The Orchid Velamen: A model system for studying patterned secondary cell wall development?

Understanding the mechanisms through which plants generate secondary cell walls is of more than academic interest: the physical properties of plant-derived materials, including timber and textiles, all depend upon secondary wall cellulose organization. Processes controlling cellulose in the secondary cell wall and their reliance on microtubules have been documented in recent decades, but this understanding is complicated, as secondary walls normally form in the plant’s interior where live cell imaging is more difficult. We investigated secondary wall formation in the orchid velamen, a multicellular epidermal layer found around orchid roots that consists of dead cells with lignified secondary cell walls. The patterns of cell wall ridges that form within the velamen vary between different orchid species, but immunolabelling demonstrated that wall deposition is controlled by microtubules. As these patterning events occur at the outer surface of the root, and as orchids are adaptable for tissue culture and genetic manipulation, we conclude that the orchid root velamen may indeed be a suitable model system for studying the organization of the plant cell wall. Notably, roots of the commonly grown orchid Laelia anceps appear ideally suited for developing this research

Asymmetric wall ingrowth deposition in Arabidopsis phloem parenchyma transfer cells is tightly associated with sieve elements

In Arabidopsis, polarized deposition of wall ingrowths in phloem parenchyma (PP) transfer cells (TCs) occurs adjacent to cells of the sieve element/companion cell (SE/CC) complex. However, the spatial relationships between these different cell types in minor veins, where phloem loading occurs, are poorly understood. PP TC development and wall ingrowth localization were compared with those of other phloem cells in leaves of Col-0 and the transgenic lines AtSUC2::AtSTP9-GFP (green fluorescent protein) and AtSWEET11::AtSWEET11-GFP that identify CCs and PP cells, respectively. The development of PP TCs in minor veins, indicated by deposition of wall ingrowths, proceeded basipetally in leaves. However, not all PP cells develop wall ingrowths, and higher levels of deposition occur in abaxial- compared with adaxial-positioned PP TCs. Furthermore, the deposition of wall ingrowths was exclusively initiated on and preferentially covered the PP TC/SE interface, rather than the PP TC/CC interface, and only occurred in PP cells that were adjacent to SEs. Collectively, these results demonstrate a tight association between SEs and wall ingrowth deposition in PP TCs and suggest the existence of two subtypes of PP cells in leaf minor veins. Compared with PP cells, PP TCs showed more abundant accumulation of AtSWEET11–GFP, indicating functional differences in phloem loading between PP and PP TCs

Whole-genome sequencing reveals host factors underlying critical COVID-19

Critical COVID-19 is caused by immune-mediated inflammatory lung injury. Host genetic variation influences the development of illness requiring critical care1 or hospitalization2,3,4 after infection with SARS-CoV-2. The GenOMICC (Genetics of Mortality in Critical Care) study enables the comparison of genomes from individuals who are critically ill with those of population controls to find underlying disease mechanisms. Here we use whole-genome sequencing in 7,491 critically ill individuals compared with 48,400 controls to discover and replicate 23 independent variants that significantly predispose to critical COVID-19. We identify 16 new independent associations, including variants within genes that are involved in interferon signalling (IL10RB and PLSCR1), leucocyte differentiation (BCL11A) and blood-type antigen secretor status (FUT2). Using transcriptome-wide association and colocalization to infer the effect of gene expression on disease severity, we find evidence that implicates multiple genes—including reduced expression of a membrane flippase (ATP11A), and increased expression of a mucin (MUC1)—in critical disease. Mendelian randomization provides evidence in support of causal roles for myeloid cell adhesion molecules (SELE, ICAM5 and CD209) and the coagulation factor F8, all of which are potentially druggable targets. Our results are broadly consistent with a multi-component model of COVID-19 pathophysiology, in which at least two distinct mechanisms can predispose to life-threatening disease: failure to control viral replication; or an enhanced tendency towards pulmonary inflammation and intravascular coagulation. We show that comparison between cases of critical illness and population controls is highly efficient for the detection of therapeutically relevant mechanisms of disease

Review: More than sweet: New insights into the biology of phloem parenchyma transfer cells in Arabidopsis

Transfer cells (TCs) develop extensive wall ingrowths to facilitate enhanced rates of membrane transport. In Arabidopsis, TCs trans-differentiate from phloem parenchyma (PP) cells abutting the sieve element/companion cell complex in minor veins of foliar tissues and, based on anatomy and expression of SWEET sucrose uniporters, are assumed to play pivotal roles in phloem loading. While wall ingrowth deposition in PP TCs is a dynamic process responding to abiotic stresses such as high light and cold, the transcriptional control of PP TC development, including deposition of the wall ingrowths themselves, is not understood. PP TC development is a trait of vegetative phase change, potentially linking wall ingrowth deposition with floral induction. Transcript profiling by RNA-seq identified NAC056 and NAC018 (NARS1 and NARS2) as putative regulators of wall ingrowth deposition, while recent single cell RNA-seq analysis of leaf vasculature identified PP-specific expression of NAC056. Numerous membrane transporters, particularly of the UmamiT family of amino acid efflux carriers, were also identified. Collectively, these findings, and the recent discovery that wall ingrowth deposition is regulated by sucrose-dependent loading activity of these cells, provide new insights into the biology of PP TCs and their importance to phloem loading in Arabidopsis, establishing these cells as a key transport hub for phloem loading

Phi thickenings in Brassica oleracea roots are induced by osmotic stress and mechanical effects, both involving jasmonic acid

Phi thickenings are peculiar secondary cell wall thickenings found in radial walls of cortical cells in plant roots. However, while thickenings are widespread in the plant kingdom, research into their development has been lacking. Here, we describe a simple system for rapid induction of phi thickenings in primary roots of Brassica. Four-day-old seedlings were transferred from control agar plates to new plates containing increased levels of osmotica. Phi thickening development occurred within a narrow region of the differentiation zone proportional to osmolarity, with cellulose deposition and lignification starting after 12h and 15h, respectively. However, osmoprotectants not only failed to induce phi thickenings, but inhibited induction when tested in combination with thickening-inducing osmotica. An independent, biomechanical pathway exists regulating phi thickening induction, with root growth rates and substrate texture being important factors in determining thickening induction. Phi thickening development is also controlled by stress-related plant hormones, most notably jasmonic acid, but also abscisic acid. Our research not only provides the first understanding of the developmental pathways controlling phi thickening induction, but also provides tools with which the functions of these enigmatic structures might be clarified