SAMARA: A crop model for
simulating rice phenotypic
plasticity
Michael Dingkuhn, Uttam Kumar,
Ma Rebecca Laza, Richard Pasco, Gemma T. Narisma, Faye A. Cruz
IRRI, CIRAD, Manila Observatory
Hypotheses
• Simulate morphological plasticity and its effect on yield
– Resources (Light and water)
– Management (crop establishment, population, water)
• => GxExM interactions on phenotype and yield
• => Ideotypes
• Competition among plants and organs drive plasticity
• Compensatory plasticity
• Plant structure: arborescent system of sinks
• Sinks compete => feedback on number/size of new sinks
Supply vs. Demand driven growth
PAR
+ H20
+ NPK
= Yield
Biomass
Soil water reserve
Agronomic angle:
Supply
(assimilation) driven system
Botanical angle:
Demand
(organogenesis) driven system
Meristem
control of
development
SAMARA
Ic, state variable driving phenological and morphological adjustments
Ic = Supply/demand
Proxy for sugar signaling
Ensures source-sink
equilibrium as the
architecture unfolds
Supply = daily [Pn-Rm]
Demand = daily aggregate
growth commitments
Ic controls:
- Tiller outgrowth (Ic>0.5)
- Tiller mortality (Ic<1)
- Leaf mortality (Ic<1)
- Reserve formation (Ic>1)
- Reserve mobilization (Ic<1)
- Organ size (Ic << 1)
SAMARA calibration/validation for
IR72: 14 environments
Calibration
Validation: Biomass, Grain yield, tillering
Also simulated:
YC1
Plants m
-2
YC3
Panicles
tiller
-1
YC4
Spikelets
panicle
-1
YC5
Grains
spikelet
-1
YC2
Tillers
plant
-1
YC6
Weight
grain
-1
YC(N) on field area basis
YC(N+1
)
/
YC(N)
Variation of
YC(N+1) to attain
constant value on
ground-area basis
o
1
0
0
%
compe
nsa
tion
YC(N) on field area basis
YC(N+1
)
/
YC(N)
o
Compen-
sation
x
Full compensation
Partial compensation
x
x
80%
105%
Tillering
Surviving,
fertile tillers
Panicle
size
Spikelet
fertility
Grain filling
Compensatory plasticity, observed and simulated:
Case of increasing population
Simulated
Panicles m-2
300
400
500
600
700
Fi
lle
d
gr
ai
ns
p
an
ic
le
-1
40
60
80
100
Dry season, 25 hills m-2
Dry season, 100 hills m-2
Wet season, 25 hills m-2
Wet season, 100 hills m-2
D S: 3 7834 g rains m -2 W S: 2 6489 g rain s m-2 Compensation WS: 115% Compensation DS: 141%
Simulated
Panicles m-2
300
400
500
600
700
S
pi
ke
le
ts
p
an
ic
le
-1
40
60
80
100
120
W S: 3 018 4 sp ikele ts m-2 DS : 50 825 sp ikele ts m-2 Compensation DS: 154% Compensation WS: 57%Simulated
Hills m-2
25
50
75
100
P
an
ic
le
s
hi
ll-1
0
5
10
15
20
DS: 4 49 panicle s m-2 WS: 3 53 panicle s m-2 Compensation DS: 96% Compensation WS: 85%Observed
Panicles m-2
300
400
500
600
700
S
pi
ke
le
ts
p
an
ic
le
-1
40
60
80
100
120
Compensation WS: 120% Compensation DS: 69% W S: 3 018 4 sp ikele ts m-2 DS : 50 825 sp ikele ts m-2Observed
Panicles m-2
300
400
500
600
700
Fi
lle
d
gr
ai
ns
p
an
ic
le
-1
40
60
80
100
D S: 3 7834 g rains m -2 W S: 2 6489 g rain s m-2 Compensation DS: 106% Compensation WS: 129%Observed
Hills m-2
25
50
75
100
P
an
ic
le
s
hi
ll-1
0
5
10
15
20
DS: 4 49 panicle s m-2 WS: 3 53 panicle s m-2 Compensation DS: 87% WS: 91%A
B
C
D
E
F
Plasticity of 12
contrasting
cultivars
Sequentially formed yield components
Ph
en
ot
yp
ic
p
la
st
ic
ity
(%
) i
nd
uc
ed
b
y
po
pu
la
tio
n
(A
C
) o
r s
ea
so
n
(B
D
)
0 20 40 60 80 100 120 V2 V3 V5 V6 V7 V9 V10V1, IR72 indica check cv. V11, trop. japonica X indica
V4, trop. japonica V8, temp. japonica X indica V10, indica Mean S.E.
Non-reproductive traits
0 20 40 60 Mean S.E. Time to F L (d ) Plan t hei ght a t FL LAI a t FL SLA at F L Flag leaf widt h Flag leaf are a Flag -1 le af a rea Leaf /ste m dw at F L 0 20 40 60 Hills m-2 Culm s hil l-1 a t PI Panic les c ulm-1 Spike lets panic le-1 Grai ns s pikele t-1 Wei ght g ra in-1 0 20 4060 Mean S.E. Mean S.E.
A
B
C
D
Sequentially formed
yield components
Non-reproductive
traits
Population
effect
Season
effect
Successive compensations converge to
minimize plasticity of last component
(mean grain weight)
Plasticity is source of yield stability
Plasticity (trait) =
Variation/mean =
Trait(T2)-Trait(T2
---
Mean Trait (T1, T2)
Pop. density effect on yield components
Cu
lm
s/h
ill a
t P
I
Cu
lm
s/h
ill a
t P
M
Cu
lm
s/a
rea
at
PM
Po
pu
lati
on
effe
ct (%)
-80
-60
-40
-20
0
20
40
60
Pla
nt
he
igh
t a
t F
L
Fla
g l
ea
f le
ng
th
LA
I a
t F
L
Le
af
dry
we
igh
t a
t F
L
a.g
. d
ry
we
igh
t a
t P
M
Gr
ain
Yi
eld
Pa
nic
les
/ar
ea
Sp
ike
let
s/p
an
icle
s
Sp
ike
let
s/a
rea
Fra
cti
on
fil
led
gr
ain
Ha
rve
st
ind
ex
Observed
Simulated
Effect of increasing
population 4-fold
(4-season means ±SE):
Example of Ic
effect on
tillering
0
20
40
60
80
100
120
Ass
imi
la
tio
n
ra
te
a
nd
R
m
(kg
h
a-1
d
-1
)
0
100
200
300
400
500
600
A (25 hills m-2)
A (100 hills m-2)
Rm (25 hills m
-2)
Rm (100 hills m
-2)
Days after sowing
0
20
40
60
80
100
120
In
de
x
of
co
mp
et
iti
on
(I
c)
0.0
0.5
1.0
1.5
2.0
C
ul
ms
hi
ll-1
0
5
10
15
20
25
30
35
Ic (25 hills m--2)
Ic (100 hills m-2)
Culms/hill
(25 hills m-2)
Culms/hill
(100 hills m-2)
Seedling
nursery
Tillering
Stem
elongation
Grain
filling
Transplanting
A
Phenotype sensitivity to model parameters:
Many traits affect phenotype but not yield due to trait-trait
compensation
Sensitivity of TilAbility
Days after sowing
0 20 40 60 80 100 120
C
ulm
s
hill
-1o
r L
A
I
0 10 20 30 40 0 5 10 15 20 IR72 (2.5) 3.5 1.5Ab
ov
eg
ro
un
d
dw
(g
re
en
)
Gr
ain
y
ield
14
%
M
C
Cu
lm
s
hill
-1LAI
Stem
reserve
Sensitivity of Phyllo (
oCd)
Days after sowing
0 20 40 60 80 100 120 0 5 10 15 20 0 10 20 30 40 IR72 (100) 115 85
Sensitivity of LeafLengthMax (mm)
Days after sowing
0 20 40 60 80 100 120 0 10 20 30 40
D
ry
m
at
te
r (
M
g
ha
-1)
0 5 10 15 20 IR72 (450) 550 350leaf
nu
mb
er
A
B
C
Sensitivity of CoeffLeafdeath
Days after sowing
0 20 40 60 80 100 120
C
ulm
s
hill
-1o
r L
A
I
0 5 10 15 20 25 30 35 40 0 2 4 6 8 10 12 14 16 18 20 IR72 (0.045) 0 (Complete staygreen)Sensitivity of RelMobiliInternodeMax
Days after sowing
0 20 40 60 80 100 120 0 5 10 15 20 25 30 35 40 0 2 4 6 8 10 12 14 16 18 20 IR72 (0.02) 0 (no mobilisation)
Ab
ov
eg
ro
un
d
dw
(g
re
en
)
Gr
ain
y
ield
14
%
M
C
LAI
Stem
reserve
Sensitivity of CoeffTillerDeath
Days after sowing
0 20 40 60 80 100 120 0 5 10 15 20 25 30 35 40 0 2 4 6 8 10 12 14 16 18 20 IR72 (0.12) 0
Cu
lm
s
hill
-1D
ry
m
at
te
r (
M
g
ha
-1)
A
B
C
Plant type optimization study
(virtual breeding)
Agdw (Mg-ha-1)
0
5
10
15
20
C
ou
nt
0
200
400
600
Virtual population
(ca. 15,000)
IR72
Grain Yield (14% MC)
0
2
4
6
8
10
12
C
ou
nt
0
100
200
300
400
• Testing virtual recombinations of traits
• Step 1: Calibrate check-cv IR72
• Step 2: Identify 8 key parameters and vary them
within realistic ranges
• Step 3: Create “recombinants” (ca. 15,000 “cvs”)
• Step 4: Simulate in 2 environments, select 60
• Step 5: Simulate under different population X
season, select 20
• Step 6: Multilocation trial (India Philippines)
• Step 7: Testing under CC scenarios (GCM, multi-side)
• Step 8: Select 4 best and compare with IR72
Effect of trait variation on yield of virtual populations
BVP
Grain Yield (14% MC)
0 2 4 6 8 10 12C
ou
nt
0 100 200 300 400 All 550 820 IR72Inter node LengthMax
Grain Yield (14% MC)
0 2 4 6 8 10 12C
ou
nt
0 100 200 300 400 All 150 225 300 IR72LeafLengthMax
Grain Yield (14% MC)
0 2 4 6 8 10 12C
ou
nt
0 100 200 300 400 All 350 450 550 IR72CoeffInternodeMass
Grain Yield (14% MC)
0 2 4 6 8 10 12C
ou
nt
0 100 200 300 400 All 0.001 0.002 0.003 IR72Tilabilty
Grain Yield (14% MC)
0 2 4 6 8 10 12C
ou
nt
0 100 200 300 400 All 0.1 0.2 0.4 0.5 2.5 IR72CoeffPanicleMass
Grain Yield (14% MC)
0 2 4 6 8 10 12C
ou
nt
0 100 200 300 400 All 0.01 0.02 0.03 IR72Effect of increased stem CH
2
O storage capacity
(from 33% to 43%)
CoeffResCapacityInternode
Grain Yield (14% MC)
0
2
4
6
8
10
12
Co
un
t
0
100
200
300
400
All
0.5
0.75
IR72
IR72
Only effective in highest
yielding backgrounds
Without it, potential yield
gains over IR72 remain
marginal
Optimized ideotype compared to IR72:
(Photosynthesis/Respiration traits not considered)
-
Slightly longer BVP (FL 85-90d)
-
Slightly reduced phyllochron
-
Partial stay-green (slightly less leaf senescence)
-
Moderate tillering
-
Increased stem [NSC] at flowering
-
Increased spikelets/panicle
S1
S2
S3
G
ra
in
Y
ie
ld
(M
g
ha
-1
)
6
8
10
12
Means, best 4
IR72
S1
S2
S3
A
gd
w
(M
g
ha
-1
)
10
12
14
16
18
20
22
S1
S2
S3
H
ar
ve
st
In
de
x
0.4
0.5
0.6
Temperature
Scenario
S1
S2
S3
o
C
20
25
30
35
Tmin
Tmax
Tmean
+1.0
o
C +1.6
o
C
1980-
2000
2010-
2029
2030-
2049
Three GCMs:
HadGEM2
CanSEM2
GFDL
Climate-change scenario
(case: Hyderabad winter crop)
Yield increases because of CO
2
fertilization (490 ppm in S3) but
some ist lost to heat damage;
Ideotype vs. IR72 gain = ca. 12%
Heat
sterility
Yield
IR72
Hyderabad winter season
Year 1981 1982 1983 1984 1985 1986 1987 1988 1989 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014 2015 2016 2017 2018 2019 2020 2021 2022 2023 2024 2025 2026 2027 2028 2029 2030 2031 2032 2033 2034 2035 2036 2037 2038 2039 2040 2041 2042 2043 2044 2045 2046 2047 2048 2049 % s te ril ity 0 10 20 30 40 50