三种芍药属植物FLOWERING LOCUS T(FT)同源基因的克隆与表达分析

doi:10.3969/j.issn.1004-1524.20240082

浙江农业学报 ›› 2025, Vol. 37 ›› Issue (1): 90-102.DOI: 10.3969/j.issn.1004-1524.20240082

三种芍药属植物FLOWERING LOCUS T(FT)同源基因的克隆与表达分析

李杏(), 刘燕, 高健洲^*()

北京林业大学园林学院,北京 100080

收稿日期:2024-01-18 出版日期:2025-01-25 发布日期:2025-02-14
作者简介:李杏(1999—),女,湖南益阳人,硕士研究生,主要从事野生芍药引种生长发育及其FT基因表达差异研究。E-mail: m15011578276@163.com
通讯作者: *高健洲,E-mail:noplenhill@163.com
基金资助:
雄安新区科技创新专项(2022XAGG0100)

Cloning and expression analysis of homologous FLOWERING LOCUS T(FT) genes from three species of Paeonia spp.

LI Xing(), LIU Yan, GAO Jianzhou^*()

College of Landscape Architecture, Beijing Forestry University, Beijing 100080, China

Received:2024-01-18 Online:2025-01-25 Published:2025-02-14

摘要/Abstract

摘要： 为探究FT基因在芍药属植物开花和种子下胚轴休眠解除中是否具有多效性,克隆得到川赤芍(Paeonia veitchii)、新疆芍药(P. anomala)和芍药凤凰涅槃(P. lactiflora ‘Fenghuangniepan’)的FT同源基因(依次命名为PvFT、PaFT和PlFT1)的编码(coding sequence, CDS)序列,对它们进行生物信息学分析,并通过qRT-PCR技术分析3种芍药种子休眠解除和成花阶段FT基因的表达模式。结果表明,3种芍药属植物FT基因序列高度保守,CDS全长均为522 bp,编码173个氨基酸。系统进化分析显示,3个FT基因编码的蛋白质序列与牡丹的PsFT蛋白亲缘关系最近。时空特异性表达分析显示,3种芍药属植物种子中FT基因在种子下胚轴伸长时均上调表达,没有与下胚轴休眠期的长短表现出一致性;相应成株中PvFT和PlFT1基因分别在能开花的川赤芍的花期和凤凰涅槃的蕾期上调表达,而在不开花的川赤芍和新疆芍药中,PvFT和PaFT基因维持低水平表达。PaFT、PlFT1和PvFT基因在芍药属植物开花和种子下胚轴休眠解除中具有多效性,在成花和种子下胚轴伸长过程中起正调控作用。

关键词: 野生芍药, FT基因, 克隆, 开花, 种子, 下胚轴

Abstract:

In order to explore whether the FT gene has pleiotropy in the flowering and seed hypocotyl dormancy release of Paeonia, the CDS (coding sequence) region of the FT homologous genes (named PvFT, PaFT, and PlFT1) of P. vetchii, P. anomala, and P. lactiflora ‘Fenghuangniepan’ were cloned and analyzed using bioinformatics, and the expression patterns of FT genes during the dormancy release and flowering stages of three peony germplasm seeds were analyzed using qRT-PCR technology. The results show that the FT gene sequences are highly conserved across the three Paeonia species, with a full CDS length of 522 bp encoding 173 amino acids. Phylogenetic analysis indicates that the proteins encoded by these three FT genes are most closely related to the PsFT protein from the peony. Temporal and spatial expression analysis reveal that in the seeds of three species of Paeonia, the expression levels of the FT genes upregulate during hypocotyl elongation, and there are no consistency with the length of the hypocotyl dormancy period. In corresponding adult plants, the expression levels of PvFT and PlFT1 genes are upregulated during the flowering stage of P. veitchii and the budding stage of P. lactiflora ‘Fenghuangnipan’, while the expression levels of PvFT and PaFT genes are expressed at a low level during the non-flowering of P. veitchii and P. anomala. PaFT, PlFT1, and PvFT genes exhibit pleiotropic effects in the flowering and seed germination processes of Paeonia species, playing a positive regulatory role during flowering and seed hypocotyl elongation.

Key words: wild peony, FT gene, clone, flowering, seed, hypocotyl

中图分类号:

S682.12
Q786

李杏, 刘燕, 高健洲. 三种芍药属植物FLOWERING LOCUS T(FT)同源基因的克隆与表达分析[J]. 浙江农业学报, 2025, 37(1): 90-102.

LI Xing, LIU Yan, GAO Jianzhou. Cloning and expression analysis of homologous FLOWERING LOCUS T(FT) genes from three species of Paeonia spp.[J]. Acta Agriculturae Zhejiangensis, 2025, 37(1): 90-102.

图/表 12

表1 本研究所用引物

Table 1 Primers used in this study

引物名称 Primer name	上游引物序列 Forward primer sequence(5’→3’)	下游引物序列 Reverse primer sequence(5’→3’)	退火温度 Annealing temperature/℃	用途 Function
F01	ATGCCTAGAAATAGGGATCC	TTATCTTCTTCCGCCTGATC	52	基因克隆Gene cloning
F02	AGAGTCCACGCCCAACTGCT	GCCACTGGTGAGCCAAGATTA	57	实时荧光定量PCR qRT-PCR
A01	CATGGTATAGTCAGCAACTGGGATG	GGCTTTGGGGTTAAGGGGTG	57	实时荧光定量PCR qRT-PCR

图1 三种芍药属植物FT基因的克隆 M,DL 2 000 DNA marker;PaFT、PlFT1和PvFT分别表示新疆芍药、凤凰涅槃和川赤芍FT基因的PCR产物。

Fig.1 Cloning of FT genes from three species of Paeonia M, DL 2 000 marker; PaFT, PlFT1, and PvFT represent PCR products of the FT genes of P. anomala, P. lactiflora ‘Fenghuangnipan’ and P. veitchii, respectively.

图2 三种芍药属植物FT基因与芍药大富贵PlFT基因的核苷酸序列比对黑色区域,相似性=100%;红色区域,相似性≥75%;蓝色区域,相似性≥50%;红色方框表示差异碱基位点。

Fig.2 Nucleotide sequence alignment of FT genes in three species of Paeonia and PlFT gene in P. lactiflora ‘Dafugui’ Black blocks, Similarity=100%; Red blocks, Similarity≥75%; Blue blocks, Similarity≥50%; Red boxes represent differential base sites.

图3 三种芍药属植物FT氨基酸序列与其他植物FT蛋白的多序列比对黑色区域,相似性=100%的氨基酸;红色区域,相似性≥75%的氨基酸;蓝色区域,相似性≥50%的氨基酸。CmFT,板栗, No.OL411873.1;CsFT1,山茶,No.AB741571.1;FcFT,日本山毛榉,No.AB775532.1;HmFT,绣球,No.MF374627.1;PnFT2b 黑杨,No.AB161108.1;PsFT,牡丹,No.KF113360.1;ZjFT,枣,No.KR8728441。

Fig.3 Multiple sequence alignment of FT amino acid sequence from three species of Paeonia with FT proteins from other plants Black blocks, Amino acids with similarity=100%; Red blocks, Amino acids with similarity ≥ 75%; Blue blocks, Amino acids with similarity ≥ 50%. CmFT, Castanea mollissima, No.OL411873.1; CsFT1, Camellia japonica, No.AB741571.1; FcFT, Fagus crenata, No.AB775532.1; HmFT, Hydrangea macrophylla, No.MF374627.1; PnFT2b, Populus nigra, No.AB161108.1; PsFT, P. suffruticosa, No.KF113360.1; ZjFT, Ziziphus jujuba, No.KR8728441.

表2 三种芍药属植物FT蛋白的理化性质

Table 2 Physical and chemical properties of FT proteins in three species of Paeonia

基因 Gene	基因ID Gene ID	分子量 Molecular mass/u	总原子个数 Total number of atoms	等电点 Isoelectric point	分子式 Molecular formula	亲水性总平均值 Total average hydrophilicity	不稳定指数 Instability index
PaFT	OQ079526	19 599.15	2 729	8.42	C₈₆₈H₁₃₄₇N₂₅₃O₂₅₅S₆	-0.41	37.16
PlFT1	OQ079527	19 613.17	2 732	8.42	C₈₆₉H₁₃₄₉N₂₅₃O₂₅₅S₆	-0.394	37.22
PvFT	OQ079529	19 599.15	2 729	8.42	C₈₆₈H₁₃₄₇N₂₅₃O₂₅₅S₆	-0.41	37.16

表3 三种芍药属植物FT蛋白的氨基酸组成差异

Table 3 Differences in amino acid composition of FT proteins in three species of Paeonia

氨基酸种类 Types of amino acid	PaFT/PvFT		PlFT1
氨基酸种类 Types of amino acid	氨基酸数量 Number of amino acids	占氨基酸总量的百分比 Percentage of total amino acids/%	氨基酸数量 Number of amino acids	占氨基酸总量的百分比 Percentage of total amino acids/%
丙氨酸Ala(A)	7	4.0	6	3.5
精氨酸Arg(R)	19	11.0	19	11.0
天冬酰胺Asn(N)	9	5.2	9	5.2
天冬氨酸Asp(D)	11	6.4	11	6.4
半胱氨酸Cys(C)	4	2.3	4	2.3
谷氨酰胺Gln(Q)	7	4.0	7	4.0
谷氨酸Glu(E)	7	4.0	7	4.0
甘氨酸Gly(G)	14	8.1	14	8.1
组氨酸His(H)	2	1.2	2	1.2
异亮氨酸lle(I)	6	3.5	6	3.5
亮氨酸Leu(L)	14	8.1	13	7.5
赖氨酸Lys(K)	1	0.6	1	0.6
蛋氨酸Met(M)	2	1.2	2	1.2
苯丙氨酸Phe(F)	9	5.2	9	5.2
脯氨酸Pro(P)	14	8.1	14	8.1
丝氨酸Ser(S)	9	5.2	9	5.2
苏氨酸Thr(T)	13	7.5	13	7.5
色氨酸Trp(W)	2	1.2	2	1.2
酪氨酸Tyr(Y)	7	4.0	7	4.0
缬氨酸Val(V)	16	9.2	18	10.4

图4 三种芍药属植物FT蛋白的二级和三级结构预测 A,PlFT1蛋白二级结构预测;B,PaFT和PvFT蛋白二级结构预测;C,3种FT蛋白的三级结构预测模板。图A和B中粉色直线为无规卷曲,蓝色箭头为β折叠,橘色波浪线为α螺旋,黑色方框为二者差异区域。

Fig.4 Prediction of the secondary and tertiary structure of FT proteins in three species of Paeonia A, Prediction of the secondary structure of PlFT1; B, Prediction of the secondary structure of PaFT and PvFT proteins; C, Prediction template of the tertiary structure of FT proteins in three species of Paeonia. The pink straight lines represent random coil, the blue arrows represent β-sheet, orange wavy lines represent α-helix, black box represent the difference area in figures A and B.

图5 三种芍药属植物FT蛋白与其他物种FT同源蛋白系统进化分析数值表示1 000次重复抽样所得40%以上的置信度。图中使用的PEBP家族成员为:At,拟南芥;Bj,芥菜;Ca,杜鹃叶山茶;Ed,台湾枇杷;Ej,枇杷;Gh,陆地棉;Hs,木槿;Md,苹果;Ms,森林苹果;Pb,白梨;Pc,西洋梨;Pf,火棘;Pp,沙梨;Ps,牡丹;Rc,月季;Rr,玫瑰;Vu,豇豆;Vv,葡萄;Zj,枣。

Fig.5 Phylogenetic relationship among FT proteins in three species of Paeonia and other FT-like proteins The number represents the percentage of 1 000 bootstrap replications, only above 40% is showed. The family members of the PEBPs used in the tree are as follows: At, Arabidopsis thaliana; Bj, Brassica juncea; Ca, Camellia azalea; Ed, Eriobotrya deflexa; Ej, Eriobotrya japonica; Gh, Gossypium hirsutum; Hs, Hibiscus syriacus; Md, Malus domestica; Ms, Malus sylvestris; Pb, Pyrus x bretschneideri; Pc, Pyrus communis; Pf, Pyracantha fortuneana; Pp, Pyrus pyrifolia; Ps, P. suffruticosa; Rc, Rosa chinensis; Rr, Rosa rugosa; Vu, Vigna unguiculata; Vv, Vitis vinifera; Zj, Ziziphus jujuba.

图6 三种芍药属植物层积沙藏时种子下胚轴休眠解除过程的解剖结构观察(×10) A、B、C分别为川赤芍、新疆芍药和凤凰涅槃下胚轴解除过程中的解剖结构;Ⅰ代表球形胚时期,Ⅱ代表心形胚时期,Ⅲ代表鱼雷胚时期,Ⅳ代表子叶胚时期;EM,胚;EN,胚乳;CP,子叶原基;时间轴表示3种芍药属植物种子发育至对应时期所需的沙藏层积时间。

Fig.6 Observation on the anatomical structure of seed hypocotyl dormancy release process during sand storage in three species of Paeonia(×10) A, B, C represent the anatomical structures during the process of releasing the hypocotyl of P. veitchii, P. anomala and P. lactiflora ‘Fenghuangnipan’, respectively; Ⅰ represent the spherical embryo stage, Ⅱ represent the heart-shaped embryo stage, Ⅲ represent the torpedo embryo stage, and Ⅳ represent the cotyledon embryo stage; EM, Embryo; EN, Endosperml; CP, Cotyledon primordia; The timelines represent the sand accumulation time required for the seed development of three species of Paeonia to reach their corresponding stages.

图7 三种芍药属植物在种子下胚轴休眠打破阶段FT基因的相对表达量柱上无相同小写字母表示差异显著(P<0.05)。下同。

Fig.7 The relative expression levels of FT genes in three species of Paeonia during the dormancy breaking stage of seed hypocotyls The bars marked without the same lowercase letter indicat significant differences at P<0.05. The same as below.

图8 引种北京后开花的川赤芍和凤凰涅槃生长发育过程观察 A、B分别为开花的川赤芍和凤凰涅槃的4个生长发育阶段;Ⅰ,展叶期;Ⅱ,显蕾期;Ⅲ,开花期;Ⅳ,花谢期。

Fig.8 Observation on the growth and development process of flowering P. veitchii and P. lactiflora ‘Fenghuangniepan’ introduced to Beijing A and B are the four growth and development stages of flowering P. veitchii and P. lactiflora ‘Fenghuangniepan’, respectively; I, Leaf spreading period; Ⅱ, Budding stage; Ⅲ, Flowering period; Ⅳ, Flower and withering period.

图9 不开花川赤芍、开花川赤芍、新疆芍药和凤凰涅槃在不同发育阶段FT基因的相对表达量

Fig.9 Relative expression levels of FT genes in different developmental stages of non-flowering P. veitchii, flowering P. veitchii, P. anomala and P. lactiflora ‘Fenghuangniepan’

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三种芍药属植物FLOWERING LOCUS T(FT)同源基因的克隆与表达分析

Cloning and expression analysis of homologous FLOWERING LOCUS T(FT) genes from three species of Paeonia spp.

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