暖温带森林木本植物展叶始期对低温和热量累积变化的响应

doi:10.17521/cjpe.2022.0036

摘要/Abstract

摘要：

为了解气候急剧变暖造成的冬春季节的低温和热量积累变化对自然环境中植物春季物候产生的影响, 利用北京东灵山暖温带森林25种木本植物2003-2019年的展叶始期数据, 采用偏最小二乘回归、动力学模型和生长度小时模型等方法模拟了各物种展叶始期所需的低温和热量累积量, 并应用线性回归分析了展叶始期对低温和热量累积变化的响应, 利用单因素方差分析对比了灌木和乔木展叶始期及其对低温和热量累积变化响应的差异。结果显示: (1) 25种木本植物展叶始期的平均低温和热量累积期分别在10月6日至次年3月17日和1月21日至4月26日之间, 平均低温和热量累积量分别为66.16冷激份额(CP)和2 933.12生长度小时(GDH)。(2)展叶始期对低温和热量累积变化的响应敏感度均值分别为每10 CP延迟3.54 d和每1 000 GDH延迟7.09 d, 各有2个和23个物种显著, 说明暖温带木本植物展叶始期主要受热量累积的影响。(3)灌木的展叶始期比乔木早3.87 d, 热量累积比乔木少543.56 GDH, 且展叶始期越早的植物, 所需热量累积也越少, 可能与其采取机会主义生存策略有关。(4)灌木和乔木展叶始期对热量累积的响应敏感度分别为每1 000 GDH延迟8.10和延迟6.13 d, 两者的差异呈边缘显著。这意味着随着气候变暖, 灌木展叶始期提前的速度可能比乔木更快。

关键词: 植物物候, 展叶始期, 气候变暖, 低温累积, 热量累积, 暖温带森林, 木本植物

Abstract:

Aims In recent decades, the rapid climate warming had affected chilling and heat accumulation during winter and spring and made profound changes in plant spring phenology. To date, most related studies focused on either a range of species grown in various gardens or on experimental research, which may be not necessarily applicable to real-world conditions.

Methods By using leaf-unfolding data of 25 woody species during 2003-2019 in warm temperate forests of the Dongling Mountain, Beijing, we simulated the daily chilling and heat accumulation by applying partial least square regression, dynamic model and growing degree hour model. We then analyzed the response of leaf-unfolding dates to the variation of chilling and heat accumulation by linear regressions. Finally, the differences of leaf-unfolding dates and their responses to the variation of chilling and heat accumulation between shrubs and trees were compared by ANOVA.

Important findings The chilling periods of 25 woody species were from October 6 to March 17 of next year, with the forcing periods from January 21 to April 26. The corresponding chilling and heat accumulation were 66.16 chill portion (CP) and 2 933.12 growing degree hour (GDH) on average. The leaf-unfolding dates were delayed 3.54 d per 10 CP and 7.09 d per 1 000 GDH as the chilling and heat accumulation changed, with 2 and 23 species significant, respectively. This indicated that leaf-unfolding dates of woody species in warm temperate zone were mainly affected by heat accumulation. Moreover, the leaf-unfolding dates of shrubs were 3.87 d earlier and required 543.56 GDH less heat than trees. Species leafed earlier required less heat accumulation than those leafed out later, presumably due to the opportunistic strategy adopted by shrubs and early-leafing species. Sensitivity of leaf-unfolding dates of shrubs to heat accumulation (delayed 8.10 d per 1 000 GDH) existed marginally significant difference with trees (delayed 6.13 d per 1 000 GDH), which implied that leaf-unfolding dates of shrubs might advance faster than trees as global warming progresses.

Key words: plant phenology, leaf-unfolding date, climate warming, chilling accumulation, heat accumulation, warm temperate forests, woody species

于海英, 杨莉琳, 付素静, 张志敏, 姚琦馥. 暖温带森林木本植物展叶始期对低温和热量累积变化的响应. 植物生态学报, 2022, 46(12): 1573-1584. DOI: 10.17521/cjpe.2022.0036

YU Hai-Ying, YANG Li-Lin, FU Su-Jing, ZHANG Zhi-Min, YAO Qi-Fu. Response of leaf-unfolding dates of woody species to variation of chilling and heat accumulation in warm temperate forests. Chinese Journal of Plant Ecology, 2022, 46(12): 1573-1584. DOI: 10.17521/cjpe.2022.0036

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2022.0036

https://www.plant-ecology.com/CN/Y2022/V46/I12/1573

图/表 7

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物种 Species	生活型 Life form	平均展叶始期 Mean leaf-unfolding date (month-day)	最早展叶始期 The earliest leaf-unfolding date (month-day)	最晚展叶始期 The latest leaf-unfolding date (month-day)	观测年数 Number of observed years
白桦 Betula platyphylla	乔木 Tree	05-04	04-27	05-17	15
大花溲疏 Deutzia grandiflora	灌木 Shrub	04-28	04-11	05-17	15
蒿柳 Salix schwerinii	灌木 Shrub	04-23	04-10	05-10	15
黑桦 Betula dahurica	乔木 Tree	05-04	04-25	05-13	11
胡桃楸 Juglans mandshurica	乔木 Tree	05-02	04-20	05-15	11
花曲柳 Fraxinus chinensis subsp. rhynchophylla	乔木 Tree	05-02	04-21	05-18	11
华北落叶松 Larix gmelinii var. principis-rupprechtii	乔木 Tree	04-22	04-09	05-08	11
金花忍冬 Lonicera chrysantha	灌木 Shrub	04-25	04-10	05-11	14
辽东栎 Quercus wutaishansea	乔木 Tree	05-03	04-20	05-15	16
裂叶榆 Ulmus laciniata	乔木 Tree	04-27	04-15	05-09	15
六道木 Zabelia biflora	灌木 Shrub	04-27	04-13	05-09	16
毛榛 Corylus mandshurica	灌木 Shrub	04-27	04-20	05-05	14
蒙椴 Tilia mongolica	乔木 Tree	04-30	04-17	05-08	15
巧玲花 Syringa pubescens	灌木 Shrub	04-19	04-03	05-06	11
青杨 Populus cathayana	乔木 Tree	04-27	04-12	05-07	15
山桃 Prunus davidiana	乔木 Tree	04-21	04-06	05-14	16
山杏 Prunus sibirica	乔木 Tree	04-26	04-10	05-10	16
土庄绣线菊 Spiraea pubescens	灌木 Shrub	04-28	04-18	05-14	11
卫矛 Euonymus alatus	灌木 Shrub	04-22	04-04	05-11	14
五角枫 Acer pictum subsp. mono	乔木 Tree	04-27	04-16	05-08	14
小花溲疏 Deutzia parviflora	灌木 Shrub	04-29	04-11	05-17	16
小叶鼠李 Rhamnus parvifolia	灌木 Shrub	04-21	04-09	05-15	15
迎红杜鹃 Rhododendron mucronulatum	灌木 Shrub	04-27	04-15	05-11	15
油松 Pinus tabuliformis	乔木 Tree	05-09	04-22	05-30	15
照山白 Rhododendron micranthum	灌木 Shrub	05-02	04-15	05-22	15

物种 Species	生活型 Life form	平均展叶始期 Mean leaf-unfolding date (month-day)	最早展叶始期 The earliest leaf-unfolding date (month-day)	最晚展叶始期 The latest leaf-unfolding date (month-day)	观测年数 Number of observed years
白桦 Betula platyphylla	乔木 Tree	05-04	04-27	05-17	15
大花溲疏 Deutzia grandiflora	灌木 Shrub	04-28	04-11	05-17	15
蒿柳 Salix schwerinii	灌木 Shrub	04-23	04-10	05-10	15
黑桦 Betula dahurica	乔木 Tree	05-04	04-25	05-13	11
胡桃楸 Juglans mandshurica	乔木 Tree	05-02	04-20	05-15	11
花曲柳 Fraxinus chinensis subsp. rhynchophylla	乔木 Tree	05-02	04-21	05-18	11
华北落叶松 Larix gmelinii var. principis-rupprechtii	乔木 Tree	04-22	04-09	05-08	11
金花忍冬 Lonicera chrysantha	灌木 Shrub	04-25	04-10	05-11	14
辽东栎 Quercus wutaishansea	乔木 Tree	05-03	04-20	05-15	16
裂叶榆 Ulmus laciniata	乔木 Tree	04-27	04-15	05-09	15
六道木 Zabelia biflora	灌木 Shrub	04-27	04-13	05-09	16
毛榛 Corylus mandshurica	灌木 Shrub	04-27	04-20	05-05	14
蒙椴 Tilia mongolica	乔木 Tree	04-30	04-17	05-08	15
巧玲花 Syringa pubescens	灌木 Shrub	04-19	04-03	05-06	11
青杨 Populus cathayana	乔木 Tree	04-27	04-12	05-07	15
山桃 Prunus davidiana	乔木 Tree	04-21	04-06	05-14	16
山杏 Prunus sibirica	乔木 Tree	04-26	04-10	05-10	16
土庄绣线菊 Spiraea pubescens	灌木 Shrub	04-28	04-18	05-14	11
卫矛 Euonymus alatus	灌木 Shrub	04-22	04-04	05-11	14
五角枫 Acer pictum subsp. mono	乔木 Tree	04-27	04-16	05-08	14
小花溲疏 Deutzia parviflora	灌木 Shrub	04-29	04-11	05-17	16
小叶鼠李 Rhamnus parvifolia	灌木 Shrub	04-21	04-09	05-15	15
迎红杜鹃 Rhododendron mucronulatum	灌木 Shrub	04-27	04-15	05-11	15
油松 Pinus tabuliformis	乔木 Tree	05-09	04-22	05-30	15
照山白 Rhododendron micranthum	灌木 Shrub	05-02	04-15	05-22	15

物种 Species	低温累积期 Chilling periods		低温累积量 Chilling accumulation (CP)	热量累积期 Forcing periods		热量累积量 Heat accumulation (GDH)
物种 Species	开始日期 Start date (month-day)	结束日期 End date (month-day)	低温累积量 Chilling accumulation (CP)	开始日期 Start date (month-day)	结束日期 End date (month-day)	热量累积量 Heat accumulation (GDH)
白桦 Betula platyphylla	09-30	01-11	43.37 ± 5.79	03-22	05-03	3 640.58 ± 963.76
大花溲疏 Deutzia grandiflora	09-30	03-23	73.26 ± 6.64	01-04	04-27	2 924.49 ± 824.72
蒿柳 Salix schwerinii	10-06	03-07	60.57 ± 6.14	03-03	04-22	2 304.28 ± 699.59
黑桦 Betula dahurica	10-31	03-23	55.55 ± 7.03	01-26	05-03	3 887.06 ± 1 105.56
胡桃楸 Juglans mandshurica	09-29	03-23	74.46 ± 7.52	12-19	05-01	3 562.93 ± 1 066.70
花曲柳 Fraxinus chinensis subsp. rhynchophylla	10-04	03-20	70.33 ± 6.99	01-04	05-01	3 562.62 ± 1 066.71
华北落叶松 Larix gmelinii var. principis-rupprechtii	10-26	03-07	48.92 ± 6.77	01-26	04-21	2 279.80 ± 775.96
金花忍冬 Lonicera chrysantha	09-30	03-23	73.28 ± 6.84	12-22	04-24	2 566.87 ± 783.05
辽东栎 Quercus wutaishansea	10-26	03-23	58.23 ± 6.29	01-26	05-02	3 669.40 ± 928.35
裂叶榆 Ulmus laciniata	09-29	03-23	73.62 ± 6.74	01-26	04-26	2 799.27 ± 797.50
六道木 Zabelia biflora	09-30	03-23	73.07 ± 6.47	12-13	04-26	2 750.16 ± 796.11
毛榛 Corylus mandshurica	09-28	03-02	61.20 ± 6.76	12-22	04-26	2 808.29 ± 931.77
蒙椴 Tilia mongolica	10-26	03-23	58.52 ± 5.89	01-23	04-29	3 200.84 ± 884.28
巧玲花 Syringa pubescens	09-02	03-24	79.72 ± 7.54	01-26	04-17	1 949.79 ± 711.70
青杨 Populus cathayana	12-05	03-24	36.79 ± 5.91	03-02	04-26	2 777.00 ± 512.17
山桃 Prunus davidiana	09-02	03-23	78.39 ± 6.43	03-20	04-20	1 879.42 ± 652.17
山杏 Prunus sibirica	09-30	03-22	72.40 ± 6.47	12-22	04-25	2 626.39 ± 773.89
土庄绣线菊 Spiraea pubescens	10-05	03-23	71.86 ± 6.97	12-17	04-27	2 995.08 ± 936.94
卫矛 Euonymus alatus	09-30	03-23	73.28 ± 6.84	03-19	04-21	2 059.65 ± 709.69
五角枫 Acer pictum subsp. mono	09-30	03-22	72.60 ± 6.84	01-22	04-26	2 808.06 ± 831.75
小花溲疏 Deutzia parviflora	09-29	03-23	73.44 ± 6.56	12-22	04-28	3 012.81 ± 844.57
小叶鼠李 Rhamnus parvifolia	09-30	03-23	73.26 ± 6.64	01-22	04-20	2 106.40 ± 666.57
迎红杜鹃 Rhododendron mucronulatum	09-29	03-23	73.62 ± 6.74	12-22	04-26	2 799.54 ± 797.67
油松 Pinus tabuliformis	10-22	03-08	51.62 ± 6.26	03-03	05-08	4 829.02 ± 1 002.10
照山白 Rhododendron micranthum	09-30	03-22	72.60 ± 6.63	12-22	05-01	3 528.25 ± 932.79
均值 Average	10-06	03-17	66.16 ± 11.26	01-21	04-26	2 933.12 ± 677.50

物种 Species	低温累积期 Chilling periods		低温累积量 Chilling accumulation (CP)	热量累积期 Forcing periods		热量累积量 Heat accumulation (GDH)
物种 Species	开始日期 Start date (month-day)	结束日期 End date (month-day)	低温累积量 Chilling accumulation (CP)	开始日期 Start date (month-day)	结束日期 End date (month-day)	热量累积量 Heat accumulation (GDH)
白桦 Betula platyphylla	09-30	01-11	43.37 ± 5.79	03-22	05-03	3 640.58 ± 963.76
大花溲疏 Deutzia grandiflora	09-30	03-23	73.26 ± 6.64	01-04	04-27	2 924.49 ± 824.72
蒿柳 Salix schwerinii	10-06	03-07	60.57 ± 6.14	03-03	04-22	2 304.28 ± 699.59
黑桦 Betula dahurica	10-31	03-23	55.55 ± 7.03	01-26	05-03	3 887.06 ± 1 105.56
胡桃楸 Juglans mandshurica	09-29	03-23	74.46 ± 7.52	12-19	05-01	3 562.93 ± 1 066.70
花曲柳 Fraxinus chinensis subsp. rhynchophylla	10-04	03-20	70.33 ± 6.99	01-04	05-01	3 562.62 ± 1 066.71
华北落叶松 Larix gmelinii var. principis-rupprechtii	10-26	03-07	48.92 ± 6.77	01-26	04-21	2 279.80 ± 775.96
金花忍冬 Lonicera chrysantha	09-30	03-23	73.28 ± 6.84	12-22	04-24	2 566.87 ± 783.05
辽东栎 Quercus wutaishansea	10-26	03-23	58.23 ± 6.29	01-26	05-02	3 669.40 ± 928.35
裂叶榆 Ulmus laciniata	09-29	03-23	73.62 ± 6.74	01-26	04-26	2 799.27 ± 797.50
六道木 Zabelia biflora	09-30	03-23	73.07 ± 6.47	12-13	04-26	2 750.16 ± 796.11
毛榛 Corylus mandshurica	09-28	03-02	61.20 ± 6.76	12-22	04-26	2 808.29 ± 931.77
蒙椴 Tilia mongolica	10-26	03-23	58.52 ± 5.89	01-23	04-29	3 200.84 ± 884.28
巧玲花 Syringa pubescens	09-02	03-24	79.72 ± 7.54	01-26	04-17	1 949.79 ± 711.70
青杨 Populus cathayana	12-05	03-24	36.79 ± 5.91	03-02	04-26	2 777.00 ± 512.17
山桃 Prunus davidiana	09-02	03-23	78.39 ± 6.43	03-20	04-20	1 879.42 ± 652.17
山杏 Prunus sibirica	09-30	03-22	72.40 ± 6.47	12-22	04-25	2 626.39 ± 773.89
土庄绣线菊 Spiraea pubescens	10-05	03-23	71.86 ± 6.97	12-17	04-27	2 995.08 ± 936.94
卫矛 Euonymus alatus	09-30	03-23	73.28 ± 6.84	03-19	04-21	2 059.65 ± 709.69
五角枫 Acer pictum subsp. mono	09-30	03-22	72.60 ± 6.84	01-22	04-26	2 808.06 ± 831.75
小花溲疏 Deutzia parviflora	09-29	03-23	73.44 ± 6.56	12-22	04-28	3 012.81 ± 844.57
小叶鼠李 Rhamnus parvifolia	09-30	03-23	73.26 ± 6.64	01-22	04-20	2 106.40 ± 666.57
迎红杜鹃 Rhododendron mucronulatum	09-29	03-23	73.62 ± 6.74	12-22	04-26	2 799.54 ± 797.67
油松 Pinus tabuliformis	10-22	03-08	51.62 ± 6.26	03-03	05-08	4 829.02 ± 1 002.10
照山白 Rhododendron micranthum	09-30	03-22	72.60 ± 6.63	12-22	05-01	3 528.25 ± 932.79
均值 Average	10-06	03-17	66.16 ± 11.26	01-21	04-26	2 933.12 ± 677.50