青海海北高寒草地12种植物花期物候对氮磷养分添加的响应
阐明氮、磷养分供给改变对高寒植物和植物群落花期物候期的影响是研究高寒草地生态系统响应全球变化的重要内容之一。为预测大气氮、磷沉降持续增加下高寒植物花期物候的演变和高寒草地植物群落花期物候格局,本研究在青海省门源县高寒草地采用定株标记法对12种高寒植物在9种不同氮、磷养分添加下的花期物候进行了观测。结果表明:1)氮、磷养分添加对整体花期物候的影响以使始花期提前、花期持续时间延长、终花期无显著变化为主(P > 0.05);2)不同氮、磷养分添加对高寒植物花期物候的影响基本一致,氮、磷养分添加对高寒植物花期物候无显著交互作用影响(P > 0.05);3)在物种水平上,不同氮、磷养分添加处理下大部分物种始花期提前、花期持续时间延长,终花期无一致变化趋势。因此,研究区域高寒草地在大气氮、磷沉降持续增加情况下,植物群落始花期倾向于提前、花期持续时间延长、终花期变化不明显;不同物种始花期、花期持续时间变化较为一致,但终花期的响应会出现一定的差异。
English
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参考文献
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[2] 李兰平. 青藏高原东部高寒草甸植物开花物候研究. 兰州: 兰州大学博士学位论文, 2017. LI L P. The study of flowering phenology of alpine plant on eastern Tibetan Plateau. PhD Thesis. Lanzhou: Lanzhou University, 2017.
[3] LI L P, LI Z K, CADOTTE W M, JIA P, CHEN G G, JIN S L, DU G Z. Phylogenetic conservatism and climate factors shape flowering phenology in alpine meadows. Oecologia, 2016, 182(2): 419-428. doi: 10.1007/s00442-016-3666-6
[4] WALTHER G R, POST E, CONVEY P, MENZEL A, PARMESAN C, BEEBEE T J C, FROMENTIN J M, HOEGH-GULDBER O, BAIRLEIN F. Ecological responses to recent climate change. Nature, 2002, 416: 389-395. doi: 10.1038/416389a
[5] CLELAND E E, CHUINE I, MENZEL A, MOONEY A H, SCHWARTZ D M. Shifting plant phenology in response to global change. Trends in Ecology and Evolution, 2007, 22(7): 357-365. doi: 10.1016/j.tree.2007.04.003
[6] 代武君, 金慧颖, 张玉红, 周志强, 刘彤. 植物物候学研究进展. 生态学报, 2020, 40(19): 6705-6719. DAI W J, JIN H Y, ZHANG Y H, ZHOU Z Q, LIU T. Advances in plant phenology. Acta Ecologica Sinica, 2020, 40(19): 6705-6719.
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[9] PAN Y P, LIU B W, CAO J, LIU J, TIAN S L, DU E Z. Enhanced atmospheric phosphorus deposition in Asia and Europe in the past two decades. Atmospheric and Oceanic Science Letters, 2021, 14(5): 100051.
[10] JIANG J, WANG Y P, YANG Y H, YU M X, WANG C, YAN J H. Interactive effects of nitrogen and phosphorus additions on plant growth vary with ecosystem type. Plant and Soil, 2019, 440(1): 523-537.
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[13] SHERRY R A, ZHOU X H, GU S L, ARNONE J A, SCHIMEL D S, VERBURG P S, WALLACE L L, LUO Y Q. Divergence of reproductive phenology under climate warming. Proceedings of the National Academy of Science, 2007, 104(1): 198-202. doi: 10.1073/pnas.0605642104
[14] 刘旭东, 章志龙, 杜国祯. 高寒草甸主要组分种开花物候对氮素添加的响应. 草业科学, 2021, 38(7): 1240-1249. LIU X D, ZHANG Z L, DU G Z. Response of dominant and common species flowering phenology to nitrogen addition in an alpine meadow. Pratacultural Science, 2021, 38(7): 1240-1249.
[15] SMITH J G, SCONIERS W, SPASOJEVIC M J, ASHTON W A, SUDING K N. Phenological changes in alpine plants in response to increased snowpack, temperature, and nitrogen. Arctic, Antarctic, and Alpine Research, 2012, 44(1): 135-142. doi: 10.1657/1938-4246-44.1.135
[16] LIU Y Z, MIAO R H, CHEN A Q, MIAO Y, LIU Y J, WU X W. Effects of nitrogen addition and mowing on reproductive phenology of three early-flowering forb species in a Tibetan alpine meadow. Ecological Engineering, 2017, 99: 119-125. doi: 10.1016/j.ecoleng.2016.11.033
[17] 杜国祯, 赵松岭. 草本群落中种多样性维持机制: 花期不同步性. 草业学报, 1997, 6(3): 1-7. DU G Z, ZHAO S L. Mechanism of diversity maintenance in herbaceous plant community: Asynchronism in flowering time. Acta Prataculturae Sinica, 1997, 6(3): 1-7.
[18] FORREST J, MILLER-RUSHING A J. Toward a synthetic understanding of the role of phenology in ecology and evolution. Philosophical Transactions of the Royal Society B: Biological Sciences, 2010, 365(1555): 3101-3112. doi: 10.1098/rstb.2010.0145
[19] CLARK C M, CLELAND E E, COLLINS S L, FARGIONE J E, GOUGH L, GROSS K L, PENNINGS S C, SUDING K N, GRACE J B. Environmental and plant community determinants of species loss following nitrogen enrichment. Ecology Letters, 2017, 10(7): 596-607.
[20] 周小龙. 高寒草甸植物群落结构组建和生产力对施肥的响应机制. 兰州: 兰州大学博士学位论文, 2016. ZHOU X L. The effect of fertilization on community assembly and production in alpine meadow community. PhD Thesis. Lanzhou: Lanzhou University, 2016.
[21] 李元恒. 内蒙古典型草原植物生殖物候对气候变化和人为干扰的响应. 兰州: 甘肃农业大学博士学位论文, 2008. LI Y H. Responses of reproductive phenology of Inner Mongolia typical steppe plants under climatic change and artificial interference. PhD Thesis. Lanzhou: Gansu Agricultural University, 2008.
[22] CLELAND E E, CHIARIELLO N R, LOARIE S R, MOONEY H A, FIELD C B. Diverse responses of phenology to global changes in a grassland ecosystem. Proceedings of the National Academy of Sciences, 2006, 103(37): 13740-13744. doi: 10.1073/pnas.0600815103
[23] BEARD K H, KELSEY K C, LEFFLER A J, WELKER J M. The missing angle: Ecosystem consequences of phenological mismatch. Trends in Ecology and Evolution, 2019, 34(10): 885-888. doi: 10.1016/j.tree.2019.07.019
[24] 施雨含, 任宗昕, 赵延会, 王红. 气候变化对植物-传粉昆虫的分布区和物候及其互作关系的影响. 生物多样性, 2021, 29(4): 495-506. doi: 10.17520/biods.2020196 SHI Y H, REN Z X, ZHAO Y H, WANG H. Effect of climate change on the distribution and phenology of plants, insect pollinators, and their interactions. Biodiversity Science, 2021, 29(4): 495-506. doi: 10.17520/biods.2020196
[25] 杨月娟, 张灏, 周华坤, 叶鑫, 姚步青, 张春辉, 马真, 赵新全. 青藏高原高寒草甸花期物候和群落结构对氮、磷、钾添加的短期响应. 草业学报, 2015, 24(8): 35-43. YANG Y J, ZHANG H, ZHOU H K, YE X, YAO B Q, ZHANG C H, MA Z, ZHAO X Q. Short-term responses of flowering phenology and community structure to nitrogen, phosphorus and potassium in an alpine meadow on the Qinghai-Tibetan Plateau. Acta Prataculturae Sinica, 2015, 24(8): 35-43.
[26] 巴雅尔塔, 贾鹏, 杨晓, 杜国祯. 青藏高原高寒草甸组分种花期物候对施肥响应. 草业学报, 2010, 19(3): 233-239. Bayaerta, JIA P, YANG X, DU G Z. Response of dominating species flowering phenology to fertilization in Qinghai-Tibetan alpine meadow. Acta Prataculturae Sinica, 2010, 19(3): 233-239.
[27] XI Y, ZHANG T, ZHANG Y J, ZHU J T, ZHANG G L, JIANG Y B. Nitrogen addition alters the phenology of a dominant alpine plant in northern Tibet. Arctic Antarctic and Alpine Research, 2015, 47(3): 511-518. doi: 10.1657/AAAR0014-054
[28] 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. doi: 10.1093/aob/mct079
[29] 叶琼丹, 任飞, 李永慧, 张子杨, 樊娜, 冶俊, 李希来, 李兰平. 不同养分添加对高寒植物功能性状的影响. 草地学报, 2022, 30(10): 2737-2744. YE Q D, REN F, LI Y H, ZHANG Z Y, FAN N, YE J, LI X L, LI L P. Effects of different nutrient additions on functional traits of alpine plant. Acta Agrestia Sinica, 2022, 30(10): 2737-2744.
[30] LIU Y Z, LI G Y, WU X W, NIKLAS K J, YANG Z L, SUN S C. Linkage between species traits and plant phenology in an alpine meadow. Oecologia, 2021, 195: 409-419. doi: 10.1007/s00442-020-04846-y
[31] 李向前, 贾鹏, 章志龙, 杜国祯. 青藏高原东缘高寒草甸植物群落的开花物候. 生态学杂志, 2009, 28(11): 2202-2207. LI X Q, JIA P, ZHANG Z L, DU G Z. Flowering phenology of alpine meadow plant community in eastern Qinghai Tibetan Plateau. Chinese Journal of Ecology, 2009, 28(11): 2202-2207.
[32] GIMENEZ-BENAVIDES L, GARCIA-CAMACHO R, IRIONDO J M, ESCUDERO A. Selection on flowering time in Mediterranean high-mountain plants under global warming. Evolutionary Ecology, 2011, 25(4): 777-794. doi: 10.1007/s10682-010-9440-z
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图 1 高寒植物花期物候对氮、磷养分添加的整体响应
不同小写字母表示不同处理间差异显著(P < 0. 05);CK,对照;N1,添加氮10 g·m−2;N2,添加氮20 g·m−2;P1,磷加氮5 g·m−2;P2,磷加氮10 g·m−2。下图同。
Figure 1. Overall response of flowering phenology of alpine plant to nitrogen and phosphorus additions
Different lowercase letters in the figure indicate a significant differences between different treatments at the 0.05 level. N1, nitrogen additions 10 g·m−2; N2, nitrogen additions 20 g·m−2; P1, phosphorus additions 5 g·m−2; P1, phosphorus additions 10 g·m−2. This is applicable for the following figures as well.
图 2 12种高寒植物始花期对不同氮、磷养分添加的响应
横坐标中正值表示与对照相比,始花期推迟,负值表示与对照相比,始花期提前。下图同。
Figure 2. Response of the first flowering date of 12 alpine species to different nitrogen and phosphorus additions
The positive value in the abscissa represents a delayed first flowering date, and the negative value represents an advanced first flowering date. This is applicable for the following figures as well.
表 1 氮、磷养分添加对高寒植物花期物候的整体影响双因素方差分析结果
Table 1 Result of the overall influence of nitrogen and phosphorus addition on flowering phenology of alpine plant based on Two-way ANOVA
处理
Treatment自由度
df始花期
First flowering date终花期
End flowering day花期持续时间
Flowering durationF P F P F P 氮添加(N) Nitrogen treatment 2 0.252 > 0.05 0.608 > 0.05 3.937 < 0.05 磷添加(P) Phosphorus treatment 2 0.417 > 0.05 0.648 > 0.05 9.966 < 0.01 N × P 4 0.006 > 0.05 0.193 > 0.05 1.044 > 0.05 -
[1] SCHWARTZ M D. Phenology: An Integrative Environmental Science. Dordrecht: Springer Netherlands, 2013.
[2] 李兰平. 青藏高原东部高寒草甸植物开花物候研究. 兰州: 兰州大学博士学位论文, 2017. LI L P. The study of flowering phenology of alpine plant on eastern Tibetan Plateau. PhD Thesis. Lanzhou: Lanzhou University, 2017.
[3] LI L P, LI Z K, CADOTTE W M, JIA P, CHEN G G, JIN S L, DU G Z. Phylogenetic conservatism and climate factors shape flowering phenology in alpine meadows. Oecologia, 2016, 182(2): 419-428. doi: 10.1007/s00442-016-3666-6
[4] WALTHER G R, POST E, CONVEY P, MENZEL A, PARMESAN C, BEEBEE T J C, FROMENTIN J M, HOEGH-GULDBER O, BAIRLEIN F. Ecological responses to recent climate change. Nature, 2002, 416: 389-395. doi: 10.1038/416389a
[5] CLELAND E E, CHUINE I, MENZEL A, MOONEY A H, SCHWARTZ D M. Shifting plant phenology in response to global change. Trends in Ecology and Evolution, 2007, 22(7): 357-365. doi: 10.1016/j.tree.2007.04.003
[6] 代武君, 金慧颖, 张玉红, 周志强, 刘彤. 植物物候学研究进展. 生态学报, 2020, 40(19): 6705-6719. DAI W J, JIN H Y, ZHANG Y H, ZHOU Z Q, LIU T. Advances in plant phenology. Acta Ecologica Sinica, 2020, 40(19): 6705-6719.
[7] STERNER R W, ELSER J J. Ecological Stoichiometry: The Biology of Elements from Molecules to the Biosphere. Princeton: Princeton University Press, 2002.
[8] MARKLEIN A R, HOULTON B Z. Nitrogen inputs accelerate phosphorus cycling rates across a wide variety of terrestrial ecosystems. New Phytologist, 2012, 193(3): 696-704. doi: 10.1111/j.1469-8137.2011.03967.x
[9] PAN Y P, LIU B W, CAO J, LIU J, TIAN S L, DU E Z. Enhanced atmospheric phosphorus deposition in Asia and Europe in the past two decades. Atmospheric and Oceanic Science Letters, 2021, 14(5): 100051.
[10] JIANG J, WANG Y P, YANG Y H, YU M X, WANG C, YAN J H. Interactive effects of nitrogen and phosphorus additions on plant growth vary with ecosystem type. Plant and Soil, 2019, 440(1): 523-537.
[11] WANG C, TANG Y J. Responses of plant phenology to nitrogen addition: A Meta-analysis. Oikos, 2019, 128(9): 1243-1253. doi: 10.1111/oik.06099
[12] 章志龙, 刘旭东, 贾鹏, 杜国祯. 氮添加对高寒草甸植物花期物候和群落结构的影响. 草业科学, 2013, 30(5): 728-735. ZHANG Z L, LIU X D, JIA P, DU G Z. Effects of nitrogen addition on flowering phenology, functional traits and community structure of alpine meadow in the eastern Qinghai-Tibet Plateau. Pratacultural Science, 2013, 30(5): 728-735.
[13] SHERRY R A, ZHOU X H, GU S L, ARNONE J A, SCHIMEL D S, VERBURG P S, WALLACE L L, LUO Y Q. Divergence of reproductive phenology under climate warming. Proceedings of the National Academy of Science, 2007, 104(1): 198-202. doi: 10.1073/pnas.0605642104
[14] 刘旭东, 章志龙, 杜国祯. 高寒草甸主要组分种开花物候对氮素添加的响应. 草业科学, 2021, 38(7): 1240-1249. LIU X D, ZHANG Z L, DU G Z. Response of dominant and common species flowering phenology to nitrogen addition in an alpine meadow. Pratacultural Science, 2021, 38(7): 1240-1249.
[15] SMITH J G, SCONIERS W, SPASOJEVIC M J, ASHTON W A, SUDING K N. Phenological changes in alpine plants in response to increased snowpack, temperature, and nitrogen. Arctic, Antarctic, and Alpine Research, 2012, 44(1): 135-142. doi: 10.1657/1938-4246-44.1.135
[16] LIU Y Z, MIAO R H, CHEN A Q, MIAO Y, LIU Y J, WU X W. Effects of nitrogen addition and mowing on reproductive phenology of three early-flowering forb species in a Tibetan alpine meadow. Ecological Engineering, 2017, 99: 119-125. doi: 10.1016/j.ecoleng.2016.11.033
[17] 杜国祯, 赵松岭. 草本群落中种多样性维持机制: 花期不同步性. 草业学报, 1997, 6(3): 1-7. DU G Z, ZHAO S L. Mechanism of diversity maintenance in herbaceous plant community: Asynchronism in flowering time. Acta Prataculturae Sinica, 1997, 6(3): 1-7.
[18] FORREST J, MILLER-RUSHING A J. Toward a synthetic understanding of the role of phenology in ecology and evolution. Philosophical Transactions of the Royal Society B: Biological Sciences, 2010, 365(1555): 3101-3112. doi: 10.1098/rstb.2010.0145
[19] CLARK C M, CLELAND E E, COLLINS S L, FARGIONE J E, GOUGH L, GROSS K L, PENNINGS S C, SUDING K N, GRACE J B. Environmental and plant community determinants of species loss following nitrogen enrichment. Ecology Letters, 2017, 10(7): 596-607.
[20] 周小龙. 高寒草甸植物群落结构组建和生产力对施肥的响应机制. 兰州: 兰州大学博士学位论文, 2016. ZHOU X L. The effect of fertilization on community assembly and production in alpine meadow community. PhD Thesis. Lanzhou: Lanzhou University, 2016.
[21] 李元恒. 内蒙古典型草原植物生殖物候对气候变化和人为干扰的响应. 兰州: 甘肃农业大学博士学位论文, 2008. LI Y H. Responses of reproductive phenology of Inner Mongolia typical steppe plants under climatic change and artificial interference. PhD Thesis. Lanzhou: Gansu Agricultural University, 2008.
[22] CLELAND E E, CHIARIELLO N R, LOARIE S R, MOONEY H A, FIELD C B. Diverse responses of phenology to global changes in a grassland ecosystem. Proceedings of the National Academy of Sciences, 2006, 103(37): 13740-13744. doi: 10.1073/pnas.0600815103
[23] BEARD K H, KELSEY K C, LEFFLER A J, WELKER J M. The missing angle: Ecosystem consequences of phenological mismatch. Trends in Ecology and Evolution, 2019, 34(10): 885-888. doi: 10.1016/j.tree.2019.07.019
[24] 施雨含, 任宗昕, 赵延会, 王红. 气候变化对植物-传粉昆虫的分布区和物候及其互作关系的影响. 生物多样性, 2021, 29(4): 495-506. doi: 10.17520/biods.2020196 SHI Y H, REN Z X, ZHAO Y H, WANG H. Effect of climate change on the distribution and phenology of plants, insect pollinators, and their interactions. Biodiversity Science, 2021, 29(4): 495-506. doi: 10.17520/biods.2020196
[25] 杨月娟, 张灏, 周华坤, 叶鑫, 姚步青, 张春辉, 马真, 赵新全. 青藏高原高寒草甸花期物候和群落结构对氮、磷、钾添加的短期响应. 草业学报, 2015, 24(8): 35-43. YANG Y J, ZHANG H, ZHOU H K, YE X, YAO B Q, ZHANG C H, MA Z, ZHAO X Q. Short-term responses of flowering phenology and community structure to nitrogen, phosphorus and potassium in an alpine meadow on the Qinghai-Tibetan Plateau. Acta Prataculturae Sinica, 2015, 24(8): 35-43.
[26] 巴雅尔塔, 贾鹏, 杨晓, 杜国祯. 青藏高原高寒草甸组分种花期物候对施肥响应. 草业学报, 2010, 19(3): 233-239. Bayaerta, JIA P, YANG X, DU G Z. Response of dominating species flowering phenology to fertilization in Qinghai-Tibetan alpine meadow. Acta Prataculturae Sinica, 2010, 19(3): 233-239.
[27] XI Y, ZHANG T, ZHANG Y J, ZHU J T, ZHANG G L, JIANG Y B. Nitrogen addition alters the phenology of a dominant alpine plant in northern Tibet. Arctic Antarctic and Alpine Research, 2015, 47(3): 511-518. doi: 10.1657/AAAR0014-054
[28] 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. doi: 10.1093/aob/mct079
[29] 叶琼丹, 任飞, 李永慧, 张子杨, 樊娜, 冶俊, 李希来, 李兰平. 不同养分添加对高寒植物功能性状的影响. 草地学报, 2022, 30(10): 2737-2744. YE Q D, REN F, LI Y H, ZHANG Z Y, FAN N, YE J, LI X L, LI L P. Effects of different nutrient additions on functional traits of alpine plant. Acta Agrestia Sinica, 2022, 30(10): 2737-2744.
[30] LIU Y Z, LI G Y, WU X W, NIKLAS K J, YANG Z L, SUN S C. Linkage between species traits and plant phenology in an alpine meadow. Oecologia, 2021, 195: 409-419. doi: 10.1007/s00442-020-04846-y
[31] 李向前, 贾鹏, 章志龙, 杜国祯. 青藏高原东缘高寒草甸植物群落的开花物候. 生态学杂志, 2009, 28(11): 2202-2207. LI X Q, JIA P, ZHANG Z L, DU G Z. Flowering phenology of alpine meadow plant community in eastern Qinghai Tibetan Plateau. Chinese Journal of Ecology, 2009, 28(11): 2202-2207.
[32] GIMENEZ-BENAVIDES L, GARCIA-CAMACHO R, IRIONDO J M, ESCUDERO A. Selection on flowering time in Mediterranean high-mountain plants under global warming. Evolutionary Ecology, 2011, 25(4): 777-794. doi: 10.1007/s10682-010-9440-z