PRINCIPLES OF CROP GROWTH

SIMULATION MODELLING

Potential production
 The Rice System and boundaries –
Potential Production Situation

Radiation, CO2, O2

Temperature
 PROCESSES OF CROP GROWTH
A. GROWTH

Photosynthesis (CO2 assimilation)

CO2 + H2O + solar energy => CH2O + O2
Respiration (maintenance, growth)
CH2O + O2 => CO2 + H2O + chemical energy
Assimilation  respiration = net growth rate

B. DEVELOPMENT
Development rate = f(temperature)
=> growth period
=> partitioning

C. BIOMASS PRODUCTION = growth rate x growth period

 A. CROP GROWTH RATE

Daily gross crop assimilation rate

(kg CH2O ha1 d1)
 CO2 assimilation

I. CO2 assimilation of a single leaf

• Individual leaf light response curve

II. Canopy CO2 assimilation

• Integration over leaves in canopy (depth, z)

• Integration over leaves in time (day, t)

III. Leaf area growth in time

I. CO2 assimilation of a single leaf
Cross section of a leaf through a stoma (a), indicating components of resistances to CO 2 diffusion (b).
 radiation
(Wm-2) PAR
per mm
50%
1.0

.5

500 1000 1500

ultra violet visible infra-red wavelength (nm)

Spectral composition of global radiation, discriminating between direct radiation (clear area)
and diffuse (i.e. indirect) radiation scattered by clouds (hatched area).
PAR = photosynthetically active radiation.
(Source: Monteith and Unsworth, 1990. Principles of environmental physics. Edward Arnold,
London, 291 p.)
Schematic representation of a leaf chamber for the
measurement of CO2 assimilation.
Leaf light response curve:
Fg (kg CO2/ha leaf/hr)
80

Fm C4 crop

60

Fm C3 crop
40

Fg C3
20
 Fg C4

APAR (J/m2/s)
0
0 100 200 300 400

Fg = gross rate of CO2 assimilation

Fm = maximum rate of CO2 assim. at high light intensity
 = initial light use efficiency
APAR = Absorbed Photosynthetic Active Radiation

Fg = Fm (1e(APAR / Fm))

C3 crops C4 crops
Potato, sugar beet, Sorghum, maize,
wheat, rye, barley, millet, sugarcane,
rice, rape, soy bean, rhodes grass, sisal,
broad beans, almost mangrove
all trees

Characteristic values of Fm and 

Fm 
C3 crops 40 (15 - 50) 0.45
C4 crops 70 (30 - 90) 0.45

Fm : kg CO2 ha leaf1 d1

 : (kg CO2 ha leaf1 h1)(J m2 s1)1
 net assimilation rate net assimilation rate net assimilation rate
[kg (CO2) ha-1 h-1] [mg (CO2) m-2 s-1] [kg (CO2) ha-1 h-1] [mg (CO2) m-2 s-1] [kg (CO2) ha-1 h-1] [mg (CO2) m-2 s-1]
100 100 100
C3 A C3 B C3 C
C4 C4 C4

75 2 75 2 75 2

50 50 50

1 1 1

25 25 25

0 0 0 0 0 0

0 200 400 600 0 200 400 600 0 10 20 30 40 50

PAR [J m-2 s-1] internal CO2 [ppm] temperature [oC]

The effect of light (a) CO2 concentration (b) and temperature (c) on net assimilation
rate of a C3 (solid line) and a C4 (dashed line) species. Assimilation rates in (b) and
(c) refer to light saturated values. In (b) CO2 concentration in the stomatal cavity is
used to avoid confounding stomatal to changes in external CO 2 concentration.
Effect of N in leaf on photosynthesis

Van Keulen and

Seligman, 1987
II. Canopy CO2 assimilation
Diffuse light
Atmospheric scattering

Canopy reflection (5%)

Direct light

Weather station: radiometer

Incident radiation
direct radiation

diffuse radiation
II.1 Integration over leaves in canopy (depth, z)
Light and single leaf

Light and multiple

leaves
 A
depth [cm]
0

10

15

20

25

30

35

40

45

50

55

60

0.0 0.2 0.4 0.6 0.0 0.2 0.4 0.6 0.8 1.0

LAI rel. light intensity

The distribution of leaf area over the depth (I.e. LAI per layer of 5 cm) and of the light intensity in the
crop, relative to the light intensity above the (sugar beet) crop (a). The relation between relative light
intensity and cumulative LAI calculated from the top downwards (b). LAI = leaf are index [in m 2 (leaf
surface) m-2 (ground surface)].
Extinction of radiation in the canopy

PARd = PAR0 e( LAId)

PARd = PAR at depth d in canopy

PAR0 = PAR above crop canopy
LAId = Leaf Area Index above d
 = extinction coeff. (0.5 - 0.7)
Canopy (leaves) architecture
II.2 Integration over leaves in time (day, t)
 PAR Agross
[J m-2 s-1] [kg (CO2) ha-1 h-1]

Agross
PAR
400 80 400 Clear day 80

Average day

300 60 300 60

200 40 200 Overcast day 40

100 20 100 20

0 0 0 0
2 6 10 14 18 22 2 6 10 14 18 22
time [h] time [h]

Time course of PAR and crop CO2 assimilation rate on an average (left) and a clear
and overcast day (right) in June in the Netherlands.
 Summary calculation of daily canopy
gross CO2 assimilation rate:

• At any depth in the canopy, at any time of

the day (and latitude, season), calculate
gross leaf CO2 assimilation rate from ‘leaf
light response curve’ and PARd.

• Integrate over depth (LAId) of the canopy

• Integrate over day length (time of day)

Canopy assimilation = t z (leaf assimilation)

So far, gross CO2 assimilation rate was
expressed in kg CO2 ha1 d1
Since:
CO2 + H2O + solar energy => CH2O + O2
Molecular weight of CO2 is 30
Molecular weight of CH2O is 44

To calculate daily gross accumulation

rate of assimilates:
1 kg CO2 ha1 d1 CO2 assimilation rate
= 30/44 kg CH2O ha1 d1
Rice canopy assimilation rate, Los Baños, Philippines

Month f Aclear Aovercast Aaverage Agross

J 0.47 616 252 445 303
F 0.34 674 281 540 368
M 0.37 734 311 577 394
A 0.28 781 334 656 447
M 0.42 800 342 608 414
J 0.52 804 342 564 384
J 0.59 803 343 532 362
A 0.62 791 339 511 348
S 0.58 757 323 505 345
O 0.54 698 293 479 327
N 0.54 633 261 432 295
D 0.54 600 244 408 278
------ kg CO2 ha1 d1 ------ kg CH2O ha1 d1
III. Leaf area growth in time
 Leaf Area Index (ha leaf ha-1 soil)

6 Leaf death

4 Linear phase

3 Seedbed

2
Exponential
phase
1

0
0 10 20 30 40 50 60 70 80 90 100 110 120
TIME
Exponential phase: leaf area growth determined by
temperature
LAIt = LAI0 e(RGRL Tsum)
RGRL = Relative Growth Rate Leaves (°Cd)-1
Tsum = Temperature sum since emergence (°Cd)

Linear phase: leaf area growth determined by assimilates

available, as expressed in weight of leaves
LAIt = SLA Wlvg,t
SLA = Specific Leaf Area (ha leaf kg-1 leaf)
Wlvg = Weight of green leaves (kg ha-1 soil surface)
NET CROP GROWTH RATE

Crop respiration
1.0

0.8

0.6
Growth respiration

0.4

Maintenance
0.2
respiration
22

18

14

10

16 20 24 4 8 12 16 20 24 4 8 12 16

26 April 27 April 28 April

Figure 2.24 CO2 efflux of an enclosed sward of perenial ryegrass, at ambient temperature
during two days of darkness.
(a) ______ : continuous record of CO 2 efflux; --------- : maintenance respiration, calculated as
the mean rate of CO2 efflux after 40-46 hours of darkness, adjusted for the effect of
temperature.
(b) continuous record of air temperature in closure.

Source : Jones, Leafe, Sites & Collett, 1978. Ann. Bot. (42):693-703.
 Maintenance respiration
• Resynthesis components (enzyme turnover: 10% d1)
• Concentration gradients
• Metabolic activity

Maintenance respiration rate: Rm = f(composition, weight)

For lowland rice (IR72):

Rm = 0.02 Wleaf + 0.015 Wstem + 0.01 Wroot + 0.03 Wstorage

(in kg CH2O (kg CH2O)1 d1 )

and at any temperature:
Rm,t = Rm Q10 [ (T Tref) /10 ]

with Q10 = 2, and Tref = 25 ºC

 Growth respiration

Sugars ( CH2O) => structural matter

• Uptake nitrogen and minerals
• Transport of sugars
• Synthesis end-product

Conversion efficiency:
Eg = f(chemical composition)
in (kg dry matter) (kg CH2O)1
Conversion efficiency for chemical compounds

Carbohydrates 0.826
N-compound from NO3 0.404
N-compound from NH+4 0.616
Organic acids 1.104
Lignin 0.465
Lipids (fats) 0.330
 Growth respiration

Composition (%) and conversion efficiency for rice:

Leaf Stem Root Panicle

Carboh. 52 62 56 76
Protein 25 10 10 12
Fats 5 2 20 2
Lignin 5 20 20 6
Org. ac 5 2 2 2
Mineral 8 4 10 2
Eg 0.75 0.75 0.75 0.68
 Net crop growth rate

Daily rate of increase in structural

dry weight of a crop (dW):

dW = Eg (Agross  Rm W)
in: kg dry matter ha1 d1

where:
• Eg = conversion efficiency (kg dry matter (kg CH2O)1)
• Agross = gross CO2 assimilation rate (kg CH2O ha1 d1)
• Rm = maintenance respiration (kg CH2O) (kg CH2O)1 d1
• W = dry weight of the crop (kg ha1)
 Net crop growth rate
Mean values of Rm and Eg, at 20 °C

Rm Eg
Root/tuber crops 0.010 0.75
Cereals (rice) 0.015 0.70
Protein-rich seed crop 0.025 0.65
Oil-rich seed crop 0.030 0.50
 For rice in Los Baños, Philippines

dW = Eg (Agross  Rm W)

Dry season – April:

dW = 0.70 (447  0.015  5000)
= 261 (kg dry matter) ha1 d1

Wet season – October:

dW = 0.70 (327  0.015  5000)
= 176 (kg dry matter) ha1 d1
 Rice IR72, IRRI farm 1991-1993, Los Baños

kg DM
25000
266 kg DM ha -1 d-1
20000 Dry season

15000

177 kg DM ha -1 d-1
10000
Wet season

5000

Day of year
0
0 50 100 150 200 250 300 350
 Rice canopy assimilation rate, Beijing, China

Month f Aclear Aovercast Aaverage Agros Anet

J 0.47 333 120 233 159 59
F 0.34 445 169 351 239 115
M 0.37 586 233 455 310 165
A 0.28 737 304 616 420 241
M 0.42 843 354 638 435 252
J 0.52 892 377 624 426 245
J 0.59 873 368 575 392 222
A 0.62 788 329 503 343 188
S 0.58 652 264 427 291 151
O 0.54 497 193 333 227 106
N 0.54 364 133 239 163 62
D 0.54 304 107 198 135 42
------ kg CO2 ha1 d1 ------- kg CH2O ha1 d1
 Rice, Changle (Beijing), 2001, China

25000 kg DM

20000 232 kg DM ha -1 d-1

JD305
15000
HD502
10000
HD297

5000

Day of year
0
0 50 100 150 200 250 300 350
B. CROP DEVELOPMENT

Development rate and

assimilate partitioning
Thermal Unit (or temperature sum):

anthesis maturity
 (T - T0) dt and  (T - T0) dt
emergence anthesis

Expressed in day degrees (°C d)

where T = average daily temperature
T0 = base temperature (0 - 10 °C)

Per crop (and cultivar), there is constant TU between

emergence-anthesis, and anthesis-maturity
 Rice variety IR72

Temperature sum (°C)

2500

2000

1500

1000

500

Flowering Maturity
0
0 0.5 1 1.5 2

Development stage
Development rate is inverse of number of degree
days between development stages (°Cd 1)

Period DVS IR72 HD502

Juvenile 0.0-0.4 0.001161 0.000506
Photoperiod- 0.4-0.65 0.000064 0.000758
sensitive
Panicle 0.65-1.0 0.001152 0.000769
formation
Grain filling 1.0-2.0 0.000504 0.001417
 Partitioning of assimilates
Assimilate partitioning is function of
development stage (DVS) of crop.
Example for fraction of assimilates (growth)
partitioned to the shoot (FSH), for rice:

DVS FSH
0.00 0.50 E.g. at development stage of
0.43 0.75 0.43 => 75% of daily growth
1.00 1.00 goes to the shoot, and 25%
to the roots.
2.00 1.00
Partitioning of assimilates; Rice IR72
Fraction daily dry matter increase
1
0.9
ROOT
0.8
0.7
SHOOT
0.6
0.5
SHOOT
0.4
0.3
0.2
0.1
0
0 0.5 1 1.5 2

Development stage
Partitioning of assimilates; Rice IR72
Fraction daily dry matter increase
1
0.9
STEM
0.8
0.7 STORAGE ORGAN
0.6
0.5
0.4
0.3
LEAF
0.2
0.1
0
0 0.5 1 1.5 2

Development stage
C. BIOMASS PRODUCTION

Total crop growth simulation

 Crop model Potential Production

Light

Temperature N leaves


Photosynthesis LAI
Development
rate

Development
stage Partitioning

Leaves

Assimilate Stems
pool Biomass

Panicles

Maintenance Growth
Roots
respiration respiration
Reduced grain yield through spikelet
sterility; effect of temperature

Horie (1993)
 Input

1. Weather data: daily temperature, radiation

=> Weather data file
2. Management: emergence date, density, etc
=> Experiment data file
3. Crop characteristics: Fm, partitioning, etc
=> Crop data file
 Output

• Time course of leaf area index, biomass

of various crop organs
• Yield, yield components
IR72; Los Baños; DS 1992; 225 kg N ha1

7
LAI simulated
6

LAI measured
5

Run 3, LAI
3 Run 3, LAI_OBS

0
0 10 20 30 40 50 60 70 80 90 100 110 120
TIME
 kg dry matter ha-1.
18000

Total biomass
16000

14000

12000
Panicle biomass
10000 Run 3, WAGT
Run 3, WAGT_OBS
Run 3, WLVG
8000 Run 3, WLVG_OBS
Run 3, WSO
Run 3, WSO_OBS
6000

4000

2000 Green leaf biomass

0
0 10 20 30 40 50 60 70 80 90 100 110 120
TIME