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The Centenary Progress of Molecular Genetics. a 100Th Anniversary of T Coll. Antropol. 34 (2010) 3: 1167–1174 Review The Centenary Progress of Molecular Genetics. A 100th Anniversary of T. H. Morgan’s Discoveries Tomislav Keros1, Fran Borove~ki2, Lorena Jemer{i}1, Dean Konjevi}3, Besi Roi}1 and Jelena Balatinec1 1 Croatian Veterinary Institute, Zagreb, Croatia 2 Department of Functional Genomics, Centre for Translation and Clinical Research, Zagreb University Hospital Centre, Zagreb, Croatia 3 Department of Game Biology, Pathology and Breeding, School of Veterinary Medicine, Zagreb University, Zagreb, Croatia ABSTRACT A century ago, Thomas Hunt Morgan, the American scientist, studied the cytogenetic changes of drosophila and came to cytogenetic explanation of Mendel’s basic laws of genetic heredity. These studies resulted in today’s Mendel-Morgan chromosomal theory of heredity. On the occasion of the hundredth anniversary of this important discovery the authors have decided to give a review of the most significant achievments in the field of molecular genetics until the completion of the Human Genome Project. The most important points concerning the technology of DNA recombination and genetic en- gineering are also presented. The final section discusses the significance of previous achievements of molecular genetics in biomedicine and other related fields. There is also a tabular presentation of the sequence of the most important find- ings in the field of molecular genetics through time. Key words: history, T. H. Morgan, molecular genetics, genetic engineering Introduction »PCR is the most important new scientific technology to come along in the last hundred years.« C.R. Hughes, 2001 The significance of heredity in inheritance of differ- lished in 1866, he states that many characteristics are ent bodily characteristics, as well as in the occurrence of transferred to offspring by certain »hereditary factors«. disease has been described from the very beginnings of Furthermore, he stated that young plants inherit one medical history. Biologists and many other scientists »factor« from each parent, who actually have a double have devoted their efforts to the understanding of the »hereditary factor«. Therefore, according to the funda- transfer of hereditary traits, as well as the physical and mental Mendel Law on Heredity, a single entity inherits chemical properties of the related processes. Gregor two »factors« for each trait, called alleles, which can be Johann Mendel (1822–1884), a highschool teacher and identical or different and are manifested in the develop- Benedictine priest, is considered the founder of tradi- ment of the entity1,2. tional genetics. Working in the St. Thomas monastery in Brno he performed numerous experiments on various Mendel’s research, published in the not so popular plants and animals. In his experiments he also used publication »Verh. Naturforsch. Verein Brünn«, passed green beans and, formally being a physicist, he applied unnoticed, and it was only in 1900 that the botanists, statistical analysis for the first time as a scientific ap- Hugo de Vries, in the Netherlands, Carl E, Correns in proach. He noticed that when cross-bred, the offspring of Germany, and Erich von Tschermak in Austria, brought plants inherit certain traits in strict numerical orders. In to wider attention Mendel’s assumptions and provided his work »Versuche über Pflanzen – Hybriden«, pub- further evidence of their significance. Namely, in 1901 Received for publication July 29, 2010 1167 T. Keros et al.: The Centenary Progress of Molecular Genetics, Coll. Antropol. 34 (2010) 3: 1167–1174 Hugo de Vries published his finding that individual or- number of chromosomes doubled and that from these ganisms had the characteristics that their ancestors did pairs the newly created cells received one segregated not have and were the variations caused by spontaneous chromosome. changes of hereditary traits, called mutations. He refer- In 1921, Theophilus Shickel Painter, the American zo- 1,3 red to the newly formed organisms as the mutants . ologist, by studying human sperm cells during meiosis The development of a microscope and laboratory discovered that each contained 24 chromosomes. Later equipment enabled the scientists to learn that all cell nu- one of his students concluded that the oocyst contained clei contained the substance called chromatin,that turns the same number of chromosomes, whereas all other hu- into denser and visible pairs of chromosomes during cell man body cells should contain 48 chromosomes. This ob- mytosis. These stained fibre-like corpuscles observed in servation remained the subject of dispute for nearly dividing living cells were described in 1882 by Walter three decades1,4. Flemming. He was also able to confirm that, during cell During almost twenty years the scientists have paid fertilisation, the offspring received the haploid number close attention to the nature of genes. Besides biologists, of chromosomes from the gametes of both parents4. many physicists and chemists have studied and clarified In 1902 Walter S. Sutton, the American scientist, fi- fundamental biochemical events occurring within cells. nally provided proof that, as assumed by Mendel, the These studies focused on amino acids, lipids, simple car- chromosomes behaved as »factors« during cell division bohydrates, but also larger so-called macromolecules such and emphasized the relation between chromosomes and as nucleic acids and proteins. The studies also included inherited traits. molecular regulation of biological processes in cells, ge- netic information contained in the studied molecules and The English mathematician Geoffrey H. Hardy and the ways in which transformation of the genetic data into German ophthalmologist and geneticist Wilhelm Wein- chemical reactions results in certain phenotypic traits berg described in 1908 the relation between the fre- and ensures biological homeostasis of the organisms3. quency of alleles and the frequency of genotype in a sin- gle site – which is today referred to as Hardy-Weinberg The American geneticists George W. Beadle and Ed- equilibrium5. ward Lawrie Tatum in 1941 revealed that single genes determined the production of enzymes and that the seg- Thomas Hunt Morgan and co-workers at the New ments of deoxyribonucleic acid (DNA) served as the ma- York Columbia University provided evidence regarding trix for the formation of protein molecules in the cell. genes as units of heredity and their strict alignment in regular sequence along the chromosome. Morgan experi- Working at the New York Rockefeller Institute in mented on fruit flies (Drosophila melanogaster), which 1944, Oswald Theodore Avery, Colin MacLeod and Me- were most appropriate for study purposes since they re- chin McCarty studied the transformation of pneumonia quire less than 14 days for new progeny. By observing he- bacillus and proved that genetic messages are contained reditary changes of white and red eyes, small and large within DNA molecules located in chromosomes that seg- wings, and similar traits in the drosophila, Morgan con- regate during mitosis, and that it is only the DNA that cluded that genes were linearly aligned in proximity one determines heredity. Consequently, these studies are to the other (linkage), but even so during the transfer considered as the beginning of the period of molecular 1–3 they could separate and get into new units individually. genetics . Thus the chromosome of a new entity contains the collec- In 1950 Erwin Chargaff in his studies of the arrange- tion of linked and independent genes which are all inter- ment of nucleotide bases showed that the DNA molecule -related (linked genes) and most often commonly inher- consist of four acid bases, showing a constant proportion ited. Furthermore, Morgan discovered that the offspring of purine and pyrimidine bases, i.e. adenine and thy- sometimes inherited only single genes from a chromo- mine, guanine and cytosine1. some and that single genes were sometimes cross-trans- Alfred Hershey and Martha Chase in 1952 showed ferred from one chromosome to the other – the transfer that viral penetration into bacterium leads to transfer of referred to as crossing over. The results of Morgan’s solely DNA molecules, followed by multiplication of many studies became essential in the construction of first ge- new viruses within the bacterium. This finding con- netic maps3,6,7. firmed the previous assumption regarding the transfer of By studying chromosome localisation and linear se- genetic informatioan via DNA, i.e. the chemical transfer quence of genes, as well as their mutational capacity, Mor- of hereditary traits. gan provided a cytogenetic explanation of Mendel’s obser- These study results stimulated many scientists to be- vations and clarified the mechanism of heredity involved gin new research of the chemical structure of genes and in fertilisation, leading to, what is nowadays referred to as soon it was established that the DNA molecule is formed Mendel-Morgan chromosomal theory of heredity. by a nitrogen base, deoxyribosis and phosphate group. Further research was aimed at discovering the posi- The British scientists Maurice Hugh Frederick Wil- tion of genes in a cell nucleus, i.e. in chromosomes. In kins and Rosalind Elsie Franklin have in 1953 studied 1913, Alfred H. Sturtevant, a student of Morgan, de- the crystallographic structure of partially dehydrated scribed linear sequence of genes in chromosomes. It was and crystalline DNA molecule by diffraction of X-rays also confirmed that during cell division (mitosis) the and have
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