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作物杂志,2023, 第6期: 1–10 doi: 10.16035/j.issn.1001-7283.2023.06.001

• 专题综述 •    下一篇

玉米叶片气孔表型鉴定及研究进展

金玉1,2,3,4(), 郭新宇1,2,3, 张颖1,2,3, 李大壮1,2,3,4, 王璟璐1,2,3()   

  1. 1数字植物北京市重点实验室，100097，北京
    2北京市农林科学院信息技术研究中心，100097，北京
    3国家农业信息化工程技术研究中心，100097，北京
    4华中农业大学，430070，湖北武汉
  • 收稿日期:2022-07-08 修回日期:2023-10-12 出版日期:2023-12-15 发布日期:2023-12-15
  • 通讯作者: 王璟璐，主要从事作物表型组学及生物信息学研究，E-mail：wangjl@nercita.org.cn
  • 作者简介:金玉，研究方向为作物表型组学，E-mail：1456368616@qq.com
  • 基金资助:
    北京市农林科学院作物表型协同创新中心项目(KJCX201917);国家自然科学基金(31801254)

Stomatal Phenotypic Identification and Research Progress in Maize Leaves

Jin Yu1,2,3,4(), Guo Xinyu1,2,3, Zhang Ying1,2,3, Li Dazhuang1,2,3,4, Wang Jinglu1,2,3()   

  1. 1Beijing Key Laboratory of Digital Plant, Beijing 100097, China
    2Research Center of Information Technology,Beijing Academy of Agriculture and Forestry Sciences, Beijing 100097, China
    3National Engineering Research Center for Information Technology in Agriculture, Beijing 100097, China
    4Huazhong Agricultural University, Wuhan 430070, Hubei, China
  • Received:2022-07-08 Revised:2023-10-12 Online:2023-12-15 Published:2023-12-15

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摘要：

叶片气孔是植物与外界联系的直接通道。气孔在玉米植株生长过程中碳、水循环调控中起着重要作用，影响着植株蒸腾和光合作用的强度。本文概述了玉米叶片气孔形态结构与主要功能特点，梳理了气孔表型破坏性与非破坏性获取方法，沿时间脉络整理了气孔表型人工―半自动化―自动化解析方法，归纳了玉米叶片气孔表型特征分析研究进展及影响气孔形态建成的相关基因研究现状，展望了叶片气孔研究可能面临的挑战与机遇，为基于气孔表型的玉米品种筛选鉴定及遗传改良提供参考。

关键词: 玉米, 叶片, 气孔, 表型, 研究进展

Abstract:

The leaf stomatas are the direct channel between plants and the outer enviroment, which play an important role in regulating the carbon and water cycles during the growth of maize plants and influence the intensity of transpiration and photosynthesis. This paper first outlined the morphological structure and main functional characteristics of maize leaf stomata. After classifying destructive and non-destructive stomatal phenotype acquisition techniques, manual, semi-automated, and automated stomatal phenotype analysis techniques were applied. The research progress of stomatal phenotype characterization in maize leaves and the status of genes affecting stomatal morphogenesis were summarized. This paper also presented the possible challenges and opportunities for stomatal research. It provides a reference for the selection, identification and genetic improvement of maize varieties based on stomatal phenotypes.

Key words: Maize, Leaves, Stomata, Phenotype, Research progress

图1

叶片气孔形态示意图 (a) 玉米叶片气孔；(b) 双子叶植物和大多数单子叶植物的气孔

表1

叶片气孔表型提取

发展阶段
Stage of
development		 参考文献
Reference		 表型解析方法
Phenotypic parsing
method		 作物
Crop		 主要表型特征
Main phenotypic characteristics		 特点
Characteristic
人工
Manpower
 [37,45-46]

 显微镜测微尺、计数室、Veeder-Root、样带法等
 菊花（Chrysanthemum morifolium）、大麦（Hordeum vulgare）、小叶杨（Populus simonii） 气孔数目、气孔密度、气孔长度、气孔宽度、气孔开度
 5min/视野，较为准确，耗时长
半自动化
Semi-
automatization		 [12]

 SMileView

 樟树（Cinnamomum camphora）

 气孔总数、张开气孔数、闭合气孔数，开放气孔比率、气孔长度、气孔宽度、气孔开度 （1）适应不同图像需求；
（2）有效降低计算复杂度；（3）2min/张，极大降低了人力投入

[53]
 Mv-1图像分析板和JAVA图像分析软件 蚕豆（Vicia faba）
 气孔孔径
[54]


 Motic Images Advance


 东栎（Quercus liaotungensis）、虎榛子（Ostryopsis davidiana）、酸枣（Ziziphus jujuba var. spinosa）和狼牙刺（Sophora viciifolia） 气孔密度、气孔长度、气孔宽度、气孔面积

[55,59]

 Auto Desk Inventor Professional 2011、Auto CAD 2010和Arc GIS10.0 冬小麦（Triticum aestivum）

 气孔密度、气孔长度、气孔宽度、气孔面积、气孔周长、气孔形状指数和气孔分布格局
[56]

 Image J

 费菜（Phedimus aizoon）、刺槐（Robinia pseudoacacia）等40种
 气孔密度、气孔长度、气孔宽度、气孔面积、气孔开度、气孔张开比
[57]
 Scope Image 9.0
 桉树（Eucalyptus robusta）
 气孔长度、气孔宽度、孔径长度、气孔面积
[58]

 UTHSCSA ImageTool

 茄子（Solanum pennelli）

 气孔密度、气孔（长度、宽度、面积）、孔隙（长度、宽度、面积）、气孔深度
自动化
Automation
 [60] 级联分类器 桉树（Eucalyptus robusta） 气孔密度、气孔分布 （1）自动化识别，平均精度可达98%以上；（2）图像数据批量、高通量处理；（3）5~10s/张，高生产率和高可重复性，节省人力投入

[61] 最大稳定外部区域法 葡萄（Vitis vinifera） 气孔孔径
[62]

 DeepStomata: 基于定向梯度的气孔检测直方图和基于卷积神经网络 白花鸭跖草（Commelina benghalensis）
 气孔孔径

[63] 基于SSD算法的目标检测模型 大豆（Glycine max） 气孔密度
[64]
 深度卷积神经网络和修复算法的混合方法 水稻（Oryza sativa）
 气孔长度、气孔宽度、气孔面积、气孔宽长比
[65]
 Gabor滤波算法下的卷积神经网络和灰度共生矩阵 姜黄（Curcuma longa）和姜
（Curcuma zanthorrhiza）		 气孔识别
[66-67]

 FPN目标检测和卷积神经网络Mask R-CNN
 裸子、蕨类、禾本科植物等12个数据集
 气孔识别、6个气孔孔隙（数目、长轴、短轴、面积、离心率、开度）性状
[39] SBOS：目标跟踪和语义分割 小麦（Triticum aestivum） 气孔开口面积
[68]
 转换学习和支持向量机
 藜麦（Chenopodium quinoa）
 气孔数量、气孔孔径、气孔面积、气孔状态（张开、闭合）
[40]
 基于FPN算法的目标检测模型和改进后的CV模型 玉米（Zea mays）
 4个气孔数量和6个气孔孔隙表型指标
[69] YOLO深度学习模型 玉米（Zea mays） 气孔密度
[70]

 Leaf Net: 分层策略下的深度卷积网络、区域合并方法、迁移学习 拟南芥（Arabidopsis thaliana）

 气孔复合体数量、尺寸、形状及表皮细胞数量、尺寸、形状等28项表型指标

图2

叶片气孔器表型性状解析发展历程

表2

叶片气孔主要表型指标

项目Item 指标Index 结构Structure 表型指标Phenotypic indicator 参考文献Reference
直接指标
Direct indicator

 气孔复合体


 由保卫细胞、副卫细胞、孔隙构成

 气孔总数、张开气孔数、闭合气孔数、气孔密度、气孔指数、张开气孔占比、气孔长度、气孔宽度、气孔周长、气孔面积、气孔偏心率、气孔长宽比、气孔圆度、气孔形状指数、气孔导度、空间分布模式、气孔深度 [40-41,55,72]


孔隙
 两个保卫细胞之间的小孔
 孔隙长度、孔隙宽度、孔隙面积、孔隙周长、孔隙开度、孔隙偏心率、孔隙面积指数 [40,73]
间接指标
Indirect indicator		 表皮细胞
 植物最外层细胞，与气孔相邻，排列紧密 表皮细胞个数、表皮细胞密度、表皮细胞面积、表皮细胞总面积 [70,73]

表3

调控玉米叶片气孔表型的功能基因

基因名称Gene name 基因位点Gene loci 基因功能Gene function 气孔表型Stomatal phenotype 参考文献Reference
ZmMUTE GRMZM2G417164 保卫细胞形成与副卫母细胞极化 气孔数目、气孔形状 [90-91]
ZmPAN1 Zm00001d031437 副卫母细胞极化 气孔形状、气孔尺寸 [84]
ZmROP2 Zm00001d053899 副卫母细胞极化 气孔形状、气孔尺寸 [85]
ZmROP9 Zm00001d015036 副卫母细胞极化 气孔形状、气孔尺寸 [85]
ZmPAN2 Zm00001d007862 副卫母细胞极化和副卫细胞形成 气孔形状、气孔尺寸 [86-87]
ZmBRK1 GRMZM5G842058 副卫母细胞极化和表皮细胞形成 气孔形状 [88]
ZmBRK3 GRMZM5G88636 副卫母细胞极化和表皮细胞形成 气孔形状 [88]
ZmNOD GRMZM2G027821 保卫细胞分化 气孔数目、气孔形状 [82]
ZmSHR1 GRMZM2G132794 保卫细胞形成 气孔数目 [83]
ZmMLKS2 Zm00001d052955 副卫细胞分化 气孔尺寸、气孔数目、气孔形状 [89]
ZmSPL10 Zm00001d015451 保卫细胞形成与副卫母细胞极化 气孔形状、气孔尺寸 [92]
ZmSPL14 Zm00001d036692 保卫细胞形成与副卫母细胞极化 气孔形状、气孔尺寸 [92]
ZmSPL26 Zm00001d053756 保卫细胞形成与副卫母细胞极化 气孔形状 [92]
ZmB73 Zm00001d042263 保卫细胞形成与副卫母细胞极化 气孔数目 [93]
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