柑橘脱落酸受体基因家族鉴定及CrPYL9调控果实成熟的作用研究

doi:10.16420/j.issn.0513-353x.2025-0791

园艺学报 ›› 2026, Vol. 53 ›› Issue (6): 1590-1612.doi: 10.16420/j.issn.0513-353x.2025-0791

• 遗传育种·种质资源·分子生物学 • 上一篇下一篇

柑橘脱落酸受体基因家族鉴定及CrPYL9调控果实成熟的作用研究

刘淑梅¹^,², 吕远达¹, 姜波¹, 刘太波²^,^*(), 闫化学¹^,^*()

¹ 广东省农业科学院果树研究所，农业农村部南亚热带果树生物学与遗传资源利用重点实验室，广东省果树科学与技术研究重点实验室，广州 510640
² 岭南现代农业科学与技术广东省实验室，广东未来作物精准育种基础研究中心，华南农业大学生命科学学院，亚热带农业生物资源保护与利用国家重点实验室，广东省农业生物发育与环境适应重点实验室，广州 510642

收稿日期:2025-11-07 修回日期:2025-12-21 出版日期:2026-06-24 发布日期:2026-06-24
通讯作者:
^* E-mail：tbliu@scau.edu.cn，
yanhuaxue@gdaas.cn
基金资助:
广东省自然科学基金项目(2024A1515010921); 广东省自然科学基金项目(2026A1515012685); 国家自然科学基金项目(32072535)

Genome-Wide Identification of ABA Receptor Gene Family in Citrus and Functional Analysis of CrPYL9 in Fruit Maturation

LIU Shumei¹^,², LÜ Yuanda¹, JIANG Bo¹, LIU Taibo²^,^*(), YAN Huaxue¹^,^*()

¹ Institute of Fruit Tree Research， Guangdong Academy of Agricultural Sciences，Key Laboratory of South Subtropical Fruit Biology and Genetic Resource Utilization，Ministry of Agriculture and Rural Affairs，Guangdong Provincial Key Laboratory of Fruit Tree Science and Technology， Guangzhou 510640， China
² Guangdong Laboratory for Lingnan Modern Agriculture， Guangdong Basic Research Center of Excellence for Precise Breeding of Future Crops，College of Life Sciences，South China Agricultural University，State Key Laboratory for Conservation and Utilization of Subtropical Agro-Bioresources，Guangdong Provincial Key Laboratory for the Development Biology and Environmental Adaptation of Agricultural Organisms， Guangzhou 510642， China

Received:2025-11-07 Revised:2025-12-21 Published:2026-06-24 Online:2026-06-24
Contact:
^* E-mail：tbliu@scau.edu.cn，
yanhuaxue@gdaas.cn

摘要/Abstract

摘要：

脱落酸（abscisic acid，ABA）受体Pyrabactin resistance 1/PYR1-Like/Regulatory component of ABA receptor（PYR/PYL/RCAR，PYL）是ABA信号转导途径的重要组分，参与调控植物的生长发育和逆境胁迫响应。分别在甜橙[Citrus sinensis（L.）Osbeck]和克里曼丁橘（C. clementina Hort. ex Tan.）中鉴定出11个和14个PYL基因。根据系统进化分析将柑橘PYL基因家族成员分成了3个亚家族。柑橘PYL表达具有明显的组织特异性，且差异响应植物生长调节剂处理。分子遗传学实验证实CrPYL9在柑橘果实的成熟和转色过程中起正向调控作用，为深入解析ABA调控非跃变型果实成熟机制提供了证据。

关键词: 柑橘, 脱落酸受体, 基因家族, 果实成熟

Abstract:

The abscisic acid（ABA）receptor Pyrabactin resistance 1/PYR1-Like/Regulatory component of ABA receptor（PYR/PYL/RCAR，PYL）is an essential component of the ABA signaling pathway，responsible for regulating various physiological processes in plants such as growth，development，and stress responses. In this study，a total of 11 and 14 PYL genes were identified in sweet orange[Citrus sinensis（L.）Osbeck]and Clementine mandarin（C. clementina Hort. ex Tan.），respectively. Through phylogenetic analysis，citrus PYL genes were categorized into three distinct subfamilies. The expression patterns of citrus PYL genes showed tissue-specific characteristics and diverse responses to external hormone treatments. The results of molecular genetic experiments further confirmed that CrPYL9 positively regulates fruit maturation and color change，providing new insights into the mechanism of ABA-mediated regulation of non-climacteric fruit maturation.

Key words: Citrus, abscisic acid receptor, gene family, fruit maturation

刘淑梅, 吕远达, 姜波, 刘太波, 闫化学. 柑橘脱落酸受体基因家族鉴定及CrPYL9调控果实成熟的作用研究[J]. 园艺学报, 2026, 53(6): 1590-1612.

LIU Shumei, LÜ Yuanda, JIANG Bo, LIU Taibo, YAN Huaxue. Genome-Wide Identification of ABA Receptor Gene Family in Citrus and Functional Analysis of CrPYL9 in Fruit Maturation[J]. Acta Horticulturae Sinica, 2026, 53(6): 1590-1612.

图/表 10

表1 引物名称和序列

Table 1 Primer name and sequence

基因 Gene	引物名称 Primer name	引物序列（5′-3′） Primer sequence	引物名称 Primer name	引物序列（5′-3′） Primer sequence
CrPYL1	PYL1qF	TTGGAGATGCTGGATGATGAG	PYL1qR	AACCGATGTGACTGAACGATAA
CrPYL2	PYL1qF	CTGGTCTGGTTCGCTACT	PYL1qR	TCGCCACATACGACTTAA
CrPYL3	PYL1qF	GGTTGGTATTGGCTGATT	PYL1qR	AATAGCATCCTCCATTGTT
CrPYL4	PYL1qF	CCTAATAACACAACGGATT	PYL1qR	TTCTCTTATGCTGCCTAC
CrPYL5	PYL1qF	ATGGCTGATAATGATGAT	PYL1qR	ATGTTGTT AGAGTTGGAA
CrPYL6	PYL1qF	ATGAACGGTGTTGATGCG	PYL1qR	GGCTTGTACTTCTGTGGC
CrPYL7-1	PYL1qF	TGGCAAGATTGAGGAGGA	PYL1qR	AGTTGTAGCGGGAAGACC
CrPYL7-2	PYL1qF	TCAACCGCAGAAGTATAAGC	PYL1qR	GACCGTAACAATAGACGAGTAG
CrPYL7-3	PYL1qF	GGCAAGATTGAGGAGGAC	PYL1qR	AGTTGTAGCGGGAAGACC
CrPYL8	PYL1qF	CGGTGGTCGTGGAGTCTTA	PYL1qR	CTGCTGATGCTGCTGTTGT
CrPYL9	PYL1qF	AACCACAACCCACAACCCTAAGT	PYL1qR	CGCCTGACCACGGACCAA
CrPYL10	PYL1qF	CTGTTCCGTCTCCGATAA	PYL1qR	CGTCCAGAATGTCAAGTC
CrPYL11	PYL1qF	ATGGAGGAGCTGAAAGTG	PYL1qR	ACATCTGCTAACGAAAGG
CrActin	ActinqF	AGAACTATGAACTGCCTGATGGC	ActinqR	GCTTGGAGCAAGTGCTGTGATT
CrPYL9	CDS F	AATCGACTCTAGAAAGCTTATGCCTGCCTCATCAATACAAG	CDS R	CTAGTATTTAAATGTCGACTTACTTGTTCGATCTAGCAGCCATG
CrPYL9	TRV-PYL9F	AAGGTTACCGAATTCTCTAGACGTTTGGTCCGTGGTCAG	TRV-PYL9R	GAGACGCGTGAGCTCGGTACCGCATGGCGAAGCGTGTAG

图1 甜橙（Cs）、克里曼丁橘（Cr）、拟南芥（At）和水稻（Os）PYL家族成员系统进化分析

Fig. 1 Phylogenetic analysis of PYL family members in Sweet orange（Cs），Clementine mandarin（Cr），Arabidopsis thaliana（At）and Oryza sativa（Os）

图2 甜橙CsPYL（A）和克里曼丁橘CrPYL（B）家族的染色体定位

Fig. 2 Chromosomal localization of PYL family genes in sweet orange（A）and Clementine mandarin（B）

表2 柑橘PYL蛋白理化性质

Table 2 The physical and chemical properties analysis of PYL proteins in Citrus

蛋白 Protein	序列ID Sequence ID	氨基酸数量 Number of amino acids	分子量/Da Molecular weight	等电点 pI	不稳定系数 Instability index	脂肪族aa 系数 Aliphatic index	亲水性指数 Hydropathy index	亚细胞定位 Subcellular localization
CsPYL1	Cs_ont_1g020400.1	216	23 982.21	4.91	32.15	87.04	-0.167	Cyt
CrPYL1	Ciclev10027552m.1	187	20 974.68	4.90	30.81	80.70	-0.327	Chl，Nu
CsPYL2	Cs_ont_4g004630.1	172	19 641.45	5.39	37.01	77.67	-0.289	Cyt
CrPYL2	Ciclev10009701m.1	172	19 627.42	5.39	36.51	77.09	-0.287	Cyt，Chl，Nu
CsPYL3	Cs_ont_4g004660.1	179	19 936.37	5.11	31.66	76.26	-0.322	Chl，Cyt，Nu
CrPYL3	Ciclev10010269m.1	179	19 969.45	5.46	31.22	77.37	-0.292	Cyt，Nu
CsPYL4	Cs_ont_4g005330.1	190	21 382.28	5.90	43.51	85.00	-0.343	Cyt
CrPYL4	Ciclev10010262m.1	190	21 382.28	5.90	43.51	85.00	-0.343	Cyt
CsPYL5	Cs_ont_4g008950.1	181	20 580.22	4.74	37.32	70.55	-0.42	Chl，Cyt，Nu
CrPYL5	Ciclev10009650m.1	181	20 589.23	4.82	43.74	70.55	-0.418	Cyt，Chl，Nu
CsPYL6	Cs_ont_5g004380.1	186	21 089.04	6.44	41.65	91.56	-0.351	Chl
CrPYL6	Ciclev10022440m.1	186	21 089.04	6.44	41.65	91.56	-0.351	Chl
CsPYL7	Cs_ont_5g049350.1	197	22 204.13	5.85	39.57	89.85	-0.374	Chl
CrPYL7-1	Ciclev10022578m.1	168	19 176.97	8.88	51.75	84.58	-0.421	Chl，Nu
CrPYL7-2	Ciclev10022206m.1	214	24 098.33	6.11	38.98	91.82	-0.293	Chl
CrPYL7-3	Ciclev10022353m.1	197	22 144.12	5.98	39.63	91.83	-0.332	Chl，Nu
CsPYL8	Cs_ont_6g013980.1	245	26 654.90	7.69	44.67	84.61	-0.284	Cyt
CrPYL8	Ciclev10012583m.1	245	26 654.90	7.69	44.67	84.61	-0.284	Cyt
CsPYL9	Cs_ont_7g002120.1	201	22 137.99	7.68	62.22	77.01	-0.344	Cyt
CrPYL9	Ciclev10032781m.1	201	22 137.99	7.68	62.22	77.01	-0.344	Cyt
CsPYL10	Cs_ont_7g010910.1	165	18 735.69	5.51	40.06	69.70	-0.656	Nu
CrPYL10	Ciclev10012769m.1	209	23 268.87	5.08	35.06	82.44	-0.362	Cyt，Nu
CsPYL11	Cs_ont_9g027270.1	189	21 335.19	6.41	43.50	96.88	-0.413	Nu
CrPYL11	Ciclev10005981m.1	189	21 335.19	6.41	43.50	96.88	-0.413	Nu
CrPYL12	Ciclev10033656m.1	158	17 571.81	4.70	35.71	80.76	-0.282	Chl

表2 柑橘PYL蛋白理化性质

Table 2 The physical and chemical properties analysis of PYL proteins in Citrus

蛋白 Protein	序列ID Sequence ID	氨基酸数量 Number of amino acids	分子量/Da Molecular weight	等电点 pI	不稳定系数 Instability index	脂肪族aa 系数 Aliphatic index	亲水性指数 Hydropathy index	亚细胞定位 Subcellular localization
CsPYL1	Cs_ont_1g020400.1	216	23 982.21	4.91	32.15	87.04	-0.167	Cyt
CrPYL1	Ciclev10027552m.1	187	20 974.68	4.90	30.81	80.70	-0.327	Chl，Nu
CsPYL2	Cs_ont_4g004630.1	172	19 641.45	5.39	37.01	77.67	-0.289	Cyt
CrPYL2	Ciclev10009701m.1	172	19 627.42	5.39	36.51	77.09	-0.287	Cyt，Chl，Nu
CsPYL3	Cs_ont_4g004660.1	179	19 936.37	5.11	31.66	76.26	-0.322	Chl，Cyt，Nu
CrPYL3	Ciclev10010269m.1	179	19 969.45	5.46	31.22	77.37	-0.292	Cyt，Nu
CsPYL4	Cs_ont_4g005330.1	190	21 382.28	5.90	43.51	85.00	-0.343	Cyt
CrPYL4	Ciclev10010262m.1	190	21 382.28	5.90	43.51	85.00	-0.343	Cyt
CsPYL5	Cs_ont_4g008950.1	181	20 580.22	4.74	37.32	70.55	-0.42	Chl，Cyt，Nu
CrPYL5	Ciclev10009650m.1	181	20 589.23	4.82	43.74	70.55	-0.418	Cyt，Chl，Nu
CsPYL6	Cs_ont_5g004380.1	186	21 089.04	6.44	41.65	91.56	-0.351	Chl
CrPYL6	Ciclev10022440m.1	186	21 089.04	6.44	41.65	91.56	-0.351	Chl
CsPYL7	Cs_ont_5g049350.1	197	22 204.13	5.85	39.57	89.85	-0.374	Chl
CrPYL7-1	Ciclev10022578m.1	168	19 176.97	8.88	51.75	84.58	-0.421	Chl，Nu
CrPYL7-2	Ciclev10022206m.1	214	24 098.33	6.11	38.98	91.82	-0.293	Chl
CrPYL7-3	Ciclev10022353m.1	197	22 144.12	5.98	39.63	91.83	-0.332	Chl，Nu
CsPYL8	Cs_ont_6g013980.1	245	26 654.90	7.69	44.67	84.61	-0.284	Cyt
CrPYL8	Ciclev10012583m.1	245	26 654.90	7.69	44.67	84.61	-0.284	Cyt
CsPYL9	Cs_ont_7g002120.1	201	22 137.99	7.68	62.22	77.01	-0.344	Cyt
CrPYL9	Ciclev10032781m.1	201	22 137.99	7.68	62.22	77.01	-0.344	Cyt
CsPYL10	Cs_ont_7g010910.1	165	18 735.69	5.51	40.06	69.70	-0.656	Nu
CrPYL10	Ciclev10012769m.1	209	23 268.87	5.08	35.06	82.44	-0.362	Cyt，Nu
CsPYL11	Cs_ont_9g027270.1	189	21 335.19	6.41	43.50	96.88	-0.413	Nu
CrPYL11	Ciclev10005981m.1	189	21 335.19	6.41	43.50	96.88	-0.413	Nu
CrPYL12	Ciclev10033656m.1	158	17 571.81	4.70	35.71	80.76	-0.282	Chl

图3 甜橙CsPYL（A1 ~ C1）和克里曼丁橘CrPYL（A2 ~ C2）家族成员基因和蛋白结构分析 A1：甜橙PYL蛋白保守基序。不同颜色代表不同基序。B1：甜橙不同PYL成员的PYR_PYL_RCAR_like保守结构域。C1：甜橙PYL基因结构。黄色部分表示外显子，黑线表示内含子，绿色部分代表非翻译区。A2：克里曼丁橘PYL蛋白保守基序。不同颜色代表不同基序。B2：克里曼丁橘不同PYL成员的PYR_PYL_RCAR_like保守结构域。C2：克里曼丁橘PYL基因结构。绿色部分表示外显子，黑线表示内含子，黄色部分代表非翻译区

Fig. 3 Analysis of gene and protein structure of CsPYL family members in sweet orange（A1-C1）and Clementine mandarin（A2-C2） A1：Conserved motifs of sweet orange PYL proteins. Different colors represent different motifs. B1：The PYR_PYL_RCAR-like conserved domains of different PYL members in sweet orange. C1：The structure of sweet orange PYL genes. Yellow regions represent exons，black lines indicate introns，and green regions represent untranslated regions（UTR）. A2：Conserved motifs of Clementine mandarin PYL proteins. Different colors represent different motifs. B2：The PYR_PYL_RCAR-like conserved domains of different PYL members in Clementine mandarin. C2：The structure of Clementine mandarin PYL genes. Green regions represent exons，black lines indicate introns，and yellow regions represent untranslated regions（UTR）

图4 CrPYL基因启动子顺式作用元件分析 A：CrPYL家族成员启动上顺式作用元件的分布，不同元件分别用不用颜色圆圈表示；B：CrPYL家族成员启动子上各顺式作用元件的数量，字母a ~ i代表不同的顺式作用元件

Fig. 4 Analysis of cis-acting elements of in the CrPYL gene promoters A：Distribution of cis-acting elements on the promoters of CrPYL family members. Different elements are represented by circles of varying colors；B：Number of each cis-acting element on the promoters of CrPYL family members. The letters a-i represent different cis-acting elements

图5 CrPYL家族基因的组织表达分析

Fig. 5 Tissue expression of CrPYL genes

图6 CrPYL在花后150 ~ 300 d沙糖橘果皮和果肉中的表达

Fig. 6 Expression analysis of CrPYL genes in the peel and pulp of Citrus reticulata‘Shatangju’ from 150 to 300 days post anthesis

图7 CrPYL基因在克里曼丁橘叶片中响应激素处理的表达分析

Fig. 7 Expression analysis of CrPYL genes in response to phytohormone treatments in the leaves of Clementine mandarin

图8 过表达和VIGS沉默CrPYL9基因对离体甜橙青果着色的影响 A：瞬时过表达CrPYL9基因加速离体甜橙青果着色。B：病毒诱导的CrPYL9沉默延缓离体甜橙青果着色。C：处理22 d后果实表面色差分析。D：处理22 d后果皮中叶绿素含量分析。t检验，**表示与对照差异显著（P < 0.05）

Fig. 8 Effects of overexpression and VIGS-mediated silencing of CrPYL9 on coloration of detached sweet orange green fruits A：Transient overexpression of CrPYL9 accelerates coloration of detached sweet orange green fruits. B：VIGS-mediated silencing of CrPYL9 delays coloration of detached sweet orange green fruits. C：Analysis of color difference on fruit surface after 22 days of treatment. D：Chlorophyll content in the fruit peel after 22 days of treatment. t-test，** indicates a significant difference（P < 0.05）

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柑橘脱落酸受体基因家族鉴定及CrPYL9调控果实成熟的作用研究

Genome-Wide Identification of ABA Receptor Gene Family in Citrus and Functional Analysis of CrPYL9 in Fruit Maturation

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柑橘脱落酸受体基因家族鉴定及CrPYL9调控果实成熟的作用研究

Genome-Wide Identification of ABA Receptor Gene Family in Citrus and Functional Analysis of CrPYL9 in Fruit Maturation

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