TEOSINTE BRANCHED1/CYCLOIDEA/PCF Transcription Factors Research Articles

TCP transcription factor identification in pecan (carya illinoensis) and salt tolerance function analysis of CiTCP8

The transcription factor TEOSINTE BRANCHED1/CYCLOIDEA/PCF(TCP) is a tiny family that is exclusive to plants, which is crucial for both abiotic stress and plant growth and development. Many different types of plants have been found to contain TCP transcription factors. Carya illinoensis is an important economic forest species, but the function of TCP transcription factors in pecan has not been reported so far. Based on the highly conserved TCP (PF03634) domain, we identified 35 TCP genes (CiTCPs) in pecan and divided them into three subfamilies. Analysis of conserved motifs and gene structure of the CiTCP gene family indicated that the gene family was highly conserved. Through bioinformatics and qRT-PCR analysis, we have identified a gene that may be related to salt stress, CiTCP8, which has typical transcription factor characteristics. Phenotypic analysis of CiTCP8 transgenic Arabidopsis indicates that CiTCP8 can enhance salt tolerance in plant. Overexpression of CiTCP8 in Arabidopsis enhance plant tolerance to salt stress, as determined through some physiological and biochemical indicators. In addition, the expressions of stress-related genes were also activated in CiTCP8 transgenic Arabidopsis.

Transcriptome-Wide Identification of TCP Transcription Factor Family Members in Pinus massoniana and Their Expression in Regulation of Development and in Response to Stress

Pinus massoniana is an important coniferous tree species for barren mountain afforestation with enormous ecological and economic significance. It has strong adaptability to the environment. TEOSINTE BRANCHED 1/CYCLOIDEA/PCF (TCP) transcription factors (TFs) play crucial roles in plant stress response, hormone signal transduction, and development processes. At present, TCP TFs have been widely studied in multiple plant species, but research in P. massoniana has not been carried out. In this study, 13 PmTCP TFs were identified from the transcriptomes of P. massoniana. The phylogenetic results revealed that these PmTCP members were divided into two categories: Class I and Class II. Each PmTCP TF contained a conserved TCP domain, and the conserved motif types and numbers were similar in the same subgroup. According to the transcriptional profiling analysis under drought stress conditions, it was found that seven PmTCP genes responded to drought treatment to varying degrees. The qRT-PCR results showed that the majority of PmTCP genes were significantly expressed in the needles and may play a role in the developmental stage. Meanwhile, the PmTCPs could respond to several stresses and hormone treatments at different levels, which may be important for stress resistance. In addition, PmTCP7 and PmTCP12 were nuclear localization proteins, and PmTCP7 was a transcriptional suppressor. These results will help to explore the regulatory factors related to the growth and development of P. massoniana, enhance its stress resistance, and lay the foundation for further exploration of the physiological effects on PmTCPs.

TCP Transcription Factors in Pineapple: Genome-Wide Characterization and Expression Profile Analysis during Flower and Fruit Development

TEOSINTE-BRANCHED1/CYCLOIDEA/PCF (TCP) transcription factors contain specific a basic helix–loop–helix structure, which is a significant factor in the regulation of plant growth and development. TCP has been studied in several species, but no pineapple TCP has been reported to date. Whether they are involved in the development of the flower and fruit in the pineapple remains unclear. In this study, nine non-redundant pineapple TCPs (AcTCPs) were identified. Chromosomal localization, phylogenetics, gene structure, motifs, multiple-sequence alignment, and covariance on AcTCP family members were analyzed. Analysis of promoter cis-acting elements illustrated that the AcTCP gene may be mainly co-regulated by light signal and multiple hormone signals. Analysis of expression characteristics showed a significant increase in AcTCP5 expression at 12 h after ethylene treatment, and significantly higher levels of AcTCP8 and AcTCP9 expression in the pistil than in other floral organs. Meanwhile, the AcTCP4, AcTCP5, AcTCP6, AcTCP7, and AcTCP9 expression levels were downregulated at later stages of fruit development. Transcription factors that may interact with TCP protein in the regulation of flower and fruit development are screened by the protein interaction prediction network, AcTCP5 interacts with AcSPL16, and AcTCP8 interacts with AcFT5 and AcFT6 proteins, verified by Y2H experiments. These findings provide a basis for further exploration of the molecular mechanisms and function of the AcTCP gene in flower and fruit development.

Systematic Characterization of TCP Gene Family in Four Cotton Species Revealed That GhTCP62 Regulates Branching in Arabidopsis.

Simple SummaryTCP transcription factors (TF) are indispensable for the normal functioning of plant growth and development. This study identified and performed phylogenetic analysis on 218 TCP genes in four cotton species. We also observed conserved exon-intron structures and protein motif distribution patterns in GhTCP genes. GhTCP62 was enriched in the axillary bud, indicating that it plays a role in branching. GhTCP62 is a nuclear-localized TF, the overexpression of which decreases the number of cauline-leaf branches and rosette-leaf branches in Arabidopsis. Additionally, the expression levels of HB21 and HB40 genes increased in plants with GhTCP62 overexpression, demonstrating that GhTCP62 could regulate branching by regulating HB21 and HB40. Collectively, the GhTCP62 TF located in the nucleus was highly enriched in the axillary buds, and GhTCP62 overexpression lines demonstrated fewer rosette-leaf branches and cauline-leaf branches, indicating that GhTCP62 regulates branching in Arabidopsis.TEOSINTE-BRANCHED1/CYCLOIDEA/PCF (TCP) transcription factors play an essential role in regulating various physiological and biochemical functions during plant growth. However, the function of TCP transcription factors in G. hirsutum has not yet been studied. In this study, we performed genome-wide identification and correlation analysis of the TCP transcription factor family in G. hirsutum. We identified 72 non-redundant GhTCP genes and divided them into seven subfamilies, based on phylogenetic analysis. Most GhTCP genes in the same subfamily displayed similar exon and intron structures and featured highly conserved motif structures in their subfamily. Additionally, the pattern of chromosomal distribution demonstrated that GhTCP genes were unevenly distributed on 24 out of 26 chromosomes, and that fragment replication was the main replication event of GhTCP genes. In TB1 sub-family genes, GhTCP62 was highly expressed in the axillary buds, suggesting that GhTCP62 significantly affected cotton branching. Additionally, subcellular localization results indicated that GhTCP62 is located in the nucleus and possesses typical transcription factor characteristics. The overexpression of GhTCP62 in Arabidopsis resulted in fewer rosette-leaf branches and cauline-leaf branches. Furthermore, the increased expression of HB21 and HB40 genes in Arabidopsis plants overexpressing GhTCP62 suggests that GhTCP62 may regulate branching by positively regulating HB21 and HB40.

TCP transcription factors suppress cotyledon trichomes by impeding a cell differentiation-regulating complex.

Trichomes are specialized epidermal cells that act as barriers against biotic and abiotic stresses. Although the formation of trichomes on hairy organs is well studied, the molecular mechanisms of trichome inhibition on smooth organs are still largely unknown. Here, we demonstrate that the CINCINNATA (CIN)-like TEOSINTE BRANCHED1/CYCLOIDEA/PCF (TCP) transcription factors inhibit the formation of trichomes on cotyledons in Arabidopsis (Arabidopsis thaliana). The tcp2/3/4/5/10/13/17 septuple mutant produces cotyledons with ectopic trichomes on the adaxial sides. The expression patterns of TCP genes are developmentally regulated during cotyledon development. TCP proteins directly interact with GLABRA3 (GL3), a key component of the MYB transcription factor/basic helix-loop-helix domain protein/WD40-repeat proteins (MYB-bHLH-WD40, MBW) complex essential for trichome formation, to interfere with the transactivation activity of the MBW complex in cotyledons. TCPs also disrupt the MBW complex-R3 MYB negative feedback loop by directly promoting the expression of R3 MYB genes, which enhance the repression of the MBW complex. Our findings reveal a molecular framework in which TCPs suppress trichome formation on adaxial sides of cotyledons by repressing the activity of the MBW complex at the protein level and the transcripts of R3 MYB genes at the transcriptional level.

TCP5 controls leaf margin development by regulating KNOX and BEL-like transcription factors in Arabidopsis.

Development of leaf margins is an important process in leaf morphogenesis. CIN-clade TCP (TEOSINTE BRANCHED1/CYCLOIDEA/PCF) transcription factors are known to have redundant roles in specifying leaf margins, but the specific mechanisms through which individual TCP genes function remain elusive. In this study, we report that the CIN-TCP gene TCP5 is involved in repressing the initiation and outgrowth of leaf serrations by activating two key regulators of margin development, the Class II KNOX factor KNAT3 and BEL-like SAW1. Specifically, TCP5 directly promotes the transcription of KNAT3 and indirectly activates the expression of SAW1. We also show that TCP5 regulates KNAT3 and SAW1 in a temporal- and spatial- specific manner that is largely in accordance with the progress of formation of serrations. This regulation might serve as a key mechanism in patterning margin morphogenesis and in sculpting the final form of the leaf.

Genome-wide identification of Ziziphus jujuba TCP transcription factors and their expression in response to infection with jujube witches’ broom phytoplasma

Phytoplasmas are plant pathogenic bacteria that inhabit the phloem sieve elements in infected plants and lack a cell wall. Infection of the woody perennial Ziziphus jujuba with phytoplasma causes jujube witches’ broom (JWB), a disease that causes heavy crop losses. The TEOSINTE BRANCHED 1/CYCLOIDEA/PCF (TCP) transcription factors (TFs) are plant-specific regulators of plant morphogenesis, development, and defense after phytoplasma infection. While the TCP gene family has been studied in many plants, there is no report on TCPs in jujube. In this study, 21 ZjTCP TFs were identified and bioinformatically analyzed based on the genome of the Dongzao cultivar. Another 2 ZjTCPs were found in the genome of the Junzao cultivar. Physicochemical properties of the ZjTCP proteins were quite varied, indicating possible versatility of functions. ZjTCP proteins ranged from 172 to 590 amino acids (aa), had isoelectric points (pIs) between 5.53 (ZjTCP1) and 9.81 (ZjTCP11), and molecular weights ranging from 19,279.98 (ZjTCP11) to 61,784.17 kDa (ZjTCP19). Interaction network mapping predicted several hubs, such as ZjTCP6, ZjTCP7, ZjTCP8, ZjTCP15, ZjTCP16, and ZjTCP19, among which ZjTCP6 and ZjTCP16 were predicted to function in plant defense and morphogenesis. Spatiotemporal expression analysis of the ZjTCPs showed that ten of the ZjTCP genes were detected after infection with ‘Candidatus Phytoplasma ziziphi’. ZjTCP6, ZjTCP7, ZjTCP16, and ZjTCP19 were up-regulated after phytoplasma infection. ZjTCP16 showed the most significant increase in transcript levels, after the emergence of disease symptoms. ZjTCP12, ZjTCP15, and ZjTCP18 were down-regulated after phytoplasma infection. We concluded that ZjTCP6 and ZjTCP16 were most likely regulatory factors with roles during the plant response to jujube witches’ broom.

Soybean TCP transcription factors: Evolution, classification, protein interaction and stress and hormone responsiveness

TEOSINTE-BRANCHED1/CYCLOIDEA/PCF (TCP) transcription factors, a family of plant-specific proteins, play crucial roles in plant growth and development and stress response. However, systematical information is unknown regarding the TCP gene family in soybean. In the present study, a total of 54 GmTCPs were identified in soybean, which were grouped into 11 groups with the typical TCP conserved domains. Phylogenetic relationship, protein motif and gene structure analyses distinguished the GmTCPs into two homology classes: Class I and Class II. Class II was then differentiated into two subclasses: CIN and CYC/TB1. Unique cis-element number and composition existed in the promoter regions which might be involved in the gene transcriptional regulation of different GmTCPs. Tissue expression analysis demonstrated the diverse spatiotemporal expression profiles of GmTCPs. Furthermore, the interaction protein of one previously functionally unknown TCP protein-GmTCP8 was investigated. Yeast two-hybrid assay showed the interaction between GmTCP8 and an abscisic acid receptor (GmPYL10). QRT-PCR assays indicated the distinct expression profiles of GmTCPs in response to abiotic stresses (heat, drought and salt) and stress-related signals (abscisic acid, brassinolide, salicylicacid and methyl jasmonate). These results will facilitate to uncover the possible roles of GmTCPs under abiotic stress and hormone signal responses in soybean.

Transcriptome – Scale characterization of salt responsive bean TCP transcription factors

TEOSINTE-BRANCHED1/CYCLOIDEA/PCF (TCP) proteins are important regulators of growth and developmental processes including branching, floral organ morphogenesis and leaf growth as well as stress response. This study identified 27 TCP genes of Phaseolus vulgaris (common bean), which were divided into three clusters based on phylogenetic relationship. In addition, this study showed that some of TCP genes such as Pvul-TCP-4 and Pvul-TCP-15 located on chromosomes 3 and 7, Pvul-TCP-7 and Pvul-TCP-20 located on chromosome 7 and 9, were segmentally duplicated. On the other hand, a total of 20 Pvul-TCP genes have predicted to be targeted by microRNAs (miRNA). Most of the miRNA-target genes were Pvul-TCP-1, -11, -13 and -27, which were targeted by 13, 17, 22 and 13 plant miRNAs, respectively. miR319 was one of the highly represented regulatory miRNAs to target TCP transcripts. Promoter region analysis of TCP genes resulted that the GT-1 motif, which was related to salt stress, was found in 14 different Pvul-TCP genes. Expression profiling of 10 Pvul-TCP genes based on RNA-sequencing data further confirmed with quantitative real-time RT-PCR measurements identified that Pvul-TCP genes under salt stress are expressed in a cultivar- and tissue-specific manner.

Genome-wide identification and expression analysis of TCP transcription factors in Gossypium raimondii.

Plant-specific TEOSINTE-BRANCHED1/CYCLOIDEA/PCF (TCP) transcription factors play versatile functions in multiple aspects of plant growth and development. However, no systematical study has been performed in cotton. In this study, we performed for the first time the genome-wide identification and expression analysis of the TCP transcription factor family in Gossypium raimondii. A total of 38 non-redundant cotton TCP encoding genes were identified. The TCP transcription factors were divided into eleven subgroups based on phylogenetic analysis. Most TCP genes within the same subfamily demonstrated similar exon and intron organization and the motif structures were highly conserved among the subfamilies. Additionally, the chromosomal distribution pattern revealed that TCP genes were unevenly distributed across 11 out of the 13 chromosomes; segmental duplication is a predominant duplication event for TCP genes and the major contributor to the expansion of TCP gene family in G. raimondii. Moreover, the expression profiles of TCP genes shed light on their functional divergence.

The Arabidopsis immune adaptor SRFR1 interacts with TCP transcription factors that redundantly contribute to effector‐triggered immunity

The plant immune system must be tightly controlled both positively and negatively to maintain normal plant growth and health. We previously identified SUPPRESSOR OF rps4-RLD1 (SRFR1) as a negative regulator specifically of effector-triggered immunity. SRFR1 is localized in both a cytoplasmic microsomal compartment and in the nucleus. Its TPR domain has sequence similarity to TPR domains of transcriptional repressors in other organisms, suggesting that SRFR1 may negatively regulate effector-triggered immunity via transcriptional control. We show here that excluding SRFR1 from the nucleus prevented complementation of the srfr1 phenotype. To identify transcription factors that interact with SRFR1, we screened an Arabidopsis transcription factor prey library by yeast two-hybrid assay and isolated six classI members of the TEOSINTE BRANCHED1/CYCLOIDEA/PCF (TCP) transcription factor family. Specific interactions were verified in planta. Although single or double T-DNA mutant tcp8, tcp14 or tcp15 lines were not more susceptible to bacteria expressing AvrRps4, the triple tcp8 tcp14 tcp15 mutant displayed decreased effector-triggered immunity mediated by the resistance genes RPS2, RPS4, RPS6 and RPM1. In addition, expression of PATHOGENESIS-RELATED PROTEIN2 was attenuated in srfr1-4 tcp8-1 tcp14-5 tcp15-3 plants compared to srfr1-4 plants. To date, TCP transcription factors have been implicated mostly in developmental processes. Our data indicate that one function of a subset of TCP proteins is to regulate defense gene expression in antagonism to SRFR1, and suggest a mechanism for an intimate connection between plant development and immunity.

TCP transcription factors interact with AS2 in the repression of class‐I KNOX genes in Arabidopsis thaliana

Leaf organogenesis occurs within the peripheral zone of the shoot apical meristem (SAM). The initiation and subsequent development of a leaf requires the stable repression of a highly conserved class of plant genes, namely class-I KNOTTED 1-like homeobox (KNOX) genes. In Arabidopsis, this class comprises four members: SHOOT MERISTEMLESS (STM); BREVIPEDICELLUS (BP); KNAT2 and KNAT6. Two transcription factors, ASYMMETRIC LEAVES 1 (AS1) and AS2, are known to form a protein complex to repress BP, KNAT2 and KNAT6. Here, we show that AS2 physically interacts with the microRNA319 (miR319)-regulated CINCINNATA-like TEOSINTE BRANCHED 1-CYCLOIDEA-PCF (TCP) transcription factors in vitro and in vivo. By chromatin immunoprecipitation, we demonstrated that AS2 and TCPs bind to similar regions of the BP and KNAT2 promoters. In addition, DNA-binding activities of the TCP proteins rely on the presence of AS2, as the activities were dramatically reduced in the as2 mutant. The jaw-D mutant, which overexpresses MIR319a to downregulate several target TCP genes, strongly enhanced the as2 phenotypes and caused more severe ectopic expression of BP, KNAT2 and KNAT6. Our results reveal that KNOX repression requires different types of transcription factors that function together to ensure normal leaf development.