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Chromosome location of bHLH genes in B. rapa (a) and B. oleracea (b). The symbols + and − indicate the gene located in sense or antisense strands, respectively. The genes in red are the newly identified bHLH genes in B. rapa

Chromosome location of bHLH genes in B. rapa (a) and B. oleracea (b). The symbols + and − indicate the gene located in sense or antisense strands, respectively. The genes in red are the newly identified bHLH genes in B. rapa

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Abstract Background The basic helix–loop–helix (bHLH) is the second largest gene family in the plant, some members play important roles in pistil development and response to drought, waterlogging, cold stress and salt stress. The bHLH gene family has been identified in many species, except for Brassica oleracea and B. napus thus far. This study aim...

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... bHLH genes were named based on their gene ID (Table S1, S2, S3 and S4). We constructed an unrooted cladogram with domain sequences of all bHLH genes of B. oleracea, B. rapa, B. napus and A. thaliana ( Figure S1), and branches of the bHLH genes in A. thaliana and B. rapa were labeled in red and green, respectively. We also constructed NJ phylogenetic trees by using the domain sequences of bHLH genes from B. oleracea, B. rapa, and B. napus, respectively ( Figure S2, Figure S3, Figure S4, Figure S5). ...
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... used three methods to determine the subfamily of bHLH genes in B. oleracea and B napus. The first one, according to the subfamiles of bHLH genes in A. thaliana and B. rapa, the subfamilies of newly identified bHLH genes in B. oleracea, B. rapa and B. napus were classified ( Figure S1). The second one, we classified the newly identified bHLH genes by the distance between the branches of their phylogenetic trees ( Figures S2, Fig- ure S3, Figure S4. Figure S5). ...
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... determine whether the number of exons and introns in different subfamilies is conserved, we analyze the intron-exon location of all bHLH members from three Brassica crops and integrated the results according to the NJ phylogenetic trees, as shown in Figure S10, Figure S11 and Figure S12. In B. oleracea, most members in some subfamilies had the similar gene structure. ...
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... determine whether the number of exons and introns in different subfamilies is conserved, we analyze the intron-exon location of all bHLH members from three Brassica crops and integrated the results according to the NJ phylogenetic trees, as shown in Figure S10, Figure S11 and Figure S12. In B. oleracea, most members in some subfamilies had the similar gene structure. ...
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... determine whether the number of exons and introns in different subfamilies is conserved, we analyze the intron-exon location of all bHLH members from three Brassica crops and integrated the results according to the NJ phylogenetic trees, as shown in Figure S10, Figure S11 and Figure S12. In B. oleracea, most members in some subfamilies had the similar gene structure. ...
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... B. rapa, 247 bHLH genes were distributed on 10 chromosomes (i.e., A01-A10) (Fig. 1a). For the 21 newly identified genes, BrabHLH237, BrabHLH242, and BrabHLH251 were distributed on A01 chromosomes; BrabHLH234 and BrabHLH248 were distributed on A02; and BrabHLH231 and BrabHLH243 were distributed on A03 and A06. Two genes were located on A04 and A10, three genes were distributed on A05 and A07, and four members were ...
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... distributed on A01 chromosomes; BrabHLH234 and BrabHLH248 were distributed on A02; and BrabHLH231 and BrabHLH243 were distributed on A03 and A06. Two genes were located on A04 and A10, three genes were distributed on A05 and A07, and four members were mapped on A09. A total of 222 BolbHLH genes were distributed in 9 chromosomes (i.e., C01-C09) (Fig. 1b). The location of all BolbHLH genes on the chromosome can be found in Table S1. The number of bHLH genes distributed on C06 was the least, with only 16 genes, while 35 genes were distributed on C04. A total of 33 genes were distributed on the C03 chromosome, 28 genes on C08, 25 genes on C01 and C07, 21 genes on C02 and C09, and 18 genes ...
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... this study, we predicted the signal peptides of 268, 251 and 440 bHLH proteins in B. oleracea, B. rapa, and B. napu, respectively. Only one and two members had signal peptides in B. oleracea and B. rapa, which were BolbHLH128 (Bol021805), BrabHLH084 (Bra014653), and BrabHLH168 (Bra029354, Figure S13). According to the C, S, and Y values, the site near serine may be a potential signal peptide shear site. ...
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... genes have randomly been chosen to conduct gene expression analysis after ABA and JA treatments. The leaves sampled at 0 h after treatments were used as the control and the relative expression of others samples were calculated (Table S10). All the relative expression results underwent logarithmic transformation and visualized with a heat map (Fig. 10). For ABA treatment, the expression levels of 24 genes decreased after treatment, including 13 BrabHLH genes, 4 BnabHLH genes and 7 BolbHLH genes. The expression levels of 16 genes were increased in, not in B. rapa, 10 in B. napus and 6 in B. olerace. The expression levels increased first and then decreased of 4 BrabHLH genes, 2 BnabHLH ...
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... selection [34,35]. Theoretically, after WGD and WGT events, the genome size of B. oleracea should be threefold larger than that of A. thaliana. However, the size was much smaller than the theoretical value. The retention rate of AtbHLH orthologous genes was 57% in B. oleracea, which was close to its retention rate (56%) in B. rapa, while the Fig. 10 Expression analysis of 20 BolbHLH, 20 BrabHLH and 20 BnabHLH genes under ABA and JA treatments actual retention rates in B. rapa and B. oleracea were 51.6 and 55.1%. The retention rate of BolbHLH was close to that of the core and random genes and slightly higher than that of the other gene families, such as the PMEI (52%) [35] and SDG ...

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... Ks and Ka values were utilized to evaluate the date of divergence within the BoBPC gene family, and Ka/Ks ratios were employed to calculate the rates of molecular evolution for every pair of paralogous genes. Assuming λ to be 6.96 × 10−9, the divergence time (T) was computed using the method T=Ks/2 [25]. To create graphical representations for the synteny study, TBtools' Multiple Synteny Plot function was used. ...
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... SSRs developed by the MADS box genes can represent a useful tool to detect polymorphisms inducing a hypertrophic curd induction (Treccarichi et al. 2021(Treccarichi et al. , 2023 Basic helix-loop-helix genes represent the second largest plants' family genes, which are involved in pistil development and in the abiotic and biotic stresses response and belong to a family of transcriptional regulators present in three eukaryotic kingdoms (Heim et al. 2003;Skinner et al. 2010). The study of Miao et al. (2020) allowed for the identification of 268 bHLH genes in B. oleracea, 440 genes in B. napus, and 251 genes in B. rapa, including 21 new bHLH members and the gene structures, Phylogeny threes and the conservative motif analysis were carried out showing similar expression patterns between B. rapa and B. napus in roots and contrasting ones in stems, leaves and in reproductive organs (Miao et al. 2020). Genome wide analysis of the bHLH transcription factor was studied for Chinese-cabbage showing the interaction network associated to the ArabidopsisbHLH genes and in cabbage, which revealed several chilling response patterns (Song et al. 2019;Shan et al. 2019). ...
... SSRs developed by the MADS box genes can represent a useful tool to detect polymorphisms inducing a hypertrophic curd induction (Treccarichi et al. 2021(Treccarichi et al. , 2023 Basic helix-loop-helix genes represent the second largest plants' family genes, which are involved in pistil development and in the abiotic and biotic stresses response and belong to a family of transcriptional regulators present in three eukaryotic kingdoms (Heim et al. 2003;Skinner et al. 2010). The study of Miao et al. (2020) allowed for the identification of 268 bHLH genes in B. oleracea, 440 genes in B. napus, and 251 genes in B. rapa, including 21 new bHLH members and the gene structures, Phylogeny threes and the conservative motif analysis were carried out showing similar expression patterns between B. rapa and B. napus in roots and contrasting ones in stems, leaves and in reproductive organs (Miao et al. 2020). Genome wide analysis of the bHLH transcription factor was studied for Chinese-cabbage showing the interaction network associated to the ArabidopsisbHLH genes and in cabbage, which revealed several chilling response patterns (Song et al. 2019;Shan et al. 2019). ...
... S5, S14, S15, and S45 appear to be the most recessive S-alleles in the stigma, while S23, S29, S35, S36, and S39 appear to be the most dominant. The importance of dominance between S-alleles in the breeding system of Brussels sprouts is discussed, and an explanation for the high frequency of S2 based on its unusual dominance behavior is proposed (Ockendon 1975 Ockendon 1982;Brace et al. 1994;Kusaba et al. 1999;Kimura et al. 2002;Sato et al. 2003;Shan et al. 2019;Miao et al. 2020 ...
... With the development of sequencing skills and bioinformatics, the genome sequences of Brassica plants, B. napus, B. rapa, and B. oleracea, have been completed at recent years (Lagercrantz 1998;Morant et al. 2002;Blanc et al. 2003;Wang et al. 2011;Cheng et al. 2014;Nguyen et al. 2020). Several family genes have been identified in Brassica species based on the genome databases (Liang et al. 2019;Wei et al. 2019;Zhao et al. 2019;Miao et al. 2020). We conducted the comparative analysis of PHO1 family genes in three Brassica species and analyzed their expression patterns under low Pi stress. ...
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... The bHLH gene family gets its name from the bHLH domain it contains. This domain is comprised of 50-60 amino acids which can be divided into basic amino acids at the Nterminus (10-15 amino acids) and the Helix-Loop-Helix (HLH) at the C-terminus (about 40 amino acids) (Li et al. 2006;Miao et al. 2020;Zhang et al. 2020). Furthermore, this region contains six basic residues and a highly conserved HER motif (His 5-Glu 9-Arg 13), both of which are believed to bind certain DNA sequences (Kavas et al. 2016;Kurt & Filiz 2018;Sun et al. 2020). ...
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The basic Helix-Loop-Helix (bHLH) superfamily is the most widespread family of transcription factors in eukaryotic organisms, which can activate the expression of genes by interacting with specific promoters in the genes. The bHLH transcription factors direct the development and metabolic process of plants, including flowering initiation and secondary metabolite production, by attaching to specific sites on their promoters. These transcription factors are essential for encouraging plant tolerance or the adjustment to harsh environmental conditions. The involvement of bHLH genes in anthocyanin formation in fleshy fruit-bearing plants, as well as the role of these genes in response to stimuli including drought, salt, and cold stress, are discussed in this article. New concepts and goals for the production of stress-tolerant fruit species are suggested. Furthermore, solid evidence for the critical role of bHLH genes in the growth and development, as well as anthocyanin biosynthesis in fleshy fruit plants, are also presented in this article. This review identifies several future research directions that can shed light on the roles of bHLH genes in fruit-bearing plants and will assist the use of these genes in efforts to breed fruit crop varieties that are more resistant to stress. Generally, there has been little research carried out on the role of bHLHs transcription factor family genes in fleshy fruit-bearing plant species and more in-depth studies are required to fully understand the diverse role of bHLH genes in these species.
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... The error bar is standard deviation (n = 3; P < 0.05; Student's t test) ◂ Content courtesy of Springer Nature, terms of use apply. Rights reserved. of homologous genes in different species are similar, but there are differences to some extent (Lohani et al. 2019;He et al. 2020;Miao et al. 2020). For example, some genes with high expression in B. napus roots, stem and leaves had low expression in B. rapa and B. oleracea, while some gene with low expression in B. napus flowers and siliques were highly expressed in B. rapa and B. oleracea (Miao et al. 2020). ...
... Rights reserved. of homologous genes in different species are similar, but there are differences to some extent (Lohani et al. 2019;He et al. 2020;Miao et al. 2020). For example, some genes with high expression in B. napus roots, stem and leaves had low expression in B. rapa and B. oleracea, while some gene with low expression in B. napus flowers and siliques were highly expressed in B. rapa and B. oleracea (Miao et al. 2020). ...
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... 162 bHLH A. thaliana and 167 rice bHLH genes were also grouped into 25 subfamilies in A. thaliana ). Besides, different other studies on this bHLH family have been reported in plant species before phylogenetic tree comparison, as described by Miao et al. (2020). In legumes, studies have been showing that transcriptomic techniques are the most positive ways to elucidate abiotic stresses (Babar et al. 2014). ...
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The basic helix loop helix (bHLH) transcription factor comprises one of the largest plant-specific transcriptional regulators in plant growth and development that response to biotic and abiotic stresses. Many members of bHLH play essential roles in the growth of root hair and response to drought, salt, and cold stresses. The family of bHLH genes has been found in many species; nevertheless, the barrel medic and alfalfa species still have a minute gap of bHLH new members thus far. This research aims to identify members of the bHLH family in barrel medic and alfalfa and elucidate their expression pattern level, network analysis, predictive 3D modeling and phylogenetic relationships. Here, we identified and characterized the bHLH gene family in both barrel medic and alfalfa plants and their genes expression response to drought, salinity, and cold stresses. A total of 159 MtbHLH and 133 MsbHLH genes were identified and characterized, divided into 18 subgroups and 17 subgroups, respectively. As a ubiquitous and popular method, neighbor-joining clustering was used. Based on the phylogenetic analyses, the VIII and IX subfamily and X subfamily were selected as the stress-related subfamily in these two species. The 154 MtbHLH genes were progressively distributed on the 8 chromosomes and 23 tandem duplicated genes, and 44 duplicated genes segments were detected in MtbHLH family. The analyses of gene ontology discovered the bHLH predominantly functions in protein and DNA binding in these two species. The results of Ka/Ks were < 1, which showed that the most orthologous of the bHLH gene values was found between A. thaliana and M. truncatula species. Remarkably, 7 MtbHLH and 10 MsbHLH genes were selected and validated with qRT-PCR after the treatment’s samples sampled under stressed abiotic conditions. The similar expression patterns between M. truncatula and M. sativa L have demonstrated identical expression patterns level in the root, and contrasting patterns in the stems and leaves were diverse. It was highlighted that the gene expression analyses of 17 bHLH genes were up-regulated to stresses, respectively, apart from some genes that were timely trended down-regulated to control (0 h). This study provided a concise understanding of the tissue specific of bHLH gene functions in genome-wide levels under drought, salt, and cold stresses. Our analyses provide the first insights onto the M. truncatula and M. sativa L evolution that contributes to molecular breeding for improving plant yield and stress tolerance.
... bHLHs are involved in regulating the synthesis of flavonoids (Ludwig et al., 1989), which play important roles in the ROS homeostasis under these stresses. As an illustration of the size of the bHLHs family in various plant species, 164 bHLH TFs have been found in Arabidopsis (Arabidopsis thaliana L.), while there are 180 in rice (Oryza sativa L.), 190 in tobacco (Nicotiana tabacum L.), 191 in grapes (Vitis vinifera L.), 102 in walnut (Juglans regia L.), 85 in Ginkgo biloba, 268 in Brassica oleracea, 440 in Brassica napus, and 251 Brassica rapa (Xiong et al., 2005;Jaillon et al., 2007;Rushton et al., 2008;Carretero-Paulet et al., 2010;Miao et al., 2020;Zhou et al., 2020;Zhao et al., 2021). ...
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Basic helix-loop-helix proteins (bHLHs) comprise one of the largest families of transcription factors in plants. They have been shown to be involved in responses to various abiotic stresses, such as drought, salinity, chilling, heavy metal toxicity, iron deficiency, and osmotic damages. By specifically binding to cis-elements in the promoter region of stress related genes, bHLHs can regulate their transcriptional expression, thereby regulating the plant’s adaptive responses. This review focuses on the structural characteristics of bHLHs, the regulatory mechanism of how bHLHs are involved transcriptional activation, and the mechanism of how bHLHs regulate the transcription of target genes under various stresses. Finally, as increasing research demonstrates that flavonoids are usually induced under fluctuating environments, the latest research progress and future research prospects are described on the mechanisms of how flavonoid biosynthesis is regulated by bHLHs in the regulation of the plant’s responses to abiotic stresses.