•The periderm of Arabidopsis is analogous to periderm in trees.•During periderm formation the endodermis undergoes programmed cell death.•Transcript profiling revealed common regulatory hubs between ...the vascular cambium and the phellogen (cork cambium).•Conserved transcription factors related to suberization act during phellem differentiation.
The periderm acts as the first line of defence for a plant, protecting wood and phloem from abiotic and biotic stresses. During secondary growth, through the increase in girth of plant organs, the periderm replaces the epidermis as the outermost tissue. The phellogen, a bifacial post-embryonic meristem, forms the phelloderm inwards (toward the vasculature) and the suberized phellem outwards (toward the environment). These three tissues are collectively referred to as the periderm. Here, we summarize recent findings on the molecular mechanisms of periderm development by describing periderm formation in connection to the fate of the surrounding tissues, by discussing common regulatory hubs between the vascular cambium and the phellogen, and by highlighting transcription factors (TFs) controlling phellem differentiation.
Plants deposit hydrophobic polymers, such as lignin or suberin, in their root cell walls to protect inner tissues and facilitate selective uptake of solutes. Insights into how individual root tissues ...contribute to polymer formation are important for elucidation of ultrastructure, function, and development of these protective barriers. Although the pathways responsible for production of the barrier constituents are established, our models lack spatiotemporal resolution—especially in roots—thus, the source of monomeric barrier components is not clear. This is mainly due to our restricted ability to manipulate synthesis of the broadly important phenylpropanoid pathway, as mutants in this pathway display lethal or pleiotropic phenotypes. Here, we overcome this challenge by exploiting highly controlled in vivo repression systems. We provide strong evidence that autonomous production of phenylpropanoids is essential for establishment of the endodermal Casparian strip as well as adherence of the suberin matrix to the cell wall of endodermis and cork. Our work highlights that, in roots, the phenylpropanoid pathway is under tight spatiotemporal control and serves distinct roles in barrier formation across tissues and developmental zones. This becomes evident in the late endodermis, where repression of phenylpropanoid production leads to active removal of suberin in pre-suberized cells, indicating that endodermal suberin depositions might embody a steady state between continuous synthesis and degradation.
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•Autonomous production of phenylpropanoids is required for Casparian strip formation•Phenylpropanoids are essential for suberin deposition in endodermis and cork•Cork differentiation requires autonomous production of phenylpropanoids•Spatiotemporal repression of phenylpropanoids can dissect barrier functions
Andersen et al. show here that cell-autonomous phenylpropanoid production is required for root barrier formation and integrity.
Peptide signaling presumably occupies a central role in plant development, yet only few concrete examples of receptor-ligand pairs that act in the context of specific differentiation processes have ...been described. Here we report that second-site null mutations in the Arabidopsis leucine-rich repeat receptor-like kinase gene barely any meristem 3 (BAM3) perfectly suppress the postembryonic root meristem growth defect and the associated perturbed protophloem development of the brews radix (brx) mutant. The roots of bam3 mutants specifically resist growth inhibition by the CLAVATA3/ENDOSPERM SURROUNDING REGION 45 (CLE45) peptide ligand. WT plants transformed with a construct for ectopic overexpression of CLE45 could not be recovered, with the exception of a single severely dwarfed and sterile plant that eventually died. By contrast, we obtained numerous transgenic bam3 mutants transformed with the same construct. These transgenic plants displayed a WT phenotype, however, supporting the notion that CLE45 is the likely BAM3 ligand. The results correlate with the observation that external CLE45 application represses protophloem differentiation in WT, but not in bam3 mutants. BAM3, BRX, and CLE45 are expressed in a similar spatiotemporal trend along the developing protophloem, up to the end of the transition zone. Induction of BAM3 expression upon CLE45 application, ectopic overexpression of BAM3 in brx root meristems, and laser ablation experiments suggest that intertwined regulatory activity of BRX, BAM3, and CLE45 could be involved in the proper transition of protophloem cells from proliferation to differentiation, thereby impinging on postembryonic growth capacity of the root meristem.
During secondary growth in most eudicots and gymnosperms, the periderm replaces the epidermis as the frontier tissue protecting the vasculature from biotic and abiotic stresses. Despite its ...importance, the mechanisms underlying periderm establishment and formation are largely unknown.
The herbaceous Arabidopsis thaliana undergoes secondary growth, including periderm formation in the root and hypocotyl. Thus, we focused on these two organs to establish a framework to study periderm development in a model organism.
We identified a set of characteristic developmental stages describing periderm growth from the first cell division in the pericycle to the shedding of the cortex and epidermis. We highlight that two independent mechanisms are involved in the loosening of the outer tissues as the endodermis undergoes programmed cell death, whereas the epidermis and the cortex are abscised. Moreover, the phellem of Arabidopsis, as in trees, is suberized, lignified and peels off. In addition, putative regulators from oak and potato are also expressed in the Arabidopsis periderm.
Collectively, the periderm of Arabidopsis shares many characteristics/features of woody and tuberous periderms, rendering Arabidopsis thaliana an attractive model for cork biology.
Tree bark is a highly specialized array of tissues that plays important roles in plant protection and development. Bark tissues develop from two lateral meristems; the phellogen (cork cambium) ...produces the outermost stem–environment barrier called the periderm, while the vascular cambium contributes with phloem tissues. Although bark is diverse in terms of tissues, functions and species, it remains understudied at higher resolution.
We dissected the stem of silver birch (Betula pendula) into eight major tissue types, and characterized these by a combined transcriptomics and metabolomics approach. We further analyzed the varying bark types within the Betulaceae family.
The two meristems had a distinct contribution to the stem transcriptomic landscape. Furthermore, inter- and intraspecies analyses illustrated the unique molecular profile of the phellem. We identified multiple tissue-specific metabolic pathways, such as the mevalonate/betulin biosynthesis pathway, that displayed differential evolution within the Betulaceae. A detailed analysis of suberin and betulin biosynthesis pathways identified a set of underlying regulators and highlighted the important role of local, small-scale gene duplication events in the evolution of metabolic pathways.
This work reveals the transcriptome and metabolic diversity among bark tissues and provides insights to its development and evolution, as well as its biotechnological applications.
A central question in developmental biology is how multicellular organisms coordinate cell division and differentiation to determine organ size. In Arabidopsis roots, this balance is controlled by ...cytokinin-induced expression of SHORT HYPOCOTYL 2 (SHY2) in the so-called transition zone of the meristem, where SHY2 negatively regulates auxin response factors (ARFs) by protein—protein interaction. The resulting down-regulation of PIN-FORMED (PIN) auxin efflux carriers is considered the key event in promoting differentiation of meristematic cells. Here we show that this regulation involves additional, intermediary factors and is spatio-temporally constrained. We found that the described cytokinin—auxin crosstalk antagonizes BREVIS RADIX (BRX) activity in the developing protophloem. BRX is an auxin-responsive target of the prototypical ARF MONOPTEROS (MP), a key promoter of vascular development, and transiently enhances PIN3 expression to promote meristem growth in young roots. At later stages, cytokinin induction of SHY2 in the vascular transition zone restricts BRX expression to down-regulate PIN3 and thus limit meristem growth. Interestingly, proper SHY2 expression requires BRX, which could reflect feedback on the auxin responsiveness of SHY2 because BRX protein can directly interact with MP, likely acting as a cofactor. Thus, cross-regulatory antagonism between BRX and SHY2 could determine ARF activity in the protophloem. Our data suggest a model in which the regulatory interactions favor BRX expression in the early proximal meristem and SHY2 prevails because of supplementary cytokinin induction in the later distal meristem. The complex equilibrium of this regulatory module might represent a universal switch in the transition toward differentiation in various developmental contexts.
Contrast-induced acute kidney injury (CI-AKI) is a serious complication during percutaneous coronary interventions (PCI). Currently, the diagnosis of CI-AKI relies on serum creatinine (SCr) that is ...however affected by several limitations potentially leading to delayed or missed diagnoses. In this study we examined the diagnostic accuracy of a "bedside" measurement of plasma Neutrophil Gelatinase-Associated Lipocalin (NGAL) in the early detection of CI-AKI in 97 patients undergoing elective PCI. The overall incidence of CI-AKI was 3%. A significant positive correlation was observed between 6-hours NGAL and post-PCI SCr (r = 0.339, p = 0.004) and a significant negative correlation between 6-hours NGAL and post-PCI CrCl (r = -0.303, p = 0.010). In patients with post-PCI SCr increase > 0.24 mg/dl (median SCr absolute increase), delta NGAL 0-6 hours and 6-hours NGAL values were higher compared with patients with SCr elevation below the defined threshold (p = 0.049 and p = 0.056). The ROC analysis showed that a 6 hours NGAL value > 96 ng/ml significantly predicted an absolute SCr increase > 0.24 mg/dl after contrast exposure with sensitivity of 53% and specificity of 74% (AUC 0.819, 95% CI: 0.656 to 0.983, p = 0.005). The use of bedside NGAL assessment may significantly hasten diagnosis and treatment of CI-AKI, with remarkable clinical prognostic consequences.
Plants are the primary producers of biomass on earth. As an almost stereotypic feature, higher plants generate continuously growing bodies mediated by the activity of different groups of stem cells, ...the meristems. Shoot and root thickening is one of the fundamental growth processes determining form and function of these bodies. Mediated by a group of cylindrical meristems located below organ surfaces, vascular and protective tissues are continuously generated in a highly plastic manner, a competence essential for the survival in an ever changing environment. Acknowledging the fundamental role of this process, which is overall designated as secondary growth, we discuss in this review our current knowledge about the evolution and molecular regulation of the vascular cambium. The cambium is the meristem responsible for the formation of wood and bast, the two types of vascular tissues important for long-distance transport of water and assimilates, respectively. Although regulatory patterns are only beginning to emerge, we show that cambium activity represents a highly rewarding model for studying cell fate decisions, tissue patterning and differentiation, which has experienced an outstanding phylogenetic diversification.
The seismic response of base-isolated structures is notably influenced by mechanical properties of isolation devices due to their essential role in structural behavior. Consequently, the variability ...of such properties should be accounted for in the design process. The current seismic codes prescribe a simplified approach based on structural analyses in two extreme situations resulting from the upper and lower bound design properties of bearings (upper and lower bound analyses). In the case that experimental data are not provided by manufacturers, seismic codes provide the so-called “property modification factors” or “λ-factors”, i.e., modification coefficients to be applied to the nominal dynamic properties of bearings to obtain their upper or lower design properties. The aim of this paper is to provide a historical review of values provided for such factors by the main seismic codes by highlighting the limits, as well as some clerical errors, present in some codes. In particular, the European seismic codes are illustrated in detail, i.e., the Eurocode for bridges (EN 1998-2) and product standard on anti-seismic devices (EN 15129). Both these codes account for different sources of variability, such as the bearings production and the environmental and behavioral effects. For all these effects, the same λ-factor values are provided by the two codes, deriving from the second version of the AASHTO guide specifications for seismic isolation of bridges (AASHTO 1999), which are based on limited and/or old data, especially for high damping rubber bearings (HDRBs), and were never updated in the successive versions. More recent standards are also illustrated, providing different perspectives that deserve attention, even though they require further investigations to be applied in the design practice.
This paper investigates the seismic design of fluid viscous dampers connecting adjacent structural systems. A simplified dampers design strategy is proposed, which relies on a linearized reduced ...order model of the coupled system. A stochastic linearization technique is adopted with the aim of extending the design method to non-linear viscous dampers. The effectiveness of the design method and of the coupling strategy are assessed via numerical analysis of two adjacent buildings with shear-type behavior connected by linear or non-linear viscous dampers and subjected to Gaussian stochastic base acceleration. Different dampers locations are analyzed. The accuracy of the reduced order model is assessed, by comparing the relevant response statistics to those provided by a refined multi degree of freedoms model. Finally, a parametric study is performed to assess the effectiveness of dissipative connection for different values of seismic intensity and dampers parameters (i.e., viscous coefficients and velocity exponents).