophytes and vascular plants emerged, but were competitively replaced by early tracheophytes. Sporogenesis always produced either dyads or tetrads, indicating strict genetic control. The long‐held consensus that tetrads were the archetypal condition in land plants is challenged.

The primary objective of this study was to identify the molecular signals present in arbuscular mycorrhizal (AM) germinated spore exudates (GSEs) responsible for activating nuclear Ca2+ spiking in the Medicago truncatula root epidermis. Medicago truncatula root organ cultures (ROCs) expressing a nuclear-localized cameleon reporter were used as a bioassay to detect AM-associated Ca2+ spiking responses and LC-MS to characterize targeted molecules in GSEs. This approach has revealed that short-chain chitin oligomers (COs) can mimic AM GSE-elicited Ca2+ spiking, with maximum activity observed for CO4 and CO5. This spiking response is dependent on genes of the common SYM signalling pathway (DMI1/DMI2) but not on NFP, the putative Sinorhizobium meliloti Nod factor receptor. A major increase in the CO4/5 concentration in fungal exudates is observed when Rhizophagus irregularis spores are germinated in the presence of the synthetic strigolactone analogue GR24. By comparison with COs, both sulphated and nonsulphated Myc lipochito-oligosaccharides (LCOs) are less efficient elicitors of Ca2+ spiking in M. truncatula ROCs. We propose that short-chain COs secreted by AM fungi are part of a molecular exchange with the host plant and that their perception in the epidermis leads to the activation of a SYM-dependent signalling pathway involved in the initial stages of fungal root colonization.

The publication of a large number of taxon names at all levels within the arbuscular mycorrhizal fungi (Glomeromycota) has resulted in conflicting systematic schemes and generated considerable confusion among biologists working with these important plant symbionts. A group of biologists with more than a century of collective experience in the systematics of Glomeromycota examined all available molecular–phylogenetic evidence within the framework of phylogenetic hypotheses, incorporating morphological characters when they were congruent. This study is the outcome, wherein the classification of Glomeromycota is revised by rejecting some new names on the grounds that they are founded in error and by synonymizing others that, while validly published, are not evidence-based. The proposed “consensus” will provide a framework for additional original research aimed at clarifying the evolutionary history of this important group of symbiotic fungi.

In disturbed or pioneer settings, spores and sclerotia of ectomycorrhizal fungi serve as the necessary inoculum for establishment of ectomycorrhizal-dependent trees. Yet, little is known about the persistence of these propagules through time. Here, live field soil was inoculated with known quantities of basidiospores from four pine-associated species of Rhizopogon; these samples were then buried in retrievable containers, and pine seedling bioassays of serially diluted spore samples were used to measure spore viability. In the first 4 yr, no evidence of loss of spore viability was found in the four Rhizopogon species tested, but all four species exhibited dormancy in which a maximum of 1-8% of their spores were initially receptive to pine roots. There were some differences between species in overall inoculum potential of their spores, but all species broke dormancy at a statistically similar rate. This result provides evidence for spore dormancy in a common ectomycorrhizal genus, but it also precludes our ability to estimate the longevity of the spores accurately. Nevertheless these results, coupled with the observed patterns of Rhizopogon spore banks, suggest that at least decade-long durations are likely. As this experiment progresses, the true longevity of the spores will eventually be revealed.

Living systems have inspired approaches to engineer cells as independent functional materials or integrate them within a natural or synthetic matrix to create engineered living materials (ELMs).To address the principal challenges arising from the ‘livingness’ of cells, bacterial spores have emerged as a game changer in the field, enabling users to ‘activate’ cells on demand and to treat ELMs with harsh conditions.The inherent physical properties of the structural components of the spores have led to diverse applications of spore-based materials.Emerging synthetic biology tools and better understanding of bacterial spores might contribute to expanding the relevance of spores in various fields, such as biosensing, biocatalysis, and data storage, among others. Inspired by biological functions of living systems, researchers have engineered cells as independent functional materials or integrated them within a natural or synthetic matrix to create engineered living materials (ELMs). However, the ‘livingness’ of cells in such materials poses serious drawbacks, such as a short lifespan and the need for cold-chain logistics. Bacterial spores have emerged as a game changer to bypass these shortcomings as a result of their intrinsic dormancy and resistance against harsh conditions. Emerging synthetic biology tools tailored for engineering spores and better understanding of their physical properties have led to novel applications of spore-based materials. Here, we review recent advances in such materials and discuss future challenges for the development of time- and cost-efficient spore-based materials with high performance. Inspired by biological functions of living systems, researchers have engineered cells as independent functional materials or integrated them within a natural or synthetic matrix to create engineered living materials (ELMs). However, the ‘livingness’ of cells in such materials poses serious drawbacks, such as a short lifespan and the need for cold-chain logistics. Bacterial spores have emerged as a game changer to bypass these shortcomings as a result of their intrinsic dormancy and resistance against harsh conditions. Emerging synthetic biology tools tailored for engineering spores and better understanding of their physical properties have led to novel applications of spore-based materials. Here, we review recent advances in such materials and discuss future challenges for the development of time- and cost-efficient spore-based materials with high performance.

Key Points The Bacillus subtilis spore coat is a multilayered protective structure composed of more than 70 different proteins. In addition to its protective role, the spore coat influences the process of spore germination and defines the type of interactions that spores can establish with various surfaces in the environment. Fluorescence microscopy in combination with high-resolution image analysis has produced a spatially scaled coat protein interaction network indicating that the coat is organized into four distinct layers. These studies led to the discovery of the outermost layer of the coat in B. subtilis , referred to as the spore crust. Time course analyses of spore coat assembly have revealed that two main steps can be distinguished in coat morphogenesis: the initial recruitment of proteins to the spore surface as a scaffold cap, followed by spore encasement in a series of successive waves. Coat assembly is regulated at the transcriptional level by the sequential expression of individual coat genes and at the protein level by a small group of coat morphogenetic proteins that coordinate both the recruitment of coat proteins to specific coat layers and spore encasement. Sporulation in Bacillus subtilis results in the formation of an endospore surrounded by a multilayered protective structure, known as the coat. In this Review, Patrick Eichenberger and colleagues describe recent studies that have illuminated the architecture of the coat and the dynamics of coat assembly. Sporulation in Bacillus subtilis involves an asymmetric cell division followed by differentiation into two cell types, the endospore and the mother cell. The endospore coat is a multilayered shell that protects the bacterial genome during stress conditions and is composed of dozens of proteins. Recently, fluorescence microscopy coupled with high-resolution image analysis has been applied to the dynamic process of coat assembly and has shown that the coat is organized into at least four distinct layers. In this Review, we provide a brief summary of B. subtilis sporulation, describe the function of the spore surface layers and discuss the recent progress that has improved our understanding of the structure of the endospore coat and the mechanisms of coat assembly.

Ash dieback is caused by an invasive pathogen Hymenoscyphus fraxineus, which emerged in Europe in the 1990s and jeopardizes the management of ash stands. Although the biological cycle of the pathogen is well understood, its dispersal patterns via airborne spores remain poorly described. We investigated the seasonal and spatial patterns of dispersal in France using both a passive spore-trapping method coupled with a real-time PCR assay and reports of ash dieback based on symptom observations. Spores detection varies from year to year with a detection ability of 30-47% depending on meteorological conditions, which affect both production of inoculum and efficiency of the trapping. Nevertheless, our results are consistent and we showed that the sporulation peak occurred from June to August and that spores were detected up to 50-100 km ahead of the disease front, proving the presence of the pathogen before any observation of symptoms. The spore dispersal gradient was steep, most of inoculum remaining within 50 m of infected ashes. Two dispersal kernels were fitted using Bayesian methods to estimate the mean dispersal distance of H. fraxineus from inoculum sources. The estimated mean distances of dispersal, either local or regional scale, were 1.4 km and 2.6 km, respectively, the best fitting kernel being the inverse power-law. This information may help to design disease management strategies.

With the increased knowledge on spore structure and advances in biotechnology engineering, the newly developed spore-surface display system confers several inherent advantages over other microbial cell-surface display systems including enhanced stability and high safety. Bacillus subtilis is the most commonly used Bacillus species for spore-surface display. The expression of heterologous antigen or protein on the surface of B. subtilis spores has now been practiced for over a decade with noteworthy success. As an update and supplement to other previous reviews, we comprehensively summarize recent studies in the B. subtilis spore-surface display technique. We focus on its benefits as well as the critical factors affecting its display efficiency and offer suggestions for the future success of this field.

The geochemical carbon cycle is strongly influenced by life on land, principally through the effects of carbon sequestration and the weathering of calcium and magnesium silicates in surface rocks and soils. Knowing the time of origin of land plants and animals and also of key organ systems (e.g. plant vasculature, roots, wood) is crucial to understand the development of the carbon cycle and its effects on other Earth systems. Here, we compare evidence from fossils with calibrated molecular phylogenetic trees (timetrees) of living plants and arthropods. We show that different perspectives conflict in terms of the relative timing of events, the organisms involved and the pattern of diversification of various groups. Focusing on the fossil record, we highlight a number of key biases that underpin some of these conflicts, the most pervasive and far-reaching being the extent and nature of major facies changes in the rock record. These effects probably mask an earlier origin of life on land than is evident from certain classes of fossil data. If correct, this would have major implications in understanding the carbon cycle during the Early Palaeozoic.

The bacterial spore, the hardiest known life form, can survive in a metabolically dormant state for many years and can withstand high temperatures, radiation, and toxic chemicals. The molecular basis of spore dormancy and resistance is not understood, but the physical state of water in the different spore compartments is thought to play a key role. To characterize this water in situ, we recorded the water ²H and ¹⁷O spin relaxation rates in D₂O-exchanged Bacillus subtilis spores over a wide frequency range. The data indicate high water mobility throughout the spore, comparable with binary protein-water systems at similar hydration levels. Even in the dense core, the average water rotational correlation time is only 50 ps. Spore dormancy therefore cannot be explained by glass-like quenching of molecular diffusion but may be linked to dehydration-induced conformational changes in key enzymes. The data demonstrate that most spore proteins are rotationally immobilized, which may contribute to heat resistance by preventing heat-denatured proteins from aggregating irreversibly. We also find that the water permeability of the inner membrane is at least 2 orders of magnitude lower than for model membranes, consistent with the reported high degree of lipid immobilization in this membrane and with its proposed role in spore resistance to chemicals that damage DNA. The quantitative results reported here on water mobility and transport provide important clues about the mechanism of spore dormancy and resistance, with relevance to food preservation, disease prevention, and astrobiology.