Effects of simulated drought on plant phenology and productivity in an alpine meadow in Northern Tibet

doi:10.11686/cyxb2020099

Abstract

Abstract:

This research aimed to elucidate the effects of drought events on the phenology and productivity of the plant community in Tibetan alpine meadow. A rainfall interception experiment comprising drought treatments imposed early or late in the growing season （SE and SL， respectively） was carried out in alpine meadow of Northern Tibet， over two years （2016-2017）. Results showed that： 1） The plant phenology had a contrasting response to SE and SL drought treatments. Plant phenology of key species was more sensitive to SE. SE significantly delayed plant green update， and led to a shorter growing season and reproductive duration （P<0.05）， while SL had no significant effect on plant phenology. 2） Biomass of the plant community and different functional groups decreased under the drought treatments. Community biomass decreased by 62.9% （P<0.05）. The height and coverage of the community was suppressed by drought， while the response of height and coverage differed between grasses and forbs. 3） Community height and biomass were significantly positively correlated with the growing season and reproductive duration （P<0.05）. The results indicate that the productivity of alpine meadow grassland was regulated by phenology in response to drought， mainly through the impact on plant height. Moreover， the responses to drought varied between different functional groups， and suggest that increasing and intensified drought may lead to change in the community structure of alpine meadow.

Key words: Northern Tibet, alpine meadow, drought events, phenological period, productivity

Wen-rong LUO, Guo-zheng HU, Ganjurjav H, Qing-zhu GAO, Yan LI, Yi-qing Ge, Yu LI, Shi-cheng HE, Luo-bu DANJIU. Effects of simulated drought on plant phenology and productivity in an alpine meadow in Northern Tibet[J]. Acta Prataculturae Sinica, 2021, 30(2): 82-92.

Figures/Tables 7

References 50

1	IPCC. Climate change 2013： The physical science basis. Cambridge： Cambridge University Press， 2013.
2	Easterling D R， Meehl G A， Parmesan C， et al. Climate extremes： Observations， modeling， and impacts. Science， 2000， 289： 2068- 2074.
3	Allen C D， Macalady A K， Chenchouni H， et al. A global overview of drought and heat-induced tree mortality reveals emerging climate change risks for forests. Forest Ecology & Management， 2010， 259（4）： 660-684.
4	Walter J， Nagy L， Hein R， et al. Do plants remember drought？ Hints towards drought-memory in grasses. Environmental & Experimental Botany， 2011， 71（1）： 34-40.
5	Thomas C D， Cameron A， Green R E， et al. Extinction risk from climate change. Nature， 2004， 427： 145-148.
6	Gao Q Z， Wan Y F， Li Y E， et al. Climate change risk and integrated adaptation management of alpine grassland ecosystem in Northern Tibet. Beijing： Low Carbon Agriculture Seminar， 2010.
	高清竹，万运帆，李玉娥，等. 藏北地区高寒草地生态系统气候变化风险及其综合适应管理. 北京：低碳农业研讨会， 2010.
7	Gao Q Z， Li Y E， Xu H M， et al. Adaptation strategies of climate variability impacts on alpine grassland ecosystems in Tibetan Plateau. Mitigation and Adaptation Strategies for Global Change， 2014， 19： 199-209.
8	Yang C Y， Shen W S， Lin N F. Climate change and its regional differences over the Tibet Plateau. Arid Land Geography， 2014， 37（2）： 290-298.
	杨春艳，沈渭寿，林乃峰. 西藏高原气候变化及其差异性. 干旱区地理， 2014， 37（2）： 290-298.
9	Sheng W P， Gao Q Z， Li Y E， et al. Characteristic of climate change in Northern Tibet and its impact. Plateau Meteorology， 2008， 27（3）： 509-516.
	盛文萍，高清竹，李玉娥，等. 藏北地区气候变化特征及其影响分析. 高原气象， 2008， 27（3）： 509-516.
10	Yuan L， Liu Y L， Ma P F. Temporal and spatial patterns of drought based on standard precipitation index （SPI） in Tibet during 1981-2013. Chinese Agricultural Science Bulletin， 2015， 31（25）： 228-234.
	袁雷，刘依兰，马鹏飞. 基于标准化降水指数的1981-2013年西藏干旱时空特征分析. 中国农学通报， 2015， 31（25）： 228-234.
11	Wang Z P， Zhang X Z， He Y T， et al. Effects of precipitation changes on the precipitation use efficiency and aboveground productivity of alpine steppe-meadow on Northern Tibetan Plateau. Chinese Journal of Applied Ecology， 2018， 29（6）： 1822-1828.
	王志鹏，张宪洲，何永涛，等. 降水变化对藏北高寒草原化草甸降水利用效率及地上生产力的影响. 应用生态学报， 2018， 29（6）： 1822-1828.
12	Ge S， Ai J F， Nima C R， et al. Preliminary analysis of drought climate change characteristics in Tibet in recent 50 years by CI Index. Tibet’s Science & Technology， 2011（5）： 63-67.
	格桑，艾俊峰，尼玛次仁，等. 初步采用CI指标分析西藏近50年干旱气候变化特征. 西藏科技， 2011（5）： 63-67.
13	Wang J S， Zhang X Z， Chen B X， et al. Causes and restoration of degraded alpine grassland in Northern Tibet. Journal of Resources and Ecology， 2013， 4（1）： 43-49.
14	Xia J， Wan S. Independent effects of warming and nitrogen addition on plant phenology in the Inner Mongolian steppe. Annals of Botany， 2013， 111（6）： 1207-1217.
15	Polley H W， Derner J D， Jackson R B， et al. Impacts of climate change drivers on C₄ grassland productivity： Scaling driver effects through the plant community. Journal of Experimental Botany， 2014， 65（13）： 3415-3424.
16	Bernal M， Estiarte M， Peñuelas J. Drought advances spring growth phenology of the Mediterranean shrub Erica multiflora. Plant Biology， 2011， 13（2）： 252-257.
17	Zhao G S， Shi P L， Zong N， et al. Decling precipitation enhances the effect of warming on phenological variation in a Semiarid Tibeten Meadow steppe. Journal of Resources and Ecology， 2017， 8（1）： 50-56.
18	Mu C X， Sun G， Luo P， et al. Flowering responses of alpine meadow plant in the Qinghai-Tibetan Plateau to extreme drought imposed in different periods. Chinese Journal of Applied & Environmental Biology， 2013， 19（2）： 272-279.
	牟成香，孙庚，罗鹏，等. 青藏高原高寒草甸植物开花物候对极端干旱的响应. 应用与环境生物学报， 2013， 19（2）： 272-279.
19	Li X H， Chen S H， Han F. Effects of drought on grassland turning green period in Inner Mongolia， China. Pratacultural Science， 2013， 30（3）： 452-456.
	李兴华，陈素华，韩芳. 干旱对内蒙古草地牧草返青期的影响. 草业科学， 2013， 30（3）： 452-456.
20	Ma G G. Responses of plant phenology to changes of precipitation regimes in a temperate steppe in Inner Mongolia， China. Kaifeng： Henan University， 2015.
	马改改. 中国内蒙古温带典型草原植物物候对降水格局改变的响应. 开封：河南大学， 2015.
21	Jentsch A， Kreyling J， Boettcher-treschkow J. Beyond gradualwarming： Extreme weather events alter flower phenology of European grassland and heath species. Global Change Biology， 2009， 15（4）： 837-849.
22	Cornelius C， Leingartner A， Hoiss B， et al. Phenological response of grassland species to manipulative snowmelt and drought along an altitudinal gradient. Journal of Experimental Botany， 2013， 64（1）： 241-251.
23	Ogaya R， Penuelas J. Tree growth， mortality， and above-ground biomass accumulation in a Holm Oak forest under a five-year experimental field drought. Plant Ecology， 2007， 189（2）： 291-299.
24	Huang C Y， Anderegg W R L. Large drought-induced aboveground live biomass losses in southern Rocky Mountain aspen forests. Global Change Biology， 2012， 18（3）： 1016-1027.
25	Ren L W， Wang X T， Liu M C， et al. Effects of drought stress on soil moisture dynamics and water use efficiency in corn. Chinese Agricultural Science Bulletin， 2015， 31（32）： 142-147.
	任丽雯，王兴涛，刘明春，等. 干旱胁迫对土壤水分动态及玉米水分利用效率影响研究. 中国农学通报， 2015， 31（32）： 142-147.
26	Tian H Q， Xu X F， Song X. Drought impacts on terrestrial ecosystem productive. Journal of Plant Ecology， 2007， 31（2）： 231-241.
	田汉勤，徐小锋，宋霞. 干旱对陆地生态系统生产力的影响. 植物生态学报， 2007， 31（2）： 231-241.
27	Zhang H， Wang X P， Zhang Y F， et al. Responses of plant growth of different life forms to rainfall amount changes in an arid desert area. Chinese Journal of Ecology， 2015， 34（7）： 1847-1853.
	张浩，王新平，张亚峰，等. 干旱荒漠区不同生活型植物生长对降雨量变化的响应. 生态学杂志， 2015， 34（7）： 1847-1853.
28	Sun Y， He M Z， Wang L. Effects of precipitation control on plant diversity and biomass in a desert region. Acta Ecologica Sinica， 2018， 38（7）： 2425-2433.
	孙岩，何明珠，王立. 降水控制对荒漠植物群落物种多样性和生物量的影响. 生态学报， 2018， 38（7）： 2425-2433.
29	Lei T J. Quantitative assessment of the impacts of droughts on grassland productivity. Acta Geodaetica et Cartographica Sinica， 2017， 46（1）： 134.
	雷添杰. 干旱对草地生产力影响的定量评估研究. 测绘学报， 2017， 46（1）： 134.
30	Chu D， Deji Y Z， Pubu C R， et al. The response of typical vegetation growth to climate conditions in North Tibetan Plateau. Journal of Applied Meteorological Science， 2007（6）： 832-839.
	除多，德吉央宗，普布次仁，等. 西藏藏北高原典型植被生长对气候要素变化的响应. 应用气象学报， 2007（6）： 832-839.
31	Zhu J T， Zhang Y J， Liu Yao J. Effects of short-term grazing exclusion on plant phenology and reproductive succession in a Tibetan Alpine Meadow. Entific Reports， 2016， 6（1）： 27781.
32	Desclaux D， Roumet P. Impact of drought stress on the phenology of two soybean （Glycine max L. Merr） cultivars. Field Crops Research， 1996， 46（1/2/3）： 61-70.
33	Yang S， Wang B X， Xu X， et al. Sex-specific responses of flowering phenology and floral morphology of Humulus scandens to drought. Plant Diversity and Resources， 2014， 36（5）： 653-660.
	杨帅，王碧霞，胥晓，等. 葎草雌雄植株开花物候和花器官对干旱的响应差异. 植物分类与资源学报， 2014， 36（5）： 653-660.
34	Borchert R， Rivera G， Hagnauer W. Modification of vegetative phenology in a tropical semi-deciduous forest by abnormal drought and rain. Biotropica， 2002， 34： 27-39.
35	Wu J G. Effects of changing in precipitation and temperature on the photosynthesis and related physiological parameters of straw-yellow gentian. Chinese Journal of Grassland， 2010， 32（5）： 73-79.
	吴建国. 降雨量和温度变化对麻花艽叶片光合作用及相关生理参数的影响. 中国草地学报， 2010， 32（5）： 73-79.
36	Song C Q， You S C， Ke L H， et al. Spatiotemporal dynamics of land cover in Northern Tibetan Plateau with responses to climate change. Chinese Journal of Applied Ecology， 2011， 22（8）： 2091-2097.
	宋春桥，游松财，柯灵红，等. 藏北高原地表覆盖时空动态及其对气候变化的响应. 应用生态学报， 2011， 22（8）： 2091-2097.
37	Holub P， Fiala K. Effects of artificially varying amounts of rainfall on two semi-natural grassland types. Journal of Vegetation Science， 2013， 24（3）： 518-529.
38	Wang C T， Wang Q J， Shen Z X， et al. Response of biodiversity and productivity to simulated rainfall on an alpine Kobresia humilis meadow. Acta Botanica Boreali-Occidentalia Sinica， 2003， 23（10）： 1713-1718.
	王长庭，王启基，沈振西，等. 高寒矮蒿草草甸群落植物多样性和初级生产力对模拟降水的响应. 西北植物学报， 2003， 23（10）： 1713-1718.
39	Zhao N W， Gong J P， Kong X P. Effect on the productive forces of grassland of phenology climatic factors in the Gonghe basin temperate steppe. Gansu Animal and Veterinary Sciences， 2010（2）： 3-9.
	赵年武，龚建平，孔祥萍. 气候因子对共和盆地温性草原物候期草地生产力的影响分析. 甘肃畜牧兽医， 2010（2）： 3-9.
40	Zhu J， Zhang Y， Jiang L. Experimental warming drives a seasonal shift of ecosystem carbon exchange in Tibetan Alpine Meadow. Agricultural & Forest Meteorology， 2017， 233 （Complete）： 242-249.
41	Bai Y F， Xu Z X. A model of above-ground biomass of aneurolepidium chinense community in response to seasonal precipitation. Acta Prataculturae Sinica， 1997， 6（2）： 1-6.
	白永飞，许志信. 降水量的季节分配对羊草草原群落地上部生物量影响的数学模型. 草业学报， 1997， 6（2）： 1-6.
42	Xin J， Zha T， Gong J， et al. Carbon and water exchange over a temperate semi-arid shrubland during three years of contrasting precipitation and soil moisture patterns. Agricultural & Forest Meteorology， 2016（228/229）： 120-129.
43	Still C J， Berry J A， Collatz G J， et al. Global distribution of C₃ and C₄ vegetation： Carbon cycle implications. Global Biogeochemical Cycles， 2003， 17（1）： 6-14.
44	Yuan W P. Study on phenological characteristics of typical grassland and its influence on ecosystem function. Beijing： Chinese Academy of Science， 2007.
	袁文平. 典型草原物候特征及其对生态系统功能的影响研究. 北京：中国科学院， 2007.
45	Bardgett R D， Wal R V D， Jónsdóttir I S， et al. Temporal variability in plant and soil nitrogen pools in a high-arctic ecosystem. Soil Biology & Biochemistry， 2007， 39（8）： 2129-2137.
46	Wipf S， Rixen C. A review of snow manipulation experiments in arctic and alpine tundra ecosystems.Polar Research， 2010， 29： 95-109.
47	Monson R K， Sparks J P， Rosenstiel T N， et al. Climatic influences on net ecosystem CO₂ exchange during the transition from wintertime carbon source to springtime carbon sink in a high-elevation， subalpine forest. Oecologia， 2005， 146（1）： 130-147.
48	Li X Q. Phenology of plant community in alpine marsh meadow on the eastern margin of Qinghai-Xizang Plateau and its relationship with other characters. Lanzhou： Lanzhou University， 2010.
	李向前. 青藏高原东缘高寒沼泽化草甸植物群落物候及其与其它性状的关系. 兰州：兰州大学， 2010.
49	Chen G G. Study on phenology and plant characters of alpine meadow flora in the eastern margin of Qinghai-Xizang Plateau at flowering stage. Lanzhou： Lanzhou University， 2016.
	陈冠光. 青藏高原东缘高寒草甸植物群落花期物候以及植物性状的研究. 兰州：兰州大学， 2016.
50	Yu H Y，Wang L. The correlation between phenology and plant height of herbaceous plants species on alpine meadow. Journal of Inner Mongolia Agricultural University （Natural Science Edition）， 2018， 39（4）： 39-46.
	于海英，王力. 高寒草甸草本植物物候期和植株高度的关系研究.内蒙古农业大学学报（自然科学版）， 2018， 39（4）： 39-46.

物候 Phenology	处理 Treatments	高山嵩草K. pygmaea		早熟禾P. pratensis		钉柱委陵菜P. saundersiana		菊叶委陵菜P. tanacetifolia
物候 Phenology	处理 Treatments	2016	2017	2016	2017	2016	2017	2016	2017
返青期 Green-up date	CK	153.0±0.0b	128.3±1.7a	156.3±1.7b	123.3±3.3a	148.3±4.7b	115.0±0.0a	146.3±1.7b	115.0±0.0a
	SE	179.0±0.0a	123.3±1.7b	183.7±2.9a	121.7±1.7a	174.0±3.2a	115.0±0.0a	173.7±10.7a	115.0±0.0a
	SL	154.7±1.7b	125.0±0.0ab	151.3±1.7b	121.7±1.7a	148.3±4.7b	115.0±0.0a	151.3±1.7b	115.0±0.0a
枯黄期 Withered date	CK	215.3±3.7a	214.3±1.7a	244.0±0.0a	223.7±1.7a	222.3±1.7b	222.0±0.0a	222.3±1.7b	221.7±3.2a
	SE	220.7±1.7a	206.0±5.0a	244.0±0.0a	216.7±10.3a	230.7±3.3a	212.7±3.3b	230.7±3.3a	216.0±0.0a
	SL	212.0±2.0a	211.0±0.0a	234.0±0.0a	214.3±1.7a	217.3±1.7b	216.0±0.0ab	211.7±3.7b	216.0±0.0a
生长季长度 Growing season duration	CK	62.3±3.7a	86.0±2.9a	87.7±1.7a	100.3±4.4a	74.0±5.7a	107.0±0.0a	76.0±0.0a	106.7±3.2a
	SE	41.7±1.7b	82.7±6.0a	60.3±2.9b	95.0±12.0a	56.7±5.0b	97.7±3.3b	57.0±7.6b	101.0±0.0a
	SL	57.3±3.7a	86.0±0.0a	82.7±1.7a	92.7±3.3a	69.0±3.0ab	101.0±0.0ab	60.3±3.2ab	101.0±0.0a
繁殖期 Reproductive duration	CK	12.0±1.0a	56.0±0.0a	53.7±3.7a	26.3±2.3a	56.0±2.9a	51.0±0.0a	50.7±3.2a	49.3±1.7a
	SE	3.3±3.3a	52.7±3.3a	32.3±3.7b	24.0±0.0a	22.3±4.5b	47.0±4.0a	20.0±2.9c	53.5±2.5a
	SL	5.7±3.2a	57.7±1.7a	45.3±3.2a	29.0±5.0a	49.3±1.7a	51.0±0.0a	37.7±4.5b	49.3±1.7a

物候 Phenology	处理 Treatments	高山嵩草K. pygmaea		早熟禾P. pratensis		钉柱委陵菜P. saundersiana		菊叶委陵菜P. tanacetifolia
物候 Phenology	处理 Treatments	2016	2017	2016	2017	2016	2017	2016	2017
返青期 Green-up date	CK	153.0±0.0b	128.3±1.7a	156.3±1.7b	123.3±3.3a	148.3±4.7b	115.0±0.0a	146.3±1.7b	115.0±0.0a
	SE	179.0±0.0a	123.3±1.7b	183.7±2.9a	121.7±1.7a	174.0±3.2a	115.0±0.0a	173.7±10.7a	115.0±0.0a
	SL	154.7±1.7b	125.0±0.0ab	151.3±1.7b	121.7±1.7a	148.3±4.7b	115.0±0.0a	151.3±1.7b	115.0±0.0a
枯黄期 Withered date	CK	215.3±3.7a	214.3±1.7a	244.0±0.0a	223.7±1.7a	222.3±1.7b	222.0±0.0a	222.3±1.7b	221.7±3.2a
	SE	220.7±1.7a	206.0±5.0a	244.0±0.0a	216.7±10.3a	230.7±3.3a	212.7±3.3b	230.7±3.3a	216.0±0.0a
	SL	212.0±2.0a	211.0±0.0a	234.0±0.0a	214.3±1.7a	217.3±1.7b	216.0±0.0ab	211.7±3.7b	216.0±0.0a
生长季长度 Growing season duration	CK	62.3±3.7a	86.0±2.9a	87.7±1.7a	100.3±4.4a	74.0±5.7a	107.0±0.0a	76.0±0.0a	106.7±3.2a
	SE	41.7±1.7b	82.7±6.0a	60.3±2.9b	95.0±12.0a	56.7±5.0b	97.7±3.3b	57.0±7.6b	101.0±0.0a
	SL	57.3±3.7a	86.0±0.0a	82.7±1.7a	92.7±3.3a	69.0±3.0ab	101.0±0.0ab	60.3±3.2ab	101.0±0.0a
繁殖期 Reproductive duration	CK	12.0±1.0a	56.0±0.0a	53.7±3.7a	26.3±2.3a	56.0±2.9a	51.0±0.0a	50.7±3.2a	49.3±1.7a
	SE	3.3±3.3a	52.7±3.3a	32.3±3.7b	24.0±0.0a	22.3±4.5b	47.0±4.0a	20.0±2.9c	53.5±2.5a
	SL	5.7±3.2a	57.7±1.7a	45.3±3.2a	29.0±5.0a	49.3±1.7a	51.0±0.0a	37.7±4.5b	49.3±1.7a

因子 Factors	返青期 Green-up date	枯黄期 Withered date	生长季长度 Growing season duration	繁殖期 Reproductive duration
Y	1063.786^***	55.914^***	317.972^***	122.336^***
T	45.236^***	10.662^***	19.911^***	37.700^***
S	12.053^***	27.827^***	20.562^***	34.392^***
Y×T	53.091^***	12.525^***	7.038^**	28.625^***
Y×S	1.108	11.621^***	5.945^**	132.034^***
T×S	0.477	0.730	0.541	2.244
Y×T×S	0.354	0.434	0.692	3.138^*

因子 Factors	返青期 Green-up date	枯黄期 Withered date	生长季长度 Growing season duration	繁殖期 Reproductive duration
Y	1063.786^***	55.914^***	317.972^***	122.336^***
T	45.236^***	10.662^***	19.911^***	37.700^***
S	12.053^***	27.827^***	20.562^***	34.392^***
Y×T	53.091^***	12.525^***	7.038^**	28.625^***
Y×S	1.108	11.621^***	5.945^**	132.034^***
T×S	0.477	0.730	0.541	2.244
Y×T×S	0.354	0.434	0.692	3.138^*

特征 Features	处理 Treatments	禾莎类草Grasses and sedges		杂类草Forbs
特征 Features	处理 Treatments	2016	2017	2016	2017
高度 Height (cm)	CK	8.5±1.1a	8.1±0.8a	1.5±0.1a	1.3±0.2a
	SE	4.4±0.6b	5.1±1.4a	1.0±0.0b	1.3±0.2a
	SL	7.4±0.8ab	4.8±1.0a	1.8±0.2a	1.5±0.1a
盖度Coverage (%)	CK	38.0±5.6a	42.6±2.3a	27.4±1.7a	19.8±0.3a
	SE	31.7±5.5a	40.0±3.3a	19.7±4.8ab	15.1±3.4a
	SL	35.7±2.4a	10.0±2.5b	14.1±1.2b	15.1±2.2a
生物量Biomass (g·m^-2)	CK	45.8±9.8a	44.1±2.7a	27.0±2.4a	20.4±3.0a
	SE	20.3±5.5b	26.0±6.8b	12.7±3.1b	15.0±3.5a
	SL	36.1±3.2ab	6.3±2.2c	16.3±1.8b	17.6±3.4a