荒漠灌木的木质部解剖结构与功能权衡——以内蒙西部18种灌木为例

doi:10.11707/j.1001-7488.LYKX20240009

摘要/Abstract

摘要：

目的: 探究荒漠灌木的木质部解剖结构特征、功能权衡关系以及系统发育和气象因子对解剖结构特征的影响，揭示荒漠灌木的水分适应策略，为退化荒漠生态系统的植被保护和恢复提供理论依据。方法: 以内蒙西部18种典型的荒漠灌木为研究对象，测定木质部各组织占比以及其他与水力相关的性状，结合来自全球的木本植物木质部解剖结构数据，分析荒漠灌木木质部典型特征；利用结构方程模型检验木质部机械安全-储存能力-水力效率间的权衡关系，结合系统发育和物种自然分布区的气候因子探讨种间变异性。结果: 1）与来自全球的木本植物平均值相比，荒漠灌木的导管组织和轴向薄壁组织占比较高、纤维组织占比较低；与来自全球的被子植物平均值相比，荒漠灌木的导管密度大、导管水力直径小，导管密度增加的比率和导管水力直径减小的比率使其最大理论导水率基本与全球平均值持平。2） 18种荒漠灌木木质部解剖特征的种间差异较大，同科植物的解剖性状在主成分分析中呈现一定聚集性，除导管密度的系统发育信号显著外，其他性状的系统发育信号均不显著。3）荒漠灌木木质部的机械安全与水力效率、机械安全与储存能力之间均具有显著权衡关系，水力效率和储存能力之间无显著权衡关系。物种自然分布区的年平均气温、最湿季度平均温度、最暖季度平均温度与导管密度负相关，与导管水力直径和导管壁厚度正相关。物种自然分布区的降水量季节性变化与导管壁厚度极显著正相关；年降水量、最湿季度降水量、最暖季度降水量与解剖解剖无显著相关性。结论: 荒漠灌木木质部总体上表现出降低机械支撑、提高储存能力、保障适当的水力效率的木质部干旱适应特征。系统发育和气象因子对导管特征有显著影响，对其他解剖特征的影响较小。

关键词: 导管组织, 纤维组织, 薄壁组织, 功能权衡, 气候因子, 系统发育

Abstract:

Objective: This study aims to uncover the water adaptation strategies of desert shrubs by exploring their xylem anatomical traits, xylem functional trade-offs, and the impact of phylogenetic and meteorological factors on their anatomical structure. Additionally, this research also aims to establish a theoretical foundation for vegetation preservation and restoration in degraded desert ecosystems. Method: In this study, 18 typical desert shrubs were used as experimental objects, and their xylem anatomical characteristics were examined and compared with those of woody plants across the globe. The typical characteristics of desert shrub xylem were analyzed. The trade-offs between the“mechanical safety, storage, and hydraulic efficiency”of the xylem were assessed using a structural equation model (PLS-PM). The interspecific variability was investigated by integrating phylogenetic analysis and climate factors in the natural distribution areas. Result: 1) Compared with the average of woody plants around the world, desert shrubs displayed a higher vessel fraction, axial parenchyma fraction, and a lower fiber fraction. Compared with the global average of angiosperms, desert shrubs exhibited a higher vessel density and smaller vessel hydraulic diameter. The combination of increased vessel density and reduced hydraulic diameter resulted in the maximum theoretical hydraulic conductivity of desert shrubs being basically consistent with the global average. 2) There were significant variations in wood anatomical characteristics among 18 desert shrubs, and the anatomical traits of plants in the same family showed a certain degree of aggregation in principal component analysis. However, apart from the significant phylogenetic signal observed in vessel density, the phylogenetic signals of other traits were not significant. 3) There was a significant trade-off between mechanical safety and hydraulic efficiency and storage capacity, and there was no significant correlation between hydraulic efficiency and storage capacity. Vessel density was negatively correlated with mean annual temperature, mean temperature of wettest quarter, and mean temperature of warmest quarter, while vessel diameter and vessel wall thickness were positively correlated with those climate factors. On the other hand, annual precipitation did not have a significant effect on anatomical traits. Precipitation seasonality exhibited a significantly positive correlation with vessel wall thickness. However, no significant correlation was observed between annual precipitation, wettest quarter precipitation, warmest quarter precipitation, and anatomical anatomy. Conclusion: The xylem of desert shrubs exhibits drought adaptation features that contribute to reduced mechanical support, increased storage capacity, and maintained hydraulic efficiency. The vessel features of these shrubs are significantly influenced by phylogeny and meteorological factors, while other anatomical traits show no significant effects.

Key words: vessel, fiber, parenchyma, functional trade-offs, climate factor, phylogeny

中图分类号:

Q945.79

谭凤森,李清河. 荒漠灌木的木质部解剖结构与功能权衡——以内蒙西部18种灌木为例[J]. 林业科学, 2025, 61(1): 81-94.

Fengsen Tan,Qinghe Li. Anatomical Structure and Functional Trade-Offs of the Xylem in Desert Shrubs in China: a Case Study with 18 Shrubs in Western Inner Mongolia[J]. Scientia Silvae Sinicae, 2025, 61(1): 81-94.

图/表 11

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表2

表1

18种荒漠灌木的木质部性状 (平均值±标准差)①"

物种 Species	RPf (%)	APf (%)	TPf (%)	Ff (%)	VLf (%)	VWf (%)	VWT (%)	Tf (%)	T/D	VD/(N·mm^?2)	D_h/μm	K_th/(kg·s^?1 MPa^?1m^?1)	WD/(g·cm^?3)
白刺Nitraria tangutorum	5.95±1.45	5.22±1.15	11.18±1.45	70.69±2.36	14.12±5.95	3.76±1.85	5.37±0.23	—	0.21±0.14	290±121.39	30.52±4.65	8.17±1.18	0.93+0.07
四合木Tetraena mongolica	7.27±2.51	4.84±1.5	12.11±3.53	77.98±5.86	6.6±1.87	3.31±0.81	6.6±0.15	—	0.29±0.03	162.5±30.95	25.32±5.05	2.03±1.10	0.93+0.05
霸王Zygophyllum xanthoxylum	16.38±3.51	6.76±4.35	23.13±6.13	58.09±8.98	14±5.28	4.78±1.95	5.4±0.24	—	0.21±0.03	275.53±126.93	31.03±4.13	7.24±4.35	0.88+0.04
黄花红砂Reaumuria trigyna	2.5±2.37	24.04±7.58	26.55±7.46	50.74±7.00	14.23±1.77	8.48±1.17	5.32±0.15	—	0.23±0.03	575.69±338.64	23.31±5.29	4.23±1.90	0.61+0.03
蒙古扁桃Prunus mongolica	19.19±6.09	2.96±0.77	22.15±5.72	45.74±3.76	23.73±6.27	8.37±1.93	6.44±0.12	—	0.41±0.01	1184.75±191.35	19.91±2.06	5.54±2.38	0.81+0.05
绵刺Potaninia mongolica	15.16±2.59	3.75±2.48	18.92±2.27	58.9±7.66	16.27±2.82	5.91±1.00	3.16±0.21	—	0.22±0.01	969.3±144.11	16±1.37	1.78±0.57	0.68+0.02
锐枝木蓼Atraphaxis pungens	2.99±2	5.98±1.17	8.97±2.11	73.3±4.93	12.71±2.92	5.01±1.16	4.07±0.11	—	0.29±0.03	901.11±393.35	17.92±3.88	3.44±2.59	0.83+0.1
沙冬青Ammopiptanthus mongolicus	29.14±4.35	1.85±1.29	31±5.50	24.23±11.39	10.98±1.75	4.72±0.77	3.93±0.16	29.07±3.35	0.35±0.03	768.99±190.85	16.1±2.26	1.42±0.54	0.8+0.03
柠条锦鸡儿Caragana korshinskii	5.39±2.42	24.21±5.78	29.59±13.94	44.7±14.39	19±5.14	6.7±1.53	5.71±0.22	—	0.23±0.01	405.21±89.46	29.53±4.16	8.89±4.43	0.62+0.02
毛刺锦鸡儿Caragana tibetica	10.9±3.55	9.97±1.3	20.87±7.23	41.48±7.76	24.55±5.81	13.1±4.55	4.56±0.16	—	0.3±0.04	1933.92±405.03	18.53±3.67	4.7±2.58	0.57+0.03
小叶锦鸡儿Caragana microphylla	9.15±4.29	16.31±2.39	25.46±3.89	56.79±6.31	10.81±3.05	6.94±1.55	5.29±0.40	—	0.34±0.06	840.03±469.46	17.77±2.46	2.16±1.08	0.56+0.03
狭叶锦鸡儿Caragana stenophylla	8.5±2.26	15.69±1.77	24.19±2.33	42.81±5.82	19.18±3.53	13.83±4.09	4.43±0.16	—	0.4±0.08	2085.01±225.71	15.02±2.36	2.91±1.34	0.79+0.04
荒漠锦鸡儿Caragana roborovskyi	7.22±2.88	14.76±1.63	21.98±2.16	45.97±6.99	20.58±5.25	11.47±2.26	4.53±0.17	—	0.28±0.04	988.99±304.37	18.74±1.22	3.46±1.16	0.66+0.03
半日花Helianthemum songaricum	22.57±3.71	7.11±2.53	29.68±2.75	50.78±6.22	11.97±3.34	7.57±2.33	3.7±0.20	—	0.33±0.02	1274.6±489.01	11.94±0.76	0.67±0.16	0.75+0.05
驼绒藜Krascheninnikovia ceratoides	—	15.27±0.85	15.27±0.85	63.27±8.31	13.96±6.12	7.5±2.68	4.73±0.13	—	0.36±0.12	1205.86±185.04	16.4±5.62	2.6±1.57	0.98+0.09
黑沙蒿Artemisia ordosica	4.31±2.84	11.92±1.11	16.23±6.32	58.91±2.69	16.51±1.43	8.35±1.18	4.24±0.17	—	0.22±0.03	614.73±136.08	21.29±2.14	3.31±0.80	0.8+0.03
紫菀木Asterothamnus alyssoides	6.68±2.87	9.06±5.19	15.74±5.31	62.04±4.81	15.09±1.89	7.13±2.25	4.7±0.16	—	0.34±0.05	1108.46±393.09	16.65±2.56	2.19±0.65	0.77+0.05
耆状亚菊Ajania achilleoides	—	21.72±3.56	21.72±8.28	60.79±4.49	11.76±3.61	5.74±1.54	4.42±0.11	—	0.31±0.03	795.54±356.4	16.23±1.94	1.37±0.36	0.93+0.03
种间变异系数Interspecific variation coefficient（%）	81.26	63.65	31.63	24.08	30.57	40.31	19.07	—	22.1	56.95	28.21	64.48	16.46

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物种 Species	科 Family	物种代码 Code	叶片习性 Leaf habit	高度 Height/m	花果期 Flowering and fruiting period（month）
白刺Nitraria tangutorum	蒺藜科Zygophyllaceae	Nt	落叶Deciduous	1~2	5—8
四合木Tetraena mongolica	蒺藜科Zygophyllaceae	Tm	落叶Deciduous	0.4~0.9	5—9
霸王Zygophyllum xanthoxylum	蒺藜科Zygophyllaceae	Zx	落叶Deciduous	0.5~1	4—8
黄花红砂Reaumuria trigyna	柽柳科Tamaricaceae	Rt	落叶Deciduous	0.1~0.3	7—9
蒙古扁桃Prunus mongolica	蔷薇科Rosaceae	Prm	落叶Deciduous	1~2	5—10
绵刺Potaninia mongolica	蔷薇科Rosaceae	Pom	落叶Deciduous	0.3~0.4	6—10
锐枝木蓼Atraphaxis pungens	廖科Polygonaceae	Ap	落叶Deciduous	0.8~1.5	5—8
沙冬青Ammopiptanthus mongolicus	豆科Fabaceae	Am	常绿Evergreen	1.5~2	4—6
柠条锦鸡儿Caragana korshinskii	豆科Fabaceae	Ck	落叶Deciduous	1~4	5—7
毛刺锦鸡儿Caragana tibetica	豆科Fabaceae	Ct	落叶Deciduous	0.2~0.3	5—8
小叶锦鸡儿Caragana microphylla	豆科Fabaceae	Cm	落叶Deciduous	1~3	5—8
狭叶锦鸡儿Caragana stenophylla	豆科Fabaceae	Cs	落叶Deciduous	0.3~0.8	4—8
荒漠锦鸡儿Caragana roborovskyi	豆科Fabaceae	Cr	落叶Deciduous	0.3~1	5—7
半日花Helianthemum songaricum	半日花科Cistaceae	Hs	落叶Deciduous	0.10~0.12	5—9
驼绒藜Krascheninnikovia ceratoides	藜科Chenopodiaceae	Kc	落叶Deciduous	0.1~1	6—9
黑沙蒿Artemisia ordosica	菊科Asteraceae	Ao	落叶Deciduous	0.5~1	7—10
紫菀木Asterothamnus alyssoides	菊科Asteraceae	Aal	落叶Deciduous	0.1~0.2	7—10
蓍状亚菊Ajania achilleoides	菊科Asteraceae	Aac	落叶Deciduous	0.1~0.2	8—10

解剖特征 Anatomical traits	Blomberg's K	p	解剖特征 Anatomical traits	Blomberg's K	p
RPf	0.056	0.775	VWT	0.163	0.527
APf	0.055	0.991	T/D	0.154	0.533
TPf	0.176	0.445	VD	0.554	0.016
Ff	0.096	0.845	D_h	0.201	0.364
VLf	0.191	0.421	K_th	0.134	0.667
VWf	0.164	0.517	WD	0.138	0.643

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