35
Integrating Crop Physiology and Modelling with Genetic Improvement Graeme Hammer Centre for Plant Science, QAAFI The University of Queensland [email protected]
Integrating Crop Physiology and
Modelling with Genetic Improvement
Graeme Hammer
Centre for Plant Science, QAAFI
The University of Queensland
Integrating Modelling with Breeding
Overview
1. Genetic improvement
– conceptual overview; prediction and complexity
2. Phenotypic distance and prediction
– crossing scales of biological organisation
3. Where does crop physiology and modelling fit?
– introducing biological insight via crop models
4. Trait dissection and phenotypic prediction
– the example of staygreen in sorghum
– the example of limited maximum transpiration in maize
5. Where to from here?
– informing future crop improvement
Integrating Modelling with Breeding
1. Genetic Improvement – process
Genotyping
Base
Germplasm
METs
- Phenotyping -
Crossing
Decisions
Selected
Lines
Phenotypic
prediction
Double haploids
Improved varieties
Inbred lines
- Phenotyping -
High thruput
phenotyping
Phenotypic prediction
Cycles of selection and evaluation in breeding
Genetic Gain – the breeder’s equation
Pij = µ + Gi + Ej + (GE)ij.
years per cycle
Genomic
prediction
The Complexity Problem in Crop Improvement
Progress in crop improvement is limited by the ability to identify favourable combinations of genotypes (G) and management practices (M) in the target population of environments (E) given the resources available to search among possible combinations.
We cannot predict (and navigate to) the desired destination on the adaptation landscape
Cooper, M. and Hammer, G.L. (1996) Plant Adaptation and Crop Improvement. CABI
1. Genetic Improvement – complexity and prediction
Integrating Modelling with Breeding
Integrating Modelling with Breeding
1. Genetic Improvement – complexity and prediction
Now Drowning in G and P Data
But starving for insight as to its
effective use for phenotypic
prediction in genetic improvement
How to scale from G to P?
Integrating Modelling with Breeding
2. Phenotypic Distance and Prediction
Crossing scales of biological organisation confounds prediction
Integrating Modelling with Breeding
2. Phenotypic Distance and Prediction
Crossing scales of biological organisation confounds prediction
- G & E context dependencies (“it depends”)
- Constrain molecular breeding and GP prediction using statistical methods
- Biological models unable to span all levels
Hammer et al 2016. in : “Crop Systems Biology“, Springer
Integrating Modelling with Breeding
2. Phenotypic Distance and Prediction
Short phenotypic distance – trait scales directly
- Few context dependencies; relatively simple traits
- Likely greater role for genetic prediction and gene network models
Hammer et al 2016. in : “Crop Systems Biology“, Springer
Integrating Modelling with Breeding
2. Phenotypic Distance and Prediction
Long phenotypic distance – complex trait does not scale directly
- G*E; complex genetic architecture and physiological interactions; scaling confounded
- Likely greater role for ecophysiology and crop models to unravel dependencies
Hammer et al 2016. in : “Crop Systems Biology“, Springer
Integrating Modelling with Breeding
3. Where does Crop Physiology & Modelling fit?
Introducing biological knowledge
“The right answer for the right reason!”
APSIM Sorghum - The tangled web of dynamic interactions & feedbacks -
Captures non-linearities and context dependencies for trait G-to-P modelling
Hammer et al. 2010. J Exp Bot, 61:2185-2202
Integrating Modelling with Breeding
3. Crop Model Introduce Biological Insight
1. Maize leaf and silk extension rate – Chenu et al. 2008 Plant Cell Environ – Chenu et al 2009 Genetics
2. Sorghum staygreen – Borrell et al. 2000 a,b,c Crop Science; 2014 New Phytol, JExpBot – van Oosterom et al 2011 Crop Science
3. Sorghum tillering – Lafarge and Hammer 2002 Ann Bot – Kim et al. 2010a,b Ann Bot ; Alam et al. 2014 New Phytol
4. Sorghum root system architecture & water capture – Singh et al. 2010 Plant and Soil – Singh et al. 2011 Crop Science – Mace et al. 2011 Theor App Gen ; Singh et al. 2012 Eur J Agron
5. Maize maximum transpiration rate – Yang et al. 2012 Env and Exp Bot – Choudhary et al. 2014 Crop Sci. – Messina et al. 2015 Agron J
Integrating Modelling with Breeding
4. Trait Dissection and Phenotypic Prediction
Sorghum staygreen example
Linking Trait Understanding with Modelling and Genetics
Integrating Modelling with Breeding
4. Trait Dissection and Phenotypic Prediction
Sorghum staygreen – effect on yield in breeding population trials
Change in Yield per staygreen unit
Mean
yie
ld (
t/h
a)
2
4
6
8
10
-0.2 0.0 0.2 0.4 0.6
Mean
Sit
e Y
ield
(t/
ha)
Yield Change per Unit of Staygreen
A generally positive
effect on yield
Especially at yield levels
< 5t/ha
Aust average yield is
about 2.5 t/ha
Integrating Modelling with Breeding
4. Trait Dissection and Phenotypic Prediction
Jordan et al. 2012. Crop Sci. 52:1153–1161
Reduced
plant 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
Sorghum staygreen – plausible mechanisms
Integrating Modelling with Breeding
van Oosterom et al 2011. Crop Sci. 51: 2728-2740 Borrell et al. 2014. New Phytologist 203:817-830
• Studies with NILs and
contrasting hybrids to seek
mechanisms conferring yield
advantage
• Basis for modelling via effects
on dynamics of water balance
through the crop cycle
4. Trait Dissection and Phenotypic Prediction
• Additional water capture at depth is
associated with narrow nodal root
angle and root system architecture
Integrating Modelling with Breeding
4. Trait Dissection and Phenotypic Prediction
Sorghum staygreen – mechanisms
Singh et al. 2011. Crop Sci 51:2011–2020; Singh et al (2012) Eur J Agron 42: 3-10.
Mace et al. 2012. Theor Appl Gen, 124: 97-109
• Additional water capture at depth is
associated with narrow nodal root
angle and root system architecture
• Develop root phenotyping system
(shorten phenotypic distance) and
explore genetic associations
Integrating Modelling with Breeding
4. Trait Dissection and Phenotypic Prediction
Sorghum staygreen – mechanisms
Singh et al. 2011. Crop Sci 51:2011–2020; Singh et al (2012) Eur J Agron 42: 3-10.
Mace et al. 2012. Theor Appl Gen, 124: 97-109
Phenotyping Root System Architecture
– Genetic architecture – High throughput phenotyping of nodal root angle for
genetic prediction/gene discovery using structured populations
4. Trait Dissection and Phenotypic Prediction
Integrating Modelling with Breeding
Root architecture variation: rapid
screening methodology
Parental lines of mapping populations, NAM
populations and breeding populations screened
Mapping population NAM populations Breeding trials
X
Identify genomic regions
and multiple alleles
List of candidate
genes Comparative
genomics
SBI-02-sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
consensus
-
R0
02
70
4
RS
29
R0
02
68
9
R0
02
69
8
R0
02
73
5
R0
02
74
7-1
R0
02
08
7
LR
91
98
R0
01
96
6-1
R0
02
04
0-1
R0
02
08
7
R0
02
04
1
R3
19
45-2
-2R
319
45-2
-2
R0
02
70
4
RS
29
R0
02
68
9
R0
02
69
8
R0
02
73
5
R0
02
74
7-1
R0
02
70
4R
002
70
4
RS
29
RS
29
R0
02
68
9R
002
68
9
R0
02
69
8R
002
69
8
R0
02
73
5R
002
73
5
R0
02
74
7-1
R0
02
74
7-1
R0
02
08
7
LR
91
98
R0
01
96
6-1
R0
02
04
0-1
R0
02
08
7
R0
02
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
-sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
consensus
-sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
consensus
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
consensus
-
R3
19
45-2
-2R
319
45-2
-2
R0
02
70
4
RS
29
R0
02
68
9
R0
02
69
8
R0
02
73
5
R0
02
74
7-1
R0
02
70
4R
002
70
4
RS
29
RS
29
R0
02
68
9R
002
68
9
R0
02
69
8R
002
69
8
R0
02
73
5R
002
73
5
R0
02
74
7-1
R0
02
74
7-1
R0
02
08
7
LR
91
98
R0
01
96
6-1
R0
02
04
0-1
R0
02
08
7
R0
02
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R3
19
45-2
-2R
319
45-2
-2
R0
02
70
4R
002
70
4
RS
29
RS
29
R0
02
68
9R
002
68
9
R0
02
73
5R
002
73
5
R0
02
74
7-1
R0
02
74
7-1
R0
02
70
4R
002
70
4
RS
29
- 1-
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1R
002
04
1
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
SBI-02R
002
08
7
LR
91
98
R0
01
96
6-1
R0
02
04
0-1
R0
02
08
7
R0
02
04
1
R0
02
08
7
LR
91
98
R0
01
96
6-1
R0
02
04
0-1
R0
02
08
7
R0
02
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R0
02
08
7
LR
91
98
R0
01
96
6-1
R0
02
04
0-1
R0
02
08
7
R0
02
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1R
002
04
1
R0
02
08
7
LR
91
98
R0
01
96
6-1
R0
02
04
0-1
R0
02
08
7
R0
02
04
1
R0
02
08
7
LR
91
98
R0
01
96
6-1
R0
02
04
0-1
R0
02
08
7
R0
02
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R0
02
08
7
LR
91
98
R0
01
96
6-1
R0
02
04
0-1
R0
02
08
7
R0
02
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1R
002
04
1
SBI-02-sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
consensus
-
R0
02
70
4
RS
29
R0
02
68
9
R0
02
69
8
R0
02
73
5
R0
02
74
7-1
R0
02
08
7
LR
91
98
R0
01
96
6-1
R0
02
04
0-1
R0
02
08
7
R0
02
04
1
R3
19
45-2
-2R
319
45-2
-2
R0
02
70
4
RS
29
R0
02
68
9
R0
02
69
8
R0
02
73
5
R0
02
74
7-1
R0
02
70
4R
002
70
4
RS
29
RS
29
R0
02
68
9R
002
68
9
R0
02
69
8R
002
69
8
R0
02
73
5R
002
73
5
R0
02
74
7-1
R0
02
74
7-1
R0
02
08
7
LR
91
98
R0
01
96
6-1
R0
02
04
0-1
R0
02
08
7
R0
02
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
-sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
consensus
-sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
consensus
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
consensus
-
R3
19
45-2
-2R
319
45-2
-2
R0
02
70
4
RS
29
R0
02
68
9
R0
02
69
8
R0
02
73
5
R0
02
74
7-1
R0
02
70
4R
002
70
4
RS
29
RS
29
R0
02
68
9R
002
68
9
R0
02
69
8R
002
69
8
R0
02
73
5R
002
73
5
R0
02
74
7-1
R0
02
74
7-1
R0
02
08
7
LR
91
98
R0
01
96
6-1
R0
02
04
0-1
R0
02
08
7
R0
02
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R3
19
45-2
-2R
319
45-2
-2
R0
02
70
4R
002
70
4
RS
29
RS
29
R0
02
68
9R
002
68
9
R0
02
73
5R
002
73
5
R0
02
74
7-1
R0
02
74
7-1
R0
02
70
4R
002
70
4
RS
29
- 1-
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1R
002
04
1
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
SBI-02R
002
08
7
LR
91
98
R0
01
96
6-1
R0
02
04
0-1
R0
02
08
7
R0
02
04
1
R0
02
08
7
LR
91
98
R0
01
96
6-1
R0
02
04
0-1
R0
02
08
7
R0
02
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R0
02
08
7
LR
91
98
R0
01
96
6-1
R0
02
04
0-1
R0
02
08
7
R0
02
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1R
002
04
1
R0
02
08
7
LR
91
98
R0
01
96
6-1
R0
02
04
0-1
R0
02
08
7
R0
02
04
1
R0
02
08
7
LR
91
98
R0
01
96
6-1
R0
02
04
0-1
R0
02
08
7
R0
02
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R0
02
08
7
LR
91
98
R0
01
96
6-1
R0
02
04
0-1
R0
02
08
7
R0
02
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1R
002
04
1
SBI-02-sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
consensus
-sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
consensus
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
consensus
-
R0
02
70
4
RS
29
R0
02
68
9
R0
02
69
8
R0
02
73
5
R0
02
74
7-1
R0
02
70
4R
002
70
4
RS
29
RS
29
R0
02
68
9R
002
68
9
R0
02
69
8R
002
69
8
R0
02
73
5R
002
73
5
R0
02
74
7-1
R0
02
74
7-1
R0
02
08
7
LR
91
98
R0
01
96
6-1
R0
02
04
0-1
R0
02
08
7
R0
02
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R3
19
45-2
-2R
319
45-2
-2
R0
02
70
4R
002
70
4
RS
29
RS
29
R0
02
68
9R
002
68
9
R0
02
69
8R
002
69
8
R0
02
73
5R
002
73
5
R0
02
74
7-1
R0
02
74
7-1
R0
02
70
4R
002
70
4
RS
29
RS
29
R0
02
68
9R
002
68
9
R0
02
69
8R
002
69
8
R0
02
73
5R
002
73
5
R0
02
74
7-1
R0
02
74
7-1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1R
002
04
1
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
-sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
consensus
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
consensus
-sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
consensus
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
consensus
-
R3
19
45-2
-2R
319
45-2
-2
R0
02
70
4R
002
70
4
RS
29
RS
29
R0
02
68
9R
002
68
9
R0
02
69
8R
002
69
8
R0
02
73
5R
002
73
5
R0
02
74
7-1
R0
02
74
7-1
R0
02
70
4R
002
70
4
RS
29
RS
29
R0
02
68
9R
002
68
9
R0
02
69
8R
002
69
8
R0
02
73
5R
002
73
5
R0
02
74
7-1
R0
02
74
7-1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1R
002
04
1
R3
19
45-2
-2R
319
45-2
-2
R0
02
70
4R
002
70
4
RS
29
RS
29
R0
02
68
9R
002
68
9
R0
02
73
5R
002
73
5
R0
02
74
7-1
R0
02
74
7-1
R0
02
70
4R
002
70
4
RS
29
- 1-
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1R
002
04
1
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -21870.0
*txp965.3
*txp639.6
sPb -419522.9
*txp2526.0
*txp29729.5
*txp5038.9
sPb -968741.7
sPb -864745.6
*txp30445.8
sPb -5643 sPb -194068.6
sPb -338678.1
sPb -970891.1
sPb -636093.8
sPb -714295.3
sPb -522497.4
sPb -036498.1
*txp499.5
sPb -8616101.8
sPb -5880 sPb -1046102.0
sPb -8992106.0
*cup74107.0
*txp55110.6
sPb -6434113.8
sPb -4970115.2
*txp3117.3
sPb -6724119.5
*txp19121.8
*txp283127.1
*txp13131.2
*txp298142.1sPb -1784143.0sPb -2075 sPb -1017
sPb -4311 sPb -6700152.0
sPb -5087157.9
sPb -2484167.8
*txp445170.9
*cup29172.4*txp430175.2*txp1183.4
*txp56187.1
sPb -3917190.4
sPb -0073193.0sPb -2568197.0
*txp286200.6*gap84202.4sPb -4862204.1
sPb -3361206.3sPb -4685212.9
sPb -1925213.7
*txp179215.2
*txp348226.7sPb -4444230.0sPb -0971 sPb -9511236.1
sPb -6009239.5
*txp315 sPb - 6663241.1
sPb -5478241.9*txp431242.7*txp429244.7
sPb -0863246.5*txp100255.5
*umc149260.8
*cup26263.3
sPb -1413 *txp207264.7*txp7265.7
sPb -7153266.0
*txp296271.8sPb -0774279.9
sPb -2350281.4
sPb -3683 sPb -5286282.0
sPb -1533294.0*cup40302.2sPb -4005306.1
*txp8309.0*cup69313.7
SBI-02R
002
08
7
LR
91
98
R0
01
96
6-1
R0
02
04
0-1
R0
02
08
7
R0
02
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1R
002
04
1
R0
02
08
7
LR
91
98
R0
01
96
6-1
R0
02
04
0-1
R0
02
08
7
R0
02
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1R
002
04
1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7R
002
08
7
LR
91
98
LR
91
98
R0
01
96
6-1
R0
01
96
6-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
04
0-1
R0
02
08
7R
002
08
7R
002
08
7R
002
08
7
R0
02
04
1R
002
04
1R
002
04
1R
002
04
1
#
#
#
###
####
####
####
#####
ROMA
DALBY
MOREE
BILOELA
EMERALD
TAMWORTH
#
#
#
###
####
####
####
#####
ROMA
DALBY
MOREE
BILOELA
EMERALD
TAMWORTH
.0 100 200 30050 Kilometers
QLD
NSW
Legend:
# EC sites
EC stations
0 - 10 000
10000 - 40000
40000 - 100000
100000 - 16000 0
Population screening
and pre-existing
performance data from
NAM populations and
breeding trials
Project onto
consensus map
and align to
WGS
Mace et al. 2012 Theor Appl Gen, 124: 97-109
Integrating Modelling with Breeding
Mace et al . 2012. Theor Appl Gen, 124:97-109
4. Trait Dissection and Phenotypic Prediction
Integrating Modelling with Breeding
SPb-80090.0SPb-5015 SPb-8773SPb-9854
11.2
SPb-495912.6SPb-049319.4SPb-375526.8SPb-446235.0
SPb-65770.0
SPb-804124.9SPb-638432.2
SPb-227073.6SPb-205893.4SPb-0232118.4SPb-8585129.8SPb-4542153.4SPb-0274168.7SPb-5358185.1SPb-2602200.8SPb-6600203.1SPb-7567 SPb-8492203.8SPb-4211223.6SPb-3311228.8SPb-2683 SPb-9203239.8SPb-2247 SPb-9497243.7SPb-4638248.1SPb-6772260.7SPb-0277271.1SPb-4593 SPb-0746273.1SPb-0383275.2SPb-2958277.3SPb-0233283.5SPb-1984302.5SPb-7611313.6SPb-8532314.2SPb-1060 SPb-3577314.9
Qsd
w1
.1
SBI-01
SPb-03330.0SPb-766819.8SPb-668923.9SPb-666631.5SPb-579333.7
SPb-379858.3SPb-946065.3
SPb-56430.0SPb-87664.4SPb-12235.7SPb-180128.3SPb-695939.9SPb-641651.9SPb-769565.1SPb-1920 SPb-643468.9SPb-122579.6SPb-788287.3SPb-2129 SPb-4561120.1SPb-8241121.4SPb-3917127.9SPb-3361132.7SPb-3124 SPb-6274134.2SPb-4111134.9SPb-1508141.6SPb-0903142.3SPb-5737 SPb-3834150.5SPb-9519156.9SPb-7100162.2SPb-4054170.2SPb-3547179.0SPb-0412195.3SPb-1671203.8SPb-4366216.8SPb-5544232.3SPb-8208 SPb-1215246.7SPb-5273252.1
Qrd
w1
.2
SBI-02
SPb-1828 SPb-14650.0SPb-79452.0SPb-404112.2SPb-090613.7SPb-238123.2SPb-288029.5
SPb-9080 SPb-5650SPb-5119
0.0
SPb-95441.6SPb-90096.3SPb-2162 SPb-261629.8SPb-526631.9SPb-589241.8SPb-6861 SPb-443352.1SPb-964252.8SPb-432359.1SPb-814965.6SPb-9312 SPb-789370.4SPb-020572.5SPb-4086101.7SPb-6323 SPb-7203107.8
Qsd
w1
.5
Qrd
w1
.5
Qra
1.5
SPb-62870.0
SPb-949036.7SPb-466441.5
SPb-341864.8SPb-789270.3SPb-139682.3SPb-0873 SPb-287483.2SPb-411992.1SPb-7639103.1
Qra
2.5
SBI-05
SPb-29560.0SPb-62488.1
SPb-6589 SPb-150321.7
SPb-7126 SPb-981843.8SPb-784345.9SPb-538748.8
SPb-93720.0SPb-4767 SPb-924223.3SPb-443225.4SPb-227133.7SPb-301446.7SPb-063349.4SPb-493452.3SPb-331365.0SPb-9700 SPb-2088SPb-1661
104.0
SPb-7889104.4SPb-7312119.2SPb-1881123.3SPb-6935130.9SPb-7864140.5SPb-0144 SPb-4546SPb-9743 SPb-0833SPb-3954
144.1
SPb-7823 SPb-0159174.8SPb-5501194.7SPb-8800199.5SPb-5572204.8SPb-6918207.5SPb-1825209.3SPb-3434212.0SPb-1291 SPb-1051217.2SPb-1272222.4SPb-7082222.7SPb-5250224.4SPb-7648224.8SPb-4385225.6SPb-9341 SPb-8160SPb-3510
227.8
SPb-3874231.6SPb-2487238.4
qrd
w1
.8
Qtla
1.8
Qtla
2.8
Qtla
3.8
Qra
1.8
SBI-08
SPb-68330.0SPb-20410.7SPb-16783.0SPb-83897.2SPb-104415.8SPb-033221.5SPb-672039.6SPb-2193 SPb-3432SPb-0248
57.7
SPb-959271.1SPb-591779.0SPb-081782.8SPb-629292.1SPb-365595.9SPb-1962103.5SPb-9275115.2SPb-9999125.5SPb-0943136.2SPb-6271142.7SPb-5841150.7SPb-9215 SPb-1722SPb-8432 SPb-3958
156.1
SPb-4480156.4SPb-8201161.8SPb-7643166.3SPb-4944 SPb-6875166.7SPb-1660202.4SPb-7058219.7SPb-6370222.7SPb-7412 SPb-1655243.0
Qra
1.1
0
SBI-10
SBI-01 SBI-02 SBI-05 SBI-08 SBI-10
R2=10%
R2=29.8%
R2=6.7%
R2=11.7%
In total, the 4 nodal root
angle QTL explained
58.2% of the phenotypic
variance
• Four major QTL explaining 58% variation in nodal root angle identified
• Root angle QTLs co-located with known QTLs for stay green
• Strong association between nodal root angle QTLs and grain yield data
from field trials; yield increase in presence of narrow root angle alleles
• Pursuing detailed genetic architecture by phenotyping NAMs
Root System Architecture - Genetics
Integrating Modelling with Breeding
Linking Trait Physiology and Genetics
with Phenotype
Sorghum staygreen - simulation
4. Trait Dissection and Phenotypic Prediction
Integrating Modelling with Breeding
Linking Trait Physiology and Genetics
with Phenotype
Sorghum staygreen - simulation
- Link genetic/QTL variation to trait variation
- Link trait variation to model to simulate dynamics
Hammer et al 2016. in : "Sorghum: State of the Art and Future Perspectives“, ASA-CSSA-SSSA
4. Trait Dissection and Phenotypic Prediction
tillering
TE
roots
Integrating Modelling with Breeding
Staygreen phenotype emerges as
consequence of model dynamics
Sorghum staygreen - simulation
Hammer et al 2016. in : "Sorghum: State of the Art and Future Perspectives“, ASA-CSSA-SSSA
4. Trait Dissection and Phenotypic Prediction
4. Trait Dissection and Phenotypic Prediction
Integrating Modelling with Breeding
Sorghum staygreen – long term simulation
Reflects observed yield advantage
pattern for staygreen
Dryland crop water balance
- Grain mass (M) is given by
M = T x TE x HI
- Simulations in sorghum suggest importance of limited max trans rate on TE and implications for HI
Yield = Resource Capture Resource Use Efficiency Partitioning
Integrating Modelling with Breeding
4. Trait Dissection and Phenotypic Prediction
Limited Max Trans Rate and
transpiration efficiency (TE) in maize
Sinclair et al 2005 Func Plant Biol
Integrating Modelling with Breeding
4. Trait Dissection and Phenotypic Prediction
Limited Max Trans Rate and transpiration efficiency (TE)
Does limited max trans rate increase TE and yield ?
Sinclair et al 2005 Func Plant Biol
Transpiration rate response of three
genotypes to increasing demand
(vpd) for sorghum genotypes.
Similar responses found in maize
Gholipoor et al (2010) Field Crops Res 119: 85-90
Integrating Modelling with Breeding
4. Trait Dissection and Phenotypic Prediction
Gholipoor et al (2013) J. Agron. Crop Sci. 199:155–160.
Duncan et al. 1967
Hammer & Wright 1994
Hammer et al. 2009
LAI=7
Jun 21
Pmax=2.11 (mg CO2 m-2 s-1)
Leaf Angle=68
Sinclair et al. 1984
k=9 kPa
vpd=3 kPa
T day = 9 mm
α T0
Integrating Modelling with Breeding
4. Trait Dissection and Phenotypic Prediction
Limited Max Trans Rate and TE in maize
- Trait evaluation for US corn belt
Messina et al (2015) Agron J 107:1978–1986
Integrating Modelling with Breeding
4. Trait Dissection and Phenotypic Prediction
Limited Max Trans Rate and TE in maize
- Trait evaluation for US corn belt
- Spatial simulation for reference hybrid
Messina et al (2015) Agron J 107:1978–1986
Integrating Modelling with Breeding
4. Trait Dissection and Phenotypic Prediction
Limited Max Trans Rate and TE in maize
- Trait evaluation for US corn belt
- Simulated yield difference with limited max trans
Messina et al (2015) Agron J 107:1978–1986
Median Best 25% Worst 25%
5. Where to from here?
Integrating Modelling with Breeding
Messina et al 2011 J Exp Bot
5. Where to from here?
Integrating Modelling with Breeding
Informing future crop improvement
- Some new thinking needed to integrate advanced technologies
5. Where to from here?
Integrating advanced technologies to hasten genetic gain
Integrating Modelling with Breeding
Advanced Research Projects Agency - Energy, US DoE
Cooper et al 2014 Crop & Past Sci; Technow et al 2015. Plos One
• Integrate data streams (genotype & phenotype) with biological insight
and predictive (modelling) capabilities – adding more power to selection
and capacity to search adaptation landscapes
“As simple as possible,
but no simpler!”
Acknowledgements
UQ QAAFI
Erik van Oosterom
David Jordan
Andrew Borrell
Vijaya Singh
Al Doherty
Kurt Deifel
CSIRO
Scott Chapman
DAF Qld
Greg McLean
Emma Mace
Pioneer Hi-Bred
Charlie Messina
Mark Cooper
Integrating Crop Physiology and
Modelling with Genetic Improvement
Graeme Hammer
Centre for Plant Science, QAAFI
The University of Queensland
