George Acquaah - Principles of Plant Genetics and Breeding

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The revised edition of the bestselling textbook, covering both classical and molecular plant breeding Principles of Plant Genetics and Breeding Now in its third edition, this essential textbook contains extensively revised content that reflects recent advances and current practices. Substantial updates have been made to its molecular genetics and breeding sections, including discussions of new breeding techniques such as zinc finger nuclease, oligonucleotide directed mutagenesis, RNA-dependent DNA methylation, reverse breeding, genome editing, and others. A new table enables efficient comparison of an expanded list of molecular markers, including Allozyme, RFLPs, RAPD, SSR, ISSR, DAMD, AFLP, SNPs and ESTs. Also, new and updated “Industry Highlights” sections provide examples of the practical application of plant breeding methods to real-world problems. This new edition:
Organizes topics to reflect the stages of an actual breeding project Incorporates the most recent technologies in the field, such as CRSPR genome edition and grafting on GM stock Includes numerous illustrations and end-of-chapter self-assessment questions, key references, suggested readings, and links to relevant websites Features a companion website containing additional artwork and instructor resources 
offers researchers and professionals an invaluable resource and remains the ideal textbook for advanced undergraduates and graduates in plant science, particularly those studying plant breeding, biotechnology, and genetics.

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A good breeder should have a keen sense of observation. Several outstanding discoveries were made just because the scientists who were responsible for these events were observant enough to spot unique and unexpected events. Luther Burbank selected one of the most successful cultivars of potato, the “Burbank potato,” from among a pool of variability. He observed a seed ball on a vine of the “Early Rose” cultivar in his garden. The ball contained 23 seeds, which he planted directly in the field. At harvest time the following fall, he dug up and kept the tubers from the plants separately. Examining them, he found two vines that were unique, bearing large smooth and white potatoes. Still, one was superior to the others. The superior one was sold to a producer who named it Burbank. The Russet Burbank potato is produced on about 50% of all lands devoted to potato production in the US.

Breeders often have to discriminate among hundreds and even tens of thousands of plants in a segregating population to select only a small fraction of promising plants to advance in the program. Visual selection is an art, but it can be facilitated by selection aids such as genetic markers(simply inherited and readily identified traits that are linked to desirable traits that are often difficult to identify). Morphological markers (not biochemical markers) are useful when visual selection is conducted. A keen eye is advantageous even when markers are involved in the selection process. As will be emphasized later in this book, the breeder ultimately adopts a holistic approach to selection, evaluating the overall worth or desirability of the genotype, not just the character targeted in the breeding program.

1.7.2 The scientific disciplines and technologies of plant breeding

The science and technology component of modern plant breeding is rapidly expanding. Whereas a large number of science disciplines directly impact plant breeding, several are closely associated with it. These are plant breeding, genetics, agronomy, cytogenetics, molecular genetics, botany, plant physiology, biochemistry, plant pathology, entomology, statistics, and tissue culture. Knowledge of the first three disciplines is applied in all breeding programs. The technologies used in modern plant breeding are summarized in Table 1.2. These technologies are discussed in varying degrees in this book. The categorization is only approximate and generalized. Some of these tools are used to either generate variability directly or to transfer genes from one genetic background to another to create variability for breeding. Some technologies facilitate the breeding process through, for example, identifying individuals with the gene(s) of interest.

GeneticsGenetics is the principal scientific basis of modern plant breeding. As previously indicated, plant breeding is about targeted genetic modification of plants. The science of genetics enables plant breeders to predict to varying extents the outcome of genetic manipulation of plants. The techniques and methods employed in breeding are determined based on the genetics of the trait of interest, regarding, for example, the number of genes coding for it and gene action. For example, the size of the segregating population to generate in order to have a chance of observing that unique plant with the desired combination of genes depends on the number of genes involved in the expression of the desired trait.

BotanyPlant breeders need to understand the reproductive biology of their plants as well as their taxonomic attributes. They need to know if their plants to be hybridized are cross‐compatible, as well as know in fine detail about flowering habits, in order to design the most effective crossing program.

Plant physiologyPhysiological processes underlie the various phenotypes we observe in plants. Genetic manipulation alters plant physiological performance, which in turn impacts the plant performance in terms of the desired economic product. Plant breeders manipulate plants for optimal physiological efficiency so that dry matter is effectively partitioned in favor of the economic yield. Plants respond to environmental factors, biotic (e.g. pathogens) and abiotic (e.g. temperature, moisture). These factors are sources of physiological stress when they occur at unfavorable levels. Plant breeders need to understand these stress relationships in order to develop cultivars that can resist them for enhanced productivity.

AgronomyPlant breeders conduct their work in both controlled (greenhouse) and field environments. An understanding of agronomy (the art and science of producing crops and managing soils) will help the breeder to provide the appropriate cultural conditions for optimal plant growth and development for successful hybridization and selection in the field. An improved cultivar is only as good as its cultural environment. Without the proper nurturing, the genetic potential of an improved cultivar would not be realized. Sometimes, breeders need to modify the plant growing environment to identify individuals to advance in a breeding program to achieve an objective (e.g. withholding water in breeding for drought resistance).

Pathology and entomologyDisease resistance breeding is a major plant breeding objective. Plant breeders need to understand the biology of the insect pest or pathogen against which resistance is being sought. The kind of cultivar to breed, the methods to use in breeding and evaluation all depend on the kind of pest or pathogen (e.g. its races or variability, pattern of spread, life cycle, and most suitable environment).

StatisticsPlant breeders need to understand the principles of research design and analysis. This knowledge is essential for effectively designing field and laboratory studies (e.g. for heritability, inheritance of a trait, combining ability), and evaluating genotypes for cultivar release at the end of the breeding program. Familiarity with computers is important for record keeping and data manipulation. Statistics is indispensable to plant breeding programs. This is because the breeder often encounters situations in which predictions about outcomes, comparison of results, estimation of response to a treatment, and many more, need to be made. Genes are not expressed in a vacuum but in an environment with which they interact. Such interactions may cause certain outcomes to deviate from the expected. Statistics is needed to analyze the variance within a population to separate real genetic effects from environmental effects. Application of statistics in plant breeding can be as simple as finding the mean of a set of data, to complex estimates of variance and multivariate analysis.

BiochemistryIn this era of biotechnology, plant breeders need to be familiar with the molecular basis of heredity. They need to be familiar with the procedures of plant genetic manipulation at the molecular level, including the development and use of molecular markers and gene transfer techniques.

Table 1.2An operational classification of technologies of plant breeding.

Classical/traditional tools; e.g. Common use of the technology/tool
Emasculation Making a completer flower female; preparation for crossing
Hybridization Crossing unidentical plants to transfer genes or achieve recombination
Wide crossing Crossing of distantly related plants
Selection The primary tool for discriminating among variability
Chromosome counting Determination of ploidy characteristics
Chromosome doubling Manipulating ploidy for fertility
Male sterility To eliminate need for emasculation in hybridization
Triploidy To achieve seedlessness
Linkage analysis For determining association between genes
Statistical tools For evaluation of germplasm
Relatively advanced tools
Mutagenesis To induce mutations to create new variability
Tissue culture For manipulating plants at the cellular or tissue level
Haploidy Used for creating extremely homozygous diploid
Isozyme markers To facilitate the selection process
In situ hybridization Detect successful interspecific crossing
More sophisticated tools
DNA markers
RFLP More effective than protein markers (isozymes)
RAPD PCR‐based molecular marker
Advanced technology
Molecular markers SSR, SNPs, ISSR, DART, etc.
Marker‐assisted selection Facilitate the selection process
DNA sequencing, NGS Ultimate physical map of an organism
Plant genomic analysis‐ Studying the totality of the genes of an organism
Bioinformatics Computer‐based technology for prediction of biological function from DNA sequence data
Microarray analysis To understand gene expression and for sequence identification
Primer design For molecular analysis of plant genome
Plant transformation For recombinant DNA work
OMICS technologies For studying various aspects of the entire genome
Genome editing For more efficient manipulation of the genome
Genome mapping For more efficient gene discovery

Whereas the training of a modern plant breeder includes these courses and practical experiences in these and other disciplines, it is obvious that one cannot be an expert in all of them. Modern plant breeding is more of a team than a solo effort. A plant breeding team will usually have experts in all these key disciplines, each one contributing to the development and release of a successful cultivar. Increasingly important disciplines in plant breeding are computer science, for their role in bioinformatics, big data, and simulations.

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