Effects of fire disturbance on seed germination of Larix gmelinii

doi:10.17521/cjpe.2023.0212

Abstract

Abstract:

Aims Forest fire is an important driver of boreal forest regeneration, and commonly generates a series of products, which play an important role in promoting or inhibiting seed germination. While seed germination in soil seed banks following forest fire is the main pathway for renewal and recovery of seed plants, relatively limited research has been conducted in China on post-fire renewal and recovery of trees compared to herbaceous and shrub plants. In this paper, the seeds of Larix gmelinii on the Da Hinggan Mountains area were selected to investigate their germination response to fire disturbance, which could provide theoretical support for the rapid renewal and recovery of L. gmelinii after fire.

Methods The individual and joint effects of heat shock, fire ash, and light on the germination rate and germination potential of L. gmelinii seeds were examined with analysis of variance.

Important findings The germination rate of L. gmelinii seeds was highest under dry thermal stimulation at 90 °C for 5 min (61.11%), 21.11% higher than the control, and the seeds were inactivated under the wet thermal stimulation at 90 °C for 5 min and above. The quality of fire ash was positively correlated with the germination rate of L. gmelinii seeds, and the germination rate of L. gmelinii seeds under the addition of 1 g of fire ash was 71.1%, 30% higher than the control; the quality of light was negatively correlated with the germination rate of L. gmelinii seeds, and the germination rate of the seeds under 100% shading increased by 14.00%, while this rate under 0 shading decreased by 16.67% compared to the control. In terms of the interactions among heat stimulation, fire ash and light, the highest seed germination rate (57%) of L. gmelinii seeds (17% higher than the control) appeared under the 1 g fire ash + 100% shading. The contributions of multiple factors to seed germination followed the order, fire ash > dry heat shock > light > wet heat shock, each explaining 33.0%, 31.8%, 20.3%, and 14.9% of the variance. Ash was the most dominant abiotic driver, and factor importance was as follows: fire ash (1.00 g) > dry heat shock (60 °C 3 min) > light (0 shading) > wet heat shock (90 °C 3 min) > wet heat shock (60 °C 3 min) > light (100% shading) > fire ash (0.25 g) > dry heat shock (120 °C 3 min) at the 95% confidence interval.

Key words: fire disturbance, seed germination, Larix gmelinii, seed

SUN Long, LI Wen-Bo, LOU Hu, YU Cheng, HAN Yu, HU Tong-Xin. Effects of fire disturbance on seed germination of Larix gmelinii[J]. Chin J Plant Ecol, 2024, 48(6): 770-779.

Add to citation manager EndNote|Ris|BibTeX

URL: https://www.plant-ecology.com/EN/10.17521/cjpe.2023.0212

https://www.plant-ecology.com/EN/Y2024/V48/I6/770

Figures/Tables 6

Table 1 Experimental treatments exposing to seed germination rate of Larix gmelinii

处理 Treatment	处理 Treatment	处理 Treatment
CK: 对照 Control	H₁₁: 105 ℃ 5 min	S₇: 90 ℃ 3 min
H₁: 60 ℃ 3 min	H₁₂: 105 ℃ 10 min	S₈: 90 ℃ 5 min
H₂: 60 ℃ 5 min	H₁₃: 120 ℃ 3 min	S₉: 90 ℃ 10 min
H₃: 60 ℃ 10 min	H₁₄: 120 ℃ 5 min	A₁: 0.25 g
H₄: 75 ℃ 3 min	H₁₅: 120 ℃ 10 min	A₂: 0.50 g
H₅: 75 ℃ 5 min	S₁: 60 ℃ 3 min	A₃: 1.00 g
H₆: 5 ℃ 10 min	S₂: 60 ℃ 5 min	L₁: 0
H₇: 90 ℃ 3 min	S₃: 60 ℃ 10 min	L₂: 25%
H₈: 90 ℃ 5 min	S₄: 75 ℃ 3 min	L₃: 50%
H₉: 90 ℃ 10 min	S₅: 75 ℃ 5 min	L₄: 75%
H₁₀: 105 ℃ 3 min	S₆: 75 ℃ 10 min	L₅: 100%

Fig. 1 Germination rate of Larix gmelinii seeds under different treatments (mean ± SD). A, Effect of dry heat shock on seed germination of L. gmelinii. B, Effect of wet heat shock on seed germination of L. gmelinii. C, Effect of fire ash on seed germination of L. gmelinii. D, Effect of light on seed germination of L. gmelinii. Information on each treatment is given in Table 1.

Table 2 Bi-factorial analysis of seed germination rate of Larix gmelinii

处理 Treatment	变异来源 Source of variation	偏平方和 SS	自由度 df	均方 Mf	F
干热激处理 Dry heat shock	温度 Temperature	0.77	4	0.19	402.31^**
	时间 Time	0.17	2	0.09	179.00^**
	温度×时间 Temperature × time	0.04	8	0.01	10.48^**
湿热激处理 Wet heat shock	温度 Temperature	0.71	2	0.36	788.62^**
	时间 Time	0.16	2	0.08	182.71^**
	温度×时间 Temperature × time	0.06	4	0.02	35.51^**

Fig. 2 Germination rate of Larix gmelinii seeds under different treatments (mean ± SD). A, Germination rate of L. gmelinii seeds under the interaction of dry heat shock, fire ash and light. B, Germination rate of L. gmelinii seeds under the interaction of wet heat shock, fire ash and light. ***, significant difference with CK (p < 0.001). Information on each treatment is given in Table 1.

Fig. 3 Relative effects of abiotic factors on seed germination rate of Larix gmelinii. The location of a point indicates relative importance of each factor for the associated 95% confidence interval; the height of a column denotes percentage of variance explained. **, p < 0.01; ***, p < 0.001. Adj.R2, adjusted R2。Information on each treatment is given in Table 1.

Fig. 4 Germination potential of Larix gmelinii seeds under different treatments. Red dashed lines indicate time point of the highest seed vigor; shaded area indicate time period when seeds reached the maximum germination potential under the 12 treatments. Information on each treatment is given in Table 1.

References 56

[1]	Abella SR (2006). Effects of smoke and fire-related cues on Penstemon barbatus seeds. The American Midland Naturalist, 155, 404-410.
[2]	Arán D, García-Duro J, Reyes O, Casal M (2013). Fire and invasive species: modifications in the germination potential of Acacia melanoxylon, Conyza canadensis and Eucalyptus globulus. Forest Ecology and Management, 302, 7-13.
[3]	Ba LN, Liu Q, Zhang YH, Wang SZ, Gu HY (2018). Effect of smoke, heat shock and ash on seed germination of Populus ussuriensis. Journal of Northeast Forestry University, 46(6), 27-32.
	[巴丽娜, 刘强, 张芸慧, 王顺忠, 谷会岩 (2018). 烟熏、热激、火烧灰对大青杨种子萌发的影响. 东北林业大学学报, 46(6), 27-32.]
[4]	Baskin JM, Baskin CC, Li XJ (2000). Taxonomy, anatomy and evolution of physical dormancy in seeds. Plant Species Biology, 15, 139-152.
[5]	Berkovich YA, Konovalova IO, Smolyanina SO, Erokhin AN, Avercheva OV, Bassarskaya EM, Kochetova GV, Zhigalova TV, Yakovleva OS, Tarakanov IG (2017). LED crop illumination inside space greenhouses. Reach Reviews in Human Space Exploration, 6, 11-24.
[6]	Bu XL, Xue JH, Wu YB, Ma WB (2020). Effect of biochar on seed germination and seedling growth of Robinia pseudoacacia L. in karst calcareous soils. Communications in Soil Science and Plant Analysis, 51, 352-363.
[7]	Carrington ME (2010). Effects of soil temperature during fire on seed survival in Florida sand pine scrub. International Journal of Forestry Research, 2010, 402346. DOI: 10.1155/2010/402346.
[8]	Castoldi E, Molina JA (2014). Effect of seed mass and number of cotyledons on seed germination after heat treatment in Pinus sylvestris L. var. iberica Svob. Forest Systems, 23, 483-489.
[9]	Cavallero L, Raffaele E (2010). Fire enhances the ‘competition- free’ space of an invader shrub: Rosa rubiginosa in northwestern Patagonia. Biological Invasions, 12, 3395-3404.
[10]	Cox RD, Chou YF, Wester DB (2017). Smoke water and heat influence emergence of shortgrass prairie species. Fire Ecology, 13, 138-148.
[11]	Chen F, Meng YJ, Shuai HW, Luo XF, Zhou WG, Liu JW, Yang WY, Shu Y (2017). Effect of plant allelochemicals on seed germination and its ecological significance. Chinese Journal of Ecological Agriculture, 25, 36-46.
	[陈锋, 孟永杰, 帅海威, 罗晓峰, 周文冠, 刘建伟, 杨文钰, 舒凯 (2017). 植物化感物质对种子萌发的影响及其生态学意义. 中国生态农业学报, 25, 36-46.]
[12]	Farooq MA, Zhang X, Zafar MM, Ma W, Zhao J (2021). Roles of reactive oxygen species and mitochondria in seed germination. Frontiers in Plant Science, 12, 781734. DOI: 10.3389/fpls.2021.781734.
[13]	Fernandes AF, Oki Y, Fernandes GW, Moreira B (2021). The effect of fire on seed germination of campo rupestre species in the South American Cerrado. Plant Ecology, 222, 45-55.
[14]	Fu JJ, Wu ZW, Yan SJ, Zhang YJ, Gu XL, Du LH (2020). Effects of climate, vegetation, and topography on spatial patterns of burn severity in the Great Xing’an Mountains. Acta Ecologica Sinica, 40, 1672-1682.
	[付婧婧, 吴志伟, 闫赛佳, 张宇婧, 顾先丽, 杜林翰 (2020). 气候、植被和地形对大兴安岭林火烈度空间格局的影响. 生态学报, 40, 1672-1682.]
[15]	Gonzalez S, Salazar CV, Ghermandi L (2022). Burning the seed bank: seed response of eight species to different fire intensities in Patagonian steppe. Applied Vegetation Science, 25, e12684. DOI: 10.1111/avsc.12684.
[16]	Gonzalez SL, Ghermandi L (2012). Fire cue effects on seed germination of six species of northwestern Patagonian grasslands. Natural Hazards and Earth System Sciences, 12, 2753-2758.
[17]	Gross N, Le Bagousse-Pinguet Y, Liancourt P, Berdugo M, Gotelli NJ, Maestre FT (2017). Functional trait diversity maximizes ecosystem multifunctionality. Nature Ecology & Evolution, 1, 132. DOI: 10.1038/s41559-017-0132.
[18]	Gu HY, Jiang KY, Zhang YH, Wang SZ, Chen XW (2017). Effect of heat shock on seed germination of three Pinaceae species in Great Hing’an Mountains. Acta Ecologica Sinica, 37, 6581-6587.
	[谷会岩, 蒋克研, 张芸慧, 王顺忠, 陈祥伟 (2017). 热激对大兴安岭三种松科树种种子萌发的影响. 生态学报, 37, 6581-6587.]
[19]	Herrero C, Kassa A, Pando V, Bravo F, Alía R (2019). Effect of heat shock on the germination of seeds of the species Acacia senegal L. and Acacia seyal Del. from sub-Saharan Africa (Ethiopia). Forest Systems, 28, e006. DOI: 10.5424/fs/2019282-14227.
[20]	Hodges JA, Price JN, Nimmo DG, Guja LK (2019). Evidence for direct effects of fire-cues on germination of some perennial forbs common in grassy ecosystems. Austral Ecology, 44, 1271-1284.
[21]	Hu HQ, Wei SJ, Sun L (2012). Estimating carbon emissions from forest fires during 2001 to 2010 in Daxing’anling Mountain. Acta Ecologica Sinica, 32, 5373-5386.
	[胡海清, 魏书精, 孙龙 (2012). 大兴安岭2001-2010年森林火灾碳排放的计量估算. 生态学报, 32, 5373-5386.]
[22]	Jeffery DJ, Holmes PM, Rebelo AG (1988). Effects of dry heat on seed germination in selected indigenous and alien legume species in South Africa. South African Journal of Botany, 54, 28-34.
[23]	Jiang KY, Zhang YH, Liu Q, Wang SZ, Gu HY (2018). Effect of heat shock on seed germination of three Betulaceae species in Daxing’an Mountains. Journal of Northeast Forestry University, 46(6), 23-26.
	[蒋克研, 张芸慧, 刘强, 王顺忠, 谷会岩 (2018). 热激对大兴安岭桦木科3个树种种子萌发的影响. 东北林业大学学报, 46(6), 23-26.]
[24]	Kin AG, Suárez CE, Chirino CC, Ávila PL, Morici EFA (2016). Impact of heat on seed germination of three perennial grasses in the semiarid region in Central Argentina. Australian Journal of Botany, 64, 451-455.
[25]	King RA, Menges EA (2018). Effects of heat and smoke on the germination of six Florida scrub species. South African Journal of Botany, 115, 223-230.
[26]	Kyereh B, Swaine MD, Thompson J (1999). Effect of light on the germination of forest trees in Ghana. Journal of Ecology, 87, 772-783.
[27]	Li MY (2020). The Effect of Forest Fire Combustion Products on Seed Germination of Ten Herbaceous Plants in Great Hing’an Mountain. Master degree dissertation, Northeast Forestry University, Harbin.
	[李明月 (2020). 林火产物对大兴安岭十种草本植物种子萌发的影响. 硕士学位论文, 东北林业大学, 哈尔滨.]
[28]	Li SY (2023). Effects of Fire Cues on Seed Germination of Typical Plants in the Songnen Grassland Under Salinity Stress and Its Mechanism. Master degree dissertation, University of Chinese Academy of Sciences, Beijing.
	[李绍阳 (2023). 火烧信号对盐碱胁迫下松嫩草地典型植物种子萌发的影响及机制研究. 硕士学位论文, 中国科学院大学, 北京.]
[29]	Li SY, Ma HY, Zhao DD, Ma MY, Qi WW (2021). Research progress on the effects of fire-related cues on seed germination. Chinese Journal of Plant Ecology, 45, 1177-1190.
	[李绍阳, 马红媛, 赵丹丹, 马梦谣, 亓雯雯 (2021). 火烧信号对种子萌发影响的研究进展. 植物生态学报, 45, 1177-1190.] DOI
[30]	Liang RF, Wu ZN, Li ZY, Kong LQ (2021). Effects of low-temperature and light on seed germination of Leymus chinensis. Chinese Journal of Grassland, 43(12), 33-39.
	[梁润芳, 武自念, 李志勇, 孔令琪 (2021). 低温和光照对羊草种子萌发的影响. 中国草地学报, 43(12), 33-39.]
[31]	Liu FL (2017). Response of four pine species to plant ash during germination and seedling growth. Acta Ecologica Sinica, 37, 5673-5680.
	[刘发林 (2017). 草木灰对四种松属种子发芽和幼苗生长的影响. 生态学报, 37, 5673-5680.]
[32]	Luo BZ, Wei SJ, Luo SS, Hu HQ, Wang ZS (2022). Effects of forest fire disturbance on carbon pools of Schima superba forest ecosystem in Guangdong Province. Acta Ecologica Sinica, 43, 3359-3369.
	[罗碧珍, 魏书精, 罗斯生, 胡海清, 王振师 (2022). 林火干扰对广东木荷林生态系统碳库的影响. 生态学报, 43, 3359-3369.]
[33]	McLauchlan KK, Higuera PE, Miesel J, Rogers BM, Schweitzer J, Shuman JK, Tepley AJ, Varner JM, Veblen TT, Adalsteinsson SA, Balch JK, Baker P, Batllori E, Bigio E, Brando P, et al. (2020). Fire as a fundamental ecological process: research advances and frontiers. Journal of Ecology, 108, 2047-2069.
[34]	Naghipour AA, Bashari H, Khajeddin SJ, Tahmasebi P, Iravani M (2016). Effects of smoke, ash and heat shock on seed germination of seven species from Central Zagros rangelands in the semi-arid region of Iran. African Journal of Range & Forage Science, 33, 67-71.
[35]	Perala DA, Alm AA (1990). Reproductive ecology of birch: a review. Forest Ecology and Management, 32, 1-38.
[36]	Qi YC, Dong Z, Li WS, Xue PP, He YN, Qi DH (2023). Effects of different light conditions on seed germination and seedling growth of Toxicodendron vernicifluum. Agriculture and Technology, 43(11), 76-81.
	[綦远才, 董智, 李文杉, 薛沛沛, 何雅楠, 齐代华 (2023). 不同光照条件对漆树种子萌发及幼苗生长的影响. 农业与技术, 43(11), 76-81.]
[37]	Reyes O, Casal M (2004). Effects of forest fire ash on germination and early growth of four Pinus species. Plant Ecology, 175, 81-89.
[38]	Ribeiro S, Gaspar MJ, Lima-Brito J, Fonseca T, Soares P, Cerveira A, Fernandes PM, Louzada J, Carvalho A (2022). Impact of fire recurrence and induced water stress on seed germination and root mitotic cell cycle of Pinus pinaster Aiton. Forests, 14, 78. DOI: 10.3390/f14010078.
[39]	Rayana DSM, Anderson CJ, José MRF, Oliveira LCD (2016). Proteomic analysis of osmoprimed and heat-shock-treated eucalyptus urophylla seeds. Trees, 1-12.
[40]	Santana VM, Bradstock RA, Ooi MKJ, Denham AJ, Auld TD, Baeza MJ (2010). Effects of soil temperature regimes after fire on seed dormancy and germination in six Australian Fabaceae species. Australian Journal of Botany, 58, 539-545.
[41]	Soto B, Basanta R, Diaz-Fierros F (1997). Effects of burning on nutrient balance in an area of gorse (Ulex europaeus L.) scrub. Science of the Total Environment, 204, 271-281.
[42]	Świeca M, Gawlik-Dziki U, Kowalczyk D, Złotek U (2012). Impact of germination time and type of illumination on the antioxidant compounds and antioxidant capacity of Lens culinaris sprouts. Scientia Horticulturae, 140, 87-95.
[43]	Tan YS (2019). Effect of Ash and Aqueous Charcoal Solution on Seed Germination of Four Herbs in Great Hing’an Mountains. Master degree dissertation, Northeast Forestry University, Harbin.
	[谭咏森 (2019). 火烧灰及木炭水提液对大兴安岭四种草本植物种子萌发的影响. 硕士学位论文, 东北林业大学, 哈尔滨.]
[44]	Tavşanoğlu C, Çatav SS, Özüdoğru B (2015). Fire-related germination and early seedling growth in 21 herbaceous species in Central Anatolian steppe. Journal of Arid Environments, 122, 109-116.
[45]	Thomas PB, Morris EC, Auld TD (2003). Interactive effects of heat shock and smoke on germination of nine species forming soil seed banks within the Sydney region. Austral Ecology, 28, 674-683.
[46]	Torres O, Calvo L, Valbuena L (2006). Influence of high temperatures on seed germination of a special Pinus pinaster stand adapted to frequent fires. Plant Ecology, 186, 129-136.
[47]	Vivar-Evans S, Barradas VL, Sánchez-Coronado ME, Gamboa de Buen A, Orozco-Segovia A (2006). Ecophysiology of seed germination of wild Dahlia coccinea (Asteraceae) in a spatially heterogeneous fire-prone habitat. Acta Oecologica, 29, 187-195.
[48]	Wang HCY, Gao ZL, Cao YF, Yu WT, Wang QH, Zhou RL, Zhao F (2023). Response mechanism of forest fire combustion products affecting plant seed germination. World Forestry Research, 36(4), 47-53.
	[王何晨阳, 高仲亮, 曹宇飞, 于闻天, 王秋华, 周汝良, 赵璠 (2023). 森林火灾产物影响植物种子萌发响应机制研究. 世界林业研究, 36(4), 47-53.]
[49]	Yan XF, Wang JL, Zhou LB (2011). Effects of light intensity on Quercus liaotungensis seed germination and seedling growth. Chinese Journal of Applied Ecology, 22, 1682-1688.
	[闫兴富, 王建礼, 周立彪 (2011). 光照对辽东栎种子萌发和幼苗生长的影响. 应用生态学报, 22, 1682-1688.]
[50]	Yang ZH (2023). Study on Seed Dormancy Mechanism of Taxodium distichum var. imbricatum. Master degree dissertation, Southwest University, Chongqing.
	[杨治华 (2023). 池杉种子休眠机理研究. 硕士学位论文, 西南大学, 重庆.]
[51]	Zenner EK, Hibbs DE (2000). A new method for modeling the heterogeneity of forest structure. Forest Ecology and Management, 129, 75-87.
[52]	Zhang L (2004). Eco-physiological characteristics of seed germination of Larix chinensis, a timberline tree. Acta Phytoecologica Sinica, 28, 579-583.
	[张玲 (2004). 林线树种太白红杉种子萌发的生理生态特性. 植物生态学报, 28, 579-583.] DOI
[53]	Zhang M, Zhu JJ (2022). Effects of light and temperature on seed germination of Pinus koraiensis with different provenances. Chinese Journal of Plant Ecology, 46, 613-623.
	[张敏, 朱教君 (2022). 光温条件对不同种源红松种子萌发的影响. 植物生态学报, 46, 613-623.] DOI
[54]	Zhou YZ (2020). Effects of Heat Shock, Smoke and Ash on Seed Germination of Seven Species in Great Hing’an Mountains. Master degree dissertation, Northeast Forestry University, Harbin.
	[周肄智 (2020). 热激、烟熏和火烧灰对大兴安岭七种植物种子萌发的影响. 硕士学位论文, 东北林业大学, 哈尔滨.]
[55]	Zou W, Chen Z, Jiang YQ, Huang MY, Zhang HN, Huang XY, Peng PH, Wang GY (2023). Responses of Pinus yunnanensis seed germination to temperature, light and GA₃. Journal of Sichuan Forestry Science and Technology, 44(1), 84-90.
	[邹雯, 陈卓, 蒋云清, 黄梦月, 张鸿南, 黄汐月, 彭培好, 王国严 (2023). 云南松种子萌发对温度、光照和GA₃的响应. 四川林业科技, 44(1), 84-90.]
[56]	Zuloaga-Aguilar S, Briones O, Orozco-Segovia A (2011). Seed germination of montane forest species in response to ash, smoke and heat shock in Mexico. Acta Oecologica, 37, 256-262.