基于PSY多层级调控的园艺作物品质形成机制研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2025-1289

摘要/Abstract

摘要：

类胡萝卜素是植物体内重要的萜类化合物，对园艺作物外观、风味和营养品质形成具有重要作用。八氢番茄红素合成酶（PSY）作为类胡萝卜素合成通路的限速酶，其编码基因PSY的表达水平直接调控园艺作物中类胡萝卜素的积累。本文系统梳理了PSY基因家族在不同作物中的成员分化与组织特异性表达模式，揭示了其功能多样性的进化基础。进而，重点阐述了PSY的多维度调控网络：在转录层面，多种转录因子通过结合PSY启动子区域精确调控其时空表达；表观遗传机制通过改变染色质状态动态调节PSY的可及性；转录后层面，可变剪接与反式剪接产生功能各异的转录本，实现了对PSY转录本丰度与翻译效率的精细调节；翻译后层面，OR伴侣蛋白与Clp蛋白酶系统拮抗调控PSY蛋白的稳定性，而泛素化修饰则精准控制其降解速率；亚细胞定位机制确保了PSY在质体中的高效催化。此外，PSY的表达与活性还受到光照、温度等环境因子以及乙烯、脱落酸等激素信号的交叉调控，形成了复杂的调控网络。当前研究仍面临诸多挑战，如研究集中于少数模式作物，不同物种间调控通路差异不明，PSY与上下游因子的互作网络及代谢通道机制有待深入解析。未来，研究需借助合成生物学工具对PSY进行理性设计，利用人工智能与多组学技术鉴定关键调控节点，并通过基因编辑与代谢工程策略，实现园艺作物类胡萝卜素含量的精准改良。

关键词: PSY1, 类胡萝卜素, 多层级调控, 园艺作物品质, 生长发育, 环境因子

Abstract:

Carotenoids are important terpenoid compounds in plants, playing a crucial role in the formation of visual, sensory, and nutritional quality in horticultural crops. As a key rate-limiting enzyme in the carotenoid biosynthetic pathway, the expression of the gene encoding phytoene synthase (PSY) plays a pivotal role in regulating carotenoid accumulation in horticultural crops. This review systematically summarizes the diversification of the PSY gene family and their tissue-specific expression patterns across different crops, revealing the evolutionary basis of their functional diversity. Furthermore, this review elaborates on the multi-level regulatory network governing PSY: At the transcriptional level, various transcription factors precisely regulate its spatiotemporal expression by binding to the PSY promoter region, epigenetic mechanisms dynamically modulate PSY accessibility by altering chromatin states; at the post-transcriptional level, alternative splicing and trans-splicing generate functionally distinct transcripts, enabling fine-tuning of PSY transcript abundance and translation efficiency; at the post-translational level, the OR chaperone and Clp protease system antagonistically regulate PSY protein stability, while ubiquitination precisely controls its degradation rate; and subcellular localization mechanism ensures efficient catalysis within plastids. In addition, PSY expression and activity are cross-regulated by environmental factors such as light and temperature, as well as hormonal signals including ethylene and abscisic acid, forming a complex regulatory network. Current research still faces several challenges. Most studies focus on a few model crops, while regulatory pathway differences among species remain poorly understood. In addition, the interaction networks between phytoene synthase (PSY) and its upstream and downstream regulatory factors, as well as the mechanisms underlying metabolic channeling, require further in-depth investigation. In the future, research should employ synthetic biology tools to rationally design PSY for improved stability and catalytic efficiency, utilize artificial intelligence and multi-omics integration to identify key regulatory nodes, and implement gene editing and metabolic engineering strategies to achieve precise enhancement of carotenoid content in horticultural crops.

Key words: PSY1, carotenoids, multi-level regulation, horticultural crop quality, growth and development, environmental factor

马莹莹, 游惠婉, 郑积荣, 汪俏梅, 刘丽红. 基于PSY多层级调控的园艺作物品质形成机制研究进展[J]. 生物技术通报, 2026, 42(3): 1-15.

MA Ying-ying, YOU Hui-wan, ZHENG Ji-rong, WANG Qiao-mei, LIU Li-hong. Advances in the Quality Formation Mechanism of Horticultural Crops Based on Multi-level Regulation of PSY[J]. Biotechnology Bulletin, 2026, 42(3): 1-15.

图/表 3

图1 PSYs作为限速酶基因在园艺作物生长发育及产品品质形成中的核心作用PSY作为限速酶参与的类胡萝卜素代谢途径与园艺产品的颜色、风味和营养品质息息相关，而且该途径衍生的GAs、ABA和SLs激素信号通路也调节了园艺作物的生长、发育和环境适应性。ABA：脱落酸；BCH：非血红素β-环羟化酶；CCDs：类胡萝卜素裂解双加氧酶；CRTISO：类胡萝卜素异构酶；CYP97A：细胞色素P450胡萝卜素β-环羟化酶；CYP97C：细胞色素P450胡萝卜素ε-环羟化酶；DMAPP：二甲基丙烯基焦磷酸；GAs：赤霉素；GGPP：香叶基香叶基焦磷酸；GGPPS：香叶基香叶基焦磷酸合成酶；IPP：异戊烯基焦磷酸；LCY-B：番茄红素β-环化酶；LCY-E：番茄红素ε-环化酶；NCEDs：9-顺式-环氧类胡萝卜素双加氧酶；NXS：新黄质合成酶；PDS：八氢番茄红素脱氢酶；PSY：八氢番茄红素合成酶；SLs：独角金内酯；VDE：紫黄质脱环氧化酶；ZDS：ζ-胡萝卜素去饱和酶；ZEP：玉米黄质环氧化酶；ZISO：ζ-胡萝卜素异构酶（该图由BioRender.com创建）

Fig. 1 PSYs, as rate-limiting enzyme genes, play a central role in the growth and development of horticultural crops and the formation of product qualityPSY, as a rate-limiting enzyme in the carotenoid biosynthesis pathway, is closely related to the color, flavor, and nutritional quality of horticultural crops. Moreover, the hormone signaling pathways of GAs, ABA, and SLs derived from this pathway also regulate the growth, development, and environmental adaptability of horticultural crops. ABA: Abscisic acid; BCH: β-carotene hydrolase; CCDs: carotenoid cleavage dioxygenases; CRTISO: carotene isomerase; CYP97A: cytochrome P450 carotene β-hydroxylase; CYP97C: cytochrome P450 carotene ε-hydroxylase; DMAPP: dimethylallyl diphosphate; GAs: gibberellins; GGPP: geranylgeranyl pyrophosphate; GGPPS: geranylgeranyl pyrophosphate synthase; IPP: isopentenyl diphosphate; LCY-B: lycopene β-cyclase; LCY-E: lycopenes ε-cyclase; NCEDs: 9-cis-epoxycarotenoid dioxygenases; NXS: neoxanthin synthase; PDS: phytoene desaturase; PSY: phytoene synthase; SLs: strigolactones; VDE: violaxanthin de-epoxidase; ZDS: ζ-carotene desaturase; ZEP: zeaxanthin epoxidase; ZISO: ζ-carotene isomerase (This image was created by BioRender.com)

表1 PSYs基因家族成员及功能

Table 1 PSYs gene family members and functions

作物名称 Crop name	PSY成员 PSY members	组织表达部位 Tissue expression sites	基因功能 Gene functions	参考文献 Reference
甘薯 Ipomoea batatas Lam.	IbPSY1、IbPSY2	IbPSY1的转录水平在幼叶中最高，在茎中最低	IbPSY1响应盐胁迫和干旱胁迫	［37］
胡萝卜 Daucus carota L.	DcPSY1、DcPSY2	主要在根、叶片部位高表达	DcPSY1、DcPSY2在根中的表达水平与类胡萝卜素含量呈正相关关系；DcPSY1在叶片的类胡萝卜素积累过程中起着主要作用	［38-39］
甜瓜 Cucumis melo L.	CmPSY1、CmPSY2	CmPSY1在果实发育早期表达量较高，随后表达量下降；CmPSY2的表达趋势相反，在果实发育后期表达增强	CmPSY1主要在绿色组织中发挥作用，而CmPSY2则可能在非绿色组织（如果实）中起主要作用	［40-41］
番茄 Solanum lycopersicum	SlPSY1、SlPSY2、SlPSY3	SlPSY1在果实中高表达；SlPSY2在绿色组织器官（尤其是叶）中高表达；SlPSY3在根中高表达	SlPSY1主导果实成熟阶段类胡萝卜素合成；SlPSY2调控绿色组织光保护相关色素积累；SlPSY3响应胁迫参与根系代谢	［29-30］
桃 Prunus persica	PpPSY1、PpPSY2、PpPSY3	PpPSY1和PpPSY2占据主导地位；PpPSY1在成熟果实中高表达，且在不同遗传背景的桃品种中均表现出果实特异性表达模式；PpPSY2主要在叶片等绿色组织中表达	PpPSY1是果实类胡萝卜素积累的核心调控因子；PpPSY2主要参与光合组织的类胡萝卜素合成，可能与光保护功能相关	［31］
柑橘 Citrus reticulata	CitPSY1、CitPSY2、CitPSY3	CitPSY2在叶片、花器官及果皮中呈低丰度转录	CitPSY1主导果实与光合组织的类胡萝卜素合成；CitPSY3仅存基因组序列，功能尚未解析	［42］
西瓜 Citrullus lanatus	ClPSY1、ClPSY2、ClPSY3	ClPSY1在果肉中显著积累；ClPSY2在叶片中表达量最高；ClPSY3则主要定位于根部	ClPSY1在西瓜果实成熟期间的类胡萝卜素生物合成中起主要作用，与果肉呈色正相关，并受复杂的光照和激素反应性网络调节	［32，43］
辣椒 Capsicum annuum	CaPSY1、CaPSY2.a、CaPSY2.b	CaPSY1在幼果中表达量较低，但在果实发育过程中持续上调表达；CaPSY2.a基因在根、茎、叶、花和果实发育第3和第7个时期中不表达；CaPSY2.b基因在所有组织中均有表达，在茎、叶和果实发育第一个时期表达量相对较高	CaPSY2基因可能补充了CaPSY1的缺失，从而导致黄色辣椒果实呈黄色	［36，44］
西葫芦 Cucurbita pepo	CpPSYA、CpPSYB、CpPSYC	CpPSYA在果实中表达量高，CpPSYB和CpPSYC在叶片中表达量高		［45］
香蕉 Musa ‘Hybrids’	MaPSY1、MaPSY2a、MaPSY2b、MaPSY2c	MaPSY2a、MaPSY2b的表达具有果实特异性		［46］
番红花 Crocus sativus L.	CsPSY1a、CsPSY1b、 CsPSY2、CsPSY3	CsPSY1a和CsPSY1b主要在叶片中表达；CsPSY2转录本主要在柱头中被检测到，而在其他组织中表达却很低；CsPSY3在根尖高表达	CsPSY1b表现出胁迫光调节，CsPSY3的表达与菌根定植和松果内酯的产生有关	［47］
菠菜 Spinacia oleracea	SoPSY1、SoPSY2、 SoPSY3、SoPSY4	SoPSY4在根中高水平表达，显著高于地上部分各组织器官；其余3个SoPSY基因在叶片中高水平表达，营养叶和薹叶中表达最高，其次是薹叶柄、短缩茎和叶柄，在根中表达量最低		［33］
苹果 Malus domestica	MdPSY1、MdPSY2、MdPSY3、MdPSY4、MdPSY5、MdPSY6	仅MdPSY1、MdPSY2和MdPSY4具有酶活性，其中MdPSY1和MdPSY2在叶片和果实成熟过程中高度表达	MdPSY1和MdPSY2与果实中类胡萝卜素合成有关	［34］
甘蓝型油菜 Brassica napus	BnaC.PSY.a BnaA.PSY.b BnaA.PSY.d BnaC.PSY.f BnaA.PSY.d BnaC.PSY.e	BnaC.PSY.a和BnaA.PSY.b在所有组织中均有表达；BnaA.PSY.d和BnaC.PSY.f在富含叶绿体的组织（如叶片）中优先表达；BnaA.PSY.d、BnaC.PSY.e在富含有色体的组织（如花瓣）中优先表达		［35］

表1 PSYs基因家族成员及功能

Table 1 PSYs gene family members and functions

作物名称 Crop name	PSY成员 PSY members	组织表达部位 Tissue expression sites	基因功能 Gene functions	参考文献 Reference
甘薯 Ipomoea batatas Lam.	IbPSY1、IbPSY2	IbPSY1的转录水平在幼叶中最高，在茎中最低	IbPSY1响应盐胁迫和干旱胁迫	［37］
胡萝卜 Daucus carota L.	DcPSY1、DcPSY2	主要在根、叶片部位高表达	DcPSY1、DcPSY2在根中的表达水平与类胡萝卜素含量呈正相关关系；DcPSY1在叶片的类胡萝卜素积累过程中起着主要作用	［38-39］
甜瓜 Cucumis melo L.	CmPSY1、CmPSY2	CmPSY1在果实发育早期表达量较高，随后表达量下降；CmPSY2的表达趋势相反，在果实发育后期表达增强	CmPSY1主要在绿色组织中发挥作用，而CmPSY2则可能在非绿色组织（如果实）中起主要作用	［40-41］
番茄 Solanum lycopersicum	SlPSY1、SlPSY2、SlPSY3	SlPSY1在果实中高表达；SlPSY2在绿色组织器官（尤其是叶）中高表达；SlPSY3在根中高表达	SlPSY1主导果实成熟阶段类胡萝卜素合成；SlPSY2调控绿色组织光保护相关色素积累；SlPSY3响应胁迫参与根系代谢	［29-30］
桃 Prunus persica	PpPSY1、PpPSY2、PpPSY3	PpPSY1和PpPSY2占据主导地位；PpPSY1在成熟果实中高表达，且在不同遗传背景的桃品种中均表现出果实特异性表达模式；PpPSY2主要在叶片等绿色组织中表达	PpPSY1是果实类胡萝卜素积累的核心调控因子；PpPSY2主要参与光合组织的类胡萝卜素合成，可能与光保护功能相关	［31］
柑橘 Citrus reticulata	CitPSY1、CitPSY2、CitPSY3	CitPSY2在叶片、花器官及果皮中呈低丰度转录	CitPSY1主导果实与光合组织的类胡萝卜素合成；CitPSY3仅存基因组序列，功能尚未解析	［42］
西瓜 Citrullus lanatus	ClPSY1、ClPSY2、ClPSY3	ClPSY1在果肉中显著积累；ClPSY2在叶片中表达量最高；ClPSY3则主要定位于根部	ClPSY1在西瓜果实成熟期间的类胡萝卜素生物合成中起主要作用，与果肉呈色正相关，并受复杂的光照和激素反应性网络调节	［32，43］
辣椒 Capsicum annuum	CaPSY1、CaPSY2.a、CaPSY2.b	CaPSY1在幼果中表达量较低，但在果实发育过程中持续上调表达；CaPSY2.a基因在根、茎、叶、花和果实发育第3和第7个时期中不表达；CaPSY2.b基因在所有组织中均有表达，在茎、叶和果实发育第一个时期表达量相对较高	CaPSY2基因可能补充了CaPSY1的缺失，从而导致黄色辣椒果实呈黄色	［36，44］
西葫芦 Cucurbita pepo	CpPSYA、CpPSYB、CpPSYC	CpPSYA在果实中表达量高，CpPSYB和CpPSYC在叶片中表达量高		［45］
香蕉 Musa ‘Hybrids’	MaPSY1、MaPSY2a、MaPSY2b、MaPSY2c	MaPSY2a、MaPSY2b的表达具有果实特异性		［46］
番红花 Crocus sativus L.	CsPSY1a、CsPSY1b、 CsPSY2、CsPSY3	CsPSY1a和CsPSY1b主要在叶片中表达；CsPSY2转录本主要在柱头中被检测到，而在其他组织中表达却很低；CsPSY3在根尖高表达	CsPSY1b表现出胁迫光调节，CsPSY3的表达与菌根定植和松果内酯的产生有关	［47］
菠菜 Spinacia oleracea	SoPSY1、SoPSY2、 SoPSY3、SoPSY4	SoPSY4在根中高水平表达，显著高于地上部分各组织器官；其余3个SoPSY基因在叶片中高水平表达，营养叶和薹叶中表达最高，其次是薹叶柄、短缩茎和叶柄，在根中表达量最低		［33］
苹果 Malus domestica	MdPSY1、MdPSY2、MdPSY3、MdPSY4、MdPSY5、MdPSY6	仅MdPSY1、MdPSY2和MdPSY4具有酶活性，其中MdPSY1和MdPSY2在叶片和果实成熟过程中高度表达	MdPSY1和MdPSY2与果实中类胡萝卜素合成有关	［34］
甘蓝型油菜 Brassica napus	BnaC.PSY.a BnaA.PSY.b BnaA.PSY.d BnaC.PSY.f BnaA.PSY.d BnaC.PSY.e	BnaC.PSY.a和BnaA.PSY.b在所有组织中均有表达；BnaA.PSY.d和BnaC.PSY.f在富含叶绿体的组织（如叶片）中优先表达；BnaA.PSY.d、BnaC.PSY.e在富含有色体的组织（如花瓣）中优先表达		［35］

图2 通过PSY的多层级调控进行类胡萝卜素的生物强化通过PSY的多层级调控可实现高效类胡萝卜素生物强化，其关键层面包括：转录调控提升PSY表达；表观调控影响染色质开放度；可变剪切产生功能差异的PSY转录本；翻译与转录后修饰调节酶的生成与活性；蛋白稳定性控制PSY的降解速率；亚细胞定位确保其在质体内高效催化，从而多维度促进类胡萝卜素积累。AC：乙酰化；AP2/ERF：AP2/ERF家族转录因子；bZIP：bZIP家族转录因子；Clp：ATP依赖性酪蛋白水解蛋白酶；DML2：DNA去甲基酶2；GGPP：香叶基香叶基焦磷酸；GGPPS：香叶基香叶基焦磷酸合成酶；HDA1：组蛋白去乙酰酶1；MADS： MADS-box家族转录因子；Me：甲基化；MYB：MYB家族转录因子；NAC：NAC家族转录因子；NUDX23：Nudix 水解酶23；OR：橙色伴侣蛋白；PPSR1：质体蛋白感知RING E3连接酶1；PSY：八氢番茄红素合成酶；SGR：叶绿素降解关键蛋白；Ub：泛素；UBC32：泛素结合酶32（该图由BioRender.com创建）

Fig. 2 Biofortification of carotenoids through multi-level regulation mechanisms of PSYThe multi-level regulation of PSY enables efficient carotenoid biofortification, with key aspects including: transcriptional regulation to enhance PSY expression, epigenetic regulation affecting chromatin accessibility, alternative splicing generating functionally distinct PSY transcripts, translational and post-transcriptional modifications regulating enzyme production and activity, control of protein stability to modulate PSY degradation rates, and subcellular localization ensuring its efficient catalysis within plastids. Together, these mechanisms promote carotenoid accumulation in a multidimensional manner. AC: Acetylation; AP2/ERF: APETALA2/ethylene-responsive factor transcription factor family; bZIP: basic leucine zipper transcription factor family; Clp: ATP-dependent caseinolytic protease; DML2: DNA demethylase 2; GGPP: geranylgeranyl pyrophosphate; GGPPS: geranylgeranyl pyrophosphate synthase; HDA1: histone deacetylase 1; MADS: MCM1, AGAMOUS, DEFICIENS, SRF-box transcription factor family; Me: methylation; MYB: myeloblastosis transcription factor family; NAC: NAM, ATAF1/2, CUC2 transcription factor family; NUDX23: nudix hydrolase 23; OR: ORANGE protein; PPSR1: plastid protein sensing RING E3 ligase 1; PSY: phytoene synthase; SGR: STAY-GREEN protein; Ub: ubiquitin; UBC32: ubiquitin-conjugating enzyme 32 (This image was created by BioRender.com)

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