Plant Breeding Core Concepts

Core Concepts and Competencies background.

All + All -

MS_Plant Breeding

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    AGRON 506 - Crop Genetics

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      1. Reproduction in crop plants - Flower morphology and distribution

      • Describe the parts of a flower and define complete, incomplete flowers, perfect and imperfect flowers

        Describe flower distribution in a plant: monoecious, dioecious and other variants

        Describe the types of reproduction in crop plants: sexual and asexual

        Describe what vegetative propagation is and the common type of propagules used for seed increase

        Describe what apomixis is and the genetic nature of an apomictic cultivar

        Describe the G1, S, G2 and M phases of a cell cycle

        Describe mitosis and meiosis, where and when does each occur and highlight the key differences between mitosis and meiosis

        Describe what homologous chromosomes are

        Describe, using diagrams, the sequence of events involving DNA in meiosis from chromosome duplication through chromosome segregation.

        Distinguish between sister chromatids and homologous chromosomes.

        Describe the process for male and female gamete formation

        Describe the double fertilization process and seed formation

        Describe different types of pollination methods and their implications on plant breeding strategies and cultivar increase

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      2. Controlled hybridization

      • Describe gametophytic and sporophytic self-incompatibility, and their use for controlled pollination and hybrid production

        Describe methods to overcome self-incompatibility in gametophytic and sporophytic systems

        Describe genetic male sterility and how sterility is maintained and used for cultivar development

        Describe cytoplasmic male sterility and how sterility is maintained and used for F1 hybrid production including the use of A, B and R lines

        Explain how sex is genetically determined in crop plants and how gynoecious cucumber or all-male asparagus cultivars are developed?

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      3. Gene segregation and recombination

      • Describe the classic definition of gene as an inherited factor that determines a characteristic

        Describe the concept of allele and the number of alleles of a diploid or polyploid individual or a population can have for a particular locus

        Describe the type of gene actions: complete, partial and overdominance and additive gene actions

        Describe genotype and phenotype and their relationship

        Describe penetrance and expressivity, two related yet different concepts

        Describe Mendelian’s first law, principle of segregation and the second law, principle of independent assortment

        Describe the use of Punnett square or rule of probability (multiplication or addition) to predict outcomes of a genetic cross at F1, F2 and beyond for monohybrid, dihybrid or multiple pairs of genes

        Explain how independent assortment of alleles during meiosis can lead to new combinations of alleles of unlinked genes

        Using pedigrees, distinguish between dominant, recessive, and cytoplasmic modes of inheritance

        Predict the transmission of phenotypes associated with maternal effect genes

        Explain why the terms “dominant” and “recessive” are context dependent and may differ at the cellular level or at the level of a pedigree

        Design genetic crosses to provide information about genes, alleles, and gene functions

        Interpret the results of experiments comparing the phenotypes that result from single mutations in two different genes with the phenotype of the double mutant, contrasting epistatic and additive interactions

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      4. Linkage and gene mapping 

      • Describe linked genes and linkage phases (repulsion or coupling)

        Describe crossovers between linked genes

        Diagram the process of homologous recombination during meiosis and explain how it can lead to new combinations of linked alleles

        Explain how a specific combination of linked alleles (haplotype) can persist through many generations (linkage disequilibrium)

        Calculate gene linkage and genetic map distances and interference from the frequencies of progeny with recombinant phenotypes from genetic crosses

        Explain how genetic distance is different from physical distance

        Calculate the probability of a particular gamete being produced from an individual, provided map distance

        Explain how to use 2 point and 3 point analysis to construct genetic maps and determine gene orders

        Describe type of DNA markers, SNP, SSR

        Describe applications of DNA markers in map construction, MAS, fingerprinting, studying genetic diversity

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      5. Chromosome variation and polyploidy

      • Explain the meaning of ploidy (haploid, diploid, aneuploid etc.) and how it relates to the number of homologues of each chromosome

        Describe various polyploids (triploids, tetraploids, hexaploids etc) and the difference between auto- and allo-polyploids

        Explain how polyploids are naturally and artificially created

        Describe the genetics of polyploids, i.e. how genes are segregated in autotetraploids

        Describe different methods of generating haploids or polyhaploids and consequent chromosome doubling for inbred development

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      6. Chemical nature of gene, DNA replication, transcription and translation

      • Explain how DNA was found to be the source of genetic information

        Describe the chemical composition and structure of DNA and explain the key characteristics that made DNA an ideal molecule for storing and transmitting genetic information

        Discuss how DNA is packaged in the chromosomes in terms of histones, nucleosomes, and chromatin

        Explain the functional significance of packaging DNA into chromosomes and the lack of correlation between chromosome size and genetic information content

        Describe how the positions of individual genes on a given chromosome are related to their positions on the homolog of that chromosome

        Differentiate between a gene and an allele, including the recognition that genes may have many alleles

        Draw a simple line diagram showing a segment of DNA from a gene and its RNA transcript, indicating which DNA strand is the template, the direction of transcription and the polarities of all DNA and RNA strands

        Draw a simple line diagram showing the important components of a gene including the promoter, coding sequence, terminator, regulatory sequences

        Describe DNA replication, transcription and translation and where each process within the cell occurs

        Describe the types of DNA regions that do not encode proteins: the general organization, possible function, and frequency of genes and non-gene DNA sequences in a typical eukaryotic genome

        Contrast the packaging of DNA into euchromatin versus heterochromatin in the context of histone modification, and DNA modification (where applicable)

        Discuss the potential roles of DNA modification, histone modification, and non-coding RNA in epigenetic inheritance, both somatic and germline

        Discuss environmental impacts on epigenetic systems

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      7. Molecular Genetic Analysis 

      • Describe the principle of PCR and how it can be used to amplify a specific DNA fragment from a genome

        Describe the importance of restriction enzymes in recombinant DNA technology and gene cloning

        Describe the principles of various DNA sequencing technologies

        + - Discuss comparative genomics as it relates to use of comparative data from multiple species to identify which regions of a protein, pathway, regulatory system etc. are critical for function

        • Describe what transcriptomics and proteomics are and how they can be used to identify candidates that affect a characteristic

      + - 8. Mutation and biotechnology - How different types of mutations affect genes and the corresponding mRNAs and proteins

      • Describe how mutations arise and how environmental factors or mutagens (chemical and physical) can increase mutation rate

        Cite examples of mutations that led to crops with novel traits such as shattering resistant cereals and seedless fruits

        Describe somatic and germinal mutations and their implications in plant breeding

        Compare point mutations with chromosome mutations

        Describe how duplications, deletions, inversions, and translocations can affect gene function, gene expression, and genetic recombination

        Describe the same for transposable elements (How transposable elements can affect gene function, gene expression, and genetic recombination?)

        Distinguish between loss of function and gain of function mutations and their potential phenotypic consequences

        Predict the most likely effects on protein structure and function of null, reduction-of-function, overexpression, dominant-negative and gain-of-function mutations

        Describe how to obtain transgenenic plants using Agrobacterium-mediated genetic transformation method and explain the importance of a selectable marker gene

        Describe targeted genome editing using CRISPR/Cas9 and how this might lead to transgene-free mutants

      + - 9. Population Genetics

      • Describe the mechanisms by which variation arises and is fixed (or lost) in a population over time

        Calculate allele frequencies based on phenotypic or genotypic data for a population, and be able to explain the assumptions that make such a calculation possible

        Model how random mating yields predicted genotype frequencies in Hardy-Weinberg Equilibrium (HWE), and how non-random mating affects allele and genotype frequencies

        Test whether HWE has been reached in a population

        Explain how inbreeding increases the number of homozygotes in comparison to HWE

        Explain how natural selection and genetic drift can affect the elimination, maintenance or increase in frequency of various types of alleles (e.g. dominant, recessive, deleterious, beneficial) in a population

        Explain how artificial selection against recessive alleles affecting allele frequencies by removing homozygous recessive individuals either before or after flowering in open pollinated species

        Explain why undesirable recessive allele will persist despite intensive selection against it

        Interpret experiments to determine the relative influences of genes and the environment on a given phenotype

        Describe how variation can be measured, and what can be done to distinguish genetic and environmental sources of variation

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      10. Quantitative genetics 

      • Explain how continuous traits are the result of many different gene combinations that can each contribute a varying amount to a phenotype

        Evaluate how genes and the environment can interact to produce a phenotype

        Describe/review basic statistical concepts that are used to analyze quantitative traits: frequency distribution, samples and populations, mean, variance and standard deviation, correlation coefficient, regression coefficient

        Describe broad-sense and narrow sense heritability

        Describe the limitations of heritability: 1) heritability does not indicate the degree to which a trait is genetically determined; 2) an individual does not have heritability; 3) there is no universal heritability for a trait; 4) heritability is specific for a given trait in a given environment

        Describe and explain how to calculate response to selection using the breeder’s equation R=h2S

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    AGRON 537 - Quantitative Analytics for Plant Breeding

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      1. Introduction and Review

      • Review and solve Linear Algebra

        Define and give examples of discrete and continuous variables

        Define sampling units and experimental units 

        Discuss different trait measurements

        Write and explain trait models

        Organize of data for analysis and presentation

        Conduct exploratory data analyses (EDA)

        List and discuss Statistical Inference parameters

        • Estimation, Prediction and Validation, Hypothesis tests
        • Precision, Power, Accuracy
        • Sensitivity, Specificity
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      2. Sampling and Experimental Designs 

      • Discuss the differences between sampling units and experimental units

        Describe the features of Population structures + -

        Discuss the differences between Treatment Design and Experimental Design

        • Factorial and incomplete factorial designs
        • + - Balanced designs
          • CRD
          • RCBD
        • + - Incomplete Block Designs
          • Lattice
          • Alpha-lattice
          • Augmented
        • + - Split Plot Designs
          • Multi-environment trials

        Translate designs to models

        • Fixed, Random and Mixed models
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      3. Exploratory Data Analyses of discrete metrics (genetic markers, genetic sequences and gene expression)

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        Describe Binary (binomial and Poisson distributions)

        • Estimates of frequency, proportions, variances, covariance
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        Distinghuish Nominal and ordinal (multinomial distributions)

        • Estimates of frequency, proportions, variances, covariance

        Write appropriate Models for EDA

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      4. Inferential Data Analyses of discrete metrics (genetic markers, genetic sequences and gene expression)

      • Write correct Expectation | model (inheritance) and Chi-Square

        Illustrate and solve Hidden Markov Models BLAST (genetic sequence alignment)

        Determine/estimate Genetic Linkage 

        Describe Population Structure (PCA, Cluster Analyses)

        Explain Generalized Linear Regression 

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      5. Exploratory Data Analyses of continuous metrics (height, yield, maturity)

      • Construct and interpret Box plots, histograms, pivot tables, biplots

        Estimates of mean, variance, standard deviation, standard error

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      6. Inferential Data Analyses of continuous metrics (height, yield, maturity)

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        Distinguish General Linear Models from generalized linear regression

        • Correlation, Regression, Analysis of covariance
        • Mean Comparisons
          • t-tests, ANOVA
        • Combined Analyses of Genotype x Environment
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        Explain difference between Mixed Linear Models: BLUEs and BLUPs and linear models

        • Fixed effects, random effects, covariates, covariance structures
        • Combined analyses of unbalanced data from multiple environments
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    AGRON 520 - Plant Breeding Methods

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      1. Plant Breeding - Inroduction

      • + - Basic Concepts/Principles
        • List the basic steps of plant breeding
        • Explain why plant breeding is important
        • Explain how plant breeding has changed from the 1950’s to the present
      • + - Breeding Objectives
        • Define what yield is
        • List traits that may contribute to yield increases/decreases in different crops
        • Compare vertical and horizontal disease resistance
      • + - Germplasm and Genetic Diversity
        • Explain how genetic variation is produced; which of these can be manipulated by a breeder

          List different sources of germplasm

          Summarize the challenges faced by plant breeders (answers should be based on a podcast by Dr. Baenzinger and the CAST Issue paper #57, Plant Breeding and Genetics)

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      2. Reproductive Biology - Review 

      • + - Floral morphology
        • Explain how different flower types (perfect vs. imperfect) and distribution (monoecious, dioecious) affect the type of cultivar (i.e., pureline, hybrid, synthetic, etc.) that is developed for different crops

          Classify floral features (i.e., cleistogamy, staminal columns, etc.) based on whether they promote/inhibit a particular mode of reproduction (i.e., self vs. cross-pollination)

      • + - Reproduction
        • Describe how the mode of reproduction (self- vs. cross-pollinations) affects breeding practices
        • Describe what apomixis is and how it could be used by breeders to produce hybrid seed
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      3. Genetics - Review

      • + - Basic concepts
        • Compare and contrast qualitative vs. quantitative inheritance

          Describe Mendelian’s first (principle of segregation) and second (principle of independent assortment) laws

          Explain how independent assortment of alleles during meiosis can lead to new combinations of alleles of unlinked genes

          Demonstrate how a Punnett square or rule of probability (multiplication or addition) is used to predict the outcome of a genetic cross during the F1, F2 and beyond for monohybrid, dihybrid, and multi-gene crosses

          Describe how epistasis occurs (via the interaction of genes) and how this alters expected phenotypic ratios

          Explain how pleiotropy and lethality alter expected phenotypic ratios

          Explain what it means when genes are linked (in repulsion or coupling) and how this affects the types of gametes produced via meiosis and how this affects decisions made by a plant breeder

          Describe how recombination frequency and the observed frequency of progeny produced by a test cross can be used by a plant breeder

          Demonstrate how to use a chi-square test to confirm the presence of linkage or epistasis

      • + - Application of Genetic Concepts to Plant Breeding
        • Demonstrate how phenotypic and genotypic frequencies of single gene and multiple (unlinked or linked) genes changes after selfing for 1 or more generations with and without selection

          Show how homozygosity and heterozygosity change after selfing

          Demonstrate how proper F terminology is used to describe individual plants, derived lines, and populations

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      4. Quantitative Genetics - Review

      • Explain how quantitative inheritance differs from qualitative (Mendelian) inheritance

        Define the concept of transgressive segregation

        Describe gene action

        List the different genetic variance components 

        Define heritability (narrow sense and broad) and demonstrate how it is calculated using variance components or regression analysis

        Describe how breeder can increase heritability of a trait and then compare which approach is best for different scenarios

        Explain how a breeder can manipulate the different components of the genetic gain equation (e.g., selection intensity, heritability, phenotypic variance) to improve their expected gain

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      5. Breeding Methods

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        Mass Selection

        • Explain what mass selection is and why is it used by plant breeders

          Outline the steps of mass selection

          Compare and contrast different examples where mass selection was implemented and discuss why certain steps were taken

          Describe the advantages and disadvantages/limitations of mass selection

          Describe a scenario where a breeder could/should use mass selection

        + - Pedigree Selection

        • Explain what pedigree selection is and why is it used by plant breeders

          Outline the steps of pedigree selection

          Compare and contrast different examples where pedigree selection was implemented and discuss why certain steps were taken

          Describe the advantages and disadvantages/limitations of pedigree selection

          Describe a scenario where a breeder could/should use pedigree selection

        + - Bulk Population

        • Explain what bulk population is and why is it used by plant breeders

          Outline the steps of bulk population

          Compare and contrast different examples where bulk population was implemented and discuss why certain steps were taken

          Describe the advantages and disadvantages/limitations of bulk population

          Describe a scenario where a breeder could/should use bulk population

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        Single Seed Descent (SSD)

        • Explain what single seed descent is and why is it used by plant breeders

          Outline the steps of single seed descent

          Compare and contrast different examples where single seed descent was implemented and discuss why certain steps were taken

          Describe the advantages and disadvantages/limitations of single seed descent 

          Describe a scenario where a breeder could/should use single seed descent

          Compare and contrast pedigree selection, bulk population, and single seed descent

        + - Doubled Haploids (DHs)

        • + - Self-pollinated Crops
          • Explain what doubled haploids are how/why they are used by plant breeders
          • Outline the steps used to create doubled haploids via anther/pollen culture and wide hybrid crosses
          • Describe the advantages and disadvantages/limitations of utilizing doubled haploids
          • Describe a scenario where a breeder could/should use doubled haploids
        • + - Maize Inducer Lines
          • Outline the steps followed to create doubled haploids of maize using haploid inducer lines

            Describe how breeders use pigment genes carried by the inducer line to differentiate between selfs, F1 hybrids, and haploids

        + - Backcrossing (BC)

        • Explain why backcrossing is utilized by plant breeders

          Outline in detail the steps that are followed when introgressing a dominant trait via backcrossing when the trait can be evaluated before flowering and when the trait can only be evaluated after flowering (e.g., seed oil content)

          Outline in detail the steps that are followed when introgressing a recessive trait via backcrossing

          Outline in detail the steps that are followed when introgressing a recessive trait via backcrossing when the BC progeny can’t be selfed and backcrossed at the same time

          Describe the advantages and disadvantages/limitations of backcrossing

          Describe a scenario where a breeder could/should use backcrossing

          Describe why backcrossing isn’t used to improve quantitative traits

          Demonstrate how a breeder can use Table 28-1 in Fehr’s book to determine the total # of progeny/lines needed to recover a certain # of progeny with the desired genotype (i.e., 1 or multiple genes)

          Describe and then compare and contrast the different approaches (e.g., step-wise, simultaneous, convergent) for introgressing multiple traits using backcrossing

        + - Mutation Breeding

        • Explain why mutation breeding is utilized by plant breeders

          Review the types of mutations generated and mutagens used

          Describe the concept of LD50 and how it could be determined experimentally

          Compare and contrast the advantages and disadvantages of mutagenizing seed, pollen, and vegetative propagules

          Describe the genotype or genotypes (homo vs. heterozygous, elite performance, demonstrating potential for improvement for the target trait – already possess low/high levels of a compound) that could/should be selected for mutagenesis

          Outline in detail the steps a breeder should follow when conducting a mutation breeding program, describe how the steps would differ if the recovered mutation was dominant vs. recessive

          Compare the advantages/disadvantages of advancing progeny via pedigreed families/progeny rows, via single seed descent, or in bulk

          Demonstrate how a breeder can use Table 28-1 in Fehr’s book to determine the total # of progeny/lines needed to recover a certain # with the desired genotype

          Based on examples provided assess/evaluate how breeders utilized mutation breeding to create or improve a new pureline variety

          Describe the advantages and disadvantages/limitations of mutation breeding

          Describe a scenario where a breeder could/should use mutation breeding

        + - Population Improvement in Cross-Pollinated Crops

        • Describe how population improvement differs when working on a self-pollinated crop vs. a cross-pollinated crop that is sold as a hybrid or synthetic variety

          Outline the steps used by plant breeders to conduct population improvement when the selection unit used is single plants (mass selection), half sib OP progeny rows, half sib tester TC rows, full sib progeny rows and selfed family rows

          Compare these different methods based on possible genetic gain/parental control, isolation needs, labor needs, advantages, disadvantages/limitations, etc.

          Demonstrate how these methods could be changed to fit different scenarios and describe how these changes may affect your predicted gain

          Outline the steps used by plant breeders to conduct population improvement using reciprocal recurrent selection

        + - Concepts related to hybrid development

        • + - Heterotic Groups
          • Define what a heterotic group is and why it’s important
          • Define what a heterotic group is and why it’s important
        • + - Combining ability
          • Define general combining ability and specific combining ability and why they matter when breeding a hybrid crop

            Describe when and how GCA and SCA are tested (include information about tester or testers used, how the TC progeny are trialed, how much the TC progeny rows will segregate)

            Illustrate when and how GCA and SCA are tested when a breeder is using pedigree selection to produce inbred lines

            Illustrate when and how CA is tested when a breeder is using doubled haploids to produce inbred lines  

        • + - Male sterility
          • Describe gametophytic and sporophytic self-incompatibility and their use for controlled pollination and hybrid production

            Describe methods to overcome self-incompatibility in gametophytic and sporophytic systems

            Describe genetic male sterility and how sterility is maintained and used for cultivar development

            Describe cytoplasmic male sterility and how sterility is maintained and used for F1 hybrid production including the use of A, B and R lines.

        • + - Types of Hybrids
          • Describe the parents used and how they are crossed to produce F1 hybrids, modified single cross hybrids, 3-way hybrids and 4-way (double cross) hybrids

            Compare and contrast the hybrids listed above for vigor, homo/heterozygosity levels, uniformity in the field and adaptability

            Formulate a scenario when each of the hybrids listed above is an appropriate choice for a farmer

        + - Recurrent Selection and Synthetic Cultivars

        • Define what a synthetic cultivar is (narrow- and broad-based)

          Describe a scenario in which a synthetic cultivar should be used vs. an F1 hybrid or an OPV, what are the advantages of using a synthetic cultivar in this scenario

          Describe in detail the steps used to select the parents/clones used to produce a synthetic cultivar

          Describe in detail the steps used produce a synthetic cultivar (when is balanced bulk used, when is harvest conducted by hand or by machinery, etc.)

          What are some techniques that can be used to ensure random pollination in a polycross nursery

        + - Molecular Plant Breeding, Marker Assisted Breeding, Marker Based Breeding

        • List ways in which markers can be used by breeders

          List characteristics that make a markers system desirable/useful to a breeder

          What are the advantages of using markers vs. or in addition to conventional selection

          Describe how/when a breeder might use MAS during line development (i.e., pedigree selection)

          Describe how MABC is conducted for a dominant trait and how it is used for a recessive trait, what are the advantages of MABC

          Define genomic selection, define the basic steps, and compare it to MAS

          What are the advantages of MAB

          What are the limitations of MAB

          Describe how Molecular PB can be used to increase genetic gain

        + - Breeding Clonal Crops

        • Summarize the main points/basic steps of a clonal breeding program

          Describe how developing a cultivar of a clonal crop differs from developing a pureline cultivar or inbred lines to create a hybrid cultivar

          Describe how backcrossing is altered when working with a clonal crop

          Describe how mutation breeding is used when working with a clonal crop

          Define tandem selection, independent culling and index selection

        + - Breeding Polyploid Crops

        • Describe the differences between and auto- and an allopolyploids and how these differences affect meiosis/fertility

          Describe how a breeder produces auto- and allopolyploids using artificial doubling agents or naturally occurring unreduced gametes (via bilateral or unilateral polyploidization)  

          Describe the ways that a breeder can identify polyploids and the incidence of unreduced gametes

          Summarize the main points/basic steps of a breeding program used to improve polyploid crops and describe why traditional methods of line development are often unsuccessful in polyploid crops

          Describe what endosperm balance number means (potato) and how a breeder might use this phenomenon in his/her breeding program

          Describe how haploids (2x) of cultivated potato (4x) are produced and why they are useful

          Describe what triploid block means (blueberry) and how a breeder might use this phenomenon in his/her breeding program

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    AGRON 524 - Applied Plant Molecular Genetics and Biotechnology

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      1. Gene: a molecular perspective

      • + - Describe the molecular nature of a gene

        • Describe the building blocks of DNA, the nucleotides
        • Describe phosphate bonds that generates the strand
        • Describe the complementary, antiparallel DNA strands
        • Describe the double helix structure of DNA molecules
        • Describe how the DNA is packaged into chromatin and chromosomes
        • Describe centromeres
        • Describe telomeres
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        Describe DNA and DNA replication

        • Semi-conservative
        • The Polymerase Chain Reaction (PCR)
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        Illustrate the process of transcription of DNA to RNA

        • RNA polymerase II
        • Transcription initiation complex
        • Template or negative DNA strand
        • Sense or positive strand
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        Discuss Gene Structure

        • Promoter
        • Terminator
        • Exon
        • Intron
        • Protein coding region
        • Untranslated region
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        Describe RNA to processing

        • Intron splicing
        • 5’ Capping
        • Poly adenylation
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        Describe translation of RNA to protein

        • Codons and the Genetic Code
        • Ribosomes, tRNA and Anti-codons
        • Peptide chain formation
        • Amino acid properties
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        Discuss the concept of protein structure and its relation to protein function

        • Primary structure
        • Secondary structure
        • Homo- and hetero-dimers
        • Non-covalent and covalent bonding among subunits
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      2. Gene expression and regulation

      • Describe the concept of gene expression

        • Expression in tissues, organs
        • In response to environmental stresses

        Discuss transcriptional regulation of gene expression

        • Differential (regulated) gene expression
        • Promoters
        • Enhancers
        • Transcription factors

        Describe epigenetic gene regulation

        • Chromatin structure and histone modification
        • DNA methylation  

        Discuss post-transcriptional regulation of gene expression (RNA level)

        • Alternative splicing  
        • RNA stability and degradation
        • Small RNAs and RNA interference
        • Translational regulation

        Discuss post-translational protein modification and regulation

        • Common types of post-translational modifications  
        • Common types of protein regulation
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      3. Mutations and Trait Variation

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        Differentiate among the types of mutation

        • DNA base substitutions
        • Insertions & deletions  
        • Epimutations
        + -

        Describe epimutation

        • DNA methylation
        • Histone modification
        • RNA interference (RNAi)
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        Discuss the effects of mutations in gene function (coding regions mutation)

        • Silent
        • Missense mutation (amino acid substitution)
        • Frameshift
        • Amino acid additions or deletions
        • Nonsense
        + -

        Discuss the effects of mutations in gene regulatory regions

        • Mutation in regulatory regions; promoters and enhancers
        • Mutation in introns and untranslated regions
        + -

        List and describe types of functional mutations

        • + - Dominant
          • Overexpression mutants
          • Neomorphic mutations
          • Dominant negative mutations
        • Recessive
        • Codominant
        + -

        Discuss how gene mutation can affect traits

        • Loss of function mutation to gain a new trait
        • Gain of function mutation
        • Semidominant traits

        Describe how mutations arise, naturally and experimentally

        • + - Spontaneous mutations
          • Mistakes in DNA replication
          • Environmental mutagens
        • + - Experimental mutagenesis
          • Chemical mutagenesis
          • Insertional mutagenesis
          • Activation tagging
      + -

      4. Genetic pathways

      • Describe how multiple genes often function together to control a trait

        • genetic pathway – many genes working together for a single trait

        Discuss the concept of a biosynthetic pathway 

        • Auxotrophic mutants
        • The anthocyanin pathway
        • Effects of mutations at different steps in anthocyanin biosynthesis
        • Anthocyanin mutants
        • Novel anthocyanin mutant

        Discuss the concept of a regulatory pathway 

        • Positive vs. negative regulators
        • Active vs. inactive states of regulators
        • Steps in a regulatory pathway
        • Signal transduction pathway for hormone signaling
        • Effects of mutation on different steps
        • Loss-of-function
        • Gain-of-function

        Discuss the concept of a network 

        • Biosynthetic pathways
        • Regulatory pathway

        Describe how mutations in different genes of a pathway can cause the same phenotype 

        • Complementary gene action in biosynthetic pathways
        • Complementary gene action in a regulatory pathway

        Describe how mutations in different genes of a pathway might cause different phenotypes

        • Epistasis in a biosynthetic pathway
        • Epistasis in a regulatory pathway
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      5. Recombinant DNA Technology

      • Describe the importance of recombinant DNA technology 

        • Learn the technology
        • History
        • Outcomes

        Describe the process of isolation of DNA and its separation on an agarose gel

        • Preparation of plant DNA
        • Determination of DNA concentration
        • Restriction endonucleases  
        • Digestion of plant DNA with restriction endonucleases
        • Separation of digested genomic DNA on agarose gel  
        • Southern blotting and hybridization

        Discuss and distinguish restriction and ligase enzymes, and their application in gene cloning

        • Cloning vector definition and requirements
        • . Ligase enzyme and gene cloning

        Describe vectors and discuss their application in gene cloning and expression

        • Types of cloning vectors
        • Protein expression in Escherichia coli  
        • Ti plasmid vector and plant transformation  
        • Gene libraries in plasmid, BAC and YAC vectors
        • Gene cloning by chromosome walking, chromosome jumping

        Describe and illustrate polymerase chain termination reaction (PCR)

        • Definition
        • History of PCR
        • DNA and RT-PCR
        • PCR in molecular mapping
        • PCR in cloning genes
        • Quantitative PCR or real-time PCR of DNA or transcripts
        • Limitations
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      6. DNA Sequencing Technology

      • Explain/Describe the importance of DNA sequence 

        Explain the principles of sequencing

        Explain/Describe to distinguish next generation (nextgen) sequencing

        Discuss application of nextgen sequencing  

      + -

      7. Biological Database Web Tools

      • List and describe some of the most commonly used databases in molecular plant breeding

        • Primary database
        • Secondary database
        • Composite database

        Use the tools for accessing and manipulating biological databases

        • NCBI
          • Information retrieval from NCBI
          • Searching NCBI by keywords through Entrez
          • NCBI Basic Local Alignment Search Tool (BLAST)

        Develop proficiency in the use of biological databases

        • Plant species sequence databases; MaizeGDB database
          • Finding map position for a specific locus
          • Using the genome browser

        Develop proficiency in use of multiple sequence alignment tools

        • Using NCBI
          • Information retrieval from NCBI
          • Searching NCBI by keywords through Entrez
          • NCBI Basic Local Alignment Search Tool (BLAST)
        • Finding polymorphisms using ClustalW
        • Developing marker assays
      + -

      8. Molecular Markers

      • Describe the general properties of DNA markers

        • Learn the properties of ideal DNA markers

        List and describe classical DNA markers  

        • Understand RFLP, SSR and AFLP as examples of classical markers

        Describe current DNA markers compared to classical markers 

        • Understand SNP and INDEL markers basics

        Discuss basic marker applications

        • Genetic Fingerprinting
        • Gene tagging
        • Use of linked markers and fingerprinting to assist backcrossing

        List and discuss prerequisites, prospects and limitations in using sequencing for genotyping

        • Sequence barcoding
        • Development of haplotype maps
        • Data Imputation
        • Tag SNPs

        Compare and contrast strengths and weaknesses of non-DNA versus DNA markers

        Explain underlying technologies of non-DNA marker systems

        • Morphological markers (e.g. flower color or seed shape), plant phenomics

          Markers based on gene products, referred to as biochemical markers

          • Markers
          • Protein-based markers
          • Metabolite-based biomarkers
      + -

      9. Comparative Mapping and Genomics 

      • Discuss/explain the difference between genetic and physical maps 

        • Genetic maps
        • Physical maps
        • Genome maps

        Develop familiarity with comparative genomics tools 

        • Colinearity and Synteny
        • Comparative mapping

        Discuss application of comparative mapping 

        • Gene prediction
        • Detecting DNA copy number variations
        • Detecting DNA copy number variations
        • Analysis of genome evolution

        Illustrate the challenges in comparative genomics 

        • Large genome size
        • Multigene family with highly similar family members
      + -

      10. Plant Transformation

      • + -

        Describe plant transformation approaches 

        • Direct DNA uptake
        • Microinjection
        • Agrobacterium-mediated gene transfer
        • Particle bombardment or biolistic transformation
        • Electroporation
        + -

        Describe the vectors required for plant transformation

        • Ti plasmid mediated gene (T-DNA) transfer
        • Binary T-DNA plasmid vector
        + -

        Discuss selectable markers and their usages 

        • Selection of transformed cells
        + -

        Describe methods of analyses and characterization of transgenic crop plants

        • Detection of DNA
        • Detection of RNA
        • Detection of protein
        • Phenotypes: visible as well biochemical

        + -

        Describe mechanism of RNAi 


        • Discovery of RNAi
        • Mechanisms of post-transcriptional gene silencing
        + -

        Develop familiarity with examples of gene silencing approaches for crop improvement 

        • Enhancing carotenoid and flavonoid content in tomatoes
        • Caffeine-free coffee
        • Increased resistance to insects
        • Increased resistance to nematodes
        + -

        Develop familiarity with examples of gene over expression approaches for crop improvement

        • Improving cotton tolerance to drought and salt stress
        • Improving rice tolerance to salinity stress
        + -

        Describe cross protection approaches

        • Transgenic virus resistant papaya
        + -

        Describe transient expression with examples

        • Virus-induced gene silencing
        • Understand the difference between stable transformation and transient expression
    + -

    AGRON 523 - Molecular Plant Breeding

    • + -

      1. Molecular Plant Breeding Concepts

      • + -

        Fundamental concepts - overview

        • List and describe the essential steps in plant breeding process, and  properties of varieties.

          Discuss at high level the toolbox available by molecular genetics and biotechnology

          Integrate plant breeding process with molecular and biotechnological tools and approaches

          Describe how use of molecular and biotechnological tools and approaches transcend traditional plant breeding

      + -

      2. Molecular and Genomic Tools

      • + -

        Bioinformatics

        • Develop markers using genome databases
        + -

        Markers

        • Discuss the use of HRM and KASP as flexible marker systems

          Describe the application of DART_seq for fingerprinting

          List and describe criteria in the classification of marker systems

        + -

        Data management and quality control

        • Demonstrate that marker data can be erroneous

          List and describe different error sources

          Identify quality control measures available

          Discuss higher level of complexity for NGS applications, error sources, big data limitations

          Describe/Explain false positives and negatives: GM testing

      + -

      3. Genetic and Gennomic Approaches

      • + -

        Modeling and data simulation

        • Discuss the fundamental principles of modelling and data simulation

          Show how to Simulate breeding populations

        + -

        Markers as genetic tools

        • Determine allele and genotype frequency changes in allo- and autogamous populations
        • Use markers to calculate heterozygosity and population differentiation
        + -

        Cluster analysis

        • Distinguish different (dis-)similarity measures

          Calculate genetic (dis-) similarities

          Describe/discuss principles of neighbor joining / cluster analysis (such as UPGMA), and PCoA

          Conduct cluster analysis

        + -

        Genetic and physical maps

        • Determine linkage between two, three, multiple markers for genetic mapping

          Describe alternative genetic mapping strategies

          Describe physical mapping and chromosome walking and illustrate their difference with genetic mapping

        + -

        QTL mapping

        • Describe the main features and importance of different QTL mapping methods (single marker, SIM, CIM)

          Identify major determinants for power of QTL detection

          Distinguish alternative QTL mapping methods: AB-QTL, eQTL, IL, Selective Genotyping

          Perform QTL mapping

        + -

        Association analysis

        • Discuss linkage disequilibrium
        • Distinguish conceptual difference of QTL and association mapping
        • Identify and describe different types of association studies
        • List and describe prerequisites for successful  association studies
        • Perform genome-wide association study
        + -

        Gene isolation

        • Describe how to isolate genes

          Illustrate position effect and variation among transgenic events

          Discuss alternative methods for stacking genes

          Explain/discuss coexistence concept

      + -

      4. Genomic Approaches for Selection

      • + -

        Marker-assisted backcrossing (MABC)

        • Describe the method of backcross breeding

          Discuss the main applications of molecular markers for BC breeding (fore-, background selection)

          List and explain factors influencing the efficiency of BC breeding for one and more genes

          Design MABC program

        + -

        Marker-assisted selection (MAS)

        • Discuss the difference(s) between MABC and MAS

          Show the relative efficiency of MAS versus phenotypic selection

          Outline factors influencing the efficiency of MAS

          List/discuss the difference between F2 improvement and MARS

          Describe alternative approaches to MAS

        + -

        Genomic selection (GS)

        • Describe the principles of GS
        • Discuss the differences between MAS and GS
        • Perform GS
        + -

        Genome construction (GC)

        • Discuss the concept of operations research in plant breeding
      + -

      5. Application of Molecular and Genomic Approaches in plant breeding

      • + -

        Marker-based management of genetic resources

        • Describe different processes involved in conservation and exploitation of plant genetic resources

          Discuss application of genomic tools, in particular DNA markers, for taxonomic classification, acquisition of genetic resources, their maintenance, characterization and utilization

        + -

        Biotechnological tools for broadening genetic variation

        • List and discuss alternative methods for inducing/broadening genetic variation using mutagenesis, TILLING, transformation, genome editing

          Describe different genome editing approaches and their application for creating genetic variation or gene replacement

        + -

        Parent selection

        • Describe the usefulness concept

          Identify and discuss alternative methods used to predict mean and genetic variance of breeding populations

        + -

        Modern tools for line development

        • Discuss alternative technologies to develop doubled haploids (DHs)
        • Describe advantages of DHs in different steps of the breeding process compared to selfed families
        + -

        Modern tools for predicting hybrid performance

        • Discuss marker applications for heterotic pool formation and assignment

          Discuss use of genomic tools to understand nature of heterosis

          Describe application of genomic tools for predicting hybrid performance

        + -

        Genomic tools for variety registration, maintenance, protection, and EDVs

        • Describe use of markers for maintenance breeding
        • Discuss use of genomic approaches for monitoring presence of transgenes
        • Discuss use of markers for patent protection
        • Discuss use of markers for DUS testing
        • Describe use of markers to identify essentially derived varieties (EDVs)
    + -

    AGRON 528 - Quantitative Genetics for Plant Breeding

    • + -

      1. Apply Foundational Concepts in Analysis 

      • + - Plant Breeding Programs for pure-line and hybrid crops

        • + - Be able to explain/compare impact of reproductive biology on
          • Genetic improvement
          • Line development
          • Cultivar placement
        • + - Be able to create/formulate/define/propose
          • Objectives
          • Cultivar Design
          • Target environments
        • + - Be able to demonstrate/explain/analyze
          • Feasibility
        • + - Be able to distinguish/evaluate/design
          • Genetic improvement
          • Line development
          • Cultivar placement

        + - Transmission Genetics (students should enter course with abilities to apply and analyze Punnet Squares)

        • + - Be able to design/develop/interpret segregation of alternate alleles at single loci through multiple generations of

          • Self-pollination
          • Inter mating
          • Backcrossing
          • Test crossing

          + - Be able to design/develop/interpret segregation of alternate alleles at multiple linked or unlinked loci through multiple generations of

          • Self-pollination
          • Inter mating
          • Backcrossing
          • Test crossing

        + - Analytic methods (students should enter course with abilities to apply and analyze using software)

        • + - Linear Algebra
          • Addition
          • Subtraction
          • Multiplication
          • Transposition
          • Inversion
        • + - Exploratory Data Analyses
          • Box plots
          • Histograms
          • Biplots
          • Filters
          • Pivot Tables
        • + - Statistics
          • Probability (binomial, multinomial and normal) distributions CLT
          • Estimator, predictor, Estimate, Validate
          • Linear models  [ EMS-ANOVA ]
            • Regression
            • Fixed, random and mixed effects
        • + - Experimental and Sampling Designs
          • Experimental units
          • CRD, RCBD, Split Plot, Incomplete block designs
          • Sampling units
          • Random, stratified random
        • + - Decision theory
          • Inference space
          • Hypothesis testing (types of errors)
          • Precision, accuracy, reliability, sensitivity, specificity
      + -

      2. Multi-environment trials and GxE  

      • Be able to design/simulate G, GxE, e/interpret MET for assumed analysis models
      • + - Be able to analyze/partition/interpret data from MET
        • Heterogeneous variance and covariance
        • Cluster analyses
        • BLUE, BLUP
      • + - Be able to design/analyze/interpret data using Mixed linear models
        • BLUE, BLUP
      + -

      3. Population Genetics 

      • + - Be able to simulate HWE and mechanisms that cause deviations
        • Reference Population
        • Estimators of allelic and genotypic variation
        • Hardy-Weinberg Equilibrium
          • Estimates and hypothesis testing
        • + - Mechanisms that cause deviations from HWE (disequilibrium)
          • Assortative mating, migration, mutation, selection, drift, inbreeding
        • Multi-Locus Disequilibrium
        • Coefficients of relationship and inbreeding
      + -

      4. Quantitative Genetic Models (Be able to simulate additive, dominance and epistatic genetic effects)

      • Phenotypic and genotypic values

        + - Average genetic effects

        • Effect of an allele, effect of an allele substitution, breeding value

        + - Deviations of average genetic effects

        • Dominance
        • Epistasis
        + -

        Variance Components

        • Genetic components
        • Phenotypic components
        • Intraclass correlation
          • Heritability

          • Covariance among relatives
      + -

      5. Selection and Genetic Gain 

      • + - Be able to use breeder’s equation to compare proposed breeding strategies

        • Selection intensity
        • Correlation (selection units, evaluation units)
        • Additive genetic variance
        • Cost
        • Time and reproductive biology

        Be able to simulate selection and compare realized genetic gains from proposed breeding strategies

        Be able to understand the impact of multiple trait selection in the context of multiple stages of evaluations on genetic gain

      + -

      6. Resource Allocation

      • CTP framework
      • Translate verbal objectives into functional objectives
      • Understand decision variables and project constraints
      • + - Optimize single objective with multiple decision variables and constraints
        • Determine/understand feasibility of meeting an objective
        • Linear programming
        • Integer programming
      • + - Optimization of Multiple objectives
        • Determine/understand feasibility of meeting objectives
        • Understand/quantify/interpret trade-offs among objectives (Pareto optimality)
    + -

    AGRON 521 - Principles of Cultivar Development

    • + -

      1.  Basic concepts in cultivar development. (What are the basic concepts and how they impact cultivar development?)

      • Define mating systems

        Recognize similarities and dissimilarities between populations and cultivars

        Identify and infer types of cultivars

        Review and explain crossing symbols, pedigree writing and selection history

        Define breeding process and breeding objectives in major crops

        Describe the principles of setting breeding objectives

        Apply knowledge of breeding objectives in cultivar development

      + -

      2. Germplasm usage in cultivar development. (How is germplasm usage important in cultivar development?)

      • Summarize natural and artificial selection
      • Illustrate plant breeding pipeline
      • Describe sources of parental material and principles in assembling genetic variation
      • Compare various mechanisms of variety protection
      • Revise material transfer agreements and germplasm sharing, and associated legal issues
      + -

      3. Review of basic population and quantitative genetics principles. (What are basic population and quantitative genetics principles and how do they impact cultivar development?)

      • Understand the different types of populations in plant breeding

        Apply different types of crossing schemes for cultivar development

        Learn to distinguish qualitative versus quantitative traits and apply in cultivar development pipeline

        Understand the concepts of variability, phenotype, genotype, and genotype x environment (G x E) interactions

        Identification of high yielding stable varieties

        Understand the concept of heritability and practice selection

        Learn selection theory and deploy for increased response to selection (breeder’s equation)

        Recognize multiple trait selection scenarios and deploy for different scenarios

        Understand mating designs, combining ability and its calculations

        Explain heterosis

        Choose program size, generation to select and population sizes

      + -

      4. Steps in cultivar development and breeding methods. (What are the critical steps in cultivar development and breeding methods?)

      • Operate field testing program with choice of statistical design and analysis for breeding trials

        Deploy principles of cultivar development to create new varieties

        Describe main steps in development of pureline, clonal, synthetic, hybrid, multi-line and blend cultivars

        Understand and deploy strategies for population improvement

        Select appropriate tools (phenomic, genomic, infrastructural) for achieving program objectives

        Describe main breeding and selection methods in cultivar development (generation advancement, selections, precommercial testing, seed production and certification, distribution)

        Evaluate cultivar development pipelines for different types of cultivars in major and minor crops

    + -

    AGRON 544 - Host-Pest Interactions

    • 1. Integrated Pest Management (IPM) Concepts 

      • + - Scope and concepts of IPM
        • Identify the essential components of a sound IPM program

          Discuss the history of pest management and how our current concepts of IPM evolved

          Demonstrate the scope of IPM and all the professional disciplines it interacts with

          Describe the basic concepts that constitutes IPM

          Analyze the economic viability of pest management tactics

          Recognize the basis of IPM theory and how it affects management strategies

          Describe and compare the basic components of the life cycles of weed, pathogen, and insect pests and why they are important to IPM strategies

          Identify and describe the major preventative and curative approaches to managing pest populations

          Discuss the concept of "pests"

      + -

      2. When insects become pests? 

      • + - Insect morphology and development
        • Explain where the class Insecta fits in evolutionary history and the extreme diversity that has evolved over geologic time

          Describe how insects make their living and why they are so successful

          Describe the structure and function of insect anatomy and relate this to their great success

          Illustrate the steps of the insect life cycle and the variation in life cycles among major groups of insects

          Describe the physiological processes involved in insect development and the environmental stimuli that regulate development

      • + - Ecological management and biological control
        • Discuss the structure of ecosystems, and how this relates to insect management

          Describe the roles that insects play within ecosystems

          Identify and discuss the factors that determine insect abundance and that knowledge of these factors allows us to predict population change through time

          List and describe the different factors that may regulate insect populations

          Describe how agroecosystems differ from natural ecosystems, and discuss the implications of these differences

          Describe how the field environment can be modified by cultural practices so that pest problems can be minimized

          Discuss how time and space interact with pest populations and describe how to reduce pest success by adopting management strategies that upset the continuity of the pest's requisite resources

          Describe how pests may be diverted away from the commodity to other suitable environments

          Describe how biocontrol methods can be incorporated into management strategies to lessen pest pressures

      • + - Insect/host interactions and insect scouting
        • Define the terms (injury, damage, damage boundary, gain threshold, etc.) upon which the EIL concept is based

          List and define the parameters in the EIL equation, and discuss how changes in the separate parameters influence the EIL

          Describe how EILs are developed and calculated

          Recognize that management decisions are based on economic thresholds and that there are limitations to the EIL concept

          Distinguish between direct and indirect sampling techniques

          List and describe the wide variety of insect sampling techniques and the sampling situations they are best suited to

          Differentiate between absolute and relative estimates, and describe how to collect statistically sound data on insect populations

          Describe criteria for designing an insect scouting and sampling program

      + -

      3. Pest management strategies

      • + - Host plant resistance
        • Describe the genetic nature of resistance in plants
        • Interpret the resistance interactions of the insect-plant relationship
        • Explain the application and integration of host plant resistance using the IPM perspective
        • Describe the selection and manipulation of resistance factors in agronomic crops
        • Recognize ecological backlash and how it develops in pest populations
        • Identify possible negative interactions of pest management tactics for multiple pests
      • + - Insecticides in pest management
        • Recognize and discuss the differences among the variety of insecticides and formulations of those active ingredients

          Distinguish between properties of classes of insecticides and describe how they function in insects and can poison humans

          Design an efficient, effective, and safe pest management strategy using insecticides

          Choose insecticides and formulations of those insecticides that are best suited to an insect pest situation

      + -

      4. Concepts of plant diseases

      • + - Concepts of abiotic disease in plants
        • Describe the concepts of disease, injury, and the signs and symptoms of plant pathogenic agents
        • Describe the basic biology of abiotic disease agents of plants
        • Describe the basic biology of the major biotic disease agents in plants
        • Discuss the interaction of pathogens, their plant hosts, and environmental factors on plant diseases
      • + - Concepts of biotic disease in plants
        • Describe the concepts of disease, injury, and the signs and symptoms of plant pathogenic agents
        • Describe the basic biology of abiotic disease agents of plants
        • Describe the basic biology of the major biotic disease agents in plants
        • Discuss the interaction of pathogens, their plant hosts, and environmental factors on plant diseases
      + -

      5. Concepts of disease management

      • + - Chemical and biological management of plant diseases
        • Describe the basic properties of fungicides, antibiotics, and nematicides
        • Discuss how these chemicals and biological agents are used in plant disease management
        • Identify the major groups of disease management chemicals and major examples of each group
      • + - Etiology, pathogenesis and epidemiology
        • Describe how pathogens incite disease in plants

          Discuss how this process relates to pathogen life cycles

          Describe how the disease is spread to other hosts

          Describe how plants protect themselves from initial infection of the spread of a disease from an infection site

          Identify and discuss about factors that can affect disease onset and spread

          Describe the basic principles of disease assessment

          Determine the effect of spatial relationships on disease spread

          Assess crop losses that are the result of disease

      • + - Recognition of major diseases in agronomic crops
        • Recognize the major diseases of corn, soybean, alfalfa, oats, and wheat in the midwestern U.S.
      • + - Disease resistance in plants
        • Describe the basic principles and concepts of disease resistance in plants

          Discuss the physical, biochemical, and genetic mechanisms that contribute to disease resistance and how these affect spatial distribution

          Describe the variability within pathogens and how these different genotypes affect management

          Discuss how resistant crop genotypes are developed in resistance breeding programs

          Identify cultural management practices that will help in the management of specific agronomic diseases

          Demonstrate management plans to reduce disease potential in specific crops

      + -

      6. Concepts of weed science and weed management

      • + - Concepts of plants as weeds – introduction to herbicide chemistry
        • Describe what weeds are and how they interact with humans and our cropping systems
        • Discuss the niches that weeds occupy and the strategies they use to survive and thrive
        • Describe the population dynamics of weed communities
        • Identify the vegetative characteristics which facilitate identification of juvenile weed species
        • Recognize the major weed species in your cropping region
      • + - Herbicide/plant relationships affecting herbicide uptake, translocation, and efficacy
        • Describe the main ways that herbicides enter plant organs or tissues

          Discuss the interaction between plant morphology and herbicides, and how this interaction affects the efficacy of herbicide action

          Describe how herbicides move through plant tissues and what factors affect translocation within a plant

          Describe how soil properties affect the movement of herbicides in solution and the efficacy of herbicide uptake in roots

          Discuss the anatomy and morphology of roots and how root architecture affects uptake of soil applied herbicides

          Describe the physical properties of chemical agents which regulate how and when they can be safely used in an environment

          Distinguish between which herbicides are competitive and noncompetitive inhibitors

          Describe the physical properties of chemical agents which regulate how and when they can be safely used in an environment

      • + - Herbicide mode of action
        • Discuss how herbicides interact with plant's biochemistry
        • Describe what effect herbicide metabolism has on the selectivity of a given herbicide group
        • Discriminate herbicide symptoms for various herbicide families  
      • + - Herbicide resistance and summary
        • List and describe what factors contribute to herbicide resistance and how this resistance develops in plant populations

          Define the physiological adaptations in plants which confer resistance or tolerance to herbicides