红树族植物次生木质部附物纹孔的电镜观测

doi:10.3724/SP.J.1259.2015.00090

植物学报 ›› 2015, Vol. 50 ›› Issue (1): 90-99.DOI: 10.3724/SP.J.1259.2015.00090

红树族植物次生木质部附物纹孔的电镜观测

邓传远^1,^*(), 辛桂亮¹, 张万超¹, 郭素枝², 薛秋华¹, 赖钟雄³, 叶露莹¹

¹福建农林大学园林学院, 福州 350002
²福建农林大学生命科学学院, 福州 350002
³福建农林大学园艺学院, 福州 350002

收稿日期:2013-12-16 接受日期:2014-11-13 出版日期:2015-01-01 发布日期:2015-04-09
通讯作者: 邓传远
作者简介:
? 共同第一作者
基金资助:
福建省教育厅项目(No.JB09295, No.B11041)和海洋公益性行业科研专项(No.201505009-4)

SEM Observations and Measurements of Vestured Pits of the Secondary Xylem in the Tribe Rhizophoreae

Chuanyuan Deng^{1, *}, Guiliang Xin¹, Wanchao Zhang¹, Suzhi Guo², Qiuhua Xue¹, Zhongxiong Lai³, Luying Ye¹

¹College of Landscape Architecture
²College of Life Sciences
³College of Horticulture Fujian Agriculture and Forestry University, Fuzhou 350002, China

Received:2013-12-16 Accepted:2014-11-13 Online:2015-01-01 Published:2015-04-09
Contact: Deng Chuanyuan
About author:
? These authors contributed equally to this paper

摘要/Abstract

摘要： 应用扫描电子显微镜详细观察了红树族4属、10种、1变种植物次生木质部管状分子附物纹孔的分布和形态, 应用Carnoy 2.0软件和扫描电镜下采集的照片, 测定了管间梯状附物纹孔丰富度指标和管间梯状纹孔数量特征指标。结果显示, 红树族植物次生木质部管状分子侧壁具附物纹孔。所观察的植物附物纹孔的分布和形态变化大。附物纹孔丰富度指标与管间梯状纹孔数量特征指标的逐步回归分析表明, 导管侧壁附物纹孔丰富度随纹孔口面积百分比的增大而增大。据此推测, 红树族植物附物纹孔丰富度与纹孔几何构造及数量特征有关。附物纹孔是红树族植物稳定存在的一个木材解剖性状。综合生态-系统演化的观点, 红树族植物具附物纹孔可能是受系统演化关系控制的生态适应结果。

Abstract: We used SEM to investigate the distribution and micromorphology of vestured pits in the secondary xylem of 10 species and one variety representing all 4 genera within the tribe Rhizophoreae. Richness indexes of intervascular vestured pits and quantitative features of scalariform intervascular bordered pits were measured on SEM images by use of Carnoy 2.0. Vestured pits were present in vessel elements and tracheary elements of secondary xylem of the specimens and varied considerably in distribution and micromorphology. Stepwise regression analysis of richness indexes and quantitative features of scalariform intervascular bordered pits indicated increased richness indexes, including frequency of vestured vessel inner aperture, outer aperture and pit chamber with increasing aperture fraction, which suggests that richness of intervascular vestured pits was associated with the geometric structure of scalariform intervascular bordered pits. We demonstrate consistent occurrence of intervascular vestured pits in the tribe Rhizophoreae from the common presence of intervascular vestured pits in our specimens. In terms of ecophylogeny, vestured pits occurring in the vessel elements and tracheary elements of the secondary xylem of the tribe Rhizophoreae might be an ecologically adaptive character controlled by phylogeny.

邓传远, 辛桂亮, 张万超, 郭素枝, 薛秋华, 赖钟雄, 叶露莹. 红树族植物次生木质部附物纹孔的电镜观测. 植物学报, 2015, 50(1): 90-99.

Chuanyuan Deng, Guiliang Xin, Wanchao Zhang, Suzhi Guo, Qiuhua Xue, Zhongxiong Lai, Luying Ye. SEM Observations and Measurements of Vestured Pits of the Secondary Xylem in the Tribe Rhizophoreae. Chinese Bulletin of Botany, 2015, 50(1): 90-99.

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链接本文: https://www.chinbullbotany.com/CN/10.3724/SP.J.1259.2015.00090

https://www.chinbullbotany.com/CN/Y2015/V50/I1/90

图/表 3

图1 红树族植物附物纹孔的分布与微形态 (A) 角果木, 管间纹孔, 导管分子侧壁部分内纹孔口具附物(a), 或不具附物(b); (B) 秋茄, 管间纹孔, 箭头示导管分子侧壁外纹孔口具附物; (C) 红海兰, 导管-薄壁细胞间纹孔, 箭头示导管分子侧壁纹孔室具附物; (D) 红茄苳, 管间纹孔, 导管分子侧壁外纹孔口和纹孔室没有附物(Dh: 纹孔膜水平直径; Dv: 纹孔膜垂直直径; LAOA: 外纹孔口长轴直径; SAOA: 外纹孔口短轴直径); (E) 澳洲角果木, 管间纹孔, 导管分子侧壁内纹孔口没有附物(LAIA: 内纹孔口长轴直径; SAIA: 内纹孔口短轴直径); (F) 红海兰, 管间纹孔, 箭头示导管侧壁内纹孔口具星散点状附物; (G) 红海兰, 导管-薄壁细胞间纹孔, 箭头示导管侧壁内纹孔口具星散点状附物; (H) 美洲大红树, 管间纹孔, 箭头示导管侧壁纹孔室具星散棒状附物; (I) 尖瓣海莲, 导管-薄壁细胞间纹孔, 箭头示导管侧壁外纹孔口具星散丝状附物; (J) 木榄, 管间纹孔, 箭头示导管侧壁外纹孔口具叉状附物; (K) 木榄, 导管-薄壁细胞间纹孔, 箭头示导管侧壁内纹孔口具叉状附物; (L) 红树, 管间纹孔, 箭头示叉状附物常与点状、棒状附物一起分布于内外纹孔口, 并向纹孔室或纹孔口中心突出, 使内外纹孔口的形状呈不规则状; (M) 木榄, 导管-薄壁细胞间纹孔, 箭头示点状与棒状附物相互连接, 聚集成团块状阻塞内纹孔口; (N) 秋茄, 管间纹孔, 箭头示导管侧壁外纹孔口和纹孔室点状(和棒状)聚集附物; (O) 木榄, 导管-薄壁细胞间纹孔, 箭头示导管侧壁内纹孔口点状(和棒状)聚集附物; (P) 海莲, 导管-薄壁细胞间纹孔, 箭头示外纹孔口和纹孔室相互交织的丝状附物, 并延伸到纹孔外; (Q) 角果木, 管间纹孔, 箭头示内纹孔口相互交织的丝状附物, 并延伸到导管内壁; (R) 澳洲角果木, 管间纹孔, 箭头示内纹孔口与导管内壁相连接的成片块状附物

Figure 1 Distribution and micromorphology of vestured pits in the study species of tribe Rhizophoreae (A) Ceriops tagal, intervascular pits in the wall of a vessel element viewed from the lumen side, the inner pit apertures are vestured (a), or non-vestured (b); (B) Kandelia candel, intervascular pits in the wall of a vessel element viewed from the outer surface, arrow shows the outer pit apertures are vestured; (C) Rhizophora stylosa, vessel-ray parenchyma pits in the wall of a vessel element viewed from the outer surface, arrow shows the pit chambers are vestured; (D) Rhizophora mucronata, non-vestured outer pit apertures and non-vestured pit chambers in the wall of a vessel element viewed from the outer surface (Dh: Horizontal pit membrane diameter; Dv: Vertical pit membrane diameter; LAOA: The longest axis of the outer aperture; SAOA: The shortest axis of the outer aperture); (E) Ceriops australis, non-vestured inner pit apertures in the wall of a vessel element viewed from the lumen side (LAIA: The longest axis of the inner aperture; SAIA: The shortest axis of the inner aperture); (F) Rhizophora stylosa, intervascular pits in the wall of a vessel element viewed from lumen side, arrow shows scattered dot-shaped vestures occur in the inner pit apertures; (G) Rhizophora stylosa, vessel-ray parenchyma pits in the wall of a vessel element viewed from the lumen side, arrow shows scattered dot-shaped vestures occur in the inner pit apertures; (H) Rhizophora mangle, intervascular pits in the wall of a vessel element viewed from the outer surface, arrow shows scattered rod-shaped vestures occur in the pit chambers; (I) Bruguiera sexangula var. rhynchopetala, vessel-ray parenchyma pits in the wall of a vessel element viewed from the outer surface, arrow shows scattered silk-like vestures occur in the outer pit apertures; (J) Bruguiera gymnorrhiza, intervascular pits in the wall of a vessel element viewed from the outer surface, arrow shows forked vestures occur in the outer pit apertures; (K) Bruguiera gymnorrhiza, vessel-ray parenchyma pits in the wall of a vessel element viewed from the lumen side, arrow shows forked vestures occur in the inner pit apertures; (L) Rhizophora apiculata, intervascular pits in the wall of a vessel element viewed from the lumen side, arrow shows forked vestures together with dot-like vestures arise from the margin of the inner pit aperture, therefore, the irregular shape of inner pit aperture develop; (M) Bruguiera gymnorrhiza, vessel-ray parenchyma pits in the wall of a vessel element viewed from the lumen side, arrow shows dot-like and rod-like vestures aggregate into a agglomerate and occlude the inner pit aperture; (N) Kandelia candel, intervascular pits in the wall of a vessel element viewed from the outer surface, arrow shows aggregate dot-shaped or rod-shaped vestures occur in the outer pit apertures and pit chambers; (O) Bruguiera gymnorrhiza, vessel-ray parenchyma pits in the wall of a vessel element viewed from the outer surface, arrow shows aggregate dot-shaped or rod-shaped vestures occur in the outer pit apertures and pit chambers; (P) Bruguiera sexangula, vessel-ray parenchyma pits in the wall of a vessel element viewed from the outer surface, arrow shows filament-like vestures in the outer pit apertures or pit chambers interweave to form into a rather compact network and extend to the outside of pits; (Q) Ceriops tagal, intervascular pits in the wall of a vessel element viewed from the lumen side, arrow shows filament-like vestures in the inner pit apertures interweave to form into a network and extend to the inner vessel wall; (R) Ceriops australis, arrow shows sheeted vestures occurred in the wall of a vessel element viewed from the lumen side and obscure the inner pit apertures

表1 红树族植物纹孔附物的分布和微形态变化

Table 1 Distribution and micromorphological variations of vestures in pits of Rhizophoreae species studied

Scientific name	Inner pit aperture							Outer pit aperture
Scientific name	T1	T2	T3	T4	T5	T6	T7		T1	T2	T3	T4	T5	T6	T7	T1	T2	T3	T4	T5	T6	T7
Bruguiera gyrmnorrhiza	+	+	+	++	++	++	+	+		+	–	++	++	+	++	+	+	-	-	++	+	+
B. parviflora	+	+	-	+	-	-	+	+		+	-	+	-	-		+	+	-	-	-	-	+
B. sexangula	+	+	+	++	++	+	+	+		+	-	+	-	+	++	+	+	-	-	-	+	+
var. rhynchuopetala	++	+	-	+	+	-	-	+		++	+	+	-	-	-	+	+	-	-	-	-	-
Ceriops australis	+	+	+	++	++	+	++	+		+	+	+	-	-	-	++	+	-	-	-	-	-
C. tagal	++	+	+	++	++	++	+	++		+	-	++	-	+	+	+	++	-	+	-	-	+
Kandelia candel	++	+	+	++	++	++	+	+		+	+	++	+	-	++	+	+	-	-	+	+	+
Rhizophora apiculata	+	+	+	++	++	+	++	+		+	-	+	-	-	+	+	-	-	-	+	-	-
R. mangle	+	+	+	++	++	+	++	+		+	+	+	-	-	++	+	+	+	-	-	-	+
R. mucronata	+	+	+	+	++	+	+	+		+	-	+	+	-	+	+	++	+	-	+	-	-
R. stylosa	++	+	+	+	++	++	+	+		+	+	+	+	-	+	+	+	-	+	+	+	+

表2 红树族10种、1变种植物管间附物纹孔丰富度和管间纹孔的数量特征(平均值±标准差)

Table 2 Richness indexes of intervascular vestured pits and quantitative features of scalariform intervascular bordered pits in 10 species and 1 variety of tribe Rhizophoreae (means±SD)

Species	FVIA (%)	FVOA (%)	FVPC (%)	Dh (μm)	Dv (μm)	LAOA (μm)	SAOA (μm)	LAIA (μm)	SAIA (μm)	AP_f	Fap (%)	PA (μm²)	OPA (μm²)	IPA (μm²)
Bruguiera gymnorrhiza	27.77± 44.57	16.40± 36.07	14.53± 34.25	23.47± 7.26	4.72± 1.90	12.56± 7.27	1.98± 1.05	18.62± 7.20	1.43± 0.62	7.56± 6.05	21.10± 9.16	95.68± 67.45	21.31± 19.68	20.94± 12.36
B. parviflora	19.09± 34.63	7.78± 23.00	5.75± 21.40	23.02± 7.50	4.57± 1.03	14.65± 5.97	1.02± 0.78	14.45± 5.62	1.02± 0.58	18.83± 11.66	14.74± 11.90	87.60± 44.24	13.02± 13.73	11.58± 10.60
B. sexangula	26.25± 39.06	15.06± 37.89	6.76± 19.60	27.47± 10.40	3.72± 1.21	21.72± 8.78	1.13± 0.90	17.94± 5.02	1.44± 0.95	24.91± 12.76	23.12± 12.37	83.49± 48.62	20.83± 18.34	20.99± 18.61
B. sexangula var. rhynchuopetala	20.54± 40.21	11.21± 39.39	4.21± 14.24	23.51± 9.00	3.70± 1.12	17.96± 7.53	0.78± 0.30	12.93± 4.34	1.28± 0.63	25.55± 12.11	16.06± 5.71	74.13± 52.31	12.11± 9.44	13.21± 8.34
Ceriops australis	21.82± 35.69	7.04± 21.32	8.83± 24.26	25.00± 9.14	3.46± 1.05	20.25± 8.07	0.79± 0.48	15.14± 5.58	0.71± 0.44	32.74± 17.60	18.23± 11.00	71.26± 40.25	12.82± 9.06	8.94± 7.87
C. tagal	25.01± 40.81	1.53± 7.97	0.48± 3.09	15.00± 3.78	2.68± 0.99	12.29± 3.63	0.59± 0.21	18.44± 7.58	1.31± 1.10	24.16± 13.64	21.84± 15.01	19.89± 20.22	18.10± 14.75	21.75± 25.90
Kandelia candel	31.56± 39.40	5.82± 18.16	1.04± 8.14	24.76± 7.75	4.98± 1.64	18.90± 5.72	1.42± 0.59	20.91± 7.75	1.33± 0.56	15.28± 6.53	22.13± 7.52	105.31± 71.20	22.89± 15.84	24.10± 18.28
Rhizophora apiculata	25.94± 35.87	8.83± 22.65	1.71± 7.14	29.84± 10.74	4.80± 1.49	21.30± 9.92	1.28± 1.05	27.98± 11.78	1.67± 0.71	21.62± 14.31	17.86± 9.33	117.43± 68.31	24.25± 34.96	35.69± 20.76
R. mangle	25.97± 37.30	5.82± 19.78	2.17± 13.38	18.03± 5.08	3.33± 0.61	13.82± 4.23	1.03± 0.46	13.32± 4.04	1.23± 1.19	16.04± 7.71	13.57± 9.76	189.41± 63.61	41.48± 13.45	13.14± 15.49
R. mucronata	26.93± 42.66	9.40± 25.61	6.22± 20.95	26.47± 5.77	4.11± 1.17	19.52± 4.08	1.03± 0.44	19.34± 4.77	1.42± 0.50	22.80± 12.01	19.24± 10.79	88.04± 42.51	16.09± 9.57	21.92± 9.58
R. stylosa	34.43± 45.00	26.97± 42.95	15.79± 37.46	18.89± 6.36	2.18± 1.32	11.99± 5.77	1.17± 0.59	18.70± 2.34	1.78± 0.32	14.47± 12.97	41.60± 19.29	33.10± 31.83	9.50± 3.39	26.34± 5.91
PI	0.45	0.94	0.97	0.50	0.56	0.45	0.70	0.54	0.60	0.77	0.67	0.89	0.77	0.75

表2 红树族10种、1变种植物管间附物纹孔丰富度和管间纹孔的数量特征(平均值±标准差)

Table 2 Richness indexes of intervascular vestured pits and quantitative features of scalariform intervascular bordered pits in 10 species and 1 variety of tribe Rhizophoreae (means±SD)

Species	FVIA (%)	FVOA (%)	FVPC (%)	Dh (μm)	Dv (μm)	LAOA (μm)	SAOA (μm)	LAIA (μm)	SAIA (μm)	AP_f	Fap (%)	PA (μm²)	OPA (μm²)	IPA (μm²)
Bruguiera gymnorrhiza	27.77± 44.57	16.40± 36.07	14.53± 34.25	23.47± 7.26	4.72± 1.90	12.56± 7.27	1.98± 1.05	18.62± 7.20	1.43± 0.62	7.56± 6.05	21.10± 9.16	95.68± 67.45	21.31± 19.68	20.94± 12.36
B. parviflora	19.09± 34.63	7.78± 23.00	5.75± 21.40	23.02± 7.50	4.57± 1.03	14.65± 5.97	1.02± 0.78	14.45± 5.62	1.02± 0.58	18.83± 11.66	14.74± 11.90	87.60± 44.24	13.02± 13.73	11.58± 10.60
B. sexangula	26.25± 39.06	15.06± 37.89	6.76± 19.60	27.47± 10.40	3.72± 1.21	21.72± 8.78	1.13± 0.90	17.94± 5.02	1.44± 0.95	24.91± 12.76	23.12± 12.37	83.49± 48.62	20.83± 18.34	20.99± 18.61
B. sexangula var. rhynchuopetala	20.54± 40.21	11.21± 39.39	4.21± 14.24	23.51± 9.00	3.70± 1.12	17.96± 7.53	0.78± 0.30	12.93± 4.34	1.28± 0.63	25.55± 12.11	16.06± 5.71	74.13± 52.31	12.11± 9.44	13.21± 8.34
Ceriops australis	21.82± 35.69	7.04± 21.32	8.83± 24.26	25.00± 9.14	3.46± 1.05	20.25± 8.07	0.79± 0.48	15.14± 5.58	0.71± 0.44	32.74± 17.60	18.23± 11.00	71.26± 40.25	12.82± 9.06	8.94± 7.87
C. tagal	25.01± 40.81	1.53± 7.97	0.48± 3.09	15.00± 3.78	2.68± 0.99	12.29± 3.63	0.59± 0.21	18.44± 7.58	1.31± 1.10	24.16± 13.64	21.84± 15.01	19.89± 20.22	18.10± 14.75	21.75± 25.90
Kandelia candel	31.56± 39.40	5.82± 18.16	1.04± 8.14	24.76± 7.75	4.98± 1.64	18.90± 5.72	1.42± 0.59	20.91± 7.75	1.33± 0.56	15.28± 6.53	22.13± 7.52	105.31± 71.20	22.89± 15.84	24.10± 18.28
Rhizophora apiculata	25.94± 35.87	8.83± 22.65	1.71± 7.14	29.84± 10.74	4.80± 1.49	21.30± 9.92	1.28± 1.05	27.98± 11.78	1.67± 0.71	21.62± 14.31	17.86± 9.33	117.43± 68.31	24.25± 34.96	35.69± 20.76
R. mangle	25.97± 37.30	5.82± 19.78	2.17± 13.38	18.03± 5.08	3.33± 0.61	13.82± 4.23	1.03± 0.46	13.32± 4.04	1.23± 1.19	16.04± 7.71	13.57± 9.76	189.41± 63.61	41.48± 13.45	13.14± 15.49
R. mucronata	26.93± 42.66	9.40± 25.61	6.22± 20.95	26.47± 5.77	4.11± 1.17	19.52± 4.08	1.03± 0.44	19.34± 4.77	1.42± 0.50	22.80± 12.01	19.24± 10.79	88.04± 42.51	16.09± 9.57	21.92± 9.58
R. stylosa	34.43± 45.00	26.97± 42.95	15.79± 37.46	18.89± 6.36	2.18± 1.32	11.99± 5.77	1.17± 0.59	18.70± 2.34	1.78± 0.32	14.47± 12.97	41.60± 19.29	33.10± 31.83	9.50± 3.39	26.34± 5.91
PI	0.45	0.94	0.97	0.50	0.56	0.45	0.70	0.54	0.60	0.77	0.67	0.89	0.77	0.75

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红树族植物次生木质部附物纹孔的电镜观测

SEM Observations and Measurements of Vestured Pits of the Secondary Xylem in the Tribe Rhizophoreae

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