园艺学报 ›› 2022, Vol. 49 ›› Issue (4): 709-722.doi: 10.16420/j.issn.0513-353x.2020-1051
• 研究论文 • 下一篇
张晓云1, 唐玉薇1, 王凯1, 张东2,*(), 杨伟伟1,*(
)
收稿日期:
2021-11-05
修回日期:
2022-01-29
出版日期:
2022-04-25
发布日期:
2022-04-24
通讯作者:
张东,杨伟伟
E-mail:afant@nwsuaf.edu.cn;yangww@shzu.edu.cn
基金资助:
ZHANG Xiaoyun1, TANG Yuwei1, WANG Kai1, ZHANG Dong2,*(), YANG Weiwei1,*(
)
Received:
2021-11-05
Revised:
2022-01-29
Online:
2022-04-25
Published:
2022-04-24
Contact:
ZHANG Dong,YANG Weiwei
E-mail:afant@nwsuaf.edu.cn;yangww@shzu.edu.cn
摘要:
为了探究苹果冠层枝梢类型对光能截获效率和光合生产力的影响,以苹果短枝型品种‘礼泉短富’和长枝型品种‘丽嘎拉’为试材,利用三维数字化技术结合枝梢和叶片尺度异速生长关系建立枝梢及冠层尺度的三维虚拟植物模型,利用RATP功能结构模型结合虚拟试验系统比较枝梢类型对枝梢形态结构、光能截获效率及枝梢相对比例对冠层光能截获效率和光合生产力的影响。结果表明:枝梢尺度、三维虚拟枝梢对各类枝梢叶片数及叶面积的模拟可满足光能截获分析的要求,但枝梢尺度光能截获模拟会低估长枝梢(> 5 cm)的光能截获效率。双参数的Beta方程可准确模拟各类枝梢叶倾角分布,果台叶片属斜生叶,其他枝梢叶片属水平叶。果台副梢节间长度及平均叶倾角高于营养枝梢,使得其光能截获效率较营养枝梢高8% ~ 14%。‘礼泉短富’叶片聚集度高于‘丽嘎拉’,二者光能截获效率无差异,但后者晴天和阴天条件下的冠层净光合速率和光合量都高于前者。在枝梢数量一定的条件下,果台副梢比例的提高可改善冠层光能截获效率,并提高‘礼泉短富’冠层在阴天条件下的冠层净光合速率及光合量。综上,对单个枝梢,花芽抽生的新梢光能截获效率显著高于叶芽抽生的新梢;花芽比例提高可显著改善冠层光能截获效率,并提高短枝型品种在阴天条件下的光合生产力。
中图分类号:
张晓云, 唐玉薇, 王凯, 张东, 杨伟伟. 苹果冠层枝梢类型对光能截获效率和光合生产力影响的模拟分析[J]. 园艺学报, 2022, 49(4): 709-722.
ZHANG Xiaoyun, TANG Yuwei, WANG Kai, ZHANG Dong, YANG Weiwei. Simulation Analysis of Effects of Shoot Type Composition on Canopy Light Interception Efficiency and Photosynthetic Productivity in Apple Trees[J]. Acta Horticulturae Sinica, 2022, 49(4): 709-722.
变量 Variable | 异速生 长关系 Allometric relation- ship | 系数 Coeffi- cient | 枝梢类型和品种Shoot type and cultivar | |||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|
果台 Bourse | 果台短副梢 Bourse short shoot | 果台长副梢 Bourse long shoot | 营养短枝 Vegetative short shoot | 营养长枝 Vegetative long shoot | ||||||||
礼泉 短富 Liquan Fuji | 丽嘎拉Regal Gala | 礼泉 短富 Liquan Fuji | 丽嘎拉 Regal Gala | 礼泉 短富 Liquan Fuji | 丽嘎拉 Regal Gala | 礼泉 短富 Liquan Fuji | 丽嘎拉 Regal Gala | 礼泉 短富 Liquan Fuji | 丽嘎拉 Regal Gala | |||
枝梢叶数—枝梢长Shoot leaf number-shoot length | SLN = aSLNSL + bSLN | aSLN | 0 | 0 | 0 | 0 | 0.42 | 0.20 | 0 | 1.40 | 0.58 | 0.34 |
bSLN | 6.00 | 6.00 | 6.00 | 8.00 | 4.95 | 8.49 | 10.00 | 6.69 | 7.22 | 9.27 | ||
r2 | ns | ns | ns | ns | 0.86** | 0.72** | ns | 0.66** | 0.93** | 0.93** | ||
枝梢叶面积—枝梢长Shoot leaf area-shoot length | SLA = aSLASL + bSLA | aSLA | 0 | 0 | 0 | 12.13 | 10.92 | 9.63 | 23.14 | 37.01 | 15.08 | 13.38 |
bSLA | 56.83 | 96.71 | 111.50 | 149.80 | 122.90 | 149.00 | 134.00 | 122.00 | 151.60 | 214.20 | ||
r2 | ns | ns | ns | 0.55** | 0.78** | 0.75** | 0.43* | 0.49* | 0.90** | 0.87** | ||
叶柄长—叶片长Petiole length- leaf length | PL = aPLLL + bPL | aPL | 0.33 | 0.36 | 0.38 | 0.21 | 0.43 | 0.17 | 0.26 | 0.24 | 0.23 | 0.16 |
bPL | 0.28 | 0.27 | 0.29 | 2.06 | 0.78 | 2.01 | 1.62 | 1.88 | 1.45 | 2.29 | ||
r2 | 0.63** | 0.55* | 0.66** | 0.43* | 0.58** | 0.48* | 0.61** | 0.59** | 0.42* | 0.31* | ||
叶面积—叶长2 Leaf area- leaf length2 | LA = aLALL² | aLA | 0.53 | 0.48 | 0.41 | 0.35 | 0.43 | 0.35 | 0.38 | 0.31 | 0.40 | 0.35 |
r2 | 0.98** | 0.98** | 0.98** | 0.97** | 0.99** | 0.97** | 0.97** | 0.97** | 0.97** | 0.98** | ||
叶宽—叶长 Leaf width- leaf length | LW = aLW LL | aLW | 0.76 | 0.69 | 0.61 | 0.52 | 0.62 | 0.54 | 0.58 | 0.50 | 0.58 | 0.53 |
r2 | 0.99** | 0.99** | 0.98** | 0.98** | 0.99** | 0.98** | 0.97** | 0.97** | 0.97** | 0.98** |
表1 枝梢和叶片异速生长关系
Table 1 Allometric relationships at shoot- and leaf-scales
变量 Variable | 异速生 长关系 Allometric relation- ship | 系数 Coeffi- cient | 枝梢类型和品种Shoot type and cultivar | |||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|
果台 Bourse | 果台短副梢 Bourse short shoot | 果台长副梢 Bourse long shoot | 营养短枝 Vegetative short shoot | 营养长枝 Vegetative long shoot | ||||||||
礼泉 短富 Liquan Fuji | 丽嘎拉Regal Gala | 礼泉 短富 Liquan Fuji | 丽嘎拉 Regal Gala | 礼泉 短富 Liquan Fuji | 丽嘎拉 Regal Gala | 礼泉 短富 Liquan Fuji | 丽嘎拉 Regal Gala | 礼泉 短富 Liquan Fuji | 丽嘎拉 Regal Gala | |||
枝梢叶数—枝梢长Shoot leaf number-shoot length | SLN = aSLNSL + bSLN | aSLN | 0 | 0 | 0 | 0 | 0.42 | 0.20 | 0 | 1.40 | 0.58 | 0.34 |
bSLN | 6.00 | 6.00 | 6.00 | 8.00 | 4.95 | 8.49 | 10.00 | 6.69 | 7.22 | 9.27 | ||
r2 | ns | ns | ns | ns | 0.86** | 0.72** | ns | 0.66** | 0.93** | 0.93** | ||
枝梢叶面积—枝梢长Shoot leaf area-shoot length | SLA = aSLASL + bSLA | aSLA | 0 | 0 | 0 | 12.13 | 10.92 | 9.63 | 23.14 | 37.01 | 15.08 | 13.38 |
bSLA | 56.83 | 96.71 | 111.50 | 149.80 | 122.90 | 149.00 | 134.00 | 122.00 | 151.60 | 214.20 | ||
r2 | ns | ns | ns | 0.55** | 0.78** | 0.75** | 0.43* | 0.49* | 0.90** | 0.87** | ||
叶柄长—叶片长Petiole length- leaf length | PL = aPLLL + bPL | aPL | 0.33 | 0.36 | 0.38 | 0.21 | 0.43 | 0.17 | 0.26 | 0.24 | 0.23 | 0.16 |
bPL | 0.28 | 0.27 | 0.29 | 2.06 | 0.78 | 2.01 | 1.62 | 1.88 | 1.45 | 2.29 | ||
r2 | 0.63** | 0.55* | 0.66** | 0.43* | 0.58** | 0.48* | 0.61** | 0.59** | 0.42* | 0.31* | ||
叶面积—叶长2 Leaf area- leaf length2 | LA = aLALL² | aLA | 0.53 | 0.48 | 0.41 | 0.35 | 0.43 | 0.35 | 0.38 | 0.31 | 0.40 | 0.35 |
r2 | 0.98** | 0.98** | 0.98** | 0.97** | 0.99** | 0.97** | 0.97** | 0.97** | 0.97** | 0.98** | ||
叶宽—叶长 Leaf width- leaf length | LW = aLW LL | aLW | 0.76 | 0.69 | 0.61 | 0.52 | 0.62 | 0.54 | 0.58 | 0.50 | 0.58 | 0.53 |
r2 | 0.99** | 0.99** | 0.98** | 0.98** | 0.99** | 0.98** | 0.97** | 0.97** | 0.97** | 0.98** |
品种 Cultivar | 枝梢类型 Shoot type | 实测叶倾角 Measured value | Beta方程参数及叶倾角估计值 Coefficients and estimates of Beta quation | 叶倾角类型 Leaf inclination type | ||||
---|---|---|---|---|---|---|---|---|
平均值/º Mean value | 标准误 Standar deviation | μ | ν | 平均值/º Mean value | 标准误 Standard deviation | |||
礼泉短富 | 果台Bourse | 44.26 a | 21.54 | 1.70 | 1.65 | 44.22 | 24.72 | 斜生叶Plagiophile |
Liquan | 果台长副梢Bourse long shoot | 35.11 c | 19.21 | 2.58 | 1.65 | 35.08 | 20.60 | 水平叶Planophile |
Fuji | 果台短副梢Bourse short shoot | 38.45 b | 22.19 | 1.73 | 1.29 | 38.41 | 18.51 | 水平叶Planophile |
营养长枝Vegetative long shoot | 27.96 e | 15.40 | 4.35 | 1.96 | 27.93 | 21.86 | 水平叶Planophile | |
营养短枝Vegetative short shoot | 32.89 d | 18.40 | 2.86 | 1.81 | 32.85 | 21.92 | 水平叶Planophile | |
丽嘎拉 | 果台Bourse | 43.19 a | 19.45 | 2.26 | 2.08 | 43.14 | 20.90 | 斜生叶Plagiophile |
Regal | 果台长副梢Bourse long shoot | 34.05 c | 16.78 | 3.58 | 2.18 | 34.01 | 18.42 | 水平叶Planophile |
Gala | 果台短副梢Bourse short shoot | 35.22 c | 17.60 | 3.18 | 2.05 | 35.19 | 20.74 | 水平叶Planophile |
营养长枝Vegetative long shoot | 30.79 d | 16.00 | 4.02 | 2.09 | 30.75 | 20.18 | 水平叶Planophile | |
营养短枝Vegetative short shoot | 32.20 d | 17.43 | 3.29 | 1.83 | 32.16 | 21.61 | 水平叶Planophile |
表2 ‘礼泉短富’和‘丽嘎拉’不同类型枝梢实测叶倾角均值和标准误及Beta方程参数和估计值
Table 2 Means and standard deviations for leaf inclination and estimates of two-parameters Beta equation for all shoot types in‘Liquan Fuji’and‘Regal Gala’apple trees,respectively
品种 Cultivar | 枝梢类型 Shoot type | 实测叶倾角 Measured value | Beta方程参数及叶倾角估计值 Coefficients and estimates of Beta quation | 叶倾角类型 Leaf inclination type | ||||
---|---|---|---|---|---|---|---|---|
平均值/º Mean value | 标准误 Standar deviation | μ | ν | 平均值/º Mean value | 标准误 Standard deviation | |||
礼泉短富 | 果台Bourse | 44.26 a | 21.54 | 1.70 | 1.65 | 44.22 | 24.72 | 斜生叶Plagiophile |
Liquan | 果台长副梢Bourse long shoot | 35.11 c | 19.21 | 2.58 | 1.65 | 35.08 | 20.60 | 水平叶Planophile |
Fuji | 果台短副梢Bourse short shoot | 38.45 b | 22.19 | 1.73 | 1.29 | 38.41 | 18.51 | 水平叶Planophile |
营养长枝Vegetative long shoot | 27.96 e | 15.40 | 4.35 | 1.96 | 27.93 | 21.86 | 水平叶Planophile | |
营养短枝Vegetative short shoot | 32.89 d | 18.40 | 2.86 | 1.81 | 32.85 | 21.92 | 水平叶Planophile | |
丽嘎拉 | 果台Bourse | 43.19 a | 19.45 | 2.26 | 2.08 | 43.14 | 20.90 | 斜生叶Plagiophile |
Regal | 果台长副梢Bourse long shoot | 34.05 c | 16.78 | 3.58 | 2.18 | 34.01 | 18.42 | 水平叶Planophile |
Gala | 果台短副梢Bourse short shoot | 35.22 c | 17.60 | 3.18 | 2.05 | 35.19 | 20.74 | 水平叶Planophile |
营养长枝Vegetative long shoot | 30.79 d | 16.00 | 4.02 | 2.09 | 30.75 | 20.18 | 水平叶Planophile | |
营养短枝Vegetative short shoot | 32.20 d | 17.43 | 3.29 | 1.83 | 32.16 | 21.61 | 水平叶Planophile |
图3 各类枝梢三维虚拟图像和基于三维虚拟冠层的‘礼泉短富’和‘丽嘎拉’冠层净光合速率模拟值示意图
Fig. 3 Representations of 3D virtual shoot for vegetative long shoot,vegetative short shoot,bourse long shoot,bourse short shoot and bourse at shoot scale and modeled canopy net photosynthesis rate based on 3D virtual canopies for‘Liquan Fuji’ and‘Regal Gala’apple trees,respectively
图4 各类枝梢模拟叶片数量、叶面积与光能截获效率STAR值与观测值比较 图d、i和o的纵坐标为对应变量的模拟值。
Fig. 4 Comparisons between simulated shoot leaf number,shoot leaf area and shoot silhouette to total area ratio(STAR)from virtual shoots and those values from observed data for all shoot types in‘Liquan Fuji’and‘Regal Gala’apple trees,respectively Tittles for y-axes for figure d,i and o were simulated values.
品种 Cultivar | 树高/cm Tree Height | 干周/cm Trunk circumstance | 叶面积/m2 Leaf area | 叶面积指数/ (m2 · m-2) LAI | 叶面积密度/(m2 · m-3)Leaf area density | 叶片分散度 Leaf dispersion | 单株枝量 Shoot number per tree |
---|---|---|---|---|---|---|---|
礼泉短富Liquan Fuji | 310.00 ± 8.15 a | 32.75 ± 2.87 a | 24.74 ± 2.74 a | 4.22 ± 0.191 a | 4.73 ± 0.22 a | 0.44 ± 0.02 a | 1 099 ± 119 a |
丽嘎拉 Regal Gala | 314.00 ± 6.26 a | 27.73 ± 2.36 b | 24.41 ± 3.27 a | 4.17 ± 0.114 a | 4.65 ± 0.21 a | 0.48 ± 0.03 b | 805 ± 120 b |
表3 ‘礼泉短富’和‘丽嘎拉’冠层结构信息
Table 3 Summary of canopy structural parameters for‘Liquan Fuji’and‘Regal Gala’apple trees,respectively
品种 Cultivar | 树高/cm Tree Height | 干周/cm Trunk circumstance | 叶面积/m2 Leaf area | 叶面积指数/ (m2 · m-2) LAI | 叶面积密度/(m2 · m-3)Leaf area density | 叶片分散度 Leaf dispersion | 单株枝量 Shoot number per tree |
---|---|---|---|---|---|---|---|
礼泉短富Liquan Fuji | 310.00 ± 8.15 a | 32.75 ± 2.87 a | 24.74 ± 2.74 a | 4.22 ± 0.191 a | 4.73 ± 0.22 a | 0.44 ± 0.02 a | 1 099 ± 119 a |
丽嘎拉 Regal Gala | 314.00 ± 6.26 a | 27.73 ± 2.36 b | 24.41 ± 3.27 a | 4.17 ± 0.114 a | 4.65 ± 0.21 a | 0.48 ± 0.03 b | 805 ± 120 b |
图6 ‘礼泉短富’和‘丽嘎拉’苹果枝类组成 P值为非参数卡方测验结果,图中同一类枝梢不同小写字母表示两品种间差异显著(P < 0.05)。
Fig. 6 Fractions of shoots number for each shoot type for‘Liquan Fuji’and‘Regal Gala’apple trees,respectively P value was obtained from Chi-square test. Different lowercase letters for the same shoot type indicate significant differences between two cultivars at 0.05 level.
品种 Cultivar | STAR值 Silhouette to total area ratio | 冠层净光合速率/(mmol · CO2 · m-2 · d-1) Canopy net photosynthesis rate | 冠层日光合量/(mol · CO2 · d-1) Canopy daily total photosynthesis | ||
---|---|---|---|---|---|
晴天 Sunny day | 阴天 Cloudy day | 晴天Sunny day | 阴天Cloudy day | ||
礼泉短富 Liquan Fuji | 0.25 ± 0.01 a | 185.41 ± 8.12 b | 32.05 ± 8.24 b | 4.91 ± 0.24 b | 0.83 ± 0.14 b |
丽嘎拉 Regal Gala | 0.26 ± 0.01 a | 202.83 ± 0.33 a | 79.11 ± 0.72 a | 5.49 ± 0.62 a | 2.15 ± 0.23 a |
表4 ‘礼泉短富’和‘丽嘎拉’苹果冠层光能截获效率STAR值、净光合速率和日光合量
Table 4 Canopy silhouette to total area ratio(STAR),canopy photosynthesis rate and canopy daily photosynthesis in sunny and cloudy days for‘Liquan Fuji’and‘Regal Gala’apple trees,respectively
品种 Cultivar | STAR值 Silhouette to total area ratio | 冠层净光合速率/(mmol · CO2 · m-2 · d-1) Canopy net photosynthesis rate | 冠层日光合量/(mol · CO2 · d-1) Canopy daily total photosynthesis | ||
---|---|---|---|---|---|
晴天 Sunny day | 阴天 Cloudy day | 晴天Sunny day | 阴天Cloudy day | ||
礼泉短富 Liquan Fuji | 0.25 ± 0.01 a | 185.41 ± 8.12 b | 32.05 ± 8.24 b | 4.91 ± 0.24 b | 0.83 ± 0.14 b |
丽嘎拉 Regal Gala | 0.26 ± 0.01 a | 202.83 ± 0.33 a | 79.11 ± 0.72 a | 5.49 ± 0.62 a | 2.15 ± 0.23 a |
图7 果台副梢和营养枝梢转换后冠层光能截获效率相对变化及晴天和阴天冠层净光合速率和日光合量的相对变化
Fig. 7 Relative changes of canopy silhouette to total area ratio(STAR)and net photosynthesis rate and total photosynthesis under sunny and cloudy conditions,by modifying relative fractions between bourse shoots and vegetative shoots, for‘Liquan Fuji’and‘Regal Gala’,respectively
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