植物生态学报 ›› 2021, Vol. 45 ›› Issue (8): 834-843.DOI: 10.17521/cjpe.2021.0100
杜军1, 王文1,2, 何志斌1,*(), 陈龙飞1, 蔺鹏飞1, 朱喜1, 田全彦1
收稿日期:
2021-03-19
修回日期:
2021-06-04
出版日期:
2021-08-20
发布日期:
2021-06-25
通讯作者:
何志斌
作者简介:
* hzbmail@lzb.ac.cn基金资助:
DU Jun1, WANG Wen1,2, HE Zhi-Bin1,*(), CHEN Long-Fei1, LIN Peng-Fei1, ZHU Xi1, TIAN Quan-Yan1
Received:
2021-03-19
Revised:
2021-06-04
Online:
2021-08-20
Published:
2021-06-25
Contact:
HE Zhi-Bin
Supported by:
摘要:
量化种群水平植物物候空间分异的遗传表达与表型可塑性, 对于合理预测气候变化对植物动态的影响具有重要实践意义。该研究以祁连山广泛分布的优势常绿针叶树种——青海云杉(Picea crassifolia)为研究对象, 通过对排露沟流域5个海拔带20个样地的156株样树进行连续两年的物候过程监测, 研究了物候表型的空间分异规律, 结合同质园移植实验, 解析了物候空间分异的遗传分化成分, 并借助岭回归和Lasso回归分析, 探讨了环境因子对非遗传分化部分(即表型可塑性)的调控作用。研究发现: 青海云杉表观物候期均随海拔升高而呈现推迟的趋势(每100 m 0.57-1.36 d), 这种空间分异体现了一定程度的种源效应(即遗传分化现象), 其所占比例平均为20.8%, 且年际之间基本维持在相对稳定的水平; 同时, 研究发现同质园内来自高海拔的种群的春季物候期要明显早于低海拔种群, 这可能与高海拔种源的芽分化需要较低的季前热量积累阈值有关; 物候格局与环境因子关系的分析结果显示, 除海拔影响外, 季前热量积累、季前降水量、冻害发生频率对于解释物候动态起到了关键的指示作用。
杜军, 王文, 何志斌, 陈龙飞, 蔺鹏飞, 朱喜, 田全彦. 祁连山青海云杉物候表型的空间分异及其内在机制. 植物生态学报, 2021, 45(8): 834-843. DOI: 10.17521/cjpe.2021.0100
DU Jun, WANG Wen, HE Zhi-Bin, CHEN Long-Fei, LIN Peng-Fei, ZHU Xi, TIAN Quan-Yan. Spatial variability of phenological phenotype of Picea crassifolia in Qilian Mountains and its internal mechanism. Chinese Journal of Plant Ecology, 2021, 45(8): 834-843. DOI: 10.17521/cjpe.2021.0100
图1 2018年阴坡样地(包括小林斑和大林斑)监测的青海云杉物候期随海拔梯度的变化(平均值±标准差)。DOY, 年序日。
Fig. 1 Variation of annual phenological stages of Picea crassifolia monitored in shady-slope plots (including small and large patches) in 2018 along the altitudinal gradient (mean ± SD). DOY, day of the year.
影响因子 Impact factor | df | 萌芽 Bud burst | 展叶 Leaf expansion | 抽枝 Branching | 开花 Flowering | 结果 Fruiting | ||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
MS | F | p | MS | F | p | MS | F | p | MS | F | p | MS | F | p | ||
海拔 Altitude (AL) | 4 | 155.23 | 38.70 | 0.00 | 216.80 | 82.50 | 0.00 | 239.76 | 72.91 | 0.00 | 63.27 | 4.44 | 0.00 | 849.72 | 744.96 | 0.00 |
坡向 Aspect (AS) | 1 | 40.19 | 10.02 | 0.00 | 39.62 | 15.08 | 0.00 | 24.09 | 7.33 | 0.01 | 16.92 | 1.19 | 0.28 | 13.14 | 11.52 | 0.00 |
斑块大小 Patch size (PS) | 1 | 0.95 | 0.24 | 0.63 | 1.83 | 0.70 | 0.41 | 1.65 | 0.50 | 0.48 | 0.07 | 0.01 | 0.94 | 0.02 | 0.01 | 0.91 |
AL × AS | 4 | 1.58 | 0.39 | 0.71 | 2.09 | 0.80 | 0.53 | 1.42 | 0.43 | 0.79 | 0.12 | 0.01 | 1.00 | 8.14 | 7.14 | 0.00 |
AL × PS | 4 | 0.39 | 0.10 | 0.98 | 0.72 | 0.28 | 0.89 | 0.61 | 0.19 | 0.95 | 0.40 | 0.03 | 1.00 | 2.35 | 2.06 | 0.12 |
AS × PS | 1 | 40.76 | 10.16 | 0.00 | 22.16 | 8.43 | 0.01 | 5.89 | 1.79 | 0.19 | 59.16 | 4.15 | 0.05 | 1.27 | 1.11 | 0.30 |
AL × AS × PS | 4 | 15.80 | 3.94 | 0.01 | 13.84 | 5.27 | 0.00 | 12.34 | 3.75 | 0.01 | 19.62 | 1.38 | 0.25 | 1.27 | 1.11 | 0.35 |
误差 Error | 60 | 4.01 | 2.63 | 3.29 | 14.26 | 1.14 |
表1 海拔、坡向及斑块大小对青海云杉种群物候空间分异的影响
Table 1 Effects of altitude, aspect and patch size on the spatial differentiation of Picea crassifolia population phenology
影响因子 Impact factor | df | 萌芽 Bud burst | 展叶 Leaf expansion | 抽枝 Branching | 开花 Flowering | 结果 Fruiting | ||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
MS | F | p | MS | F | p | MS | F | p | MS | F | p | MS | F | p | ||
海拔 Altitude (AL) | 4 | 155.23 | 38.70 | 0.00 | 216.80 | 82.50 | 0.00 | 239.76 | 72.91 | 0.00 | 63.27 | 4.44 | 0.00 | 849.72 | 744.96 | 0.00 |
坡向 Aspect (AS) | 1 | 40.19 | 10.02 | 0.00 | 39.62 | 15.08 | 0.00 | 24.09 | 7.33 | 0.01 | 16.92 | 1.19 | 0.28 | 13.14 | 11.52 | 0.00 |
斑块大小 Patch size (PS) | 1 | 0.95 | 0.24 | 0.63 | 1.83 | 0.70 | 0.41 | 1.65 | 0.50 | 0.48 | 0.07 | 0.01 | 0.94 | 0.02 | 0.01 | 0.91 |
AL × AS | 4 | 1.58 | 0.39 | 0.71 | 2.09 | 0.80 | 0.53 | 1.42 | 0.43 | 0.79 | 0.12 | 0.01 | 1.00 | 8.14 | 7.14 | 0.00 |
AL × PS | 4 | 0.39 | 0.10 | 0.98 | 0.72 | 0.28 | 0.89 | 0.61 | 0.19 | 0.95 | 0.40 | 0.03 | 1.00 | 2.35 | 2.06 | 0.12 |
AS × PS | 1 | 40.76 | 10.16 | 0.00 | 22.16 | 8.43 | 0.01 | 5.89 | 1.79 | 0.19 | 59.16 | 4.15 | 0.05 | 1.27 | 1.11 | 0.30 |
AL × AS × PS | 4 | 15.80 | 3.94 | 0.01 | 13.84 | 5.27 | 0.00 | 12.34 | 3.75 | 0.01 | 19.62 | 1.38 | 0.25 | 1.27 | 1.11 | 0.35 |
误差 Error | 60 | 4.01 | 2.63 | 3.29 | 14.26 | 1.14 |
图2 2018年同质园内青海云杉种群物候期与原海拔的关系(平均值±标准差)。DOY, 年序日。
Fig. 2 Phenological pattern of Picea crassifolia populations in the common garden experiment versus elevation of provenance sites in 2018 (mean ± SD). DOY, day of the year.
阶段 Phase | 2018 | 2019 | ||||
---|---|---|---|---|---|---|
D | F | p | D | F | p | |
萌芽期 Bud burst | 13.4 | 2.669 | 0.043 | 19.9 | 3.636 | 0.011 |
展叶期 Leaf expansion | 23.2 | 4.283 | 0.005 | 20.4 | 3.700 | 0.010 |
抽枝期 Branching | 29.2 | 5.512 | 0.001 | 19.3 | 3.495 | 0.013 |
表2 青海云杉种群各物候阶段遗传分化所占比例
Table 2 Genetic differentiation in phenological stages among populations of Picea crassifolia
阶段 Phase | 2018 | 2019 | ||||
---|---|---|---|---|---|---|
D | F | p | D | F | p | |
萌芽期 Bud burst | 13.4 | 2.669 | 0.043 | 19.9 | 3.636 | 0.011 |
展叶期 Leaf expansion | 23.2 | 4.283 | 0.005 | 20.4 | 3.700 | 0.010 |
抽枝期 Branching | 29.2 | 5.512 | 0.001 | 19.3 | 3.495 | 0.013 |
变量 Variable | 展叶 Leaf expansion (Scaled) | Lar序列 Lar sequence | 开花 Flowering (Scaled) | Lar序列 Lar sequence | 结果 Fruiting (Scaled) | Lar序列 Lar sequence |
---|---|---|---|---|---|---|
海拔 Altitude | 4.80 (0.72)*** | 2 | 2.34 (0.69) *** | 1 | 6.85 (0.23) *** | 5 |
CH0(Oct.-Feb.) | -4.07 (0.38) *** | 8 | -1.42 (0.51) ** | 7 | -7.00 (0.22) *** | 10 |
CH5(Oct.-Feb.) | -2.52 (0.31) *** | - | -1.09 (0.43) * | - | -4.69 (0.18) *** | - |
CDD0(Mar.-May)1 | -5.93 (0.63) *** | 5 | -1.90 (0.57) *** | 5 | -9.49 (0.29) *** | - |
CDD5(Mar.-May)1 | -6.83 (0.79) *** | 1 | -2.13 (0.65) ** | 2 | -10.65 (0.36) *** | 1 |
CDD0(Mar.-May)2 | 0.74 (0.88) | 6 | -0.90 (0.94) | 9 | 2.34 (0.45) *** | 7 |
CDD5(Mar.-May)2 | - | - | - | - | - | - |
TR(Mar.-May) | 1.01 (1.68) | 9 | 1.27 (1.41) | 6 | 4.27 (0.30) *** | 9 |
RD(Mar.-May) | 4.33 (1.27) *** | 3 | 0.84 (1.11) | 4 | 9.87 (0.80) *** | 2 |
FD(May) | 6.47 (0.83) *** | 4 | 2.13 (0.67) ** | 3 | 10.11 (0.41) *** | 3 |
HD(May) | 3.60 (1.29) ** | - | 0.32 (0.97) | - | 4.26 (0.62) *** | 8 |
PSD(Mar.-May) | 0.02 (1.15) | 7 | 0.93 (1.21) | 8 | -1.68 (0.51) ** | 6 |
FSD(May) | -4.78 (0.60) *** | 10 | -1.98 (0.64) ** | 10 | -6.50 (0.38) *** | 4 |
表3 青海云杉种群物候与环境要素的岭回归及Lasso回归分析结果
Table 3 Ridge regression and Lasso regression analysis of the phenology of Picea crassifolia populations and environmental factors
变量 Variable | 展叶 Leaf expansion (Scaled) | Lar序列 Lar sequence | 开花 Flowering (Scaled) | Lar序列 Lar sequence | 结果 Fruiting (Scaled) | Lar序列 Lar sequence |
---|---|---|---|---|---|---|
海拔 Altitude | 4.80 (0.72)*** | 2 | 2.34 (0.69) *** | 1 | 6.85 (0.23) *** | 5 |
CH0(Oct.-Feb.) | -4.07 (0.38) *** | 8 | -1.42 (0.51) ** | 7 | -7.00 (0.22) *** | 10 |
CH5(Oct.-Feb.) | -2.52 (0.31) *** | - | -1.09 (0.43) * | - | -4.69 (0.18) *** | - |
CDD0(Mar.-May)1 | -5.93 (0.63) *** | 5 | -1.90 (0.57) *** | 5 | -9.49 (0.29) *** | - |
CDD5(Mar.-May)1 | -6.83 (0.79) *** | 1 | -2.13 (0.65) ** | 2 | -10.65 (0.36) *** | 1 |
CDD0(Mar.-May)2 | 0.74 (0.88) | 6 | -0.90 (0.94) | 9 | 2.34 (0.45) *** | 7 |
CDD5(Mar.-May)2 | - | - | - | - | - | - |
TR(Mar.-May) | 1.01 (1.68) | 9 | 1.27 (1.41) | 6 | 4.27 (0.30) *** | 9 |
RD(Mar.-May) | 4.33 (1.27) *** | 3 | 0.84 (1.11) | 4 | 9.87 (0.80) *** | 2 |
FD(May) | 6.47 (0.83) *** | 4 | 2.13 (0.67) ** | 3 | 10.11 (0.41) *** | 3 |
HD(May) | 3.60 (1.29) ** | - | 0.32 (0.97) | - | 4.26 (0.62) *** | 8 |
PSD(Mar.-May) | 0.02 (1.15) | 7 | 0.93 (1.21) | 8 | -1.68 (0.51) ** | 6 |
FSD(May) | -4.78 (0.60) *** | 10 | -1.98 (0.64) ** | 10 | -6.50 (0.38) *** | 4 |
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