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Alfalfa (Medicago sativa L.) forage quality is adversely affected by lignin deposition in cell walls at advanced maturity stages. Reducing lignin content through RNA interference or antisense approaches has been shown to improve alfalfa forage quality and digestibility. We employed a multiplex CRISPR/Cas9-mediated geneediting system to reduce lignin content and alter lignin composition in alfalfa by targeting the COUMARATE 3-HYDROXYLASE (MsC3H) gene, which encodes a key enzyme in lignin biosynthesis. Four guide RNAs (gRNAs) targeting the first exon of MsC3H were designed and clustered into a tRNA-gRNA polycistronic system and introduced into tetraploid alfalfa via Agrobacterium-mediated transformation. Out of 130 transgenic lines, at least 73 lines were confirmed to contain gene-editing events in one or more alleles of MsC3H. Fifty-five lines were selected for lignin content/composition analysis. Amongst these lines, three independent tetra-allelic homozygous lines (Msc3h-013, Msc3h-121, and Msc3h-158) with different mutation events in MsC3H were characterized in detail. Homozygous mutation of MsC3H in these three lines significantly reduced the lignin content and altered lignin composition in stems. Moreover, these lines had significantly lower levels of acid detergent fiber and neutral detergent fiber as well as higher levels of total digestible nutrients, relative feed values, and in vitro true dry matter digestibility. Taken together, these results showed that CRISPR/Cas9-mediated editing of MsC3H successfully reduced shoot lignin content, improved digestibility, and nutritional values without sacrificing plant growth and biomass yield. These lines could be used in alfalfa breeding programs to generate elite transgene-free alfalfa cultivars with reduced lignin and improved forage quality. [ABSTRACT FROM AUTHOR]

Nitrogen can be taken up by trees in the form of nitrate, ammonium and amino acids, but the influence of the different forms on tree growth and development is poorly understood in angiosperm species like Populus. We studied the effects of both organic and inorganic forms of nitrogen on growth and wood formation of hybrid aspen trees in experimental conditions that allowed growth under four distinct steady‐state nitrogen levels. Increased nitrogen availability had a positive influence on biomass accumulation and the radial dimensions of both xylem vessels and fibers, and a negative influence on wood density. An optimal level of nitrogen availability was identified where increases in biomass accumulation outweighed decreases in wood density. None of these responses depended on the source of nitrogen except for shoot biomass accumulation, which was stimulated more by treatments complemented with nitrate than by ammonium alone or the organic source arginine. The most striking difference between the nitrogen sources was the effect on lignin composition, whereby the abundance of H‐type lignin increased only in the presence of nitrate. The differential effect of nitrate is possibly related to the well‐known role of nitrate as a signaling compound. RNA‐sequencing revealed that while the lignin‐biosynthetic genes did not significantly (FDR <0.01) respond to added NO3−, the expression of several laccases, catalysing lignin polymerization, was dependent on N‐availability. These results reveal a unique role of nitrate in wood formation and contribute to the knowledge basis for decision‐making in utilizing hybrid aspen as a bioresource. [ABSTRACT FROM AUTHOR]

Alfalfa (Medicago sativa L.) forage quality is adversely affected by lignin deposition in cell walls at advanced maturity stages. Reducing lignin content through RNA interference or antisense approaches has been shown to improve alfalfa forage quality and digestibility. We employed a multiplex CRISPR/Cas9-mediated gene-editing system to reduce lignin content and alter lignin composition in alfalfa by targeting the COUMARATE 3-HYDROXYLASE (MsC3H) gene, which encodes a key enzyme in lignin biosynthesis. Four guide RNAs (gRNAs) targeting the first exon of MsC3H were designed and clustered into a tRNA-gRNA polycistronic system and introduced into tetraploid alfalfa via Agrobacterium-mediated transformation. Out of 130 transgenic lines, at least 73 lines were confirmed to contain gene-editing events in one or more alleles of MsC3H. Fifty-five lines were selected for lignin content/composition analysis. Amongst these lines, three independent tetra-allelic homozygous lines (Msc3h-013, Msc3h-121, and Msc3h-158) with different mutation events in MsC3H were characterized in detail. Homozygous mutation of MsC3H in these three lines significantly reduced the lignin content and altered lignin composition in stems. Moreover, these lines had significantly lower levels of acid detergent fiber and neutral detergent fiber as well as higher levels of total digestible nutrients, relative feed values, and in vitro true dry matter digestibility. Taken together, these results showed that CRISPR/Cas9-mediated editing of MsC3H successfully reduced shoot lignin content, improved digestibility, and nutritional values without sacrificing plant growth and biomass yield. These lines could be used in alfalfa breeding programs to generate elite transgene-free alfalfa cultivars with reduced lignin and improved forage quality.

Soil organic carbon (SOC) stocks have been greatly depleted across the globe by conversion of wetlands to croplands, agroforestry, and urban areas. Here, we investigated SOC distribution and turnover in four land use types: a wetland, cropland, forestland, and construction land in the Baiyangdian Wetland, Northern China. The C : N ratios were up to 1.70‐times larger in cropland, forestland, and construction land than in the original wetland because of faster N losses compared to C following wetland conversion. The δ13C values of SOC increased with depth in wetland, and showed an overall depletion compared with the other three land use types. Acid‐to‐Aldehyde ratios of syringyl in wetland were 0.72–1.14‐, 0.72–1.72‐, and 1.18–1.43‐times, and cinnamyl/vanillyl ratios were 0.56–1.05‐, 0.22–0.48‐, and 0.40–0.76‐times those of cropland, forestland, and construction land, which reflects faster lignin decomposition rate in wetlands. The β value was defined by the slope of the linear regression between the logarithm of SOC and δ13SOC values, and decreased from cropland over construction land and forestland to wetland, reflecting the faster SOC turnover with the lower β values. However, SOC content and storage were up to 2.29‐ and 2.07‐times higher in wetlands than in soils of other land use types. The combination of C : N ratios, δ13C, and lignin monomer composition can explain the decrease of β value (corresponding to faster SOC turnover), and can be used as effective proxies to evaluate the sources and turnover of SOC in response to land use changes. [ABSTRACT FROM AUTHOR]

The shells of three important food nuts, walnut, almond, and pine nut, were studied in view of valorization as residues. The shells differed chemically: walnut shells had 10.6% extractives, 30.1% lignin, and 49.7% polysaccharides; almond shells 5.7% extractives, 28.9% lignin, and 56.1% polysaccharides; and pine nut shells 4.5% extractives, 40.5% lignin, and 48.7% polysaccharides. The polysaccharide composition also differed, e.g., glucose/xylose ratio of 1.12, 0.94, and 2.29 for walnut, almond, and pine nut shells, respectively. Walnut and almond shells have a SG lignin (S/G 1.6 and 1.0, respectively) and pine nut shell a G lignin. The lipophilic extracts contained mostly saturated and unsaturated alkanoic acids. The ethanol-water extracts contained total phenolics, flavonoids, and condensed tannins. The antioxidant activity was moderate (IC50 15.2, 7.9, and 8.2 μg/mL for walnut, almond, and pine nut). The three nut shells fractured easily with little formation of fines. [ABSTRACT FROM AUTHOR]

Abstract Plants respond to the loss of vertical growth re-orientating their affected organs. In trees, this phenomenon has received the scientific attention due to its importance for the forestry industry. Nowadays it is accepted that auxin distribution is involved in the modulation of the tilting response, but how this distribution is controlled is not fully clear. Auxin transporters that determine the spatio-temporal auxin distribution in radiate pine seedlings exposed to 45° of tilting were identified. Additionally, based on indications for an intimate plant hormone crosstalk in this process, IAA and JA contents were evaluated. The experiments revealed that expression of the auxin transporters was down-regulated in the upper half of the tilted stem, while being induced in the lower half. Moreover, transporter-coding genes were first induced at the apical zone of the stem. IAA was consistently redistributed toward the lower half, which is in accordance with the expression profile of the auxin transporters. In contrast, JA was mainly accumulated in the upper half of tilted stems. Finally, lignin content and monomeric composition were analyzed in both sides of stem and along the time course of tilting. As expected, lignin accumulation was higher at the lower half of stem at longer times of tilting. However, the most marked difference was the accumulation of the H-lignin monomer in the lower half, while the G-lignin unit was more dominant in the upper half. Here, we provide detailed insight in the distribution of IAA and JA, affecting the lignin composition during the tilting response in Pinus radiata seedlings. Graphical abstract Image 1 Highlights • Six auxin transporter transcripts were identified in stems of radiata pine after tilting. • Differential transcript abundance was recorded along the stem. • Auxin and jasmonic acid content change in a spatio-temporal manner in tilted stems. • Lignin monomer composition is differentially accumulated along the tilted stems. [ABSTRACT FROM AUTHOR]

Abstract Tropical peatlands are very different from boreal and temperate regions, because they have particular climatic characteristics and different vegetation types. Similarly, high mountain peatlands differ significantly from those found at low altitudes. Lignin components were investigated as phyto-physiological proxies of landscape evolution in two big national environmental conservation parks of Southeastern Brazil. Lignin phenols showed a high degree of humification and indicated the same land-use for both forest parks, suggesting minimal anthropic intervention over the last 150 years, predominating native grasslands. However, total organic carbon (TOC) and total nitrogen (TN) parameters evidenced the influence of indirect effects of human activities, mainly from atmospheric deposition since 1950s, associated to the early Brazilian industrial development. This study reinforces the importance of protection and conservation to those pristine areas. Graphical abstract Unlabelled Image Highlights • Lignin was a useful biomarker of land-use change over time. • Lignin phenols indicated a high degree of humification in tropical peatlands. • C and lignin storage changes were due to industrial activities and/or environmental changes. [ABSTRACT FROM AUTHOR]

Summary: A reduction in the lignin content in transgenic plants induces the ectopic expression of defense genes, but the importance of altered lignin composition in such phenomena remains unclear. Two Arabidopsis lines with similar lignin contents, but strikingly different lignin compositions, exhibited different quantitative and qualitative transcriptional responses. Plants with lignin composed primarily of guaiacyl units overexpressed genes responsive to oomycete and bacterial pathogen attack, whereas plants with lignin composed primarily of syringyl units expressed a far greater number of defense genes, including some associated with cis‐jasmone‐mediated responses to aphids; these plants exhibited altered responsiveness to bacterial and aphid inoculation. Several of the defense genes were differentially induced by water‐soluble extracts from cell walls of plants of the two lines. Glycome profiling, fractionation and enzymatic digestion studies indicated that the different lignin compositions led to differential extractability of a range of heterogeneous oligosaccharide epitopes, with elicitor activity originating from different cell wall polymers. Alteration of lignin composition affects interactions with plant cell wall matrix polysaccharides to alter the sequestration of multiple latent defense signal molecules with an impact on biotic stress responses. [ABSTRACT FROM AUTHOR]

Main conclusion: CAD-deficient poplars enabled studying the influence of altered lignin composition on mechanical properties. Severe alterations in lignin composition did not influence the mechanical properties.Wood represents a hierarchical fiber-composite material with excellent mechanical properties. Despite its wide use and versatility, its mechanical behavior has not been entirely understood. It has especially been challenging to unravel the mechanical function of the cell wall matrix. Lignin engineering has been a useful tool to increase the knowledge on the mechanical function of lignin as it allows for modifications of lignin content and composition and the subsequent studying of the mechanical properties of these transgenics. Hereby, in most cases, both lignin composition and content are altered and the specific influence of lignin composition has hardly been revealed. Here, we have performed a comprehensive micromechanical, structural, and spectroscopic analysis on xylem strips of transgenic poplar plants, which are downregulated for cinnamyl alcohol dehydrogenase (CAD) by a hairpin-RNA-mediated silencing approach. All parameters were evaluated on the same samples. Raman microscopy revealed that the lignin of the hpCAD poplars was significantly enriched in aldehydes and reduced in the (relative) amount of G-units. FTIR spectra indicated pronounced changes in lignin composition, whereas lignin content was not significantly changed between WT and the hpCAD poplars. Microfibril angles were in the range of 18°-24° and were not significantly different between WT and transgenics. No significant changes were observed in mechanical properties, such as tensile stiffness, ultimate stress, and yield stress. The specific findings on hpCAD poplar allowed studying the specific influence of lignin composition on mechanics. It can be concluded that the changes in lignin composition in hpCAD poplars did not affect the micromechanical tensile properties. [ABSTRACT FROM AUTHOR]

BACKGROUND The relationship between the chemical and molecular properties - in particular the (acid detergent) lignin (ADL) content and composition expressed as the ratio between syringyl and guaiacyl compounds (S:G ratio) - of maize stems and in vitro gas production was studied in order to determine which is more important in the degradability of maize stem cell walls in the rumen of ruminants. Different internodes from two contrasting maize cultivars (Ambrosini and Aastar) were harvested during the growing season. RESULTS The ADL content decreased with greater internode number within the stem, whereas the ADL content fluctuated during the season for both cultivars. The S:G ratio was lower in younger tissue (greater internode number or earlier harvest date) in both cultivars. For the gas produced between 3 and 20 h, representing the fermentation of cell walls in rumen fluid, a stronger correlation ( R2 = 0.80) was found with the S:G ratio than with the ADL content ( R2 = 0.68). The relationship between ADL content or S:G ratio and 72-h gas production, representing total organic matter degradation, was weaker than that with gas produced between 3 and 20 h. CONCLUSION The S:G ratio plays a more dominant role than ADL content in maize stem cell wall degradation. © 2017 The Authors. Journal of The Science of Food and Agriculture published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry. [ABSTRACT FROM AUTHOR]

Analysis of thin slivers of juvenile (2-year-old) pine wood by pyrolysis-gas chromatography-mass spectrometry (Py-GC/MS) was explored as a fast and highthroughput method to assess compression wood (CW) via the ratio of the p-hydroxyphenyl to guaiacyl units (H/G ratio). Various chemical parameters such as contents of lignin, galactose, and p-hydroxyphenyl (H) units have been related to the CW in the tissue. The H/G ratio was calculated from all identified H and G lignin-derived pyrolysis products. The H/G ratio ranged between 0.019 and 0.052 for normal wood (NW) and between 0.133 and 0.227 for CW. There was a weak correlation between CW H/G ratio and Klason lignin content (R2 = 0.4), but it was not correlated to the galactose content. Lignin- and carbohydrate-derived pyrolysis products were subjected to principal component (PC) analysis. The first two PCs discriminate the CW from the NW, accounting for about 41% of the total variance in the dataset. The model developed based on lignin-derived peaks explained 54% of the variance. [ABSTRACT FROM AUTHOR]

Water bamboo shoots (WBS) are popular hydrophytic plant in China with a high nutritive value. However, WBS is susceptible to postharvest lignification. The firmness, lignin content, lignin composition, activity of lignin biosynthesis-related enzymes, and expression of lignin biosynthesis-related genes in WBS were determined after postharvest storage at 20 °C or 2 °C for 15 d or 63 d, respectively. The results indicated that lignin present in WBS was composed of G (vanillin and vanillic acid), H (p-hydroxybenzaldehyde and p-hydroxybenzoic acid), and S (syringaldehyde and syringic acid) monomers. During storage, the activities of phenylalanine ammonia-lyase, cinnamyl alcohol dehydrogenase, and peroxidase also increased. Moreover, the upregulated gene expression levels of lignin biosynthesis-related enzymes positively correlated with increasing lignin content, which were the key contributors to the lignification of WBS during storage. We observed that low temperatures delayed lignification by repressing the expression of lignification-related genes and suppressing the activity of lignin biosynthesis-related enzymes. • Low temperature regulated lignin-related enzyme activity and gene expression. • Temperature and storage time could affect lignin composition in WBS during storage. • Lignin has a positive relationship with PAL and CAD activities and gene expression. [ABSTRACT FROM AUTHOR]

As the attraction of creating biofuels and bio-based chemicals from lignocellulosic biomass has increased, researchers have been challenged with developing a better understanding of lignin structure, quantity and potential uses. Lignin has frequently been considered a waste-product from the deconstruction of plant cell walls, in attempts to isolate polysaccharides that can be hydrolyzed and fermented into fuel or other valuable commodities. In order to develop useful applications for lignin, accurate analytical instrumentation and methodologies are required to qualitatively and quantitatively assess, for example, what the structure of lignin looks like or how much lignin comprises a specific feedstock׳s cellular composition. During the past decade, various diverse strategies have been employed to elucidate the structure and composition of lignin. These techniques include using two-dimensional nuclear magnetic resonance to resolve overlapping spectral data, measuring biomass with vibrational spectroscopy to enable modeling of lignin content or monomeric ratios, methods to probe and quantify the linkages between lignin and polysaccharides, or refinements of established methods to provide higher throughput analyses, less use of consumables, etc. This review seeks to provide a comprehensive overview of many of the advancements achieved in evaluating key lignin attributes. Emphasis is placed on research endeavored in the last decade. [ABSTRACT FROM AUTHOR]

Background: Interest in the detailed lignin and polysaccharide composition of plant cell walls has surged within the past decade partly as a result of biotechnology research aimed at converting biomass to biofuels. High-resolution, solution-state 2D 1H-13C HSQC NMR spectroscopy has proven to be an effective tool for rapid and reproducible fingerprinting of the numerous polysaccharides and lignin components in unfractionated plant cell wall materials, and is therefore a powerful tool for cell wall profiling based on our ability to simultaneously identify and comparatively quantify numerous components within spectra generated in a relatively short time. However, assigning peaks in new spectra, integrating them to provide relative component distributions, and producing color-assigned spectra, are all current bottlenecks to the routine use of such NMR profiling methods. Results: We have assembled a high-throughput software platform for plant cell wall profiling that uses spectral deconvolution by Fast Maximum Likelihood Reconstruction (FMLR) to construct a mathematical model of the signals present in a set of related NMR spectra. Combined with a simple region of interest (ROI) table that maps spectral regions to NMR chemical shift assignments of chemical entities, the reconstructions can provide rapid and reproducible fingerprinting of numerous polysaccharide and lignin components in unfractionated cell wall material, including derivation of lignin monomer unit (S:G:H) ratios or the so-called SGH profile. Evidence is presented that ROI-based amplitudes derived from FMLR provide a robust feature set for subsequent multivariate analysis. The utility of this approach is demonstrated on a large transgenic study of Arabidopsis requiring concerted analysis of 91 ROIs (including both assigned and unassigned regions) in the lignin and polysaccharide regions of almost 100 related 2D 1H-13C HSQC spectra. Conclusions: We show that when a suitable number of replicates are obtained per sample group, the correlated patterns of enriched and depleted cell wall components can be reliably and objectively detected even prior to multivariate analysis. The analysis methodology has been implemented in a publicly-available, cross-platform (Windows/ Mac/Linux), web-enabled software application that enables researchers to view and publish detailed annotated spectra in addition to summary reports in simple spreadsheet data formats. The analysis methodology is not limited to studies of plant cell walls but is amenable to any NMR study where ROI segmentation techniques generate meaningful results. [ABSTRACT FROM AUTHOR]

Abstract: Samples of cork tissues separated from the barks of Betula pendula, Quercus suber and Quercus cerris were characterized by Py–GC–MS/FID in respect to composition of lignin. The pyrolysis temperature influenced the yield of pyrolysis products, and in the conditions usually applied to wood pyrolysis (550°C) suberin is only partially degraded into products. The lignin composition of the three corks showed predominantly a guaiacyl-based lignin: guaiacyl (G) units represented 85.7%, 96.4% and 93.7% of lignin respectively in B. pendula, Q. suber and Q. cerris corks, while syringyl (S) units amounted respectively to 11.9%, 2.5% and 2.7% and p-hydroxyphenyl (H) units to 2.4%, 1.1% and 3.6%. Pyrolysis of the woods of these same species confirmed the GS character of their lignins in contrast to the corks G-lignin type. The influence of ferulic acid in the results and its integration in a heterogeneous cork lignin structure was discussed. [Copyright &y& Elsevier]

We have recently described a hitherto unsuspected catechyl lignin polymer (C-lignin) in the seed coats of Vanilla orchid and in cacti of one genus, Melocactus (Chen et al., Proc. Natl. Acad. Sci. USA. 2012, 109, 1772-1777.). We have now determined the lignin types in the seed coats of 130 different cactus species. Lignin in the vegetative tissues of cacti is of the normal guaiacyl/syringyl (G/S) type, but members of most genera within the subfamily Cactoidae possess seed coat lignin of the novel C-type only, which we show is a homopolymer formed by endwise β-O-4-coupling of caffeyl alcohol monomers onto the growing polymer resulting in benzodioxane units. However, the species examined within the genera Coryphantha, Cumarinia, Escobaria and Mammillaria (Cactoideae) mostly had normal G/S lignin in their seeds, as did all six species in the subfamily Opuntioidae that were examined. Seed coat lignin composition is still evolving in the Cactaceae, as seeds of one Mammillaria species (M. lasiacantha) possess only C-lignin, three Escobaria species (E. dasyacantha, E. lloydii and E. zilziana) contain an unusual lignin composed of 5-hydroxyguaiacyl units, the first report of such a polymer that occurs naturally in plants, and seeds of some species contain no lignin at all. We discuss the implications of these findings for the mechanisms that underlie the biosynthesis of these newly discovered lignin types. [ABSTRACT FROM AUTHOR]

Heartwood and sapwood samples from Eucalyptus globulus were char-acterized by Py-GC/MS and GC-FID in respect to composition and content of lignin. The pyrolysis lignin-derived compounds were assembled by gmups: "syringol," "S-aldehydes," "S-ketones," "S-alcohols," and "C11H12O3" (S-units); "guaiacol," "eugenol," "G-aldehydes," "G-ketones," "G-alcohols," and "others" (G-units); "phe-nol" and "H-aldehydes" (H-units). Heartwood and sapwood had similar lignin content in an extractive-free basis (23.7% and 23.0%, respectively) and in lignin composition (S-units, 76.0% vs. 76.3%; G-units, 22.0% vs. 21.0%; H-units, 1.9% vs. 2.7%; S/G ratio 3.5 and 3.6). The wood samples were kraft pulped under isothermal conditions at 130°C, 150°C and 170°C and several cooking times. Heartwood and sapwood behaved similarly. Al 130°C llie delignification was slow willi no significant selectivity in respect to lignin composition. At 150°C and 170°C the S-units were more susceptible to reaction and comparatively more removed, inducing a decrease of S/G ratio to 0.6. The main prod-ucts to be extracted belong to "syringol" and "S-aldehydes," while the residual lignin in pulps was enriched in "guaiacol," "eugenol" (G-units), and "phenol" (H-units). [ABSTRACT FROM AUTHOR]

The lignin structural ramifications of coumarate 3-hydroxylase (C3H) downregulation have not been addressed in hardwoods. Such information is required to accompany an assessment of the digestibility and bioenergy performance characteristics of poplar, in particular. Structurally rich 2D NMR methods were applied to the entire lignin fraction to delineate lignin p-hydroxyphenyl:guaiacyl:syringyl (H:G:S) levels and linkage distribution changes (and to compare with traditional degradative analyses). C3H downregulation reduced lignin levels by half and markedly increased the proportion of H units relative to the normally dominant G and S units. Relative stem H unit levels were up by ∼ 100-fold to ∼ 31 %, almost totally at the expense of G units; differences in the lignin interunit linkage distributions were more subtle. The H level in the most drastically C3H-downregulated transgenic poplar falls well beyond the H:G:S compositional bounds of normal angiosperms. The response observed here, in poplar, differs markedly from that reported for alfalfa where the S:G ratio remained almost constant even at substantial H levels, highlighting the often differing responses among plant species. [ABSTRACT FROM AUTHOR]

Abstract: Smoke taint in wines from bushfire smoke exposure has become a concern for wine producers. Smoke taint compounds are primarily derived from pyrolysis of the lignin component of fuels. This work examined the influence of the lignin composition of pyrolysed vegetation on the types of putative smoke taint compounds that accrue in wines. At veraison, Merlot vines were exposed to smoke generated from five vegetation types with differing lignin composition. Smoke was generated under pyrolysis conditions that simulated bushfire temperature profiles. Lignin and smoke composition of each fuel type along with putative smoke taint compounds in wines were determined. The results showed that, regardless of fuel type, the commonly reported guaiacyl lignin derived smoke taint compounds, guaiacol and 4-methylguaiacol, represented about 20% of the total phenols in wines. Quantitatively, syringyl lignin derived compounds dominated the total phenol pools in both free and bound forms. The contributions of p-hydroxyphenyls were generally similar to the guaiacyl sources. A further unexpected outcome of the study was that pine smoke affected wines had significantly elevated levels of syringols compared to the controls although pine fuel and its smoke emission lacked syringyl products. [Copyright &y& Elsevier]

Abstract: Eucalyptus globulus sapwood and heartwood showed no differences in lignin content (23.0% vs. 23.7%) and composition: syringyl-lignin (17.9% vs. 18.0%) and guaiacyl-lignin (4.8% vs. 5.2%). Delignification kinetics of S- and G-units in heartwood and sapwood was investigated by Py-GC–MS/FID at 130, 150 and 170°C and modeled as double first-order reactions. Reactivity differences between S and G-units were small during the main pulping phase and the higher reactivity of S over G units was better expressed in the later pulping stage. The residual lignin composition in pulps was different from wood or from samples in the initial delignification stages, with more G and H-units. S/G ratio ranged from 3 to 4.5 when pulp residual lignin was higher than 10%, decreasing rapidly to less than 1. The S/H was initially around 20 (until 15% residual lignin), decreasing to 4 when residual lignin was about 3%. [Copyright &y& Elsevier]

The quantity and quality of plant litter in the soil play an important role in the soil organic matter balance. Besides other pedo-climatic aspects, the content of recalcitrant molecules of plant residues and their chemical composition play a major role in the preservation of plant residues. In this study, we report that intrinsically resistant alkali-soluble bio-macromolecules extracted from maize plant (plant-humic acid) (plant-HA) contribute directly to the soil organic matter (OM) by its addition and conservation in the soil. Furthermore, we also observed that a high syringyl/guaiacyl (S/G) ratio in the lignin residues comprising the plant tissue, which modifies the microscopic structure of the alkali-soluble plant biopolymers, enhances their recalcitrance because of lower accessibility of molecules to degrading enzymes. These results are in agreement with a recent study, which showed that the humic substance of soil consists of a mixture of identifiable biopolymers obtained directly from plant tissues that are added annually by maize plant residues. [Copyright &y& Elsevier]

Lignin is the cell wall component most frequently associated with hardening. Its characterization and quantification are very important to understand the biochemical modifications related to the changes in texture of vegetables such as asparagus ( Asparagus officinalis), in which this organoleptic attribute is a very important quality factor. In this study, asparagus lignin from the basal sections of fresh and stored spears was analyzed using 2 methods, the traditional (Klason lignin) and the recently developed derivatization, followed by reductive cleavage (DFRC) method. The latter is a simple and reproducible technique for lignin characterization based on a degradation procedure that produces analyzable monomers and dimers by cleaving α- and β-aryl ethers in lignins. The primary monomers derived from DFRC degradation of lignins are essentially p-coumaryl peracetate, coniferyl peracetate, and sinapyl peracetate. To evaluate the efficiency of the DFRC method, our investigations have been carried on distinct sample types, including wood (data not shown), straw, and asparagus samples. The results have confirmed that lignin composition is affected by plant nature. It has been found that whereas wood samples mostly contain coniferyl units, plant foods, such as straw and asparagus, contain both coniferyl and guaiacyl units. [ABSTRACT FROM AUTHOR]

Abstract: Many aspects of the interactions between host wood structure and fungal activity can be revealed by high resolution light microscopy, and this technique has provided much of the information discussed here. A wide range of different types of decay can result from permutations of host species, fungal species and conditions within wood. Within this spectrum, three main types are commonly recognised: brown rot, white rot and soft rot. The present review explores parts of the range of variation that each of these encompasses and emphasizes that degradation modes appear to reflect a co-evolutionary adaptation of decay fungi to different wood species or the lignin composition within more primitive and advanced wood cell types. One objective of this review is to provide evidence that the terms brown rot, white rot and soft rot may not be obsolete, but rigid definitions for fungi that are placed into these categories may be less appropriate than thought previously. Detailed knowledge of decomposition processes does not only aid prognosis of decay development in living trees for hazard assessment but also allows the identification of wood decay fungi that can be used for biotechnology processes in the wood industry. In contrast to bacteria or commercial enzymes, hyphae can completely ramify through solid wood. In this review evidence is provided that wood decay fungi can effectively induce permeability changes in gymnospermous heartwood or can be applied to facilitate the identification of tree rings in diffuse porous wood of angiosperms. The specificity of their enzymes and the mild conditions under which degradation proceeds is partly detrimental for trees, but also make wood decay fungi potentially efficient biotechnological tools. [Copyright &y& Elsevier]

A Chinese kenaf (Hibiscus cannabinus L.), variety Sekko-ichi, was planted in May 1996, and harvested at 76, 116, 152, 185, and 226 days after planting. The hand-separated bast fibers were analyzed for lignin content and in situ lignin composition to clarify their variations with maturity. Bast fibers increased in lignin until 152 days after planting, and did not increase thereafter. Variations in in situ lignin compositions with maturity were monitored using pyrolysis-gas chromatography. The volatile pyrolysis products released at 500 °C for 4 s were identified by gas chromatography-mass spectrometry. Pyrolysis products derived from guaiacyl and syringyl lignin units were revealed. The yield of guaiacyl lignin-derived pyrolysis products decreased with maturity, while the yield of the syringyl analogous increased with maturity. The ratio of yield of syringyl lignin-derived products to that of guaiacyl ones (S/G) increased from 1.86 to 3.16 with maturity, but remained constant at 152 days after planting. High S/G ratio showed that mature bast lignin is of syringyl type. Quantitative pyrolysis results suggested that harvesting of kenaf at age of 5-months growth period provides a raw material with an ease of delignification. [Copyright &y& Elsevier]

Leaf litters from beech (Fagus sylvatica L.) and oak (Quercus robur L.) trees were collected from mixed, deciduous woodlands growing on three soil types that varied in mineral nutrient concentrations and N mineralisation potential. Litter quality, including %N, %Mn, %P, acid detergent fibre, cellulose, Klason lignin, phenylpropanoid constituents of lignin, hexose and pentose sugar (mainly from hemicelluloses) varied within species according to soil type. However, oak and beech showed the opposite responses to soil nutrient status for most of these variables. The litters were incubated in the laboratory for 12 months (at 18 °C and constant moisture) on beds of forest floor material from two soils of contrasting high nutrient material (HNM) or low nutrient material (LNM) nutrient status to investigate litter quality and substrate interactions. At 4, 8 and 12 months there were significant differences in mass losses from oak and beech litters from all sites, and for each litter type exposed to the HNM and LMN soils. At 12 months mean mass losses were higher for HNM treatment (38.7% oak, 27.8% beech) than for the LNM treatment (30.6% oak, 25.5% beech). However, the beech and oak litters from the different sites consistently responded in opposite ways on the same soil treatment reflecting site-related effects on litter quality. Initial concentration of Klason lignin was the best predictor for mass losses from litter species and litter types. Intra-specific variation in rates of litter decomposition of beech and oak litters from different sites, and differences in their interactions with the two forest floor materials, illustrate the complexities of proximate controls on decomposition that are often masked in system-level studies. [Copyright &y& Elsevier]

• We evaluated larch tree-ring δ2H LM and δ13C LM in individual and pooled, bulk and degreased wood from permafrost forest. • The total lignin content, but not G-lignin and methoxy isotopes, shifted at the boundary between sapwood and heartwood. • The May climate strongly affected the heartwood lignin content and tree growth. • δ2H LM series showed great potential as growing season temperature and VPD proxies. The hydrogen and carbon stable isotope ratios (δ2H LM and δ13C LM values) of tree-ring lignin methoxy groups have recently been recognized as valuable palaeoclimate indicators. However, the environmental factors but also sample preparation processes that might cause variations of δ2H LM and δ13C LM values have been not fully explored. Furthermore, the temporal dynamics of wood lignin content on both isotopes hasn't been investigated. To investigate the effects of total lignin content, removal of lipids and differences between individual and pooled tree-ring series on isotopic values, we analyzed ring samples of dominant Larix gmelinii trees from a typical permafrost region of northeastern China. We also examined relationships between δ2H LM and δ13C LM values with annual-resolution lignin content and climatic parameters over the common period of overlap 1901 to 2013. We found that the inter-tree variability between the individual and pooled isotopic chronologies was consistent, and the removal of lipids did not change the results of stable isotope measurements. The total lignin content shifted between the heartwood and sapwood, whereas the content of guaiacyl monomers (G-lignin), the only donor of methoxy groups, remained constant. Correlations analysis between lignin proxies and climate variables indicated that the heartwood total lignin content was positively correlated with the intrinsic water-use efficiency, temperature and vapor-pressure deficit (VPD), but negatively with the early-growing-season standardized precipitation evaporation index. Whilst δ2H LM values were positively correlated with the growing season temperature (May to August) and VPD, particularly in August, δ13C LM series showed insignificant correlation. Therefore, we suggest that the total lignin content and δ2H LM values of tree rings (both individual and pooled series) are suitable for tracking environment dynamics, as they can reveal the climate responses of tree growth. [ABSTRACT FROM AUTHOR]

Phytohormones are crucial molecules regulating plant development and responses to environmental challenges, including abiotic stresses, microbial and insect attacks. Most notably, phytohormones play important roles in the biosynthesis of lignocellulosics. Jasmonates are involved in secondary growth and secondary metabolism, such as phenylpropanoids and lignin biosyntheses. At the physiological and molecular levels, the actions of phytohormones depend on subtle concentration changes, as well as antagonistic equilibria between two or more of these molecules. In this article, we investigate the consequences of jasmonic acid (JA) spraying on young hemp hypocotyls. First, we show that JA application results in changes in the monomeric composition of lignin. Second, we highlight that, five days after application, JA leads to an increase in salicylic acid (SA) content in hemp hypocotyls. These results are discussed in the light of the known antagonism between JA and SA at both the physiological and molecular levels.

The derivatization followed by reductive cleavage (DFRC) method is a well-established tool to characterize the lignin composition of plant materials. However, the application of the original procedure, especially the chromatographic determination of the DFRC monomers, is problematic for low-lignin foods. To overcome these problems a modified sample cleanup and a stable-isotope dilution approach were developed and validated. To quantitate the diacetylated DFRC monomers, their corresponding hexadeuterated analogs were synthesized and used as internal standards. By using the selected-ion monitoring mode, matrix-associated interferences can be minimized resulting in higher selectivity and sensitivity. The modified method was applied to four low-lignin samples. Lignin from carrot fibers was classified as guaiacyl-rich whereas the lignins from radish, pear, and asparagus fibers where classified as balanced lignins (guaiacyl/syringyl ratio=1-2).