蕨类植物孢子囊形态 I. 鳞始蕨科

doi:10.11983/CBB16104

植物学报 ›› 2017, Vol. 52 ›› Issue (3): 322-330.DOI: 10.11983/CBB16104

蕨类植物孢子囊形态 I. 鳞始蕨科

周喜乐¹, 金冬梅¹, 刘以诚², 商辉¹, 严岳鸿^1,^*()

¹上海辰山植物园, 中国科学院上海辰山植物科学研究中心, 上海 201602
²嘉义大学生物资源系, 嘉义 600004

收稿日期:2016-05-06 接受日期:2016-07-12 出版日期:2017-05-01 发布日期:2017-05-27
通讯作者: 严岳鸿
作者简介:
# 共同第一作者
基金资助:
国家自然科学基金(No.31370234)和泛喜马拉雅地区植物综合考察与植物志编研(No.2013FY112100)

Sporangia Morphology of Ferns I. Lindsaeaceae

Xile Zhou¹, Dongmei Jin¹, Yicheng Liu², Hui Shang¹, Yuehong Yan^1*

¹Shanghai Chenshan Plant Science Research Center, Shanghai Chenshan Botanical Garden, Chinese Academy of Sciences, Shanghai 201602, China
²Department of Biological Resources, National Chiayi University, Chiayi 600004, China

Received:2016-05-06 Accepted:2016-07-12 Online:2017-05-01 Published:2017-05-27
Contact: Yan Yuehong
About author:
# Co-first authors

摘要/Abstract

摘要： 孢子囊是蕨类植物的繁殖器官, 其形态在蕨类植物的分类和系统发育研究上具有重要意义。用次氯酸钠溶液处理新鲜成熟的孢子囊, 在光学显微镜下获得清晰的孢子囊图像, 系统研究了中国鳞始蕨科4属13种孢子囊的形态特征。结果表明, 鳞始蕨科孢子囊呈椭球形, 孢子囊柄由3列细胞构成, 环带类型为垂直环带。通过分析孢子囊形态数据探讨了中国鳞始蕨科属内及属间差异。结果表明, 乌蕨属(Odontosoria)、香鳞始蕨属(Osmolindsaea)、达边蕨属(Tapeinidium)和鳞始蕨属(Lindsaea)的孢子囊环带细胞数依次减少, 囊蒴体积、唇细胞数和囊壁细胞数的变化由大(多)到小(少)依次为乌蕨属、达边蕨属、香鳞始蕨属和鳞始蕨属。孢子囊属内差异最大的是阔片乌蕨(Odontosoria biflora)与乌蕨(O. chinensis)以及香鳞始蕨(Osmolindsaea odorata)与日本鳞始蕨(O. japonica); 而达边蕨属和鳞始蕨属的属内差异则很小。研究结果为揭示鳞始蕨科系统发育关系提供了形态基础, 特别是提出阔片乌蕨和乌蕨以及香鳞始蕨和日本鳞始蕨在孢子囊形态上的差异值得进一步研究。

Abstract: Sporangia are propagative organs of ferns and their morphology has great significance for fern taxonomy and phylogeny. In this study, we used sodium hypochlorite solution to observe fern sporangia. By this process, we could obtain sporangia photos under light microscopy. We studied the sporangia morphology of 13 species belonging to four genera of the fern family Lindsaeaceae and found that the shape of the capsule is ellipsoidal, with a vertical annulus; the pedicel is composed of three rows of cells. The cell number annulus is reduced in Odontosoria, Osmolindsaea, Tapeinidium and Lindsaea, whereas the volume of capsule and cell number of stomium and capsule are reduced in Odontosoria, Tapeinidium, Osmolindsaea and Lindsaea. As well, the same genus shows more differences between species, such as Odontosoria biflora and O. chinensis as well as Osmolindsaea odorata and Os. japonica, but less difference between the genera of Tapeinidium and Lindsaea. Studies of sporangia morphology will be useful for further research in other groups of ferns.

周喜乐, 金冬梅, 刘以诚, 商辉, 严岳鸿. 蕨类植物孢子囊形态 I. 鳞始蕨科. 植物学报, 2017, 52(3): 322-330.

Xile Zhou, Dongmei Jin, Yicheng Liu, Hui Shang, Yuehong Yan. Sporangia Morphology of Ferns I. Lindsaeaceae. Chinese Bulletin of Botany, 2017, 52(3): 322-330.

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https://www.chinbullbotany.com/CN/Y2017/V52/I3/322

图/表 9

图1 孢子囊环带类型(Foster et al., 1983)(A) 侧生环带; (B) 顶生环带; (C)-(F) 斜生环带; (G) 垂直环带

Figure 1 Type of sporangia annulus (Foster et al., 1983) (A) Lateral; (B) Apical; (C)-(F) Oblique; (G) Vertical

表1 鳞始蕨科中的属

Table 1 The genera of Lindsaeaceae

Associated with Lindsaeaceae in the past	Recognized today	Unsolved
Lindsaea Dryand. ex Sm. Odontosoria Fée Ormoloma Maxon Sphenomeris Maxon Tapeinidium (C.Presl) C.Chr. Schizoloma Gaud. Stenoloma Fée Xyropteris Kramer Cystodium J. Sm. Lonchitis L.	Lindsaea Dryand. ex Sm. (Synonym: Ormoloma; Schizoloma) Odontosoria Fée Sphenomeris Maxon (Synonym: Stenoloma) Tapeinidium (C.Presl) C.Chr. Osmolindsaea (K. U. Kramer) Lehtonen & Christenh. Nesolindsaea Lehtonen & Christenh.	Xyropteris Kramer Cystodium J. Sm. Lonchitis L.

表2 材料来源

Table 2 Source of materials

Species	Locality	Vouchers
Odontosoria biflora	Lanyu, Taiwan	Y. C. Liu et al., 6003
O. chinensis	Xiuning, Anhui	X. L. Zhou et al., 1-20
Osmolindsaea odorata	Wuming, Guangxi	X. L. Zhou et al., 5161
Os. japonica	Zihuai, Sichuan	X. L. Zhou et al., 5680
Lindsaea eberhardtii	Lingshui, Hainan	Y. H. Liu et al., 1-55
L. merrillii subsp. yaeyamensis	Lanyu, Taiwan	Y. C. Liu et al., 6012
L. javanensis	Pingbian, Yunnan	X. L. Zhou et al., 5887
L. orbiculata	Shangsi, Guangxi	X. L. Zhou et al., 5093a
L. cultrata	Lanyu, Taiwan	Y. C. Liu et al., 6014
L. heterophylla	Shanghai Chenshan Botanical Garden, Shanghai	X. L. Zhou, 1-B2
L. ensifolia	Menghai, Yunnan	X. L. Zhou et al., 5801
Tapeinidium pinnatum	Lanyu, Taiwan	Y. C. Liu et al., 6011
T. pinnatum var. biserratum	Lanyu, Taiwan	Y. C. Liu et al., 6015

图2 已开裂的具纵行环带的孢子囊示意图下面未着色的为孢子囊柄, 上面着色部分为孢子囊蒴。棕色部分为环带。粗黑线为加厚的细胞壁。绿色部分为唇细胞, 唇细胞中间位置壁较厚的为唇细胞加厚细胞。黄色部分为孢子囊壁细胞。

Figure 2 Sporangium with vertical annulus The pedicel is uncolored, the colored part is capsule. The annulus in brown color. Bold black line shows thick-wall. The stomium in green color, and with thick-walled stomium in centre. The wall of capsule in yellow color.

图3 鳞始蕨科孢子囊形态(A) 阔片乌蕨; (B) 乌蕨; (C) 香鳞始蕨; (D) 日本鳞始蕨; (E) 线片鳞始蕨; (F) 攀缘鳞始蕨; (G) 爪哇鳞始蕨; (H) 团叶鳞始蕨。Bar=100 μm。图中囊柄有断裂情况, 故其长度不做比较。

Figure 3 Sporangia morphology of Lindsaeaceae(A) Odontosoria biflora; (B) O. chinensis; (C) Osmolindsaea odorata; (D) Os. japonica; (E) Lindsaea eberhardtii; (F) L. merrillii subsp. yaeyamensis; (G) L. javanensis; (H) L. orbiculata. Bar=100 μm. The length of pedicel can not be used for comparing since some of it was broken.

图4 鳞始蕨科孢子囊形态(A) 网脉鳞始蕨; (B) 异叶鳞始蕨; (C) 剑叶鳞始蕨; (D) 达边蕨; (E) 二羽达边蕨。Bar=100 μm。图中囊柄有断裂情况, 故其长度不做比较。

Figure 4 Sporangia morphology of Lindsaeaceae(A) Lindsaea cultrata; (B) L. heterophylla; (C) L. ensifolia; (D) Tapeinidium pinnatum; (E) T. pinnatum var. biserratum. Bar=100 μm. The length of pedicel can not be used for comparing since some of it was broken.

表3 中国鳞始蕨科孢子囊的形态特征

Table 3 Sporangia morphology of Lindsaeaceae

Taxon	Cell No. of annulus	Length of pedicel (μm)	Length of capsule (μm)	Width of capsule (μm)	Volume of capsule (mm³)	Cell No. of sto- mium	Thick- walled stomium	Cell No. of cap- sule
Lindsaeaceae	12.9	326.0	195.0	158.8	0.003	8.4	2.4	15.9
Odontosoria	17.9	309.1	240.6	199.0	0.006	10.2	2.7	19.1
Osmolindsaea	16.5	313.2	213.0	172.4	0.004	7.9	2.0	17.1
Tapeinidium	13.6	388.2	231.1	180.1	0.005	9.1	2.7	17.4
Lindsaea	10.2	293.5	166.5	137.4	0.002	7.8	2.3	14.3
Odontosoria biflora	21.6	331.0	252.6	199.7	0.006	11.1	3.3	20.0
O. chinensis	14.2	287.2	228.7	198.2	0.005	9.3	2.0	18.2
Osmolindsaea odorata	19.4	329.7	270.6	215.5	0.008	7.9	2.0	18.2
Os. japonica	13.5	296.8	155.3	129.2	0.002	7.8	2.0	16.0
Tapeinidium pinnatum	13.0	359.5	223.7	174.8	0.004	8.6	2.0	16.8
T. pinnatum var. biserratum	14.2	416.9	238.5	185.4	0.005	9.6	3.4	18.0
Lindsaea eberhardtii	11.5	333.7	183.1	141.5	0.002	8.0	2.0	15.0
L. merrillii subsp. yaeyamensis	9.7	319.9	146.2	139.0	0.002	7.0	2.9	13.0
L. javanensis	10.2	408.1	171.0	132.6	0.002	7.4	2.0	14.0
L. orbiculata	10.0	348.0	172.7	140.8	0.002	7.9	2.0	13.8
L. cultrata	9.7	213.6	150.4	132.6	0.001	7.0	3.1	15.2
L. heterophylla	10.2	250.3	168.3	142.6	0.002	8.6	2.0	14.6
L. ensifolia	10.2	180.8	174.2	132.9	0.002	8.8	2.0	14.4

图5 鳞始蕨科4属孢子囊形态数据比较(A) 囊蒴体积; (B) 环带细胞数; (C) 囊壁细胞数; (D) 唇细胞数。无共同字母表示属间差异显著(Turkey’s多重比较, P<0.05)。在每个箱线图中, 箱体显示25%、中位数和75%的四分位数。箱体外的虚线延伸至距箱体1.5倍四分位距(箱体长度)范围内的极值, 超过此限的极值以散点表示。

Figure 5 Comparisons of sporangia morphology among four genera in Lindsaeaceae(A) Volume of capsule; (B) Cell number of annulus; (C) Cell number of capsule; (D) Cell number of stomium. Genera with no letters in common are significantly different (multiple comparisons using Turkey’s method, P<0.05). For each box-and-whisker plot, the box shows 25%, median and 75% quantile of the given values. The whiskers extend to the most extreme data points that are not more than 1.5 times the interquartile range (length of the box) from the box.

图6 鳞始蕨科属间系统关系(Lehtonen et al., 2010)

Figure 6 Phylogenetic relationships within Lindsaeaceae (Lehtonen et al., 2010)

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蕨类植物孢子囊形态 I. 鳞始蕨科

Sporangia Morphology of Ferns I. Lindsaeaceae

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