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Dynamics of change
1.1
nx
d xn
n 1
dx
x n1
C , n 1
n 1
where both n and C are constants. But what is the point of calculus? More specifically, how
do we use or apply calculus in agriculture?
Consider a hypothetical growing crop. Table .1 shows that the plants growth rate
remained constant at 10 g day-1 from day 10 to 20. This means that in this period the plant
would consistently gain weight by 10 g per day. So after ten days, the plant must gain (10 g
day-1 10 days) or 100 g. Hence, provided the plant is growing at a constant rate, there is a
simple relationship between the cumulative (total) weight change (W) and the rate of growth
(wr):
W wr t
where t is the time interval. In Table .1, we can confirm that in ten days from day 10 to 20
(t = 10) the plant weight increased by 100 g.
And if we plot the growth rate (which is constant) against time, we will get a
horizontal line at 10 g day-1 (Fig. .1a). Furthermore, the area under this line is actually the
cumulative weight change. As mentioned earlier, the cumulative weight change is the growth
rate multiplied by the time interval. The area under the line in Fig. .1a is a rectangle with a
height of 10 g day-1 (i.e., the growth rate) and a width of (20 10) days (i.e., the time
interval). Consequently, this gives the area under the line from day 10 to 20 as the area of the
rectangle:
(a)
Day
10
10
300
11
10
310
12
10
320
13
10
330
14
10
340
15
10
350
16
10
360
17
10
370
18
10
380
19
10
390
20
10
400
(b)
Fig. 2.1. Plotted constant growth rate and plant weight against time
Since the growth rate remained constant, the plant weight must increase linearly with
time (Fig. .1b). And if we take the slope of this linear line, we will obtain 10 g day -1, which is
the rate of weight change or growth. Recall that for a linear line, its slope (m) is calculated as
y2 y1
x2 x1
where (x1, y1) and (x2, y2) are two pairs of points along the linear line. From Table .1, let us
take the two pairs of points as (15, 350) and (20, 400), so that we determine the slope of the
line as
m
400 350 g 50
10 g day -1
20 15 days 5
In other words, the instantaneous rate of weight change is the slope of the line (or curve) for
the graph of plant weight against time.
To generalise: the slope of a curve gives us the instantaneous rate of change, whereas
the area under a rate of change curve gives us the cumulative change. As we will later
discuss, the slope of a curve and the area under a curve are concerns of differential calculus
and integral calculus, respectively.
1.2
In the previous section, calculations were simple because the rate of plant growth was
constant. But when the rates of change are not constant, the use of calculus then becomes
essential. Consider a second example using the same hypothetical crop but it grew instead at
a variable (non-constant) rate.
Table .2 shows the measured daily plant weight from day 10 to 20. Using the data in
Table .2, we can show that the plant weight is related to time by the following quadratic
function:
f (t ) t 2 t 190
where f (t) is the weight of the plant (g) at time t (day). For example:
at day 10: f (10) 102 10 190 300 g
at day 11: f (11) 112 11 190 322 g
10
300
11
322
12
346
13
372
14
400
15
430
16
462
17
496
18
532
19
570
20
610
Fig. 2.2. Plotted plant weight against time (variable growth rate)
But when the growth rate is variable, as in this second example, the weight of the
plant will increase in a non-linear manner with time. Because this curve is non-linear, its
slope will vary from point-to-point along the curve, in contrast to a linear curve. Finding the
slope of a curve at, say, point (a, b) is, by definition, finding the slope of the tangent line at
(a, b). But what exactly is a tangent line?
Fig. .3 shows point P on two curves. We have drawn an enlarged version of the box
around each point P. Notice that the portion of each curve within the boxed region looks
almost straight. With increasing magnification, the portion of the curve near P becomes
increasingly more exactly like a straight line. This straight line is called the tangent line to the
curve at point P.
f x h f x f x h f x
h
x h x
To let the slope of secant line approach the slope of tangent line, we move point Q
increasingly closer to point P, thereby h becomes increasingly smaller (but h is never zero).
In other words, by taking h sufficiently small, the slope of the secant line can be taken as the
slope of the tangent line with the desired accuracy. Mathematically, we write this as
f a
df a
f a h f a
lim
h 0
dt
h
where h0 means h approaches but never reaches zero, and f (a) is the derivative of the
function f (x) at x = a. In other words, f (a) is the slope of the tangent line at x = a. As
mentioned in the previous chapter, the derivative of the function f (x) is sometimes written as
dy/dx, df (x)/dx or D(x).
Fig. 2.4. Tangent line to the curve at point P approximated by the secant line through points P
and Q
Let us return to the example where the plant growth rate is variable. What is the rate
of plant growth at, say, day 15? To do this, we need to find the slope of the tangent line at t =
15. In other words, we differentiate the plant weight function f (t) = t2 + t + 190 at t = 15:
df 15
f 15 h f 15
lim
h 0
dt
h
2
15 h (15 h) 190 152 15 190
lim
h 0
h
lim h 31
f 15
h 0
31
where we see that as h0, f (15) approaches 31. Consequently, we can take 31 g day-1 as the
plant growth rate at day 15. Likewise, to determine the growth rate at day 20 is
df 20
f 20 h f 20
lim
h 0
dt
h
20 h 2 (20 h) 190 202 20 190
lim
h 0
h
lim h 41
f 20
h 0
41
where the growth rate at day 20 is 41 g day-1. To generalise, the growth rate of the plant at any
given time t is:
df t
f t h f t
lim
h 0
dt
h
t h 2 (t h) 190 t 2 t 190
lim
h 0
h
lim h 2t 1
f t
h 0
2t 1
So the derivative of the plant weight function f (t) = t2 + t + 190 gives the growth rate
function as f (t) = 2t + 1. Table .3 expands on the earlier Table .2 by showing the measured
daily plant weight as well as the calculated daily growth rates (i.e., 2t + 1) from day 10 to 20.
Table 2.3. Measured plant weight and its variable (calculated) rate of increase
Day
10
21
300
11
23
322
12
25
346
13
27
372
14
29
400
15
31
430
16
33
462
17
35
496
18
37
532
19
39
570
20
41
610
Fig. .5 shows that the growth rate increased linearly with time, where the line is
described by the function 2t + 1. We saw in the previous section that if we took the area under
the rate of change curve, we would obtain the cumulative change. In Fig. .5, the area under
the curve from day 10 to 20 is the combined area of the triangle (Area A) and rectangle (Area
B):
Area Triangle area (A) + Rectangle area (B)
1
2
20 10 41-21
20 10 21
= 310 g
which agrees with our measured weight gain of 310 g from day 10 to 20 (Table .3).
instantaneous rate of
du
change from (a, b) change at u
f u du
a
In our first example, the instantaneous rate of change was constant at 10 g day -1. Hence,
integrating
20
10
gives us the cumulative change of 100 g from day 10 to 20 (Table .1). But in our second
example, the rate of growth varied according to the function 2t + 1 so integrating this
function
20
2t 1 dt t
10
20
10
gives us a cumulative change of 310 g in the same ten-day period (Table .3).
1.3
In Fig. .6, the area under the curve ABCD can be approximated by dividing this area
into several rectangular strips, where each strip has a width of x (the symbol is
interpreted as very small). The area of each strip is approximately equal to f (x)x (i.e.,
height width for a rectangle area). And the sum area for all the strips gives the approximate
area of ABCD, or
S f (a ) x f (a x) x f (a 2 x ) x ... f (b) x
b
[2.1]
f ( x) x
xa
Notice that the last term f (b)x is the area of the last rectangle which lies just outside the
region ABCD, and that, in Fig. .6, the areas of the rectangles tend to underestimate the area
under the curve. The area of the first rectangle, for example, underestimates the area APRD
by the area APQ.
Fig. 2.6. Approximating the area under the curve by a series of equal width rectangles
Eq. .1, however, becomes increasingly more accurate if we take the width of each
strip to be increasingly smaller and so increasing the number of strips to cover ABCD. In
other words, as x approaches zero (x 0), the accuracy of Eq. .1 increases.
Mathematically, we write this as
S lim
x 0
xa
f ( x) x
We have learnt that the area under a curve ABCD is determined exactly by integrating
the function f (x) from x = a to x = b:
b
S f ( x)dx
a
S f ( x)dx lim
x 0
xa
f ( x) x
[2.2]
y1 y2 ... yn
n
n
n
y lim
f ( x)
lim
n
xa
f ( x)
y lim x a
lim
n 0
x 0
lim f ( x) x
x 0
xa
lim b a x
f ( x) x
x a
n x
x 0
b
1
f ( x)dx
b a a
1.4
xa
f ( x) x f ( x )dx .
Numerical integration
There are integrals that are impossible to evaluate analytically such as
2
exp x dx
There are no exact solutions to such integrals, and we must resort to numerical integration to
obtain their approximate solutions. There are many numerical methods available, but in this
book, we will discuss three common methods: midpoint, trapezoidal and Gauss.
1.4.1
Midpoint method
We have actually discussed a slight variation of this method in the previous section
(Eq. .2). As described previously, we approximate the area under the curve by rectangles. The
area under curve is divided into n number of rectangular strips, each having an interval width
of x (Fig. .7). What is different, however, we select the midpoint for each interval, ck, which
it is determined by
ck
xk 1 xk
2
so that the area for a rectangle strip is determined by f (ck) x (note: k = 1, 2, , n). The sum
of all the rectangle areas is then the area under the curve, or
b
f ( x)dx f (ck ) x
k 1
Since (b a) = n x,
b
f ( x)dx
ba n
f (ck )
n k 1
[2.3]
Fig. 2.7. The midpoint method to approximate the area under the curve
The more rectangle strips we use (i.e., larger n and smaller x), the accuracy of the
midpoint method increases.
1.4.2
Trapezoidal method
Instead of using rectangles to approximate the area under the curve, trapezoids are
used instead in this method. We again divide the area under the curve into n equal intervals
with width x (Fig. .8).
Fig. 2.8. The trapezoidal method to approximate the area under the curve
We see that the sum of all the trapezoidal areas gives us the area under the curve.
From Fig. .9, the area of a trapezoid is
h h
area w 1 2
2
where w is the width of the interval x; and h1 and h2 are f (xi) and f (xi+1), respectively (i = 0,
1, , n). Consequently, the area under the curve is determined by
b
f ( xn1 ) f ( xn )
f ( x0 ) f ( x1 ) f ( x1 ) f ( x2 )
...
2
2
2
f ( x)dx x
a
f ( xn )
f ( x0 )
f ( x1 ) f ( x2 ) ... f ( xn1 )
2
2
f ( x)dx
a
f ( xn )
b a f ( x0 )
f ( x1 ) f ( x2 ) ... f ( xn1 )
n 2
2
[2.4]
Like the previous method, the accuracy of the trapezoidal method increases by
increasing the number of trapezoidal strips and decreasing the interval width x for each strip.
Gauss method
One of the most accurate numerical method is the Gauss-Legendre (or simply known
as Gauss) method. This method is also one the most common methods used in agriculture
work.
Consider the function y = f (t) which we wish to integrate over the interval t = -1 to 1.
We will approximate its solution as
1
f (t )dt wi f (ti )
[2.5]
i 1
where the coefficients w and t are the weight and abscissa, respectively. We wish to determine
all the weights wi and abscissas ti in such a way so that the linear additions of wi f (ti) for i =
1 to N is the exact solution of the integral of f (t) over [-1, 1]. A two-point Gaussian
integration means N = 2, and Eq. .5 is expanded to
1
[2.6]
Let us further approximate the function f (t) as a cubic polynomial which has a polynomial
degree (or order) of three:
f (t ) a0 a1 x a2t 2 a3t 3
Substitution into Eq. .6 produces
1
a0 a1t a2t
1
1
wt
3
11
w2t23 a3
2
3
w1t1 w2t2 0
w1 w2 2
We now have four unknowns (the weights w1, w2, and abscissas t1, t2) in four independent
equations which means they can be solved. Thus, it can be shown that for a two-point
Gaussian integration method:
t1 0.5773503, t2 0.5773503
w1 1.0, w2 1.0
At hindsight, approximating the function f (t) as a cubic polynomial is exactly right
for a two-point Gaussian integration. This is because it produces four unknowns in four
independent equations. So for a three-point Gaussian integration the function f (t) must be
approximated by a polynomial curve having a degree of five. This would produce six
unknowns (weights w1, w2, w3, and abscissas t1, t2, t3) in six independent equations.
To generalize this rule, an N-point Gaussian integration requires that the function f (t)
be approximated by a polynomial curve with 2N-1 degrees. As expected, the higher the N
points of integration the higher the accuracy of solution. This is because we are
approximating the function f (t) by an increasingly more flexible polynomial curve (i.e.,
increasingly higher polynomial degree). Nonetheless, the higher the N points of integration,
the higher the number of required computations as well. In agriculture, the three- or fivepoint Gaussian integration are often used. The abscissa xN,i and weights wN,i up to eight points
are tabulated in Table .4.
If we had the integral over an interval [a, b] instead of [-1, 1] we must normalize the
interval [a, b] to [-1, 1] so that if a x b and that -1 t 1 this means, using linear
transformation (Fig. .10), x is related to t by
x a bt
[2.7]
f ( x)dx
f ( a bt ) b dt
[2.8]
b wN ,i f ( a bt N ,i )
i 1
Table 2.4. Abscissas and weights for the N-point Gaussian integration
N
Abscissas, tN,i
Weights, wN,i
Abscissas, tN,i
Weights, wN,i
0.57735027
1.00000000
0.93246951
0.17132449
0.57735027
1.00000000
0.66120939
0.36076157
0.77459667
0.55555556
0.23861919
0.46791393
0.00000000
0.88888888
0.94910791
0.12948497
0.86113631
0.34785485
0.74153119
0.27970539
0.33998104
0.65214515
0.40584515
0.38183005
0.90617985
0.23692689
0.00000000
0.41795918
0.53846931
0.47862867
0.96028986
0.10122854
0.00000000
0.56888888
0.79666648
0.22238103
0.52553241
0.31370665
0.18343464
0.36268378
3
4
5
x dx
2
Solution
Midpoint method:
The interval width for each rectangle strip is given by
b a 1 0
0.25
n
4
Consequently, the midpoints ck for the four rectangle strips are:
Interval, (xk-1, xk)
Midpoint, ck
f (ck) = ck2
(0, 0.25)
0.125
0.015625
(0.25, 0.5)
0.375
0.140625
(0.5, 0.75)
0.625
0.390625
(0.75, 1.0)
0.875
0.765625
TOTAL
1.312500
Trapezoidal method:
As before, the interval width for each strip (trapezoidal) is 0.25, and applying Eq. .4 to
obtain the integral approximation as:
1
f ( x4 )
f ( x0 )
f ( x1 ) f ( x2 ) f ( x3 )
2
2
x dx 0.25
2
f (1.0)
f (0)
f (0.25) f (0.50) f (0.75)
2
2
0.25
02
1.02
0.252 0.502 0.752
2
2
0.25
0.3438
Gauss method:
Using Eq. .7 to normalize the interval from [1, 0] to [-1, 1] means
x a bt
1 0 1 0 t 1 1 t
2
2
x dx
0
1 0
1 1
2 2 t 2 dt
1
1 1 2
1 1 2
1
1 1
w
f
w
t3,2
w3,3
t3,3
3,1
3,1
3,2 f
2
2
2
2
2
2
2
1
0.5
2 0.5556
2
0.5556 0.1127 2 0.8889
0.8873
2
0.3334
Analytical solution:
The exact solution of the integral is
1
1 3
x dx 3 x
0
2
1
3
The error for the midpoint method is an underestimation by 1.562% of the exact solution,
whereas the trapezoidal method overestimates by 3.125%. The highest accuracy obtained is
by the Gauss method: a very small overestimation by 0.02% of the exact solution.
Example
Leaf photosynthesis can be described by the following equation, as given by
Goudriaan and van Laar (1994):
AL
Am kI 0 exp kL
Am kI 0 exp kL
[2.9]
where AL is the leaf photosynthesis (that is, the assimilation rate of CO 2 by a single leaf; g
CO2 m-2 leaf area s-1); Am is the maximum leaf photosynthesis rate (g CO2 m-2 leaf area s-1);
is the solar radiation conversion factor (g CO2 J-1); k is the canopy extinction coefficient for
solar radiation (unitless); I0 is the solar irradiance above canopy (W m-2 ground area or J m-2
ground area s-1); and L is the cumulative leaf area index from the canopy top to the leaf being
considered (m2 leaf area m-2 ground area).
Determine the canopy photosynthesis (that is, determine the cumulative
photosynthesis for all the leaves together). Use the following values: k = 0.5, total leaf area
index (LAI) = 3 m2 leaf area m-2 ground area, I0 = 200 W m-2 ground area, Am = 1500 g CO2
m-2 leaf area s-1, and = 12 g CO2 J-1.
Solution
Eq. .9 gives the photosynthetic rate of a single leaf. To determine the canopy
photosynthesis (all leaves), we need to integrate Eq. .9 over the entire canopy; that is,
integration is over the interval [0, LAI], where LAI is the total leaf area index. To integrate
complex equations like Eq. .9, it is convenient to use mathematical software to perform the
integration. The analytical solution to integrating Eq. .9 is
LAI
AT
LAI
AL dL
Am kI 0 exp kL
dL
Am kI 0 exp kL
[2.10]
A
Am kI 0
m ln
k
Am kI 0 exp kL
Substituting the given parameter values into Eq. .10 gives the canopy photosynthesis as
1270.6220 g CO2 m-2 leaf area s-1.
Let us try the 3-point Gauss integration method to integrate Eq. .9. We first need to
normalise the integration interval [0, LAI] to [-1, 1]. Using Eq. .7, we obtain
x a bt
so that
LAI
AL dL
LAI 0
dt
2
LAI
AL (t )dt
2 1
LAI
w3,1 AL t3,1 w3,2 AL t3,2 w3,3 AL t3,3
2
[2.11]
where
AL (t )
From Table .4, we obtain the values for the three abscissas t3,1, t3,2 and t3,3, and for each
abscissa, we determine the leaf photosynthetic rate AL (t):
Abscissa, t3,i
AL (t3,i)
Weight, w3,i
w3,i AL (t3,i)
-0.7746
604.784
0.5556
336.018
0.0000
411.3816
0.8889
365.6771
0.7746
261.7437
0.5556
145.4248
TOTAL
847.1199
LAI
AL dL
LAI
w3,1 AL t3,1 w3,2 AL t3,2 w3,3 AL t3,3
2
3
847.1199
2
1270.6800 g CO2 m -2 leaf area s -1
which is very close the analytical solution (an error less than 0.01%).
1.5
In this section, we will discuss how calculus can be applied to describe the growth of
a generic plant. We will begin by assuming that the plant has adequate supply of water and
nutrients (i.e., no water stress or nutrient deficiency), and the plant is free from pests and
diseases. The only limiting factor to plant growth is solar radiation which is the main source
of energy for various plant processes including photosynthesis.
Photosynthesis, aided with solar radiation energy, converts carbon dioxide and water
into carbohydrates (sugars). These carbohydrates are plant food; that is, substrates which
the plant will use for two purposes: maintenance and growth. Maintenance is a process
whereby the plant uses the substrates for its continual survival. In other words, the substrates
are used to maintain or sustain existing plant parts. Growth, on the other hand, is a process
whereby the substrates are used to build or synthesise new materials; thus, enlarging existing
plant parts or creating new ones. Consequently, the plant becomes bigger and heavier.
The plant growth model is as shown in Fig. .11. The plant weight consists of two
components: the storage weight and the plant structural weight. The storage weight, Ws, is
supported by the addition of newly produced substrates via the photosynthesis process. A
portion of the plants storage, kgWs, will be utilised for maintenance and growth, where of this
total (kgWs), Yg of it will be utilised for growth, and the remainder (1 Yg) for maintenance.
Of the total plant structural weight, Wg, kd of it will degrade and the substrates returned to the
storage, and ks of it will senesce (i.e., loss of materials due to increasing age).
In plant growth studies we are often interested in the plant dry weight (without its
water content) per unit ground area. A large portion of the dry matter of the plant is in the
form of organic compounds whose primary constituent is carbon which is assimilated in the
form of carbon dioxide through the plant leaves. Consequently, we need to express the plant
weight with respect to its carbon content (g C m -2 ground area). On average, the carbon
content in a plant is about 45% of the plant dry weight. So in the absence of carbon analysis,
plant dry weight equivalent to its carbon weight is 45% of the total dry weight.
Moreover, as shown earlier, Eq. .10 gives us the gross canopy photosynthesis, AT, in
units of g CO2 m-2 leaf area s-1 which should be expressed with respect to carbon; that is, in
units g C m-2 leaf area s-1. This conversion is done by multiplying AT by 12/44 (or about
0.273) since the atomic weight of carbon and molecular weight of carbon dioxide are 12 and
44 g, respectively. In other words, every 44 g of CO 2 has 12 g of C, so AT g CO2 will have
(12/44)AT g of C.
Fig. 2.11. A simple plant growth model (adapted from Thornley, 1977)
From Fig. .11, the rates of change in the plant storage and structure dry weight at time
t (in unit hour) are
dWs
12
kdWg k gWs 3600 LAI AT
dt
44
dWg
dt
[2.12]
[2.13]
where Ws and Wg are the dry weights for the substrates storage and plant structure,
respectively (g C m-2 ground area). Since time t is in unit hour, we need to multiply AT by
3600 (i.e., 60 s 60 mins) to express AT in units g CO2 m-2 leaf area hour-1. Further
multiplication by LAI is to convert AT into units g CO2 m-2 ground area hour-1; hence,
making it compatible with the dry weight units (which are expressed as g C per unit ground
area):
m-2 leaf area
g CO 2
3600 LAI AT -2
-2
g CO 2
-2
dW dWs dWg
dt
dt
dt
k gWs Yg 1 k sWg
10800
LAI AT
11
where W is the total plant dry weight (g C m-2 ground area). The cumulative increase in the
total plant dry weight, W, from time t1 to t2 is then obtained by integrating this rate of
change function over the period [t1, t2]:
t2
10800
W k gWs Yg 1 ksWg
LAI AT dt
11
t1
2
10800
k gWs Yg 1 k sWg t
LAI AT dt
11
t
[2.14]
where t is the time interval (t2 - t1), and kg ks and Yg are taken as constants. Calculations are
straightforward with the exception of the integration of the canopy photosynthesis AT. From
Eq. .10, canopy photosynthesis is
AT
Am
Am kI 0
ln
k
Am kI 0 exp k LAI
where both the leaf area index, LAI, and the solar irradiance above canopy, I0, are not
constants but both vary with time. For short periods (such as a day, or t = 24 hours), it is
reasonable to assume LAI remains constant. Consequently, we will determine the daily
increment in the total dry weight W, with t1 = 0 and t2 = 24.
The instantaneous solar irradiance I0 (J m-2 ground area s-1) is taken to vary with time t
(hour) according to a simple equation adapted from France and Thornley (1984) as
I0 t
I t ,d
1800 t ss tsr
t tsr
tss tsr
sin 2
t sr t t ss
[2.15]
where It,d is the daily total solar irradiance (J m-2 ground area day-1); and tsr and tss are the
times of sunrise and sunset, respectively (hour). Since there is no photosynthesis (AL = 0) for
periods before sunrise and after sunset, integration of canopy photosynthesis over the whole
day is equivalent to its integration over the period between sunrise and sunset; that is,
24
tss
tsr
AT dt AT dt
Hence, substituting Eq. .15 into Eq. .10 and its integration gives the daily canopy
photosynthesis, AT,d, as
AT ,d
10800 LAI Am
11 k
tss
Am k I 0 t
dt
Am kI 0 t exp k LAI
ln
tsr
[2.16]
recalling that LAI is unchanged during the day (thus, LAI is not a function of t for that
period). Unfortunately, Eq. .16 cannot be integrated analytically, and we must rely on
numerical integration methods.
For this case, we will choose the 3-point Gauss integration method to determine AT,d,
and Table .5 lists the parameters and their values to be used in the plant growth model. The
first step in the Gauss method is to normalise the integration interval [tsr = 6, tss = 18] to [-1,
1]. Using Eq. .7, we obtain
a bt
18 6 18 6 t 12 6t
2
so that
AT , d
18 6
Am k I 0 12 6t
10800 LAI Am
ln
1
1
64800 LAI Am
A(t )dt
11 k
1
64800 LAI Am
w3,1 A t3,1 w3,2 A t3,2 w3,3 A t3,3
11 k
[2.17]
where
A t ln
Am k I 0 12 6t
Am kI 0 12 6t exp k LAI
From Table .4, we obtain the values for the three abscissas t3,1, t3,2 and t3,3, and for each
abscissa, we determine A(t) using Eq. .15 with its relevant parameter values from Table .5:
Abscissa, t3,i
A (t3,i)
Weight, w3,i
w3,i A (t3,i)
-0.7746
0.1223
0.5556
0.0679
0.0000
0.5284
0.8889
0.4697
0.7746
0.1223
0.5556
0.0679
TOTAL
0.6055
64800 LAI Am
w3,1 A t3,1 w3,2 A t3,2 w3,3 A t3,3
11 k
64800 2 1500
0.6055
11 0.5
21.4017 g C m -2 ground area day-1
AT , d
[2.18]
From Eq. .14, the daily increment (t = 1) in the total plant dry weight is
W k gWs Yg 1 k sWg t AT ,d
1.98 20
0.75 1 0.05 130
21.4017
Wg Ws W 130 20 5.0017
[2.19]
Units
Value
Am
g CO2 J-1
LAI
kg
1500
12
0.5
-2
-1
1.98
-1
day
kd
day
0.10
ks
day-1
0.05
Yg
0.75
Ws
20
Wg
130
tsr
time of sunrise
hour
tss
time of sunset
hour
18
It,d
10000000
A more rigorous modeling approach to plant growth would be to include the death
rates of leaves (whether due to self shading or age) and roots, as well as the partitioning of
substrates to the various plant organs such as roots, stem, leaves and storage organs. In
particular, the partitioning of substrates to the leaves is important as it would allow us to work
out weight of the leaves, and in turn, the leaf area index, LAI; that is,
LAI Wleaves SLA
where Wleaves is the weight of green leaves (g m -2 ground area); and SLA is the specific leaf
area which is the leaf area per unit leaf weight (m 2 leaf area g-1). The specific leaf area varies
with the plant growth stage and it typically ranges between 0.005 to 0.03 m2 leaf area g-1.
In depth details about plant growth modeling are discussed in Goudriaan and van Laar
(1994), Teh (2006), and Thornley and France (2006).
1.5.1
Growth functions
We saw in the previous section that the rate of weight change, dW/dt, is often used to
indicate growth rate. dW/dt is known as the absolute growth rate (AGR). A more useful
measure of growth rate, however, is the relative or specific growth rate (SGR), which is given
by
SGR
AGR 1 dW
W
W dt
which shows that SGR is the increase in weight per unit time per unit weight (or the absolute
growth rate per unit weight). SGR is a better indicator of growth rate than AGR because SGR
expresses growth rate as the rate of weight increase per unit weight. Hence, SGR is a measure
of the efficiency of the plant or animal to produce new mass material.
If SGR remains constant throughout the plant growth period, it means
SGR
1 dW
u
W dt
where u is a growth constant; in this case, u is also SGR. When SGR is constant, the rate of
weight increase, dW/dt, is proportional only to the weight W. Integrating the above equation,
we obtain
1
W dW udt
W W0 exp ut
[2.20]
where W0 is the initial plant weight at time t = 0. Eq. .20 is the exponential function for
growth (Fig. .12).
Exponential growth functions are seldom used to describe long periods of growth
because as plants become larger, their specific growth rates tend to decrease until zero. In
other words, SGR is not a constant value. So let us formulate SGR as a function of the plant
weight in such a way so that as the plant weight increases, SGR will decrease. SGR is now
SGR
1 dW
W
u 1
W dt
Wm
where Wm is the maximum plant weight. We can see that as W approaches Wm, the quotient
W/Wm approaches 1, so SGR approaches zero. As done previously, we will integrate the
above equation to give
1
W 1 W Wm dW udt
W
W0Wm
W0 Wm W0 exp ut
which is known as the logistic or sigmoid function for growth (Fig. .12). Logistic growth
functions are typified by S-shaped curves.
Fig. 2.12. Plant growth as described by three types of growth functions: exponential (u = 0.2,
W0 = 1), logistic (u = 0.3, W0 = 1, Wm = 100), and Gompertz (u0 = 0.5, W0 = 1, D = 0.1086)
Another frequently used function for growth is the Gompertz function (Fig. .12). For
this function, SGR also decreases with increasing plant age, but the growth coefficient u is no
longer a constant but it varies exponentially according to:
u u0 exp Dt
where u0 is the initial u at time t = 0; and D is the decay constant rate of u. Hence, SGR is
now
SGR=
1 dW
u0 exp Dt W
W dt
W dW u0 exp Dt dt
1 exp Dt
W W0 exp u0
Fig. .12 shows the increase in plant weight as described by the exponential, logistic
and Gompertz functions. Of the three functions, the exponential function is the least
representative of plant growth. It often shows a very slow initial growth rate, followed by an
increasingly rapid growth rate. As mentioned in the previous chapter, exponential growth
functions are typically limited to describe the growth of very young seedlings or the growth
of embryos. The logistic growth function typically shows an S-shaped curve: growth rate is
initially slow, then increases for long periods before slowing down again towards the end of
the growth period. Compared to the logistic growth, Gompertz growth typically shows a
faster early growth rate, followed by a slower, almost linear approach to the asymptote.
1.6
Multivariable calculus
When a function depends on two independent variables, say, x and y, we express the
function as f (x, y). Suppose we want to know how the function f (x, y) changes when x and y
change. Instead of changing both variables simultaneously, we could change one variable (x)
while keeping the other (y) constant. In turn, we could now change the y variable, while
keeping the x variable constant. In this way, we will have a better understanding on how the
function f (x, y) depends on x and y. This is the idea behind the concept of partial derivatives.
The function f (x, y) has two partial derivatives (as it has two independent variables).
f ( x, y )
The partial derivative of f (x, y) with respect to x, written as
, is the derivative of f (x,
x
y) where y is kept constant, and the function f (x, y) depends solely on x. In the same way, the
f ( x, y )
partial derivative of f (x, y) with respect to y, written as
, is the derivative of f (x, y)
y
where x is kept constant, and the function f (x, y) depends solely on y. Note that for partial
derivatives, we use the symbol instead of d.
We can extend these principles to three or more independent variables. The function
f ( x, y, z )
f (x, y, z) depends on three variables: x, y, z, and the partial derivative
means the
x
partial derivative of the function f (x, y, z) with respect to x depends solely on x, with both
f ( x, y, z )
f ( x, y, z )
variables y and z kept constant. The other two partial derivatives:
and
y
z
are interpreted using the same principles.
Finding partial derivatives is no different from finding the derivatives of functions of
one variable, since by keeping all but one variable constant, determining a partial derivative
is the same as finding the derivative of one variable.
Example
Let f (x, y) = 5x3y2. Determine the two partial derivatives.
Solution
To determine
constant. So
5 x3 y 2
x
f ( x, y )
, we keep the y variable constant. In other words, 5y2 is a
x
3 5x
31
y2
15 x 2 y 2
To determine
f ( x, y )
, we keep the x variable constant instead; hence, 5x3 is now a constant.
y
Finally,
5 x3 y 2
y
2 5x y
3
21
10 x3 y
Recall that the converse of differentiation is integration. When a function f (x) is
integrated over an interval x = a and x = b, we obtained the cumulative change in that period.
The same principle applies for a multivariable function like f (x, y). To determine the
cumulative change, we need to perform a double integration over intervals [x = a, x = b] and
[y = p, y = q]. Mathematically, this is expressed as
qb
f ( x, y)dxdy
pa
The integrals are called iterated integrals because they are evaluated in more than a single
step, from inside out; that is,
q b
p a
f ( x, y )dx dy
f ( x, y)dx
The result of this first integration is a function of the y variable, and in the second step, this
resulting function is integrated over the limits as indicated by the outside integral sign (i.e.,
over y = p and y = q). The order of the differentials dx and dy indicates which integration is
performed first.
Example
43
Evaluate
2x y dxdy .
21
Solution
The order of the differentials is dx then dy, indicating that the integration is over the
x-interval first [1, 3] (the inside integral) then over the y-interval [2, 4] (the outside integral).
In other words,
43
4 3
21
2 x y dxdy
2 x y dx
1
dy
where the first integration step is with respect to x (the y variable is treated as a constant):
3
2 x y dx
1
x 2 xy
3
1
8 2y
and the resulting function is for the next integration step which is now with respect to y:
4
2
8 2 y dy 8 y y
2
4
2
28
43
Hence,
2 x y dxdy 28 .
21
We have actually used double integration in the previous section (1.5) on plant growth
modeling. We learned that to determine the daily canopy photosynthesis, we first need to
integrate the leaf photosynthesis function (Eq. .9) over the entire total leaf area [0, LAI]
which will give us the canopy photosynthesis function (Eq. .10). Eq. .10, however, will only
give us the canopy photosynthesis at an instantaneous moment. To determine the canopy
photosynthesis for the entire day, we must integrate Eq. .10 over the whole day, which is
equivalent to integrating the function from the time of sunrise to sunset (since there is no
photosynthesis for periods before sunrise and after sunset). The daily canopy photosynthesis
function is as shown in Eq. .16 which can be expressed in units g CO2 m-2 leaf area day-1 as
tss LAI
tsr 0
Am kI 0 t exp kL
dL dt
Am kI 0 t exp kL
where I0(t) is from Eq. .15, and the multiplication by 3600 is because time t is in unit hour.