模拟增温改变川西高山草甸优势植物繁殖物候序列特征

doi:10.17521/cjpe.2017.0133

摘要/Abstract

摘要：

以西南横断山区高山草甸优势植物种珠芽拳参(Polygonum viviparum)和银叶委陵菜(Potentilla leuconota)为研究对象, 将其物候分为花芽期、开花期、凋谢期和种子成熟期4个阶段, 每个阶段又分为开始、峰值和结束3个状态。采用开顶式增温箱进行模拟增温, 连续增温4年后, 于增温第5年的2016年生长季跟踪调查了模拟增温对珠芽拳参和银叶委陵菜的繁殖物候序列的影响, 以探讨高山植物群落对气候变化的响应过程。结果显示, 模拟增温后: 1)珠芽拳参各物候阶段的持续时间缩短; 除凋谢阶段起始、结束期延迟外, 其他状态均有不同程度的提前; 各阶段的过渡期有不同程度的缩短, 繁殖周期缩短; 2)银叶委陵菜各物候阶段的持续时间延长; 凋谢期结束前各状态(除开花峰值外)表现为不同程度的提前; 各阶段过渡期对增温的响应不一致, 繁殖周期延长。结果表明: 完整的繁殖物候序列能更准确地反映植物物候对气候变暖的响应; 植物对环境变化的响应和应对策略存在种间差异, 这种差异可能会进一步改变植物群落组成和结构。

关键词: 高山草甸, 气候变化, 开顶式增温箱, 珠芽拳参, 银叶委陵菜, 繁殖物候

Abstract:

Aims We studied phenological sequences of two dominant plants (Polygonum viviparum and Potentilla leuconota) in an alpine meadow of the Hengduan Mt., western of Sichuan to explore the alpine plants responses on climate change.

Methods Open-top chambers (OTCs) chosen by ITEX were used to monitor the warming in the field. After a four-year experimental warming, in the 5th growing season we recorded the phenological sequences of two dominant species, focusing on plant responses on warming. The sequence was divided into four stages: budding, flowering, withering and ripe seeds. Each stage had three events: first, peak, and last.

Important findings Our results showed that: 1) For P. viviparum, experimental warming elicited a shortening of the duration of each stage, advanced all of the phenological events but the first of withering and ripe seeds, shortened the period of each stage and reduced the duration of entire reproduction. 2) For P. leuconota, experimental warming extended the duration of every stage. All phenological events before the end of withering occurred earlier on experimental warming but the peak of flowering. The period of each stage had inconsistent responses on warming and warming prolonged the duration of entire reproduction. The present results indicated that not all phenological events were equally responsive to experimental warming and an entire sequence could be a more accurate way to evaluate the responses on environmental variation. Therefore, the plastic responses to warming of different species would have effects on community composition and structure.

Key words: alpine meadow, climate change, open-top chamber, Polygonum viviparum, Potentilla leuconota, reproductive phenology

张莉, 王根绪, 冉飞, 彭阿辉, 肖瑶, 杨阳, 杨燕. 模拟增温改变川西高山草甸优势植物繁殖物候序列特征. 植物生态学报, 2018, 42(1): 20-27. DOI: 10.17521/cjpe.2017.0133

ZHANG Li, WANG Gen-Xu, RAN Fei, PENG A-Hui, XIAO Yao, YANG Yang, YANG Yan. Experimental warming changed plants’ phenological sequences of two dominant species in an alpine meadow, western of Sichuan. Chinese Journal of Plant Ecology, 2018, 42(1): 20-27. DOI: 10.17521/cjpe.2017.0133

导出引用管理器 EndNote|Ris|BibTeX

链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2017.0133

https://www.plant-ecology.com/CN/Y2018/V42/I1/20

图/表 6

图1 生长季开顶式增温箱内外空气和土壤的月平均温度、月平均含水量。A, 月平均气温。B, 土壤5 cm深月平均温度。C, 土壤20 cm深月平均温度。D, 土壤5 cm深月平均含水量。E, 土壤20 cm深月平均含水量。

Fig. 1 Monthly mean air temperature, soil temperature, and soil water content inside and outside the open-top chambers during the growing season. A, Monthly mean air temperature. B, Monthly mean soil temperature at 5 cm soil depth. C, Monthly mean soil temperature at 20 cm soil depth. D, Monthly mean soil water content at 5 cm soil depth. E, Monthly mean soil water content at 20 cm soil depth.

图2 珠芽拳参(A)和银叶委陵菜(B)物候序列变化。■、▲和●分别表示各个阶段的起始、峰值和结束状态。负值表示处理与对照相比提前的天数, 正值表示与对照相比延迟的天数。

Fig. 2 Phenological shifts at the sequence of Polygonum viviparum (A) and Potentilla leuconota (B). ■, ▲ and ● symbol represent a phenological shift of first, peak, and last of the four stages, respectively. Negative value represents earlier stations than control in days, and the positive value represents delayed stations than control in days. OTCs, open-top chambers.

表1 各物候指标对模拟增温响应的参数统计

Table 1 Parameter estimates of GLME models investigating phenological sequences responses to experimental warming

阶段 Stage	状态 Station	银叶委陵菜 Potentilla leuconota			珠芽拳参 Polygonum viviparum
阶段 Stage	状态 Station	N	截距 Intercept	OTCs	N	截距 Intercept	OTCs
花芽期 Budding	开始 First	6	5.11^***	-0.03	13	5.19^***	0.01
	峰值 Peak	6	5.17^***	-0.05	13	5.19^***	0.01
	结束 Last	6	5.23^***	-0.01	13	5.28^***	0.02
开花期 Flowering	开始 First	8	5.17^***	-0.06	10	5.28^***	-0.02
	峰值 Peak	8	5.18^***	0.02	10	5.30^***	-0.01
	结束 Last	8	5.27^***	-0.02	10	5.33^***	-0.02
凋谢期 Withering	开始 First	8	5.18^***	-0.05	13	5.25^***	0.02
	峰值 Peak	8	5.27^***	-0.03	13	5.30^***	0.01
	结束 Last	8	5.36^***	0.04	13	5.33^***	0.01
种子成熟期 Ripe seeds	开始 First	8	5.23^***	0.02	12	5.36^***	-0.02
	峰值 Peak	8	5.37^***	0.02	12	5.38^***	-0.02
	结束 Last	8	5.47^***	0.01	12	5.41^***	-0.03

图3 开顶式增温箱模拟增温对珠芽拳参(A)和银叶委陵菜(B)各阶段持续时间长度的影响(平均值±标准误差)。

Fig. 3 Effects of open-top chambers (OTCs) warming on the duration of each stage of Polygonum viviparum (A) and Potentilla leuconota (B)(mean ± SE).

图4 开顶式增温箱模拟增温对珠芽拳参(A)和银叶委陵菜(B)相邻物候阶段峰值期相差天数的影响(平均值±标准误差)。

Fig. 4 Effects of open-top chambers (OTCs) warming on the period between the peak time of the neighboring stages of Polygonum viviparum (A) and Potentilla leuconota (B) (mean ± SE).

附件I 增温后物种盖度和垂直高度的变化(平均值±标准误差)

Appendix I Changes in coverage and height of the two species under experimental warming (mean ± SE). *, p < 0.05

参考文献 36

[1]	Amano T, Smithers RJ, Sparks TH, Sutherland WJ ( 2010). A 250-year index of first flowering dates and its response to temperature changes. Proceedings of the Royal Society of London B: Biological Sciences, 277, 2451-2457. DOI URL PMID
[2]	Arft A, Walker M, Gurevitch J, Alatalo J, Bret-Harte M, Dale M, Diemer M, Gugerli F, Henry G, Jones M ( 1999). Responses of tundra plants to experimental warming: Meta-analysis of the international tundra experiment. Ecological Monographs, 69, 491-511. DOI URL
[3]	Badeck FW, Bondeau A, B?ttcher K, Doktor D, Lucht W, Schaber J, Sitch S ( 2004). Responses of spring phenology to climate change. New Phytologist, 162, 295-309. DOI URL
[4]	Beaubien E, Freeland H ( 2000). Spring phenology trends in Alberta, Canada: Links to ocean temperature. International Journal of Biometeorology, 44, 53-59. DOI URL PMID
[5]	CaraDonna PJ, Iler AM, Inouye DW ( 2014). Shifts in flowering phenology reshape a subalpine plant community. Proceedings of the National Academy of Sciences of the United States of America, 111, 4916-4921. DOI URL
[6]	Cleland EE, Chiariello NR, Loarie SR, Mooney HA, Field CB ( 2006). Diverse responses of phenology to global changes in a grassland ecosystem. Proceedings of the National Academy of Sciences of the United States of America, 103, 13740-13744. DOI URL PMID
[7]	Ding YH, Wang HJ ( 2015). Newly acquired knowledge on the scientific issues related to climate change over the recent 100 years in China. Chinese Science Bulletin, 61, 1029-1041.
	[ 丁一汇, 王会军 ( 2015). 近百年中国气候变化科学问题的新认识. 科学通报, 61, 1029-1041.]
[8]	Dorji T, Totland ?, Moe SR, Hopping KA, Pan J, Klein JA ( 2013). Plant functional traits mediate reproductive phenology and success in response to experimental warming and snow addition in Tibet. Global Change Biology, 19, 459-472. DOI URL PMID
[9]	Dudgeon SR, Steneck RS, Davison IR, Vadas RL ( 1999). Coexistence of similar species in a space-limited intertidal zone. Ecological Monographs, 69, 331-352. DOI URL
[10]	Forrest J, Miller-Rushing AJ ( 2010). Toward a synthetic understanding of the role of phenology in ecology and evolution. The Royal Society, 365, 3101-3112. DOI URL PMID
[11]	Gugger S, Kesselring H, Stocklin J, Hamann E ( 2015). Lower plasticity exhibited by high-versus mid-elevation species in their phenological responses to manipulated temperature and drought. Annals of Botany, 116, 953-962. DOI URL PMID
[12]	Hollister RD, Webber PJ, Bay C ( 2005). Plant response to temperature in northern Alaska: Implications for predicting vegetation change. Ecology, 86, 1562-1570. DOI URL
[13]	Iler AM, H?ye TT, Inouye DW, Schmidt NM ( 2013). Nonlinear flowering responses to climate: Are species approaching their limits of phenological change? Philosophical Transactions of the Royal Society of London B: Biological Sciences, 368, 20120489, doi: 10.1098/rstb.2012.0489. DOI URL PMID
[14]	Inouye DW ( 2008). Effects of climate change on phenology, frost damage, and floral abundance of montane wildflowers. Ecology, 89, 353-362. DOI URL
[15]	IPCC (Intergovernmental Panel on Climate Change) ( 2013) : Contribution of Working Group I to the Fifth Assessment Report of the Intergovernmental Panel on climate change. In: Stocker TF, Qin DH, Plattner GK, Tignor M, Allen SK, Boschung J, Nauels A, Xia Y, Bex V, Midgley PM eds. Climate Change in 2013: The Physical Science Basis. Cambridge University Press, Cambridge, UK.
[16]	Jonasson S, Havstr?m M, Jensen M, Callaghan TV ( 1993). In situ mineralization of nitrogen and phosphorus of arctic soils after perturbations simulating climate change. Oecologia, 95, 179-186.
[17]	Klein JA, Harte J, Zhao XQ ( 2004). Experimental warming causes large and rapid species loss, dampened by simulated grazing, on the Tibetan Plateau. Ecology Letters, 7, 1170-1179. DOI URL
[18]	Li ZX, He YQ, Xin HJ, Wang CF, Jia WX, Zhang W, Liu J ( 2010). Spatio-temporal variations of temperature and precipitation in Mts. Hengduan Region during 1960-2008. Acta Geographica Sinica, 65, 563-579. DOI URL
	[ 李宗省, 何元庆, 辛惠娟, 王春凤, 贾文雄, 张蔚, 刘婧 ( 2010). 我国横断山区1960-2008年气温和降水时空变化特征. 地理学报, 65, 563-579.] DOI URL
[19]	Liu YZ, Reich PB, Li GY, Sun SC ( 2011). Shifting phenology and abundance under experimental warming alters trophic relationships and plant reproductive capacity. Ecology, 92, 1201-1207. DOI URL
[20]	Memmott J, Craze PG, Waser NM, Price MV ( 2007). Global warming and the disruption of plant-pollinator interactions. Ecology Letters, 10, 710-717. DOI URL
[21]	Meng FD, Cui SJ, Wang SP, Duan JC, Jiang LL, Zhang ZH, Luo CY, Wang Q, Zhou Y, Li XN, Zhang LR, Dorji T, Li YN, Du MY, Wang GJ ( 2016). Changes in phenological sequences of alpine communities across a natural elevation gradient. Agricultural and Forest Meteorology, 224, 11-16. DOI URL
[22]	Menzel A, Sparks TH, Estrella N, Koch E, Aasa A, Ahas R, Alm-Kubler K, Bissolli P, Braslavská O, Briede A ( 2006). European phenological response to climate change matches the warming pattern. Global Change Biology, 12, 1969-1976. DOI URL
[23]	Pe?uelas J, Filella I ( 2001). Responses to a warming world. Science, 294, 793-795. DOI URL PMID
[24]	Pe?uelas J, Filella I ( 2009). Phenology feedbacks on climate change. Science, 324, 887-888. DOI URL PMID
[25]	Pe?uelas J, Filella I, Comas P ( 2002). Changed plant and animal life cycles from 1952 to 2000 in the Mediterranean region. Global Change Biology, 8, 531-544. DOI URL
[26]	Pepin N, Bradley RS, Diaz HF, Baraer M, Caceres EB, Forsythe N, Fowler H, Greenwood G, Hashmi MZ, Liu XD, Miller JR, Ning L, Ohmura A, Palazzi E, Rangwala I, Sch?ner W, Severskiy I, Shahgedanova M, Wang MB, Williamson SN, Yang DQ ( 2015). Elevation-dependent warming in mountain regions of the world. Nature Climate Change, 5, 424-430. DOI URL
[27]	Post ES, Pedersen C, Wilmers CC, Forchhammer MC ( 2008). Phenological sequences reveal aggregate life history response to climatic warming. Ecology, 89, 363-370. DOI URL
[28]	Price MV, Waser NM ( 1998). Effects of experimental warming on plant reproductive phenology in a subalpine meadow. Ecology, 79, 1261-1271. DOI URL
[29]	Root TL, Price JT, Hall KR, Schneider SH, Rosenzweig C, Pounds JA ( 2003). Fingerprints of global warming on wild animals and plants. Nature, 421, 57-60. DOI URL PMID
[30]	Schwartz MD, Reiter BE ( 2000). Changes in North American spring. International Journal of Climatology, 20, 929-932. DOI URL
[31]	Thomas CD, Cameron A, Green RE, Bakkenes M, Beaumont LJ, Collingham YC, Erasmus BF, de Siqueira MF, Grainger A, Hannah L ( 2004). Extinction risk from climate change. Nature, 427, 145-148. DOI URL PMID
[32]	Tilman D, Lehman CL, Thomson KT ( 1997). Plant diversity and ecosystem productivity: Theoretical considerations. Proceedings of the National Academy of Sciences of the United States of America, 94, 1857-1861. DOI URL PMID
[33]	Totland ?, Schulte-Herbrüggen B ( 2003). Breeding system, insect flower visitation, and floral traits of two alpine Cerastium species in Norway. Arctic, Antarctic, and Alpine Research, 35, 242-247.
[34]	Wang SP, Meng FD, Duan JC, Wang YF, Cui XY, Piao SL, Niu HS, Xu GP, Luo CY, Zhang ZH ( 2014). Asymmetric sensitivity of first flowering date to warming and cooling in alpine plants. Ecology, 95, 3387-3398. DOI URL
[35]	Wolkovich EM, Cook BI, Allen JM, Crimmins TM, Betancourt JL, Travers SE, Pau S, Regetz J, Davies TJ, Kraft NJ ( 2012). Warming experiments under predict plant phenological responses to climate change. Nature, 485, 494-497. DOI URL PMID
[36]	Yang Y, Wang GX, Klanderud K, Wang JF, Liu GS ( 2015). Plant community responses to five years of simulated climate warming in an alpine fen of the Qinghai-Tibetan Plateau. Plant Ecology & Diversity, 8, 211-218. DOI URL