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Summary: Branch length is an important plant architecture trait in cotton (Gossypium) breeding. Development of cultivars with short branch has been proposed as a main object to enhance cotton yield potential, because they are suitable for high planting density. Here, we report the molecular cloning and characterization of a semi‐dominant quantitative trait locus, Short Branch Internode 1(GhSBI1), which encodes a NAC transcription factor homologous to CUP‐SHAPED COTYLEDON 2 (CUC2) and is regulated by microRNA ghr‐miR164. We demonstrate that a point mutation found in sbi1 mutants perturbs ghr‐miR164‐directed regulation of GhSBI1, resulting in an increased expression level of GhSBI1. The sbi1 mutant was sensitive to exogenous gibberellic acid (GA) treatments. Overexpression of GhSBI1 inhibited branch internode elongation and led to the decreased levels of bioactive GAs. In addition, gene knockout analysis showed that GhSBI1 is required for the maintenance of the boundaries of multiple tissues in cotton. Transcriptome analysis revealed that overexpression of GhSBI1 affects the expression of plant hormone signalling‐, axillary meristems initiation‐, and abiotic stress response‐related genes. GhSBI1 interacted with GAIs, the DELLA repressors of GA signalling. GhSBI1 represses expression of GA signalling‐ and cell elongation‐related genes by directly targeting their promoters. Our work thus provides new insights into the molecular mechanisms for branch length and paves the way for the development of elite cultivars with suitable plant architecture in cotton. [ABSTRACT FROM AUTHOR]

Growth-regulating factor (GRF) is a plant-specific family of transcription factors crucial for meristem development and plant growth. Sorghum (Sorghum bicolor L. Moench) is a cereal species widely used for food, feed and fuel. While sorghum stems are important biomass components, the regulation of stem development and the carbohydrate composition of the stem tissues remain largely unknown. Here, we identified 11 SbGRF-encoding genes and found the SbGRF expansion driven by whole-genome duplication events. By comparative analyses of GRFs between rice and sorghum, we demonstrated the divergence of whole-genome duplication (WGD)-derived OsGRFs and SbGRFs. A comparison of SbGRFs' expression profiles supports that the WGD-duplicated OsGRFs and SbGRFs experienced distinct evolutionary trajectories, possibly leading to diverged functions. RNA-seq analysis of the internode tissues identified several SbGRFs involved in internode elongation, maturation and cell wall metabolism. We constructed co-expression networks with the RNA-seq data of sorghum internodes. Network analysis discovered that SbGRF1, 5 and 7 could be involved in the down-regulation of the biosynthesis of cell wall components, while SbGRF4, 6, 8 and 9 could be associated with the regulation of cell wall loosening, reassembly and/or starch biosynthesis. In summary, our genome-wide analysis of SbGRFs reveals the distinct evolutionary trajectories of WGD-derived SbGRF pairs. Importantly, expression analyses highlight previously unknown functions of several SbGRFs in internode elongation, maturation and the potential involvement in the metabolism of the cell wall and starch during post-anthesis stages. [ABSTRACT FROM AUTHOR]

Dwarfing is a pivotal agronomic trait affecting both yield and quality. Citrus species exhibit substantial variation in plant height, among which internode length is a core element. However, the molecular mechanism governing internode elongation remains unclear. Here, we unveiled that the transcriptional cascade consisting of B‐BOX DOMAIN PROTEIN 22 (BBX22) and ELONGATED HYPOCOTYL 5 (HY5) finely tunes plant height and internode elongation in citrus. Loss‐of‐function mutations of BBX22 in an early‐flowering citrus (Citrus hindsii "SJG") promoted internode elongation and reduced pigment accumulation, whereas ectopic expression of BBX22 in SJG, sweet orange (C. sinensis), pomelo (C. maxima) or heterologous expression of BBX22 in tomato (Solanum lycopersicum) significantly decreased internode length. Furthermore, exogenous application of gibberellin A3 (GA3) rescued the shortened internode and dwarf phenotype caused by BBX22 overexpression. Additional experiments revealed that BBX22 played a dual role in regulation internode elongation and pigmentation in citrus. On the one hand, it directly bound to and activated the expression of HY5, GA metabolism gene (GA2 OXIDASE 8, GA2ox8), carotenoid biosynthesis gene (PHYTOENE SYNTHASE 1, PSY1) and anthocyanin regulatory gene (Ruby1, a MYB DOMAIN PROTEIN). On the other hand, it acted as a cofactor of HY5, enhancing the ability of HY5 to regulate target genes expression. Together, our results reveal the critical role of the transcriptional cascade consisting of BBX22 and HY5 in controlling internode elongation and pigment accumulation in citrus. Unraveling the crosstalk regulatory mechanism between internode elongation and fruit pigmentation provides key genes for breeding of novel types with both dwarf and health‐beneficial fortification in citrus. [ABSTRACT FROM AUTHOR]

Complete submergence, especially deep submergence, poses a serious threat to the growth and survival of plants. One study previously showed that Alternanthera philoxeroides (a herbaceous perennial plant) submerged at depth of 2 m presented fast stem elongation and reduced stem elongation as water depth increased. In the present study, we aimed to figure out from the morphological and anatomical perspective how the differential growth response of the plant to water depth was achieved. We investigated the elongation of different stem parts and the relationship of stem elongation to cell size and number in A. philoxeroides by conducting experiments using a series of submergence depths (0 m, 2 m, 5 m, and 9 m). The results showed that, in comparison with unsubmerged plants, completely submerged plants exhibited enhanced elongation at depths of 2 m and 5 m but suppressed elongation at depth of 9 m in immature stem internodes, and displayed very little elongation in mature stem internodes at any depths. The stem growth of A. philoxeroides at any submergence depth was chiefly caused by the elongation of the basal parts of immature internodes. The elongation of the basal parts of immature internodes was highly correlated to both cell proliferation and cell enlargement, but the elongation of the middle and upper parts of immature internodes correlated nearly only with cell enlargement. This study provided new information on the growth responses of A. philoxeroides to heterogeneous submergence environments and deepened our understanding of the growth performance of terrestrial plants in habitats prone to deep floods. [ABSTRACT FROM AUTHOR]

Autumn grazing management aims to accumulate herbage for defoliation prior to a decrease in growth rates for the extension of the grazing season. The current study investigated the impact of building different target herbage masses (THMs) in autumn and imposing one of three different defoliation dates (DDs) between mid-October and late November on light transmitted to the base of the sward, free leaf lamina (FLL), leaf stage and internode elongation. Four THMs (low, medium, high and very high) and three DDs (DD1—15 October, DD2—7 November and DD3—21 November) were assigned to a 4 × 3 split plot design over two years. Light transmitted to the base of the sward was greatest in the low THM and decreased in all other THMs. Internode elongation increased in tillers in the medium to the very high THMs. Defoliation of the medium, high and, in particular, the very high THMs earlier in autumn reduced the effect of decreased light transmission on internode elongation. This study highlights that, as light transmitted to the base of the sward decreases, internode elongation increases, and this could negatively impact sward structure. [ABSTRACT FROM AUTHOR]

Drought stress impairs plant growth and other physiological functions. MeHDZ14, a homeodomain-leucine zipper I transcription factor, is strongly induced by drought stress in various cassava cultivars. However, the role of MeHDZ14 in cassava growth regulation has remained unclear. Here we report that MeHDZ14 affected plant height, such that a dwarf phenotype and altered internode elongation were observed in transgenic cassava lines. MeHDZ14 was found to negatively regulate the biosynthesis of lignin. Its overexpression resulted in abaxially rolled leaves. The morphogenesis of leaf epidermal cells was inhibited by overexpression of MeHDZ14, with decreased auxin and gibberellin and increased cytokinin contents. MeHDZ14 was found to regulate many drought-responsive genes, including genes involved in cell wall synthesis and expansion. MeHDZ14 bound to the promoter of caffeic acid 3-O-methyltransferase 1 (MeCOMT1), acting as a transcriptional repressor of genes involved in cell wall development. MeHDZ14 appears to act as a negative regulator of internode elongation and epidermal cell morphogenesis during cassava leaf development.

Growth-regulating factor (GRF) is a plant-specific family of transcription factors crucial for meristem development and plant growth. Sorghum (Sorghum bicolor L. Moench) is a cereal species widely used for food, feed and fuel. While sorghum stems are important biomass components, the regulation of stem development and the carbohydrate composition of the stem tissues remain largely unknown. Here, we identified 11 SbGRF-encoding genes and found the SbGRF expansion driven by whole-genome duplication events. By comparative analyses of GRFs between rice and sorghum, we demonstrated the divergence of whole-genome duplication (WGD)-derived OsGRFs and SbGRFs. A comparison of SbGRFs’ expression profiles supports that the WGD-duplicated OsGRFs and SbGRFs experienced distinct evolutionary trajectories, possibly leading to diverged functions. RNA-seq analysis of the internode tissues identified several SbGRFs involved in internode elongation, maturation and cell wall metabolism. We constructed co-expression networks with the RNA-seq data of sorghum internodes. Network analysis discovered that SbGRF1, 5 and 7 could be involved in the down-regulation of the biosynthesis of cell wall components, while SbGRF4, 6, 8 and 9 could be associated with the regulation of cell wall loosening, reassembly and/or starch biosynthesis. In summary, our genome-wide analysis of SbGRFs reveals the distinct evolutionary trajectories of WGD-derived SbGRF pairs. Importantly, expression analyses highlight previously unknown functions of several SbGRFs in internode elongation, maturation and the potential involvement in the metabolism of the cell wall and starch during post-anthesis stages.

Context: Tall bunch-type tropical forage grasses are known for their rapid (true) stem elongation late in the regrowth cycle, even during the vegetative phase. Aims: This study aimed to evaluate the stem and pseudostem growth pattern of 'Mombaça' guineagrass [ Megathyrsus maximus (Jacq.) BK Simon & SWL Jacobs] grown in field conditions and how this relates to its high biomass accumulation during long regrowth cycles. Methods: A guineagrass field was managed from December 2017 to January 2019, in three regrowth cycles of 10, 14 and 12 weeks duration. Individual tillers were assessed weekly to measure stem, pseudostem and leaf elongation, leaf appearance; and angle of insertion of the leaves. Aboveground biomass samples were taken weekly for biomass accumulation and leaf area index assessment. Key results: At the beginning of each regrowth cycle, the pseudostem elongated while the stem length remained constant. Subsequently, the pseudostem length reached a plateau, while the stem length increased at a constant rate. Because of a positive relationship between pseudostem and leaf laminae lengths, the long pseudostem ensured the formation of long leaves in tall tillers and positive net leaf elongation in long regrowth cycles. Conclusions: The high biomass accumulation reflected the continuous positive net leaf elongation by the tillers and was underpinned by the younger leaves being more erect than the older ones, allowing for lower self-shading of the older leaves and greater leaf tissue retention. Implications: The high production of tropical forage grasses in late regrowth should be utilised with caution, as it is underpinned by stem elongation and meristem elevation. Tropical grasses are used as forage for ruminants and for biofuel production from biomass. Stem elongation improves light distribution in the canopy and photosynthetic rates of tropical grasses. Nevertheless, stem has lower nutritive value than leaves, and frequently animals are not able to harvest it. The aim of this study was to understand the relationship between stem elongation and biomass production of tropical grasses. Our results may help improve animal protein and biofuel production from tropical grasses. [ABSTRACT FROM AUTHOR]

Autumn grazing management aims to accumulate herbage for defoliation prior to a decrease in growth rates for the extension of the grazing season. The current study investigated the impact of building different target herbage masses (THMs) in autumn and imposing one of three different defoliation dates (DDs) between mid-October and late November on light transmitted to the base of the sward, free leaf lamina (FLL), leaf stage and internode elongation. Four THMs (low, medium, high and very high) and three DDs (DD1—15 October, DD2—7 November and DD3—21 November) were assigned to a 4 × 3 split plot design over two years. Light transmitted to the base of the sward was greatest in the low THM and decreased in all other THMs. Internode elongation increased in tillers in the medium to the very high THMs. Defoliation of the medium, high and, in particular, the very high THMs earlier in autumn reduced the effect of decreased light transmission on internode elongation. This study highlights that, as light transmitted to the base of the sward decreases, internode elongation increases, and this could negatively impact sward structure.

Rice plants that form ventilated tissues, such as aerenchyma in the leaves, stems, and roots, allow for growth in waterlogged conditions (paddy fields), but they cannot breathe and drown in flooded environments where the whole plant body is submerged. However, deepwater rice plants grown in flood-prone areas of Southeast Asia survive in prolonged flooded environments by taking in air through an elongated stem (internode) and leaves that emerge above the water surface, even if the water level is several meters high and flooding continues for several months. Although it has been known that plant hormones, such as ethylene and gibberellins, promote internode elongation in deepwater rice plants, the genes that control rapid internode elongation during submergence have not been identified. We recently identified several genes responsible for the quantitative trait loci involved in internode elongation in deepwater rice. Identification of the the genes revealed a molecular gene network from ethylene to gibberellins in which internode elongation is promoted by novel ethylene-responsive factors and enhances gibberellin responsiveness at the internode. In addition, elucidation of the molecular mechanism of internode elongation in deepwater rice will help our understanding of the internode elongation mechanism in normal paddy rice and contribute to improving crops through the regulation of internode elongation. [ABSTRACT FROM AUTHOR]

Xylogen-like arabinogalactan protein (XYLP) is an atypical lipid transport protein. In this study, 23 Phyllostachys edulis XYLPs were identified, and their proteins contain characteristic structures of AGP and nsLTP domain. All PeXYLPs can be divided into four clades, and their genes were unevenly distributed on 11 chromosome scaffolds. Collinear analysis revealed that segmental duplication was the main driver for PeXYLP family expansion. The c is-acting elements presented in the promoter are involved in various regulations of PeXYLPs expression. G.O. annotation revealed that PeXYLPs are mainly interested in lipid transport and synthesis and primarily function at the plasma membrane. Transcriptome analysis revealed that PeXYLPs were spatiotemporally expressed and displayed significant variability during various tissue development. Besides that, some PeXYLP s also respond to multiple phytohormones and abiotic stresses. By semi-quantitative RT-PCR, the response of some PeXYLPs to MeJA was confirmed, and the proteins were shown to localize to the plasma membrane mainly. WGCNA in defined regions of fast-growing bamboo shoots revealed that 5 PeXYLPs in 4 gene co-expression modules showed a positive module-trait relationship with three fast-growing regions. This systematic analysis of the PeXYLP family will provide a foundation for further insight into the functions of individual PeXYLP in a specific tissue or organ development, phytohormone perception, and stress responses in the future. • Twenty-three XYLP genes (PeXYLPs) were identified in the genome of Phyllostachys edulis and were classified into four evolutionary clades. • Segmental duplication plays a dominant role in expanding the PeXYLP family in Phyllostachys edulis. • Large-scale expansion of the XYLP family could have occurred after the segregation of monocotyledons and dicotyledons and then expanded independently. • Expression of some PeXYLP genes showed positive correlations with fast growth and elongation of internode in Phyllostachys edulis. [ABSTRACT FROM AUTHOR]

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Main conclusion: CG and CHG methylation levels in the rapid shoot growth stages (ST2–ST4) of woody bamboos were obviously decreased, which might regulate the internode elongation during rapid shoot growth, while CHH methylation was strongly associated with shoot developmental time or age. DNA methylation plays a critical role in the regulation of plant growth and development. Woody bamboos have a unique trait of rapid stem growth resulted from internode elongation at the shooting period. However, it is still unclear whether DNA methylation significantly controls the bamboo rapid stem growth. Here we present whole-genome DNA methylation profiles of the paleotropical woody bamboo Bonia amplexicaulis at five newly defined stages of shoot growth, named ST1–ST5. We found that CG and CHG methylation levels in the rapid shoot growth stages (ST2–ST4) were significantly lower than in the incubation (ST1) and plateau stages (ST5). The changes in methylation levels mainly occurred in flanking regions of genes and gene body regions, and 23647 differentially methylated regions (DMRs) were identified between ST1 and rapid shoot growth stages (ST2–ST4). Combined with transcriptome analysis, we found that DMR-related genes enriched in the auxin and jasmonic acid (JA) signal transduction, and other pathways closely related to plant growth. Intriguingly, CHH methylation was not involved in the rapid shoot growth, but strongly associated with shoot developmental time by gradually accumulating in transposable elements (TEs) regions. Overall, our results reveal the importance of DNA methylation in regulating the bamboo rapid shoot growth and suggest a role of DNA methylation associated with development time or age in woody bamboos. [ABSTRACT FROM AUTHOR]

The present study carried out with a field plot experiment and light environment simulation experiment in 2017 to assess the effect of spectral distribution on internode elongation in soybean canopy. Henong 60 and Heinong 48 were used as the experimental materials. The spectral distribution in the soybean canopy was studied under four planting densities 200,000(D20), 300,000(D30), 400,000(D40) and 500,000(D50) plant ha−1. Meanwhile, a pot experiment of light environment simulation with light-emitting diode (LED) was carried out to further analyze the effects of light quantity and quality on the elongation of soybean seedlings. The results showed that the intensity of PAR, blue light, red light and far-red light in soybean canopy decreased by varying degrees with density and canopy depth increasing, and the decrease of PAR and R/FR led to significant elongation of internodes and plant height in high density. The spectral intensity in the middle and base of the canopy was strongly reduced, resulting in the strongest internode elongation in the middle of soybean stem. The increase in soybean plant height was also mainly due to the internode elongation in the middle at high density. In addition, the mono-light of far-red light could promote the internode elongation, while the red light and blue light acted as inhibitors and the inhibitory effect of blue light was even greater.

The non-structural carbohydrate (NSC) stored in rice stems (culm plus leaf sheaths) at pre-heading stage is one of the major carbon sources for grain filling. In this study, we conducted field experiments in 2018 and 2019 using two varieties, Momiroman (japonica-dominant) and Teqing (indica), which have different accumulation and remobilization patterns of NSC in the stems. We analyzed NSC dynamics of each of the five nodal segments of internodes and leaf sheaths at pre- and post-heading stages. The dynamics of NSC at pre- and post-heading stages differed among segments. The lower segments of internodes and leaf sheaths accumulated more NSC at pre-heading stage compared to the upper segments in both varieties. The lower internodes in Teqing were morphologically larger than in Momiroman and would be able to store more NSC. However, the upper parts of the stems required biomass for development and formation at the stage of NSC accumulation in stems. Momiroman had longer upper internodes and leaf sheaths and required more biomass than Teqing. The NSC remobilization was later in the upper than the lower internodes for both varieties, which led to the delay of NSC remobilization in the entire culm after heading. These results indicated that ideal morphology of stems, to maximize the potential contribution of stem NSC as the carbon source for grain filling, is large lower internodes and short upper internodes and leaf sheaths.

The adaptation and survival of boreal and temperate perennials relies on the precise demarcation of the growing season. Seasonal growth and development are defined by day length and temperature signals. Under long-day conditions in spring, poplar FLOWERING LOCUS T2 (FT2) systemically induces shoot growth. In contrast, FT2 downregulation induced by autumnal short days triggers growth cessation and bud set. However, the molecular role of FT2 in local and long-range signaling is not entirely understood. In this study, the CRISPR/Cas9 editing tool was used to generate FT2 loss of function lines of hybrid poplar. Results indicate that FT2 is essential to promote shoot apex development and restrict internode elongation under conditions of long days. The application of bioactive gibberellins (GAs) to apical buds in FT2 loss of function lines was able to rescue bud set. Expression analysis of GA sensing and metabolic genes and hormone quantification revealed that FT2 boosts the 13-hydroxylation branch of the GA biosynthesis pathway in the shoot apex. Paclobutrazol treatment of WT leaves led to limited internode growth in the stem elongation zone. In mature leaves, FT2 was found to control the GA 13-hydroxylation pathway by increasing GA2ox1 and reducing GA3ox2 expression, causing reduced GA1 levels. We here show that in poplar, the FT2 signal promotes shoot apex development and restricts internode elongation through the GA 13-hydroxylation pathway. [ABSTRACT FROM AUTHOR]

The non-structural carbohydrate (NSC) stored in rice stems (culm plus leaf sheaths) at pre-heading stage is one of the major carbon sources for grain filling. In this study, we conducted field experiments in 2018 and 2019 using two varieties, Momiroman (japonica-dominant) and Teqing (indica), which have different accumulation and remobilization patterns of NSC in the stems. We analyzed NSC dynamics of each of the five nodal segments of internodes and leaf sheaths at pre- and post-heading stages. The dynamics of NSC at pre- and post-heading stages differed among segments. The lower segments of internodes and leaf sheaths accumulated more NSC at pre-heading stage compared to the upper segments in both varieties. The lower internodes in Teqing were morphologically larger than in Momiroman and would be able to store more NSC. However, the upper parts of the stems required biomass for development and formation at the stage of NSC accumulation in stems. Momiroman had longer upper internodes and leaf sheaths and required more biomass than Teqing. The NSC remobilization was later in the upper than the lower internodes for both varieties, which led to the delay of NSC remobilization in the entire culm after heading. These results indicated that ideal morphology of stems, to maximize the potential contribution of stem NSC as the carbon source for grain filling, is large lower internodes and short upper internodes and leaf sheaths. [ABSTRACT FROM AUTHOR]

The present study carried out with a field plot experiment and light environment simulation experiment in 2017 to assess the effect of spectral distribution on internode elongation in soybean canopy. Henong 60 and Heinong 48 were used as the experimental materials. The spectral distribution in the soybean canopy was studied under four planting densities 200,000(D20), 300,000(D30), 400,000(D40) and 500,000(D50) plant ha−1. Meanwhile, a pot experiment of light environment simulation with light-emitting diode (LED) was carried out to further analyze the effects of light quantity and quality on the elongation of soybean seedlings. The results showed that the intensity of PAR, blue light, red light and far-red light in soybean canopy decreased by varying degrees with density and canopy depth increasing, and the decrease of PAR and R/FR led to significant elongation of internodes and plant height in high density. The spectral intensity in the middle and base of the canopy was strongly reduced, resulting in the strongest internode elongation in the middle of soybean stem. The increase in soybean plant height was also mainly due to the internode elongation in the middle at high density. In addition, the mono-light of far-red light could promote the internode elongation, while the red light and blue light acted as inhibitors and the inhibitory effect of blue light was even greater. [ABSTRACT FROM AUTHOR]

As a member of the small GTPases family, Rab GTPases play a key role in specifying transport pathways in the intracellular membrane trafficking system and are involved in plant growth and development. By quantitative trait locus (QTL) mapping, PdRabG3f was identified as a candidate gene associated with shoot height in a hybrid offspring of Populus deltoides 'Danhong' × Populus simonii 'Tongliao1'. PdRabG3f localized to the nucleus, endoplasmic reticulum and tonoplast and was primarily expressed in the xylem and cambium. Overexpression of PdRabG3f in Populus alba × Populus glandulosa (84 K poplar) had inhibitory effects on vertical and radical growth. In the transgenic lines, there were evident changes in the levels of 15 gibberellin (GA) derivatives, and the application of exogenous GA 3 partially restored the phenotypes mediated by GAs deficiency. The interaction between PdRabG3f and RIC4, which was the GA-responsive factor, provided additional explanation for PdRabG3f's inhibitory effect on poplar growth. RNA-seq analysis revealed differentially expressed genes (DEGs) associated with cell wall, xylem, and gibberellin response. PdRabG3f interfering endogenous GAs levels in poplar might involve the participation of MYBs and ultimately affected internode elongation and xylem development. This study provides a potential mechanism for gibberellin-mediated regulation of plant growth through Rab GTPases. • PdRabG3f inhibitd the vertical elongation of poplar by suppressing internode elongation. • PdRabG3f inhibited xylem development by suppressing cambium division and differentiation. • PdRabG3f interfered with gibberellin-mediated internode elongation and xylem developing in poplar. [ABSTRACT FROM AUTHOR]

Abstract Background Short internodes contribute to plant dwarfism, which is exceedingly beneficial for crop production. However, the underlying mechanisms of internode elongation are complicated and have been not fully understood. Results Here, we report a maize dwarf mutant, dwarf2014 (d2014), which displays shortened lower internodes. Map-based cloning revealed that the d2014 gene is a novel br2 allele with a splicing variation, resulting in a higher expression of BR2-T02 instead of normal BR2-T01. Then, we found that the internode elongation in d2014/br2 exhibited a pattern of inhibition-normality-inhibition (transient for the ear-internode), correspondingly, at the 6-leaf, 12-leaf and 14-leaf stages. Indeed, BR2 encodes a P-glycoprotein1 (PGP1) protein that functions in auxin efflux, and our in situ hybridization assay showed that BR2 was mainly expressed in vascular bundles of the node and internode. Furthermore, significantly higher auxin concentration was detected in the stem apex of d2014 at the 6-leaf stage and strictly in the node region for the ear-internode at the 14-leaf stage. In such context, we propose that BR2/PGP1 transports auxin from node to internode through the vascular bundles, and excessive auxin accumulation in the node (immediately next to the intercalary meristem) region suppresses internode elongation of d2014. Conclusions These findings suggest that low auxin levels mediated by BR2/PGP1 in the intercalary meristem region are crucial for internode elongation.

Abstract Background Appropriate plant architecture can improve the amount of cotton boll opening and allow increased planting density, thus increasing the level of cotton mechanical harvesting and cotton yields. The internodes of cotton fruiting branches are an important part of cotton plant architecture. Thus, studying the molecular mechanism of internode elongation in cotton fruiting branches is highly important. Results In this study, we selected internodes of cotton fruiting branches at three different stages from two cultivars whose internode lengths differed significantly. A total of 76,331 genes were detected by transcriptome sequencing. By KEGG pathway analysis, we found that DEGs were significantly enriched in the plant hormone signal transduction pathway. The transcriptional data and qRT-PCR results showed that members of the GH3 gene family, which are involved in auxin signal transduction, and CKX enzymes, which can reduce the level of CKs, were highly expressed in the cultivar XLZ77, which has relatively short internodes. Genes related to ethylene synthase (ACS), EIN2/3 and ERF in the ethylene signal transduction pathway and genes related to JAR1, COI1 and MYC2 in the JA signal transduction pathway were also highly expressed in XLZ77. Plant hormone determination results showed that the IAA and CK contents significantly decreased in cultivar XLZ77 compared with those in cultivar L28, while the ACC (the precursor of ethylene) and JA contents significantly increased. GO enrichment analysis revealed that the GO categories associated with promoting cell elongation, such as cell division, the cell cycle process and cell wall organization, were significantly enriched, and related genes were highly expressed in L28. However, genes related to the sphingolipid metabolic process and lignin biosynthetic process, whose expression can affect cell elongation, were highly expressed in XLZ77. In addition, 2067 TFs were differentially expressed. The WRKY, ERF and bHLH TF families were the top three largest families whose members were active in the two varieties, and the expression levels of most of the genes encoding these TFs were upregulated in XLZ77. Conclusions Auxin and CK are positive regulators of internode elongation in cotton branches. In contrast, ethylene and JA may act as negative regulators of internode elongation in cotton branches. Furthermore, the WRKY, ERF and bHLH TFs were identified as important inhibitors of internode elongation in cotton. In XLZ77(a short-internode variety), the mass synthesis of ethylene and amino acid conjugation of auxin led to the inhibition of plant cell elongation, while an increase in JA content and degradation of CKs led to a slow rate of cell division, which eventually resulted in a phenotype that presented relatively short internodes on the fruiting branches. The results of this study not only provide gene resources for the genetic improvement of cotton plant architecture but also lay a foundation for improved understanding of the molecular mechanism of the internode elongation of cotton branches.

Abstract Background Internode elongation is an important agronomic trait in rice that determines culm length, which is related to lodging, panicle exsertion, and biomass. sui4 (shortened uppermost internode 4) mutants show reduced internode length and a dwarf phenotype due to shortened internodes; the uppermost internode is particularly severely affected. The present study was performed to identify the molecular nature and function of the SUI4 gene during internode elongation. Results Our previous study showed that the SUI4 gene was mapped to a 1.1-Mb interval on chromosome 7 (Ji et al. 2014). In order to isolate the gene responsible for the sui4 phenotype, genomic DNA resequencing of sui4 mutants and wild-type plants and reciprocal transformation of wild-type and mutant alleles of the putative SUI4 gene was performed. The data revealed that the causative mutation of sui4 was a T to A nucleotide substitution at the microRNA172 binding site of Os07g0235800, and that SUI4 is a new allele of the previously reported gene SUPERNUMERARY BRACT (SNB), which affects flower structure. In order to understand the effect of this mutation on expression of the SUI4/SNB gene, SUI4/SNB native promoter-fuzed GUS transgenics were examined, along with qRT-PCR analysis at various developmental stages. In sui4 mutants, the SUI4/SNB gene was upregulated in the leaves, culms, and panicles, especially when internodes were elongated. In culms, SUI4/SNB was expressed in the nodes and the lower parts of elongating internodes. In order to further explore the molecular nature of SUI4/SNB during internode elongation, RNA-seq and qRT-PCR analysis were performed with RNAs from the culms of sui4 mutants and wild-type plants in the booting stage. The data showed that in sui4 mutants, genes deactivating bioactive gibberellins and cytokinin were upregulated while genes related to cell expansion and cell wall synthesis were downregulated. Conclusion In summary, this paper shows that interaction between SUI4/SNB and microRNA172 could determine internode elongation during the reproductive stage in rice plants. Due to a mutation at the microRNA172 binding site in sui4 mutants, the expression of SUI4/SNB was enhanced, which lowered the activities of cell expansion and cell wall synthesis and consequently resulted in shortened internodes.

High crop yields are generally associated with high nitrogen (N) fertilizer rates. A growing tendency that is urgently demanding the adoption of precision technologies that manage N more efficiently, combined with the advances of crop genetics to meet the needs of sustainable farm systems. Among the plant traits, stem architecture has been of paramount importance to enhance harvest index in the cereal crops. Nonetheless, the reduced stature also brought undesirable effect, such as poor N-uptake, which has led to the overuse of N fertilizer. Therefore, a better understanding of how N signals modulate the initial and late stages of stem development might uncover novel semi-dwarf alleles without pleiotropic effects. Our attempt here is to review the most recent advances on this topic.

High crop yields are generally associated with high nitrogen (N) fertilizer rates. A growing tendency that is urgently demanding the adoption of precision technologies that manage N more efficiently, combined with the advances of crop genetics to meet the needs of sustainable farm systems. Among the plant traits, stem architecture has been of paramount importance to enhance harvest index in the cereal crops. Nonetheless, the reduced stature also brought undesirable effect, such as poor N-uptake, which has led to the overuse of N fertilizer. Therefore, a better understanding of how N signals modulate the initial and late stages of stem development might uncover novel semi-dwarf alleles without pleiotropic effects. Our attempt here is to review the most recent advances on this topic. [ABSTRACT FROM AUTHOR]

MicroRNAs are crucial regulators of gene expression in maize. However, the mechanisms through which miRNAs control internode elongation remain poorly understood. This study engineered varying levels of internode elongation inhibition, revealing that dwarfing treatments diminished gibberellin levels, curtailed cell longitudinal growth, and slowed the rate of internode elongation. Comprehensive transcriptome and miRNA profiling of the internode elongation zone showed gene expression changes that paralleled the extent of the internode length reduction. We identified 543 genes and 29 miRNAs with significant correlations to internode length, predominantly within families, including miR164 and miR396. By incorporating target gene expression levels, we pinpointed nine miRNA-mRNA pairs that are significantly associated with the regulation of the internode elongation. The inhibitory effects of these miRNAs on their target genes were confirmed through dual-luciferase reporter assays. Overexpression of miR164h in maize resulted in increased internode and cell length, suggesting a novel genetic avenue for manipulating plant stature. These miRNAs may also serve as precise spatiotemporal regulators for in vitro plant development.

The physiological function of bamboo shoot sheaths is still unclear. In the present study, we investigated the anatomical and physiological influences of bamboo shoot sheaths on internode elongation by longitudinally striping parts of sheaths. The internodes would bend toward the bare sides during night. The results showed that amounts of water leaked at the cut of shoot sheaths during night, which impeded the increase of water, water pressure and assimilate transport rates, and decreased starch and soluble sugar catabolism in the bare side of the internodes. A higher level of water pressure and sugar metabolism increased the vacuole expansion and promoted the cell expansion in the outer sides as compared to the bare sides. The bending growth of internodes was mainly due to the significant differences in cell expansion, which was led by the difference in water pressure and sugar hydrolysis levels between the inner and outer sides. Bamboo internode elongation mainly relied on the increase of water pressure and soluble sugar concentration. Shoot sheaths played an important role in the rapid growth of bamboo shoots as a controller in water and assimilate transportation. This study gave a new insight into understanding the rapid growth mechanism of bamboo plants.

SNORKEL1 (SK1) and SNORKEL2 (SK2) are ethylene responsive factors that regulate the internode elongation of deepwater rice in response to submergence. We previously reported that normal cultivated rice lacks SK genes because the Chromosome 12 region containing SK genes was deleted from its genome. However, no study has analyzed how the genome defect occurred in that region by comparing normal cultivated rice and deepwater rice. In this study, comparison of the sequence of the end of Chromosome 12, which contains SK genes, between normal and deepwater rice showed that complicated genome changes such as insertions, deletions, inversions, substitutions, and translocation occurred frequently in this region. In addition to SK1 and SK2 of deepwater rice, gene prediction analysis identified four genes containing AP2/ERF domains in normal cultivated rice and six in deepwater rice; we called these genes SK-LIKE (SKL) genes. SKs and SKLs were present in close proximity to each other, and the SKLs in normal cultivated rice were in tandem. These predicted genes belong to the same AP2/ERF subfamily and were separated into four types: SK1, SK2, SKL3, and SKL4. Sequence comparison indicated that normal cultivated rice possesses a gene with high homology to SK2, which we named SKL1. However, none of the predicted SKLs except for SKL3s were expressed during submergence. Although SKL3s were expressed in both normal and deepwater rice, normal rice does not undergo internode elongation, suggesting that its expression does not contribute to internode elongation. Plants overexpressing SKL1, which showed the most homology to SK2, underwent internode elongation similar to plants overexpressing SK1 and SK2 under normal growth conditions. A yeast one-hybrid assay showed that the C-end of SKL1 has transcription activity, as do the C-ends of SK1 and SK2. Our results suggested that SKLs were derived via gene duplication, but were not expressed and pseudogenized in normal cultivated rice during sequence evolution.

A study was conducted in four compartments of a polycarbonate greenhouse at Gainesville, FL, USA to investigate how a soybean (Glycine max L. Merr.) cultivar, Maverick (maturity group III, indeterminate), responded to three elevated temperatures, ELT, (day/night of 34/26 °C, 38/30 °C, and 42/34 °C) in comparison to a control growth temperature (30/22 °C). Carbon dioxide (CO2) concentration was maintained at 700 μmol mol−1 in each compartment by a processor controlled air-sampling and CO2-injection system. Three sequential experiments were conducted at different times of year (summer, autumn, and early spring) to investigate the effect of intensity, timing, and duration of ELT on soybean node number, internode elongation, mainstem length, and number of pods set per plant. At the control temperature, the soybean plants grown in the polycarbonate greenhouse were taller than field-grown plants. When plants were grown under continuous ELT applied soon after sowing or at initial flowering, the number of nodes increased with increasing ELT intensity, whereas the length of individual internodes decreased. When ELT treatment was applied during the beginning of flowering stage (R1–R2) or earlier, more nodes were produced and the length of affected internodes was decreased. When the ELT was imposed later at reproductive stage R5+ just before the beginning of seed filling, effects on node numbers and internode lengths were negligible. Short-term (10-day) duration of ELT applied at four stages from V3 to R5+ did not significantly affect final mean numbers of nodes or mean mainstem lengths. Possible mechanisms of elevated temperature effects on soybean internode elongation and node number (internode number) are discussed. Total pod numbers per plant increased linearly with mainstem node numbers and mainstem length. Furthermore, total pod numbers per plant were greatest at 34/26 °C rather than at the control temperature of 30/22 °C (and remained high at 38/30 °C). Mild increases in temperature might not threaten, but actually increase, yields of soybean in northerly zones where this crop is currently grown at slightly suboptimal temperatures. However, a sustained increase in ambient temperature would likely threaten soybean yields.

The physiological function of bamboo shoot sheaths is still unclear. In the present study, we investigated the anatomical and physiological influences of bamboo shoot sheaths on internode elongation by longitudinally striping parts of sheaths. The internodes would bend toward the bare sides during night. The results showed that amounts of water leaked at the cut of shoot sheaths during night, which impeded the increase of water, water pressure and assimilate transport rates, and decreased starch and soluble sugar catabolism in the bare side of the internodes. A higher level of water pressure and sugar metabolism increased the vacuole expansion and promoted the cell expansion in the outer sides as compared to the bare sides. The bending growth of internodes was mainly due to the significant differences in cell expansion, which was led by the difference in water pressure and sugar hydrolysis levels between the inner and outer sides. Bamboo internode elongation mainly relied on the increase of water pressure and soluble sugar concentration. Shoot sheaths played an important role in the rapid growth of bamboo shoots as a controller in water and assimilate transportation. This study gave a new insight into understanding the rapid growth mechanism of bamboo plants. [ABSTRACT FROM AUTHOR]

In traditional opinions, no radial transportation was considered to occur in the bamboo internodes but was usually considered to occur in the nodes. Few studies have involved the phloem unloading and post-phloem transport pathways in the rapid elongating bamboo shoots. Our observations indicated a symplastic pathway in phloem unloading and post-unloading pathways in the culms of Fargesiayunnanensis Hsueh et Yi , based on a 5,6-carboxyfluorescein diacetate tracing experiment. Significant lignification and suberinization in fiber and parenchyma cell walls in maturing internodes blocked the apoplastic transport. Assimilates were transported out of the vascular bundles in four directions in the inner zones but in two directions in the outer zones via the continuum of parenchyma cells. In transverse sections, assimilates were outward transported from the inner zones to the outer zones. Assimilates transport velocities varied with time, with the highest values at 0):00 h, which were affected by water transport. The assimilate transport from the adult culms to the young shoots also varied with the developmental degree of bamboo shoots, with the highest transport velocities in the rapidly elongating internodes. The localization of sucrose, glucose, starch grains and the related enzymes reconfirmed that the parenchyma cells in and around the vascular bundles constituted a symplastic pathway for the radial transport of sugars and were the main sites for sugar metabolism. The parenchyma cells functioned as the 'rays' for the radial transport in and between vascular bundles in bamboo internodes. These results systematically revealed the transport mechanism of assimilate and water in the elongating bamboo shoots. [ABSTRACT FROM AUTHOR]

As a member of the CLAVATA3 (CLV3)/EMBRYO SURROUNDING REGION-related (CLE) peptide family, tracheary element differentiation inhibitory factor (TDIF) plays crucial roles in vascular meristem maintenance by promoting cell proliferation and inhibiting xylem cell differentiation. In Populus trichocarpa , six TDIF-encoding genes are all expressed in vascular tissues, and in Arabidopsis PtTDIFpro:GUS lines, the expression driven by PtTDIF promoters were predominantly detected in stem vascular bundles, initiating leaves and leaf veins. Although exogenous application of two poplar TDIF peptides did not evidently affect the shoot growth in vitro , overexpression of PtTDIF genes in hybrid poplar severely retarded the internodal elongation by upregulating the expression of GA2ox and GA20ox genes and thus decreasing the level of endogenous gibberellins (GAs), which phenotypic defect could be rescued by exogenously applied GA3. In addition, TDIF overexpression unexpectedly induced a more complex venation pattern in poplar leaves, which was underpinned by the elevated expression of WOX4 and WOX13 genes. Upon TDIF treatment, the DR5:GUS poplar leaves revealed a higher GUS activity and in TDIF -overexpressing leaves, the transcript abundances of several PIN-FORMED (PIN) genes, especially that of PIN1 , were increased, which implied an integration of TDIF and auxin in mediating this process. Collectively, data of this work presented novel activities of TDIF involved in internode elongation and leaf vein formation, thus revealing the divergent functions of TDIF in perennial tree species from those in annual herbaceous Arabidopsis. [ABSTRACT FROM AUTHOR]

The SWR1 complex (SWR1-C) is important for the deposition of histone variant H2A.Z into chromatin to regulate gene expression. Characterization of SWR1-C subunits in Arabidopsis thaliana has revealed their role in variety of developmental processes. Oryza sativa actin related protein 6 (OsARP6) is a subunit of rice SWR1-C. Its role in rice plant development is unknown. Here, we examined the subcellular localization, expression patterns, and loss of function phenotypes for this protein and found that OsARP6 is a nuclear localized protein, and is broadly expressed. OsARP6 interacted with OsPIE1, a central ATPase subunit of rice SWR1-C. The osarp6 knockout mutants displayed pleiotropic phenotypic alterations in vegetative and reproductive traits, including semi-dwarf phenotype, lower tillers number, short leaf length, changes in spikelet morphology, and seed abortion. Microscopic thin sectioning of the top internode revealed that the dwarf phenotype of osarp6 was due to reduced number of cells rather than reduced cell length. The altered transcript level of genes involved in cell division suggested that OsARP6 affects cell cycle regulation. In addition, H2A.Z levels were reduced at the promoters and transcription start sites (TSS) of the regulated genes in osarp6 plants. Together, these results suggest that OsARP6 is involved in rice plant development, and H2A.Z deposition.

Complete submergence, especially deep submergence, poses a serious threat to the growth and survival of plants. One study previously showed that Alternanthera philoxeroides (a herbaceous perennial plant) submerged at depth of 2 m presented fast stem elongation and reduced stem elongation as water depth increased. In the present study, we aimed to figure out from the morphological and anatomical perspective how the differential growth response of the plant to water depth was achieved. We investigated the elongation of different stem parts and the relationship of stem elongation to cell size and number in A. philoxeroides by conducting experiments using a series of submergence depths (0 m, 2 m, 5 m, and 9 m). The results showed that, in comparison with unsubmerged plants, completely submerged plants exhibited enhanced elongation at depths of 2 m and 5 m but suppressed elongation at depth of 9 m in immature stem internodes, and displayed very little elongation in mature stem internodes at any depths. The stem growth of A. philoxeroides at any submergence depth was chiefly caused by the elongation of the basal parts of immature internodes. The elongation of the basal parts of immature internodes was highly correlated to both cell proliferation and cell enlargement, but the elongation of the middle and upper parts of immature internodes correlated nearly only with cell enlargement. This study provided new information on the growth responses of A. philoxeroides to heterogeneous submergence environments and deepened our understanding of the growth performance of terrestrial plants in habitats prone to deep floods., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. The author(s) declared that they were an editorial board member of Frontiers, at the time of submission. This had no impact on the peer review process and the final decision., (Copyright © 2024 Jing, Ren, Lin, Niu, Ayi, Wan, Zeng and Zhang.)

Background: Short internodes contribute to plant dwarfism, which is exceedingly beneficial for crop production. However, the underlying mechanisms of internode elongation are complicated and have been not fully understood. Results: Here, we report a maize dwarf mutant, dwarf2014 (d2014), which displays shortened lower internodes. Map-based cloning revealed that the d2014 gene is a novel br2 allele with a splicing variation, resulting in a higher expression of BR2-T02 instead of normal BR2-T01. Then, we found that the internode elongation in d2014/br2 exhibited a pattern of inhibition-normality-inhibition (transient for the ear-internode), correspondingly, at the 6-leaf, 12-leaf and 14-leaf stages. Indeed, BR2 encodes a P-glycoprotein1 (PGP1) protein that functions in auxin efflux, and our in situ hybridization assay showed that BR2 was mainly expressed in vascular bundles of the node and internode. Furthermore, significantly higher auxin concentration was detected in the stem apex of d2014 at the 6-leaf stage and strictly in the node region for the ear-internode at the 14-leaf stage. In such context, we propose that BR2/PGP1 transports auxin from node to internode through the vascular bundles, and excessive auxin accumulation in the node (immediately next to the intercalary meristem) region suppresses internode elongation of d2014. Conclusions: These findings suggest that low auxin levels mediated by BR2/PGP1 in the intercalary meristem region are crucial for internode elongation. [ABSTRACT FROM AUTHOR]

Internode elongation determines plant height, which in turn affects the cane yield in sugarcane. This study aimed to explore the mechanisms of sugarcane internode elongation regulation by plant growth regulators, gibberellic acid (GA3) and its inhibitor 1, 1-dimethylpiperidinium chloride (DPC, also called mepiquat chloride). We sprayed GA3 or DPC on the leaves at the early shoot elongation stage with water as the control. The contents of endogenous hormones and the expression of GA20 ox1, GID1, and GAI in sugarcane leaves were measured at days 3, 6, 12, and 24 after treatment. It was found that the height of sugarcane plants treated with GA3 showed a significant increase compared to the control, whereas an opposite trend was observed in the DPC-treated plants. Compared to the control, the content of gibberellins (GAs) in leaves was increased significantly, while the indole acetic acid (IAA) content showed an initial increase, then a decline, and eventually remained stable in the GA3-treated plants. The contents of cytokinins (CTK), abscisic acid (ABA), and ethylene (ETH) in leaves exhibited a significant downward trend in the GA3-treated plants compared to the control. In the DCP-treated plants, however, the levels of ABA and ETH in leaves were significantly increased, IAA and GAs significantly decreased, and CTK showed a trend of an initial decline followed by an increase. After GA3 treatment, the gene expression levels of GA20 ox1 and GID1 in sugarcane increased first then decreased, whereas the gene expression level of GAI first decreased and then stabilized. After DPC treatment, the expression of GA20 ox1 in sugarcane showed an initial decrease followed by an increase, while the expression of GID1 decreased gradually and then stabilized, and the expression of GAI showed a trend of an initial increase followed by stabilization. The results of the present study indicate that exogenous GA3 treatment stimulated the internode elongation in sugarcane mainly by increasing the endogenous levels of GAs, decreasing the levels of ABA and ETH, and regulating the expression of GA20 ox1, GID1 and GAI genes; the DPC treatment mainly increased the contents of ABA and ETH and reduced the contents of GAs and IAA, thereby inhibiting the expression of GID1, promoting the expression of GAI, and ultimately inhibiting the internode elongation in sugarcane. [ABSTRACT FROM AUTHOR]

Limited plant stem elongation is hypothesized to be the primary cause of decreased plant growth in response to long‐term heavy grazing (LHG). However, it remains largely unknown how LHG‐induced changes in stem elongation are mediated by internode traits. Accordingly, we experimentally investigated the effects of LHG (defined as twice the stocking rate recommended by the local government) on plant stem elongation and internode allometry in Leymus chinensis in Xilinhot, Inner Mongolia, China. Plant size and internode traits, including internode length (IL), internode diameter (ID), and internode number (IN), from plant individuals were measured according to their phytomer position from base to apex. The results revealed that IN and their distributions were changed in response to LHG. Although LHG negatively affected IL and ID of each L. chinensis internode from base to apex and eventually stem length, the plasticity index of IL and ID increased initially and then decreased from the lowermost to the uppermost internode in response to LHG. These contrasting responses of internode traits were result of allometry between internode traits determined by both LHG and internode position. Ultimately, IL and IN were identified to be key traits influencing stem length reduction under LHG conditions and explaining 52.51% and 47.49% of variation in stem length, respectively. Overall, our findings suggest that the limitation of L. chinensis stem elongation induced by LHG is mediated by the plasticity and allometry of internode traits. This result emphasizes the sensitivity of individual plants to LHG and is potentially valuable for optimal grassland management. [ABSTRACT FROM AUTHOR]

Bolting refers to the rapid lengthening of the plant stem, and is due to the coordinated effects of developmental and environmental factors. The shoot apical meristem differentiates into the inflorescence meristem and eventually the floral meristem, which then develops into various floral organs during the bolting period. The timing of bolting is critical for plant propagation and reproductive success because it influences seed production, species survival, and crop yield. Therefore, analyses of the causes of bolting and how it occurs may enable researchers to regulate bolting. In this review, two different bolting phenotypes are described, namely inflorescence axis elongation and stem internode elongations. Various factors, such as light, temperature, and endogenous hormones, are integrated to mediate bolting. Moreover, the mechanism underlying bolting, including signal cascades and epigenetic inheritance, are discussed, and appropriate future research directions are proposed. [ABSTRACT FROM AUTHOR]

Deepwater rice has a remarkable shoot elongation response to partial submergence. Shoot elongation to maintain air-contact enables 'snorkelling' of O2 to submerged organs. Previous research has focused on partial submergence of deepwater rice. We tested the hypothesis that leaf gas films enhance internode O2 status and stem elongation of deepwater rice when completely submerged. Diel patterns of O2 partial pressure (pO2) were measured in internodes of deepwater rice when partially or completely submerged, and with or without gas films on leaves, for the completely submerged plants. We also took measurements for paddy rice. Deepwater rice elongated during complete submergence and the shoot tops emerged. Leaf gas films improved O2 entry during the night, preventing anoxia in stems, which is of importance for elongation of the submerged shoots. Expressions of O2 deprivation inducible genes were upregulated in completely submerged plants during the night, and more so when gas films were removed from the leaves. Diel O2 dynamics showed similar patterns in paddy and deepwater rice. We demonstrated that shoot tops in air enabled 'snorkelling' and increased O2 in internodes of both rice ecotypes; however, 'snorkelling' was achieved only by rapid shoot elongation by deepwater rice, but not by paddy rice. [ABSTRACT FROM AUTHOR]

Summary: A network of environmental inputs and internal signaling controls plant growth, development and organ elongation. In particular, the growth‐promoting hormone gibberellin (GA) has been shown to play a significant role in organ elongation. The use of tomato as a model organism to study elongation presents an opportunity to study the genetic control of internode‐specific elongation in a eudicot species with a sympodial growth habit and substantial internodes that can and do respond to external stimuli. To investigate internode elongation, a mutant with an elongated hypocotyl and internodes but wild‐type petioles was identified through a forward genetic screen. In addition to stem‐specific elongation, this mutant, named tomato internode elongated ‐1 (tie‐1) is more sensitive to the GA biosynthetic inhibitor paclobutrazol and has altered levels of intermediate and bioactive GAs compared with wild‐type plants. The mutation responsible for the internode elongation phenotype was mapped to GA2oxidase 7, a class III GA 2‐oxidase in the GA biosynthetic pathway, through a bulked segregant analysis and bioinformatic pipeline, and confirmed by transgenic complementation. Furthermore, bacterially expressed recombinant TIE protein was shown to have bona fide GA 2‐oxidase activity. These results define a critical role for this gene in internode elongation and are significant because they further the understanding of the role of GA biosynthetic genes in organ‐specific elongation. Significance Statement: Gibberellins (GAs) are a class of phytohormones with a significant role in organ elongation. A tomato mutant with elongated internodes but wild‐type petioles was identified from a forward screen and found to have a disruption in a class III GA 2‐oxidase, a gene in the GA biosynthetic pathway, and concomitant changes to GA catabolism. [ABSTRACT FROM AUTHOR]

Plant growth regulators (PGRs) are commonly used in cereal cropping systems to restrict plant height and control lodging. Ethephon has been reported to shorten internodes and increase grain yield of maize. To analyze the transcriptomic profiles of maize internode elongation following ethephon treatment, differentially expressed genes were compared between the treatment and control samples of inbred line Zong 31 using the Affymetrix Maize Genome Array. According to the microarray data, 326 probe sets showed significant change in expression. Further research revealed that the most remarkable effects of ethephon on maize internodes elongation occurred during a 48 h period, when 89 differentially expressed genes were detected. There were dramatic change in transcript levels at 24 h and six Auxin transport genes and four gibberellin biosynthesis pathway genes were differentially expressed in Zong 31 in response to ethephon treatment. In summary, we showed that gaseous ethylene release is involved in internode meristem cell elongation through the regulation of plant hormone signaling in maize. This work provides a platform for studies in which candidate genes will be functionally tested for involvement in internode elongation.

Internode elongation is one of the key agronomic traits determining a plant’s height and biomass. However, our understanding of the molecular mechanisms controlling internode elongation is still limited in crop plant species. Here, we report the functional identification of an atypical basic helix-loop-helix transcription factor (OsbHLH073) through gain-of-function studies using overexpression (OsbHLH073-OX) and activation tagging (osbhlh073-D) lines of rice. The expression of OsbHLH073 was significantly increased in the osbhlh073-D line. The phenotype of osbhlh073-D showed semi-dwarfism due to deficient elongation of the first internode and poor panicle exsertion. Transgenic lines overexpressing OsbHLH073 confirmed the phenotype of the osbhlh073-D line. Exogenous gibberellic acid (GA3) treatment recovered the semi-dwarf phenotype of osbhlh073-D plants at the seedling stage. In addition, quantitative expression analysis of genes involving in GA biosynthetic and signaling pathway revealed that the transcripts of rice ent-kaurene oxidases 1 and 2 (OsKO1 and OsKO2) encoding the GA biosynthetic enzyme were significantly downregulated in osbhlh073-D and OsbHLH073-OX lines. Yeast two-hybrid and localization assays showed that the OsbHLH073 protein is a nuclear localized-transcriptional activator. We report that OsbHLH073 participates in regulating plant height, internode elongation, and panicle exsertion by regulating GA biosynthesis associated with the OsKO1 and OsKO2 genes.

In many grain crops, the length of internodes below ears is related to lodging resistance in the field. To clarify the relationship between internode morphological differentiation and internode proteins during primary elongation stages in maize (Zea mays L.), we used proteomics analysis to explore factors regulating internodes in eight elite inbred maize lines: Zong3, Yu87-1, Xun9058, Xun928, Chang7-2, Zheng58, P2, and A50—the parents of four commercial hybrids in China (Yuyu22, Xundan20, Zhengdan958, and Jinsai6850). A total of 66 protein spots corresponding to 48 non-redundant proteins were identified in developing seventh to ninth leaf internodes. Of these spots, seven spots corresponding to six non-redundant proteins were related to the gibberellin (GA) pathway. Nineteen protein spots corresponding to 13 non-redundant proteins were related to the auxin (IAA) pathway, and 31 protein spots corresponding to 20 non-redundant proteins were associated with ethylene biosynthesis. A correlation analysis revealed that GA and IAA contents are negatively correlated with internode length, with the first hormone more strongly length-correlated than the second. [ABSTRACT FROM AUTHOR]

Auxin is important in regulating bamboo growth and development. Morphological and anatomical analyses of naturally growing Moso bamboo treated with auxin revealed that auxin-mediated regulation of bamboo height mainly increased internode length rather than nodes number. For individual internodes, auxin regulated internode elongation mainly by promoting cell elongation. Targeted metabolomic assays of endogenous hormones found that exogenous auxin treatment affected the dynamic balance of endogenous hormones, resulting in increased levels of gibberellin and cytokinin. Transcriptome sequencing indicated that the regulation of internode length by auxin is mainly through phytohormone signaling pathway, amino acid synthesis and protein processing pathway, and energy metabolism pathway. The transcription factors that play major regulatory roles included the ethylene response factor, WRKY and MYB transcription factor, NAC structural domain protein and basic leucine zipper protein. By constructing a gene regulatory network, we found that auxin may crosstalk with multiple plant hormones. Yeast two-hybrid and transgenic experiments demonstrated that the auxin signaling factor PheAUX/IAA34 and the jasmonic acid signaling factor PheTIFY28 do interact and regulate growth in response to phytohormones. Overall we have obtained good quality bamboo internode material for industry through the auxin treatment and have preliminarily investigated the mechanism, which will provide a reference for bamboo plant height regulation. • Auxin can promote internode elongation of moso bamboo. • Auxin regulates moso bamboo growth through multiple pathways and levels. • Auxin treatment affect the dynamic balance of endogenous phytohormones. • PheAUX/IAA34 is an important auxin signaling pathway gene. [ABSTRACT FROM AUTHOR]

Three theories of apical dominance, direct, diversion, and indirect, were proposed in the 1930s to explain how auxin synthesized in the shoot apex might inhibit axillary bud outgrowth, and thus shoot branching. The direct and diversion theories of apical dominance have been investigated in detail, and they are replaced with the current auxin transport canalization and second messenger theories, respectively. These two current theories still cannot entirely explain the phenomenon of apical dominance. Although there is ample evidence that the inhibition of bud outgrowth by auxin from the shoot apex is linked to stem elongation and highly branched auxin biosynthesis or signaling mutants are dwarf, the third theory proposed in the 1930s, the indirect theory, that explains apical dominance as auxin-induced stem growth indirectly inhibits bud outgrowth has been overlooked. The indirect theory did not propose how a growing stem might inhibit bud outgrowth. Recent discoveries indicate bud dormancy (syn. quiescence, paradormancy) in response to intrinsic and environmental factors in diverse species is linked to enhanced growth of the main shoot and reduced sugar level in the buds. Since a growing stem is a strong sink for sugars, and sugar is indispensable for shoot branching, the indirect theory of apical dominance might now be explained as auxin-induced stem growth inhibits bud outgrowth by diverting sugars away from buds. Detailed study of the indirect theory and the effect of source–sink status on dormancy and outgrowth of axillary buds will advance our knowledge of apical dominance and shoot branching in plants.

MicroRNAs (miRNAs), the post-transcriptional gene regulators, are known to play an important role in plant development. The identification of differentially expressed miRNAs could better help us understand the post-transcriptional regulation that occurs during maize internode elongation. Accordingly, we compared the expression of MIRNAs between fixed internode and elongation internode samples and classified six differentially expressed MIRNAs as internode elongation-responsive miRNAs including zma-MIR160c, zma-MIR164b, zma-MIR164c, zma-MIR168a, zma-MIR396f, and zma-MIR398b, which target mRNAs supported by transcriptome sequencing. Functional enrichment analysis for predictive target genes showed that these miRNAs were involved in the development of internode elongation by regulating the genes respond to hormone signaling. To further reveal how miRNA affects internode elongation by affecting target genes, the miRNA−mRNA−PPI (protein and protein interaction) network was constructed to summarize the interaction of miRNAs and these target genes. Our results indicate that miRNAs regulate internode elongation in maize by targeting genes related to cell expansion, cell wall synthesis, transcription, and regulatory factors.