玉米叶片气孔表型鉴定及研究进展

doi:10.16035/j.issn.1001-7283.2023.06.001

Abstract

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

Jin Yu, Guo Xinyu, Zhang Ying, Li Dazhuang, Wang Jinglu. Stomatal Phenotypic Identification and Research Progress in Maize Leaves[J].Crops, 2023, 39(6): 1-10.

Figures/Tables 5

Fig.1

Table 1

Phenotypic extraction of leaf stomata"

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
自动化 Automation	[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

Table 1

Fig.2

Table 2

Table 3

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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]

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]

Stomatal Phenotypic Identification and Research Progress in Maize Leaves

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