光温条件对不同种源红松种子萌发的影响

doi:10.17521/cjpe.2021.0474

摘要/Abstract

摘要：

种子萌发受种源和环境条件(如光照和温度条件等)的影响。种子萌发决定红松(Pinus koraiensis)的天然更新能力, 因此也是恢复地带性顶极群落阔叶红松林的关键。该研究以我国3个主要种源地(辽宁清原、吉林长白山和黑龙江伊春)当年的红松种子为对象, 监测自然光、温(不同季节、林窗和林下)和控制光、温(3个光照强度: 200、20和0 μmol·m^-2·s^-1, 记为L200、L20和L0; 2个温度: 25和15 ℃)条件下红松种子的萌发特征。结果表明, 自然越冬后, 春季(5月)所有种源种子均未萌发, 夏季(7月)和秋季(9月)各种源种子萌发率(GP)均较低(1.8%-33.7%), 但夏季林窗内各种源种子GP均显著高于林下。对林下各季节未萌发的种子, 带回室内给予适宜的光温条件(L200、25/15 ℃)后发现: 春季和夏季种子均能大量萌发(32%-77%), 而秋季种子GP极低(<2%)。25 ℃下, 3个种源地种子GP和萌发指数(GV)均呈现在L200条件下显著高于L20和L0处理。15 ℃条件下, 清原种子GP和GV对光照的响应与25 ℃条件下相一致, 长白山和伊春种子GP和GV分别在L0和L20处理下最高。所有光照强度下, 3个种源地种子GP和GV均为25 ℃处理显著高于15 ℃处理。可见, 适宜的温度是红松种子萌发的必要先决条件, 长白山和伊春红松种子可能需要更高的积温解除休眠; 适宜温度下(25 ℃), 红松种子表现出萌发需光性; 红松种子萌发对光、温的响应存在种源特异性, 长白山和伊春红松种子在相对低温下萌发需光性弱或消失。林下光、温条件不适宜红松种子萌发是造成红松林天然更新慢或更新差的重要原因。

关键词: 红松林, 天然林, 种子, 萌发特性, 更新抚育

Abstract:

Aims Seed germination is affected by both provenance and habitat conditions (light, temperature, etc.). Then natural regeneration capacity of Korean pine (Pinus koraiensis) is determined by its seed germination, which is the key to restore the broadleaved Korean pine forest—a regional climax community. This study aimed to reveal the effects of light and temperature on seed germination of Korean pine with different provenances.

Methods Germinations of the P. koraiensisseeds produced in the year were investigated in both field and lab from three provenances in China (i.e. Qingyuan, Liaoning Province; Changbai Mountain, Jilin Province; and Yichun, Heilongjiang Province). Specifically, for field survey, seed germination data were obtained from both forest canopy gap and forest understory in different seasons. While for lab experiment, the seed germination in growth chamber were recorded under three light intensities (i.e. 200, 20 and 0 μmol·m^-2·s^-1, L200, L20 and L0 hereafter) and two temperature conditions (i.e. 25 and 15 °C).

Important findings Results showed that no seeds germinated in spring (May) under natural conditions. In comparison, the seed germination percentages (GP) from three provenances were relatively low ranging from 1.8% in understory to 33.7% in canopy gaps in summer (July) and autumn (September). However, we found that the GP were significantly higher in the canopy gap than in the understory in summer. Besides, lab experiments showed that the GP were relatively high (ranged from 32% to 77%) for spring and summer seeds under suitable light and temperature conditions (L200, 25/15 °C) in growth chamber, but the GP were very low for autumn seeds (<2%). At 25 °C condition, the GP and germination values (GV) were significantly higher under L200 than those under L20 and L0 conditions for all the three provenances. At 15 °C condition, the response of GP and GV was consistent with that of 25 °C for seeds from Qingyuan, while the GP and GV were the highest under L0 and L20 conditions for seeds from Changbai Mountain and Yichun, respectively. Under all light intensities, GP and GV from three provenances were all higher under 25 °C than those under 15 °C. In conclusion, temperature is essential for seed germination of P. koraiensis, and the seeds from Changbai Mountain and Yichun might require higher accumulated temperature to break dormancy. Moreover, light helps promote seed germination at relatively high temperature condition (25 °C). However, the impacts of light and temperature were provenance-dependent. More specifically, the light impacts on germination were reduced at lower temperature conditions for seeds collected from Changbai Mountain and Yichun. Our study indicated that the poor regeneration of Korean pine natural forests might attribute to the unsuitable light and temperature conditions for seed germination.

Key words: Pinus koraiensis forest, natural forests, seed, germination characteristics, regeneration tending

张敏, 朱教君. 光温条件对不同种源红松种子萌发的影响. 植物生态学报, 2022, 46(6): 613-623. DOI: 10.17521/cjpe.2021.0474

ZHANG Min, ZHU Jiao-Jun. Effects of light and temperature on seed germination of Pinus koraiensis with different provenances. Chinese Journal of Plant Ecology, 2022, 46(6): 613-623. DOI: 10.17521/cjpe.2021.0474

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2021.0474

https://www.plant-ecology.com/CN/Y2022/V46/I6/613

图/表 8

图1 不同透光林分鱼眼镜头照片(A、B)、林冠开阔度(C)和土壤温度(D)的季节变化。

Fig. 1 Fisheye photographs (A, B), seasonal variations of canopy openness (C), and daily soil temperature (D) in forest stands with different transmittances.

表1 不同透光林分下红松种子萌发率的季节、种源和透光率三因素方差分析

Table 1 Three-way ANOVA of seasons, provenance, and light condition effects on germination percentage of Pinus koraiensis seeds under different canopy openness conditions

源 Source	III类平方和 Type III sum of squares	自由度 df	F	p
模型 Model	6 942.93^a	17	20.66	0.00
季节 Season	2 066.04	1	195.69	0.00
种源 Provenance	2 859.23	2	72.34	0.00
透光率 Light	106.96	1	5.41	0.03
季节×种源 Season × Provenance	1 475.41	4	18.66	0.00
季节×透光率 Season × Light	222.48	2	5.63	0.01
种源×透光率 Provenance × Light	98.84	2	2.50	0.10
种源×透光率×季节 Provenance × Light × Season	113.96	4	1.44	0.24

图2 三个种源红松种子在不同透光林分下不同季节的自然萌发率(平均值±标准误)。红松种子在春季均未萌发, 因此仅显示夏、秋季数据。不同大写字母表示同一林分不同季节间差异显著(p ≤ 0.05), 不同小写字母表示同一季节不同透光林分间差异显著(p ≤ 0.05)。

Fig. 2 Germination percentage of Pinus koraiensis seeds from three provenances under different light conditions between different seasons (mean ± SE). The seeds did not germinate in spring, thus only summer and autumn data were displayed. Different uppercase letters indicate significant differences between different seasons in the forest stands with the same light transmittance (p ≤ 0.05). Different lowercase letters indicate significant differences between forest stands with different light transmittances in the same season (p ≤ 0.05).

表2 不同透光林分下越冬储藏的3个种源地红松种子室内萌发率(%) (平均值±标准误)

Table 2 Germination percentages (%) of Pinus koraiensis seeds from three provenances cultivated in growth chamber after overwintering under different forest stands (mean ± SE)

种源地 Provenance	种子取出季节 Seed-collection season	林窗 Gap	林下 Understory
清原 Qingyuan	春 Spring	76.67 ± 1.45^a	61.00 ± 8.19^a
	夏 Summer	56.00 ± 5.29^a	58.33 ± 7.54^a
	秋 Autumn	2.00 ± 1.00^a	0.67 ± 0.33^a
长白山 Changbai Mountain	春 Spring	34.00 ± 8.50^a	32.00 ± 4.93^a
	夏 Summer	53.00 ± 1.15^a	45.00 ± 10.69^a
	秋 Autumn	1.67 ± 0.89^a	0.33 ± 0.33^a
伊春 Yichun	春 Spring	57.33 ± 5.90^a	56.00 ± 11.50^a
	夏 Summer	62.33 ± 5.04^a	50.00 ± 10.69^a
	秋 Autumn	0.67 ± 0.33^b	0^a

表3 不同光照、温度和种源地对红松种子萌发率(GP)、平均萌发时间(MGT)、萌发指数(GV)的三因素方差分析

Table 3 Three-way ANOVA for light, temperature, and provenance effects on the germination percentage (GP), mean germination time (MGT) and germination value (GV) of Pinus koraiensis seeds

指标 Index	源 Source	III类平方和 Type III sum of squares	自由度 df	F	p
GP	模型 Model	27 581.179^a	17	56.175	0.00
	种源 Provenance	6 191.542	2	107.189	0.00
	光 Light	2 987.890	2	51.727	0.00
	温度 Temp	12 454.163	1	431.217	0.00
	种源×光 Provenance × Light	1 289.040	4	11.158	0.00
	种源×温度 Provenance × Temperature	1 448.825	2	25.082	0.00
	光×温度 Light × Temperature	877.690	2	15.195	0.00
	种源×光×温度 Provenance × Light × Temperature	1 090.786	4	9.442	0.00
MGT	模型 Model	4 888.740^b	17	6.396	0.00
	种源 Provenance	686.604	2	7.636	0.00
	光 Light	542.808	2	6.037	0.00
	温度 Temp	1 355.676	1	30.154	0.00
	种源×光 Provenance × Light	178.422	4	0.992	0.42
	种源×温度 Provenance × Temperature	679.857	2	7.561	0.00
	光×温度 Light × Temperature	371.673	2	4.133	0.02
	种源×光×温度 Provenance × Light × Temperature	743.805	4	4.136	0.00
GV	模型 Model	57.775^c	17	47.783	0.00
	种源 Provenance	17.631	2	123.943	0.00
	光 Light	0.556	2	3.911	0.03
	温度 Temperature	28.159	1	395.914	0.00
	种源×光 Provenance × Light	1.801	4	6.331	0.00
	种源×温度 Provenance × Temperature	6.471	2	45.491	0.00
	光×温度 Light × Temperature	0.869	2	6.106	0.00
	种源×光×温度 Provenance × Light × Temperature	2.266	4	7.964	0.00

图3 光照培养箱不同光照和温度条件下红松种子的萌发率(平均值±标准误)。L0, L20, L200代表光照强度分别为0, 20, 200 μmol·m-2·s-1。不同大写字母表示相同光照下不同温度之间萌发率差异显著(p ≤ 0.05); 不同小写字母表示相同温度条件下不同光照之间萌发率差异显著(p ≤ 0.05)。

Fig. 3 Germination percentage (GP) of Pinus koraiensis seeds from three provenances under different light and temperature conditions in growth chamber (mean ± SE). L0, L2 and L200 represent 0, 20, 200 μmol·m-2·s-1 light intensities, respectively. Different uppercase letters indicate significant differences between different temperature conditions under the same light intensity (p ≤0.05); different lowercase letters indicate significant differences between different light intensities under the same temperature condition (p ≤ 0.05).

图4 不同光照和温度条件下红松的平均萌发时间(平均值±标准误)。L0、L20、L200代表光照强度分别为0、20、200 μmol·m-2·s-1。不同大写字母表示相同光照下不同温度之间萌发率差异显著(p ≤ 0.05); 不同小写字母表示相同温度条件下不同光照之间萌发率差异显著(p ≤ 0.05)。

Fig. 4 Mean germination time (MGT) of Pinus koraiensis seeds from three provenances under different light and temperature conditions (mean ± SE). L0, L2 and L200 represent 0, 20, 200 μmol·m-2·s-1 light intensities, respectively. Different uppercase letters indicate significant differences between different temperature conditions under the same light intensity (p ≤ 0.05); different lowercase letters indicate significant differences between different light intensities under the same temperature condition (p ≤ 0.05).

图5 不同光照和温度条件下红松种子萌发指数(平均值±标准误)。L0、L20、L200代表光照强度分别为0、20、200 μmol·m-2·s-1。不同大写字母表示相同光照下不同温度之间萌发率差异显著(p ≤ 0.05); 不同小写字母表示相同温度条件下不同光照之间萌发率差异显著(p ≤ 0.05)。

Fig. 5 Germination value (GV) of Pinus koraiensis seeds from three provenances under different light and temperature conditions (mean ± SE). L0, L2 and L200 represent 0, 20, 200 μmol·m-2·s-1 light intensities, respectively. Different uppercase letters indicate significant differences between different temperature conditions under the same light intensity (p ≤ 0.05); different lowercase letters indicate significant differences between different light intensities under the same temperature condition (p ≤ 0.05).

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