黑沙蒿应对降水变化的木质部与韧皮部协同响应机制

doi:10.17521/cjpe.2023.0103

植物生态学报 ›› 2024, Vol. 48 ›› Issue (7): 903-914.DOI: 10.17521/cjpe.2023.0103 cstr: 32100.14.cjpe.2023.0103

黑沙蒿应对降水变化的木质部与韧皮部协同响应机制

张富崇¹^,²^,⁴, 于明含¹^,³^,⁴^,^*(), 张建玲¹^,³^,⁴, 王平¹^,²^,⁴, 丁国栋¹^,³^,⁴, 何莹莹¹^,³^,⁴, 孙慧媛¹^,³^,⁴

¹北京林业大学水土保持学院, 北京 100083
²山西吉县森林生态系统国家野外科学观测研究站, 山西临汾 042200
³宁夏盐池毛乌素沙地生态系统国家定位观测研究站, 宁夏盐池 751500
⁴北京林业大学水土保持国家林业和草原局重点实验室, 北京 100083

收稿日期:2023-04-14 接受日期:2023-10-09 出版日期:2024-07-20 发布日期:2023-10-10
通讯作者: * 于明含(ymh_2012tai@163.com)
基金资助:
中央高校基本科研业务费专项资金(QNTD202303)

Synergistic response mechanisms in xylem and phloem of Artemisia ordosica to changes in precipitation

ZHANG Fu-Chong¹^,²^,⁴, YU Ming-Han¹^,³^,⁴^,^*(), ZHANG Jian-Ling¹^,³^,⁴, WANG Ping¹^,²^,⁴, DING Guo-Dong¹^,³^,⁴, HE Ying-Ying¹^,³^,⁴, SUN Hui-Yuan¹^,³^,⁴

¹School of Soil and Water Conservation, Beijing Forestry University, Beijing 100083, China
²Jixian National Forest Ecosystem Observation and Research Station, Linfen, Shanxi 042200, China
³Yanchi Ecology Research Station, Yanchi, Ningxia 751500, China
⁴Key Laboratory of State Forestry and Grassland Administration on Soil and Water Conservation, Beijing Forestry University, Beijing 100083, China

Received:2023-04-14 Accepted:2023-10-09 Online:2024-07-20 Published:2023-10-10
Supported by:
Fundamental Research Funds for the Central Universities(QNTD202303)

摘要/Abstract

摘要：

探究不同降水情景下荒漠植物茎干解剖学结构的适应性调节, 可以更好地理解未来降水格局变化下荒漠植物水和碳运输间的协调机制。该研究以毛乌素沙地黑沙蒿(Artemisia ordosica)种群为对象, 通过野外人工控制降水的方法, 模拟半干旱气候区降水变化趋势, 设置3个降水量水平(减水30%、自然降水、增水30%)以及2个降水间隔水平(降水间隔5 d、降水间隔15 d)开展双因素完全随机实验, 测定了黑沙蒿茎木质部与韧皮部解剖结构在不同降水情境下的轴向与径向变异。结果表明: 1)在降水改变的情况下, 黑沙蒿并未产生更抗栓塞的轴向木质部结构及传导效率更高的轴向韧皮部结构来适应环境; 2)降水变化通过改变40-60 cm土层含水率对黑沙蒿的木质部、韧皮部径向解剖性状产生影响。在低水分生境下, 黑沙蒿减小导管直径和增大导管壁厚度以保证水分运输的安全性, 并且通过增大韧皮部筛管面积来维持韧皮部导度保证碳的有效运输, 以此保证黑沙蒿进行正常的生理活动; 3)黑沙蒿木质部导管和韧皮部筛管具有等标度的轴向缩放规律, 二者协同关联共同维持水力功能, 且这种相关关系不受降水变化的影响。该研究表明, 黑沙蒿通过改变径向茎干结构而不是轴向结构来适应降水的改变。

关键词: 降水变化, 黑沙蒿, 木质部, 韧皮部, 异速生长

Abstract:

Aims Investigating the adaptive regulation of stem anatomy in desert plants under different precipitation scenarios will lead to a better understanding of the coordination mechanisms between water and carbon transport in desert plants under future precipitation patterns.
Methods In this study, a two-factor completely randomized experiment was conducted to determine the axial and radial variation in the xylem and phloem anatomy of Artemisia ordosica stems under different precipitation conditions by manipulating precipitation in the field in a semi-arid climate zone, with three precipitation treatments in amounts (30% precipitation reduction, natural precipitation and 30% precipitation increase) and two precipitation intervals (5 d precipitation interval and 15 d precipitation interval).
Important findings The results indicate: 1) Under altered precipitation, A. ordosica did not develop more conductive axial xylem structures and more conductive axial phloem structures to adapt to the changes; 2) Precipitation changes affected the radial anatomical traits of xylem and phloem of A. ordosica by altering the moisture content of the 40-60 cm soil layer. Under low moisture habitats, A. ordosica reduced the conduit diameter and increased the conduit wall thickness to ensure the safety of water transport, and maintained the phloem conductivity by increasing the lumen area of phloem sieve cells to ensure the effective transport of carbon, thus ensuring the normal physiological activities of A. ordosica; 3) The xylem conduit and phloem lumen of A. ordosica had an equal scaling axial scaling pattern, and the two were synergistically related to each other to maintain the hydraulic function, and this correlation was not affected by changes in precipitation. This study showed that A. ordosica adapted to changes in precipitation by altering the radial stem structure rather than the axial. This study is a valuable addition to the anatomical knowledge of the hydraulic structure of desert shrubs and provides a theoretical basis for future management of vegetation stability maintenance under changing precipitation patterns in semi-arid desert areas.

Key words: precipitation change, Artemisia ordosica, xylem, phloem, allometry

张富崇, 于明含, 张建玲, 王平, 丁国栋, 何莹莹, 孙慧媛. 黑沙蒿应对降水变化的木质部与韧皮部协同响应机制. 植物生态学报, 2024, 48(7): 903-914. DOI: 10.17521/cjpe.2023.0103

ZHANG Fu-Chong, YU Ming-Han, ZHANG Jian-Ling, WANG Ping, DING Guo-Dong, HE Ying-Ying, SUN Hui-Yuan. Synergistic response mechanisms in xylem and phloem of Artemisia ordosica to changes in precipitation. Chinese Journal of Plant Ecology, 2024, 48(7): 903-914. DOI: 10.17521/cjpe.2023.0103

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2023.0103

https://www.plant-ecology.com/CN/Y2024/V48/I7/903

图/表 10

表1 宁夏盐池实验样地降水量与降水间隔时间设置

Table 1 Experimental setting of the precipitation amounts and precipitation intervals in experimental plots in Yanchi, Ningxia

月份 Month	平均月降水量 Average monthly precipitation (mm)	降水间隔 Precipitation interval	平均单次降水量 Average precipitation per event (mm)			降水频次 Precipitation frequency
月份 Month	平均月降水量 Average monthly precipitation (mm)	降水间隔 Precipitation interval	W-	W	W+	降水频次 Precipitation frequency
5月 May	33.09	T	3.86	5.52	7.17	6
5月 May	33.09	T++	11.58	16.55	21.51	2
6月 June	41.08	T	4.79	6.85	8.90	6
6月 June	41.08	T++	14.38	20.54	26.70	2
7月 July	72.39	T	8.45	12.07	15.68	6
7月 July	72.39	T++	25.34	36.20	47.05	2
8月 August	63.51	T	7.41	10.59	13.76	6
8月 August	63.51	T++	22.23	31.76	41.28	2
9月 September	52.71	T	6.15	8.79	11.42	6
9月 September	52.71	T++	18.45	26.36	34.26	2

图1 宁夏盐池实验样地及植物样本示意图。A, 样地俯视图。B, 遮雨棚内黑沙蒿植物。C, 黑沙蒿茎轴向取样示意图。

Fig. 1 Schematic diagram of the experimental plots and plants in Yanchi, Ningxia. A, Overview of the plot. B, Artemisia ordosica plants in the rain shelter. C, Schematic diagram illustrating the axial sampling of Artemisia ordosica stems. DA, length of sampling point from stem tip.

表2 不同降水处理对宁夏盐池黑沙蒿群落不同土层土壤质量含水率(%)的影响(平均值±标准误)

Table 2 Effect of precipitation treatments on soil water content (%) in different soil layers of the Artemisia ordosica communities in Yanchi, Ningxia (mean ± SE)

降水处理Precipitation treatment	土壤层次 Soil layer (cm)
降水处理Precipitation treatment	0-10	10-20	20-30	30-40	40-50	50-60
W-T	1.61 ± 0.69^a	1.91 ± 0.34^a	2.07 ± 0.35^a	2.68 ± 0.11^a	2.87 ± 0.18^b	2.69 ± 0.27^b
WT	2.22 ± 1.04^a	2.55 ± 0.66^a	2.87 ± 0.74^a	3.17 ± 0.10^a	3.39 ± 0.28^b	3.18 ± 0.57^b
W+T	2.12 ± 0.83^a	2.51 ± 0.56^a	2.52 ± 0.40^a	3.07 ± 0.51^a	3.87 ± 0.38^ab	3.60 ± 0.44^b
W-T++	1.36 ± 0.20^a	1.84 ± 0.46^a	2.15 ± 0.29^a	2.85 ± 0.10^a	3.60 ± 0.65^b	4.40 ± 0.68^ab
WT++	1.46 ± 0.17^a	2.33 ± 0.55^a	2.58 ± 0.54^a	3.89 ± 0.33^a	4.83 ± 0.36^ab	4.44 ± 0.76^ab
W+T++	1.52 ± 0.26^a	2.88 ± 0.96^a	3.92 ± 1.62^a	4.88 ± 1.92^a	6.33 ± 1.41^a	5.81 ± 0.53^a
双因素方差分析结果(F值) Results of Two-Way ANOVA (F-values)
W	0.394	1.824	1.917	2.213	7.434^**	4.612^*
T	2.160	0.006	0.783	3.566	15.373^***	27.910^***
W × T	0.173	0.250	1.243	1.017	1.607	0.709

图2 不同降水处理下黑沙蒿茎各个解剖性状随距离茎尖长度(DA)的轴向变化。A, 导管直径(Dc)的轴向变化。B, 水力直径(Dh)的轴向变化。C, 导管壁厚度(Tc)的轴向变化。D, 韧皮部筛管面积(PA)的轴向变化。T, 降水间隔5天; T++, 降水间隔15天; W-, 减水30%; W, 自然降水量; W+, 增水30%。CS, 共同斜率。

Fig. 2 Variations in the anatomical characteristics of stems of Artemisia ordosica under precipitation treatments with respect to the axial changes in stem tip length (DA). A, Axial changes in conduit diameter (Dc). B, Axial changes in hydraulically weighted diameter of xylem conduits (Dh). C, Axial changes in conduit wall thickness (Tc). D, Axial changes in lumen area of phloem sieve cells (PA). T, precipitation interval 5 days; T++, precipitation interval 15 days; W-, precipitation reduce by 30%; W, natural precipitation; W+, precipitation increase by 30%. CS, common slope.

表3 不同降水处理下黑沙蒿各木质部与韧皮部解剖特征与距离茎尖长度的幂函数模型输出结果

Table 3 Power function model outputs for each xylem and phloem anatomical feature and distance from stem tip length for different precipitation treatments of Artemisia ordosica

模型 Model	处理 Treatment	斜率(下限-上限) Slope (lower limit-upper limit)	截距(下限-上限) Intercept (lower limit-upper limit)
lg DA VS lg D_c	W-T	0.178 (0.135-0.234)^a	1.059 (1.004-1.115)^B
	WT	0.154 (0.120-0.197)^a	1.105 (1.058-1.152)^B
	W+T	0.154 (0.122-0.193)^a	1.147 (1.106-1.188)^A
	W-T++	0.167 (0.119-0.236)^a	1.083 (1.025-1.141)^B
	WT++	0.199 (0.137-0.288)^a	1.077 (0.999-1.156)^B
	W+T++	0.182 (0.140-0.236)^a	1.073 (1.016-1.129)^B
lg DA VS lg D_h	W-T	0.162 (0.117-0.226)^a	1.156 (1.094-1.217)^B
	WT	0.154 (0.110-0.218)^a	1.165 (1.099-1.230)^B
	W+T	0.179 (0.146-0.219)^a	1.179 (1.137-1.222)^B
	W-T++	0.202 (0.135-0.302)^a	1.134 (1.105-1.217)^B
	WT++	0.213 (0.129-0.351)^a	1.137 (1.021-0.254)^B
	W+T++	0.183 (0.140-0.238)^a	1.137 (1.080-1.195)^B
lg DA VS lg T_c	W-T	0.253 (0.173-0.371)^a	-0.092 (-0.104-0.020)^A
	WT	0.215 (0.143-0.323)^a	-0.112 (-0.222- -0.003)^A
	W+T	0.249 (0.173-0.360)^a	-0.221 (-0.330- -0.113)^B
	W-T++	0.237 (0.167-0.336)^a	-0.066 (-0.151-0.020)^A
	WT++	0.202 (0.135-0.303)^a	-0.097 (-0.185- -0.008)^A
	W+T++	0.235 (0.163-0.338)^a	-0.156 (-0.259- -0.054)^A
lg DA VS lg PA	W-T	0.277 (0.177-0.435)^a	1.043 (0.888-1.198)^A
	WT	0.243 (0.180-0.328)^a	0.948 (0.856-1.041)^B
	W+T	0.231 (0.181-0.294)^a	0.967 (0.898-1.036)^B
	W-T++	0.236 (0.188-0.295)^a	1.064 (1.006-1.123)^A
	WT++	0.236 (0.177-0.315)^a	0.995 (0.917-1.073)^AB
	W+T++	0.243 (0.190-0.311)^a	0.943 (0.871-1.014)^B

图3 不同降水处理下黑沙蒿茎同一轴向位置各个解剖性状的差异。T, 降水间隔5天; T++, 降水间隔15天; W-, 减水30%; W, 自然降水; W+, 增水30%。Dc, 导管直径; Dh, 水力直径; Tc, 导管壁厚度; PA, 韧皮部筛管面积。不同大写字母表示在同一降水间隔期下不同降水量具有显著差异(p < 0.05); 不同小写字母表示同一降水量下不同降水间隔期具有显著差异(p < 0.05), 最小显著差异(LSD)事后检验在α = 0.05水平下进行的。

Fig. 3 Differences in individual anatomical traits at the same axial position of the Artemisia ordosica stem under precipitation treatments. T, precipitation interval 5 days; T++, precipitation interval 15 days; W-, precipitation reduce by 30%; W, natural precipitation; W+, precipitation increase by 30%. Dc, conduit diameter; Dh, hydraulically weighted diameter of xylem conduits; Tc, conduit wall thickness; PA, lumen area of phloem sieve cells. Different uppercase letters indicate significant differences (p < 0.05) between different amount of precipitation at the same precipitation interval; different lowercase letters indicate significant differences (p < 0.05) among different precipitation intervals at the same amount of precipitation, least significant difference (LSD) post hoc test at α = 0.05 level.

表4 同一轴向位置黑沙蒿茎解剖特征的双因素方差分析结果(F值)

Table 4 Results of two-way ANOVA (F-values) for anatomical characteristics of Artemisia ordosica stems in the same axial position

降水处理 Precipitation treatment	解剖特征 Anatomical characteristics
降水处理 Precipitation treatment	D_c	D_h	T_c	PA
W	24.932^***	2.968	29.155^***	70.233^***
T	5.079^*	2.527	0.203	1.816
W × T	0.347	0.645	0.104	0.775

图4 黑沙蒿茎同一轴向位置导管直径(Dc)、水力直径(Dh)、导管壁厚度(Tc)与韧皮部筛管面积(PA)的相关关系。***, p < 0.001。

Fig. 4 Correlation between conduit diameter (Dc), hydraulically weighted diameter of xylem conduits (Dh), conduit wall thickness (Tc) and lumen area of phloem sieve cells (PA) in the same axial position of the stem of Artemisia ordosica. ***, p < 0.001.

表5 不同降水处理下黑沙蒿茎同一轴向位置木质部解剖特征与韧皮部筛管面积线性拟合模型斜率的差异性(平均值±标准误)

Table 5 Differences in the slope of the linear fit model between xylem anatomical features and sieve tube area of the bast at the same axial position of Artemisia ordosica stems under different precipitation treatments (mean ± SE)

降水处理 Precipitation treatment	解剖特征 Anatomical characteristics
降水处理 Precipitation treatment	D_c	D_h	T_c
W-T	-1.89 ± 0.68^a	-0.71 ± 0.39^a	8.85 ± 2.62^a
WT	-0.68 ± 0.88^a	-0.19 ± 0.45^a	7.74 ± 4.14^a
W+T	-0.76 ± 0.74^a	-0.50 ± 0.45^a	5.99 ± 4.05^a
W-T++	-0.62 ± 0.79^a	-0.47 ± 0.43^a	9.92 ± 3.93^a
WT++	-0.85 ± 0.72^a	-0.46 ± 0.42^a	13.90 ± 6.17^a
W+T++	-0.69 ± 0.75^a	-0.47 ± 0.49^a	4.34 ± 3.61^a

图5 不同土层含水率与黑沙蒿各个解剖特征相关关系。Dc, 导管直径; Dh, 水力直径; Tc, 导管壁厚度; PA, 韧皮部筛管面积。

Fig. 5 Correlation between water content of different soil layers and individual anatomical features of Artemisia ordosica. Dc, conduit diameter; Dh, hydraulically weighted diameter of xylem conduits; Tc, conduit wall thickness; PA, lumen area of phloem sieve cells.

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黑沙蒿应对降水变化的木质部与韧皮部协同响应机制

Synergistic response mechanisms in xylem and phloem of Artemisia ordosica to changes in precipitation

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