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This is a linear, autonomous differential equation since the right-hand side is linear in N
and does not depend explicitly on time t. This can be generalized to a system of linear,
autonomous differential equations. In matrix form, a system of differential equations can
be written as
(1)
x1 (t )
a11
x (t )
a
d 2
21
dt
x n (t )
a n1
a12
a 22
an2
a1n x1 (t )
a 2 n x 2 (t )
a nn x n (t )
Here, the matrix A aij is an n n square matrix with constant and real coefficients.
We denote by x(t) the vector consisting of the functions xi(t), for i=1,2,,n.
The carbon and nitrogen model by gren and Bosatta
gren and Bosatta1 (1996) published a simple compartment model of carbon and
nitrogen dynamics that describes the change of carbon and nitrogen content of litter that
is decomposed. The litter has carbon content, denoted by C, and nitrogen content,
denoted by N. The decomposer takes up food (litter) and its biomass, B(t), grows
according to the equation
dB
production death P (t ) M (t )
dt
Not all of the food the decomposer takes up is converted into biomass; a certain fraction
is respired. We define the production efficiency e0 as
e0
production
0,1
uptake
We assume that the decomposer has fixed carbon and nitrogen concentrations, denoted by
fC (gC g-1) and fN (gN g-1), respectively. We assume that carbon and nitrogen are
incorporated into biomass at rates f C P and f N P , respectively. If we denote the rate of
1
gren, G.I. and E. Bosatta (1996) Theoretical Ecosytem Ecology. Cambridge University Press.
-1-
respiration
litter
decomposer
uptake
production
B
death
mineralization/
immobilization
decomposer
litter
uptake
production
B
death
We assume that nitrogen uptake follows carbon uptake passively, that is, it is equal to the
carbon uptake rate times the nitrogen to carbon ratio (N:C) of the substrate.
Nitrogen uptake
fC P N
e0 C
The difference between nitrogen uptake and production can be positive or negative. If it
is positive, decomposers release nitrogen (mineralization); if it is negative, decomposers
need to import nitrogen from the surroundings (immobilization).
To complete the dynamics, we need to include death of the decomposer. We
assume that both carbon and nitrogen of the decomposer are lost to the litter pool at rates
f C M and f N M , respectively.
This yields the following set of equations for the carbon and nitrogen dynamics of
the litter
-2-
We make the following assumptions: (1) mortality rate is equal to production rate, that is,
M P , and (2) production rate is proportional to the carbon content of the litter, that is,
P uC , where u is the constant of proportionality. Then,
1 e0
dC
f C uC
dt
e0
(2)
f u
dN
C N f N uC
dt
e0
If we define
(3)
fCu
, f N u , and 1 e0 , then this can be written
e0
dC
C
dt
dN
N C
dt
-3-
we approximate
with y y 0 when x x0
y ( x x ) y ( x )
dy
by the difference quotient
. This yields
dx
x
y ( x x) y ( x)
dy
x y ( x) g ( y )x
dx
C (t t ) C (t ) C (t ) t
N (t t ) N (t ) N (t )t C (t )t
d 2
21 a 22 a 2 n x 2 (t )
(6)
dt
x n (t )
a n1 a n 2 a nn x n (t )
-4-
dx
Ax ? Lets compute the left-hand side and the
dt
and Ax e t Aw
After equating the two expressions and canceling e t on both sides, we find
Aw w
x(t )
ci e t wi
i
i 1
-5-
Note that we assume that the eigenvector is different from the zero vector. An eigenvalue
though can be 0, it can even be a complex number. To find eigenvectors and eigenvalues,
we need to solve the equation
Aw w
Solving this equation will yield values for . For each value of , we can then find an
eigenvector w using Equation (8).
We assume now that all eigenvalues of the matrix A are distinct and real. In this
case, all the eigenvectors can be computed by solving Aw w for each eigenvalue; the
eigenvectors are then real.
Task 1:
Show that the eigenvalues of A are 1 and 2 with corresponding
eigenvectors
w1
and w2
1
Use (7) to show that the solution with initial conditions C(0) and N(0) is given by
C (t ) C (0)e t
N (t ) N (0)e t
C (0) t
e
e t
(1 )
-6-
-7-
f C uC
dt
e0
f u
dN
C N f N uC
dt
e0
1 e0
f C uC (t )t
e0
N (t t ) N (t )
fCu
N (t ) t f N uC (t )t
e0
-8-
A9: =A8+$H$4
B9: =B8-(1-$A$4)*$B$4*$D$4*B8*$H$4/$A$4
C9: =C8-$B$4*$D$4*C8*$H$4/$A$4+$C$4*$D$4*$H$4*B8
D9: =C9/$G$4
E9: =B9/$F$4
F9: =C9/B9
Drag Row 9 down to about 3000 time steps and graph N/N0, C/C0, and N/C as functions
of time in one coordinate system where N0 and C0 are the nitrogen and carbon litter
contents at time 0.
Step 3
Solve the system of differential equations numerically using the Euler method for the
following set of N:C ratios: (a) N:C=0.01, (b) N:C=0.1, and (c) N:C=0.3. Graph N/N0,
C/C0, and N/C as functions of time in one coordinate system for each of the three cases.
Homework (due ____________________)
Step 4
Explain the following statements using your graphs from Step 3: (A) When N:C ratio of
the food is less than that of the microbes, the microbes take up nitrogen from the
surrounding soil. (B) When the N:C ratio of the litter is equal to that of the microbes, the
decline of N/N0 and C/C0 are identicalthe food is perfectly balanced. When N:C ratio
of the food is higher than that of the microbes, the microbes need not take up any
nitrogen from the surrounding environment.
-9-
dX
L kX
dt
Where X (t ) denotes the amount of organic carbon in dead organic matter per square
meter of ground surface (measured in grams). Equation (6) in Olsons paper is identical
dC
C ).
to the carbon decay model in this module (Equation 3,
dt
(a) Set L 0 in Equation (8) and solve the resulting equation with the initial condition
X (0) X 0 (use paper and pencil). What is the fraction X (t ) / X (0) that remains at time t?
1
to solve Equation (8)
365
numerically in EXCEL for two cases: (i) continuous litter fall (case 1) and (ii) discrete
annual litter fall (case 2). In each case assume that the total amount of litter that is added
per year per square meter of ground surface is 200 grams. Assume that k=1 and X(0)=0,
and run each model long enough so that the model has time to stabilize (i.e., the longterm pattern becomes clear). In your write-up produce a graph of X (t ) for each case.
(b) Use the Euler approximation with time step t
- 10 -