光信号调控园艺作物果实发育的研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2025-1286

摘要/Abstract

摘要：

光是调控园艺作物果实发育的核心环境信号。目前，光信号调控果实发育的研究已经系统揭示了光受体（光敏色素、隐花色素、UVR8）通过核心转录因子（HY5、PIFs、COP1、BBX等）网络介导的分子通路，以及光质、光强、光周期对果实色泽、糖酸比、香气和重量等品质性状的特异性调控机制。本文综述了光信号在果实发育关键阶段的调控作用，重点涵盖其对果实早期发育过程的调控，以及在果实成熟进程中对果实品质形成的调控机制；同时重点总结了光信号通过调控乙烯、脱落酸和茉莉酸的相关通路，并与生长素和赤霉素等激素形成协同与拮抗调控网络，从而协调果实成熟进程与品质形成的分子与生理机制；讨论了不同光质、光强与光周期对果实发育的调控效应具有物种特异性与剂量依赖性。当前研究虽已揭示光信号调控果实发育的部分核心分子机制，但对于果实不同发育阶段的分子调控开关、多种光因子之间的协同量化模型、非模式园艺作物的独特机制及光信号与温度、水分和养分等其他环境因子协同调控等方面仍缺乏系统解析。未来可结合基因编辑与智能光控等技术，深入解析光信号整合多环境因子的分子机制，为设施园艺果实品质精准调控及作物遗传改良提供理论支撑，推动光调控技术标准化与规模化应用于优质果实生产。

关键词: 光信号, 光受体, 果实发育, 果实品质, 转录因子, 激素互作, 基因编辑, 智能光控

Abstract:

Light is a core environmental signal regulating fruit development in horticultural crops. Research of light regulating fruit development has revealed molecular pathways of photoreceptors (phytochromes, cryptochromes, and UVR8) mediated by the network of core transcription factors (HY5, PIFs, COP1, BBX, etc.), and mechanism of light quality, light intensity, and photoperiod specifically regulating traits such as color, sugar-acid balance, aroma, and weight. This review summarizes the regulatory roles of light signaling at key stages of fruit development, with a particular focus on its regulation of early fruit developmental processes and its control of fruit quality formation during ripening. It further highlights how light signaling regulates ethylene, abscisic acid, and jasmonic acid pathways and forms synergistic and antagonistic networks with hormones such as auxin and gibberellin, thereby coordinating the molecular and physiological mechanisms underlying fruit ripening and quality formation. It also notes that regulating effects of different light-quality, light intensity and photoperiod on fruit development are species-specific and dose-dependent. Although key molecular mechanisms of light signal regulating partially have been uncovered, it is still lack for system analysis on stage-specific molecular switches, quantitative models of multi-factor light interactions, unique mechanisms in non-model horticultural crops, and synergistical regulation of light with temperature, moisture, nutrients and other environmental factors. In future, integrating gene editing with intelligent light-control technologies, the molecular mechanism of light integrates multiple environmental signals can be dissected, which provides theoretical support for the precise regulation of fruit quality and crop genetic improvement in protected horticulture, and promote the standardization and large-scale application of light regulation technology in the production of high-quality fruits.

Key words: light signaling, photoreceptors, fruit development, fruit quality, transcription factors, hormonal crosstalk, genome editing, smart light regulation

马世杰, 李铮, 李蔚, 郭仰东, 张娜. 光信号调控园艺作物果实发育的研究进展[J]. 生物技术通报, doi: 10.13560/j.cnki.biotech.bull.1985.2025-1286.

MA Shi-jie, LI Zheng, LI Wei, GUO Yang-dong, ZHANG Na. Research Progress in Light Signaling Regulation of Fruit Development in Horticultural Crops[J]. Biotechnology Bulletin, doi: 10.13560/j.cnki.biotech.bull.1985.2025-1286.

图/表 3

图1 光受体介导的果实花青素与类胡萝卜素合成途径的调控机制光受体（PhyA/PhyB/PhyE、cry1/2、UVR8）感知不同波长的光后，通过信号传导调控果实花青素与类胡萝卜素合成途径。光照激活的光受体诱导光敏色素互作因子（PIF1/PIF3/PIF4/PIF5）降解，解除其对下游合成基因的抑制；同时通过COP1-HY5-BBX20/21/22信号轴正向调控花青素合成途径（PAL→CHS→......→UFGT）及类胡萝卜素合成途径（PSY→PDS→...→番茄红素/胡萝卜素）。箭头表示正调控，钝头箭头表示负调控

Fig. 1 Photoreceptor-mediated regulating mechanism of anthocyanin and carotenoid biosynthetic pathways in fruitsPhotoreceptors (PhyA/PhyB/PhyE, CRY1/2, and UVR8) perceive light of different wavelengths and regulate anthocyanin and carotenoid biosynthetic pathways in fruits through downstream signaling cascades. Light-activated photoreceptors induce the degradation of phytochrome‐interacting factors (PIF1, PIF3, PIF4, and PIF5), thereby releasing their repression on downstream biosynthetic genes; meanwhile, the COP1-HY5-BBX20/21/22 signaling module positively regulates the anthocyanin biosynthetic pathway (PAL→CHS→…→UFGT) and the carotenoid biosynthetic pathway (PSY→PDS→……→lycopene/carotene). Arrows indicate positive regulation, whereas blunt-ended arrows indicate negative regulation

图2 光信号与激素信号协同调控的分子网络该图由4个模块组成，分别表示光信号对乙烯（ETH）、脱落酸（ABA）、茉莉酸（JA）、生长素（IAA）和赤霉素（GA）的调控，每个模块用A-D标注，箭头表示正调控，钝头箭头表示负调控。A：光受体（PhyB、CRY1）感知光信号（红光、蓝光、白光），通过EIN3、HY5等元件调控乙烯的生物合成与信号；B：光信号通过ABI1/2、NCED3、ABI5、PIFs等元件调控ABA合成及信号；C：光信号通过LOX、AOC、ST2a等元件影响茉莉酸生物合成；D：光信号通过YUC8/9、GA3ox1/2、IAA19、GA2ox等元件调节生长素和赤霉素的生物合成与信号传导

Fig. 2 Molecular network underlying the crosstalk between light signaling and hormone signalingThe figure consists of four modules (A-D) representing the regulation of ethylene (ETH), abscisic acid (ABA), jasmonic acid (JA), auxin (IAA), and gibberellin (GA) by light signaling. Arrows indicate positive regulation, and blunt-ended arrows indicate negative regulation. A: Photoreceptors (PhyB and CRY1) perceive light signals (red, blue, and white) and regulate ethylene biosynthesis and signaling via EIN3, HY5, and related components. B: Light signaling regulates ABA biosynthesis and signaling through ABI1/2, NCED3, ABI5, and PIFs. C: Light signaling influences jasmonic acid biosynthesis via LOX, AOC, and ST2a. D: Light signaling modulates auxin and gibberellin biosynthesis and signaling through YUC8/9, GA3ox1/2, IAA19, and GA2ox

表1 不同光质对不同果实商品品质的调控效应

Table 1 Regulatory effects of different light spectra on commercial quality traits of fruits

光质 Light quality	园艺作物 Horticultural crops	效果 Effect	参考文献Reference
红光 Red light	草莓 Fragaria × ananassa Duch， ‘Benihoppe’	果实亮度显著提升；维生素C含量显著增加；货架期延长	［95］
	辣椒 Capsicum annuum L.	维生素C含量显著增加；货架期延长	［96］
	金桔 F. crassifolia Swingle	加速果实褪绿，果实色泽均匀性得到改善	［79］
	西瓜 Citrullus lanatus Thunb	瓜瓤着色提前5~10天，采收时间显著提前	［97］
	甜瓜 Cucumis melo L.	酯类挥发性风味化合物含量显著提高；果实成熟时间显著缩短；可溶性固形物含量显著提升	［98］
	柑橘 Citrus sinensis L. Osbeck	类胡萝卜素含量显著提高	［99］
	番茄 Solanum lycopersicum L.	果实成熟时间显著缩短；可溶性固形物含量显著提高；	［81］
	香蕉 Musa nana Lour	加速果实软化；提前达到糖度峰值；果实成熟时间显著提前	［100］
	芒果 Mangifera indica L.	果实转色更快；可溶性糖积累高峰提前；后熟软化进程加快	［83］
	香梨 Pyrus sinkiangensis Yu	果实转色加快；后熟软化进程加快	［83］
	冬枣 Ziziphus jujuba Mill.	果实转色加快；后熟软化进程加快	［83］
	青皮桔 Citrus reticulata Blanco	果实转色加快；后熟软化进程加快	［83］
	羊角蜜甜瓜 Cucumis melo L.	货架期出现“褪白转绿”现象	［83］
远红光 Far-red light	番茄 Solanum lycopersicum L.	可溶性固形物含量积累显著提高；维生素含量显著提高；果实大小略有增加；部分品种果实产量略有增加	［84］
远红光 Far-red light	辣椒 Capsicum annuum L.	果实产量显著减少；果实变小；	［86］
蓝光 Blue light	油桃 Prunus persica var. nectarine	类胡萝卜素含量显著提高	［89］
	柑橘 Citrus sinensis L. Osbeck	果实对绿霉病抗性提高，果实色泽更加均匀	［101］
	梨 Pyrus ussuriensis	延缓果实成熟	［87］
	杨梅 Myrica rubra Sieb. and Zucc. cv. Biqi	花青素含量显著提高	［88］
	辣椒 Capsicum annuum L.	辣椒素含量显著提高	［102］
紫外光 UV-B	苹果 Malus × domestica	单果重显著提高；果实成熟进程加快	［103］
	桃 Prunus persica L. Batsch	萜类物质含量显著增加	［104］
	蓝莓 Vaccinium spp.	花青素含量显著增加	［91］
	葡萄 Vitis vinifera L.	多酚类物质显著增加	［105］
	山竹 Garcinia mangostana L.	货架期延长	［92］

表1 不同光质对不同果实商品品质的调控效应

Table 1 Regulatory effects of different light spectra on commercial quality traits of fruits

光质 Light quality	园艺作物 Horticultural crops	效果 Effect	参考文献Reference
红光 Red light	草莓 Fragaria × ananassa Duch， ‘Benihoppe’	果实亮度显著提升；维生素C含量显著增加；货架期延长	［95］
	辣椒 Capsicum annuum L.	维生素C含量显著增加；货架期延长	［96］
	金桔 F. crassifolia Swingle	加速果实褪绿，果实色泽均匀性得到改善	［79］
	西瓜 Citrullus lanatus Thunb	瓜瓤着色提前5~10天，采收时间显著提前	［97］
	甜瓜 Cucumis melo L.	酯类挥发性风味化合物含量显著提高；果实成熟时间显著缩短；可溶性固形物含量显著提升	［98］
	柑橘 Citrus sinensis L. Osbeck	类胡萝卜素含量显著提高	［99］
	番茄 Solanum lycopersicum L.	果实成熟时间显著缩短；可溶性固形物含量显著提高；	［81］
	香蕉 Musa nana Lour	加速果实软化；提前达到糖度峰值；果实成熟时间显著提前	［100］
	芒果 Mangifera indica L.	果实转色更快；可溶性糖积累高峰提前；后熟软化进程加快	［83］
	香梨 Pyrus sinkiangensis Yu	果实转色加快；后熟软化进程加快	［83］
	冬枣 Ziziphus jujuba Mill.	果实转色加快；后熟软化进程加快	［83］
	青皮桔 Citrus reticulata Blanco	果实转色加快；后熟软化进程加快	［83］
	羊角蜜甜瓜 Cucumis melo L.	货架期出现“褪白转绿”现象	［83］
远红光 Far-red light	番茄 Solanum lycopersicum L.	可溶性固形物含量积累显著提高；维生素含量显著提高；果实大小略有增加；部分品种果实产量略有增加	［84］
远红光 Far-red light	辣椒 Capsicum annuum L.	果实产量显著减少；果实变小；	［86］
蓝光 Blue light	油桃 Prunus persica var. nectarine	类胡萝卜素含量显著提高	［89］
	柑橘 Citrus sinensis L. Osbeck	果实对绿霉病抗性提高，果实色泽更加均匀	［101］
	梨 Pyrus ussuriensis	延缓果实成熟	［87］
	杨梅 Myrica rubra Sieb. and Zucc. cv. Biqi	花青素含量显著提高	［88］
	辣椒 Capsicum annuum L.	辣椒素含量显著提高	［102］
紫外光 UV-B	苹果 Malus × domestica	单果重显著提高；果实成熟进程加快	［103］
	桃 Prunus persica L. Batsch	萜类物质含量显著增加	［104］
	蓝莓 Vaccinium spp.	花青素含量显著增加	［91］
	葡萄 Vitis vinifera L.	多酚类物质显著增加	［105］
	山竹 Garcinia mangostana L.	货架期延长	［92］

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