水毛花种子萌发特性的差异及休眠解除方法

doi:10.17521/cjpe.2022.0523

摘要/Abstract

摘要：

水毛花(Schoenoplectiella triangulata)是一种广泛应用于湿地植被恢复中的挺水植物, 探究不同地区水毛花种群种子的萌发特性与休眠特性对其种质收集和科学利用至关重要。该研究测定了水毛花种子形态特性、萌发特性的种群间差异, 并对其休眠机制进行探索, 结果显示: (1)水毛花的种子存在生理休眠, 其形态、千粒质量和吸水特性在种群间存在显著差异。(2)光照和变温条件有助于水毛花种子的萌发。(3)低温沙藏、低温水藏和常温沙藏有助于水毛花种子休眠的解除; 氟啶酮和赤霉素处理均有助于打破水毛花种子的休眠, 但在不同地区水毛花种群间存在差异。(4)不同地区水毛花种群间种子形态特性、萌发特性以及休眠特性的差异与采集地的环境因子相关。该研究结果表明水毛花种子应用于湿地生态修复时应考虑种源的问题, 在播种前对其进行适当处理可以提高发芽率和生态修复成效。

关键词: 莎草科, 水毛花, 种子萌发, 种子休眠, 层积

Abstract:

Aims Schoenoplectiella triangulata is an emergent plant species widely used in wetland vegetation restoration. It is important to study the germination and dormancy characteristics of S. triangulata seeds from different populations for germplasm collection and scientific utilization.

Methods In this study, differences in the morphological characteristics and germination characteristics of S. triangulata seeds were investigated, and the mechanisms of seed dormancy were explored.

Important findings (1) The seeds of S. triangulata showed physiological dormancy (PD), and there were significant differences in morphology, PD, 1 000-grain mass and water absorption characteristics among populations. (2) Light and variable temperature were conducive to seed germination of S. triangulata.(3) Sand storage at low temperatures, water storage at low temperatures and sand storage at normal temperatures helped to break the dormancy of S. triangulata seeds. Both fluridone and gibberellin treatments also helped to break the dormancy of S. triangulata seeds, but these effects varied among different populations. (4) Among different populations of S. triangulata, seed morphological characteristics, germination characteristics and dormancy characteristics were correlated with environmental conditions of the collection site. These results indicate that provenance should be considered when S. triangulata seeds were applied in wetland ecological restoration and that proper seed treatments before sowing could improve seed germination and the effectiveness of ecological restoration.

Key words: Cyperaceae, Schoenoplectiella triangulata, seed germination, seed dormancy, stratification

袁涵, 钟爱文, 刘送平, 彭焱松, 徐磊. 水毛花种子萌发特性的差异及休眠解除方法. 植物生态学报, 2024, 48(5): 638-650. DOI: 10.17521/cjpe.2022.0523

YUAN Han, ZHONG Ai-Wen, LIU Song-Ping, PENG Yan-Song, XU Lei. Differences in the germination characteristics of Schoenoplectiella triangulata seeds and methods for breaking seed dormancy. Chinese Journal of Plant Ecology, 2024, 48(5): 638-650. DOI: 10.17521/cjpe.2022.0523

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2022.0523

https://www.plant-ecology.com/CN/Y2024/V48/I5/638

图/表 13

表1 水毛花种源地信息

Table 1 Provenance site information of Schoenoplectiella triangulata

原生地 Site of collection	种群编号 Population ID	位置 Location	经纬度 Latitude and longitude	年平均气温 Mean annual air temperature (℃)	年降水量 Annual precipitation (mm)
海南 Hainan	HN1	海口昌旺 Changwang, Haikou	19.80° N, 110.25° E	25.4	1 639
	HN2	海口羊山 Yangshan, Haikou	19.94° N, 110.31° E	25.4	1 639
贵州 Guizhou	GZ1	草海 Caohai	26.89° N, 104.23° E	10.5	950
	GZ2	草海 Caohai	26.99° N, 104.24° E	10.5	950
江西 Jiangxi	JX	上饶余干 Yugan, Shangrao	28.91° N, 116.49° E	19.0	1 700

图1 水毛花江西种群种子形态特征。

Fig. 1 Seed morphological characteristics of Schoenoplectiella triangulata in Jiangxi.

表2 水毛花种子形态和千粒质量的种群间差异 (平均值±标准误)

Table 2 Differences in the morphology and 1 000-grain mass of Schoenoplectiella triangulata seeds among populations (mean ± SE)

种群编号 Population ID	长 Length (mm)	宽 Width (mm)	高 Height (mm)	千粒质量 Thousand grain mass (g)
HN1	2.12 ± 0.12^a	1.57 ± 0.11^a	0.81 ± 0.08^b	1.24 ± 0.03^a
HN2	1.94 ± 0.11^b	1.47 ± 0.09^b	0.71 ± 0.07^d	0.85 ± 0.02^d
GZ1	1.89 ± 0.16^c	1.44 ± 0.09^c	0.75 ± 0.06^c	1.04 ± 0.06^bc
GZ2	1.86 ± 0.11^c	1.23 ± 0.14^d	0.63 ± 0.09^e	0.99 ± 0.03^c
JX	1.84 ± 0.24^c	1.48 ± 0.10^b	0.86 ± 0.09^a	1.04 ± 0.03^b

图2 不同地区种群水毛花种子吸水率和吸水速率变化(平均值±标准误)。不同小写字母表示差异显著(p < 0.05)。HN1, 海口昌旺种群; HN2, 海口羊山种群; GZ1、GZ2, 贵州草海不同种群; JX, 上饶余干种群。

Fig. 2 Water uptake percentage and water uptake rate of Schoenoplectiella triangulata seeds from different populations (mean ± SE). Different lowercase letters indicate significant differences (p < 0.05). HN1, Changwang population in Haikou; HN2, Yangshan population in Haikou; GZ1 and GZ2, different populations in Caohai, Guizhou; JX, Yugan population in Shangrao.

表3 不同温度及光照条件对水毛花种子萌发的影响 (平均值±标准误)

Table 3 Effects of different light and temperature conditions on the germination of Schoenoplectiella triangulate seeds (mean ± SE)

温度 Temperature (℃)	光照条件 Light condition	发芽率 Germination percentage (%)
温度 Temperature (℃)	光照条件 Light condition	HN1	HN2	JX	GZ1	GZ2
15/10	光照 Light	8.50 ± 1.50^Ba	9.50 ± 2.50^Ca	0 ± 0^Ba	0 ± 0^Ba	0 ± 0^Aa
15/10	黑暗 Dark	0 ± 0^Ab	0 ± 0^Ab	0 ± 0^Aa	0 ± 0^Aa	0 ± 0^Aa
20/15	光照 Light	32.50 ± 5.50^Aa	53.00 ± 8.50^Aa	6.00 ± 5.00^Aa	1.50 ± 0.50^Ba	0 ± 0^Aa
20/15	黑暗 Dark	0 ± 0^Ab	1.00 ± 0.50^Ab	0 ± 0^Ab	0 ± 0^Aa	0 ± 0^Aa
25/20	光照 Light	7.75 ± 3.25^Ba	22.00 ± 4.00^Ba	10.0 ± 8.00^Aa	9.00 ± 1.00^Aa	0 ± 0^Aa
25/20	黑暗 Dark	0 ± 0^Ab	0 ± 0^Ab	0.50 ± 0.25^Ab	0 ± 0^Ab	0 ± 0^Aa
30/25	光照 Light	3.50 ± 2.50^Ca	3.50 ± 4.50^Da	2.00 ± 1.50^Ba	7.50 ± 4.50^Aa	0 ± 0^Aa
30/25	黑暗 Dark	0 ± 0^Ab	0 ± 0^Aa	0.50 ± 0.50^Aa	0.50 ± 0.50^Ab	0 ± 0^Aa
35/30	光照 Light	0 ± 0^Ca	3.50 ± 2.50^Da	0.75 ± 0.50^Ba	1.00 ± 0.50^Ba	0.75 ± 0.50^Aa
35/30	黑暗 Dark	0 ± 0^Aa	0 ± 0^Aa	0 ± 0^Aa	0 ± 0^Aa	0 ± 0^Aa
15	光照 Light	0 ± 0^Ca	1.50 ± 1.25^Da	0 ± 0^Ba	0 ± 0^Ba	0 ± 0^Aa
15	黑暗 Dark	0 ± 0^Aa	0 ± 0^Aa	0 ± 0^Aa	0 ± 0^Aa	0 ± 0^Aa
20	光照 Light	1.75 ± 0.75^Ca	2.75 ± 1.25^Da	0.75 ± 0.50^Ba	0.50 ± 0.50^Ba	0 ± 0^Aa
20	黑暗 Dark	0 ± 0^Aa	0 ± 0^Aa	0 ± 0^Aa	0 ± 0^Aa	0 ± 0^Aa
25	光照 Light	2.50 ± 1.00^Ca	0.50 ± 1.00^Da	0.50 ± 1.00^Ba	0.25 ± 0.50^Ba	0 ± 0^Aa
25	黑暗 Dark	0 ± 0^Aa	0 ± 0^Aa	0 ± 0^Aa	0 ± 0^Aa	0 ± 0^Aa
30	光照 Light	0 ± 0^Ca	0 ± 0^Da	0 ± 0^Ba	0 ± 0^Ba	0.25 ± 0.50^Aa
30	黑暗 Dark	0 ± 0^Aa	0 ± 0^Aa	0 ± 0^Aa	0 ± 0^Aa	0 ± 0^Aa

图3 不同温度条件下水毛花种子萌发动态(平均值±标准误)。不同小写字母表示不同种群间差异显著(p < 0.05)。HN1, 海口昌旺种群; HN2, 海口羊山种群; GZ1、GZ2, 贵州草海不同种群; JX, 上饶余干种群。

Fig. 3 Germination dynamics of Schoenoplectiella triangulate seeds under different temperature conditions (mean ± SE). Different lowercase letters indicate significant differences among different populations (p < 0.05). HN1, Changwang population in Haikou; HN2, Yangshan population in Haikou; GZ1 and GZ2, different populations in Caohai, Guizhou; JX, Yugan population in Shangrao.

表4 基于广义线性分析的种群、存储方式、存储时间、存储温度及其交互作用对水毛花种子萌发的影响

Table 4 Effects of population, storage manner, storage time, storage temperature and their interaction on the germination of Schoenoplectiella triangulate seeds based on generalized linear analysis

因子 Factor	Wald’s χ²	df	p	因子 Factor	Wald’s χ²	df	p
种群 Population (P)	664.030	4	<0.001	M × Te	88.044	2	<0.001
存储方式 Storage manner (M)	190.189	2	<0.001	M × Ti	170.335	6	<0.001
存储温度 Storage temperature (Te)	142.543	1	<0.001	Ti × Te	58.457	3	<0.001
存储时间 Storage time (Ti)	515.916	3	<0.001	P × M × Te	107.331	8	<0.001
P × M	193.495	8	<0.001	P × M × Ti	165.427	24	<0.001
P × Te	139.538	4	<0.001	P × Ti × Te	109.271	12	<0.001
P × Ti	553.715	12	<0.001	M × Ti × Te	65.653	6	<0.001

图4 存储条件对水毛花种子的累积发芽率的影响。A, 海口昌旺种群。B, 海口羊山种群。C、D, 贵州草海不同种群。E, 上饶余干种群。

Fig. 4 Effects of storage conditions on the cumulative germination percentage of Schoenoplectiella triangulate seeds. A, Changwang population in Haikou. B, Yangshan population in Haikou. C, D, Different populations in Caohai, Guizhou. E, Yugan population in Shangrao.

图5 存储时间对水毛花种子发芽率的影响(平均值±标准误)。A, 冷沙。B, 冷水。C, 冷干。D, 常沙。E, 常水。F, 常干。不同小写字母表示不同存储时间间差异显著(p < 0.05)。HN1, 海口昌旺种群; HN2, 海口羊山种群; GZ1、GZ2, 贵州草海不同种群; JX, 上饶余干种群。

Fig. 5 Effects of storage time on the germination percentage of Schoenoplectiella triangulate seeds (mean ± SE). A, Low temperature plus sand reservoir. B, Low temperature plus water reservoir. C, Low temperature plus dry reservoir. D, Normal temperature plus sand reservoir. E, Normal temperature plus water reservoir. F, Normal temperature plus dry reservoir. Different lowercase letters indicate significant differences among different storage time (p < 0.05). HN1, Changwang population in Haikou; HN2, Yangshan population in Haikou; GZ1 and GZ2, different populations in Caohai, Guizhou; JX, Yugan population in Shangrao.

图6 氟啶酮(FL)对水毛花种子萌发的影响(平均值±标准误)。 A, 发芽率。B, 萌发指数。C, 萌发时滞。D, 幼苗形态, 从左到右依次为FL处理JX种子萌发后第2、4、5、6、7、9、11、14天幼苗形态。不同小写字母表示不同处理间差异显著(p < 0.05)。HN1, 海口昌旺种群; HN2, 海口羊山种群; GZ1、GZ2, 贵州草海不同种群; JX, 上饶余干种群。CK, 对照。

Fig. 6 Effects of fluridone (FL) on the germination of Schoenoplectiella triangulate seeds (mean ± SE). A, Germination percentage. B, Germination index. C, Germination time-lag. D, Seedling morphology (from left to right, on day 2, 4, 5, 6, 7, 9, 11 and 14 after JX seeds treated by FL germinated). Different lowercase letters indicate significant differences between different treatments (p < 0.05). HN1, Changwang population in Haikou; HN2, Yangshan population in Haikou; GZ1 and GZ2, different populations in Caohai, Guizhou; JX, Yugan population in Shangrao. CK, control.

图7 赤霉素(GA3)对水毛花种子萌发的影响(平均值±标准误)。A, 发芽率。B, 萌发指数。C, 萌发时滞。D, 幼苗形态, 从左到右依次为JX种子萌发后第2、4、6、8、10、12天幼苗形态。不同小写字母表示不同处理间差异显著(p < 0.05)。HN1, 海口昌旺种群; HN2, 海口羊山种群; GZ1、GZ2, 贵州草海不同种群; JX, 上饶余干种群。

Fig. 7 Effects of Gibberellins on the germination of Schoenoplectiella triangulate seeds (mean ± SE). A, Germination percentage. B, Germination index. C, Germination time-lag. D, Seedling morphology (from left to right, on day 2, 4, 6, 8, 10 and 12 after JX seeds treated by GA3 germinated). Different lowercase letters indicate significant differences among different treatments (p < 0.05). HN1, Changwang population in Haikou; HN2, Yangshan population in Haikou; GZ1 and GZ2, different populations in Caohai, Guizhou; JX, Yugan population in Shangrao.

图8 水毛花不同地区种群种子的聚类分析。Length、Width、Height、Mass、WA为种子的长、宽、高、千粒质量、吸水率; GRT1、GRT2、GRT3、GRT4、GRT5依次为5个变温下发芽率。HN1、HN2、GZ1、GZ2、JX同表1。

Fig. 8 Cluster analysis of Schoenoplectiella triangulata seeds from different populations. Length, Width, Height, Mass, and WA are the length, width, height, 1 000-grain mass, and water absorption rate of seeds, respectively; GRT1, GRT2, GRT3, GRT4 and GRT5 are the germination percentages under five variable temperatures, respectively. HN1, HN2, GZ1, GZ2 and JX were the same as those in Table 1.

图9 不同地区种群水毛花种子的形态特性、发芽率与环境因子的相关性分析。GRT1、GRT2、GRT3、GRT4、GRT5依次为5个变温下发芽率; Lat, 纬度; Length、Width、Height、Mass、WA依次为种子的长、宽、高、千粒质量、吸水率; Long, 经度; MAAT, 年平均气温; MAP, 年降水量。*, p < 0.05; **, p < 0.01。

Fig. 9 Correlation analyses among morphological characteristics, germination percentage and environmental factors of Schoenoplectiella triangulata seeds from different populations. GRT1, GRT2, GRT3, GRT4, GRT5, the germination percentages under five variable temperatures, respectively; Lat, latitude; Length, Width, Height, Mass, and WA are the length, width, height, 1 000-grain mass, and water absorption rate of seeds, respectively; Long, longitude; MAAT, mean annual air temperature; MAP, mean annual precipitation. *, p < 0.05; **, p < 0.01.

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