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Physiological determination of oats yield: Physiological determination of oats yield: thoughts on promissory traits for further thoughts on promissory traits for further
genetic gainsgenetic gains
Daniel Daniel CalderiniCalderini
Universidad Austral de ChileUniversidad Austral de Chile
International Oat ConferenceInternational Oat ConferenceMinneapolis, 2008Minneapolis, 2008
This presentation has the objective of:
1. Review few aspects of the effect of plant breeding on oat grain yield and associated traits
2. Analyze major grain yield components (i.e., grain number and grain weight)
3. Propose traits that could be useful to continueincreasing yield potential
4. Propose a model of GW determination for temperatecereals
IntroductionIntroduction
Grain crops are facing important challenges:
- Expected increase of food demand
- Alternative uses (biofuels)
- Little opportunities for expanding arable land and
irrigated areas
- High grain production with low environmental impact
1.4
1.6
1.8
2
2.2
2.4
2.6
2.8
80 90 2000
Gra
in y
ield
(M
g ha
Gra
in y
ield
(M
g ha
-1))
OatOat
85 95
Years
WheatWheat
BarleyBarley RyeRye
Slafer and Peltonen-Sainio (2001)
In addition, some evidences are showingthat grain yield of temperate cereals areleveling off
IntroductionIntroduction
Plant breeding is the most cost effective, and environmentally safe, way of increasing grain yield
The knowledge of physiological bases of grainyield determination could provide useful tools toplant breeding programs aimed at increasingyield potential
1
2
3
4
5
6
1910 1930 1950 1970 1990 2010 1910 1930 1950 1970 1990 2010 1910 1930 1950 1970 1990 2010
Minnesota Finland Italy
Gra
in y
ield
(M
g ha
-1)
Year of release
y = 0.036x – 67.4(r = 0.71; p<0.01)
y = 0.099x – 15.7(r = 0.66; p<0.05)
y = 0.018x – 31.9(r = 0.96; p<0.001)
Oat breeding effect on grain yieldOat breeding effect on grain yield
BreedingBreeding
Adapted from Wych and Stuthman (1983);
Peltonen-Sainio (1990); Redaelli et al. (2008)
0
1
2
3
4
20 25 30 35 400
1
2
3
4
5
45 50 55 60
Gra
in y
ield
(M
g ha
-1)
Harvest index (%)
Minnesota Finland
y = 0.099x – 0.36(r = 0.90; p<0.001)
Relationship between grain yield and harvest indexRelationship between grain yield and harvest index
y = 0.114x – 2.2(r = 0.54; p<0.05)
BreedingBreeding
Adapted from Wych and Stuthman (1983);
Peltonen-Sainio (1990)
0
2
4
6
8
10
0 10 20 30 40 50 60
Oat
Wheat
Harvest index (%)
Gra
in y
ield
(M
g h
a-1)
Theoretical maximum HITheoretical maximum HI
Relationship between grain yield and harvest indexRelationship between grain yield and harvest indexin wheat and oatin wheat and oat
BreedingBreeding
DD SD SH
Plant height (cm)
Gra
in y
ield
70 100
Plant height optimumPlant height optimum
Richards (1992)
Miralles and Slafer (1995)
BreedingBreeding
Grain yield Grains m-2=Grain
Weightx
Grain yield will be increased by improvingGrain yield will be increased by improvingmajor yield componentsmajor yield components
ArgentinaAustraliaIndiaItalyMexicoUK
Relationship between grain yield and grain numberRelationship between grain yield and grain number
Calderini et al. (1999)
Variability of grain number, grain weight and grain yieldVariability of grain number, grain weight and grain yield(comparison between years)(comparison between years)
0.00
0.02
0.04
0.06
0.08
0.10
0.12
0.14
0.16
0.18
Spring
barley
Springoat
Spring
wheat
Winter
wheat
Winterrye
Grain Number
Grain Weight
Yield
Sta
ndar
d de
viat
ion/
mea
n
Adapted from Peltonen-Sainio et al. (2007)
Grain yield
Grain weightGrains m-2
Sowing
EmergenceAnthesis Physiological
MaturityBooting HarvestTillering
Adapted from Slafer and Rawson (1994)
Major yield components and crop cycleMajor yield components and crop cycle
Emergence Anthesis PhysiologicalMaturity
Booting HarvestTillering
Adapted from Slafer and Rawson (1994)
Gra
in n
um
ber
un
der
str
ess
Grain numberGrain number Growth of reproductive
organs
Gro
wth
of
rep
rod
uct
ive
org
ans
How to improve the growth of reproductive organsaimed at increasing grain number?
1.Higher partitioning of crop biomass to reproductive organs at pre-heading
2. Higher biomass production
3. Longer duration of the period when reproductive organs are growing
Grain numberGrain number
1.Higher partitioning of crop biomass to reproductive
organs at pre-heading
Gonzalez et al. (2005)
Grain numberGrain number
Year of release Grain yield Stem weight (g m-2) (g m-2)
1920 319.9 839.11990 649.1 496.4
Difference 329.3 -342.7
Calderini et al. (1995)
1. Higher partitioning of crop biomass
Grain numberGrain number
Muurinen & Peltonen-Sainio (2006)
Radiation interception
Radiation use efficiency
RUE at pre-heading
ab abbc
0.0
0.5
1.0
1.5
2.0
2.5
3.0
Jama Aslak Suomi
Cultivar
Rad
iatio
n us
e ef
ficie
ncy
(g M
J-1 m
-2)
2. Higher biomass production
Grain numberGrain number
WheatRUE at pre-anthesis
Amador & Calderini (unpublished)
Rad
iatio
n us
e ef
ficie
ncy
(g M
J-1 m
-2)
0.0
0.5
1.0
1.5
2.0
2.5
3.0
3.5
4.0
Otto Quijote
b
a
Cultivar
Grain numberGrain number
0
100
200
300
400
500
600
700
Otto Quijote
a b
0.00.51.01.52.02.53.03.54.0
ba
Wheat
Cultivar
RU
ER
I
Amador & Calderini (unpublished)
Grain numberGrain number
(Slafer et al., 1996)
Emergence Anthesis PhysiologicalMaturity
Booting HarvestTillering
Adapted from Slafer and Rawson (1994)
Gro
wth
of
rep
rod
uct
ive
org
ans
3. Longer duration of the period when reproductive organs are growing
Grain numberGrain number
Whitechurch et al., (2007)
In wheat and barley, clear differences between In wheat and barley, clear differences between duration of pheno-phases have been found duration of pheno-phases have been found
Grain numberGrain number
Relative duration of pheno-phases in oat(14 cultivars and 6 lines from Finland)
Vegetative / whole crop cycle
Generative / whole crop cycle
Range of relative duration(%)
14.3 – 17.4
32.0 – 36.5
Phenological phases
Peltonen-Sainio & Rajala (2007)
Grain numberGrain number
Major yield components: Grain WeightMajor yield components: Grain Weight
Although more conservative than GN, GW could be aAlthough more conservative than GN, GW could be auseful trait for continuing increasing yield potential ofuseful trait for continuing increasing yield potential oftemperate cereals, especially if future plant breedingtemperate cereals, especially if future plant breedingprograms set more grains with lower GW potentialprograms set more grains with lower GW potential
Due to higher stability of GW, the knowledge of Due to higher stability of GW, the knowledge of physiological bases of GW determination canphysiological bases of GW determination canhelp breeding programs to increase grain yieldhelp breeding programs to increase grain yieldby improving GWby improving GW
Major yield components: Grain WeightMajor yield components: Grain Weight
In oat, there are clear evidences that this trait has beenIn oat, there are clear evidences that this trait has beenincreased by plant breedingincreased by plant breeding (e.g., Wych and Stuthmann, 1983;(e.g., Wych and Stuthmann, 1983;Peltonene-Sainio, 1994)Peltonene-Sainio, 1994)
Grain yield
Grain weightGrains m-2
Sowing
EmergenceAnthesis Physiological
MaturityBooting HarvestTillering
Adapted from Slafer and Rawson (1994)
Assumptions on GN and GW determinationAssumptions on GN and GW determination
Temperature was increased (5.5°C) at differentTemperature was increased (5.5°C) at differentdevelopmental phases in wheat, barley and triticaledevelopmental phases in wheat, barley and triticale
S Em Anthesis PM Heading Harvest
Time DR TS Booting
B-AH-A
SE-B
Ugarte et al. (2007)
Effect of temperature on major yield componentsEffect of temperature on major yield components
Grain weightGrain weight
Effect of increased temperature on GN and GWEffect of increased temperature on GN and GWAveraged across crops and experiments (3 years)Averaged across crops and experiments (3 years)
H-A -7%
B-A -15% -15%
SE-B -7%
GNGN GWGW
-8%
-42%
Grain yield
S Em Anthesis PM Head Harvest Time DR FI TE
Grain weightGrains m-2
Booting
This has important consequences for the study of This has important consequences for the study of grain weight determination grain weight determination
All these evidences are showing that flowers,All these evidences are showing that flowers,which grow at the pre-anthesis period, are which grow at the pre-anthesis period, are
involved in GW determinationinvolved in GW determination
Grain weightGrain weight
New evidences of the involvement of the pre-anthesis period in GW determination gives support to promising molecular traits for increasing GW in temperate cereals
spikelet
Different traits associatedwith GW were measuredat central spikelets of thewheat spike
G1G2
G3G4
rachis
Grain weightGrain weight
36.351.055.552.7Huayun
45.357.557.956.3Pandora2
41.751.658.455.9Huanil
42.752.657.953.5Pandora12
42.565.172.167.8Kambara
35.350.356.952.5Bacanora3
36.357.866.261.5Kambara
37.451.656.351.6Bacanora2
41.860.365.262.3Kambara
31.643.949.844.9Bacanora11
G4G3G2G1
Grain weight (mg)CultivarSeasonExperiment
Grain weight in two experimentsGrain weight in two experiments
Grain weightGrain weight
y = 86.18x + 3.97
R2 = 0.790
1020
30
40
5060
70
80
0,0 0,2 0,4 0,6 0,8 1,0
y = 126.93x – 5.93
R2 = 0.860
10
20
3040
50
60
70
80
0,0 0,2 0,4 0,6 0,8 1,0
Carpel weight at pollination, W10 (mg)Carpel weight at pollination, W10 (mg)
Gra
in w
eigh
t (m
g)G
rain
wei
ght
(mg) year 2006year 2006 year 2007year 2007
Relationship between grain weight and carpel weightRelationship between grain weight and carpel weight
Grain weightGrain weight
Dynamic of grain dimensionsDynamic of grain dimensionsPM
0
2
4
6
8
10
0 10 20 30 40 50
days after anthesisdays after anthesis
Gra
in d
imen
sion
s (m
m)
Gra
in d
imen
sion
s (m
m)
Length
WidthHeight
Grain weightGrain weight
y = 1.12x + 4.06
R2 = 0.870
20
40
60
80
100
0 20 40 60 80 100
Grain volume (mmGrain volume (mm 33))
Gra
in w
eigh
t (m
g)G
rain
wei
ght
(mg)
y = 17.89x - 81.53
R2 = 0.860
20
40
60
80
5 6 7 8 9 10
Grain length (mm)Grain length (mm)G
rain
vol
ume
(mm
Gra
in v
olum
e (m
m33))
Relationship between grain weight, grain volume andRelationship between grain weight, grain volume andgrain lengthgrain length
Grain weightGrain weight
0
10
20
30
40
50
60
70
0 10 20 30 40 50
days after anthesisdays after anthesis
Gra
in d
ry m
atte
r an
d w
ater
G
rain
dry
mat
ter
and
wat
er
cont
ent
(mg)
cont
ent
(mg)
Dynamic of dry matter and water content of grainsDynamic of dry matter and water content of grains
Dry matter
Water
Maximum water contentMaximum water content
Grain weightGrain weight
Relationship between grain weight and maximumRelationship between grain weight and maximumwater content of grainswater content of grains
Gra
in w
eigh
t (m
g)G
rain
wei
ght (
mg)
Maximum water content (mg)Maximum water content (mg)
Experiment 1Experiment 1 Experiment 2Experiment 2
y = 1,39x - 2,45
R2 = 0,950
20
40
60
80
20 30 40 50 60
y = 0,93x + 15,89
R2 = 0,92
0
20
40
60
80
20 30 40 50 60
Grain weightGrain weight
010203040506070
0 20 40 60 0 20 40 60 80 0 20 40 60 800
2
4
6
8
10
Days after flowering
Gra
in w
ater
co
nte
nt
(mg
)G
rain len
gth
(mm
)
Pandora Bacanora Kambara
Dynamics of water content and enlargement of grainsDynamics of water content and enlargement of grains
Grain weightGrain weight
0
20
40
60
80
5 6 7 8 9 10
Grain length (mm)Grain length (mm)
Gra
in w
eig
ht o
r M
axim
um
Gra
in w
eig
ht o
r M
axim
um
W
ate
r C
ont
en
t (m
g)W
ate
r C
ont
en
t (m
g)
Relationship between grain weight or maximum waterRelationship between grain weight or maximum watercontent and grain lengthcontent and grain length
GW
MWC
(r= 0.91; p<0.001)
(r= 0.95; p<0.001)
Grain weightGrain weight
Grain length is associated with carpel weightGrain length is associated with carpel weight
4
5
6
7
8
9
10
0.20 0.30 0.40 0.50 0.60 0.70
r = 0.75; p<0.01
Carpel weight at pollination (mg)Carpel weight at pollination (mg)
Gra
in le
ngth
(m
g)G
rain
leng
th (
mg) year 2007year 2007
year 2006year 2006
Grain weightGrain weight
Recurrent selection for grain yield in oat increased GW, grain area, grain length and grain width (De Koeyer et al., 1993)
In the poster session of this conference, Hu et al. (2008) show data of GW and grain dimensions
Relationship between GW and grain length:
r2= 0.74; p<0.05
Grain weightGrain weight
As a consequence of the relationship between final grain weight and carpel weight at anthesis, is proposed that the pericarp of grains is involved in grain weight determination. In other words, the final size of the pericarp is setting the potential weight of grainsPericarp growth is the result of the number of pericarp cellsand the enlargement of these cells. Plant cell enlargementis controlled by proteins called expansins, which are involved in cell wall loosening (McQueen-Mason et al., 1992)
Recently, expansin expression has been found in growing grains of wheat (Calderini et al., 2006; Liu et al, 2007)
Grain weightGrain weight
Preliminarily, we found the expression of 6 expansins in pericarp of grains at 10 days after anthesis from the experiment 2
- ExpA 2- ExpA 4- ExpA 6- ExpA 8- Exp novel 1 (similar to Festuca
pratensis)- Exp novel 2 (similar to Oriza sativa)
After this first step, ExpA 6 has been separated in 3 different goups: ExpA 6 a, ExpA 6 b and ExpA 6 c
ExpansinsExpansinsGrain weightGrain weight
0
2
4
6
8
0 5 10 15 20 25
Days after anthesis
Gra
in le
nght
(m
m)
1,0
1,2
1,4
1,6
1,8
2,0
Expression of E
XP
A 6c (ID
)
Lenght
Expansin
Dynamics of grain length and ExpA 6c expresion in Dynamics of grain length and ExpA 6c expresion in cultivar Bacanoracultivar Bacanora
Grain weightGrain weight
Antisense Sense Negative control
Hybridization of Expansin A 6c in grains at Hybridization of Expansin A 6c in grains at 5 days5 days after anthesisafter anthesis
Grain weightGrain weight
Antisense Sense Negative control
Hybridization of Expansin A 6c in grains at Hybridization of Expansin A 6c in grains at 10 days10 days after anthesisafter anthesis
Grain weightGrain weight
Proposed Model of Potential Grain Weight DeterminationProposed Model of Potential Grain Weight Determination
Carpelsgrowth
Lag Phase
Carpel weightat pollination
Endosperm
Endospermcell number
PhysiologicalMaturityPollinationBooting
Pericarpcells
Water ingrain
Maximum watercontent
Pericarp elongation
Expansin expression
ConclusionsConclusions
- The overlapping of both GN and GW The overlapping of both GN and GW determination in temperate cereals is determination in temperate cereals is longer than generally assumed longer than generally assumed
- The sensitivity of major yield components The sensitivity of major yield components to environmental constraints during this to environmental constraints during this overlapping has been similaroverlapping has been similar
- There is not evidences of trade-off betweenThere is not evidences of trade-off between GN and GW as a consequence of theGN and GW as a consequence of the overlapping between booting and anthesisoverlapping between booting and anthesis
ConclusionsConclusions
- There are opportunities for increasing GNThere are opportunities for increasing GN of temperate cereals. Differences between of temperate cereals. Differences between pheno-phases found in oats is a starting pointpheno-phases found in oats is a starting point for using this trait in plant breeding programsfor using this trait in plant breeding programs
- The involvement of expansins in grain The involvement of expansins in grain enlargement could be used for increasingenlargement could be used for increasing GW in temperate cerealsGW in temperate cereals
Mrs. Carolina Lizana Ph.D. Student (U. Austral de Chile)
Dr.Ricardo Riegel Adjunt Professor (U. Austral de Chile)
Dr. Simon McQueen-Mason Professor (U. of York)
Thank youThank you!!
Time (days after anthesis)
G
rain
we
igh
t (m
g)
0
10
20
30
40
50
60
0 10 20 30 40 50
Grain growth rate
Grain filling duration
Usual approach for studying GW Usual approach for studying GW
Grain weightGrain weight
Response of sunflower Response of sunflower yield components to yield components to shadingshading
Cantagallo et al. (2004)
0
20
40
60
80
15.5 24.5Pre-anthesis temperature (°C)
K
ern
el w
eig
ht
(mg
)
Wardlaw (1994)
Wheat
Other evidences of Other evidences of the involvement of the involvement of the pre-anthesis the pre-anthesis period in grain weight period in grain weight determinationdetermination
Grain weightGrain weight
Carpel weight at anthesis (mg)
Gra
in w
eig
ht
(mg
)
0
20
40
60
80
0 1 2 3 4 0 1 2 3 4 0 1 2 3 4
G1 G2 G3
Control
De-graining
Buenos Aires
Relationship between GW and Relationship between GW and carpel weight at anthesis in carpel weight at anthesis in
wheatwheat
(data from different experiments)
Calderini & Reynolds (2000)
Grain weightGrain weight
Grain number reduction relative to the controlGrain number reduction relative to the control
H-A +1%
B-A -13% -13%
WheaWheatt
SE-B -40%
Exp. 1 Exp. 2
H-A +3%
B-A +4% -18%
BarleyBarley SE-B -36%
H-A -2%
B-A -15% -16%
TriticaleTriticale SE-B -48%
Ugarte et al. (2007)
Grain weight reduction relative to the controlGrain weight reduction relative to the control
H-A -9%
B-A -14% -19%
WheaWheatt
SE-B -7%
Exp. 1 Exp. 2
H-A -8%
B-A -13% -21%
BarleyBarley SE-B -8%
H-A -9%
B-A -15% -23%
TriticaleTriticale SE-B -6%
Ugarte et al. (2007)
Grain weightGrain weight
y = 1.12x + 4.06
R2 = 0.870
20
40
60
80
100
0 20 40 60 80 100
Grain volume (mmGrain volume (mm 33))
Gra
in w
eigh
t (m
g)G
rain
wei
ght
(mg)
Relationship between grain weight and grain volumeRelationship between grain weight and grain volume
Grain weightGrain weight