Formation of silica aggregates in sorghum root endodermis is predetermined by cell wall architecture and development

Background and Aims Deposition of silica in plant cell walls improves their mechanical properties and helps plants to withstand various stress conditions. Its mechanism is still not understood and silica-cell wall interactions are elusive. The objective of this study was to investigate the effect of silica deposition on the development and structure of sorghum root endodermis and to identify the cell wall components involved in silicification. MethodsSorghum bicolor seedlings were grown hydroponically with (Si+) or without (Si-) silicon supplementation. Primary roots were used to investigate the transcription of silicon transporters by quantitative RT-PCR. Silica aggregation was induced also under in vitro conditions in detached root segments. The development and architecture of endodermal cell walls were analysed by histochemistry, microscopy and Raman spectroscopy. Water retention capability was compared between silicified and non-silicified roots. Raman spectroscopy analyses of isolated silica aggregates were also carried out. Key Results Active uptake of silicic acid is provided at the root apex, where silicon transporters Lsi1 and Lsi2 are expressed. The locations of silica aggregation are established during the development of tertiary endodermal cell walls, even in the absence of silicon. Silica aggregation takes place in non-lignified spots in the endodermal cell walls, which progressively accumulate silicic acid, and its condensation initiates at arabinoxylan-ferulic acid complexes. Silicification does not support root water retention capability; however, it decreases root growth inhibition imposed by desiccation. Conclusion A model is proposed in which the formation of silica aggregates in sorghum roots is predetermined by a modified cell wall architecture and takes place as governed by endodermal development. The interaction with silica is provided by arabinoxylan-ferulic acid complexes and interferes with further deposition of lignin. Due to contrasting hydrophobicity, silicification and lignification do not represent functionally equivalent modifications of plant cell walls

Silicon-mediated herbivore defence in a pasture grass under reduced and Anthropocene levels of CO2

The uptake and accumulation of silicon (Si) in grass plants play a crucial role in alleviating both biotic and abiotic stresses. Si supplementation has been reported to increase activity of defence-related antioxidant enzyme, which helps to reduce oxidative stress caused by reactive oxygen species (ROS) following herbivore attack. Atmospheric CO2 levels are known to affect Si accumulation in grasses; reduced CO2 concentrations increase Si accumulation whereas elevated CO2 concentrations often decrease Si accumulation. This can potentially affect antioxidant enzyme activity and subsequently insect herbivory, but this remains untested. We examined the effects of Si supplementation and herbivory by Helicoverpa armigera on antioxidant enzyme (catalase, CAT; superoxide dismutase, SOD; and ascorbate peroxidase, APX) activity in tall fescue grass (Festuca arundinacea) grown under CO2 concentrations of 200, 410, and 640 ppm representing reduced, ambient, and elevated CO2 levels, respectively. We also quantified foliar Si, carbon (C), and nitrogen (N) concentrations and determined how changes in enzymes and elemental chemistry affected H. armigera relative growth rates and plant consumption. Rising CO2 concentrations increased plant mass and foliar C but decreased foliar N and Si. Si supplementation enhanced APX and SOD activity under the ranging CO2 regimes. Si accumulation and antioxidant enzyme activity were at their highest level under reduced CO2 conditions and their lowest level under future levels of CO2. The latter corresponded with increased herbivore growth rates and plant consumption, suggesting that some grasses could become more susceptible to herbivory under projected CO2 conditions

COBRA-LIKE2, a Member of the Glycosylphosphatidylinositol-Anchored COBRA-LIKE Family, Plays a Role in Cellulose Deposition in Arabidopsis Seed Coat Mucilage Secretory Cells ,

Differentiation of the maternally derived seed coat epidermal cells into mucilage secretory cells is a common adaptation in angiosperms. Recent studies identified cellulose as an important component of seed mucilage in various species. Cellulose is deposited as a set of rays that radiate from the seed upon mucilage extrusion, serving to anchor the pectic component of seed mucilage to the seed surface. Using transcriptome data encompassing the course of seed development, we identified COBRA-LIKE2 (COBL2), a member of the glycosylphosphatidylinositol-anchored COBRA-LIKE gene family in Arabidopsis (Arabidopsis thaliana), as coexpressed with other genes involved in cellulose deposition in mucilage secretory cells. Disruption of the COBL2 gene results in substantial reduction in the rays of cellulose present in seed mucilage, along with an increased solubility of the pectic component of the mucilage. Light birefringence demonstrates a substantial decrease in crystalline cellulose deposition into the cellulosic rays of the cobl2 mutants. Moreover, crystalline cellulose deposition into the radial cell walls and the columella appears substantially compromised, as demonstrated by scanning electron microscopy and in situ quantification of light birefringence. Overall, the cobl2 mutants display about 40% reduction in whole-seed crystalline cellulose content compared with the wild type. These data establish that COBL2 plays a role in the deposition of crystalline cellulose into various secondary cell wall structures during seed coat epidermal cell differentiation

The Hygroscopic Opening of Sesame Fruits Is Induced by a Functionally Graded Pericarp Architecture

To enhance the distribution of their seeds, plants often utilize hygroscopic deformations that actuate dispersal mechanisms. Such movements are based on desiccation-induced shrinkage of tissues in predefined directions. The basic hygroscopic deformations are typically actuated by a bi-layer configuration, in which shrinking of an active tissue layer is resisted by a stiff layer, generating a set of basic movements including bending, coiling, and twisting. In this study, we investigate a new type of functionally graded hygroscopic movement in the fruit (capsule) of sesame (Sesamum indicum L.). Microscopic observations of the capsules showed that the inner stiff endocarp layer is built of a bilayer of transverse (i.e. circumferential) and longitudinal fiber cells with the layers positioned in a semi-circle, one inside the other. The outer mesocarp layer is made of soft parenchyma cells. The thickness of the fibrous layers and of the mesocarp exhibits a graded architecture, with gradual changes in their thickness around the capsule circumference. The cellulose microfibrils in the fiber cell walls are lying parallel to the cell long axis, rendering them stiff. The outer mesocarp layer contracted by 300% as it dried. Removal of this outer layer inhibited the opening movement, indicating that it is the active tissue. A biomechanical hygro-elastic model based on the relative thicknesses of the layers successfully simulated the opening curvature. Our findings suggest that the sesame capsules possess a functionally graded architecture, which promotes a non-uniform double-curvature hygroscopic bending movement. In contrast to other hygroscopic organs described in the literature, the sesame capsule actuating and resisting tissues are not uniform throughout the device, but changing gradually. This newly described mechanism can be exploited in bio-inspired designs of novel actuating platforms

Spectroscopic Discrimination of Sorghum Silica Phytoliths

Grasses accumulate silicon in the form of silicic acid, which is precipitated as amorphous silica in microscopic particles termed phytoliths. These particles comprise a variety of morphologies according to the cell type in which the silica was deposited. Despite the evident morphological differences, phytolith chemistry has mostly been analysed in bulk samples, neglecting differences between the varied types formed in the same species. In this work, we extracted leaf phytoliths from mature plants of Sorghum bicolor (L.) Moench. Using solid state NMR and thermogravimetric analysis, we show that the extraction methods alter greatly the silica molecular structure, its condensation degree and the trapped organic matter. Measurements of individual phytoliths by Raman and synchrotron FTIR microspectroscopies in combination with multivariate analysis separated bilobate silica cells from prickles and long cells, based on the silica molecular structures and the fraction and composition of occluded organic matter. The variations in structure and composition of sorghum phytoliths suggest that the biological pathways leading to silica deposition vary between these cell types.Peer Reviewe

Multimodal Imaging of Silicified Sorghum Leaves

The plant cell wall is a complex composite material made of polysaccharides, polyphenols, proteins, and minerals. In this work, a multimodal imaging approach was taken, using Raman and Fourier-transform infrared (FTIR) microspectroscopy along with fluorescence imaging, scanning electron microscopy (SEM), and elemental mapping by energy dispersive X-ray spectroscopy (EDX). We characterized the chemical composition of sorghum leaf cross-sections extracted from fresh tissue as well as after paraffin embedding. The complementary vibrational information of Raman and FTIR spectra related a silica deposition to a specific organic composition in the epidermis, specifically with respect to lignin. Moreover, the data enable in situ correlation of autofluorescence with a specific lignin structure. Our results showed that lignin 5–5’ linkages that produce biphenyl structures are important determinants of the cell wall fluorescence properties. The reported multimodal approach will help to clarify the process of biosilica formation and related questions regarding cell wall biochemistry.Peer Reviewe

An early Iron Age assemblage of faience beads from Ashkelon, Israel: chemical composition and manufacturing process

International audienceThe microstructure and chemical composition of eight faience beads from an early Iron Age (12th centuryBCE) assemblage found in the ancient city port of Ashkelon (Israel) are determined by means of FTIRspectrometry, pXRF, microRaman and SEM-EDS analysis. The results are compared with published dataof Egyptian and Near Eastern artifacts. Each sample exhibits a hue which is obtained by adding a specificcolorant to the glazing mixture. A new gray chalcopyrite-manganese-based colorant was identified.Cementation glazing was most likely used in the manufacturing process of the specimens analyzed,except for the blue bead, which is an Egyptian blue frit. The results suggest that these objects represent aunique assemblage, quite different from contemporary Egyptian and Near Eastern materials, and providenew information regarding the Iron Age faience evidence in the southern Levant