KEGG(Kyoto Encyclopedia of Genes and Genomes)〔和文〕 (特集 ゲノム医学の現在と未来--基礎と臨床) -- (データベース)

Nature’s hierarchical materials

Lignins are complex natural polymers resulting from oxidative coupling of, primarily, 4-hydroxyphenylpropanoids. An understanding of their nature is evolving as a result of detailed structural studies, recently aided by the availability of lignin-biosynthetic-pathway mutants and transgenics. The currently accepted theory is that the lignin polymer is formed by combinatorial-like phenolic coupling reactions, via radicals generated by peroxidase-H2O2, under simple chemical control where monolignols react endwise with the growing polymer. As a result, the actual structure of the lignin macromolecule is not absolutely defined or determined. The ``randomness'' of linkage generation (which is not truly statistically random but governed, as is any chemical reaction, by the supply of reactants, the matrix, etc.) and the astronomical number of possible isomers of even a simple polymer structure, suggest a low probability of two lignin macromolecules being identical. A recent challenge to the currently accepted theory of chemically controlled lignification, attempting to bring lignin into line with more organized biopolymers such as proteins, is logically inconsistent with the most basic details of lignin structure. Lignins may derive in part from monomers and conjugates other than the three primary monolignols (p-coumaryl, coniferyl, and sinapyl alcohols). The plasticity of the combinatorial polymerization reactions allows monomer substitution and significant variations in final structure which, in many cases, the plant appears to tolerate. As such, lignification is seen as a marvelously evolved process allowing plants considerable flexibility in dealing with various environmental stresses, and conferring on them a striking ability to remain viable even when humans or nature alter ``required'' lignin-biosynthetic-pathway genes/enzymes. The malleability offers significant opportunities to engineer the structures of lignins beyond the limits explored to date.

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Lignins: Natural polymers from oxidative coupling of 4-hydroxyphenyl- propanoids

L-tryptophan, L-phenylalanine, and L-tyrosine are aromatic amino acids (AAAs) that are used for the synthesis of proteins and that in plants also serve as precursors of numerous natural products, such as pigments, alkaloids, hormones, and cell wall components. All three AAAs are derived from the shikimate pathway, to which ≥30% of photosynthetically fixed carbon is directed in vascular plants. Because their biosynthetic pathways have been lost in animal lineages, the AAAs are essential components of the diets of humans, and the enzymes required for their synthesis have been targeted for the development of herbicides. This review highlights recent molecular identification of enzymes of the pathway and summarizes the pathway organization and the transcriptional/posttranscriptional regulation of the AAA biosynthetic network. It also identifies the current limited knowledge of the subcellular compartmentalization and the metabolite transport involved in the plant AAA pathways and discusses metabolic engineering efforts aimed at improving production of the AAA-derived plant natural products.

The shikimate pathway and aromatic amino acid biosynthesis in plants.

SECONDARY WALL-ASSOCIATED NAC DOMAIN PROTEIN1 (SND1) is a master transcriptional switch activating the developmental program of secondary wall biosynthesis. Here, we demonstrate that a battery of SND1-regulated transcription factors is required for normal secondary wall biosynthesis in Arabidopsis thaliana. The expression of 11 SND1-regulated transcription factors, namely, SND2, SND3, MYB103, MYB85, MYB52, MYB54, MYB69, MYB42, MYB43, MYB20, and KNAT7 (a Knotted1-like homeodomain protein), was developmentally associated with cells undergoing secondary wall thickening. Of these, dominant repression of SND2, SND3, MYB103, MYB85, MYB52, MYB54, and KNAT7 significantly reduced secondary wall thickening in fiber cells. Overexpression of SND2, SND3, and MYB103 increased secondary wall thickening in fibers, and overexpression of MYB85 led to ectopic deposition of lignin in epidermal and cortical cells in stems. Furthermore, SND2, SND3, MYB103, MYB85, MYB52, and MYB54 were able to induce secondary wall biosynthetic genes. Direct target analysis using the estrogen-inducible system revealed that MYB46, SND3, MYB103, and KNAT7 were direct targets of SND1 and also of its close homologs, NST1, NST2, and vessel-specific VND6 and VND7. Together, these results demonstrate that a transcriptional network consisting of SND1 and its downstream targets is involved in regulating secondary wall biosynthesis in fibers and that NST1, NST2, VND6, and VND7 are functional homologs of SND1 that regulate the same downstream targets in different cell types.

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A battery of transcription factors involved in the regulation of secondary cell wall biosynthesis in Arabidopsis.

Abstract The present study is aimed at the detection of mechanical stresses generated in the cellulose microfibril (CMF) crystals situated in the secondary wall (S2) of living cells. Green wood specimens were boiled in water to release the internal stress in the CMF by the hygrothermal softening of the lignin-hemicellulose matrix (MT). Thereafter, the changes in d200 and d004 lattice spacings of crystalline cellulose were observed in boiled and nonboiled samples by wide-angle X-ray diffraction. The d200 lattice spacing increased, whereas d004 lattice spacing decreased. The results show that a mechanical stress still remained in the CMF and the MT region in the green cell wall, even after releasing the macroscopic surface growth stresses by removal of the wood block from the living stem. The interpretation is that CMF generates tensile stress in the longitudinal direction, and surrounding MT substances generate compressive stress in the living cell wall, which compresses the CMF in the lateral direction. The results confirm the “unified hypothesis” for explaining the mechanism of growth stress generation.

Micromechanical detection of growth stress in wood cell wall by wide-angle X-ray diffraction (WAX)

Effects of the lateral growth rate on wood quality parameters of Eucalyptus grandis from different latitudes in Brazil and Argentina

Tree forms and internal stresses III. Growth stresses of branches

In order to clarify the growth stress mechanism of tree species in tropical forest, peripheral and internal longitudinal released strains in 39 year-old teak (Tectona grandis) trees planted in West Java, Indonesia were investigated using the strain-gauge technique. Results showed that both of the peripheral and internal longitudinal released strains were not affected by the rate of growth of the trees. However, released strains distribution inside tree stem, from pith to periphery, were significantly affected by growth rate; slow growing trees generate steeper released strain gradients.

Relationship between released strain and growth rate in 39 year-old Tectona grandis planted in Indonesia

Plasticity of root growth in response to environmental cues and stresses is a fundamental characteristic of plants, in accordance with their sessile lifestyle. This is linked to the balance between plasticity and rigidity of cells in the root apex, and thus is coordinated with the control of cell wall properties. However, mechanisms underlying such harmonization are not well understood, in particular under stressful conditions. We have recently demonstrated that RICE SALT SENSITIVE3 (RSS3), a nuclear factor that mediates restrictive expression of jasmonate-induced genes, plays an important role in root elongation under saline conditions. In this study, we report that loss-of-function of RSS3 results in changes in cell wall properties such as lignin deposition and sensitivity to a cellulose synthase inhibitor, concomitant with altered expression of genes involved in cell wall metabolism. Based on these and previous phenotypic observations of the rss3 mutant, we propose that RSS3 plays a role in the coordin...

Masato Yoshida

Papers

Micromechanical detection of growth stress in wood cell wall by wide-angle X-ray diffraction (WAX)

Effects of the lateral growth rate on wood quality parameters of Eucalyptus grandis from different latitudes in Brazil and Argentina

Tree forms and internal stresses III. Growth stresses of branches

Relationship between released strain and growth rate in 39 year-old Tectona grandis planted in Indonesia

RICE SALT SENSITIVE3 binding to bHLH and JAZ factors mediates control of cell wall plasticity in the root apex.