• 综述与专论 • 下一篇
崔之瀚1,2(
), 魏庆镇2, 胡娜2, 包崇来2(
), 王华森1(
)
收稿日期:2025-11-14
出版日期:2026-03-02
通讯作者:
包崇来,男,硕士,研究员,研究方向 :蔬菜遗传育种;E-mail: baocl@zaas.ac.cn作者简介:崔之瀚,男,硕士研究生,研究方向 :蔬菜遗传育种;E-mail: 17306395131@163.com
基金资助:
CUI Zhi-han1,2(
), WEI Qing-zhen2, HU Na2, BAO Chong-lai2(
), WANG Hua-sen1(
)
Received:2025-11-14
Published:2026-03-02
摘要:
植物特异性IQ67结构域(IQ67-domain, IQD)基因家族作为编码钙调蛋白结合蛋白与钙信号支架蛋白的关键因子,构成了植物钙信号调控网络的核心枢纽。该家族成员具备独特的结构特征,即N末端IQ67保守域特异性募集,并通过C末端DUF4005结构域锚定微管。在园艺作物中,IQD家族的核心功能多聚焦于果实形态建成,其通过改变微管排列方向决定细胞分裂面,直接控制果形;同时作为多维信号整合节点,深度介导钙信号与生长素、赤霉素、细胞分裂素及脱落酸等激素通路,以及病原菌侵染、盐、旱胁迫等逆境响应的交互作用。尽管拟南芥(Arabidopsis thaliana)中IQD的分子互作与细胞调控网络已获系统解析,但针对园艺作物的研究在机制解析深度与系统性上仍存在明显差距,番茄(Solanum lycopersicum)、黄瓜(Cucumis sativus)等园艺作物虽已鉴定出多个关键IQD基因,但多数研究仍局限于基因与表型的关联分析,对细胞尺度微管动力学及多信号整合机制的解析相对不足。本文在系统梳理主要园艺作物IQD基因研究进展的基础上,重点整合其在果实形态调控、细胞骨架重塑及信号级联中的核心分子机制,规范基因命名,提出当前研究存在的局限性。未来研究需聚焦于IQD蛋白结合钙信号、激素信号、微管骨架等方面的分子功能解析,以期为完善园艺作物生长发育调控网络、推动优质抗逆品种的分子设计育种提供理论参考。
崔之瀚, 魏庆镇, 胡娜, 包崇来, 王华森. 园艺作物IQD基因研究进展[J]. 生物技术通报, doi: 10.13560/j.cnki.biotech.bull.1985.2025-1232.
CUI Zhi-han, WEI Qing-zhen, HU Na, BAO Chong-lai, WANG Hua-sen. Research Progress in IQD Genes in Horticultural Crops[J]. Biotechnology Bulletin, doi: 10.13560/j.cnki.biotech.bull.1985.2025-1232.
图2 园艺作物IQD基因家族调控果实形态网络模型IQD基因通过两条通路调控果实形态建成:MAPs通路:IQD蛋白可与SlMAP70/SlMAP65-1蛋白互作,改变微管皮层(CMT)的排列方向,进而决定细胞的扩张方向;激素通路:钙调蛋白-IQD复合物调控钙离子信号传导及生长素(IAA)/赤霉素(GA3)/脱落酸(ABA)的含量水平,以此促进细胞纵向分裂与果实坐果。表型效应:IQD基因低表达会使果实形态更短粗,而高表达则可促进果实伸长(该现象在番茄、黄瓜中均有验证)
Fig. 2 Network model of the IQD gene family regulating fruit morphology in horticultural cropsThe IQD gene modulates fruit morphology via two synergistic pathways: MAPs pathway: IQD interacts with SlMAP70/SlMAP65-1 to alter cortical microtubule (CMT) orientation, directing the axis of cell expansion.Hormonal pathway: The CaM-IQD complex modulatesCa2+ signaling and IAA/GA3/ABA levels, promoting longitudinal cell division and fruit set. Phenotypic outcomes: Low IQD expression results in rounder fruits, while high expression promotes fruit elongation (e.g., in tomato and cucumber)
基因名称 Gene name | 基因ID Gene ID | 物种 Species | 基因功能 Gene function | 参考文献 Reference |
|---|---|---|---|---|
| AtIQD1 | At3g09710 | 拟南芥 Arabidopsis thaliana | 参与激素信号通路调控,调控植物防御反应; | [ |
| 参与钙信号通路调控,调控植物防御反应,连接钙信号与基因表达; | [ | |||
| 调控芥子油苷生物合成、整合钙信号与防御响应,增强抗虫与抗病性,与维管组织中高表达; | [ | |||
| 调控细胞内货物运输,与驱动蛋白轻链相关蛋白KLCR1发生互作 | [ | |||
| AtIQD20 | At3g51380 | 拟南芥 Arabidopsis thaliana | 作为钙调蛋白结合靶标 | [ |
| AtIQD13 | At3g59690 | 拟南芥 Arabidopsis thaliana | 调控木质部血管次生细胞壁形状,调节微管动力学,协同强化微管‒质膜连接 | [ |
| AtIQD2 | At5g03040 | 拟南芥 Arabidopsis thaliana | 介导细胞骨架桥接与网络协调,维持EPCS结构与ER形态,调控细胞形态 | [ |
| AtIQD5 | At3g22190 | 拟南芥 Arabidopsis thaliana | 调控皮质微管(MT)动态与组装,影响微管组织与细胞形态建成,支撑细胞生长与形状控制; | [ |
| 调控路面细胞(PC)形态,介导钙信号与微管阵列,作为植物特异性微管相关蛋白发挥作用; | [ | |||
| 微管结合的结构基础,调控植物生长发育与细胞形态 | [ | |||
| AtIQD11 | At5g13460 | 拟南芥 Arabidopsis thaliana | 对植物表型与细胞形态的调控,对MT排列方向的影响 | [ |
| AtIQD14 | At2g43680 | 拟南芥 Arabidopsis thaliana | 对植物表型与细胞形态的调控,对MT排列方向的影响 | [ |
| AtIQD16 | At4g10640 | 拟南芥 Arabidopsis thaliana | 调控皮质微管排序与细胞扩张,参与乙烯反应与差异生长调控,分子互作与募集功能; | [ |
| 作为植物特异性微管相关蛋白发挥作用\调控植物发育与细胞形态建成; | [ | |||
| 调控皮质微管的组织与方向,调控植物气生器官形态与细胞形态; | [ | |||
| 调控下胚轴伸长,与微管切断酶KTN1互作 | [ | |||
| AtIQD21 | At3g49260 | 拟南芥 Arabidopsis thaliana | 铺板细胞凹陷形成的关键调控,铺板细胞形态建成的微管介导作用 | [ |
| AtIQD26 | At3g16490 | 拟南芥 Arabidopsis thaliana | 钙调蛋白结合功能,调控逆境响应 | [ |
| AtIQD22 | At4g23060 | 拟南芥 Arabidopsis thaliana | 调控植物对缺氧的耐受性,与ADH1关联激活无氧呼吸 | [ |
| AtIQD15 | At3g49380 | 拟南芥 Arabidopsis thaliana | 增强微管稳定性并促进横向平行排列,可响应生长素诱导的钙瞬变 | [ |
| AtIQD17 | At4g00820 | 拟南芥 Arabidopsis thaliana | 受生长素‒细胞分裂素协同调控,影响铺板细胞“叶”状突起形成 | [ |
| AtIQD18 | At1g01110 | 拟南芥 Arabidopsis thaliana | 促进微管纵向排列并增强稳定性,介导生长素调控根伸长 | [ |
| SlSUN1/SlIQD12 | Solyc10g079240 | 番茄 Solanum lycopersicum | 调控果实伸长; | [ |
| 调控果实发育,参与植物激素信号通路; | [ | |||
| 调控营养器官的形态,调控种子发育,调控番茄果实形状; | [ | |||
| 调控番茄果实早期形态,调控核心代谢通路; | [ | |||
| 调控果实伸长,调控营养器官与生殖器官形态,与ovate、fs8.1存在协同作用; | [ | |||
| 调控子房与果实伸长,影响基因表达; | [ | |||
| 调控番茄果实形状 | [ | |||
| SlSUN24 | Solyc08g083240.2.1 | 番茄 Solanum lycopersicum | 激素网络调控,诱导种子萌发,钙调蛋白互作 | [ |
| SlSUN18/SlIQD26 | Solyc06g066430.2.1 | 番茄 Solanum lycopersicum | 调控细胞分裂模式,控制果实形状,调控微管动态骨架 | [ |
| SlSUN10/SlIQD21 | Solyc03g083100.2.1 | 番茄 Solanum lycopersicum | 调控细胞形态与微管动态,调控发育早期的形态,协同调控果实形状 | [ |
| CsSUN2 | Csa1G575000 | 黄瓜 Cucumis sativus | 调控果实形态 | [ |
| CsSUN5 | CsaV3_2G029380 | 黄瓜 Cucumis sativus | 对逆境的响应 | [ |
| CsSUN | Csa1G575000、Cucsa.288770 | 黄瓜 Cucumis sativus | 钙信号‒细胞骨架‒调控通路 | [ |
| CsSUN | Csa1G575000,Cucsa.288770 | 黄瓜 Cucumis sativus | 与CsLNG协同调控黄瓜果实形状,正向调控黄瓜果实纵向伸长,响应激素信号通路 | [ |
| CsSUN | CsaV3_1G039870 | 黄瓜 Cucumis sativus | 调控果实纵向伸长,调控细胞分裂方向,影响激素相关基因表达 | [ |
| CsSUN | CsaV3_1G039870 | 黄瓜 Cucumis sativus | 异源表达调控果实伸长,调控细胞分裂和大小 | [ |
| CsSUN | CsaV3_1G039870 | 黄瓜 Cucumis sativus | 蛋白稳定性受CsTON2正向调控,通过促进细胞纵向分裂,抑制横向分裂来正向调控果实伸长 | [ |
| Smechr0301963 | Smechr0301963 | 茄子 Solanum melongena L. | 潜在预测调控果实的长度 | [ |
| SmeFL | Smechr0302217 | 茄子 Solanum melongena L. | 初步定位SUN基因,预测调控茄子果实长度,调控微纤丝的取向来影响茄子细胞的扩张方向 | [ |
| CaIQD1 | 辣椒 Capsicum annuum | 调控多蛋白互作网络,激素信号的交叉调控 | [ | |
| BrIQD5 | Bra023414 | 白菜 Brassica rapa | 微管动态调控与细胞结构重塑,脱落酸信号通路的协同调控,干旱响应的分子开关 | [ |
| CmSUN23-24、CmSUN25-26-27c | MELO3C006884、MELO3C013004 | 甜瓜 Cucumis melo | 调控果实伸长,导致甜瓜果实形状变异 | [ |
| CmSUN7、CmSUN12 | MELO3C013004、MELO3C006884 | 甜瓜 Cucumis melo | 果实伸长调控,信号通路参与 | [ |
| CmIQD1、CmIQD31 | MELO3C025505、MELO3C002201 | 甜瓜 Cucumis melo | 调控果实形态、叶片形态,调控果实重量 | [ |
| MdIQD17、MdIQD18 | MD07G1178400、MD07G1265200 | 苹果 Malus domestica | 果实参与冷胁迫的适应性调控 | [ |
| PeIQD13、PeIQD9 | POPTR_0005s05310、POPTR_0003s04240 | 毛竹 Phyllostachys edulis | 调节干旱胁迫、MeJA胁迫 | [ |
| ClIQD24 | Cla021081 | 西瓜 Citrullus lanatus | 调控果实生长发育 | [ |
表1 IQD基因家族生物学功能研究进展汇总表
Table 1 Summary table of research progress on the biological functions of the IQD gene family
基因名称 Gene name | 基因ID Gene ID | 物种 Species | 基因功能 Gene function | 参考文献 Reference |
|---|---|---|---|---|
| AtIQD1 | At3g09710 | 拟南芥 Arabidopsis thaliana | 参与激素信号通路调控,调控植物防御反应; | [ |
| 参与钙信号通路调控,调控植物防御反应,连接钙信号与基因表达; | [ | |||
| 调控芥子油苷生物合成、整合钙信号与防御响应,增强抗虫与抗病性,与维管组织中高表达; | [ | |||
| 调控细胞内货物运输,与驱动蛋白轻链相关蛋白KLCR1发生互作 | [ | |||
| AtIQD20 | At3g51380 | 拟南芥 Arabidopsis thaliana | 作为钙调蛋白结合靶标 | [ |
| AtIQD13 | At3g59690 | 拟南芥 Arabidopsis thaliana | 调控木质部血管次生细胞壁形状,调节微管动力学,协同强化微管‒质膜连接 | [ |
| AtIQD2 | At5g03040 | 拟南芥 Arabidopsis thaliana | 介导细胞骨架桥接与网络协调,维持EPCS结构与ER形态,调控细胞形态 | [ |
| AtIQD5 | At3g22190 | 拟南芥 Arabidopsis thaliana | 调控皮质微管(MT)动态与组装,影响微管组织与细胞形态建成,支撑细胞生长与形状控制; | [ |
| 调控路面细胞(PC)形态,介导钙信号与微管阵列,作为植物特异性微管相关蛋白发挥作用; | [ | |||
| 微管结合的结构基础,调控植物生长发育与细胞形态 | [ | |||
| AtIQD11 | At5g13460 | 拟南芥 Arabidopsis thaliana | 对植物表型与细胞形态的调控,对MT排列方向的影响 | [ |
| AtIQD14 | At2g43680 | 拟南芥 Arabidopsis thaliana | 对植物表型与细胞形态的调控,对MT排列方向的影响 | [ |
| AtIQD16 | At4g10640 | 拟南芥 Arabidopsis thaliana | 调控皮质微管排序与细胞扩张,参与乙烯反应与差异生长调控,分子互作与募集功能; | [ |
| 作为植物特异性微管相关蛋白发挥作用\调控植物发育与细胞形态建成; | [ | |||
| 调控皮质微管的组织与方向,调控植物气生器官形态与细胞形态; | [ | |||
| 调控下胚轴伸长,与微管切断酶KTN1互作 | [ | |||
| AtIQD21 | At3g49260 | 拟南芥 Arabidopsis thaliana | 铺板细胞凹陷形成的关键调控,铺板细胞形态建成的微管介导作用 | [ |
| AtIQD26 | At3g16490 | 拟南芥 Arabidopsis thaliana | 钙调蛋白结合功能,调控逆境响应 | [ |
| AtIQD22 | At4g23060 | 拟南芥 Arabidopsis thaliana | 调控植物对缺氧的耐受性,与ADH1关联激活无氧呼吸 | [ |
| AtIQD15 | At3g49380 | 拟南芥 Arabidopsis thaliana | 增强微管稳定性并促进横向平行排列,可响应生长素诱导的钙瞬变 | [ |
| AtIQD17 | At4g00820 | 拟南芥 Arabidopsis thaliana | 受生长素‒细胞分裂素协同调控,影响铺板细胞“叶”状突起形成 | [ |
| AtIQD18 | At1g01110 | 拟南芥 Arabidopsis thaliana | 促进微管纵向排列并增强稳定性,介导生长素调控根伸长 | [ |
| SlSUN1/SlIQD12 | Solyc10g079240 | 番茄 Solanum lycopersicum | 调控果实伸长; | [ |
| 调控果实发育,参与植物激素信号通路; | [ | |||
| 调控营养器官的形态,调控种子发育,调控番茄果实形状; | [ | |||
| 调控番茄果实早期形态,调控核心代谢通路; | [ | |||
| 调控果实伸长,调控营养器官与生殖器官形态,与ovate、fs8.1存在协同作用; | [ | |||
| 调控子房与果实伸长,影响基因表达; | [ | |||
| 调控番茄果实形状 | [ | |||
| SlSUN24 | Solyc08g083240.2.1 | 番茄 Solanum lycopersicum | 激素网络调控,诱导种子萌发,钙调蛋白互作 | [ |
| SlSUN18/SlIQD26 | Solyc06g066430.2.1 | 番茄 Solanum lycopersicum | 调控细胞分裂模式,控制果实形状,调控微管动态骨架 | [ |
| SlSUN10/SlIQD21 | Solyc03g083100.2.1 | 番茄 Solanum lycopersicum | 调控细胞形态与微管动态,调控发育早期的形态,协同调控果实形状 | [ |
| CsSUN2 | Csa1G575000 | 黄瓜 Cucumis sativus | 调控果实形态 | [ |
| CsSUN5 | CsaV3_2G029380 | 黄瓜 Cucumis sativus | 对逆境的响应 | [ |
| CsSUN | Csa1G575000、Cucsa.288770 | 黄瓜 Cucumis sativus | 钙信号‒细胞骨架‒调控通路 | [ |
| CsSUN | Csa1G575000,Cucsa.288770 | 黄瓜 Cucumis sativus | 与CsLNG协同调控黄瓜果实形状,正向调控黄瓜果实纵向伸长,响应激素信号通路 | [ |
| CsSUN | CsaV3_1G039870 | 黄瓜 Cucumis sativus | 调控果实纵向伸长,调控细胞分裂方向,影响激素相关基因表达 | [ |
| CsSUN | CsaV3_1G039870 | 黄瓜 Cucumis sativus | 异源表达调控果实伸长,调控细胞分裂和大小 | [ |
| CsSUN | CsaV3_1G039870 | 黄瓜 Cucumis sativus | 蛋白稳定性受CsTON2正向调控,通过促进细胞纵向分裂,抑制横向分裂来正向调控果实伸长 | [ |
| Smechr0301963 | Smechr0301963 | 茄子 Solanum melongena L. | 潜在预测调控果实的长度 | [ |
| SmeFL | Smechr0302217 | 茄子 Solanum melongena L. | 初步定位SUN基因,预测调控茄子果实长度,调控微纤丝的取向来影响茄子细胞的扩张方向 | [ |
| CaIQD1 | 辣椒 Capsicum annuum | 调控多蛋白互作网络,激素信号的交叉调控 | [ | |
| BrIQD5 | Bra023414 | 白菜 Brassica rapa | 微管动态调控与细胞结构重塑,脱落酸信号通路的协同调控,干旱响应的分子开关 | [ |
| CmSUN23-24、CmSUN25-26-27c | MELO3C006884、MELO3C013004 | 甜瓜 Cucumis melo | 调控果实伸长,导致甜瓜果实形状变异 | [ |
| CmSUN7、CmSUN12 | MELO3C013004、MELO3C006884 | 甜瓜 Cucumis melo | 果实伸长调控,信号通路参与 | [ |
| CmIQD1、CmIQD31 | MELO3C025505、MELO3C002201 | 甜瓜 Cucumis melo | 调控果实形态、叶片形态,调控果实重量 | [ |
| MdIQD17、MdIQD18 | MD07G1178400、MD07G1265200 | 苹果 Malus domestica | 果实参与冷胁迫的适应性调控 | [ |
| PeIQD13、PeIQD9 | POPTR_0005s05310、POPTR_0003s04240 | 毛竹 Phyllostachys edulis | 调节干旱胁迫、MeJA胁迫 | [ |
| ClIQD24 | Cla021081 | 西瓜 Citrullus lanatus | 调控果实生长发育 | [ |
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