基于微观力学的黄瓜叶片形态变化模型研究
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摘要
植物形态建模和形态模型的可视化是近年来计算机图形学和农业工程领域的研究热点,研究人员试图通过数字化技术实现农林植物形态变化过程的可视化表达。
现有植物形态变化模型主要有两类,一类是从图形学出发进行植物形态变化的模型研究,该领域的研究能得到逼真的形态图像,但建模时,末考虑作物生长的真实性,该类模型不能用于实际生产指导；另一类是农业工程领域研究人员试图结合植物生理学建立符合植物自身特性的虚拟植物模型,并在虚拟环境中模拟植物对环境的响应情况,这部分模型大部分是基于宏观实验的统计模型,虽然能在一定程度上反映作物的形态变化,但并未对作物形态发生变化的机理进行探究,无法获得植物对环境的响应机制,因此在实际应用中存在局限性。
针对现有模型的不足,本文从“作物形态的变化过程就是其内部组织细胞的变形过程”这个观点出发,对黄瓜叶片宏观形态变化及其内部细胞的变化情况进行实验研究,在此基础上建立了基于微观力学的宏观形态变化模型,并与真实形态进行了比较,模拟结果和真实形态在一定范围内具有较好的吻合度。借助该模型可以了解特定宏观形态下,作物本身对水分、光温的需求情况和自身的生命特征,从而为生产实际中水分等环境因子的调控提供理论依据；对实现我国设施农业的高效和可持续发展具有十分重要的理论意义和实用价值。
1、考查了叶片含水量变化时,叶片表面气孔、叶片内部细胞及胞间结构变化情况。叶片含水量为89.9%时,所有的叶面气孔呈展开状态,细胞间隙明显,细胞形态饱满,细胞内部叶绿体片层清晰；当叶片含水量为86.7%时,只有1/3的表面气孔完全张开,细胞形态开始变得不饱满,细胞间隙减小,细胞内部叶绿体肿胀并出现脂类小球；叶片含水量为75.4%时,表面气孔基本处于完全闭合状态,叶片表面出现收缩,细胞呈现收缩,细胞间隙进一步减小,细胞内部叶绿体进一步肿胀甚至解体,叶绿体内部嗜锇颗粒增多增大；叶片水分含量为60%左右时,叶片完全萎蔫,表面气孔完全闭合,部分气孔的保卫细胞出现坏死现象,表面皱褶进一步加剧,细胞变形严重,细胞间隙消失,细胞内部大多数细胞器解体。
2、利用切片技术和纳米压痕技术对黄瓜叶肉细胞的细胞壁力学特性进行了测试,并对加载-卸载曲线、以及应力-应变曲线进行了分析。结果表明,当应力小于1Mpa时,细胞壁呈现非线性弹性特征；当应力小于0.4Mpa时,细胞壁的非线性特征减弱,可以视为是线性弹性材料。本文所得到细胞壁力学参数是在对黄瓜叶片组织进行脱水处理后测最得到的,而植物体内细胞壁含有少量水分,因此两材存在一定的差异,在应用细胞壁的力学参数时需要考虑水分的影响。
3、采用FEM有限元方法,建立了符合生理学特征的叶肉细胞力学模型—一内部充满液体的球形细胞力学模型,根据水分代谢方程,计算出于蒸腾作用而产生肉细胞内膨压变化量,对单个细胞进行受力分析,得到细胞壁上任一点在膨压随时间变化的情况。
4、对叶脉结构进行多尺度分析的基础上,建立了细胞结构、分层结构、膨压对叶脉力学性能影响的计算模型。
5、在植物组织力学性能方面,首次借助均匀化方法,得到了叶肉组织等效材料属性的计算模型,并比较了叶片在不同水分条件下力学特性的变化。其计算结果为计算叶片的宏观变形模型提供了有效的的依据。
6、首次借助三维激光扫描与重建技术、显微技术,结合生物力学,建立了叶片的变形模型,根据不同的水分情况,计算得到不同的宏观形态模型。将模拟结果和真实叶片的三维扫描数据进行了比较,结果发现两者有较好的吻合度。借助该模型计算了各种形态黄瓜叶片的含水量,叶片含水量子啊70%-90%之间变化时,模型的计算值与实测值之间基于1:1上直线的决定系数R2为0.823, RMSE为3.4223。
7、根据光温因子和黄瓜叶片的生长情况,建立了以光温因子为驱动变量,黄瓜叶特征参数、叶柄直径和长度的模拟模型。在坫此基础上,对不同光温因子条件下生长的叶片进行缺水情况的模拟,得到光温因子影响下的叶片缺水形态变化。
Plant morphologic modeling and visualizing is the research focus in computer graphics and agricultural engineering in recent years, researchers try to model and visualize the process of plant morphology variation by digital technology.
The existing plant morphologic model mainly can be divided into two types:one type model is built by computer graphics scientist, from which lively morphologic images can be obtained. However, the lack of understanding about plant morphology change mechanism lead to this kind of model cannot be used in practical production.Another one is virtual plant model built by researchers in the agricultural engineering, which combined with plant growth process and based on macroscopic experimental, could simulate plant's response to environment to a certain extent. This kind of model also have not made thorough research about plant morphology change mechanism, so response mechanism of plant to environment can't be obtained. but it has limitation in practice.Aim at these limitations, this thesis try to establish the macro-morphological change models from the view of "morphologic changes of crop depend on the alternation of cells in it" by using micromechanics. The simulation results were compared with authenticity, they correspond with each other. With the help of the model, requirements of the crop in water, light, temperature can be learnt. The model also can provide a basis of how to control the environmental factors such as water and light. So the research is of great theoretical significance to realize the facility agriculture of high efficiency and sustainable development.
1. The alteration of stomason the surface and cells in leaves were examined when leaf water content decreased. When the leaf water content was89.9%, stomas were openmainly, the intercellular space was obvious, the shape of the cells was perfect, chloroplast lamellar was visible. The opened stomas are decreased as leaf water content reduced, when itwas86.5%, only1/3stomaswerein the open state, at the same time leaves began to fold;the cellswere atrophy, the intercellular space became small, chloroplasts were swollen,osmiophilic globulesappeared in them.When leaf water content was75%around, all the stomas on the surface of leaves closedcompletely, the leaf folded further, the intercellular space deceased further,chloroplasts were swollen much more even some of them disintegrated, osmiophilic globule became more and bigger.When the leaf water content decreased to60%around, all the stomas on the surface of leaves closedcompletely, some of them were necrotic, the leaf folded further, the cells shape changed seriously, intercellular space disappeared, and cells glued together, most organelles in the cellsdisintegrated.With the further lack of water, organelles within cells gradually disintegrated, then part of the cell walls disintegrated too.
2. The mechanical property of cue μ mber's mosophyll cell wall was tested by using slicing technique and nano-indentation technology. The results (loading-unloading curve, andstress-strain curve) were analyzed.Results show that when the stress is less than1Mpa, the cell-wall has non-linear elastic characteristics; when the stress is less than0.4Mpa, the nonlinear characteristics of the cell walls weaken, which can be considered as a linear elastic material. In this article, the mechanical property of cell wall was measured after dehydration. Actually cell wall containing a amount of water, so there are certain differences between the two. So in the application of mechanical parameters of the cell wall, the effects of water need to be considered.
3.A mechanical model of cue μ mber mosophyll cell (spherical ball filled with liquid)was established by using FEM (finite element method).According to the water metabolism equation, the turgor pressure change caused by transpiration have been calculated. Through mechanical analysis of single cell, dynamics equation of turgor pressure changes over time was obtained.
4. A computation model was established to analyzethe impact of cell structure and turgor pressure on the mechanical property of vein, according to the hierarchy structure of vein.
5.A computation model was establishedto get mesophyll tissue equivalent mechanical properties by using homogenization method for the first time. The result shows the relation between mechanical property of leaf and water in it. The result also provides an effective basis to calculate the leaf's macroscopic deformation model.
6.By using of three dimensional laser scanning, reconstruction technique, micro technologies, combined with bio-mechanics, aleaf deformation model under different water conditions is established at the first time. Simulation results of leaf shap and scanned data of real leaf correspond well.The water contents in differentleave which were water stressed were calculated with the model, and the result was compared with the measured data. When the leafwater content change between70%and90%, the coefficient of determination (R2)of independent validation set was0.823,and RMSE is3.4223.
7According to light, temperature factor and growth of cucμmber leaf, themodel in which light temperature factor was the driving variables was built to simulate the grow of cucμmber leaf, petiole length and diameter.The leafparameters under different light temperature conditions were calculated by the model. Then,the shape of the leave water stressed were obtained.
现有植物形态变化模型主要有两类,一类是从图形学出发进行植物形态变化的模型研究,该领域的研究能得到逼真的形态图像,但建模时,末考虑作物生长的真实性,该类模型不能用于实际生产指导；另一类是农业工程领域研究人员试图结合植物生理学建立符合植物自身特性的虚拟植物模型,并在虚拟环境中模拟植物对环境的响应情况,这部分模型大部分是基于宏观实验的统计模型,虽然能在一定程度上反映作物的形态变化,但并未对作物形态发生变化的机理进行探究,无法获得植物对环境的响应机制,因此在实际应用中存在局限性。
针对现有模型的不足,本文从“作物形态的变化过程就是其内部组织细胞的变形过程”这个观点出发,对黄瓜叶片宏观形态变化及其内部细胞的变化情况进行实验研究,在此基础上建立了基于微观力学的宏观形态变化模型,并与真实形态进行了比较,模拟结果和真实形态在一定范围内具有较好的吻合度。借助该模型可以了解特定宏观形态下,作物本身对水分、光温的需求情况和自身的生命特征,从而为生产实际中水分等环境因子的调控提供理论依据；对实现我国设施农业的高效和可持续发展具有十分重要的理论意义和实用价值。
1、考查了叶片含水量变化时,叶片表面气孔、叶片内部细胞及胞间结构变化情况。叶片含水量为89.9%时,所有的叶面气孔呈展开状态,细胞间隙明显,细胞形态饱满,细胞内部叶绿体片层清晰；当叶片含水量为86.7%时,只有1/3的表面气孔完全张开,细胞形态开始变得不饱满,细胞间隙减小,细胞内部叶绿体肿胀并出现脂类小球；叶片含水量为75.4%时,表面气孔基本处于完全闭合状态,叶片表面出现收缩,细胞呈现收缩,细胞间隙进一步减小,细胞内部叶绿体进一步肿胀甚至解体,叶绿体内部嗜锇颗粒增多增大；叶片水分含量为60%左右时,叶片完全萎蔫,表面气孔完全闭合,部分气孔的保卫细胞出现坏死现象,表面皱褶进一步加剧,细胞变形严重,细胞间隙消失,细胞内部大多数细胞器解体。
2、利用切片技术和纳米压痕技术对黄瓜叶肉细胞的细胞壁力学特性进行了测试,并对加载-卸载曲线、以及应力-应变曲线进行了分析。结果表明,当应力小于1Mpa时,细胞壁呈现非线性弹性特征；当应力小于0.4Mpa时,细胞壁的非线性特征减弱,可以视为是线性弹性材料。本文所得到细胞壁力学参数是在对黄瓜叶片组织进行脱水处理后测最得到的,而植物体内细胞壁含有少量水分,因此两材存在一定的差异,在应用细胞壁的力学参数时需要考虑水分的影响。
3、采用FEM有限元方法,建立了符合生理学特征的叶肉细胞力学模型—一内部充满液体的球形细胞力学模型,根据水分代谢方程,计算出于蒸腾作用而产生肉细胞内膨压变化量,对单个细胞进行受力分析,得到细胞壁上任一点在膨压随时间变化的情况。
4、对叶脉结构进行多尺度分析的基础上,建立了细胞结构、分层结构、膨压对叶脉力学性能影响的计算模型。
5、在植物组织力学性能方面,首次借助均匀化方法,得到了叶肉组织等效材料属性的计算模型,并比较了叶片在不同水分条件下力学特性的变化。其计算结果为计算叶片的宏观变形模型提供了有效的的依据。
6、首次借助三维激光扫描与重建技术、显微技术,结合生物力学,建立了叶片的变形模型,根据不同的水分情况,计算得到不同的宏观形态模型。将模拟结果和真实叶片的三维扫描数据进行了比较,结果发现两者有较好的吻合度。借助该模型计算了各种形态黄瓜叶片的含水量,叶片含水量子啊70%-90%之间变化时,模型的计算值与实测值之间基于1:1上直线的决定系数R2为0.823, RMSE为3.4223。
7、根据光温因子和黄瓜叶片的生长情况,建立了以光温因子为驱动变量,黄瓜叶特征参数、叶柄直径和长度的模拟模型。在坫此基础上,对不同光温因子条件下生长的叶片进行缺水情况的模拟,得到光温因子影响下的叶片缺水形态变化。
Plant morphologic modeling and visualizing is the research focus in computer graphics and agricultural engineering in recent years, researchers try to model and visualize the process of plant morphology variation by digital technology.
The existing plant morphologic model mainly can be divided into two types:one type model is built by computer graphics scientist, from which lively morphologic images can be obtained. However, the lack of understanding about plant morphology change mechanism lead to this kind of model cannot be used in practical production.Another one is virtual plant model built by researchers in the agricultural engineering, which combined with plant growth process and based on macroscopic experimental, could simulate plant's response to environment to a certain extent. This kind of model also have not made thorough research about plant morphology change mechanism, so response mechanism of plant to environment can't be obtained. but it has limitation in practice.Aim at these limitations, this thesis try to establish the macro-morphological change models from the view of "morphologic changes of crop depend on the alternation of cells in it" by using micromechanics. The simulation results were compared with authenticity, they correspond with each other. With the help of the model, requirements of the crop in water, light, temperature can be learnt. The model also can provide a basis of how to control the environmental factors such as water and light. So the research is of great theoretical significance to realize the facility agriculture of high efficiency and sustainable development.
1. The alteration of stomason the surface and cells in leaves were examined when leaf water content decreased. When the leaf water content was89.9%, stomas were openmainly, the intercellular space was obvious, the shape of the cells was perfect, chloroplast lamellar was visible. The opened stomas are decreased as leaf water content reduced, when itwas86.5%, only1/3stomaswerein the open state, at the same time leaves began to fold;the cellswere atrophy, the intercellular space became small, chloroplasts were swollen,osmiophilic globulesappeared in them.When leaf water content was75%around, all the stomas on the surface of leaves closedcompletely, the leaf folded further, the intercellular space deceased further,chloroplasts were swollen much more even some of them disintegrated, osmiophilic globule became more and bigger.When the leaf water content decreased to60%around, all the stomas on the surface of leaves closedcompletely, some of them were necrotic, the leaf folded further, the cells shape changed seriously, intercellular space disappeared, and cells glued together, most organelles in the cellsdisintegrated.With the further lack of water, organelles within cells gradually disintegrated, then part of the cell walls disintegrated too.
2. The mechanical property of cue μ mber's mosophyll cell wall was tested by using slicing technique and nano-indentation technology. The results (loading-unloading curve, andstress-strain curve) were analyzed.Results show that when the stress is less than1Mpa, the cell-wall has non-linear elastic characteristics; when the stress is less than0.4Mpa, the nonlinear characteristics of the cell walls weaken, which can be considered as a linear elastic material. In this article, the mechanical property of cell wall was measured after dehydration. Actually cell wall containing a amount of water, so there are certain differences between the two. So in the application of mechanical parameters of the cell wall, the effects of water need to be considered.
3.A mechanical model of cue μ mber mosophyll cell (spherical ball filled with liquid)was established by using FEM (finite element method).According to the water metabolism equation, the turgor pressure change caused by transpiration have been calculated. Through mechanical analysis of single cell, dynamics equation of turgor pressure changes over time was obtained.
4. A computation model was established to analyzethe impact of cell structure and turgor pressure on the mechanical property of vein, according to the hierarchy structure of vein.
5.A computation model was establishedto get mesophyll tissue equivalent mechanical properties by using homogenization method for the first time. The result shows the relation between mechanical property of leaf and water in it. The result also provides an effective basis to calculate the leaf's macroscopic deformation model.
6.By using of three dimensional laser scanning, reconstruction technique, micro technologies, combined with bio-mechanics, aleaf deformation model under different water conditions is established at the first time. Simulation results of leaf shap and scanned data of real leaf correspond well.The water contents in differentleave which were water stressed were calculated with the model, and the result was compared with the measured data. When the leafwater content change between70%and90%, the coefficient of determination (R2)of independent validation set was0.823,and RMSE is3.4223.
7According to light, temperature factor and growth of cucμmber leaf, themodel in which light temperature factor was the driving variables was built to simulate the grow of cucμmber leaf, petiole length and diameter.The leafparameters under different light temperature conditions were calculated by the model. Then,the shape of the leave water stressed were obtained.
引文
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