花楸属直脉组7种/变种基因组大小及叶表皮微形态特征的分类学意义

doi:10.12302/j.issn.1000-2006.2021100096

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南京林业大学学报（自然科学版） ›› 2023, Vol. 47 ›› Issue (3): 77-86.doi: 10.12302/j.issn.1000-2006.2021100096

花楸属直脉组7种/变种基因组大小及叶表皮微形态特征的分类学意义

邱靖¹(), 李嘉宝(), 朱大海³, 陈昕¹^,^*()

1.南京林业大学,南方现代林业协同创新中心,南京林业大学生物与环境学院,江苏南京 210037
2.三江学院,江苏南京 210012
3.龙溪-虹口国家级自然保护区,四川成都 611830

收稿日期:2021-10-09 修回日期:2021-11-08 出版日期:2023-05-30 发布日期:2023-05-25
通讯作者: 陈昕
基金资助:
2021年江苏省高校 “青蓝工程”优秀青年骨干教师培养计划

Taxonomic implications of genome sizes and micromorphological characteristics of leaf epidermis of species in Sorbus Sect. Alnifoliae

QIU Jing¹(), LI Jiabao(), ZHU Dahai³, CHEN Xin¹^,^*()

1. Co-Innovation Center for Sustainable Forestry in Southern China, College of Biology and the Environment, Nanjing Forestry University, Nanjing 210037, China
2. Sanjiang University, Nanjing 210012, China
3. Longxi-Hongkou National Nature Reserve,Chengdu 611830,China

Received:2021-10-09 Revised:2021-11-08 Online:2023-05-30 Published:2023-05-25
Contact: CHEN Xin

摘要/Abstract

摘要：

【目的】通过对花楸属直脉组(Sorbus Sect. Alnifoliae)5种及具有分类争议的2变种的基因组大小和叶表皮微形态研究,探讨其分类学意义以及基因组大小和细胞大小之间的相关性。【方法】基于流式细胞术、光学和电子显微镜分别对研究对象的基因组大小、叶表皮细胞和气孔的质量和数量性状进行测定、分析。【结果】基因组大小测定显示所有分类群均为极小基因组,大小介于神农架花楸(S. yuana)的1.348 pg至长果花楸(S. zahlbruckneri)的1.586 pg之间;虽均为二倍体,但分类群间基因组大小差异显著。光学显微镜下叶表皮细胞形状兼有多边形和无规则形,或者仅无规则形,垂周壁式样有平直状、弧状和波状;扫描电镜下表皮纹饰有近光滑、局部加厚、脊状、网纹、褶皱等类型,蜡质纹饰有壳状、颗粒状、线状、片状等;4个分类群的下表皮具带状单毛。气孔器均为无规则型,分布于下表皮脉间区;保卫细胞具明显的环形外缘或无。表皮细胞面积、气孔宽度及面积、气孔密度、气孔指数等指标在分类群间差异显著,基因组大小同表皮细胞面积、气孔长宽及面积呈正相关,同气孔密度和气孔指数呈负相关。【结论】基因组大小、叶表皮微形态质量和数量性状可为7个分类群的鉴定提供依据,不支持将棱果花楸(S. alnifolia var. angulata)和裂叶水榆花楸(S. japonica)作为日本花楸(S. alnifolia var. lobulata)的异名处理,建议恢复棱果花楸和裂叶水榆花楸的名称。

关键词: 基因组大小, 细胞大小, 微形态特征, 叶表皮, 花楸属直脉组

Abstract:

【Objective】Genome sizes and micromorphological characteristics of leaf epidermis in five species representing Sorbus Sect. Alnifoliae together with two taxonomically controversial varieties were studied. The purpose of this study was to explore the taxonomic significance and the correlation between genome size and cell size. 【Method】The genome size was estimated by flow cytometry. Qualitative and quantitative traits of leaf epidermis were studied by optical and scanning electron microscopy【Result】The flow cytometric genome size estimation revealed that seven investigated taxa possessed very small genomes. The nuclear DNA content varied from 1.348 pg in S. yuana to 1.586 pg in S. zahlbruckneri. Though all are diploids, significant variation of genome size was observed among taxa. Polygonal-irregular or irregular epidermal cell shape, straight, arched or undulate of anticlinal walls, were determined using light microscopy. Patterns of cuticular ornamentation such as smooth, thickened areas, ridges, reticulate and folds and epicuticular wax types such as crusts, granules, threads and platelets were observed by scanning electron microscopy. Simple single ribbon-like trichomes are located on the lower epidermis of four taxa. Stomatal apparatuses are anomocytic and confined to areole on abaxial, not adaxial, epidermis. Guard cells with or without obvious annular outer edges. Significant variations in epidermal cell area, stomatal width, area, density and stomatal index, were observed among the taxa. There was a positive relationship between genome size and stomatal length, width, area and epidermal cell area and a negative relationship with stomatal density and stomatal index. 【Conclusion】Genome sizes, qualitative and quantitative traits of leaf epidermis allow clear delimitation and identification of the seven taxa. The treatments of S. alnifolia var. angulata and S. alnifolia var. lobulata as synonyms of S. japonica are not supported and the latter two names are suggested to be reinstated here.

Key words: genome size, cell size, micromorphological characteristics, leaf epidermis, Sorbus Sect. Alnifoliae

中图分类号:

S792.25
Q943

邱靖,李嘉宝,朱大海,等. 花楸属直脉组7种/变种基因组大小及叶表皮微形态特征的分类学意义[J]. 南京林业大学学报（自然科学版）, 2023, 47(3): 77-86.

QIU Jing, LI Jiabao, ZHU Dahai, CHEN Xin. Taxonomic implications of genome sizes and micromorphological characteristics of leaf epidermis of species in Sorbus Sect. Alnifoliae[J].Journal of Nanjing Forestry University (Natural Science Edition), 2023, 47(3): 77-86.DOI: 10.12302/j.issn.1000-2006.2021100096.

图/表 8

表1

图1

表2

图2

图3

表3

花楸属直脉组植物叶表皮微形态特征(定性性状)"

种/变种 taxon	上表皮/ 下表皮 Ad or Ab	表皮细胞形状 epidermal cell shape	垂周壁 anticlinal wall	平周壁 periclinal wall	表皮毛 trichome	表面纹饰 cuticular ornamentation	表皮蜡质 epicuticular wax	气孔外缘 outer stomatal rim	气孔纹饰 stomatal ornamentation	图 figure
水榆花楸 S. alnifolia var. alnifolia	Ad	多边形、不规则形	平直、弧形; 隆起	凹陷	-	局部加厚	光滑层、颗粒状、片状、线状	-	-	图2Aa 图3Aa、Ab
	Ab	多边形、不规则形	平直、弧形; 隆起	凹陷	-	局部加厚	光滑层、颗粒状、片状、线状	不完全环形外缘; 两极T状加厚	颗粒状、片状	图2Ab 图3 Ac-Ae
棱果花楸 S. alnifolia var. angulata	Ad	多边形、不规则形	平直、弧状; 凹陷	隆起	-	近光滑	壳状、颗粒状	-	-	图2Ba 图3Ba、3Bb
	Ab	多边形、不规则形	平直、弧状; 凹陷	隆起	无	局部加厚	壳状、颗粒状、线状	不完全环形外缘; 两极T状加厚	壳状	图2Bb 图3 Bc-Be
裂叶水榆花楸 S. alnifolia var. lobulata	Ad	不规则形	波状、弧形; 隆起	凹陷	无	局部加厚	壳状、颗粒状、片状、线状	-	-	图2Ca 图3Ca、3Cb
	Ab	不规则形	波状、弧形; 隆起	凹陷	无	局部加厚	壳状、颗粒状	光滑	壳状	图2Cb 图3 Cc-Ce
石灰花楸 S. folgneri	Ad	多边形、不规则形	平直、弧形、波状;凹陷	隆起	无	脊状隆起	壳状、颗粒状、线状	-	-	图2Da 图3Da、3Db
	Ab	多边形、不规则形	平直、弧形、波状;不可见	不可见	带状	不可见	不可见	不完全环形外缘; 两极T状加厚	壳状	图2Db 图3 Dc-De
日本花楸 S. japonica	Ad	多边形、不规则形	平直、弧形、波状;隆起	凹陷	无	网纹、局部加厚	壳状、颗粒状	-	-	图2Ea 图3 Ea、3Eb
	Ab	多边形、不规则形	平直、弧形、波状;隆起	凹陷	带状	褶皱	壳状、颗粒状	环形外缘	壳状	图2Eb 图3 Ec-Ee
神农架花楸 S. yuana	Ad	多边形、不规则形	平直、弧形; 隆起	凹陷	无	局部加厚	壳状、颗粒状、片状	-	-	图2Fa 图3Fa、3Fb
	Ab	多边形、不规则形	平直、弧形; 隆起	凹陷	带状	褶皱	壳状、颗粒状	环形外缘	壳状	图2Fb 图3 Fc-Fe
长果花楸 S. zahlbruckneri	Ad	不规则形	波状; 隆起	凹陷	-	局部加厚	平滑层、颗粒状、片状、线状	-	-	图2Ga 图3 Ga、3Gb
	Ab	不规则形	波状; 隆起	凹陷	带状	褶皱	壳状、颗粒状	环形外缘	近光滑	图2Gb 图3 Gc-Ge

表3

表4

表5

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种/变种 taxon	采集地 location	地理坐标 geographical coordinates	海拔/m altitude	采集人及凭证标本号 collector and voucher No.	采集时间(年-月) collection time (year-month)
水榆花楸 S. alnifolia var. alnifolia	江西三清山	118°3'27.69″E, 28°54'31.46″N	1 563	陈昕、朱凯琳、李嘉宝 1443	2019-09
棱果花楸 S. alnifolia var. angulata	山东鲁山	118°03'09.00″E, 36°17'45.31″N	1 047	陈昕、邱靖 0140	2015-10
裂叶水榆花楸 S. alnifolia var. lobulata	山东崂山	120°36'13.76″E, 36°12'29.14″N	393	陈昕 0038	2014-10
石灰花楸 S. folgneri	贵州梵净山	108°43'20.57″E, 27°53'48.05″N	1 109	陈昕、朱凯琳、李嘉宝1582	2019-09
日本花楸 S. japonica	日本武奈ヶ岳山	135°53'48.95″E, 35°15'52.61″N	1 018	王丰 0257	2015-10
神农架花楸 S. yuana	湖北神农架温水林场桥洞沟	110°19'17.76″E, 31°34'21.30″N	2 173	陈云、赵洋 0817	2016-10
长果花楸 S. zahlbruckneri	湖北兴山龙门河国家森林公园	110°29'42.12″E, 31°19'14.93″N	1 539	陈昕、洑香香、刘清亮 0500	2016-05

种/变种 taxon	2C值/pg 2C value	C_x值/pg C_x value
水榆花楸Sorbus alnifolia var. alnifolia	1.435±0.019 c	0.717±0.010 c
棱果花楸S. alnifolia var. angulata	1.434±0.002 c	0.717±0.001 c
裂叶水榆花楸S. alnifolia var. lobulata	1.441±0.007 bc	0.72±0.004 bc
石灰花楸S. folgneri	1.460±0.029 b	0.730±0.014 b
日本花楸S. japonica	1.393±0.010 d	0.696±0.005 d
神农架花楸S. yuana	1.348±0.032 e	0.674±0.016 e
长果花楸S. zahlbruckneri	1.586±0.007 a	0.793±0.004 a

种/变种 taxon	上表皮细胞面积/μm² cell area of adaxial epidermis	下表皮细胞面积/μm² cell area of abaxial epidermis	气孔器长/μm stomatal length	气孔器宽/μm stomatal width	气孔器面积/μm stomatal area	气孔密度/ 个/mm² stomatal density	气孔指数/% stomatal index
水榆花楸 S. alnifolia var. alnifolia	678.64±148.30 c	635.24±143.75 d	28.76±3.26 a	21.36±2.55 a	564.17±115.56 a	170.64±36.39 d	11.44±2.67 ab
棱果花楸 S. alnifolia var. angulata	606.77±147.80 e	632.83±115.64 d	28.46±3.36 a	22.87±3.79 a	564.55±150.01 a	265.02±36.01 a	17.08±2.10 a
裂叶水榆花楸 S. alnifolia var. lobulata	797.55±210.44 b	701.2±227.46 c	24.41±3.68 a	19.58±2.56 a	457.59±110.74 e	190.16±56.45 c	12.31±2.81 ab
石灰花楸 S. folgneri	662.84±147.18 d	496.84±118.16 f	24.98±3.18 a	20.98±2.34 a	510.12±89.97 c	205.04±29.34 b	12.89±1.83a b
日本花楸 S. japonica	798.03±182.78 b	776.79±184.5 b	26.91±2.79 a	20.52±1.83 a	514.63±102.16 c	101.82±21.86 f	7.64±1.61 b
神农架花楸 S. yuana	605.05±101.58 e	613.19±121.32 e	26.3±3.92 a	20.69±2.51 a	493.03±123.16 d	189.7±35.53 c	10.51±1.78 ab
长果花楸 S. zahlbruckneri	967.63±230.32 a	826.62±164.36 a	30.81±3.02 a	21.5±2.23 a	552.81±105.21 b	117.17±13.48 e	8.74±1.08 b

指标 index	基因组大小 genome size	上表皮细胞面积 cellular area of adaxial epidermis	下表皮细胞面积 cellular area of abaxial epidermis	气孔器长 stomata length	气孔器宽 stomata width	气孔器面积 stomata length	气孔密度 stomata density	气孔指数 stomata index
基因组大小genome size	1
上表皮细胞面积 cellular area of adaxial epidermis	0.737^*	1
下表皮细胞面积 cellular area of abaxial epidermis	0.413	0.826^*	1
气孔器长stomata length	0.562	0.379	0.524	1
气孔器宽stomata width	0.241	-0.274	-0.116	0.663	1
气孔器面积stomata length	0.379	-0.017	0.101	0.852^**	0.874^**	1
气孔密度stomata density	-0.260	-0.748^*	-0.705^*	-0.273	0.434	0.048	1
气孔指数stomata index	-0.099	-0.623	-0.593	-0.152	0.532	0.185	0.966^**	1

花楸属直脉组7种/变种基因组大小及叶表皮微形态特征的分类学意义

Taxonomic implications of genome sizes and micromorphological characteristics of leaf epidermis of species in Sorbus Sect. Alnifoliae

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