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Modelling the ecophysiological and genetic control of the staygreen trait in sorghum Graeme Hammer, Erik van Oosterom, Greg McLean, Al Doherty, David Jordan, Emma Mace and Andrew Borrell QAAFI, The University of Queensland DAFF Qld Working together with the Queensland Government

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Page 1: Modelling the ecophysiological and genetic control of the ... Workshop... · Modelling the ecophysiological and genetic control of the staygreen trait ... txp 84t xp211 sPb ... Modelling

Modelling the ecophysiological and genetic control of the staygreen trait

in sorghum

Graeme Hammer, Erik van Oosterom, Greg McLean, Al Doherty, David Jordan, Emma Mace and Andrew Borrell

QAAFI, The University of QueenslandDAFF Qld

Working together with theQueensland Government

Page 2: Modelling the ecophysiological and genetic control of the ... Workshop... · Modelling the ecophysiological and genetic control of the staygreen trait ... txp 84t xp211 sPb ... Modelling

1. Staygreen – What is it? Consequences on yield?

2. What is the genetic regulation?

3. What mechanisms might underpin the SG phenotype?

4. Can we model the mechanisms and simulate the phenotype as an emergent property?

5. What are the consequences on yield across sites and seasons?

6. So What?

Outline

Page 3: Modelling the ecophysiological and genetic control of the ... Workshop... · Modelling the ecophysiological and genetic control of the staygreen trait ... txp 84t xp211 sPb ... Modelling

• Hybrids with the stay-green trait have the capacity to maintain more green functional leaves when subjected to drought during the grain filling period

1. Staygreen – What is it?

Reduced lodging

Increased grain yield

Increased grain size

Senescent Impact Staygreen

Page 4: Modelling the ecophysiological and genetic control of the ... Workshop... · Modelling the ecophysiological and genetic control of the staygreen trait ... txp 84t xp211 sPb ... Modelling

• Retrospective analysis of 4 years of breeding trials to quantify the relationship between stay-green and yield

• Small number of male parents contrasting in staygreen crossed with many females with varying levels of staygreen

• Examine yield-staygreen association for each site by male combination

1. Staygreen – Effect on yield?

Male parent Qualitative stay-green Rating

Qualitative ranking of yield potential in high yielding environments

R931945-2-2 High Moderate

R986087-2-4-1 High Moderately high

R933396 Moderately low Moderately high

R940386 Moderately low Moderately high

R995248 Low Moderately high

Jordan et al. 2012. Crop Sci. 52:1153–1161

Page 5: Modelling the ecophysiological and genetic control of the ... Workshop... · Modelling the ecophysiological and genetic control of the staygreen trait ... txp 84t xp211 sPb ... Modelling

LSN

Yiel

d (t

/ha)

-2-10123

-2 -1 0 1 2

R931945-2-22007Bil

-2 -1 0 1 2

R986087-2-4-12007Bil

-2 -1 0 1 2

R9403862007Bil

-2 -1 0 1 2

R9933962007Bil

-2 -1 0 1 2

R9952482007Bil

-2-10123

R931945-2-22007Clermont2

R986087-2-4-12007Clermont2

R9403862007Clermont2

R9933962007Clermont2

R9952482007Clermont2-2

-10123

R931945-2-22008Bil

R986087-2-4-12008Bil

R9403862008Bil

R9933962008Bil

R9952482008Bil

-2-10123

R931945-2-22008DalBox

R986087-2-4-12008DalBox

R9403862008DalBox

R9933962008DalBox

R9952482008DalBox

Low Yield environmentsSt

anda

rdis

ed Y

ield

Standardised Senescence Score

High SG Low SG

Page 6: Modelling the ecophysiological and genetic control of the ... Workshop... · Modelling the ecophysiological and genetic control of the staygreen trait ... txp 84t xp211 sPb ... Modelling

LSN

Yiel

d (t

/ha)

-2-1012

-2 -1 0 1 2

R931945-2-22006CecPl

-2 -1 0 1 2

R986087-2-4-12006CecPl

-2 -1 0 1 2

R9403862006CecPl

-2 -1 0 1 2

R9933962006CecPl

-2 -1 0 1 2

R9952482006CecPl

-2-1012

R931945-2-22006HRS

R986087-2-4-12006HRS

R9403862006HRS

R9933962006HRS

R9952482006HRS

-2-1012

R931945-2-22008HRS

R986087-2-4-12008HRS

R9403862008HRS

R9933962008HRS

R9952482008HRS

-2-1012

R931945-2-22008LivPl

R986087-2-4-12008LivPl

R9403862008LivPl

R9933962008LivPl

R9952482008LivPl

High Yield EnvironmentsSt

anda

rdis

ed Y

ield

Standardised Senescence Score

High SG Low SG

Page 7: Modelling the ecophysiological and genetic control of the ... Workshop... · Modelling the ecophysiological and genetic control of the staygreen trait ... txp 84t xp211 sPb ... Modelling

1. Staygreen – Effect on yield?

Change in Yield per staygreen unit

Mea

n yi

eld

(t/h

a)

2

4

6

8

10

-0.2 0.0 0.2 0.4 0.6

Mea

n Si

te Y

ield

(t/h

a)

Yield Change per Unit of Staygreen Jordan et al. 2012. Crop Sci. 52:1153–1161

A generally positive effect on yield

Especially at yield levels < 5t/ha

Aust average yield is about 2.5 t/ha

Page 8: Modelling the ecophysiological and genetic control of the ... Workshop... · Modelling the ecophysiological and genetic control of the staygreen trait ... txp 84t xp211 sPb ... Modelling

2. What is the genetic regulation?

SBI-01 SBI-02 SBI-03 SBI-04 SBI-05 SBI-06 SBI-07 SBI-08 SBI-09 SBI-1

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t x p5 4 1 tx p 4 97

s P b -1 0 40

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M 18 8 42 7

M 18 8 70 3 s P b- 4 44 8

M 18 8 39 4 M 3 4 28 8 6

M 34 0 87 5

s P b -7 0 15

S b - id s1

c d o 45 7

s P b -3 6 35 s P b -1 4 35

s P b -5 1 60

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t x p6 8 u mc 8 4t x p7 8 s P b -0 93 9

c u p 01 sP b -6 0 1 8

M 34 1 99 9 M 3 4 18 8 3M 34 1 95 7 s P b- 4 49 2

s P b -6 1 60 s P b -5 6 11

s P b -8 2 37 s P b -5 6 68M 34 2 69 8 M 3 4 26 9 9

M 34 2 72 4 M 3 4 27 7 2

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s P b -2 0 14 s P b -0 6 02s P b -1 5 24 S T 1 3 80

s P b -6 1 44 P l as t oc h ro n 3 /Go li at h

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s P b -3 6 97 s P b -1 7 60M 34 0 43 3 M 1 8 74 5 8

M 18 8 43 4 H d 6

s P b -2 5 83 s P b -0 6 23s P b -5 5 21 M 18 7 99 7

M 18 7 55 6 t xp 2 0 8

s P b -2 5 74 s P b -4 3 68s P b -4 7 55 u mc 1 4 7

S b R P G 8 37

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s P b -3 7 74 s P b -7 4 81

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u m c 9 0

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c u p 08 cu p 2 7

t x p5 2 3p h y A s Pb - 56 0 5

c d o 12 1 s P b -1 0 03

t x p3 0 2 H V H GA 1H V GA D 1 u mc 1 07

m sb c ir 34 7 ms b c ir2 4 2

M 34 0 37 0 T b 1

t x p4 8 2 s Pb - 38 9 1D s e n hs b m 6 4

M 18 8 41 1 M 3 4 19 7 1

M 18 7 06 2 M 1 8 70 9 9m sb c ir 22 8

t x p1 1 9

M 34 0 90 4M 34 0 42 0 M 3 4 04 9 1

M 34 2 63 1

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t x p1 3 2

s b A GF O8

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s P b -6 3 84 s P b -0 3 15

GE 5 2

t x p5 5 1 tx p 1 57

s P b -7 3 98

s P b -0 1 50 s P b -2 2 41

u m c 1 28 sP b -6 1 7 6

s P b -3 6 18

u m c 1 66

s P b -8 2 61

s P b -2 0 70

u m c 8 3

s P b -9 8 06

t x p3 5 7

s P b -4 7 93

s P b -6 7 80

g p s b 08 9s P b -0 4 22

S T 1 70 0

s P b -2 8 23 s P b -8 7 94

u m c 1 15 sP b -6 0 6 1s P b -9 4 30 s P b -0 1 29

s P b -1 7 04

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s P b -3 7 27s P b -1 1 91

s P b -3 5 25 s P b -6 0 07

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s P b -4 2 27 s P b -4 2 34

M 18 7 88 0

u m c 1 67

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s P b -2 6 84

s P b -8 9 47 s P b -1 8 90

s P b -3 8 50 s P b -4 5 94

s P b -2 4 12 s P b -9 2 16s P b -0 8 10 C S U 6 5 3

s P b -8 6 53 t x p3 2 9

M 18 8 40 4t x p4 3

s P b -5 8 08 s P b -9 5 14

s P b -6 8 83 s P b -8 4 91s P b -8 8 68 s P b -2 0 58

s P b -0 0 90 s P b -3 3 86

s P b -1 1 70 R G2 9 3 1aS S C IR 1 0 7 S S C IR 2 17

S S C IR 7 4 S S C IR 7 4 I

S T 5 25 ST 5 5

t x S3 6 1 s Pb - 44 2 0s P b -9 0 50

S T 3 44

s P b -2 5 67 s P b -9 7 95t x p8 8

t x p1 4 9 P S B 33

s P b -7 2 29 s P b -9 9 30

i su 7 3

s P b -0 2 32

t x p1 7 0

M 34 3 35 6

t x S1 5 37

C r tr e t xS 1 57 3s P b -8 5 85 s P b -4 5 42

t x s1 0 60

s P b -0 2 74

t x s4 9 0

s P b -5 3 58

t x p3 2 t xp 1 1E S T 1 4

c d o 20

p S B 0 62

t x p8 6

s P b -2 6 02

s P b -9 2 81

c d o 45 4s P b -6 6 00

i su 1 6 5

s P b -7 5 67

s P b -8 4 92

s P b -9 5 60

S T 1 27 7 S T 1 84 7

M 18 8 58 2

t x p3 7 s P b -0 50 9s P b -3 9 30 s P b -6 9 71

s P b -0 1 23 s P b -0 2 07

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s P b -4 0 83

t x p1 8 2

m sb c ir 30 4 b

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s P b -1 9 35 t x p3 3 5

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s P b -7 0 31 r z 25 1

m sb c ir 28 6 g p s b1 7 3

t x s1 1 29

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g a p 57

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S T 1 59 5s P b -3 8 01 s P b -9 2 03

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s P b -8 8 18s P b -8 4 23

S t r4 8 s P b- 5 90 5

s P b -7 9 58s P b -4 6 38 s P b -7 8 33

s P b -3 1 03 s P b -5 4 72

s P b -3 2 64 c d o 9 38s P b -6 0 83

g p s b 05 1i su 5 1

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s P b -2 1 47OP - K 20 C

M 18 7 23 8

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Y

M T1 7

s P b -4 0 29 t x p2 7 9

c u p 22 tx p 20 4M 34 3 24 1 M 1 8 76 1 6

t x s1 7 65

i su 1 3 3 g ap 3 6

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c u p 60 sP b -4 0 3 7s P b -3 3 45 t x p1 5 4

m sb c ir 33 6

s P b -7 4 22 s P b -9 1 56

s P b -6 7 72 s P b -8 1 62

c u p 30

c d o 13 8 7

s P b -0 6 29

t x p5 2 2t x p4 3 3

t x p2 8 4

t x p4 3 2s P b -0 1 31 s P b -3 7 97

t x p6 1 i su 4 9

s P b -0 2 77 s P b -0 7 46

E S T 1 5 E ST 4 0s P b -0 3 83 s P b -4 5 93

t x s3 0 0 sP b - 04 1 4

s P b -2 9 58 s P b -7 1 54

b n l5 .0 9S T 1 83 6 -1

b n l1 4 .2 8

c u p 44

s P b -0 2 33

v r s1

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s P b -6 7 46

S b R P G 8 0

t x s1 6 25

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c u p 62

t x p5 1 5

t x p5 1 9

r z 32 9 t xp 1 80t x p3 4 0

s P b -5 9 60 s P b -1 9 02

u m c 1 57 M 34 2 9 04

M 34 0 36 4

s P b -1 9 84

s P b -4 5 82t x s1 1 53 s P b -5 5 27

s P b -5 8 32 s P b -8 0 66

s P b -8 2 75 s P b -5 9 75s P b -9 3 14 s P b -5 4 31

s P b -1 0 20

s P b -0 4 94 s P b -6 0 66

C c d 1 a cu p 1 5s P b -7 6 11 s P b -6 3 66

s P b -9 4 99 s P b -8 5 32

s P b -1 0 60

S t r3 7

s P b -7 3 31

s P b -3 5 77

S T 1 42 1

r z 27 3 s P b- 3 86 1

m sb c ir 30 6

t x p5 2 4 s Pb - 78 5 0C r tr b 1

t x p2 4 8

s P b -9 0 65s P b -6 0 44 s o c 1

s P b -3 0 43

s P b -2 7 29

s P b -4 4 35g p s b 12 9 M 3 4 03 7 3

F C - 6 G E 32

S S C IR 7 3 S S C IR 8 7S T 1 20 0 -2

s P b -8 7 43 t x p3 1 6

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t x p1 5 0

t x p4 6

s P b -5 2 92

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3_1Q

STG4_1

QSTG

5_1

QS

TG8_1

QS

TG10_1

QS

TG11_1

QS

TG13_1

sP b - 75 4 9 S o c1

H d 6 tx p 9 6

tx p 18 8 s P b -4 8 55sP b - 76 5 7 s Pb - 79 8 4

tx s 12 6 7 tx p 1 97

M3 41 6 62 M 20 0 04 2

M3 41 5 92 m sb c ir 26 6

tx p 63

g ps b 0 11 sP b -2 7 54

cu p 6 4 g p sb 1 7 4

M1 87 7 72

sP b - 16 3 1

sP b - 03 3 3 s Pb - 09 0 9

sP b - 82 1 9 s Pb - 78 0 1

sP b - 31 0 3

tx p 25

M3 42 9 27 gp s b 17 8M3 42 7 11 sP b -8 3 45

M3 40 9 57

M3 41 7 41

sP b - 12 0 9

tx p 80 tx p 29 7

sP b - 09 6 3

ms bc ir 2 23 sP b -7 6 68

tx p 84 tx p 21 1

sP b - 66 8 9 s Pb - 48 6 4tx p 50 is u 67

tx p 65 5 s P b -2 2 04

tx p 65 4 S b 0 2QGM 15 8

S b0 2 QG M1 59 tx i5 2S b0 2 QG M1 60 R f2

S b0 2 QG M1 61 Sb 0 2 QGM 1 62

tx p 73 tx p 61 7tx p 61 6

OP A7

tx i5 1

sP b - 17 6 4 tx p 6 14

sP b - 30 8 6

M3 43 2 89

S SC I R 21 4

S bR P G1 3 6

B 2

tx p 30 4

sP b - 11 2 7 s Pb - 96 8 7

sP b - 95 4 7 tx p 6 13

tx p 61 2 t x p6 1 1tx p 61 0 t x p6 0 9

tx p 60 8 s P b -5 5 11

tx p 61 5 M 34 0 41 6M3 42 0 90 M 34 2 32 9

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tx p 23 9 s P b -3 7 98

sP b - 12 1 8 s Pb - 67 9 4sP b - 11 1 9

R f2 _ ne w

sP b - 83 7 6 s Pb - 53 4 7

S b0 2 QG M1 27 Sb 0 2 QGM 1 28

S b0 2 QG M1 29 Sb 0 2 QGM 1 30

sP b - 34 4 7

tx s 18 9 1

M3 42 3 80

sP b - 44 5 3 s Pb - 88 9 7

sP b - 89 6 7

u m c5 5 s P b -8 6 47

is u 71sP b - 56 4 3 s Pb - 19 4 0

tx s 11 6 4 s Pb - 92 5 3

sP b - 01 4 6sP b - 00 9 0 s Pb - 33 8 6

tx s 13 8 1

sP b - 87 6 6

sP b - 97 0 8 s Pb - 71 7 7

sP b - 10 4 6 s Pb - 58 8 0

tx p 31 8 s P b -1 2 23

sP b - 52 2 4

sP b - 03 6 4 s Pb - 63 6 0sP b - 71 4 2

sP b - 40 8 1

tx p 4

ms bc ir 2 38sP b - 44 1 9 s Pb - 18 0 1

tx p 63 8 s P b -8 6 00

sP b - 86 1 6S T1 3 09

g ps b 0 87

is u 15 1

tx S 91 7 b s P b -8 99 2

sP b - 26 8 5 s Pb - 46 8 9

sP b - 69 5 9M3 40 4 31 M 34 2 47 1

sP b - 76 9 5 s Pb - 64 1 6

M1 87 3 82S T1 6 64 S T9 0 2

sP b - 19 2 0

sP b - 64 3 4 s Pb - 62 8 8

sP b - 42 6 3 tx p 2 01tx p 72 tx p 3

F C A s Pb - 0 15 8

sP b - 49 7 0 s Pb - 16 1 7cu p 7 4 s P b- 7 55 5

tx p 55

sP b - 66 9 3rz 4 13

tx S 64 5 c

tx s 16 3 9 s Pb - 67 2 4M3 43 0 67 M 34 0 88 8

M3 43 1 20

sP b - 41 4 5

F C 40 btx p 19

tx p 28 3 s P b -1 2 62

sP b - 58 9 8 s Pb - 72 0 5

tx p 13

sP b - 78 8 2

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M1 88 4 24 M 34 2 35 8

M3 43 0 86

M1 88 6 43

M1 88 4 79 M 34 2 67 3

M3 42 7 59

sP b - 03 5 0

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sP b - 60 8 0 E f4M3 42 4 98

Z

S bA GA B 0 3

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u m c1 3 9

M3 41 1 08 M 18 8 17 4

sP b - 10 1 7

M1 88 3 48 M 18 7 69 7

sP b - 14 4 2

sP b - 20 7 5 s Pb - 26 3 9

sP b - 43 1 1 s Pb - 67 0 0

ra 1 s P b- 4 16 4sP b - 50 8 8

sP b - 19 7 0

sP b - 13 6 5M1 87 8 71 M 18 7 89 4

sP b - 50 8 7 s Pb - 43 1 2

sP b - 65 3 0 s Pb - 56 8 3

sP b - 34 1 9sP b - 16 8 4

M1 88 3 35

sP b - 24 8 4

sP b - 52 7 2

P SB 4 1 6 S tr 2 9

g am m a- k af ir in

tx p 44 5

ms bc ir 3 39 M18 9 02 0g K AF _ S S R 14 cu p 2 9

C B 92 6 47 3 s P b -8 8 95

sP b - 09 8 2 d ep 1sP b - 21 2 9 s Pb - 26 7 8

u m c5 tx p 43 0

tx p 56 is u 86

tx s 11 1 1 u m c 13 6

tx p 1

S G -3

sP b - 77 0 2

M3 42 7 62

S tr 66

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S T1 9 02

u m c1 1 6

sP b - 25 6 8

sP b - 86 8 8 s Pb - 36 4 6

sP b - 70 6 6 s Pb - 00 7 3

tx p 28 6

g ap 8 4 s P b- 3 89 3

S T6 2 3

F C 28 s P b -7 5 38

sP b - 33 6 1 M 3 40 4 14

M3 40 5 89 M 34 2 40 9sP b - 48 6 2

sP b - 22 2 9

sP b - 09 0 3 s Pb - 15 0 8

is u 11 7

S T1 3 00 sP b - 44 4 4S oc 1 s P b -3 0 64

F t tx p 17 9

sP b - 64 2 4

is u 43

sP b - 33 9 7sP b - 19 2 5 s Pb - 46 8 5

tx s 28 4

T AC 1

M3 41 5 04

M1 87 3 57

sP b - 38 3 4 s Pb - 57 3 7

D f1

sP b - 05 4 9

M1 89 0 02

sP b - 03 1 0 s Pb - 73 1 1tx S 18 5 5 S T 33 6

tx s 44 3

sP b - 14 5 3 is u 66

tx s 15 2 7M3 41 9 68

sP b - 11 5 4 s Pb - 20 8 0

sP b - 23 6 0 tx S 2 04 2 bsP b - 71 0 0 s Pb - 36 3 7

sP b - 08 8 7 tx p 3 48

sP b - 08 6 3 s Pb - 09 7 1cd o 1 50 8 s P b -9 5 11

cd o 3 85

tx p 42 8

sP b - 60 0 9

sP b - 21 3 1 s Pb - 24 2 4

sP b - 52 2 5 S T 10 9M3 42 8 47

sP b - 66 6 3 tx p 3 15

sP b - 45 8 0 tx s 20 4 2tx s 36 0 s P b -5 47 8

tx p 43 1

sP b - 66 4 8

tx p 42 9

Ma 5

M3 42 9 72

tx s 19 2 5sP b - 27 0 0

MT 49

sP b - 61 4 7

u m c4

sP b - 40 5 4

S G 38

cd o 5 9

tx S 11 4 3

sP b - 86 7 7 tx p 1 00

sP b - 35 4 7

sP b - 20 3 9

sP b - 26 2 8cu p 2 6 tx p 7

u m c1 4 9 t xp 2 07

tx s 18 4 5 s Pb - 14 1 3

M1 87 9 35

sP b - 38 3 5 s Pb - 51 4 6

sP b - 71 5 3

tx p 29 6

M3 41 1 72

u m c1 2 5

sP b - 98 6 9 s Pb - 04 1 2sP b - 07 7 4 s Pb - 61 1 3

sP b - 86 0 2

sP b - 23 5 0u m c1 2 2 F C 1 0

v rs 1 S T 35 9

sP b - 52 8 6 s Pb - 56 3 0

sP b - 36 8 3 b n l8 .3 7

sP b - 14 2 3sP b - 86 9 7

rz 3 95sP b - 51 3 8 s Pb - 32 6 9

sP b - 09 9 0

sP b - 80 9 4

sP b - 15 3 3 s Pb - 83 9 9

sP b - 25 8 8 is u 11 3

tx s 29 0 s P b -1 67 1tx s 28 3

cu p 4 0

sP b - 40 0 5 s Pb - 76 4 7

sP b - 68 9 2 s Pb - 50 8 4sP b - 68 0 3

S T1 6 71tx p 8

F P

cu p 6 9

sP b - 43 6 6 s Pb - 55 4 4

tx s 16 1 0

u m c1 6 8

M3 42 8 79

S b- b d 1

b nl 16 .0 6 s P b -8 0 75

sP b - 88 9 6

sP b - 91 9 0

sP b - 28 3 3

sP b - 24 6 5 s Pb - 82 0 8

sP b - 70 1 5

sP b - 04 3 0 s Pb - 73 3 4

sP b - 31 0 9

QSTG

1_2Q

STG2_2

QSTG

3_2

QS

TG5_2

QSTG

8_2

s P b -8 7 06 tx s3 3 3

S S ms b c ir7 8

t x p4 5 6 t xp 4 96

s P b -0 9 65 sP b -9 6 73

s P b -2 2 49 FTs P b -5 3 93 sP b -7 7 95

t x p4 9 4 s P b- 8 56 8

s P b -0 3 63 tx p 45 7

s P b -5 4 54 um c1 2 1s P b -4 3 26 sP b -3 9 67

F T

t x p2 6 6t x p2 2 8 t xs 1 17 8

t x s3 1

t x p5 1 8

M 18 9 27 0 M 3 4 27 0 0

M 19 9 86 1M 34 0 96 3

t x p4 5 4

t x p4 9 2

M 34 2 20 0 M 3 4 07 4 7

M 18 8 91 7 M 3 4 22 8 3

t x p4 9 1

g p s b 01 2M 34 1 40 8 s P b -2 30 9

s P b -5 5 94 sP b -6 7 14

s P b -2 4 61M 34 2 80 8 u mc 1 24

s P b -3 9 40

S b - ra 2 r a2

s P b -1 9 06 sP b -4 9 15s P b -5 4 91

s P b -6 3 11

s P b -1 6 69 sP b -2 0 55s P b -0 6 56 M18 8 93 3

s P b -2 9 63

M 18 8 68 1 M 1 8 86 5 7M 34 0 76 3

M 34 1 54 0

t x s1 0 92

S B R P G8 4 6

M 34 2 54 7

s P b -1 3 64

t x p4 8 9 s P b- 0 31 9

s P b -9 3 24 M34 1 22 3

t x p4 5 1 s P b- 4 97 5s P b -5 4 09

t x p4 5 2

s P b -4 4 31 tx p 21 5M 18 7 27 6

P l a

s P b -3 6 32 sP b -1 0 12

t x p9 tx p 48 8R Z 5 4 3 s Pb - 66 3 3

s P b -6 4 72

s P b -6 8 57

t x p4 2 3 s P b- 9 42 0

s P b -6 5 53

M 34 0 73 2

s P b -9 3 88

t x s1 0 53S b R P G1 06 tx p 48 5

s P b -8 8 32 sP b -6 9 82

F P sP b - 42 2 6S S C IR 16 5 S T 69 8

B H 2 4 5 38 9

s P b -6 6 90

g p s b 08 6

s P b -4 6 39

s P b -2 5 62 tx s5 7 8t x p5 0 0

s P b -2 6 49

S T 1 53 7

i su 1 4 8s P b -4 3 03

P S B 5 1 7

s P b -9 0 76

t x s5 0 3

S T 1 03 1

B H 2 4 6 08 2

S G 2 8r z 24 4 t xp 4 6 1

t x s1 4 38 um c1 0

s P b -1 3 62 sP b -5 1 08s P b -0 5 29 sP b -6 2 76

s P b -1 9 01 sP b -6 1 22

s P b -0 9 52 sP b -1 8 18s P b -8 4 87 sP b -9 4 83

s P b -7 6 88 sP b -1 1 37

t x s1 1 62s P b -5 9 46

s P b -3 5 84 Ef 3

s P b -9 1 35 cd o 9 20

E p s 1s P b -7 6 75 SG- 5 1

m sb c ir 22 4 t xp 3 3

g a p 23 6

s P b -1 2 94

L c ye sP b - 60 2 4

s P b -9 7 79 M18 7 54 6

i su 1 1 4

s P b -5 9 22

t x p2 0 5

P S B 6 0 9

c s u 58M 34 1 88 5 M 3 4 26 6 3

s P b -0 3 57 sP b -1 8 00

M 34 0 38 4 M 3 4 08 5 2

M 34 0 80 2 M 1 8 81 1 4M T3 6

t x p3 1

s P b -1 7 00 sP b -1 4 47

t x p3 3 6 s P b- 0 87 5s P b -8 7 38

t x p1 8 3

m sb c ir 27 6

s P b -8 7 87

s P b -5 3 00t x p4 4 4

t x p5 4 3 t xp 5 44

s P b -5 2 02 sP b -9 8 64

t x s5 8 4

t x p5 1 3

t x p5 0 3t x S1 4 3 9 tx S 53 6

u m c 6 3

R

s P b -4 4 23t x p1 2 0 u m c 93

t x p4 3 5

t x p5 1 4 t xp 5 12

s P b -2 2 09

t x p2

s P b -2 2 46

s P b -0 3 98 tx p 53 0

s P b -3 7 99

c d o 11 6 0

t x p2 3 1 t xp 4 36

s P b -3 9 65

t x p5 9

s P b -1 4 96

S T 1 17 6

t x S3 0 7

W G8 89

t x S5 4 5

b c d 82 8

s P b -9 5 24 cd o 4 70

t x p2 1 8 t xs 1 17 5

u m c 1 6 u m c 17i su 1 2 1

m sb c ir 31 4 s P b -9 13 9

s P b -7 1 61

t x S5 4 5 ls

t x p1 1 4

t x p4 3 7

s P b -1 7 25

S T 4 58

s P b -6 6 82 sP b -6 6 69

s P b -6 1 21 sP b -6 4 37

s P b -5 1 94 sP b -5 6 55

s P b -6 4 66

s b A G E O1s P b -0 0 38 sP b -6 7 78

S b 0 3Q GM 10 0 S b 0 3QG M 10 1

S b 0 3Q GM 10 2 S b 0 3QG M 10 3S b 0 3Q GM 10 4 S b 0 3QG M 10 5

F T

s P b -0 7 85 M34 1 44 6

M 34 1 57 1 s P b -4 13 9s P b -9 9 73

t x s1 9 27 tx s4 2 2

B H 2 4 5 19 1

t x p4 3 8 t xp 4 39

s P b -1 4 80 tx p 44 0

s P b -8 3 49 tx p 54 2b n l1 5 .2 0 t xp 5 63

P I N 5 tx p 4 41

b c d 73 8

s P b -9 4 29 tx p 58 1

M 34 2 70 4 s P b -8 32 3t x p5 8 7 M1 8 83 1 1

t x p4 4 6

S b 0 3Q GM 10 6

t x p4 4 2

S 1 3 29S b 0 3Q GM 10 7 S b 0 3QG M 10 8

s P b -4 9 21 sP b -5 2 67

t x p5 6 4 t xp 5 65S b 0 3Q GM 10 9 t xp 5 8 0

S b 0 3Q GM 12 0 S b 0 3QG M 12 1

S b 0 3Q GM 11 0 S b 0 3QG M 12 2S b 0 3Q GM 11 1 R Z 4 44

s P b -6 8 82

S b 0 3Q GM 12 3

i su 1 6 6 S b 03 Q GM1 16S b 0 3Q GM 11 7 S b 0 3QG M 11 8

S b 0 3Q GM 11 9

s P b -1 0 68 tx p 57 2s P b -8 0 40 tx p 58 3

t x p4 4 7

t x s1 2 26m sb c ir 22 5

t x p2 8 5

S T 2 03 2 -2

t x p4 2 1t x p3 8

t x p4 4 8

R Z 5 3 8

t x p4 4 9

s P b -7 1 86 M18 8 84 6

s P b -2 6 82 sP b -3 6 70s P b -6 0 13 sP b -7 8 66

t x p4 2 2 s P b- 9 97 7

t x p4 2 0

t x S3 0 1

s P b -4 6 29

t x p3 4

S b 0 3Q GM 11 2 S b 0 3QG M 11 3

S b 0 3Q GM 11 4 S b 0 3QG M 11 5

t x p4 2 4b n l5 .3 7

s P b -6 9 25 sP b -3 2 28

s P b -6 6 49 is u6 8M 34 2 77 9 M 3 4 09 7 3

v p 1

M 18 8 34 7 M 3 4 23 0 5M 34 2 56 7 A lt S B

M 18 8 92 0

s P b -9 8 94 cu p 3 2

t x S1 0 7 5t x S1 0 7 5l s

m s3

s P b -0 2 08 sP b -4 8 27

s P b -6 7 70F P

t x p4 2 7

A

F T - lik e t xp 6 9

S b - ba 1

t x p4 2 5

S t r5 8

C U a 1 s P b -5 8 71

t x p4 2 6

s P b -1 6 42 sP b -5 5 80

s P b -8 8 02

QS

TG1_3

QS

TG2_3 Q

STG

3_3

QSTG

4_3 QSTG

12_3

QS

TG9_3

QS

TG11_3

QS

TG13_3

QSTG

15_3

QSTG

18_3

t xp 5 06

t xp 5 04 sP b - 94 3 2

S o c1 sP b - 90 0 6

s P b- 3 75 4 s P b -6 20 8

s P b- 6 46 5

s P b- 2 52 1 s P b -8 88 7

c d o9 6 2 is u 4 7

c u p4 8

t xs 7 54

s P b- 4 07 4

s P b- 6 64 0

s P b- 7 49 3 s P b -8 44 6

s P b- 2 73 9

U n g n h sb m3 2

ms b ci r3 4 0

s P b- 4 71 8 u m c 1 11

s P b- 2 55 8

s P b- 1 58 4

s P b- 4 23 3 S T 1 16 3 -1

s P b- 2 50 0 s P b -1 08 0

c u p0 5 s P b -3 0 78

s P b- 8 35 5 s P b -7 04 2s P b- 5 82 6

S G3 0

g p sb 0 5 0

GE 8

s P b- 1 18 3 s P b -1 96 9

t xp 2 6 b mr 6

s P b- 9 44 1 s P b -5 15 1

s P b- 6 74 1

is u 5 6

F C 2 9a FC 2 9 c

F C 2 9b

N E W - 7 s P b -3 34 3

s P b- 6 55 9

ms b ci r3 2 3 M 1 87 7 63

s P b- 1 85 3s P b- 2 00 9 s P b -9 46 8

s P b- 6 46 9

s P b- 7 25 6

s P b- 1 93 2

M1 8 79 8 5

s P b- 9 58 0 J H 1 4

s P b- 6 95 8

M1 8 83 2 3

GE 2

M2 0 04 1 3

s P b- 5 66 2

M1 8 80 0 5 M1 88 9 03

R Z 5 99

s P b- 6 54 7 s P b -2 94 4

F T s P b -3 8 73

s P b- 6 75 9 GW D

s P b- 2 69 9 s P b -4 03 9

s P b- 4 92 8 s P b -5 61 8s P b- 2 13 8 s P b -9 63 5

s P b- 0 25 8 s P b -2 44 9

s P b- 0 54 3 s P b -5 27 6s P b- 1 81 1 s P b -6 23 1

s P b- 1 84 1 s P b -8 13 8

s P b- 8 37 8 s P b -7 21 0s P b- 8 52 1 s P b -9 39 3

s P b- 5 74 2 s P b -0 86 7

s P b- 1 29 5 s P b -1 79 8s P b- 3 90 3 s P b -4 50 6

s P b- 7 27 4 s P b -7 31 0

s P b- 9 03 3 s P b -9 35 9

s P b- 9 98 7 s P b -8 47 0s P b- 2 73 6 s P b -9 30 0

s P b- 0 27 9 s P b -1 57 4

s P b- 5 53 0 s P b -8 29 6s P b- 0 70 3

M3 4 09 3 0

s P b- 2 84 2

s P b- 4 66 1

s P b- 5 22 0

M1 8 77 1 6

s P b- 4 85 1

s P b- 3 93 9

M1 8 75 0 2 s Pb - 43 4 3s P b- 8 48 8

s P b- 1 63 7

t xp 1 2 t xp 3 43s P b- 1 14 7

s P b- 0 72 4

M3 4 30 0 7

s P b- 7 53 4

M1 8 76 6 1

F C 2 6

S T 97 7

s P b- 6 53 5

t xS 1 11 3 t x S1 1 13 ls

s P b- 4 82 5

s P b- 5 14 3 s P b -5 44 2

t xs 1 09 0 s P b -2 31 3

s P b- 6 94 5

c u p2 8

s P b- 8 01 3 s P b -6 11 5

s P b- 9 30 3

E S T 33

S T 51 7

s P b- 9 35 5

s P b- 0 45 9

S B 1 -1 0

s P b- 0 14 2 s P b -3 98 1

s P b- 6 59 9 s P b -7 79 4

s P b- 8 28 0s P b- 7 32 4

s P b- 9 99 5

s b AG G O2R Z 1 66 M34 0 88 6

M3 4 18 5 1 M3 40 4 82

s P b- 1 93 9 s P b -6 38 8s P b- 7 21 6 s P b -3 83 8

s P b- 4 73 4 s P b -6 13 4

F T -l ik e s Pb - 25 3 7

s P b- 8 33 7 s P b -1 36 8s P b- 9 68 4

M3 4 09 9 0 s Pb - 20 8 4

s P b- 9 04 1 s P b -3 52 0s P b- 3 68 6

S b 5- 2 14

s P b- 7 81 1 s P b -4 55 2s P b- 0 85 4 c d o 13 8 0

s P b- 9 71 3 i su 4 2

s P b- 2 68 9 s P b -7 31 5

s P b- 2 85 2t xp 2 5

t xs 1 07 2s P b- 9 30 4 s P b -4 19 8

s P b- 4 47 3

u mc 54 sP b - 06 1 5s P b- 8 80 6

s P b- 5 44 6 s P b -0 63 7

s P b- 0 92 8 s P b -9 37 4s P b- 9 43 1

s P b- 4 07 0

u mc 68

s P b- 4 64 9is u 1 32

M3 4 06 6 4 M3 42 4 64

M3 4 24 7 0 M1 87 5 39

g p sb 0 7 3 s Pb - 38 5 8t xp 1 77 sP b - 10 3 1

t xp 2 4 t xp 4 1

s P b- 3 45 2 s P b -4 51 7s P b- 9 25 2 C O ?

s P b- 2 71 7

M3 4 03 7 5 M3 41 9 49c su 2 6

is u 3 5

M3 4 29 2 8M3 4 20 1 0

M3 4 20 6 5

t xp 3 27 C C D 8

t xs 6 04

s P b- 9 56 8

s P b- 5 36 7

t xp 6 0

s P b- 4 26 2 s P b -2 46 8

ms b ci r2 3 2

s P b- 3 70 3 s P b -5 23 1

s P b- 7 40 2

s P b- 8 10 0

M1 8 71 7 0 s Pb - 62 6 5

M1 8 72 2 3 M3 42 9 16M1 8 72 0 7 M3 40 9 52

M1 8 83 0 5

t xp 5 1

M1 8 87 1 0

t xp 2 12

t xs 1 90 1

s P b- 3 69 4 s P b -0 09 5s P b- 8 20 7

g p sb 0 4 5

b c d1 4 2 7

s P b- 2 83 2

t xs 1 10 3

s P b- 8 01 6

t xp 2 7

t xp 2 1

s P b- 6 73 1

M3 4 19 0 2 M3 42 3 20M3 4 21 9 1 M3 42 6 43

M1 8 74 6 1

M1 8 71 1 8 tx p 2 65

s P b- 0 97 6 s P b -0 72 9s P b- 8 25 5 s P b -1 71 1

s P b- 4 03 5

s P b- 4 69 2 s P b -8 21 8s P b- 5 34 3 s P b -6 09 8

s P b- 7 43 4

s P b- 5 97 6

u mc 10 4

s P b- 5 55 0

s P b- 0 11 0 s P b -2 34 3s P b- 9 92 9 s P b -6 26 9

QSTG

1_4Q

STG2_4

QSTG

6_4

t xs 2 03 1 s P b -8 5 99

m s b c ir3 2 9 s P b- 8 18 6

M 1 8 84 5 1

g p s b1 5 9

s P b -4 04 1 M 34 1 69 3

s P b -0 90 6

s P b -2 38 1

t xS 9 1 7

s P b -2 88 0

s P b -5 14 8 P S B 5 17 b

s P b -5 89 0

c u ti cu la r wa x

t xp 6 5

F C 4 a

S B 0 5- 2 13 6 71 3s P b -0 38 1

s P b -7 04 7 s P b -5 2 56

S B 0 5- 2 42 9 98 9 s P b -7 30 8

s P b -6 56 5

g p s b8 4 M 1 8 8 10 2s P b -8 27 8 t x p9 4

M 3 4 09 1 6 M3 4 28 6 5

S B 0 5- 3 69 3 45 9 s P b -5 69 2g p s b0 0 5

s P b -4 13 1

s P b -3 97 7

s P b -2 94 1s P b -4 31 3 s P b -8 1 64

s P b -5 39 0 s P b -1 2 27

s P b -2 53 9 s P b -6 6 27s P b -7 63 8 s P b -9 8 15

s P b -1 28 2 s P b -5 3 17

M 1 8 88 9 8 ms b ci r2 4 8u mc 8 g p sb 0 1 7

g p s b1 6 5 s P b- 0 51 0

s P b -8 24 7 s P b -3 1 55

s P b -6 13 7t xs 7 22

t xs 2 07 2 S b - tf l1

t xp 1 1 5s P b -3 29 0 s P b -2 6 26

t xp 3 0 S B 0 5- 4 79 1 64 8

s P b -8 05 3 s P b -6 0 39s P b -7 54 8

s P b -8 88 1

S B 0 5- 4 96 4 87 6 s P b -5 69 8S B 0 5- 5 35 2 10 5 s P b -5 01 7

s P b -7 29 3 s P b -9 0 80

s P b -5 40 8

P S B 6 00 S T 4 5 3s P b -5 65 0

R G5 8 04

s P b -3 99 5s P b -6 00 6

s P b -9 54 4 s P b -5 1 19

t xp 3 0 3

s P b -5 80 5 s P b -9 0 09

s P b -8 31 6

u mc 5 2

s P b -8 39 3 s P b -9 8 47

s P b -0 11 5 s P b -1 5 58s P b -2 87 5 s P b -9 9 14

s P b -2 75 2

s P b -0 65 2

s P b -0 93 2

s P b -8 11 6 s P b -5 8 39

s P b -1 98 9 s P b -2 1 62

s P b -2 61 6 s P b -1 2 53

t xS 3 8 7c

s P b -1 23 6

M 1 8 73 6 5

t xp 2 2 5

C D S C 1 46

i su 1 20

s P b -5 07 7

o p r

s P b -5 89 2

s P b -6 78 7 s P b -1 2 49

s P b -3 16 4 s P b -6 5 05s P b -1 26 0 s P b -9 0 59

s P b -5 39 9 s P b -0 3 68

M 1 8 90 5 8 M3 4 04 6 4C C D 8 s P b -1 0 47

i su 6 5 M3 40 3 9 7

M 3 4 04 4 4 is u 5 9

s P b -4 80 6 s P b -5 2 09s P b -2 48 0 t x p2 9 9

s P b -8 10 8 s P b -4 7 52

M 3 4 09 5 3 M3 4 15 2 1s P b -8 30 6 s P b -7 8 71

s P b -6 86 1 t x p1 4

t xp 1 5 t xs 3 87s P b -3 84 5 t x s7 1 3

M 3 4 05 0 2 F C 4 b

s P b -4 43 3

s P b -9 64 2s P b -0 73 8 s P b -6 5 08

s P b -3 03 1 s P b -3 7 66

s P b -5 19 4s P b -6 34 1 s P b -4 3 23

M 1 8 85 1 2 s P b- 8 14 9

M 3 4 11 5 5M 3 4 15 1 7 s P b- 7 89 3

s P b -9 31 2

s P b -9 05 4s P b -1 50 1

s P b -7 16 7

s P b -9 34 7

s P b -6 63 6s P b -8 67 7

s P b -1 10 4 s P b -6 3 47

s P b -0 20 5t xS 6 4 5b

M 3 4 30 6 5 s P b- 4 76 6

M 1 8 88 7 8s P b -3 86 3 s P b -6 6 04

s P b -8 97 3 s P b -2 1 87

s P b -8 47 0s P b -4 08 6 M 34 2 39 8

c s u1 6 6

s P b -7 20 3 s P b -6 3 23

s P b -6 85 5s P b -7 10 6

s P b -7 03 0

s P b -1 76 6s P b -3 81 7 s P b -9 5 35

s P b -3 52 9

g p s b0 0 3s P b -1 45 4

P S B 2 62

t xp 2 3c d o 52 0

s P b -1 98 5 s P b -7 7 24

R G. S1

P S B 3 02s P b -6 28 7

M 1 8 88 3 4

g p s b1 7 6

s P b -3 49 4

s P b -4 86 1

S b R P G60 8

s P b -9 49 0 S b K A F GK 1_ I

s P b -1 74 0

s P b -7 74 5 S o c 1

s P b -6 83 2 s P b -0 8 14S B K A F G K 1 a lp h ak a fi rin

t xp 1 2 3 al p h ak a fir in

s P b -9 85 8 s P b -2 5 70

t xp 2 6 2 al p h ak a fir in ?E f 4

s P b -1 18 1

t xp 1 3 6

s P b -1 37 3 s P b -2 1 67

s P b -8 88 8 S b 0 5QG M 16 3

M 3 4 20 4 6 M3 4 05 4 7

s P b -3 30 4 M 34 2 03 3

M 3 4 12 7 4 M3 4 33 1 2

s P b -3 41 8

s P b -7 89 2m s b c ir2 2 2

S S c ir1 2 0

g p s b0 3 2

s P b -1 39 6 s P b -0 8 73s P b -2 87 4

S b 0 5QGM 1 6 5

s P b -1 63 1

s P b -5 56 2 s P b -3 8 77

s P b -9 10 7 S b 0 5QG M 16 6

s P b -1 79 4 s P b -9 9 90s P b -7 46 9 s P b -9 7 82

S b 0 5QGM 1 6 7

QS

TG1_5

QSTG

2_5

QSTG

3_5

QS

TG4_5

QS

TG5_5

s Pb - 16 0 2

s Pb - 41 8 3

s Pb - 97 7 2 s P b- 1 67 7

s Pb - 49 0 2 tx p 5 21

s Pb - 14 9 1

s Pb - 49 3 7 s P b- 0 52 0

s Pb - 12 7 9 s P b- 6 49 0

M 3 42 6 40 ms bc ir 2 11

s Pb - 09 8 7

s Pb - 51 8 2

s Pb - 87 9 3 s P b- 9 18 8s Pb - 83 4 2 s P b- 5 37 1

s Pb - 49 9 2

s Pb - 36 4 4

s Pb - 25 5 1 MT 2

c do 4 5 6

s Pb - 35 3 9

S T 18 0 7

s Pb - 89 6 3 s P b- 9 51 1

D s en h s bm 2

tx p 6 s P b- 1 38 7

s Pb - 40 3 6 s P b- 1 63 5

M 3 40 4 48

s Pb - 23 2 2

s Pb - 84 2 2

c do 7 1 8F P

M 1 87 8 05

s Pb - 81 9 8 s P b- 0 03 3

s Pb - 24 0 4 F T

s Pb - 33 4 2 s P b- 8 36 2s Pb - 55 6 4 s P b- 6 01 7

s Pb - 71 6 9 s P b- 1 39 5

d w2B G0 50 4 02

M a 1g ap 7 g a p 72

M 1 87 5 89 sP b - 76 6 0

s Pb - 55 0 6 s P b- 2 11 4s Pb - 60 5 9

M 3 42 0 89 SS m sb c ir 92

M 3 41 5 30

is u 14 2

s Pb - 23 6 2 s P b- 5 41 2

tx p 5 68 s P b -6 7 67s Pb - 26 3 0 s P b- 4 03 0

s Pb - 49 4 7

s Pb - 45 0 3 s P b- 3 01 9

s Pb - 80 6 0

s Pb - 38 3 7 s P b- 2 45 7

s Pb - 84 2 5

s Pb - 91 4 6

u mc 11 9

M 3 41 6 35

S tr 1

M T 8

s Pb - 56 6 1

c up 3 6

s Pb - 46 3 2

s Pb - 50 3 0

S G2 tx S 1 20 6

s Pb - 11 6 9 is u 1 38

s Pb - 24 6 3

g c

s Pb - 81 9 6 s P b- 4 83 3

s Pb - 21 0 3 s P b- 5 40 2

s Pb - 15 4 3

s Pb - 64 4 5

s Pb - 43 2 4 tx p 1 45

u mc 53

d

s Pb - 89 2 8 tx s 1 03 0s Pb - 84 3 6 s P b- 9 06 6

s Pb - 65 7 3 F T -l ik e

s Pb - 83 6 1 s P b- 9 95 0s Pb - 15 7 2

tx p 2 65 -1

is u 58tx p 3 17 s P b -1 9 21

s Pb - 85 3 0 s P b- 1 75 5

tx p 2 65tx p 2 74 s P b -7 7 77

tx S 1 16 1 t xS 1 4 39 b

S T 13 8 1 S T7 2

tx p 1 04 t x p2 1 9

tx p 9 7 sP b - 75 4 4

s Pb - 97 5 2

u mc 34

g p sb 1 27 M1 88 0 23

tx s 15 4 6

s Pb - 01 8 8

s Pb - 72 9 0

D s en h s bm 37

s Pb - 88 5 1

P S B 14 0

M 3 40 9 45 M3 41 0 47M 3 41 7 12

M 3 42 5 81

M 3 42 3 91

M 3 42 7 64

g p sb 0 69

M 3 41 1 92

R s 1

s Pb - 58 0 2 s P b- 7 58 1

tx s 11 7 3

tx p 4 84 F Ls Pb - 77 5 3 s P b- 5 23 0

tx s 19 0 6 c up 1 2

u mc 44 b n l1 0 .1 3 .2

S b -lf y

b n l1 0. 13

s Pb - 53 7 4s Pb - 02 5 5 S o c1

L F Y

tx p 9 5

s Pb - 17 2 1

K S 1

M 3 43 1 62 sP b - 13 7 2

M 1 87 8 24

M 1 89 1 79

s Pb - 39 6 2

tx s 11 3 9

tx p 1 76 s P b -5 4 03

tx s 18 6 8

s Pb - 79 6 0 s P b- 5 76 5

L g

S T 98 2

tx p 5 7

s Pb - 00 1 7 s P b- 1 48 6

s Pb - 47 3 2 s P b- 5 60 3

s Pb - 65 2 6 tx s 2 06 3

s Pb - 97 3 6

M 3 42 0 82

P

is u 14 7

tx s 15 3 3

M 3 40 4 15 sP b - 50 8 6

s Pb - 89 5 4

s Pb - 52 1 2 s P b- 6 07 6

s Pb - 94 2 2 s P b- 5 10 1tx p 1 7 sP b - 05 8 1

s Pb - 16 5 7 s P b- 7 62 7

s Pb - 43 3 4 P d s

tx s 10 8 5 is u 5 3

tx s 11 2 4s Pb - 84 1 0

S G36

s Pb - 96 6 7

is u 14 4

M 1 87 3 62 sP b - 74 2 8

s Pb - 35 1 6 s P b- 7 84 0E S T1 sP b -1 6 2 3

g p sb 1 69

S b R PG 2 2 5

s Pb - 68 0 1 s P b- 9 24 9

s Pb - 04 7 7 s P b- 3 77 3

M 3 41 1 10

M 3 41 3 92 M3 42 4 92M 3 43 3 04 gp s b 0 19

M 1 88 0 96 M1 88 2 51

M 3 40 4 83 cu p 3 7

M 3 42 6 78

QSTG

1_6Q

STG3_6

t xp 4 1 6 tx p 6 60

t xp 6 5 9 tx p 4 18

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t xp 5 9 4 tx p 5 96t xp 1 1 8 H H U 2 0

t xi 46 tx p 66 5

s P b -6 5 45 t x i4 5t xp 6 5 7 tx i4 7

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t xp 3 6 t xp 4 1 4

s P b -4 7 40 s P b -7 0 64s P b -9 5 95 s P b -9 3 09

s P b -3 9 95 s P b -8 3 16

s P b -2 0 34 s P b -0 1 38s P b -4 3 88 s P b -5 7 21

s P b -7 1 05 s P b -7 6 98

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t xs 1 67 4 s P b -0 9 07

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s P b -9 9 78 s P b -8 7 97

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s P b -3 9 46 s P b -6 9 42s P b -3 0 17

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s P b -3 7 08 s P b -7 5 31

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s P b -3 8 43 s P b -6 3 98

s P b -5 4 85 s P b -8 2 58

s P b -8 0 19g p s b0 1 0 s P b- 7 67 8

s P b -7 2 80

p lw8 s P b -0 9 60s P b -4 8 74 s P b -6 4 11

s P b -9 5 08 s P b -7 9 44

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s P b -2 7 74 s P b -3 4 86s P b -1 8 30

s P b -6 9 06

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s P b -6 5 77s P b -8 6 73

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s P b -6 8 63 s P b -7 6 87s P b -3 5 41 s P b -9 2 06

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I

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s P b -5 3 97 s P b -9 3 65

s P b -9 3 51

t xp 2 9 5 s Pb - 61 8 0

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c u p 52

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t xp 9 9 s P b -6 32 6s P b -5 6 88

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s b A GA B 02 s P b -9 7 50t xs 1 55 4 t x s1 1 85 .1

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s P b- 2 03 3 s P b -2 71 6

s P b- 2 74 1 s P b -3 12 3s P b- 3 78 3 s P b -4 25 0

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s P b- 6 91 9 s P b -7 34 3s P b- 8 49 4 s P b -8 50 8

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t xp 2 92 sP b - 25 6 8

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s P b- 3 31 2s P b- 3 18 9 s P b -9 70 0

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t xp 3 54s P b- 1 66 1

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t xs 2 06 5 s P b -1 43 3s P b- 4 14 4 c d o 34 4 .1

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s P b- 7 31 2

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s P b- 1 88 1 t xp 3 2 1m s b ci r3 0 4a R f1

s P b- 6 93 5

s P b- 1 59 5 s P b -1 85 0t xp 4 00

t xp 4 06 tx s5 6 0

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t xs 1 22 0

g p sb 0 0 2

s P b- 5 68 9 s P b -3 50 9

S h s1

s P b- 4 54 6 s P b -0 14 4

s P b- 0 03 1 s P b -2 22 8

s P b- 9 74 3s P b- 0 83 3 s P b -3 95 4

R G1 27 4

t xp 1 05

g p sb 1 2 3

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s P b- 7 82 3 s P b -3 72 4

s P b- 0 15 9 s P b -8 87 8

t xs 1 29 4 s P b -2 49 8s P b- 7 54 2

s P b- 5 57 2

s P b- 1 82 5 s P b -6 91 8

M 1 8 84 0 8s P b- 6 96 0 s P b -4 07 3

s P b- 8 93 1 is u 1 46

s P b- 2 84 6M 1 8 85 2 3

s P b- 7 44 9

s P b- 7 36 9 s P b -9 58 4M 3 4 28 8 0 M3 40 4 54

s P b- 3 43 4

s P b- 7 22 0 s P b -5 40 1s P b- 1 05 1 s P b -1 29 1

s P b- 8 36 3 B H 2 4 57 7 3

c u p3 1 g a p 34

t xs 1 35 3s P b- 8 38 4 s P b -0 28 9

s P b- 1 27 2 s P b -7 08 2

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s P b- 5 53 4 s P b -0 37 6

s P b- 3 63 9

s P b- 8 99 3

s P b- 7 64 8

s P b- 4 38 5 s P b -9 34 1s P b- 3 51 0 s P b -8 16 0

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s P b- 2 29 0s P b- 5 42 0

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s P b- 0 08 7

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s P b- 1 32 3

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s P b -5 4 73

b e ta - ka f iri n s P b- 3 27 4

s P b -1 9 97

s P b -9 4 49

s P b -2 7 70

s P b -8 0 90

s P b -8 5 90

s P b -4 0 11

s P b -1 8 95

t x p2 8 9 S b 09 QG M1 2 6

b K A F _ S SR 1 4 b K A F _S S R 8

s P b -8 8 59 um c1 2

s P b -8 3 66

g a p 85

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s P b -0 0 31

r z 39 0

s P b -6 5 06t x s1 7 03 tx p 35 8

s P b -9 9 62 sP b -4 7 87

s P b -9 4 10 sP b -8 3 68

s P b -7 9 55 sP b -9 0 91s P b -7 3 67

J H 7

M 34 0 96 4

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t x p4 5 9

s P b -0 3 03

s P b -4 7 36 sP b -5 0 56

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s P b -5 0 46 tx p 41 0A P 2

c d o 58 0

t x s1 1 50 sP b -9 3 49

s P b -6 5 15 tx p 41 2

u mc 6 4

M 34 1 11 5M 34 2 28 0

M 34 0 58 7

C D S 0 9 4

s P b -7 4 63

s P b -4 1 56 sP b -9 7 65s P b -3 0 02 sP b -2 1 79

s P b -7 0 13

s P b -0 0 05

t x p4 1 1

g a p 42

s P b -4 8 53

t x p2 5 8 t xp 2 87

S b 4 -3 2

s P b -7 1 93

s P b -3 9 71

t x p2 3 0

t x p6 7

t x p3 5 5

s P b -6 6 67

F T ga p 1 5s P b -5 1 42 sP b -8 3 38

s P b -1 2 39 sP b -0 7 71

M 18 8 20 7 ms b c ir3 1 7

m sb c ir 33 7 b

M 18 7 90 2

M 34 2 49 4 M3 4 27 4 1M 34 1 02 1

M 34 1 25 1

M 34 2 54 3s P b -7 9 05

s B A C 1 20 sP b -5 8 87

s P b -0 3 26

t x s9 4 3 c up 0 2A W 67 9 2 55 s P b -9 1 58

S b R P G 8 21

s P b -5 0 55 sP b -8 6 40

M 18 8 50 1 s P b -6 28 0s P b -9 9 36 M 34 2 02 8

M 34 1 81 7 s P b -9 27 1

s P b -7 6 20 sP b -9 5 38s P b -3 2 53 sP b -3 2 10

s P b -9 2 72 sP b -5 3 12

S G3 4 G E 5

t x S1 6 2 8 S T1 7 1bS T 1 29 6

s P b -9 3 06 sP b -3 1 58

t x s1 3 83 dh n 2s P b -1 3 24 sP b -8 8 73

s P b -6 8 52 sP b -4 7 86

s P b -6 0 43 sP b -9 0 03s P b -4 5 87 tx p 10

s P b -8 7 64

M 18 8 56 0

s P b -9 6 88s P b -6 7 48

s P b -3 2 98

S T 1 37 4

s P b -6 1 46 M 34 0 53 6

s P b -8 7 77

s P b -8 5 42 bc d 4 54S b 0 9Q GM15 3 s P b -1 81 4

s P b -2 1 71 sP b -4 4 16

s P b -0 9 75s P b -9 9 89

M 34 0 97 4

M 18 8 41 2 S b R P G74 3

M 34 1 48 0

F C 7

s P b -0 5 33 sP b -0 0 16

s P b -3 5 18 sP b -8 7 88S b 0 9Q GM15 4 s P b -3 07 2

s P b -2 9 49

s P b -4 9 33

s P b -9 2 43 sP b -9 6 66

s P b -2 5 30 Sb 0 9 QGM 15 5

s P b -6 4 14

S b 0 9Q GM15 6

s P b -8 3 19 sP b -6 4 03

s P b -2 6 70

S T 3 29 c

u mc 3 8 t xp 3 39

s P b -9 2 27 sP b -8 8 12

s P b -2 3 68

s P b -1 9 91c d o 89

s P b -6 0 89 M 34 0 81 0

S b 0 9Q GM15 7 M 3 4 07 3 0

g p s b 07 9M 34 2 83 9

s P b -1 7 18 sP b -5 0 41

s P b -8 8 53

S T 1 05 4 E S T 2 2

M 34 2 54 0 M1 8 86 5 2M 18 9 11 2

F T sP b - 40 0 6

s P b -8 7 16 um c1 3 2

s P b -1 7 32 sP b -0 8 52M 34 3 23 1 s P b -2 96 8

s P b -7 8 55

s P b -6 3 90

g p s b 02 0 s P b -7 68 6

s P b -7 4 60

M 18 8 06 1

t x S3 0 7 b S b -H T 9 .1

s P b -4 6 54 sP b -4 9 20s P b -7 5 99

M 34 3 36 3

t x S1 0 1 5

M 18 7 49 4u mc 6 2

s b A GE O3

s P b -5 0 20 tx p 10 7

s P b -0 4 45

S T 7 15

S T 1 01 7

S b R P G 9 50

t x s1 1 02 rz 20 6

c d o 20 2

s P b -8 2 85 sP b -7 1 04

s P b -4 6 14 EF 3

s P b -7 6 42

s P b -4 5 22 sP b -6 6 78

u mc 1 3 4 cd o 5 42c d o 39 3

i su 1 4 0

QSTG

1_9

QSTG

1_9Q

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QS

TG2_9

S G33

sP b - 39 1 2

cd o 5 90 a X s b ar s lb k1 0 .4 8

b mX s b ar sl b k1 0 .5 7

cu p 4 9 P S B 30 5sP b - 20 4 1 rl f

sP b - 50 7 9 sP b - 41 2 9

sP b - 68 3 3 sP b - 03 1 1sP b - 06 0 0 D L T

cd o 5 90 sP b - 10 4 4

A P2

sP b - 44 2 7

tx s 10 7 8

A D 12

cd o 4 75

S G19

tx s 11 6 3

sP b - 83 0 6

S bR P G9 1 7

sP b - 52 8 1S T2 0 72

M 3 40 7 04

sP b - 60 1 3

sP b - 19 4 5

m s bc ir 3 31 m sb c ir 32 4

M 1 89 1 36

sP b - 68 7 3 wx

sP b - 59 6 5

m a 4

M 3 42 5 57

S tr 30 b

is u 13 6M 1 87 9 40 s P b -6 7 20

g ps b 0 27

R s2 sP b -8 7 7 5F T S o c1

S bR P G9 2 1

sP b - 00 8 8S tr 14

sP b - 95 1 7 sP b - 88 5 8

sP b - 83 5 7

u mc1 1 3

sP b - 34 3 2 sP b - 02 4 8sP b - 21 9 3 sP b - 50 0 5

M 3 41 2 94 s P b -9 5 92

sP b - 38 2 8sP b - 11 0 4

tx S 31 0

sP b - 59 1 7

sP b - 08 1 7 sP b - 32 8 7

sP b - 05 6 2 sP b - 53 9 1sP b - 40 4 5

sP b - 21 4 9sP b - 62 9 2 sP b - 36 5 5

sP b - 22 2 2 tx s 55 8

sP b - 04 9 4

sP b - 28 3 6sP b - 81 5 0 sP b - 84 9 7

sP b - 35 4 9

rz 1 44

u mc1 1 0

sP b - 93 8 1 sP b - 74 7 3

sP b - 19 6 2M 3 40 7 74 M 34 2 24 4

M 1 88 9 83 M 34 0 87 1

sP b - 45 2 1sP b - 15 3 8

sP b - 63 3 1

G E 37

tx p 20H d 1

G E 30 S T 4 03

sP b - 55 1 2 sP b - 62 8 8sP b - 12 4 4 sP b - 00 6 9

sP b - 92 7 5

cd o 1 7sP b - 99 9 9 sP b - 97 9 6

tx p 27 0 t xp 3 0 9

u mc2 1 S b 1 0Q GM 12 4

S b1 0 QGM 1 25 C Otx p 33 1

sP b - 30 3 7 sP b - 71 4 5

sP b - 99 2 1M 3 41 6 06

M 3 41 1 23

sP b - 98 2 6sP b - 09 4 3

b nl 7. 24 Sv P E P C A A

m s bc ir 2 83 s P b -6 2 71sP b - 91 4 0 sP b - 06 6 8

sP b - 58 4 1 tx p 2 17

d K AF tx p 13 0

tx s 19 2 3 sP b - 82 0 1sP b - 27 3 5 sP b - 68 8 9

sP b - 71 8 9 sP b - 41 2 1

tx s 19 2 9 sP b - 90 6 4sP b - 35 2 1 M1 87 1 42

sP b - 17 2 2 sP b - 92 1 5

sP b - 84 3 2 sP b - 39 5 8sP b - 44 8 0

sP b - 68 7 5 sP b - 49 4 4

sP b - 76 4 3sP b - 18 3 9 sP b - 38 8 6

sP b - 97 3 1

sP b - 72 8 4 sP b - 71 7 3

sP b - 80 3 5sP b - 19 3 8 tx s 11 0 6

F T

M 3 40 3 33sP b - 79 4 8

sP b - 36 9 6

sP b - 58 3 4rz 4 76

sP b - 96 8 2

sP b - 03 7 4tx s 44 3

sP b - 32 3 3

sP b - 95 5 5

sP b - 23 1 8sP b - 80 1 9

S T4 8 6- 1

cd o 5 90 b s P b -8 5 27sP b - 19 4 1

JH 2 3

sP b - 14 7 2sP b - 29 3 5 sP b - 82 3 2

sP b - 10 9 5 sP b - 47 8 1

sP b - 79 8 5b nl 5. 04

sP b - 08 3 0

S bR P G7 4 2

cd o 7 8 S E 5, H Y 1S SC I R 63

sP b - 57 0 3

M 3 42 8 42

M 3 40 8 56 M 34 2 95 4

M 3 40 7 12 M 34 0 49 5g ap 1

cd o 4 00

sP b - 35 6 3 sP b - 65 3 2X nh s b m1 00 8 X n h sb m 1 0 11

X nh s b m1 01 3 s P b -4 81 9

sP b - 47 7 2 sP b - 97 4 8

sP b - 97 6 7

T rit

u mc1 5 0

sP b - 08 5 9 tx s 23 6

tx s 16 8 4

m s bc ir 2 62 m sb c ir 22 7sP b - 01 6 2

X nh s b m1 03 3

sP b - 09 3 9 tx S 1 69 4u mc2 1 8

X nh s b m1 04 3

sP b - 45 3 9

sP b - 30 0 3 X n hs b m 1 04 4

sP b - 17 0 1

X nh s b m1 04 8S T1 7 50 - 2

F C 40

M 3 42 5 66 M 34 2 05 3

M 3 43 2 52

tx S 30 9

sP b - 16 6 0

F C 48 S T 1 84 5

R Z 14 3

tx s 75 8

sP b - 70 5 8 cu p 4 3

sP b - 45 8 8 sP b - 58 1 4

g ps b 1 45

P SB 6 1 9

tx p 14 1 S b 6 -3 2 5

S T7 4 5

is u 16 2

sP b - 58 0 9is u 45

tx s 66 4

sP b - 16 5 5 sP b - 74 1 2sP b - 13 0 3 sP b - 76 5 5

sP b - 56 7 8

cu p 0 7 P sy 1

Stg3bStg1

Stg2

Stg4

Stg3a

Crasta et al 1999Hausmann et al 2002

Kebede et al 2001Srinivas et al 2009Subudhi et al 2000

Tao et al 2000Xu et al 2000 Mace and Jordan 2011. Theor Appl Genet. 123: 169-191

All Published stay-green QTL - 87 QTL; 27 unique regions

Four major regions of interest have received detailed study

Page 9: Modelling the ecophysiological and genetic control of the ... Workshop... · Modelling the ecophysiological and genetic control of the staygreen trait ... txp 84t xp211 sPb ... Modelling

Reducedplant size

Reduced water use at anthesis

Reduced tillering

Reduced leaf size

Deeper rooting OR

lateral spread

Root architecture

Increased water availability anthesis

Increased water accessibility

Better water

extraction

Increased growth rate and panicle mass

Increased HI

Increased grain number

Early anthesis

Fig. 1

Increased TE

Increased grain mass

Increased yield

• Studies with NILs and contrasting hybrids to seek mechanisms conferring yield advantage (Borrell, van Oosterom)

• Focus on canopy dynamics (water demand); root system architecture (water capture); transpiration efficiency

• Basis for modelling via effects on dynamics of water balance through the crop cycle

• Understanding supports gene discovery

3. What mechanisms underpin staygreen? • Staygreen is a complex trait – not all mechanisms are desirable

(eg reduced grain number)

van Oosterom, Borrell et al 2011. Crop Sci. 51: 2728-2740

Page 10: Modelling the ecophysiological and genetic control of the ... Workshop... · Modelling the ecophysiological and genetic control of the staygreen trait ... txp 84t xp211 sPb ... Modelling

Reducedplant size

Reduced water use at anthesis

Reduced tillering

Reduced leaf size

Deeper rooting OR

lateral spread

Root architecture

Increased water availability anthesis

Increased water accessibility

Better water

extraction

Increased growth rate and panicle mass

Increased HI

Increased grain number

Early anthesis

Fig. 1

Increased TE

Increased grain mass

Increased yield

• Staygreen regions associated with reduced tillering and canopy leaf area

• QTLs for tillering co-locate with some staygreen QTLs

3. What mechanisms underpin staygreen?

• Reduced tillering is associated with staygreen

van Oosterom, Borrell et al 2011. Crop Sci. 51: 2728-2740

Page 11: Modelling the ecophysiological and genetic control of the ... Workshop... · Modelling the ecophysiological and genetic control of the staygreen trait ... txp 84t xp211 sPb ... Modelling

Reducedplant size

Reduced water use at anthesis

Reduced tillering

Reduced leaf size

Deeper rooting OR

lateral spread

Root architecture

Increased water availability anthesis

Increased water accessibility

Better water

extraction

Increased growth rate and panicle mass

Increased HI

Increased grain number

Early anthesis

Fig. 1

Increased TE

Increased grain mass

Increased yield

3. What mechanisms underpin staygreen?

van Oosterom, Borrell et al 2011. Crop Sci. 51: 2728-2740

• Staygreen regions associated with smaller leaves and reduced canopy leaf area

• Reduced leaf size is associated with staygreen

0

200 

400 

600 

800 

2 4 6 8 10 12 14 16  18

Leaf length ( mm )

Leaf number

‐NIL (Stg2)RTx7000Le

af S

ize

(cm

2)

Page 12: Modelling the ecophysiological and genetic control of the ... Workshop... · Modelling the ecophysiological and genetic control of the staygreen trait ... txp 84t xp211 sPb ... Modelling

Reducedplant size

Reduced water use at anthesis

Reduced tillering

Reduced leaf size

Deeper rooting OR

lateral spread

Root architecture

Increased water availability anthesis

Increased water accessibility

Better water

extraction

Increased growth rate and panicle mass

Increased HI

Increased grain number

Early anthesis

Fig. 1

Increased TE

Increased grain mass

Increased yield

3. What mechanisms underpin staygreen?

van Oosterom, Borrell et al 2011. Crop Sci. 51: 2728-2740

• Staygreen regions associated with enhanced total water extraction

• QTLs for narrow root angle co-locate with staygreen QTLs

• Additional water capture is associated with staygreen

0

50

100

150

200

250

300

350

0 20 40 60 80 100 120 140 160

Days after emergence

Cum

ulat

ive

crop

wat

er u

se (m

m)

Stg2Tx7000

Mace et al. 2012. Theor Appl Gen, 124: 97-109

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Reducedplant size

Reduced water use at anthesis

Reduced tillering

Reduced leaf size

Deeper rooting OR

lateral spread

Root architecture

Increased water availability anthesis

Increased water accessibility

Better water

extraction

Increased growth rate and panicle mass

Increased HI

Increased grain number

Early anthesis

Fig. 1

Increased TE

Increased grain mass

Increased yield

• Staygreen regions associated with enhanced TE

• May be linked to leaf size (SLA, SLN) and/or limited maximum transpiration rate mechanism

3. What mechanisms underpin staygreen?

• Enhanced TE is associated with staygreen

Sincalir et al 2005. Funct Plant Biol. , 32: 945-952.van Oosterom, Borrell et al 2011. Crop Sci. 51: 2728-2740

Gholipoor et al 2010. Field Crops Res. . 119:85-90

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4. Modelling staygreen - mechanisms Incorporate effects in APSIM – sorghum model

• Modelling the tangled web of interactions and feedbacks

Hammer et al. 2010. J Exp Bot, 61:2185-2202

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4. Modelling staygreen - mechanisms

• Tillering – reduce propensity to tiller (0.5 tillers/plant)• Leaf size – reduce potential leaf area per leaf by 10%• Root architecture – increase potential extractable soil water by a

maximum of 15mm over soil depth from 1-2m• TE – increase intrinsic TE from 9 to 9.5Pa in combination with

leaf size reduction− impose maximum transpiration rate of 1mm/hr

• Does this generate simulated staygreen phenotype under terminal stress?

• What are consequences on yield across environments?• Simulation study for conventional agronomy and hybrid at key

locations in NE Australia (with 100 years historical weather)

Incorporate estimates of known effects

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5. Modelling staygreen - phenotype Staygreen phenotype emerges as consequence of dynamics

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5. Modelling staygreen - phenotype Staygreen phenotype emerges as consequence of dynamics

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6. Simulating consequences on grain yield Yield consequences reflect trends in field data (eg Dalby)

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6. Simulating consequences on grain yield Yield consequences relate to Environ Type (eg Dalby)

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7. So What?

• Model captures dynamics and consequences of mechanisms • Staygreen phenotype an emergent consequence• Shifting a small amount of water from pre- to post-anthesis

makes a big difference• Different mechanisms generate different consequences and

GxE • Opportunity for targeted phenotyping and molecular breeding• Opportunity for plant design/improvement simulation (G-to-P

models & G(QTL)*M*E)• Confidence to pursue use of model in molecular breeding• Support for gene function targets in gene discovery

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Modelling the ecophysiological and genetic control of the staygreen trait

in sorghum

Graeme Hammer, Erik van Oosterom, Greg McLean, Al Doherty, David Jordan, Emma Mace and Andrew Borrell

QAAFI, The University of QueenslandDAFF Qld

Working together with theQueensland Government