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Cell walls in plants, particularly forest trees, are the major carbon sink of the terrestrial ecosystem. Chemical and biosynthetic features of plant cell walls were revealed early on, focusing mostly on herbaceous model species. Recent developments in genomics, transcriptomics, epigenomics, transgenesis, and associated analytical techniques are enabling novel insights into formation of woody cell walls. Here, we review multilevel regulation of cell wall biosynthesis in forest tree species. We highlight current approaches to engineering cell walls as potential feedstock for materials and energy and survey reported field tests of such engineered transgenic trees. We outline opportunities and challenges in future research to better understand cell type biogenesis for more efficient wood cell wall modification and utilization for biomaterials or for enhanced carbon capture and storage. This review highlights recent discoveries in the biogenesis and function of three xylem cell types and multilevel regulation of cell wall biosynthesis for wood formation. Moreover, it provides a comprehensive summary of current understanding of the biosynthesis and modification of cell wall components as well as utilization of plant cell walls. Finally, opportunities and challenges for future research of cell walls in forest tree species are discussed. [ABSTRACT FROM AUTHOR]

The outer wall of pollen and spores, namely the exine, is composed of sporopollenin, which is highly resistant to chemical reagents and enzymes. In this study, we demonstrated that phenylpropanoid pathway derivatives are essential components of sporopollenin in seed plants. Spectral analyses showed that the autofluorescence of Lilium and Arabidopsis sporopollenin is similar to that of lignin. Thioacidolysis and NMR analyses of pollen from Lilium and Cryptomeria further revealed that the sporopollenin of seed plants contains phenylpropanoid derivatives, including p -hydroxybenzoate (p -BA), p -coumarate (p -CA), ferulate (FA), and lignin guaiacyl (G) units. The phenylpropanoid pathway is expressed in the tapetum in Arabidopsis , consistent with the fact that the sporopollenin precursor originates from the tapetum. Further germination and comet assays showed that this pathway plays an important role in protection of pollen against UV radiation. In the pteridophyte plant species Ophioglossum vulgatum and Lycopodium clavata , phenylpropanoid derivatives including p -BA and p -CA were also detected, but G units were not. Taken together, our results indicate that phenylpropanoid derivatives are essential for sporopollenin synthesis in vascular plants. In addition, sporopollenin autofluorescence spectra of bryophytes, such as Physcomitrella and Haplocladium , exhibit distinct characteristics compared with those of vascular plants, indicating the diversity of sporopollenin among land plants. The spectral and biochemical characteristics of sporopollenin are distinct among seed plants, pteridophytes, and bryophytes. This study discovers that the phenylpropanoid pathway is required for the synthesis of both lignin and sporopollenin in vascular plants. Moreover, this study shows that this pathway plays an important role in pollen protection against UV radiation. [ABSTRACT FROM AUTHOR]

Lignin is specifically deposited in plant secondary cell walls, and initiation of lignin biosynthesis is regulated by a variety of developmental and environmental signals. However, the mechanisms governing the regulation of lignin biosynthesis remain to be elucidated. In this study, we identified a lignin biosynthesis-associated transcription factor (LTF) from Populus , LTF1, which binds the promoter of a key lignin biosynthetic gene encoding 4-coumarate-CoA ligase (4CL). We showed that LTF1 in its unphosphorylated state functions as a regulator restraining lignin biosynthesis. When LTF1 becomes phosphorylated by PdMPK6 in response to external stimuli such as wounding, it undergoes degradation through a proteasome pathway, resulting in activation of lignification. Expression of a phosphorylation-null mutant version of LTF1 led to stable protein accumulation and persistent attenuation of lignification in wood cells. Taken together, our study reveals a mechanism whereby LTF1 phosphorylation acts as a sensory switch to regulate lignin biosynthesis in response to environmental stimuli. The discovery of novel modulators and mechanisms modifying lignin biosynthesis has important implications for improving the utilization of cell-wall biomass. This study reports that an MYB transcription factor, LTF1, regulates lignin biosynthesis in Populus. LTF1 binds the promoters of lignin biosynthesis genes to repress the lignin biosynthesis pathway, but this repression can be released when LTF1 is phosphorylated and degraded in response to external stimuli. Thus, LTF1 acts as a sensory switch modulating lignin biosynthesis in wood cells. [ABSTRACT FROM AUTHOR]

Abstract Breeding semi-dwarf varieties to improve lodging resistance has been proven to be enormously successful in increasing grain yield since the advent of the "green revolution." However, the breeding of the majority of semi-dwarf rice varieties in Asia has been dependent mainly on genetic introduction of the mutant alleles of SD1 , which encodes a gibberellin (GA) 20-oxidase, OsGA20ox2, for catalyzing GA biosynthesis. Here, we report a new rice lodging-resistance gene, Shortened Basal Internodes (SBI), which encodes a gibberellin 2-oxidase and specifically controls the elongation of culm basal internodes through deactivating GA activity. SBI is predominantly expressed in culm basal internodes. Genetic analyses indicate that SBI is a semi-dominant gene affecting rice height and lodging resistance. SBI allelic variants display different activities and are associated with the height of rice varieties. Breeding with higher activity of the SBI allele generates new rice varieties with improved lodging resistance and increased yield. The discovery of the SBI provides a desirable gene resource for producing semi-dwarf rice phenotypes and offers an effective strategy for breeding rice varieties with enhanced lodging resistance and high yield. We report that Shortened Basal Internodes (SBI) encodes a gibberellin 2-oxidase catalyzing the conversion of bioactive GA into inactive GA molecules and contributing to lodging resistance in rice. SBI regulates the elongation of culm basal internodes and can be used in breeding to generate elite rice varieties with an ideal culm structure for improved lodging resistance and increased grain production. [ABSTRACT FROM AUTHOR]

The plant vascular system is an elaborate network of conducting and supporting tissues that extends throughout the plant body, and its structure and function must be orchestrated with different environmental conditions. Under high temperature, plants display thin and lodging stems that may lead to decreased yield and quality of crops. However, the molecular mechanism underlying high-temperature-mediated regulation of vascular development is not known. Here, we show that Arabidopsis plants overexpressing the basic-helix-loop-helix (bHLH) transcription factor PHYTOCHROME INTERACTING FACTOR 4 (PIF4), a central regulator of high-temperature signaling, display fewer vascular bundles (VBs) and decreased secondary cell wall (SCW) thickening, mimicking the lodging inflorescence stems of high-temperature-grown wild-type plants. Rising temperature and elevated PIF4 expression reduced the expression of MIR166 and, concomitantly, elevated the expression of the downstream class III homeodomain leucine-zipper (HD-ZIP III) family gene HB15. Consistently, knockdown of miR166 and overexpression of HB15 led to inhibition of vascular development and SCW formation, whereas the hb15 mutant displayed the opposite phenotype in response to high temperature. Moreover, in vitro and in vivo assays verified that PIF4 binds to the promoters of several MIR166 genes and represses their expression. Our study establishes a direct functional link between PIF4 and the miR166-HB15 module in modulating vascular development and SCW thickening and consequently stem-lodging susceptibility at elevated temperatures. [Display omitted] • At high temperature, Arabidopsis exhibits lodging stem of inhibited vascular system • PIF4 inhibits vascular development and SCW thickening • PIF4 directly binds and suppresses MIR166 expression, releasing downstream HB15 • Upregulated HB15 results in the inhibition of vascular development and SCW formation Wei et al. report a mechanism underlying high-temperature-induced inflorescence stem lodging in Arabidopsis. PIF4 is important in high-temperature-induced inhibition of the vascular system. PIF4 directly represses MIR166 genes to release the downstream target HB15 , in turn inhibiting vascular development and secondary cell wall formation. [ABSTRACT FROM AUTHOR]

Crosstalk among phytohormones is crucial for balancing plant growth and adjustment to various environments. Abscisic acid (ABA) and brassinosteroids (BRs) exhibit antagonistic interactions during many plant development processes, but little is known about the molecular mechanism mediating those interactions. Here, we identified a rice (Oryza sativa) remorin gene, OsREM4.1, whose expression is upregulated by ABA through the transcriptional activator OsbZIP23. OsREM4.1, in return, negatively regulates BR signaling output. We discovered that OsREM4.1 interacts with OsSERK1 to inhibit its interaction with rice BR receptor OsBRI1. Moreover, OsBRI1 could phosphorylate OsREM4.1 to reduce the binding affinity of OsREM4.1 to OsSERK1. These results demonstrate that OsREM4.1 is transcriptionally regulated by ABA and functions as an OsBRI1 substrate and OsSERK1-interacting protein to inhibit the formation and subsequent activation of the OsBRI1-OsSERK1 receptor complex. Our findings provide insight into the mechanism by which the antagonistic interactions between ABA and BRs are coordinated in rice., (Copyright © 2016 Elsevier Inc. All rights reserved.)

Endo-1,4-β-glucanases (EGases) are involved in many aspects of plant growth. Our previous study found that an EGase, PtrCel9A6, is specifically expressed in differentiating xylem cells during Populus secondary growth. In this study, the xylem-specific PtrCel9A6 was characterized for its role in xylem differentiation. The EGase is localized on the plasma membrane with catalytic domain toward the outside cell wall, hydrolyzing amorphous cellulose. Suppression of PtrCel9A6 expression caused secondary cell wall defects in xylem cells and significant cellulose reduction in Populus. Heterologous expression of PtrCel9A6 in Arabidopsis enhanced plant growth as well as increased fiber cell length. In addition, introduction of PtrCel9A6 into Arabidopsis resulted in male sterility due to defects in anther dehiscence. Together, these results demonstrate that PtrCel9A6 plays a critical role in remodeling the 1,4-β-glucan chains in the wall matrix and is required for cell wall thickening during Populus xylem differentiation.

A key question in the secondary growth of trees is how differentiation of the vascular cambium cells is directed to concurrently form two different tissues: xylem or phloem. class III homeodomain-leucine zipper (HD-Zip III) genes are known to play critical roles in the initiation, patterning, and differentiation of the vascular system in the process of primary and secondary growth. However, the mechanism of how these genes control secondary vascular differentiation is unknown. Here, we show that a Populus class III HD-Zip gene, PtrHB7, was preferentially expressed in cambial zone. PtrHB7-suppressed plants displayed significant changes in vascular tissues with a reduction in xylem but increase in phloem. Transcriptional analysis revealed that genes regulating xylem differentiation were down-regulated, whereas genes regulating phloem differentiation were up-regulated. Correspondingly, PtrHB7 overexpression enhanced differentiation of cambial cells toward xylem cells but inhibited phloem differentiation. PtrHB7 regulation on cambial cell differentiation was associated with its transcript abundance. Together, the results demonstrated that PtrHB7 plays a critical role in controlling a balanced differentiation between secondary xylem and phloem tissues in the process of Populus secondary growth in a dosage-dependent manner.