植物避荫综合征的分子机制与育种应用：从光信号感知到作物改良

doi:10.13560/j.cnki.biotech.bull.1985.2025-0985

生物技术通报 ›› 2026, Vol. 42 ›› Issue (6): 149-163.doi: 10.13560/j.cnki.biotech.bull.1985.2025-0985

• 综述与专论 • 上一篇

植物避荫综合征的分子机制与育种应用：从光信号感知到作物改良

张丽¹^,²^,³(), 张玉¹^,³, 刘斌⁴, 聂峰杰¹^,³, 巩檑⁵, 何泽学¹^,³, 罗莉洁¹, 刘丽丽¹, 司怀军²^,⁶()

^1.宁夏农林科学院农业生物技术研究中心，银川 750002
^2.甘肃农业大学农学院，兰州 730070
^3.植物生物育种宁夏重点实验室，银川 750002
^4.中国农业科学院作物科学研究所，北京 100081
^5.宁夏农林科学院固原分院，固原 756000
^6.甘肃农业大学干旱生境作物学国家重点实验室，兰州 730070

收稿日期:2025-09-13 出版日期:2026-06-26 发布日期:2026-07-11
通讯作者: 司怀军，男，博士，教授，研究方向：马铃薯遗传育种；E-mail: hjsi@gsau.edu.cn
作者简介:张丽，女，硕士，副研究员，研究方向：马铃薯生物育种；E-mail: lesley119@163.com
第一联系人：张玉同为本文第一作者
基金资助:
国家自然科学基金项目(32260529);宁夏农业科技自主创新专项(DW-X-2023004);宁夏农业科技自主创新专项(NKYG-25-14);宁夏自然科学基金项目(2024AAC03379)

Molecular Mechanisms and Breeding Applications of Plant Shade Avoidance Response： From Light Signal Perception to Crop Improvement

ZHANG Li¹^,²^,³(), ZHANG Yu¹^,³, LIU Bin⁴, NIE Feng-jie¹^,³, GONG Lei⁵, HE Ze-xue¹^,³, LUO Li-jie¹, LIU Li-li¹, SI Huai-jun²^,⁶()

^1.Research Center of Agricultural Biotechnology, Ningxia Academy of Agriculture and Forestry Sciences, Yinchuan 750002
^2.College of Agriculture, Gansu Agricultural University, Lanzhou 730070
^3.Ningxia Key Laboratory of Plant Biobreeding, Yinchuan 750002
^4.Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, Beijing 100081
^5.Guyuan Branch of Ningxia Academy of Agriculture and Forestry Sciences, Guyuan 756000
^6.State Key Laboratory of Aridl and Crop Science, Gansu Agricultural University, Lanzhou 730070

Received:2025-09-13 Published:2026-06-26 Online:2026-07-11

摘要/Abstract

摘要：

避荫综合征（shade avoidance syndrome, SAS）是植物为了适应竞争性遮荫环境，通过感知红光∶远红光比值（R∶FR）降低及光强减弱等光信号，启动的一系列形态与生理适应性响应。SAS显著影响作物群体的光能利用效率及高密度种植条件下的产量潜力。植物感知环境光质变化主要依赖以光敏色素（phytochromes, PHYs）和隐花色素（cryptochromes, CRYs）为核心的光受体网络。遮荫条件下，这两种主要光受体活性的降低，协同解除了对光敏色素互作因子（phytochrome-interacting factors, PIFs）的抑制，促使其大量积累并激活下游SAS相关基因的转录。作为信号整合的核心枢纽，PIFs一方面正向调控生长素、赤霉素等促生长激素的合成与信号传导，驱动细胞快速伸长；另一方面则拮抗免疫防御信号，体现植物在竞争光照资源时的适应性权衡策略。此外，PIFs还介导SAS与生物钟、碳氮代谢、温度及盐胁迫等信号紧密整合，且调控模块在不同作物中表现出保守性与物种特异性。本文综述了SAS分子机制的研究进展，阐明了PHYs和CRYs感知光质机理，以及PIFs整合激素、表观遗传修饰及生物钟信号的复杂调控网络。重点比较了C₃与C₄作物在光受体系统演化及信号传导上的差异，深入探讨基于基因组编辑和启动子编辑的耐密植分子育种策略。此外，讨论了SAS调控与碳氮代谢及非生物胁迫响应的平衡机制。未来可借助单细胞多组学技术精准解析SAS信号的时空特异性调控网络，并通过智能设计耐密植的理想株型，为作物高产育种提供坚实的理论指导与技术支撑。

关键词: 避荫综合征, 光敏色素, 光敏色素互作因子, 多环境信号整合, 分子育种

Abstract:

Shade avoidance syndrome (SAS) encompasses a series of morphological and physiological adaptive responses initiated by plants to cope with competitive shade environments. This is triggered by perceiving light signals such as a decreased red to far-red light ratio (R:FR) and reduced light intensity. SAS significantly impacts the light utilization efficiency of crop populations and their yield potential under high-density planting conditions. Plants perceive changes in environmental light quality primarily through a photoreceptor network centered on phytochromes (PHYs) and cryptochromes (CRYs). Under shade, the reduced activity of these two main photoreceptors cooperatively relieve inhibition on phytochrome-interacting factors (PIFs), leading to large PIF accumulation and the activation of downstream SAS-related gene transcription. As the core hub of signal integration, PIFs positively regulate the synthesis and signal transduction of growth-promoting hormones like auxins and gibberellins, driving cell elongation, while simultaneously suppressing defense signaling, reflecting the adaptive trade-off strategies of plants when competing for light resources. Furthermore, PIFs also mediate the close integration of SAS with signals such as circadian rhythm, carbon and nitrogen metabolism, temperature and salt stress, with its regulatory modules exhibiting both conservation and species-specificity across different crops. This review summarizes advances in SAS molecular mechanisms, systematically elucidating how PHYs and CRYs perceive light quality, and the complex regulatory networks where PIFs act as central hubs integrating hormonal, epigenetic, and circadian clock signals. It specifically compares differences in photosensory system evolution and signal transduction between C₃ and C₄ crops, and thoroughly discusses molecular breeding strategies, including genome and promoter editing. In addition, the review discusses the balance mechanism between SAS regulation, carbon-nitrogen metabolism, and abiotic stress responses. Future research can leverage single-cell multi-omics technologies to precisely dissect the spatiotemporally specific regulatory networks of SAS signals. This will facilitate the intelligent design of dense planting-tolerant ideotypes, providing robust theoretical guidance and technical support for high-yield crop breeding.

Key words: shade avoidance syndrome, phytochromes, phytochrome-interacting factors, multi-environmental signal integration, molecular breeding

张丽, 张玉, 刘斌, 聂峰杰, 巩檑, 何泽学, 罗莉洁, 刘丽丽, 司怀军. 植物避荫综合征的分子机制与育种应用：从光信号感知到作物改良[J]. 生物技术通报, 2026, 42(6): 149-163.

ZHANG Li, ZHANG Yu, LIU Bin, NIE Feng-jie, GONG Lei, HE Ze-xue, LUO Li-jie, LIU Li-li, SI Huai-jun. Molecular Mechanisms and Breeding Applications of Plant Shade Avoidance Response： From Light Signal Perception to Crop Improvement[J]. Biotechnology Bulletin, 2026, 42(6): 149-163.

图/表 2

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基因类型 Gene type	基因名称 Gene name	作物种类 Crop type	主要功能 Main function	改良方法 Improvement method	改良效果与应用价值 Improvement effect and application value	参考文献 References
光受体	OsPHYB	水稻	感知R∶FR比值变化，调控SAS	过表达	增强稻瘟病抗性，增加分蘖数和籽粒大小，提高单株产量	［113］
	AtPHYB	马铃薯	感知R∶FR比值变化，调控SAS	过表达	在高密度种植条件下显著提高块茎产量和数量，增加分枝数，提高所有冠层叶片的最大光合速率和气孔导度	［114］
	AtPHYA	水稻	感知FR，介导远红光高辐照响应（FR-HIR）	过表达	株高、节间长度和茎粗显著降低，每株穗数增加，谷物产量提高6%-21%	［115］
	ZmPHYC	玉米	调节开花时间和植株高度	过表达	适度降低株高和穗位高度，改善株型	［116］
	GmCRY1s	大豆	感知蓝光强度并抑制PIFs介导的伸长生长	过表达	形成紧凑株型，提高高密度条件下单株产量	［117］
	SlUVR8	番茄	感知紫外辐射调控光形态建成	过表达	增强UV-B耐受性，促进果实叶绿体发育，提升在UV-B环境下抗逆性与营养品质	［118-119］
转录因子	OsPIL1	水稻	调控节间伸长和抗逆性	过表达	增强水稻生长和稻瘟病抗性；异源转化提高柳枝稷生物量和糖化效率，用于纤维生物燃料乙醇生产	［120-123］
	ZmSPL12	玉米	D1的负调节因子，催化GA生物合成的最后一步	过表达	适度降低株高，显著增强抗倒伏性；在高密度种植条件下提高产量	［124］
	OsSPL14	水稻	调控分蘖和穗部发育	分子标记辅助选择	形成“少分蘖、大穗型”株型，已在多个商业品种上实现应用，适合高密度栽培	［125-126］
	ZmLG1 / ZmLG2	玉米	调控叶片-叶鞘边界和叶角	CRISPR编辑	减小叶夹角，改良冠层结构，提高光能利用效率	［127］
	ZmTAC1	玉米	调控分枝/叶片角度	定向选择育种	形成更直立叶片和紧凑株型，实验条件下耐密性增强，产量增加19%-22%	［128］
	GmCOP1b	大豆	调控光形态建成和抗倒伏能力	基因编辑	在密植条件下增强抗倒伏能力，提高农艺性状与产量	［129］
	ZmPIF4/5	玉米	SAS核心转导因子	定点突变	降低SAS反应强度；在高密度种植条件下增产15%-18%	［130］
	ZmDBB2	玉米	与HY5互作抑制GA2ox表达，调控GA稳态及SAS反应	RNAi抑制	调控植株高度，减弱SAS，有利于高密度种植条件下株型优化	［131］
	AtHFR1	小麦	与PIFs形成非DNA结合异源二聚体	异源过表达	降低株高，增加穗状花序数量；在种子萌发期间显著提高了其对渗透胁迫的耐受性，转基因品系籽粒产量比非转基因小麦增加18.2%-48.1%	［132］
	BBX7/8	拟南芥	调控PIF4活性	过表达	优化SAS时间调控；提高环境适应性，参与光信号调控、发育与应答	［73］
	MED25	番茄	Mediator复合体亚基，作为转录共激活因子介导 PIF4对遮荫响应基因的激活	RNAi抑制	抑制遮荫诱导的下胚轴伸长，增强高密度种植适应性	［133］
	SlANT1	番茄	调控花青素积累并响应遮荫	启动子编辑	引发发育、生理和代谢重编程，潜在提升果实营养价值和环境适应性	［102］
激素合成酶/蛋白激酶	AtSOS2	拟南芥	协同调控盐和遮荫双重胁迫响应	过表达	在光照和黑暗条件下均能提高植物对盐胁迫的耐受性	［82］
	SD1（OsGA20ox2）	水稻	调控GA合成与降解	基因编辑	株高降低15%-20%，穗粒数增加；抗倒伏性提高30%-35%，收获指数改善	［134］
	ZmGA20ox3/ZmGA20ox5	玉米	调控GA合成与降解	基因编辑	产生了半矮化玉米植株，对耐高密度种植的品种选育具有应用价值	［135-136］
	Rht-1	小麦	编码DELLA蛋白的Rht-B1b和Rht-D1b等位基因，GA信号负调控因子，与PIFs拮抗抑制SAS	基因编辑	增强GA敏感性，调节植株高度，通过调控DELLA-PIF平衡优化群体结构	［137-138］
生物钟组分	GmLHY1a/1b	大豆	调控昼夜节律	CRISPR编辑	解除耐旱抑制，显著提高干旱胁迫存活率	［139］

基因类型 Gene type	基因名称 Gene name	作物种类 Crop type	主要功能 Main function	改良方法 Improvement method	改良效果与应用价值 Improvement effect and application value	参考文献 References
光受体	OsPHYB	水稻	感知R∶FR比值变化，调控SAS	过表达	增强稻瘟病抗性，增加分蘖数和籽粒大小，提高单株产量	［113］
	AtPHYB	马铃薯	感知R∶FR比值变化，调控SAS	过表达	在高密度种植条件下显著提高块茎产量和数量，增加分枝数，提高所有冠层叶片的最大光合速率和气孔导度	［114］
	AtPHYA	水稻	感知FR，介导远红光高辐照响应（FR-HIR）	过表达	株高、节间长度和茎粗显著降低，每株穗数增加，谷物产量提高6%-21%	［115］
	ZmPHYC	玉米	调节开花时间和植株高度	过表达	适度降低株高和穗位高度，改善株型	［116］
	GmCRY1s	大豆	感知蓝光强度并抑制PIFs介导的伸长生长	过表达	形成紧凑株型，提高高密度条件下单株产量	［117］
	SlUVR8	番茄	感知紫外辐射调控光形态建成	过表达	增强UV-B耐受性，促进果实叶绿体发育，提升在UV-B环境下抗逆性与营养品质	［118-119］
转录因子	OsPIL1	水稻	调控节间伸长和抗逆性	过表达	增强水稻生长和稻瘟病抗性；异源转化提高柳枝稷生物量和糖化效率，用于纤维生物燃料乙醇生产	［120-123］
	ZmSPL12	玉米	D1的负调节因子，催化GA生物合成的最后一步	过表达	适度降低株高，显著增强抗倒伏性；在高密度种植条件下提高产量	［124］
	OsSPL14	水稻	调控分蘖和穗部发育	分子标记辅助选择	形成“少分蘖、大穗型”株型，已在多个商业品种上实现应用，适合高密度栽培	［125-126］
	ZmLG1 / ZmLG2	玉米	调控叶片-叶鞘边界和叶角	CRISPR编辑	减小叶夹角，改良冠层结构，提高光能利用效率	［127］
	ZmTAC1	玉米	调控分枝/叶片角度	定向选择育种	形成更直立叶片和紧凑株型，实验条件下耐密性增强，产量增加19%-22%	［128］
	GmCOP1b	大豆	调控光形态建成和抗倒伏能力	基因编辑	在密植条件下增强抗倒伏能力，提高农艺性状与产量	［129］
	ZmPIF4/5	玉米	SAS核心转导因子	定点突变	降低SAS反应强度；在高密度种植条件下增产15%-18%	［130］
	ZmDBB2	玉米	与HY5互作抑制GA2ox表达，调控GA稳态及SAS反应	RNAi抑制	调控植株高度，减弱SAS，有利于高密度种植条件下株型优化	［131］
	AtHFR1	小麦	与PIFs形成非DNA结合异源二聚体	异源过表达	降低株高，增加穗状花序数量；在种子萌发期间显著提高了其对渗透胁迫的耐受性，转基因品系籽粒产量比非转基因小麦增加18.2%-48.1%	［132］
	BBX7/8	拟南芥	调控PIF4活性	过表达	优化SAS时间调控；提高环境适应性，参与光信号调控、发育与应答	［73］
	MED25	番茄	Mediator复合体亚基，作为转录共激活因子介导 PIF4对遮荫响应基因的激活	RNAi抑制	抑制遮荫诱导的下胚轴伸长，增强高密度种植适应性	［133］
	SlANT1	番茄	调控花青素积累并响应遮荫	启动子编辑	引发发育、生理和代谢重编程，潜在提升果实营养价值和环境适应性	［102］
激素合成酶/蛋白激酶	AtSOS2	拟南芥	协同调控盐和遮荫双重胁迫响应	过表达	在光照和黑暗条件下均能提高植物对盐胁迫的耐受性	［82］
	SD1（OsGA20ox2）	水稻	调控GA合成与降解	基因编辑	株高降低15%-20%，穗粒数增加；抗倒伏性提高30%-35%，收获指数改善	［134］
	ZmGA20ox3/ZmGA20ox5	玉米	调控GA合成与降解	基因编辑	产生了半矮化玉米植株，对耐高密度种植的品种选育具有应用价值	［135-136］
	Rht-1	小麦	编码DELLA蛋白的Rht-B1b和Rht-D1b等位基因，GA信号负调控因子，与PIFs拮抗抑制SAS	基因编辑	增强GA敏感性，调节植株高度，通过调控DELLA-PIF平衡优化群体结构	［137-138］
生物钟组分	GmLHY1a/1b	大豆	调控昼夜节律	CRISPR编辑	解除耐旱抑制，显著提高干旱胁迫存活率	［139］

植物避荫综合征的分子机制与育种应用：从光信号感知到作物改良

Molecular Mechanisms and Breeding Applications of Plant Shade Avoidance Response： From Light Signal Perception to Crop Improvement

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