Applying Molecular Marker information and Data Mining to a Commercial Breeding Pipeline

Transcription

1 Applying Molecular Marker information and Data Mining to a Commercial Breeding Pipeline David V. Butruille, Ph.D. Data Mining Lead - Breeding Technology Monsanto Company Ankeny, IA ISU Plant Breeding Lecture Series May 19,

2 Overview Background Key processes and tools Examples of data mining and marker applications 2

3 A company focused on agriculture Headquartered in St. Louis, Missouri CEO Hugh Grant Monsanto 2005 Net Sales More than 14,000 employees More than 320 locations worldwide in four primary regions Europe/Africa, Asia Pacific, Latin America and North America Seeds & Genomics Ag Productivity $6.3 billion in annual sales NYSE Symbol: MON A leader in the field of plant breeding, agricultural biotechnology and genomics. 3

4 Our technology platforms Monsanto uses its scientific capabilities in a number of arenas to develop new products. These capabilities form a toolkit used throughout our product pipeline, as each tool can be used on its own or in combination to help study, test and refine new traits, germplasm and other characteristics in plants. GENOMICS This talk s focus Genomics involves the mapping of the genes of plants to understand their structure and the role they play in how the plant functions. CONVENTIONAL BREEDING Conventional breeding is the process of cross-pollinating plants with desirable qualities to develop improved plants in successive generations MOLECULAR BREEDING Molecular breeding involves the use of DNA markers for genes in combination with physical measurement of traits to manage plant breeding programs. Molecular breeding significantly accelerates the efficiency in bringing new varieties to market. CROP ANALYTICS Crop analytics involves the application of advanced analytical methods and technologies to identify the composition of food and feed traits. BIOTECHNOLOGY Biotechnology is the application of scientific knowledge to transfer beneficial genetic traits to enhance plants growth or to provide nutritional or other benefits to farmers, food and feed processors, or consumers. 4

5 Average U.S. corn yields and kinds of corn, Civil War to 2004 Source: Troyer, AF Adaptedness and Heterosis in Corn and Mule Hybrids. Crop Science 46:

6 Technologies applied to breeding evolves over time Impact of Technology on Plant Breeding by Decade Technology Statistics, Experimental Design Low Mod Mod High High High High Computer Technologies None None Low Mod Mod High High Quantitative Genetics Mod Mod High High High High High Plot Mechanization None None Low Low Mod High High Winter Nurseries None None Low Mod High High High Analytic Technologies Low Low Low Low Mod High High Doubled Haploids Low Mod Mod Low Mod High High Molecular Breeding None None None Low Mod High High Genomics/Proteomics None None None None None Low High Bioinformatics None None None None None Low High Transgenic Traits None None None None Mod High High # Highs Statistics and Quantitative Genetics have been important for a long time, and that will not change any time soon 6

7 Traditional breeding process Year 1 Choose parents based on data, judgment X Recombine genes Years 2-6 Select best based on phenotype Advance Inbreeding Advance Inbreeding Advance Inbreeding Advance Inbreeding Make hybrids, Yield test Select, Make Hybrids Yield test Select, Make Hybrids Yield test 6- to 8-year process Advance Inbreeding Select, Make Hybrids Year 7 Success rate <.01% Advanced Trials, Commercialize 7

8 Ranges for key parameters of a global corn breeding program. Pipeline Step Breeding Populations Segregating Lines Finished Lines Test Crosses Finished Hybrids Nursery Rows Yield Trial Plots Number of Items 2,000 5, , ,000 10,000 30, , ,000 20,000 50,000 1,000,000 2,000,000 2,000,000 4,000,000 8

9 Some of the key components: -Phenotype -Cycle time -Genotype -Information technology 9

10 Phenotypic data Quality, accuracy, and throughput 10

11 State-of-the-art field plot equipment mimics best farming practices. Increase in plot testing capacity High quality phenotypic data 11

12 Properly identify and screen for performance reducers Stewarts Wilt Gray Leaf Spot Green Snap Diplodia Ear Rot Northern Leaf Blight FAW Etc.. ECB Common Rust Southern Leaf Blight BCW CRW Southern Rust Anthracnose Leaf Blight Anthracnose Stalk Rot 12

13 Quality Traits: Phenotype is not measured in the field only 13

14 MSP Multi Season Program 14

15 MSP dramatically accelerates the product development Breeding Corn Soybean Sunflower Sorghum Winter Breeding Corn Soybean Sunflower Sorghum Historically breeding programs used MSP Sites to grow a second generation during the winter season decreasing development time by 50% 15

16 Major MSP sites are now focal points for many areas and conduct research year round Breeding Southern Hemisphere Several Crops Biotechnology Yieldguard CB Yieldguard CRW Roundup Ready Etc. Multi Gen Single Gen Winter Spring Year Round Nurseries Genomics Summer Multi Gen Trait Integration Multi Gen/Yrs Markers Multi Gen Fall Single Gen Breeding Northern Hemisphere, Several Crops 16

17 DNA Markers Getting to the Genotype 17

18 18 We have identified and developed assays for thousands of corn SNPs X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC X-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC X-NC X-NC X-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC X-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC X-NC X-NC X-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC X-NC X-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC X-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC X-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC X-NC Q-NC X-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC X-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC X-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC 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Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC X-NC X-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC 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Q-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC X-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC X-NC X-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC X-NC X-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC X-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC X-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC X-NC X-NC X-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC X-NC X-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC X-NC X-NC X-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC X-NC X-NC X-NC X-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC X-NC X-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC X-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC X-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC X-NC X-NC X-NC X-NC X-NC Q-NC X-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC X-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC Q-NC Q-NC Q-NC X-NC Q-NC X-NC Q-NC X-NC Q-NC Q-NC X-NC Q-NC Q-NC Utilized high resolution mapping populations Large percentage of SNPs are in genes

19 RFLP SSR SNP? Key Factors Affecting Volume and Cost SNPs Automation Reagent cost reduction IT systems Genotyping platform changes enable growth in marker assisted breeding applications Data points (M) '98 '99 '00 '01 '02 '03 '04 '05 '06 Calendar year Cost per DP ($) VIC FAM 19

20 Marker Assisted Selection; Replacing phenotyping with genotyping Ol d Susceptible Resistant New Latest 1.2 Susceptible Resistant VIC FAM 20

21 Ranges for key parameters of a global corn breeding program. Pipeline Step Breeding Populations Segregating Lines Finished Lines Test Crosses Finished Hybrids Nursery Rows Yield Trial Plots Number of Items 2,000 5, , ,000 10,000 30, , ,000 20,000 50,000 1,000,000 2,000,000 2,000,000 4,000,000 Marker Applications Parental Selection Screen Lines at Specific Loci Select on Marker & Phenotype Choice of Testers Hybrid Prediction Quality Assurance 21

22 Multiple Impacts of Molecular Markers on Plant Breeding Marker Assisted Backcrossing Quality Assurance: Parent-offspring tests Genetic purity tests Event tests Germplasm analysis Fingerprinting germplasm Arrangement of diversity (clustering, PCA, etc.) Protection of intellectual property. Trait Analysis Association of traits with genomic regions ( QTL-mapping ) Understanding of relationship among traits (linkage vs. pleiotropy?) Understanding the causes of variation (aid in gene cloning) Breeding Alternative or support to selection for qualitative and quantitative traits Increase rate of genetic gain: Selection during off-season cycles Selection of hybrid traits on inbred individuals Early selection (e.g. pre-flowering) 22

23 Breeding IT Putting it all together 23

24 Global field breeding IT system: collection of phenotypic information All crops & all world regions Manage breeding process Create breeding pops Organize and execute field trials QC and analysis of phenotypic data Global standardization Pedigree system Data attributes Centralized database Every plant is uniquely identified Global laboratory IT systems: collection of genotypic information All genotyping labs Manage genotyping lab Marker inventory Marker selection Sample tracking QC and analysis of genotypic data Global standardization Methodologies Nomenclature Centralized database Every sample is uniquely identified 24

25 Created an integrated data analysis and decision making system Field System: Phenotypic data collection, and analysis Genotyping System: Genotypic data collection, and analysis Data Decisions Analysis System: Integrate and analyze phenotypic and genotypic data Data 25

26 Monsanto designed the organization to support the breeding strategy New Technology BREEDING TECHNOLOGY LINE DEVELOPMENT BREEDING COMMERCIAL BREEDING CHANNEL DEPLOYMENT Marker Labs Statistics Pathology Winter Nurseries Discovery Breeding Trait Integration F1 to National Testing Technology Optimization Product Characterization Channel Tech Support Testing QA/QC PRODUCT MANAGEMENT Partitioned the process into components to enable optimal execution 26

27 Data mining: The environment 27

28 Being alert for, understanding and exploring patterns in the environment at multiple granularities Detecting plot level anomalies and interactions Exploring patterns of variability and interaction in a target population of environments Leveraging commonalities and accounting for differences at a global scale 28

29 Understanding of what happens in the field Cause: Anhydrous ammonia applied diagonally, malfunction for one pass through the field, reduced the grain yields in that strip to harvest From planting time Stuck TW bucket door, grain dribbling through bucket. The two column pattern shows this problem with one side of the combine 29

30 Some systematic field variation can be accounted for Yield Trial Residual Maps Residuals With Field Trend BAD Adjusted Data GOOD 30

31 Corn testing site map 31

32 Leveraging environmental data Published Pioneer Hi-Bred work, using GIS data (mostly precipitation, temperature and day length) to classify environments and better understand underlying reasons for of observed genotype- by-environment interactions. Environment-standardized GGE biplot of grain yield of 18 maize hybrids (H1-H18) grown in 266 environments over three years, stratified by environment class Geographic distribution and variability of five abiotic environment classes (EC) in the United States for CRM 110 maize in Source: Loffler, CM, &al. & Cooper, M Classification of Maize Environment Using Crop Simulation and Geographic Information Systems. Crop Science 46:

33 Global corn germplasm l l l l Tropical Subtropical Temperate High Lands Temperate Subtropical Tropical Subtropical Temperate 33

34 Traits under selection vary based on environment Trait Temperate Sub-Tropical Tropical Yield Standability Dry down Maturity Stability Tolerance to abiotic stress Disease Tolerance Grain Quality Key traits under selection in a NA corn breeding programs Yield, Yield, Yield Agronomic traits based on breeding target market. Tolerance to root lodging, stalk lodging, dry down, early vigor, etc. Disease Traits Rusts, stalk rots, ear rots, etc. Consistency over diverse growing conditions. Production characteristics (COGs). 34

35 Identify global QTLs Directional consistency of QTLs between US and Europe 20 Europe effect Parent A favorable Selection index score (heavily weighted by yield) -10 Parent B favorable U.S. effect -24/35 loci signif. in both continents -8 U.S. locations and 3 E.U. locations 35

36 Data mining: The base germplasm 36

37 Mining information from base germplasm Compilations from the literature Linkage disequilibrium Association tests 37

38 Lineage of elite Lancaster, Oh07-Midland, and Oh43 corn inbred lines Data Mining is not always about applying of sophisticated algorithms. Carefully compiling available information can lead to interesting insights. Refereed papers are one source, not the only one. Example of other sources: - PVP certificates, - Patent applications, - Public disclosure to regulatory agencies. Source: Mikel, MA and Dudley, JW Evolution of North American Dent Corn from Public to Proprietary Germplasm. Crop Science 46:

39 Linkage disequilibrium in haploids extracted from old open-pollinated maize varieties and synthetics GOALS: Assess amount of linkage disequilibrium within Old OP material? Use haploid plants to avoid linkage phase estimation. Compare doubling efficiency for different germplasm types. Key originating germplasm Reid Krug Minnesota 13 Lancaster Leaming Midland Cultivars of local importance Alph Golden Glow Goldmine Hays Jellicorse Johnson County White Longfellow Murdock Osterland Synthetics (from inbreds) Corn Borer Synthetic (BCBS) Stiff Stalk Synthetic (BSSS) Pioneer Long Ear Two Ear Synthetic Long Ear Synthetic 39

40 LD of different Group of OP germplasm R-squared (200 obs moving avg) Group 1 Pop's Group 3 Pop's Standard F2 Group 2 Pop's Pairwise Distance (cm) Population name Group for R2 analysis Alph 1 Pion Long Ear 1 Two Ear Synth 1 Osterland 1 BSSS 1 Hays 1 Johnson County White 1 Long Ear Synth 1 BCBS 2 Longfellow 2 Murdock 2 Goldmine 2 Golden Glow 2 Jellicorse 2 Midland 3 Krug 3 Leaming 3 Minn 13 3 Lancaster 3 Reid Yellow dent 3 40

41 Table 1.Segment Genotyped Chr. Segment Begin pos. End pos. # loci Population Type Population name # Haploid plants # Loci # Polym. loci % seg. loci Avg 2p(1-p) across polym. loci Main OP Krug % 4% Main OP Lancaster % 35.9% Main OP Leaming % 36.1% Main OP Midland % 35.2% Main OP Minn % 37.8% Main OP Reid Yellow dent % 37.0% 2ary OP Alph % 29.3% 2ary OP Golden Glow % 37.2% 2ary OP Goldmine % 35.3% 2ary OP Hays % 35.6% 2ary OP Jellicorse % 38.4% 2ary OP Johnson County White % 36.7% 2ary OP Longfellow % 33.6% 2ary OP Murdock % 36.8% 2ary OP Osterland % 34.8% Synthetic BCBS % 37.9% Synthetic BSSS % 38.0% Synthetic Long Ear Synth % 35.4% Synthetic Pion Long Ear % 31.4% Synthetic Two Ear Synth % 32.8% If we define overall efficiency as: Ear with putative di-haploid seed obtained / Total number of kernel screened. The ratio was [1:4:24] for [OP varieties : Synthetics : Process checks]. 41

42 Finding marker-trait associations in the base germplasm A1 A2 B1 B2 C1 C2 D1a D1b D2 E F G H I J K L M N O 10 cm 658 soybean lines genotyped (either Roundup Ready or conventional) Phenotypic data was tolerance to Roundup herbicide (h 2 = 1) Marker association at p-value < 1 x associations on 16 chromosomes 42

43 Finding marker-trait associations in the base germplasm A1 A2 B1 B2 C1 C2 D1a D1b D2 E F G H I J K L M N O RR 10 cm Marker association at p-value < 1 x soybean lines genotyped (either Roundup Ready or conventional) 47 associations on 16 chromosomes Phenotypic data was tolerance to Roundup herbicide (h 2 = 1) Identified only the true association when analysis accounted for population structure 43

44 Data mining: Segregating populations 44

45 Segregating populations How much of the variation do we truly explain with QTL? 45

46 Watch out: we see more than what we get! Schön et al. (2004) Very large population (975 F5 testcrosses in 19 environments) and simulated populations QTL analyses (PLABQTL) 2 R adj Obtained proportion of phenotypic variance explained by QTLs ( ) Derived proportion of genotypic variance explained by all detected QTLs: pˆ = Rˆ 2 hˆ adj 2 Data subdivided to verify impact of number of progenies and number of environments. Used resampling techniques to estimate the amount of bias in detecting QTL (comparing R 2 and P in estimation and test data sets (ES vs. TS). Source: Schön, C.C. et al Quantitative Trait Locus Mapping Based on Resampling in a Vast Maize Testcross Experiment and Its Relevance to Quantitative Genetics for Complex Traits. Genetics 167:

47 Figure 1. Proportion of the genotypic variance explained by detected QTL in estimation sets averaged over all data sets (ES) for 12 combinations of experimental data PED (N, E), using fivefold standard cross-validation and two significance levels for grain yield, grain moisture, and plant height. Individual columns are partitioned into the genotypic variance explained in test sets ( TS, solid bottom) and the bias calculated as the difference ES TS (shaded top). Source: Schön, C.C. et al Quantitative Trait Locus Mapping Based on Resampling in a Vast Maize Testcross Experiment and Its Relevance to Quantitative Genetics for Complex Traits. Genetics 167:

48 Figure 2. Mean ( ), median (o), and 12.5 and 87.5% quantiles of the proportion of the genotypic variance explained in test sets calculated for individual data sets for 12 combinations of experimental data PED (N, E) using fivefold standard cross-validation and LOD 2.5 for grain yield, grain moisture, and plant height. Figure 3-left. Mean ( ), median (o), and 12.5 and 87.5% quantiles of the bias (ES TS) calculated for individual data sets for 12 combinations of experimental data PED (N, E), using fivefold standard cross-validation for grain yield, grain moisture, and plant height. Source: Schön, C.C. et al Quantitative Trait Locus Mapping Based on Resampling in a Vast Maize Testcross Experiment and Its Relevance to Quantitative Genetics for Complex Traits. Genetics 167:

49 Main conclusions of Schön et al. Adding more genotypes is more efficient than replicating the same genotypes (provided that a minimum number of environments are sampled) Knowledge about the power of detection of these experiments (previous discussion) + low values of unbiased estimates of genotypic variance explained: confirm large number of genes with small effects model. The use of this tool may have to be trait specific Source: Schön, C.C. et al Quantitative Trait Locus Mapping Based on Resampling in a Vast Maize Testcross Experiment and Its Relevance to Quantitative Genetics for Complex Traits. Genetics 167: What power a large breeding program can afford us (under very simplistic assumptions)? 49

50 Power calculations taken to a different scale Using the formula that relates sample size N needed, at a α and β error level, to detect an effect explaining a given proportion r 2 of the phenotypic variance: Fixing α=1 and β=5 With 200 lines With 1,000 lines With 10,000 lines With 100,000 lines r 2 = 8.6% r 2 = 1.8% r 2 = 0.18% r 2 = 18% Percent Phenotypic variance 1000% 100% 1.000% 0.100% 10% 01% ( 2 1 r ) z 1 [ α 2] Number of Lines ( ) = 2 + z 2 1 r ,000 10, ,000 1,000,000 10,000,000 Source: Walsh, B. and Lynch, M Genetics and Analysis of Quantitative Traits. N (for r 2 small N~1/r 2 ) r PhD Breeding program Entire company ( 1 β ) 50 2

51 Segregating populations Leveraging previously acquired knowledge on marker-trait associations 51

52 We continue to expand our knowledge Continue to add to this information each year ~No. of Marker-Phenotype Associations Over 40 Fold Increase * *estimate 52

53 Selection based on predictive associations Proof of concept Selection for grain moisture at harvest Favorable Unfavorable Change in Grain Moisture (%) QTL identified in a breeding population Parent 1 best Parent 2 best % Change 3.9% Change Hybrid Inbred 53

54 New data views combined with predictive breeding methodologies create new opportunities to drive commercial success Multiple data cuts on phenotypic information Create new designer germplasm New breeding paradigm Explore genetic relationship of germplasm Genome wide trait QTL knowledge 54

55 Segregating populations Epistatic decay Using the data available without doing the right experiment 55

56 Rationale (this is a simplistic generation means analysis) One can expect that elite breeding lines accumulate favorable effects (additive or dominant) that act additively across loci. If these are the only ones then: In a Breeding Cross: Mean(Parent x Tester)=Mean(Progeny x Tester) Can we expect that these lines also accumulate a greater amount of positive epistatic interactions than negative ones? In a Breeding Cross: Mean(Parent x Tester)>Mean(Progeny x Tester) If a positive epistatic balance is detected can we explore this further? 56

57 Possible experiment: Several parents (total: n) Randomly paired Derived two random progenies per cross (total: n) Create a 2n-entry replicated yield trial, at multiple locations and over multiple years, and common tester(s), RCB or Split- Plot. Analyze. 57

58 Alternative approach through data mining Query company data base. Find first generation yield trials with parents included in trial, assuming: No intra-plot competition issues. No negative selection in nursery. Compile difference between mid-parent and mid-progeny on a breeding cross basis. Average difference over crosses. Analyze (overall mean and multiple data cuts). 58

59 The Data Data used: 479 projects Year Count Total 479 Cross Heterotic Group Count BC1 F 50 M 48 F2 F 191 M 190 Analysis: Ep = C 0 MP C 0 = C0 MP = Mid mean = n i= 1 n P1+ P 2 parent = P1+ l for F 2 RP + Factor considered: Year Heterotic Group Cross Type for BC Similarity between parents 59

60 Results I Proc glm data=test; class hg year cross_type; model p_yld =year hg cross_type simil/ss1; quit; R-Square Coeff Var Root MSE P_YLD Mean Source DF Type I SS Mean Square F Value Pr > F year HG cross_type Simil <

61 Results II Cross Type Year C0-P (Bu/acre) BC BC1 Total F F2 Total Grand Total