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Elements of Genetics GENETICS AND PLANT BREEDING Elements of Genetics Dr. B. M. Prasanna National Fellow Division of Genetics Indian Agricultural Research Institute New Delhi-110012 (12-06- 2007) CONTENTS Introduction History Cell Cell Division Special Chromosomes Dominance Relationships Gene Interactions Multiple Alleles Sex Determination Sex Linkage Linkage and Crossing Over Genetic Mapping Structural Changes in Chromosomes Numerical Changes in Chromosomes Nature of the Genetic Material Gene Regulation Operon Concept Gene Concept Mutation Polygenic and Quantitative Inheritance Extrachromosomal Inheritance Plant Tissue Culture Keywords Mitosis, Meiosis 1 Introduction In biology, heredity is the passing on of characteristics from one generation to the next. It is the reason why offspring look like their parents. It also explains why cats always give birth to kittens and never puppies. The process of heredity occurs among all living organisms, including animals, plants, bacteria, protists and fungi. Genetic variation refers to the variation in a population or species. Genetics is the study of heredity and variation in living organisms. Two research approaches were historically important in helping investigators understand the biological basis of heredity. The first of these approaches, ‘transmission genetics’, involved crossing organisms and studying the offsprings' traits to develop hypotheses about the mechanisms of inheritance. The second approach involved using cytological techniques to study the machinery and processes of cellular reproduction. This approach laid a solid foundation for the more conceptual understanding of inheritance that developed as a result of transmission genetics. Ever since 1970s, with the advent of molecular tools and techniques, geneticists are able to intensively analyze genetic basis of trait variation in various organisms, including plants, animals and humans. History Pre-Mendelian and Post-Mendelian Concepts of Heredity It was apparent to ancient humans that offspring resembled their parents. The Greek philosophers had a variety of ideas about heredity: Theophrastus proposed that male flowers caused female flowers to ripen; Hippocrates speculated that "seeds" were produced by various body parts and transmitted to offspring at the time of conception, and Aristotle thought that male and female semen mixed at conception. Aeschylus, in 458 BC, proposed the male as the parent, with the female as a "nurse for the young life sown within her". Various hereditary mechanisms were envisaged without being properly tested or quantified. These included “blending inheritance” and the “inheritance of acquired traits”. Nevertheless, people were able to develop domestic breeds of animals as well as crops through artificial selection. During the 1700s, Dutch microscopist Anton van Leeuwenhoek (1632-1723) discovered "animalcules" in the sperm of humans and other animals. Some scientists speculated they saw a "little man" (homunculus) inside each sperm. These scientists formed a school of thought known as the "spermists". They contended the only contributions of the female to the next generation were the womb in which the homunculus grew, and prenatal influences of the womb. An opposing school of thought, the “ovists”, believed that the future human was in the egg, and that sperm merely stimulated the growth of the egg. Ovists thought women carried eggs containing boy and girl children, and that the gender of the offspring was determined well before conception. “Pangenesis” was an idea that males and females formed "pangenes" in every organ. These pangenes subsequently moved through their blood to the genitals and then to the children. The concept originated with the ancient Greeks and influenced biology until little over 100 years ago. The terms "blood relative", "full-blooded", and "royal blood" are relics of pangenesis. Francis Galton, Charles Darwin's cousin, experimentally tested and disproved pangenesis during the 1870s. 2 Charles Darwin proposed a theory of evolution in 1859 and one of its major problems was the lack of an underlying mechanism for heredity. Darwin believed in a mix of blending inheritance and the inheritance of acquired traits (pangenesis). Blending inheritance would lead to uniformity across populations in only a few generations and thus would remove variation from a population on which natural selection could act. Darwin's initial model of heredity was adopted by, and then heavily modified by Francis Galton, who laid the framework for the biometric school of heredity. Galton rejected the aspects of Darwin's pangenesis model which relied on acquired traits. The inheritance of acquired traits was shown to have little basis in the 1880s when August Weismann cut the tails off many generations of mice to find that their offspring did continue to develop tails. The idea of particulate inheritance of genes can be attributed to Gregor Mendel who presented his work on pea plants in 1865. The year 1900 gave birth to a new discipline that soon came to be called ‘genetics’. During that year, three botanists, Hugo de Vries, Carl Correns, and Erich Tschermak, reported on their breeding experiments of the late 1890s and claimed to have confirmed the regularities in the transmission of characters from parents to offspring that Mendel had already presented in his seminal paper of 1865. The additional observation that sometimes several elements behaved as if they were linked, contributed to the assumption soon promoted by Walter Sutton and Theodor Boveri that these elements were located in groups on the different chromosomes of the nucleus. Thus, the ‘chromosome theory of inheritance’ was proposed. Toward the end of the first decade of the 20th century, after Bateson had coined the term genetics for the emerging new field of transmission studies in 1906, Wilhelm Johannsen codified this distinction by introducing the notions of genotype and phenotype, respectively. In addition, for the elements of the genotype, he proposed the notion of gene. This terminology was gradually taken up by the genetics community. By the early 1900s, cytologists had demonstrated that heredity is the consequence of the genetic continuity of cells by cell division, had identified the gametes as the vehicles that transmit genetic information from one generation to another, and had collected strong evidence for the central role of the nucleus and the chromosomes in heredity. It was initially assumed the Mendelian inheritance only accounted for large (qualitative) differences, such as those seen by Mendel in his pea plants — and the idea of additive effect of (quantitative) genes was not realized until R.A. Fisher's (1918) classic paper on The Correlation Between Relatives on the Supposition of Mendelian Inheritance. In the 1930s, work by Fisher and others resulted in a combination of Mendelian and biometric schools into the modern synthesis of evolution. From the early 1910s right into the 1930s, the growing community of researchers around Thomas Hunt Morgan used mutants of the fruit fly Drosophila, constructed in ever more sophisticated ways, in order to produce a map of the fruit fly’s genotype in which genes, and alleles thereof, figured as genetic markers occupying a particular locus on one of the four homologous chromosome pairs of the fly. Meanwhile, cytological work added credence to the materiality of genes-on-chromosomes. 3 In 1941, George Beadle and Edward Tatum showed that mutations in genes caused errors in certain steps in metabolic pathways. This showed that specific genes code for specific proteins, leading to the "one gene-one enzyme" hypothesis. Oswald Avery, Collin Macleod, and Maclyn McCarty showed in 1944 that DNA holds the gene's information. In 1953, James D. Watson and Francis Crick demonstrated the molecular structure of DNA, the genetic material in all organisms, except some viruses. Mendelian Principles of Heredity The principles of inheritance were derived by a 19th century monk, Gregor Johann Mendel (born in Moravia, which was then a part of Austria, and now in Czeck Republic), who conducted plant hybridization experiments in his monastery. Between 1856 and 1863, he cultivated and tested some 28,000 garden pea (Pisum sativum) plants. His experiments brought forth two generalizations which later were known as Mendel’s principles of heredity or Mendelian inheritance. These were described in his essay "Experiments on Plant Hybridization" that was read to the Natural History Society of Brunn on February 8 and March 8, 1865, and was published in 1866. Gregor Mendel is now regarded as the ‘Father of Genetics’, as he introduced a new formal tool for the analysis of heredity and variation in organisms, initiated the hybridization experiments that were based on a new experimental regime: the selection of discrete character pairs. His findings allowed other scientists to simplify the emergence of traits to mathematical probability. A large portion of Mendel's findings can be traced to his choice to start his experiments only with true breeding plants. He measured only absolute characteristics such as colour, shape, and Gregor Mendel and the first page of his classical position of the offspring. His data was paper expressed numerically and subjected to statistical analysis. This method of data reporting and the large sampling size he used gave credibility to his data. He also had the foresight to look through several successive generations of his pea plants and record their variations. Without his careful attention to procedure
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