中国传统大菊叶片形态的数量化定义与分类

doi:10.11983/CBB20014

植物学报 ›› 2021, Vol. 56 ›› Issue (1): 10-24.DOI: 10.11983/CBB20014

中国传统大菊叶片形态的数量化定义与分类

宋雪彬¹^,², 高康¹, 黄河¹, 刘芷兰¹, 戴思兰¹^,^*(), 嵇彧¹

¹北京市林木分子设计育种高精尖创新中心; 北京林业大学园林学院, 国家花卉工程技术研究中心, 花卉种质创新与分子育种北京市重点实验室, 城乡生态环境北京实验室, 北京 100083
²青岛农业大学园林与林学院, 青岛 266109

收稿日期:2020-02-02 接受日期:2020-11-11 出版日期:2021-01-01 发布日期:2021-01-15
通讯作者: 戴思兰
作者简介:*E-mail: silandai@sina.com
基金资助:
北京市科技计划(Z191100008519002);国家自然科学基金(31530064);国家重点研发计划(2018YFD1000405)

Quantitative Definition and Classification of Leaves in Large- flowered Chinese Chrysanthemum Based on the Morphological Traits

Xuebin Song¹^,², Kang Gao¹, He Huang¹, Zhilan Liu¹, Silan Dai¹^,^*(), Yu Ji¹

¹Beijing Advanced Innovation Center for Tree Breeding by Molecular Design; Beijing Laboratory of Urban and Rural Ecological Environment, Beijing Key Laboratory of Flower Germplasm Innovation and Molecular Breeding, National Flower Engineering Technology Research Center, College of Landscape Architecture, Beijing Forestry University, Beijing 100083, China
²College of Landscape Architecture and Forestry, Qingdao Agricultural University, Qingdao 266109, China

Received:2020-02-02 Accepted:2020-11-11 Online:2021-01-01 Published:2021-01-15
Contact: Silan Dai

摘要/Abstract

摘要： 中国传统大菊品种叶片形态变异丰富, 然而至今仍未对其进行科学的定义和分类, 无法有效利用这些形态性状进行品种鉴定和叶形遗传解析。利用数量化分析方法对植物形态进行定义和分类, 是植物性状遗传解析的前提。对436个中国传统大菊品种的24个叶形性状进行重新定义及观测, 通过相关性分析确定了8个相对独立的性状, 用变异系数及主成分分析等数量化分析方法筛选出叶长/叶宽、叶片最宽处所在位置/叶长、右下裂片长/右下叶脉长、右下裂片长/右下裂片宽及叶柄长/全叶长5个相对独立且关键的叶部性状。利用这5个性状, 通过Q聚类分析, 最终将菊花(Chrysanthemum × morifolium)叶片分为16种叶型。研究结果为菊花品种鉴定提供了有效的叶部评价标准, 并建立了中国传统菊花品种叶片数量化定义和分类体系, 也为观赏植物复杂性状的解析提供了新方法。

关键词: 中国传统大菊, 叶片形态, 形态特征, 数量化定义, 分类

Abstract: The leaf morphology of Chinese traditional chrysanthemum varieties is very variable, but it has not been scientifically defined and classified, so it is impossible to make effective use of these morphological traits for variety identification and genetic analysis. The definition and classification of plant morphology by quantitative analysis is the premise of genetic analysis of plant traits. In this study, 24 leaf traits of 436 chrysanthemum varieties were re-defined and measured. The correlation analysis identified 8 relatively independent traits, and principal component analysis further focused on 5 key traits, including the leaf blade length/leaf blade width, widest part length/leaf blade length, right lower leaf lobe length/leaf vein length of right lower lobe, right lower leaf lobe length/right lower leaf lobe width, leaf petiole length/leaf blade length. The leaf shapes were classified into 16 types by Q clustering analysis. This study established a quantitative definition and classification system for the leaves of Chinese traditional chrysanthemum varieties. It provided an effective leaf evaluation standard for the identification of chrysanthemum varieties, and a new method for the analysis of complex traits of ornamental plants.

Key words: Chinese chrysanthemum, leaf shape, morphological traits, quantitative definition, classification

宋雪彬, 高康, 黄河, 刘芷兰, 戴思兰, 嵇彧. 中国传统大菊叶片形态的数量化定义与分类. 植物学报, 2021, 56(1): 10-24.

Xuebin Song, Kang Gao, He Huang, Zhilan Liu, Silan Dai, Yu Ji. Quantitative Definition and Classification of Leaves in Large- flowered Chinese Chrysanthemum Based on the Morphological Traits. Chinese Bulletin of Botany, 2021, 56(1): 10-24.

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https://www.chinbullbotany.com/CN/Y2021/V56/I1/10

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Trait No.	Traits and measuring methods
C1	Leaf blade length (T₁B₁)
C2	Leaf blade width
C3	Widest part length (the length from the leaf apex to the widest part of the leaf blade)
C4	Leaf vein length of left upper lobe (T₂B₂)
C5	Leaf vein length of right upper lobe (T₃B₃)
C6	Leaf vein length of left lower lobe (T₄B₄)
C7	Leaf vein length of right lower lobe (T₅B₅)
C8	Top leaf lobe length (T₁M₁)
C9	Left upper leaf lobe length (T₂M₂)
C10	Right upper leaf lobe length (T₃M₃)
C11	Left lower leaf lobe length (T₄M₄)
C12	Right lower leaf lobe length (T₅M₅)
C13	Top leaf lobe width
C14	Left upper leaf lobe width
C15	Right upper leaf lobe width
C16	Left lower leaf lobe width
C17	Right lower leaf lobe width
C18	Angle of leaf vein in the left upper lobe (α₁)
C19	Angle of leaf vein in the right upper lobe (α₂)
C20	Angle of leaf vein in the left lower lobe (β₁)
C21	Angle of leaf vein in the right lower lobe (β₂)
C22	Leaf petiole length
C23	Leaf base shape
C24	Leaf margin shape

Trait No.	Traits and measuring methods
C1	Leaf blade length (T₁B₁)
C2	Leaf blade width
C3	Widest part length (the length from the leaf apex to the widest part of the leaf blade)
C4	Leaf vein length of left upper lobe (T₂B₂)
C5	Leaf vein length of right upper lobe (T₃B₃)
C6	Leaf vein length of left lower lobe (T₄B₄)
C7	Leaf vein length of right lower lobe (T₅B₅)
C8	Top leaf lobe length (T₁M₁)
C9	Left upper leaf lobe length (T₂M₂)
C10	Right upper leaf lobe length (T₃M₃)
C11	Left lower leaf lobe length (T₄M₄)
C12	Right lower leaf lobe length (T₅M₅)
C13	Top leaf lobe width
C14	Left upper leaf lobe width
C15	Right upper leaf lobe width
C16	Left lower leaf lobe width
C17	Right lower leaf lobe width
C18	Angle of leaf vein in the left upper lobe (α₁)
C19	Angle of leaf vein in the right upper lobe (α₂)
C20	Angle of leaf vein in the left lower lobe (β₁)
C21	Angle of leaf vein in the right lower lobe (β₂)
C22	Leaf petiole length
C23	Leaf base shape
C24	Leaf margin shape

Trait No.	Mean square	Standard deviation	Intravarietal coefficient variation (%)	Intervarietal coefficient variation (%)
C18	33.43	7.12	14.26	21.29
C19	33.60	7.07	13.76	21.03
C20	45.49	11.35	15.55	24.94
C21	46.46	11.12	14.89	23.94
C22	3.85	1.16	11.34	29.99
C25	1.32	0.18	4.23	13.32
C26	0.68	0.09	7.56	13.70
C27	0.35	0.10	12.61	27.56
C28	0.31	0.10	14.64	33.27
C29	0.32	0.10	13.95	32.97
C30	0.46	0.13	11.44	27.96
C31	0.45	0.13	11.83	28.47
C32	1.36	0.40	13.03	29.39
C33	0.76	0.26	11.33	33.94
C34	0.78	0.27	11.34	34.29
C35	1.01	0.33	9.73	32.84
C36	0.98	0.32	9.43	32.67
C37	0.25	0.05	8.89	21.34

Trait No.	Mean square	Standard deviation	Intravarietal coefficient variation (%)	Intervarietal coefficient variation (%)
C18	33.43	7.12	14.26	21.29
C19	33.60	7.07	13.76	21.03
C20	45.49	11.35	15.55	24.94
C21	46.46	11.12	14.89	23.94
C22	3.85	1.16	11.34	29.99
C25	1.32	0.18	4.23	13.32
C26	0.68	0.09	7.56	13.70
C27	0.35	0.10	12.61	27.56
C28	0.31	0.10	14.64	33.27
C29	0.32	0.10	13.95	32.97
C30	0.46	0.13	11.44	27.96
C31	0.45	0.13	11.83	28.47
C32	1.36	0.40	13.03	29.39
C33	0.76	0.26	11.33	33.94
C34	0.78	0.27	11.34	34.29
C35	1.01	0.33	9.73	32.84
C36	0.98	0.32	9.43	32.67
C37	0.25	0.05	8.89	21.34

Traits	Left upper and right upper	Left lower and right lower	Left upper and left lower	Right upper and right lower	Left upper and top lobe	Right upper and top lobe	Left lower and top lobe	Right lower and top lobe
Leaf vein length (cm)	0.851**	0.889**	0.656**	0.689**	0.729**	0.746**	0.784**	0.792**
Leaf lobe length (cm)	0.794*	0.846**	0.537**	0.502**	0.722**	0.712**	0.543**	0.534**
Leaf lobe width (cm)	0.821**	0.833**	0.579**	0.560**	0.763**	0.718**	0.369**	0.363**
Leaf lobe length/leaf vein length	0.696**	0.789**	0.338**	0.358**	0.375**	0.400**	0.420**	0.408**
Leaf lobe length/leaf lobe width	0.676**	0.744**	0.372**	0.424**	0.521**	0.428**	0.323**	0.468**
Angle of leaf vein	0.540**	0.621**	0.436**	0.337**	-	-	-	-

中国传统大菊叶片形态的数量化定义与分类

Quantitative Definition and Classification of Leaves in Large- flowered Chinese Chrysanthemum Based on the Morphological Traits

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	C21	C23	C24	C25	C26	C31	C36	C37
C21	1
C23	0.122*	1
C24	0.087	0.132**	1
C25	-0.185**	-0.156**	-0.230**	1
C26	-0.005	0.194**	0.051	-0.203**	1
C31	-0.007	0.083	-0.264**	-0.008	0.226**	1
C36	-0.053	-0.069	-0.334**	0.120*	0.051	0.782**	1
C37	-0.190**	-0.144**	-0.148**	-0.170**	-0.178**	-0.084	0.020	1

Traits No.	Principal component			Principal component
Traits No.	1	2	3	1	2	3
C21	-0.193	0.426	0.025	-	-	-
C23	-0.071	0.489	-0.206	-	-	-
C24	-0.573	0.404	-0.013	-	-	-
C25	0.256	-0.495	-0.717	0.054	0.734	-0.514
C26	0.197	0.656	0.087	0.407	-0.736	-0.156
C31	0.863	0.336	0.085	0.938	0.032	0.100
C36	0.892	0.066	0.079	0.879	0.308	0.172
C37	0.008	-0.476	0.736	-0.175	0.234	0.878
Total	2.015	1.604	1.119	1.852	1.232	1.099
Contribution rate (%)	25.19	20.05	13.99	37.04	24.64	21.97
Cumulative (%)	25.19	45.24	59.23	37.04	61.68	83.65

No.	Leaf blade shape	Leaf lobe shape/crack depth	Leaf length/leaf width	Widest part length/leaf length	Leaf lobe length/leaf vein length	Leaf lobe length/leaf lobe width
I-1	Broad ovate	Shallow/wide	(0.80, 1.25]	-	(0.15, 0.30]	(0.40, 1.00]
I-2		Medium/wide	(0.80, 1.25]	-	(0.30, 0.50]	(0.40, 1.00]
I-3		Deep/medium	(0.80, 1.25]	-	>0.50	(1.00, 1.50]
I-4		Deep/narrow	(0.80, 1.25]	-	>0.50	>1.50
II-1	Ovate	Shallow/wide	(1.25, 1.40]	-	(0.15, 0.30]	(0.40, 1.00]
II-2		Medium/wide	(1.25, 1.40]	-	(0.30, 0.50]	(0.40, 1.00]
II-3		Deep/medium	(1.25, 1.40]	-	>0.50	(1.00, 1.50]
II-4		Deep/narrow	(1.25, 1.40]	-	>0.50	>1.50
III-1	Obovate	Shallow/wide	>1.40	≤0.55	(0.15, 0.30]	(0.40, 1.00]
III-2		Medium/wide	>1.40	≤0.55	(0.30, 0.50]	(0.40, 1.00]
III-3		Deep/medium	>1.40	≤0.55	>0.50	(1.00, 1.50]
III-4		Deep/narrow	>1.40	≤0.55	>0.50	>1.50 not found
IV-1	Long ovate	Shallow/wide	>1.40	>0.55	(0.15, 0.30]	(0.40, 1.00]
IV-2		Medium/wide	>1.40	>0.55	(0.30, 0.50]	(0.40, 1.00]
IV-3		Deep/medium	>1.40	>0.55	>0.50	(1.00, 1.50]
IV-4		Deep/narrow	>1.40	>0.55	>0.50	>1.50

	Types	Identifiable fitness (%)	Number of varieties of different leaf types				The total number of varieties
	Types	Identifiable fitness (%)	1	2	3	4	The total number of varieties
Leaf blade shape	1	96.2	51	2	0	0	53
	2	96.9	0	31	1	0	32
	3	100	0	0	6	0	6
	4	65.5	0	7	3	19	29
Leaf lobe shape	1	100	15	0	0	0	15
	2	92.8	1	64	3	1	69
	3	100	0	0	27	0	27
	4	88.9	0	0	1	8	9
Leaf petiole	1	100	20	0	0	-	20
	2	87.7	5	71	5	-	81
	3	100	0	0	19	-	19

Types	Identifiable fitness (%)	1	2	3	5	6	7	8	9	10	13	14	15	16	Total
1	90.0	9	0	0	1	0	0	0	0	0	0	0	0	0	10
2	97.1	0	33	0	0	0	1	0	0	0	0	0	0	0	34
3	100.0	0	0	8	0	0	0	0	0	0	0	0	0	0	8
4	0.0	0	0	0	0	0	0	1	0	0	0	0	0	0	1
5	50.0	1	0	0	1	0	0	0	0	0	0	0	0	0	2
6	87.5	0	0	0	1	14	0	0	0	0	0	0	0	1	16
7	100.0	0	0	0	0	0	10	0	0	0	0	0	0	0	10
8	75.0	0	0	0	0	0	1	3	0	0	0	0	0	0	4
9	0.0	0	0	0	1	0	0	0	0	0	0	0	0	0	1
10	100.0	0	0	0	0	0	0	0	0	4	0	0	0	0	4
11	0.0	0	0	0	0	0	0	0	0	0	0	0	1	0	1
13	50.0	0	0	0	0	0	0	0	1	0	1	0	0	0	2
14	93.3	0	0	0	0	1	0	0	0	0	0	14	0	0	15
15	87.5	0	0	0	0	0	1	0	0	0	0	0	7	0	8
16	75.0	0	0	0	0	0	0	1	0	0	0	0	0	3	4

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