A noticeable proportion of low transcribed genes involved in wood formation in conifers may have been missed in previous transcriptomic studies. This could be the case for genes related to less abundant cell types, such as axial parenchyma and resin ducts, and genes related to juvenile wood. In this study, two normalized libraries have been obtained from the cambial zone of young individuals of Pinus canariensis C. Sm. ex DC, a species in which such cells are comparatively abundant. These two libraries cover earlywood (EW) and latewood (LW) differentiation, and reads have been de novo meta-assembled into one transcriptome. A high number of previously undescribed genes have been found. The transcriptional profiles during the growing season have been analyzed and several noticeable differences with respect to previous studies have been found. This work contributes to a more complete picture of wood formation in conifers. The genes and their transcription profiles described here provide a useful molecular tool for further studies focused on relevant developmental issues, such as wound response and the formation of traumatic wood, re-sprouting, etc., presumably related to those cells.
BackgroundWounds caused by fire, herbivorism, rock impacts, etc. cause the direct loss of photosynthetic, storage and/or vascular tissue. In addition, they may entail other damages, such as desiccation of the exposed internal parts, or become a gateway to infection by fungi and other pathogens. To successfully overcome such injuries, plants must reorganize their meristems or even differentiate new ones, producing new traumatic tissues to cover the wound and restore the vascular connection.ResultsIn this work we analyse the anatomical growth response in conifers after debarking and injuring the vascular cambium, using Pinus canariensis as model species, due to its high wound recovery ability. Conversely to angiosperm woody species, this process is initiated and largely driven by the damaged vascular cambium and not by proliferation in the wound surface. We have detected alterations and switches in the divisions of cambial cells, associated to their position relative to the surface and edges of the wound, resulting in disordered traumatic xylem. We also describe the formation of column-like structures, after girdling, which are in part formed by the proliferation of xylem parenchymatous cells, associated to axial resin ducts.ConclusionsAbundant resinosis on the wound surface, typical of conifers, is an efficient barrier against opportunistic fungi, insects, etc. but it also hinders the healing process directly from the surface. Thus, wound closure must be largely carried out from the wound margins, being a much slower process, which very often remains unconcluded for long years. This work also describes for the first time the proliferation of inner parenchymatous cells to form column-like structures, which accelerates wound closure in girdled P. canariensis. Irregularities in the surface of the healing edge or column-like structures result in the production of disordered vascular tissues, compromising their future functionality, and which must be overcome through the fast restoration of the proper polarity in vascular cambium.
BackgroundWoody plants, especially trees, usually must face several injuries caused by different agents during their lives. Healing of injuries in stem and branches, affecting the vascular cambium and xylem can take several years. In conifers, healing takes place mainly from the remaining vascular cambium in the margin of the wound. The woundwood formed in conifers during healing usually presents malformed and disordered tracheids as well as abundant traumatic resin ducts. These characteristics affect its functionality as water conductor and its technological properties.ResultsIn this work we analyze for the first time the transcriptomic basis of the formation of traumatic wood in conifers, and reveal some differences with normal early- and late-wood. Microarray analysis of the differentiating traumatic wood, confirmed by quantitative RT-PCR, has revealed alterations in the transcription profile of up to 1408 genes during the first period of healing. We have grouped these genes in twelve clusters, according to their transcription profiles, and have distinguished accordingly two main phases during this first healing.ConclusionsWounding induces a complete rearrangement of the transcriptional program in the cambial zone close to the injuries. At the first instance, radial growth is stopped, and a complete set of defensive genes, mostly related to biotic stress, are induced. Later on, cambial activity is restored in the lateral borders of the wound, even at a high rate. During this second stage certain genes related to early-wood formation, including genes involved in cell wall formation and transcription factors, are significantly overexpressed, while certain late-wood related genes are repressed. Additionally, significant alterations in the transcription profile of abundant non annotated genes are reported.Electronic supplementary materialThe online version of this article (10.1186/s12870-017-1183-3) contains supplementary material, which is available to authorized users.
The increase of sequencing capacity provided by high‐throughput platforms has made it possible to routinely obtain large sets of genomic and transcriptomic sequences from model and non‐model organisms. Subsequent genomic analysis and gene discovery in next‐generation sequencing experiments are, however, bottlenecked by functional annotation. One common way to perform functional annotation of sets of sequences obtained from next‐generation sequencing experiments, is by searching for homologous sequences and accessing the related functional information deposited in genomic databases. Functional annotation is especially challenging for non‐model organisms, like many plant species. In such cases, existing free and commercial general‐purpose applications may not offer complete and accurate results. We present TOA (Taxonomy‐oriented annotation), a Python‐based user‐friendly open source application designed to establish functional annotation pipelines geared towards non‐model plant species that can run in Linux/Mac computers, HPCs and cloud servers. TOA performs homology searches against proteins stored in the PLAZA databases, NCBI RefSeq Plant, Nucleotide Database and Non‐Redundant Protein Sequence Database, and outputs functional information from several ontology systems: Gene Ontology, InterPro, EC, KEGG, Mapman and MetaCyc. The software performance was validated by comparing the runtimes, total number of annotated sequences and accuracy of the functional information obtained for several plant benchmark data sets with TOA and other functional annotation solutions. TOA outperformed the other software in terms of number of annotated sequences and accuracy of the annotation and constitutes a good alternative to improve functional annotation in plants. TOA is especially recommended for gymnosperms or for low quality sequence data sets of non‐model plants.
The shoot apical meristem is responsible of seasonal length increase in plants. In woody plants transition from primary to secondary growth is also produced during seasonal apical growth. These processes are controlled by different families of transcription factors. Levels of transcriptomic activity during apical growth were measured by means of a cDNA microarray designed from sequences related to meristematic activity in Pinus canariensis. The identification of differentially expressed genes was performed using a time‐course analysis. A total of 7170 genes were differentially expressed and grouped in six clusters according to their expression profiles. We identified master regulators, such as WUSCHEL‐like HOMEOBOX (WOX), to be involved in the first stages of apical development, i.e. growth of primary tissues, while other transcription factors, such as Class III HOMEODOMAIN‐LEUCINE ZIPPER (HD‐ZIP III) and KNOTTED‐like (KNOX) and BEL1‐like (BELL) HOMEODOMAIN proteins, were found to be induced during last stages of apical seasonal development, already with secondary growth. Our results reveal the main expression patterns of these genes during apical development and the transition from primary to secondary stem growth. In particular, the regulatory factors identified play key roles in controlling stem architecture and constitute candidate genes for the study of other development processes in conifers.
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