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Introduction
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that chromosomes are linear arrays of genes. In addition, the foundations of modern evolutionary and molecular genetics were derived. In 1900, three biologists working independently Hugo de Vries, Carl Correns, and Erich von Tschermak rediscovered Mendel s landmark work on the rules of inheritance, published in 1866, thus beginning our era of modern genetics. In 1903, Walter Sutton hypothesized that the behavior of chromosomes during meiosis explained Mendel s rules of inheritance, thus leading to the discovery that genes are located on chromosomes. In 1913, Alfred Sturtevant created the rst genetic map, using the fruit y. He showed that genes existed in a linear order on chromosomes. In 1927, L. Stadler and H. J. Muller showed that genes can be mutated arti cially by X rays. Between 1930 and 1932, R. A. Fisher, S. Wright, and J. B. S. Haldane developed the algebraic foundations for our understanding of the process of evolution. In 1943, S. Luria and M. Delbr ck demonstrated that bacteria have normal genetic systems and thus could serve as models for studying genetic processes.
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1944-Present
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The period from 1944 to the present is the era of molecular genetics, beginning with the demonstration that DNA is the genetic material and culminating with our current explosion of knowledge due to recombinant DNA technology. In 1944, O. Avery and colleagues showed conclusively that deoxyribonucleic acid DNA was the genetic material. James Watson and Francis Crick worked out the structure of DNA in 1953. Between 1968 and 1973, W. Arber, H. Smith, and D. Nathans, along with their colleagues, discovered and described restriction endonu-
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cleases, the enzymes that opened up our ability to manipulate DNA through recombinant DNA technology. In 1972, Paul Berg was the rst to create a recombinant DNA molecule. Since 1972, geneticists have cloned numerous genes. Scientists now have the capability to create transgenic organisms, organisms with functioning foreign genes. For example, we now have farm animals that produce pharmaceuticals in their milk that are harvested easily and inexpensively for human use. In 1997, the rst mammal was cloned, a sheep named Dolly. The sequence of the entire human genome was determined in 2000; we will spend the next century mining its information in the newly created eld of genomics, the study of the complete genetic complement of an organism. Although no inherited disease has yet been cured by genetic intervention, we are on the verge of success in numerous diseases, including cancer. The material here is much too brief to convey any of the detail or excitement surrounding the discoveries of modern genetics. Throughout this book, we will expand on the discoveries made since Darwin rst published his book on evolutionary theory in 1859 and since Mendel was rediscovered in 1900.
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THE THREE GENERAL AREAS OF GENETICS
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Historically, geneticists have worked in three different areas, each with its own particular problems, terminology, tools, and organisms. These areas are classical genetics, molecular genetics, and evolutionary genetics. In classical genetics, we are concerned with the chromosomal theory of inheritance; that is, the concept that genes are
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Table 1.1 The Three Major Areas of Genetics_Classical, Molecular, and Evolutionary_
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and the Topics They Cover
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Classical Genetics Mendel s principles Meiosis and mitosis Sex determination Sex linkage Chromosomal mapping Cytogenetics (chromosomal changes) Molecular Genetics Structure of DNA Chemistry of DNA Transcription Translation DNA cloning and genomics Control of gene expression DNA mutation and repair Extrachromosomal inheritance Evolutionary Genetics Quantitative genetics Hardy-Weinberg equilibrium Assumptions of equilibrium Evolution Speciation
Tamarin: Principles of Genetics, Seventh Edition
I. Genetics and the Scientific Method
1. Introduction
The McGraw Hill Companies, 2001
How Do We Know
located in a linear fashion on chromosomes and that the relative positions of genes can be determined by their frequency in offspring. Molecular genetics is the study of the genetic material: its structure, replication, and expression, as well as the information revolution emanating from the discoveries of recombinant DNA techniques (genetic engineering, including the Human Genome Project). Evolutionary genetics is the study of the mechanisms of evolutionary change, or changes in gene frequencies in populations. Darwin s concept of evolution by natural selection nds a rm genetic footing in this area of the study of inheritance (table 1.1). Today these areas are less clearly de ned because of advances made in molecular genetics. Information coming from the study of molecular genetics allows us to understand better the structure and functioning of chromosomes on the one hand and the mechanism of natural selection on the other. In this book we hope to bring together this information from a historical perspective. From Mendel s work in discovering the rules of inheritance (chapter 2) to genetic engineering (chapter 13) to molecular evolution (chapter 21), we hope to present a balanced view of the various topics that make up genetics.
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