SummaryMany scholars conclude that leaf shape is highly related with the veins. Based on this theory, we assume the leaf growth in each direction satisfies a function. For the leaves in the same tree, the parameters are different; for those of separate trees, the function mode is different. Thus the shape of leaf differs from that of another. In the end of section 3, we simulate one growing period and depict the leaf shape.Through thousands of years of evolution, the leaves find various wa ys to make a full use of natural resources, including minimizing overlapping individual shadows. In order to find the main factors promoting the evolution of leaves, we analyze the distribution of adjacent leaves and the equilibrium point of photosynthesis and respiration. Besides, we also make a coronary hierarchical model and transmission model of the solar radiation to analyze the influence of the branches.As to the tree structure and the leaf shape, first we consider one species. Different tree shapes have different space which is built up by the branch quality and angle, effect light distribution, ventilation and humidity and concentration of CO2 in the tree crown. These are the factors which affect the leaf shape according to the model in section 1. Here we analyze three typical tree shapes: Small canopy shape, Open center shape and Freedom spindle shape, which can be described by BP network and fractional dimension model. We find that the factors mainly affect the function of Sthat affects the additional leaf area. Factors are assembled in different ways to create different leaf shapes. So that the relationship between leaf shape and tree profile/branching structure is proved.Finally we develop a model to calculate the leaf mass from the basic formula of . By adjusting the crown of a tree to a half ellipsoid, we first define thefunction of related factors,such as the leaf density and the effective ratio of leaf area. Then we develop the model using calculus. With this model, weapproximately evaluate the leaf mass of a middle-sized tree is 141kg.Dear editor,How much the leaves on a tree weigh is the focus of discussion all the time. Our team study on the theme following the current trend and we find something interesting in the process.The tree itself is component by many major elements. In our findings, we analyze the leaf mass with complicated ones, like leaf shape, tree structure and branch characteristics, which interlace with each other.With the theory that leaf shape is highly related with the veins, we assume theleaf growth in each direction satisfies a function . For the leaves in thesame tree, the parameters are different; for those of separate trees, the function mode is di fferent. That’s why no leaf shares the same shape. Also, we simulate one growing period and depict the leaf shape.In order to find the main factors promoting the evolution of leaves, we analyze the distribution of adjacent leaves and the equilibrium point of photosynthesis and respiration. Besides, we also make a coronary hierarchical model and transmission model of the solar radiation to analyze the influence of the branches.As to the tree structure and the leaf shape, different tree shapes have differe nt space which is built up by the branch quality and angle, effect light distribution, ventilation and humidity and concentration of CO2 in the tree crown that affect leaf shapes. Here we analyze three typical tree shapes which can be described by BP network and fractional dimension model. We find that the factors mainly affect the function of Sthat affects the additional leaf area. Factors are assembled in different ways to create different leaf shapes. So that the relationship between leaf shape and tree profile or branching structure is proved.Finally we develop the significant model to calculate the leaf mass from thebasic formula of . By adjusting the crown of a tree to a half ellipsoid,we first define the function of related factors and then we develop the model using calculus. With this model, we approximately evaluate the leaf mass of a middle-sized tree is 141kg.We are greatly appreciated that if you can take our findings into consideration. Thank you very much for your precious time for reading our letter.Yours sincerely,Team #14749Contents1. Introduction (4)2. Parameters (4)3. Leaves have their own shapes (5)3.1 Photosynthesis is important to plants (5)3.2 How leaves grow? (6)3.3 Build our model (7)3.4 A simulation of the model (10)4. Do the shapes maximize exposure? (14)4.1 The optimum solution of reducing overlapping shadows (14)4.1.1 The distribution of adjacent leaves (14)4.1.2 Equilibrium point of photosynthesis and respiration (15)4.2 The influence of the “volume” of a tree and its branches (17)4.2.1 The coronary hierarchical model (17)4.2.2 Spatial distribution model of canopy leaf area (18)4.2.3 Transmission model of the solar radiation (19)5. Is leaf shape related to tree structure? (20)5.1 The experiment for one species (21)5.2 Different tree shapes affect the leaf shapes (23)5.2.1 The light distribution in different shapes (23)5.2.2 Wind speed and humidity in the canopy (23)5.2.3 The concentration of carbon dioxide (24)5.3 Conclusion and promotion (25)6. Calculus model for leaf mass (26)6.1 How to estimate the leaf mass? (26)6.2 A simulation of the model (28)7. Strengths and Weakness (29)7.1 Strengths (29)7.2 Weaknesses (30)8. Reference (30)1. IntroductionHow much do the leaves on a tree weigh? Why do leaves have the various shapes that they have? How might one estimate the actual weight of the leaves? How might one classify leaves?We human-beings have never stopped our steps on exploring the natural world. But, as a matter of fact, the answer to those questions is still unresolved. Many scientists continue to study on this area. Recently , Dr. Benjamin Blonder (2010) achieved a new breakthrough on the venation networks and the origin of the leaf econo mics spectrum. They defined a standardized set of traits – density , distance and loopiness and developed a novel quantitative model that uses these venation traits to model leaf-level physiology .Now, it is commonly thought that there are four key leaf functional traits related to leaf economics: net carbon assimilation rate, life span, leaf mass per area ratio and nitrogen content.2. Parametersthe area a leaf grows decided by photosynthesisthe additional leaf area in one growing periodthe leaf growing obliquity Pthe total photosynthetic rate 0d R the dark respiration rate of leavesn P the net photosynthetic rateh the height of the canopyd the distance between two branchesdi the illumination intensity of scattered light from a given directionthe solar zenith angleh the truck highh the crown high13. Leaves have their own shapes3.1 Photosynthesis is important to plantsIt is widely accepted that two leaves are different, no matter where they are chosen from; even they are from the very tree. To understand how leaves grow is helpful to answer why leaves have the various shapes that they have.The canopy photosynthesis and respiration are the central parts of most biophysical crop and pasture simulation models. In most models, the acclamatory responses of protein and the environmental conditions, such as light, temperature and CO2 concentration, are concerned[1].In 1980, Farquhar et al developed a model named FvCB model to describe photosynthesis[2]:The FvCB model predicts the net assimilation rate by choosing the minimum between the Rubisco-limited net photosynthetic rate and the electron transport-limited net photosynthetic rate.Assume A n, A c, A j are the symbols for net assimilation rate, the Rubisco-limited net photosynthetic rate and the electron transport-limited net photosynthetic rate respectively, and the function can be described as:(1)(2)where and are the intercellular partial pressures of CO2 and O2,respectively, and are the Michaelis–Menten coefficients ofRubisco for CO2and O2, respectively, is the CO2compensation point inthe absence of (day respiration in andis the photosystem II electron transport rate that is used for CO2fixation and photorespiration[3][4].We apply the results of this model to build the relationship between the photosynthesis and the area a leaf grows during a period of time. It can be released as:(3)and are the area of the target leaf and the period of time it grows.is a function which can transfer the amount ofCO2into the area the leaf grows and the are parameters which affect S p. S p can be used as a constraint condition in our model.3.2 How leaves grow?As the collocation of computer hardware and software develops, people can refer to bridging biology, morphogenesis, applied mathematics and computer graphics to simulate living organisms[5], thus how to model leaves is of great challenge. In 2001, Dengler and Kang[6]brought up the thought that leaf shape is highly related to venation patterns. Recently, Runions[7] brought up a method to portray the leaf shape by analyzing venation patterns. Together with the Lindenmayer system (L-system), an advanced venation model can adjust the growth better that it solved the problem occurred in the previous model that the secondary veins are retarded.We knowleaves have various shapes.For example, leaves can be classified in to simple leaves which have an undivided blade and compound leaves whose blade is divided into two or more distinctleaflets such as the Fabaceae. As to the shape of a leaf, it may have marginal dentations of the leaf blades or not, and like a palm with various fingers or an elliptical cake. Judd et al defined a set of terms which describe the shape of leaves as follows [8]:We chose entire leaves to produce this model as a simplification. What’s more, they confirmed again that the growth of venations relates with that of the leaf.To disclose this relationship, Relative Elementary Rate of Growth (RERG) can be introduced to depict leaves growth [9]. RERG is defined as the growth rate per distance, in the definitive direction l at a point p of the growing object, yielding(4)Considered RERG , the growth patterns of leaves are also different. Roth-Nebelsick et al brought up four styles in their paper [10]:3.3 Build our modelWe chose marginal growth to build our model. Amid all above-mentioned studies, weFigure 3.1 Terms pertinent to the description of leaf shapes.Figure 3.2 A sample leaf (a) and the results of its: (b) marginal growth, (c) uniform isotropic (isogonic) growth, (d) uniform anisotropic growth, and (e) non-uniform anisotropic growth.Figure 3.3 The half of a leaf is settled in x-y plane like this with primary vein overlapping x-axis. The leaf grows in the direction of .assume that the leaf produce materials it needs to grow by photosynthesis to expand its leaf area from its border and this process is only affected by what we have discussed in the previous section about photosynthesis. The border can be infinitesimally divided into points. Set as the angle between the x-axis and thestraight line connecting the grid origin and one point on the curve,andas the growth distance in the direction of . To simplify the model, we assume that the leaf grows symmetrical. We put half of the leaf into the x-y plane and make the primary vein overlap x-axis.This is how we assume the leaf grows.In one circle of leaf growth, anything that photosynthesis provided transfers into theadditional leaf area, which can be described as:(5)while in the figure.In this case, we can simulate leaf growth thus define the leaf shape by using iterative operations the times N a leaf grow in its entire circle ①.① For instance, if the vegetative circle of a leaf is 20 weeks on average, the times of iterative operations N can beset as 20 when we calculate on a weekly basis.Figure 3.4 The curves of the adjacent growing period and their relationship.First, we pre-establish the border shape of a leaf in the x-y plane, yielding . where , the relevant satisfies:(6)(7) In the first growing period, assume , the growth distance in the direction of ,satisfies:(8)(9) It releases the relationship of the coordinates in the adjacent growing period. In this case, we can use eq.(9)to predict the new border of the leaf after one period of growth②:(10)And the average simple recursions are③:②That means, in the end of period 1.③As we both change the x coordinate and the y coordinate, in the new period, these two figures relate through those in the last period in the functions.(11)After simulate the leaf borders of the interactive periods, use definite integral ④ to settle parametersin the eq.(8) then can be calculated in each direction of , thus the exact shape of a leaf in the next period is visible.When the number of times N the leaf grows in its life circle applies above-mentioned recursions to iterate N times and the final leaf shape can be settled.By this model, we can draw conclusions about why leaves have different shapes. For the leaves on the same tree, they share the same method of expansion which can be described as the same type of function as Eq.(8). The reason why they are different, not only in a sense of big or small, is that in each growing period they acquire different amount of materials used to expand its own area. In a word, the parameters in the fixedly formed eq.(8) are different for any individual leaf on the same tree. For the leaves of different tree species, the corresponding forms of eq.(8) are dissimilar. Some are linear, some are logarithmic, some are exponential or mixtures of that, which settle the totally different expansion way of leaf, are related with the veins. On that condition, the characters can be divided by a more general concept such as entire or toothed.3.4 A simulation of the modelWe set the related parameters by ourselves to simulate the shape of a leaf and to express the model better.First we initialize the leaf shape by simulating the function of a leaf border at the④The relationship must meet eq.(5).Figure 3.5 and the recurrence relations.beginning of growing period 1 in the x-y plane. By observation, we assume that themovement of the initial leaf border satisfied:(12)Suppose the curve goes across the origin of coordinates, then the constraint conditionscan be:(13)Thus the solution to eq.(13) is:(14)By using Mathematica we calculatewhereandthe area of the half leaf is: (15)Wesettleaccording to the research by S. V . Archontoulisin et al [11] in eq.(3). On a weekly basis, theparameter. In eq.(15), we have .Figure 3.6When assume ,thus constraint condition eq.(5) becomes: (16)When eq.(8) is linear and after referring to Runions’s paper, we assume that Y -valuedecreases when X-value increases, which means the leaf grows faster at the end of theprimary vein. If the grow rate at the end of the primary vein is 0,as ,eq.(8) can be described as: (17)where b is decided by eq.(16).To settle the value of b , we calculate multiple sets of data by Excel then use a planecurve to trace them and get the approximation of b . In this method, we use grid toapproximate .Apparently,is monotone.When b is 2: The square of each square is 0.0139cm 2, and the total number of the squares in theadditional area is about 240. SoFigure 3.7When b is 2.5:Thesquare of each squareis 0.0240cm 2, and the total number of the squares in theadditional area is about 201. SoWhen b is 3:The square of each square is 0.0320cm 2, and the total number of the squares in the additional area is about 193. SoAfter comparing, we can draw a conclusion that fit eq.(16)best; accordingly, in period 1: (18)In each growing period, may be different for the amount of material produced isFigure 3.9Figure 3.8related with various factors, such as the change of relative location and CO 2 or O 2concentration, and other reasons. By using the same method, the leaf shape in period2 or other period can be generated on the basis of the previous growing period untilthe end of its life circle.What’s more, when eq.(8) is remodeled, the corresponding leaf shape can be changed.With , the following figure shows the transformationof a leaf shape in the firstgrowing period with the samesettled above and we can see that the shape will be dissected in the end.4. Do the shapes maximize exposure?4.1 The optimum solution of reducing overlapping shadows4.1.1 The distribution of adjacent leavesV ein is the foundation of the leaves. With the growing of veins, the leaves graduallyexpand around. The distribution of main and lateral veins plays an important decisiverole in the shapes of leaves. The scientists created a mathematical model which usesthree decisive factors - the relationship between the rate of photosynthesis, leaf life,carbon consumption or nitrogen consumption, to simulate the leaves’ shape. Becauseof carbon consumption is a constant for one tree, and we take the neighboring leavesin the same growth cycle to observe. So, we can only focus on one factor - the rate ofphotosynthesis.Figure 3.10The shape differs from figure 3.7-3.9, as the kind offunction of is different. In this case, it is a cubic model while a linear model in figure 3.7-3.9.Through the observation of dicotyledon, leaves on a branch will grow in a staggered way that can reduce the overlapping individual shadows of adjacent leaves and make them get more sunlight (Figure 4.1).Figure 4.1 The rotation distribution of leavesBase on the similar environment, we assume that adjacent leaves nearly have the same shape. From the perspective of looking down, the leaves grow from a point on the branch. So, we can simplify the vertical view of leaves as a circle of which the radius is the length of a vein which is represented with r. The width of the leaf is represented with w. The angle between two leaves is represented with β(Figure4.2).Figure 4.2 The vertical view of leavesThe leaves should use the space as much as possible, and for the leaf with one main vein, oval is the best choice. In general, βis between 15°and 90°. In this way, effectively reduce the direct overlapping area. According to the analysis of the first question, r and w are determined by the rate of photosynthesis and respiration. Besides, the width of leaf is becoming narrower when the main vein turns to be thinner.4.1.2 Equilibrium point of photosynthesis and respirationThe organism produced by photosynthesis firstly satisfies needs of leaf itself. Then the remaining organism delivered to the root to meet the growth needs of the tree. As we know, respiration needs to consume organism. If the light is not sufficient, organism produced by photosynthesis may no longer be able to afford the materials required for the growth of leaves. There should be an equilibrium point so as toprevent the leaf is behindhand in its circumstances.The formula [12] that describes the photosynthetic rate in response to light intensity with gradual exponential growth index can be expressed as:(19) Where P is total photosynthetic rate, m ax P is maximum photosynthetic rate ofleaves, a is initial solar energy utilization and I is photosynthetic photon quanta flux density .The respiration rate is affected by temperature ，using the formal of index to describe as follows:025102T d d R R -=⋅ (20)Where T is temperature and 0d R is the dark respiration rate of leaves, when the temperature is 25 degrees. As a model parameter, 0d R can be determined bynonlinear fitting. Therefore, the net photosynthetic rate can be expressed in the indexform as follows:(21)Where n P is the net photosynthetic rate, which does not include the concentration ofcarbon dioxide and other factors. The unit of n P is 21m ol m s μ--⋅⋅.We assume that the space for leaf growth is limited, the initial area of the leaf is 0S . At this point, the entire leaf happens to be capable of receiving sunlight. If the leaf continue to grow, some part of the leaf will be in the shadows and the area in the shadows is represented with x . Ignore the fluctuation cycle of photosynthesis and respiration, on average, the duration of photosynthesis is six hours per day . In the meanwhile, the respiration is ongoing all the time. When the area increased to S , we established an equation as follows:(22)where μ is the remaining organism created by the leaf per day . And from where 0μ=, which means the organism produced by photosynthesis has all been broken down completely in the respiration, we get the equilibrium point.4n n d S P x P R ⋅=+ (23) The proportion of the shaded area in the total area:4nn d P xS P R =+ (24)Cite an example of oak trees, we found the following data (Figure 4.3), which shows the relationship between net photosynthesis and dark respiration during 160 days. Thehabitat of the trees is affected by the semi-humid monsoon climate.Figure 4.3 The diurnal rate of net photosynthesis and dark respiration [13]In general, 217n P m ol m s μ--=⋅⋅，214d R m ol m s μ--=⋅⋅. Taking the given numbersinto the equation, we can get the result.0025xS =From the result, we can see that in the natural growth of leaves, with the weakening of photosynthesis, the leaves will naturally stop growing once they come across the blade between blocked. Therefore, the leaves can always keep overlapping individual shadows about 25%, so as to maximize exposure.4.2 The influence of the “volume” of a tree and its branches4.2.1 The coronary hierarchical modelMr. Bōken and Dr. J. Fischer (1987) found that in order to adequate lighting, leaves have different densities and the branches is distributed according to certain rules. They observed tropical plants in Miami, found that the ratio of main branch and two side branches is 1:0.94:0.87, and the angles between them are 24.4°and 36.9°. According to the computer simulation, the two angles can maximize the exposure of leaves.For simply, we use hemisphere to simulate the shape of the canopy. According to the light transmittance rate, we can divide the canopy into outer and inner two layers(Figure 4.3). The volume of the hemisphere depends on the size and distribution of branches.Figure 4.3 The coronary hierarchical modelThe angle between the branches will affect the depth of penetration of sun radiation and the distribution of leaves. Besides, thickness of the branches will affect the transfer of nutrients to the leaves.4.2.2 Spatial distribution model of canopy leaf areaAssume that the distribution of leaves is uniform in the section xz but not uniform in the y direction, as the figure 4.3. Then, we can get the formula of leaf area index (LAI) as follows [14]:/20/21(,)h d d dz x z dx LAI d -∂=⎰⎰(25)Where h is the height of the canopy , d is the distance between two branches, and (,)x z ∂ is the leaf area density function of the micro-body at the point (,)x z . x a is the function of leaf area which means the distribution of cross-section of the X direction, and Its value is a dimensionless, defined as follows: /2/21()d x d a x dx LAI d -=⎰ (26)()()x s a x d LAI C x =⋅⋅ (27)For the canopy which is not uniform in the horizontal direction, the distribution of its leaves is not entirely clear. We use s C to represent the distribution function of thedensity of leaves. s C can be expressed as a quadratic function or a Gaussiandistribution function.4.2.3 Transmission model of the solar radiationAssume that the attenuation of the solar radiation accords with the law of Beer-Lambert. The attenuation value at a point of canopy where the light arrives with a certain angle of incidence and azimuth is proportional to the length of the path, and the length can be calculated by Goudriaan function. Approximate function of the G function is shown as follows [15]：00(12)cos G G k G θ=+- (28)where k is a parameter decided by different plants.Take a micro unit in the canopy . Direct sunlight intercepted in this micro unit can be expressed in the following form:()(,)b b L dI I z G z dLAI =-Θ (29)Where b I means the direct sunlight, Θ is the solar zenith angle and L dL A I is the LAI on the path of light.Figure 4.4 Micro unit of the canopyThrough the integral and the chain rule, we can get the transmittance at the point (',')x z as follows:(,)[()]tan sin ()(',')exp()()h b b z G z a x z a z I x z I h ΘΘ∆Φ+=-⎰ (30)Average direct transmission rate:/2/21(')(',')'d b b d t z t x z dx d --=⎰(31)Different with the direct light, the scattered light in all directions is intercepted by theleaf surface from the upper hemisphere. The irradiance d dI of scattering at the point(',')x z can be expressed as follows:'cos d d b dI i t d ωθ=⋅⋅ (32) where d i is the illumination intensity of scattered light from a given direction.The transmission rate of the sun scattered light is as follows:2/2'00(',')1sin cos ()d b d I x z t d d I h ππθθθϕπ=⎰⎰ (33) Average scattering transmission rate:/2/21(')(',')'d d d d t z t x z dx d --=⎰(34)Global solar radiation reaching the canopy with a given depth z :(')()[(1)(')(')]d b d d I z I h k t z k t z ---=-+ (35)According to eq.(30) and eq.(33), the maximum depth the solar radiation can reach has a major link with h and θ. The shape of leaves have a relationship with photosynthesis. So, the h and θ of branches do have an influence on the leaves. Results showed that the light level and light utilization of high stem and open ce nter shape as well as small and sparse canopy shape were better than others ．Double canopy shape ，spindle shape and center shape took second place ，while big canopy shape had the lowest light distribution [16]．5. Is leaf shape related to tree structure?Due to internal and external factors, there are many kinds of tree shapes in the nature. For example, the apple's tree shape is semi-ellipsoidal, the willow's is hemispherical, the peach's likes a cup, the pine's likes cone and so on. The shape of their leaves varies. The leaf shape of apple is oval, pine's is needle, and the Indus's is palm. Is leaf shape related to tree shape? Even for the same species, there are many kinds of tree shapes. For instance, Small canopy shape, Open center shape, Freedom spindle shape, high stem and open center shape, Double canopy shape and so on. The sizes of leaves are different. Does the tree profile/branching structure effects the leaf shape?5.1 The experiment for one speciesTo solve this problem, we first consider the relationship of one tree species such as apple, which is semi-ellipsoidal. According to the second question, the hemispherical model is further extended to the semi-ellipsoidal model. Every tree individual shows irregularly because of natural and man-made factors, which reflects on the diversity of the crown in the canopy height direction of the changing relationship between the crown and canopy height. Figure 5.1 shows the relationship.Figure5.1 Schematic diagram of tree crown contourWhere 0h means the truck high, 1h means the crown high, 8710,,,,d d d d ⋅⋅⋅ meanscrown diameter. Tree shape with the scale change can be characterized in fractal dimension. Across the same scales, a fixed fractal dimension indicates the boundary shape's self-similarity; on different scales, the change of fractal dimension means that different processes or limiting factor has superiority. (Wiens, 1989)[17] According to the application of BP network and fractional dimension, we describe the tree shapes.[18]According to Li Guodong and Zhang Junke's work which detected and evaluated in different tree shapes of ‘Fuji’ apple. We choose three kinds of tree shapes to study; they are Small canopy shape, Open center shape and Freedom spindle shape as figure5.2 shows.。
