红海榄木材结构的生态解剖

doi:10.17521/cjpe.2015.0058

摘要/Abstract

摘要：

红树林是海岸重要生态关键区, 在维护生物多样性和沿海地区生态安全等方面发挥着重要作用。红海榄(Rhizophora stylosa)是我国红树林生态系统的建群种之一。为了揭示红海榄次生木质部解剖特征可塑性的生态适应意义, 该文测定了海南省东寨港红树林自然保护区6个红海榄种群18个采样点的土壤理化性质, 应用显微镜和电镜详细观测了各采样点生长的18株红海榄植株次生木质部的形态解剖特征, 并测量了红海榄次生木质部的数量解剖特征。不同样地的红海榄次生木质部都具有纤维状导管和环管管胞、螺旋雕纹和附物、生长轮、薄壁细胞(含淀粉粒)等结构。这些特化结构具有生态适应意义, 在潮间带高盐生境中能促进水分输导的安全性。不同生境中红海榄次生木质部数量解剖特征可塑性大, 利用逐步回归分析方法分析了土壤理化因子与红海榄次生木质部数量特征的关系。结果表明: 土壤全盐含量、土壤Mn²⁺含量、土壤Na⁺含量、土壤Cl^-含量、土壤Ca²⁺含量、土壤有机质含量、土壤全磷含量和土壤pH值对次生木质部数量特征的影响达到极显著水平, 不同样地红海榄次生木质部数量特征的变化是红海榄适应异质生境的结果。

关键词: 生态解剖, 红海榄, 土壤理化因子, 木材结构

Abstract: Aims

Mangrove forest is desirable for studying variations in wood structure along an ecological gradient because mangroves are subjected to considerable habitat changes apart from salt stress within a small area. To elucidate the adaptive capability of wood structures towards fluctuating environmental conditions, variations in wood structures were investigated in 18 individuals of Rhizophora stylosa representing 6 populations along a natural soil physicochemical gradient in the National Nature Reserve of Dongzhai Harbor, Hainan Province.

Methods

Soil physicochemical properties were determined at the sites of 18 sampling trees in six R. stylosa populations. The anatomical characteristics of the secondary xylem were studied in details in the 18 trees by means of light microscopy, laser scanning confocal microscopy, scanning electron microscopy and transmission electron microscopy. Variations in the quantitative wood anatomical features in R. stylosa were assessed in details. Relationships between soil physicochemical variables and the quantitative wood anatomical features were analyzed by means of statistical methods.

Important findings

Some common specialized wood structures were observed in R. stylosa growing in different habitats, suggesting that these features may function for safely conducting water under high negative pressure and are thus adaptive to intertidal habitats. These common features include the occurrence of: 1) some fibriform vessel elements and a few vasicentric tracheids; 2) vesturing in pits of vessels and helical structures on vessel walls; 3) growth rings; 4) starch grains in ray cells and septate fibers. The quantitative anatomical characteristics have great plasticity in response to heterogeneous habitats. Stepwise regression analyses revealed that the total salt, contents of Mn²⁺, Na⁺, Cl^-, Ca²⁺, organic matters, and total phosphorus of soils, and soil pH all have significant effects on quantitative wood anatomical features. Variations in the quantitative wood anatomical features in R. stylosa growing at different sites are adaptive to fluctuating environmental conditions in the intertidal areas.

Key words: ecological anatomy, Rhizophora stylosa, soil physicochemical factors, wood structure

邓传远, 郑俊鸣, 张万超, 郭素枝, 薛秋华, 叶露莹, 孙建文. 红海榄木材结构的生态解剖. 植物生态学报, 2015, 39(6): 604-615. DOI: 10.17521/cjpe.2015.0058

DENG Chuan-Yuan,ZHENG Jun-Ming,ZHANG Wan-Chao,GUO Su-Zhi,XUE Qiu-Hua,YE Lu-Ying,SUN Jian-Wen. Ecological wood anatomy of Rhizophora stylosa. Chinese Journal of Plant Ecology, 2015, 39(6): 604-615. DOI: 10.17521/cjpe.2015.0058

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2015.0058

https://www.plant-ecology.com/CN/Y2015/V39/I6/604

图/表 5

参考文献 34

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采样点Sampling sites		土壤中离子含量 Ion content in soil							TSC (g·kg^-1)	pH	TP (g·kg^-1)	OM (g·kg^-1)	TN (g·kg^-1)
采样点Sampling sites		Cl^- (g·kg^-1)	K⁺ (g·kg^-1)	Ca²⁺ (g·kg^-1)	Na⁺ (g·kg^-1)	Mg²⁺ (g·kg^-1)	Al³⁺ (µg·g^-1)	Mn²⁺ (µg·g^-1)	TSC (g·kg^-1)	pH	TP (g·kg^-1)	OM (g·kg^-1)	TN (g·kg^-1)
1	A	4.67 ± 1.15	0.725 ± 0.075	0.15 ± 0.043	4.01 ± 0.13	0.14 ± 0.012	73.59 ± 8.57	557.76 ± 88.64	11.23 ± 1.07	3.02 ± 0.04	0.39 ± 0.04	4.29 ± 0.63	0.75 ± 0.17
	B	4.38 ± 1.51	0.700 ± 0.053	0.16 ± 0.012	3.77 ± 0.09	0.16 ± 0.019	71.82 ± 6.34	504.34 ± 63.24	12.15 ± 1.65	3.50 ± 0.06	0.31 ± 0.04	4.07 ± 0.82	0.88 ± 0.19
	C	4.21 ± 1.26	0.752 ± 0.082	0.18 ± 0.029	4.23 ± 0.19	0.13 ± 0.023	66.57 ± 8.91	556.32 ± 75.52	10.12 ± 2.01	4.01 ± 0.07	0.48 ± 0.06	5.40 ± 0.67	0.95 ± 0.12
2	A	5.36 ± 1.11	0.843 ± 0.098	0.35 ± 0.015	4.10 ± 0.23	0.17 ± 0.025	63.24 ± 10.02	499.72 ± 101.22	11.49 ± 1.32	7.24 ± 0.05	1.15 ± 0.06	8.42 ± 0.89	1.54 ± 0.37
	B	5.42 ± 1.21	0.792 ± 0.029	0.29 ± 0.019	3.27 ± 0.15	0.12 ± 0.014	64.25 ± 9.58	510.07 ± 97.25	11.81 ± 1.43	6.92 ± 0.04	1.14 ± 0.09	8.31 ± 0.48	1.49 ± 0.31
	C	4.85 ± 1.03	0.834 ± 0.092	0.27 ± 0.021	3.73 ± 0.11	0.11 ± 0.010	58.61 ± 9.12	505.36 ± 107.45	11.49 ± 1.52	7.65 ± 0.11	1.11 ± 0.06	7.72 ± 0.64	1.26 ± 0.28
3	A	6.87 ± 1.06	0.670 ± 0.095	0.31 ± 0.012	4.34 ± 0.09	0.15 ± 0.022	49.87 ± 6.33	621.67 ± 94.68	12.36 ± 1.09	7.12 ± 0.08	1.16 ± 0.10	7.68 ± 0.73	1.56 ± 0.43
	B	6.98 ± 0.96	0.540 ± 0.084	0.37 ± 0.024	4.17 ± 0.22	0.25 ± 0.017	54.55 ± 7.45	687.69 ± 99.32	13.86 ± 1.17	6.72 ± 0.08	1.05 ± 0.09	7.31 ± 0.82	1.72 ± 0.32
	C	6.52 ± 1.24	0.610 ± 0.076	0.33 ± 0.019	4.05 ± 0.16	0.16 ± 0.021	47.41 ± 4.13	630.21 ± 103.25	14.37 ± 1.58	7.02 ± 0.07	0.91 ± 0.08	7.17 ± 0.77	1.11 ± 0.24
4	A	7.87 ± 1.32	0.727 ± 0.085	0.42 ± 0.016	4.68 ± 0.24	0.12 ± 0.022	43.54 ± 8.41	505.61 ± 75.65	17.75 ± 1.35	6.94 ± 0.14	0.86 ± 0.05	6.75 ± 0.69	0.59 ± 0.21
	B	7.85 ± 1.22	0.746 ± 0.046	0.49 ± 0.025	4.44 ± 0.29	0.10 ± 0.018	38.81 ± 7.59	545.32 ± 98.64	16.36 ± 1.87	6.05 ± 0.09	0.77 ± 0.02	7.50 ± 0.72	0.49 ± 0.19
	C	6.94 ± 1.19	0.790 ± 0.091	0.45 ± 0.028	4.95 ± 0.13	0.13 ± 0.029	41.21 ± 5.39	571.88 ± 82.63	19.38 ± 1.96	6.79 ± 0.03	0.91 ± 0.03	6.21 ± 0.77	0.65 ± 0.28
5	A	8.63 ± 0.98	0.521 ± 0.102	0.34 ± 0.014	3.41 ± 0.12	0.15 ± 0.031	44.74 ± 11.25	569.21 ± 97.54	12.70 ± 1.23	6.85 ± 0.07	1.06 ± 0.14	6.69 ± 0.34	1.27 ± 0.41
	B	8.31 ± 1.09	0.485 ± 0.079	0.37 ± 0.015	3.77 ± 0.17	0.15 ± 0.027	37.22 ± 7.73	528.96 ± 102.57	12.12 ± 1.54	5.64 ± 0.04	1.25 ± 0.13	5.70 ± 0.45	1.12 ± 0.26
	C	9.23 ± 1.13	0.519 ± 0.099	0.33 ± 0.021	4.28 ± 0.14	0.12 ± 0.022	35.98 ± 10.36	584.21 ± 85.95	13.23 ± 2.01	6.64 ± 0.06	1.30 ± 0.21	6.69 ± 0.38	1.13 ± 0.33
6	A	10.95 ± 0.97	0.347 ± 0.086	0.06 ± 0.011	5.16 ± 0.15	0.07 ± 0.013	28.93 ± 6.12	129.43 ± 64.57	14.86 ± 1.82	6.87 ± 0.10	0.30 ± 0.04	9.39 ± 0.66	1.20 ± 0.25
	B	10.36 ± 1.13	0.358 ± 0.056	0.08 ± 0.016	5.90 ± 0.26	0.09 ± 0.014	33.65 ± 3.25	102.57 ± 81.47	15.42 ± 1.52	6.53 ± 0.05	0.37 ± 0.02	10.80 ± 0.47	1.48 ± 0.31
	C	12.24 ± 1.28	0.301 ± 0.092	0.05 ± 0.013	5.81 ± 0.32	0.05 ± 0.011	26.35 ± 4.99	122.33 ± 57.39	14.52 ± 1.69	6.93 ± 0.06	0.27 ± 0.04	10.20 ± 0.52	1.34 ± 0.23

采样点Sampling sites		土壤中离子含量 Ion content in soil							TSC (g·kg^-1)	pH	TP (g·kg^-1)	OM (g·kg^-1)	TN (g·kg^-1)
采样点Sampling sites		Cl^- (g·kg^-1)	K⁺ (g·kg^-1)	Ca²⁺ (g·kg^-1)	Na⁺ (g·kg^-1)	Mg²⁺ (g·kg^-1)	Al³⁺ (µg·g^-1)	Mn²⁺ (µg·g^-1)	TSC (g·kg^-1)	pH	TP (g·kg^-1)	OM (g·kg^-1)	TN (g·kg^-1)
1	A	4.67 ± 1.15	0.725 ± 0.075	0.15 ± 0.043	4.01 ± 0.13	0.14 ± 0.012	73.59 ± 8.57	557.76 ± 88.64	11.23 ± 1.07	3.02 ± 0.04	0.39 ± 0.04	4.29 ± 0.63	0.75 ± 0.17
	B	4.38 ± 1.51	0.700 ± 0.053	0.16 ± 0.012	3.77 ± 0.09	0.16 ± 0.019	71.82 ± 6.34	504.34 ± 63.24	12.15 ± 1.65	3.50 ± 0.06	0.31 ± 0.04	4.07 ± 0.82	0.88 ± 0.19
	C	4.21 ± 1.26	0.752 ± 0.082	0.18 ± 0.029	4.23 ± 0.19	0.13 ± 0.023	66.57 ± 8.91	556.32 ± 75.52	10.12 ± 2.01	4.01 ± 0.07	0.48 ± 0.06	5.40 ± 0.67	0.95 ± 0.12
2	A	5.36 ± 1.11	0.843 ± 0.098	0.35 ± 0.015	4.10 ± 0.23	0.17 ± 0.025	63.24 ± 10.02	499.72 ± 101.22	11.49 ± 1.32	7.24 ± 0.05	1.15 ± 0.06	8.42 ± 0.89	1.54 ± 0.37
	B	5.42 ± 1.21	0.792 ± 0.029	0.29 ± 0.019	3.27 ± 0.15	0.12 ± 0.014	64.25 ± 9.58	510.07 ± 97.25	11.81 ± 1.43	6.92 ± 0.04	1.14 ± 0.09	8.31 ± 0.48	1.49 ± 0.31
	C	4.85 ± 1.03	0.834 ± 0.092	0.27 ± 0.021	3.73 ± 0.11	0.11 ± 0.010	58.61 ± 9.12	505.36 ± 107.45	11.49 ± 1.52	7.65 ± 0.11	1.11 ± 0.06	7.72 ± 0.64	1.26 ± 0.28
3	A	6.87 ± 1.06	0.670 ± 0.095	0.31 ± 0.012	4.34 ± 0.09	0.15 ± 0.022	49.87 ± 6.33	621.67 ± 94.68	12.36 ± 1.09	7.12 ± 0.08	1.16 ± 0.10	7.68 ± 0.73	1.56 ± 0.43
	B	6.98 ± 0.96	0.540 ± 0.084	0.37 ± 0.024	4.17 ± 0.22	0.25 ± 0.017	54.55 ± 7.45	687.69 ± 99.32	13.86 ± 1.17	6.72 ± 0.08	1.05 ± 0.09	7.31 ± 0.82	1.72 ± 0.32
	C	6.52 ± 1.24	0.610 ± 0.076	0.33 ± 0.019	4.05 ± 0.16	0.16 ± 0.021	47.41 ± 4.13	630.21 ± 103.25	14.37 ± 1.58	7.02 ± 0.07	0.91 ± 0.08	7.17 ± 0.77	1.11 ± 0.24
4	A	7.87 ± 1.32	0.727 ± 0.085	0.42 ± 0.016	4.68 ± 0.24	0.12 ± 0.022	43.54 ± 8.41	505.61 ± 75.65	17.75 ± 1.35	6.94 ± 0.14	0.86 ± 0.05	6.75 ± 0.69	0.59 ± 0.21
	B	7.85 ± 1.22	0.746 ± 0.046	0.49 ± 0.025	4.44 ± 0.29	0.10 ± 0.018	38.81 ± 7.59	545.32 ± 98.64	16.36 ± 1.87	6.05 ± 0.09	0.77 ± 0.02	7.50 ± 0.72	0.49 ± 0.19
	C	6.94 ± 1.19	0.790 ± 0.091	0.45 ± 0.028	4.95 ± 0.13	0.13 ± 0.029	41.21 ± 5.39	571.88 ± 82.63	19.38 ± 1.96	6.79 ± 0.03	0.91 ± 0.03	6.21 ± 0.77	0.65 ± 0.28
5	A	8.63 ± 0.98	0.521 ± 0.102	0.34 ± 0.014	3.41 ± 0.12	0.15 ± 0.031	44.74 ± 11.25	569.21 ± 97.54	12.70 ± 1.23	6.85 ± 0.07	1.06 ± 0.14	6.69 ± 0.34	1.27 ± 0.41
	B	8.31 ± 1.09	0.485 ± 0.079	0.37 ± 0.015	3.77 ± 0.17	0.15 ± 0.027	37.22 ± 7.73	528.96 ± 102.57	12.12 ± 1.54	5.64 ± 0.04	1.25 ± 0.13	5.70 ± 0.45	1.12 ± 0.26
	C	9.23 ± 1.13	0.519 ± 0.099	0.33 ± 0.021	4.28 ± 0.14	0.12 ± 0.022	35.98 ± 10.36	584.21 ± 85.95	13.23 ± 2.01	6.64 ± 0.06	1.30 ± 0.21	6.69 ± 0.38	1.13 ± 0.33
6	A	10.95 ± 0.97	0.347 ± 0.086	0.06 ± 0.011	5.16 ± 0.15	0.07 ± 0.013	28.93 ± 6.12	129.43 ± 64.57	14.86 ± 1.82	6.87 ± 0.10	0.30 ± 0.04	9.39 ± 0.66	1.20 ± 0.25
	B	10.36 ± 1.13	0.358 ± 0.056	0.08 ± 0.016	5.90 ± 0.26	0.09 ± 0.014	33.65 ± 3.25	102.57 ± 81.47	15.42 ± 1.52	6.53 ± 0.05	0.37 ± 0.02	10.80 ± 0.47	1.48 ± 0.31
	C	12.24 ± 1.28	0.301 ± 0.092	0.05 ± 0.013	5.81 ± 0.32	0.05 ± 0.011	26.35 ± 4.99	122.33 ± 57.39	14.52 ± 1.69	6.93 ± 0.06	0.27 ± 0.04	10.20 ± 0.52	1.34 ± 0.23

解剖性状 Anatomical characteristics	生态因子 Ecological factor	回归方程 Regression equation	校正决定系数Adjusted R²
VEL	土壤全盐含量 Total soil salt content	Y _VEL = 344.761 + 12.039X_TSC	0.386^**
RD	土壤Mn²⁺含量 Soil Mn²⁺ content、土壤Na⁺含量 Soil Na⁺ content	Y _RD= 101.890 - 0.042X _Mn2+- 5.777X_Na+	0.726^**
RD50	土壤Mn²⁺含量 Soil Mn²⁺ content、土壤Na⁺含量 Soil Na⁺ content	Y_RD= 128.047 - 0.050X _Mn2+- 5.634X_Na+	0.587^**
TD	土壤全磷含量 Soil total phosphorus content、土壤Mn²⁺含量 Soil Mn²⁺ content、土壤Na⁺含量 Soil Na⁺ content、土壤有机质含量 Soil organic matter content、土壤Cl-含量 Soil Cl^-content	Y_TD= 71.773 + 12.836 X_TP- 0.033X _Mn2+- 3.589X_Na+- 0.901X_{OM +} 0.547 X_Cl-	0.896^**
PA	土壤Mn²⁺含量 Soil Mn²⁺ content、土壤Na⁺含量 Soil Na⁺ content、土壤全磷含量 Soil total phosphorus content、土壤有机质含量 Soil organic matter content	Y_PA= 5262.683 - 3.836 X _Mn2+ + 916.810X_TP-315.615X_Na+ -75.814X_OM	0.900^**
PA50	土壤Mn²⁺含量 Soil Mn²⁺ content、土壤Na⁺含量 Soil Na⁺ content、土壤全磷含量 Soil total phosphorus content、土壤有机质含量 Soil organic matter content	Y_PA50= 8978.891 - 6.142X_Mn2+- 454.463X_Na++ 1191.560X_TP- 177.127X_OM	0.815^**
CA	土壤pH值 Soil pH value	Y_CA= 5.695 + 0.453X_pH	0.385^**
PD	土壤Ca²⁺含量、土壤全盐量 Total soil salt content	Y_PD= 10.615 + 40.737X_Ca2++ 1.608X_TSC	0.762^**
RH	土壤Ca²⁺含量 Soil Ca²⁺ content	Y_RH= 967.204 + 754.122X_Ca2+	0.422^**
RW	土壤Ca²⁺含量 Soil Ca²⁺ content、土壤全磷含量 Soil total phosphorus content、土壤Mn²⁺含量 Soil Mn²⁺ content、土壤Na⁺含量 Soil Na⁺ content	Y_RW= 24.786 + 20.811X_Ca2+- 8.091X_TP+ 0.024X _Mn2++ 3.194X_Na+	0.723^**
RF	土壤全磷含量 Soil total phosphorus content、土壤Ca²⁺含量 Soil Ca²⁺ content	Y_RF= 9.331 - 2.759X_TP + 6.038X_Ca2+	0.562^**
FL	土壤Mn²⁺含量 Soil Mn²⁺ content、土壤Ca²⁺含量 Soil Ca²⁺ content	Y_FL = 1287.519 - 0.687X_Mn2+ + 491.239X_Ca2+	0.819^**
FWT	土壤Mn²⁺含量 Soil Mn²⁺ content	Y_FWT= 6.868 - 0.002X	0.441^**

解剖性状 Anatomical characteristics	生态因子 Ecological factor	回归方程 Regression equation	校正决定系数Adjusted R²
VEL	土壤全盐含量 Total soil salt content	Y _VEL = 344.761 + 12.039X_TSC	0.386^**
RD	土壤Mn²⁺含量 Soil Mn²⁺ content、土壤Na⁺含量 Soil Na⁺ content	Y _RD= 101.890 - 0.042X _Mn2+- 5.777X_Na+	0.726^**
RD50	土壤Mn²⁺含量 Soil Mn²⁺ content、土壤Na⁺含量 Soil Na⁺ content	Y_RD= 128.047 - 0.050X _Mn2+- 5.634X_Na+	0.587^**
TD	土壤全磷含量 Soil total phosphorus content、土壤Mn²⁺含量 Soil Mn²⁺ content、土壤Na⁺含量 Soil Na⁺ content、土壤有机质含量 Soil organic matter content、土壤Cl-含量 Soil Cl^-content	Y_TD= 71.773 + 12.836 X_TP- 0.033X _Mn2+- 3.589X_Na+- 0.901X_{OM +} 0.547 X_Cl-	0.896^**
PA	土壤Mn²⁺含量 Soil Mn²⁺ content、土壤Na⁺含量 Soil Na⁺ content、土壤全磷含量 Soil total phosphorus content、土壤有机质含量 Soil organic matter content	Y_PA= 5262.683 - 3.836 X _Mn2+ + 916.810X_TP-315.615X_Na+ -75.814X_OM	0.900^**
PA50	土壤Mn²⁺含量 Soil Mn²⁺ content、土壤Na⁺含量 Soil Na⁺ content、土壤全磷含量 Soil total phosphorus content、土壤有机质含量 Soil organic matter content	Y_PA50= 8978.891 - 6.142X_Mn2+- 454.463X_Na++ 1191.560X_TP- 177.127X_OM	0.815^**
CA	土壤pH值 Soil pH value	Y_CA= 5.695 + 0.453X_pH	0.385^**
PD	土壤Ca²⁺含量、土壤全盐量 Total soil salt content	Y_PD= 10.615 + 40.737X_Ca2++ 1.608X_TSC	0.762^**
RH	土壤Ca²⁺含量 Soil Ca²⁺ content	Y_RH= 967.204 + 754.122X_Ca2+	0.422^**
RW	土壤Ca²⁺含量 Soil Ca²⁺ content、土壤全磷含量 Soil total phosphorus content、土壤Mn²⁺含量 Soil Mn²⁺ content、土壤Na⁺含量 Soil Na⁺ content	Y_RW= 24.786 + 20.811X_Ca2+- 8.091X_TP+ 0.024X _Mn2++ 3.194X_Na+	0.723^**
RF	土壤全磷含量 Soil total phosphorus content、土壤Ca²⁺含量 Soil Ca²⁺ content	Y_RF= 9.331 - 2.759X_TP + 6.038X_Ca2+	0.562^**
FL	土壤Mn²⁺含量 Soil Mn²⁺ content、土壤Ca²⁺含量 Soil Ca²⁺ content	Y_FL = 1287.519 - 0.687X_Mn2+ + 491.239X_Ca2+	0.819^**
FWT	土壤Mn²⁺含量 Soil Mn²⁺ content	Y_FWT= 6.868 - 0.002X	0.441^**