东北碱化草甸芦苇匍匐型分株超速生长过程及生理机制

doi:10.17521/cjpe.2023.0158

摘要/Abstract

摘要： 芦苇(Phragmites australis)是长根茎型克隆植物, 是世界广布种, 具有随着环境的变化改变其形态甚至生长型的可塑性与适应性。芦苇匍匐型分株是根茎伸出碱斑地面的一种特殊生长形式, 该研究旨在探讨匍匐型分株的生长规律及形成机制。采用挂标签定期对分株生长跟踪测量、不同叶龄叶片光合生理测定和¹⁵N同位素转移测定等方法, 测定并分析了芦苇匍匐型分株超速生长的节律与格局、光合特性及分株间生理整合指标。结果表明: 在碱斑极端生境, 芦苇匍匐型分株与对照(直立型分株)有着不同的生长节律和格局。经过120 d生长, 芦苇匍匐样本分株长度平均为(685.25 ± 118.75) cm, 在整个观测期间平均生长速率为(6.64 ± 3.51) cm·d^-1, 是对照分株的15.4倍, 呈先快速再减缓的对数异速生长过程; 而对照呈相对稳定的线性同速生长过程。芦苇匍匐型分株顶部幼叶具有与壮龄级功能叶相同的最大光合生产能力, 随着叶序增加, 其叶片的净光合速率呈逻辑斯蒂曲线变化, 对照则呈先增加再下降的二次曲线变化。匍匐型分株净光合速率的理论最大值也比对照高19.4%。经¹⁵N同位素处理的匍匐型分株各器官¹⁵N丰度均显著高于未处理的匍匐型分株。匍匐型分株是广布种芦苇在碱斑极端严酷生境形成的一种新的适应特征, 顶端幼叶的高光合速率以及丛生化基生分株与匍匐型分株之间的生理整合过程, 是匍匐型分株得以快速生长的物质基础。该研究拓展了对芦苇在极端生境适应的新认识, 提供了以物质生产和生理整合为佐证解析匍匐分株快速生长的研究方法, 具有重要的理论意义。

关键词: 克隆植物, 生理整合, ¹⁵N同位素, 分株生长型, 同化物转移, 极端生境, 植物生态学

Abstract:

Aims The reed (Phragmites australis) is a long-rhizomed clonal plant, which is distributed worldwide and has the plasticity and adaptability to change its morphology, even growth form with environmental change. The creeping ramets of reeds are a special growth form generated from their rhizomes extending out of alkaline soil patches. This study aims to explore the growth of creeping ramets and their underlying mechanisms.

Methods Using methods such as regular tracking of ramet growth by hanging tags, measurement of photosynthetic physiology of leaves of different ages, and determination of ¹⁵N isotope transfer, we measured and analyzed the growth rhythms and patterns of creeping reed ramets, their photosynthetic characteristics, and indicators of physiological integration between ramets.

Important findings We found that in highly alkaline areas, reeds’ creeping ramets displayed different growth patterns compared to control, upright ramets. After 120 days of growth, the creeping ramets had an average length of (685.25 ± 118.75) cm, and their average growth rate during the observation period was (6.64 ± 3.51) cm·d^-1. This was 15.4 times faster than the control, upright ramets, indicating a logarithmic allometry growth process that started quickly but then slowed down. In contrast, the control ramets showed a relatively stable linear isogonic growth process. Young leaves at the top of the creeping ramets had the same maximum photosynthetic capacity as mature functional leaves. The net photosynthetic rate of leaves on the creeping ramets varied in a logistic curve with increasing leaf order, while the control ramets varied in a quadratic curve that first increased and then decreased. Moreover, the theoretical maximum net photosynthetic rate of creeping ramets was 19.4% higher than that of the control ramets. Creeping ramets treated with ¹⁵N isotopes had significantly higher levels of ¹⁵N abundance in various organs compared to untreated creeping ramets. Creeping ramets are a new adaptive feature of this widespread plant in extremely harsh alkaline habitat patches. The superior growth of creeping ramets is attributed to the high photosynthetic rate of young apical leaves and the physiological integration between tufted basal ramets and creeping ramets. This study provides new insights into the adaptation of reeds to extreme habitats and offers a method for analyzing the superior growth of creeping ramets based on matter production and physiological integration, with important theoretical implications.

Key words: clonal plant, physiological integration, ¹⁵N isotope, ramet growth form, assimilate transfer, extreme habitat, plant ecology

韩大勇, 李海燕, 张维, 杨允菲. 东北碱化草甸芦苇匍匐型分株超速生长过程及生理机制. 植物生态学报, 2025, 49(2): 320-330. DOI: 10.17521/cjpe.2023.0158

HAN Da-Yong, LI Hai-Yan, ZHANG Wei, YANG Yun-Fei. Superior growth process of creeping ramets of Phragmites australis and its physiological mechanisms in an alkaline meadow in Northeast China. Chinese Journal of Plant Ecology, 2025, 49(2): 320-330. DOI: 10.17521/cjpe.2023.0158

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2023.0158

https://www.plant-ecology.com/CN/Y2025/V49/I2/320

图/表 9

图1 7月中旬羊草草甸围栏样地裸斑(A)和丛生的芦苇生长型组成(B)。CR, 匍匐型分株; LR, 伏生型分株; NR, 节生型分株; OR, 斜生型分株; UR, 直立型分株。

Fig. 1 Bare patches (A) and composition of the tufted Phragmites australis growth form (B) in fenced plots in a salinized Leymus chinensis meadow during mid-July. CR, creeping ramet; LR, lying ramet; NR, nodal ramet; OR, oblique ramet; UR, upright ramet.

表1 不同时间段芦苇种群匍匐型分株生长速率的基本统计参数(cm·d-1)

Table 1 Basic statistics on the growth rate of the creeping ramets of Phragmites australis at different time periods (cm·d-1)

时间段(月-日) Time period (month-day)	样本数 n	最大值 Max	最小值 Min	平均值 Mean	标准差 SD	变异系数 CV (%)
4-25-6-04	7	4.20	1.26	2.94^c	1.12	38.0
6-05-6-14	7	15.72	2.09	8.59^ab	4.72	54.9
6-15-6-24	7	15.08	7.92	11.46^a	2.67	23.3
6-25-7-04	7	16.22	8.21	10.74^a	2.78	25.9
7-05-7-14	7	12.95	5.27	8.51^ab	2.88	33.9
7-15-7-24	7	10.94	1.70	6.06^b	3.50	57.8
7-25-8-04	7	11.06	1.70	6.66^b	4.41	66.2
8-05-8-14	7	8.39	0.33	3.17^bc	2.75	86.6
8-15-8-24	7	6.71	0.04	1.64^c	2.29	139.8

图2 不同时间段芦苇种群匍匐型与直立型对照分株之间生长速率(平均值±标准差)差异的比较。*, 相同时间段不同分株之间的差异显著(p < 0.05); ns, 差异不显著(p > 0.05)。

Fig. 2 Comparison of differences in growth rate (mean ± SD) between the creeping ramets and the control upright ramets of Phragmites australis at different time periods. *, significant difference (p < 0.05) between different ramets at the same period; ns, no significant difference (p > 0.05).

表2 芦苇种群匍匐型和对照直立型分株长度(y, cm)与生长时间(x, d)关系的拟合方程参数及显著性检验

Table 2 Parameters and significance tests of the fitting relationships between the length (y, cm) of the creeping (CR) and control upright ramets (CK), and growth time (x, d) of Phragmites australis

分株 Ramet	样本号 No.	样本数 n	函数类型 Function type	方程参数 Equation parameters		R²	p
分株 Ramet	样本号 No.	样本数 n	函数类型 Function type	a	b	R²	p
匍匐型 CR	1	9	对数 Logarithmic	-1 322.8	378.34	0.92	<0.01
	2	9	对数 Logarithmic	-1 686.9	502.44	0.99	<0.01
	3	9	对数 Logarithmic	-1 727.7	522.14	0.93	<0.01
	4	9	对数 Logarithmic	-2 316.3	635.67	0.98	<0.01
	5	9	对数 Logarithmic	-1 682.3	514.24	0.95	<0.01
	6	9	线性 Linear	-238.1	8.33	0.99	<0.01
	7	9	对数 Logarithmic	-2 577.9	709.60	0.96	<0.01
	平均值 Mean	9	对数 Logarithmic	-1 946.3	556.36	0.99	<0.01
直立型对照 CK	1	9	线性 Linear	17.80	0.27	0.99	<0.01
	2	9	对数 Logarithmic	-57.88	23.10	0.98	<0.01
	3	9	线性 Linear	23.25	0.32	0.94	<0.01
	4	9	线性 Linear	13.50	0.44	0.99	<0.01
	5	9	线性 Linear	24.51	0.37	0.99	<0.01
	6	9	线性 Linear	17.38	0.45	0.99	<0.01
	7	9	对数 Logarithmic	-97.72	35.43	0.99	<0.01
	平均值 Mean	9	线性 Linear	0.38	19.18	0.97	<0.01

图3 芦苇种群匍匐型(A)和直立型对照(B)平均分株长度与生长时间的观测值和拟合曲线/直线(平均值±标准差)。

Fig. 3 Observed values and fitting curves on the relationship between the growth time and the average length of creeping ramets (A) and control upright ramets (B) of Phragmites australis (mean ± SD).

图4 芦苇匍匐型分株不同叶序光合速率(Pn)和蒸腾速率(Tr)差异的比较(平均值±标准差)。叶序间不同小写字母表示差异显著(p < 0.05)。

Fig. 4 Comparison of photosynthetic rate (Pn) and transpiration rate (Tr) (mean ± SD) of leaves at different orders of creeping Phragmites australis. Different lowercase letters indicate significant difference (p < 0.05) between leaf orders.

图5 芦苇种群匍匐型分株(A1、B1)、节生型分株(A2、B2)和直立型分株(对照A3、B3)的光合速率与蒸腾速率随着叶序变化的观测值及拟合曲线。

Fig. 5 Observed values and fitting curves of photosynthetic rate (Pn) and transpiration rate (Tr) of Phragmites australis leaves with varying leaf order on creeping ramets (A1, B1), nodal ramets (A2, B2) and control upright ramets (A3, B3).

表3 芦苇种群匍匐型分株(CR)、节生分株(NR)和直立型分株(对照, CK)的光合生理参数(y)与叶序(x)之间关系的拟合方程参数及显著性检验

Table 3 Parameters and significance tests of the relationships between the photosynthetic physiological indices (y) and leaf order (x) of Phragmites australis creeping ramets (CR), nodal ramets (NR) and control upright ramets (CK)

因变量 Dependent variable (y)	分株 Ramet	样本数 n	方程参数 Equation parameter				R²	p
因变量 Dependent variable (y)	分株 Ramet	样本数 n	A₁/a	A₂/b	x₀/c	m		p
光合速率 P_n (μmol·m^-2·s^-1)	匍匐型分株 CR	15	14.40	22.83	11.62	10.21	0.96	<0.01
	节生型分株 NR	8	2.89	4.81	-0.35		0.88	<0.01
	直立型分株 CK	8	-8.50	6.59	-0.40		0.94	<0.01
蒸腾速率 T_r (µmol·m^-2·s^-1)	匍匐型分株 CR	15	5.04	7.21	12.09	15.63	0.94	<0.01
	节生型分株 NR	8	1.33	2.18	-0.19		0.93	<0.01
	直立型分株 CK	8	-4.71	2.90	-0.19		0.92	<0.01

图6 芦苇匍匐型分株不同器官和部位的15N丰度在示踪元素处理与对照(A)及示踪元素处理的各器官(B)之间的差异比较(平均值±标准差)。L, 叶; LS, 叶鞘; S, 茎。1、2、3代表上中下部位。*表示示踪元素处理与对照差异显著(p < 0.05)。不同小写字母表示器官间差异显著(p < 0.05)。

Fig. 6 Comparison of 15N abundance between tracer-treated and control Phragmites australis creeping ramets in different organs and sections (A), and among tracer-treated organs (B) (mean ± SD). L, leaf; LS, leaf sheath; S, stem. 1, 2 and 3 represent upper, middle and lower sections. *, significant difference (p < 0.05) between tracer treated (15N) and control treatments. Different lowercase letters indicate significant difference (p < 0.05) among organs.

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