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A biológia története az élővilág tanulmányozásának történetét követi végig az ókortól napjainkig. Bár a biológi, mint önálló tudományág csak a 19. században alakult ki, a biológiai tudományok a korábbi orvostudományi és természetrajzi hagyományokból alakultak ki, melyek egészen az ókori görög Galenusig és Arisztotelészig vezethetőek vissza. A reneszánsz korában és a korai újorban a biológiai gondolkodásmódot forradalmasította az újonnan  felélénkülő érdeklődés az empirizmus iránt és számos új szervezet felfedezése. A mozgalom élén járt  Vesalius és William Harvey, akik élettani kutatásaikban  kísérleti módszereket  alkalmazták és gondos megfigyeléseket tettek; valamint  Carl Linnaeus és Buffon természettudósok, akik megkezdték az élő és fosszilis szervezetek tudományos osztályozását és az organizmusok fejlődésének és viselkedésének tanulmányozását. A mikroszkópia felfedezése a mikroorganizmusok korábban ismeretlen világa felé nyitott kaput és lefektette a sejtelmélet alapjait. A természeti teológia növekvő fontossága -- részben a mechanikai filozófiára válaszként --elősegítette a természetrajz fejlődését (bár segítette a teológiai érvelés berögzülését is.

A 18-19. század folyamán a biológiai tudományok (növény és állattan) egyre elismertebb tudományterületté váltak. Lavoisier és más természettudósok megindították azt a folyamatot, melynek során az élő és élettelen világ egyre közelebb került egymáshoz a közös fizikai és kémiai törvények miatt. Explorer-naturalists such as Alexander von Humboldt és más kutató természettudósok kutatni kezdték a szervezetek és környezetük közötti kölcsönhatásokat és  azt, hogy ezek hogyan függenek a fölrajzi adottságoktól – ezzel lefektették a   biogeográfia (természetföldrajz), ökológia és etológia   tudományának alapjait. A természetkutatók már egyre kevésbé fogadták el az eszencializmust és fontosabbnak találták a kihalás és a fajok mutabilitását. A sejtelmélet új perspektívába helyezte az élet alapjait. Ezek a fejlődések, valamint az embriológiai és paleontológiai ismeretek bővülése vezetett el ahhoz, hogy Charles Darwin felállította az evolúció és a természetes kiválasztódás elméletét. A 19. század végére a spontán képződés eszméje elbukott és felváltotta a betegségek csíraelmélete, bár a biológiai öröklődés mikéntje egyelőre ismeretlen maradt.

A 20. század elején Gregor Mendel  életművének ismertté válásával  a genetika gyors fejlődésnek indult --   Thomas Hunt Morgannek és tanítványainak köszönhetően -- és az  1930-as évekre a populációgenetika és a természetes kiválasztódás kombinációjával kialakult az új-Darwinizmus eszméje. Egyre újabb tudományágak indultak fejlődésnek, különösen miután James D. Watson és Francis Crick  megfejtették a DNS szerkezetét. A közpomti dogma  (Central Dogma) felállítása és a genetikai kód feltörése után  a biológiai tudomány kettévált a  szervezeti szintű biológia  – az a terület, amely teljes élő szervezetekkel és ezek csoportjaiaval foglalkozik  –  és a  sejt- és molekuláris biológia területeire. A 20. század végére a genomika és proteomika előretörésével a két nagy terület újra közeledni kezdett egymáshoz, mivel a szervezet-biológusok molekuláris technikákat kezdtek használni; a molekuláris és sejtbiológusok pedig vizsgálni kezdték a gének és a környezet kölcsönhatásait valamint a szervezetek természeti populációinak genetikáját.

Ókori görög biológiai hagyomány
 The Pre-Socratic philosophers asked many questions about life but produced little systematic knowledge of specifically biological interest&mdash;though the attempts of the atomists to explain life in purely physical terms would recur periodically through the history of biology. However, the medical theories of Hippocrates and his followers, especially humorism, had a lasting impact.

The philosopher Aristotle was the most influential scholar of the living world from antiquity. Though his early work in natural philosophy was speculative, Aristotle's later biological writings were more empirical, focusing on biological causation and the diversity of life. He made countless observations of nature, especially the habits and attributes of plants and animals in the world around him, which he devoted considerable attention to categorizing. In all, Aristotle classified 540 animal species, and dissected at least 50. He believed that intellectual purposes, formal causes, guided all natural processes.

Aristotle, and nearly all scholars after him until the 18th century, believed that creatures were arranged in a graded scale of perfection rising from plants on up to humans: the scala naturae or Great Chain of Being. Aristotle's successor at the Lyceum, Theophrastus, wrote a series of books on botany&mdash;the History of Plants&mdash;which survived as the most important contribution of antiquity to botany, even into the Middle Ages. Many of Theophrastus' names survive into modern times, such as carpos for fruit, and pericarpion for seed vessel. Pliny the Elder was also known for his knowledge of plants and nature, and was the most prolific compiler of zoological descriptions.

A few scholars in the Hellenistic period under the Ptolemies&mdash;particularly Herophilus of Chalcedon and Erasistratus of Chios&mdash;amended Aristotle's physiological work, even performing experimental dissections and vivisections. Claudius Galen became the most important authority on medicine and anatomy. Though a few ancient atomists such as Lucretius challenged the teleological Aristotelian viewpoint that all aspects of life are the result of design or purpose, teleology (and after the rise of Christianity, natural theology) would remain central to biological thought essentially until the 18th and 19th centuries. In the words of Ernst Mayr, "Nothing of any real consequence happened in biology after Lucretius and Galen until the Renaissance." The ideas of the Greek traditions of natural history and medicine survived, but they were generally taken unquestioningly.

17-18. század


Extending the work of Vesalius into experiments on still living bodies (of both humans and animals), William Harvey and other natural philosophers investigated the roles of blood, veins and arteries. Harvey's De motu cordis in 1628 was the beginning of the end for Galenic theory, and alongside Santorio Santorio's studies of metabolism, it served as an influential model of quantitative approaches to physiology.

In the early 17th century, the micro-world of biology was just beginning to open up. A few lensmakers and natural philosophers had creating crude microscopes since the late 16th century, and Robert Hooke published the seminal Micrographia based on observations with his own compound microscope in 1665. But it was not until Antony van Leeuwenhoek's dramatic improvements in lensmaking beginning in the 1670s&mdash;ultimately producing up to 200-fold magnification with a single lens&mdash;that scholars discovered spermatozoa, bacteria, infusoria and the sheer strangeness and diversity of microscopic life. Similar investigations by Jan Swammerdam led to new interest in entomology and built the basic techniques of microscopic dissection and staining.



As the microscopic world was expanding, the macroscopic world was shrinking. Botanists such as John Ray worked to incorporate the flood of newly discovered organisms shipped from across the globe into a coherent taxonomy, and a coherent theology (natural theology). Debate over another flood, the Noachian, catalyzed the development of paleontology; in 1669 Nicholas Steno published an essay on how the remains of living organisms could be trapped in layers of sediment and mineralized to produce fossils. Although Steno's ideas about fossilization were well known and much debated among natural philosophers, an organic origin for all fossils would not be accepted by all naturalists until the end of the 18th century due to philosophical and theological debate about issues such as the age of the earth and extinction.

Systematizing, naming and classifying dominated natural history throughout much of the 17th and 18th centuries. Carl Linnaeus published a basic taxonomy for the natural world in 1735 (variations of which have been in use ever since), and in the 1750s introduced scientific names for all his species. While Linnaeus conceived of species as unchanging parts of a designed hierarchy, the other great naturalist of the 18th century, Georges-Louis Leclerc, Comte de Buffon, treated species as artificial categories and living forms as malleable&mdash;even suggesting the possibility of common descent. Though he was opposed to evolution, Buffon is a key figure in the history of evolutionary thought; his work would influence the evolutionary theories of both Lamarck and Darwin.

The discovery and description of new species and the collection of specimens became a passion of scientific gentlemen and a lucrative enterprise for entrepreneurs; many naturalists traveled the globe in search of scientific knowledge and adventure.

Az evolúció és a biogeográfia
 The most significant evolutionary theory before Darwin's was that of Jean-Baptiste Lamarck; based on the inheritance of acquired characteristics (an inheritance mechanism that was widely accepted until the 20th century), it described a chain of development stretching from the lowliest microbe to humans. The British naturalist Charles Darwin, combining the biogeographical approach of Humboldt, the uniformitarian geology of Lyell, Thomas Malthus's writings on population growth, and his own morphological expertise, created a more successful evolutionary theory based on natural selection; similar evidence lead Alfred Russel Wallace to independently reach the same conclusions.

The 1859 publication of Darwin's theory in On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life is often considered the central event in the history of modern biology. Darwin's established credibility as a naturalist, the sober tone of the work, and most of all the sheer strength and volume of evidence presented, allowed Origin to succeed where previous evolutionary works such as the anonymous Vestiges of Creation had failed. Most scientists were convinced of evolution and common descent by the end of the 19th century. However, natural selection would not be accepted as the primary mechanism of evolution until well into the 20th century, as most contemporary theories of heredity seemed incompatible with the inheritance of random variation.

Wallace, building on earlier work by Humbolt and Darwin, made major contributions to biogeography by focusing on the distribution of closely allied species with particular attention to the effects of geographical barriers during his research in the Amazon basin and the Malay archipelago. He discovered the Wallace line dividing the fauna of the Malay archipelago between a zone allied with Asia and a zone allied with Australia. The ornithologist Philip Sclater, drawing on the work of Wallace and others, proposed a system of 6 major geographical regions to describe the distribution of bird species in the world. Wallace and others would in turn extend Sclater's system from birds to animals of all kinds.

The scientific study of heredity grew rapidly in the wake of Darwin's Origin of Species with the work of Francis Galton and the biometricians. The origin of genetics is usually traced to the 1866 work of the monk Gregor Mendel, who would later be credited with the laws of inheritance. However, his work was not recognized as significant until 35 years afterward. In the meantime, a variety of theories of inheritance (based on pangenesis, orthogenesis, or other mechanisms) were debated and investigated vigorously. Embryology and ecology also became central biological fields, especially as linked to evolution and popularized in the work of Ernst Haeckel. Most of the 19th century work on heredity, however, was not in the realm of natural history, but that of experimental physiology.

Sejtelmélet, embriológia és csíraelmélet
<!-- Advances in microscopy also had a profound impact on biological thinking. In the early 19th century, a number of pointed to the central importance of the cell. In 1838 and 1839, Schleiden and Schwann began promoting the the ideas that (1) the basic unit of organisms is the cell and (2) that individual cells have all the characteristics of life, though they opposed the idea that (3) all cells come from the division of other cells. Thanks to the work of Robert Remak and Rudolf Virchow, however, by the 1860s most biologists accepted all three tenets of what came to be known as cell theory.

Cell theory led biologists to re-envision individual organisms as interdependent assemblages of individual cells. Scientists in the rising field of cytology, armed with increasingly powerful microscopes and new staining methods, soon found that even single cells were far more complex than the homogeneous fluid-filled chambers of described by earlier microscopists. Robert Brown had described the nucleus in 1831, and by the end of the 19th century cytologists identified many of the key cell components: chromosomes, centrosomes mitochondria, chloroplasts, and other structures made visible through staining. Between 1874 and 1884 Walther Flemming described the discrete stages of mitosis, showing that they were not artifacts of staining but occurred in living cells, and moreover, that chromosomes doubled in number just before the cell divided and a daughter cell was produced. Much of the research on cell reproduction came together in August Weismann's theory of heredity: he identified the nucleus (in particular chomosomes) as the hereditary material, proposed the distinction between somatic cells and germ cells (arguing that chromosome number must be halved for germ cells, a precursor to the concept of meiosis), and adopted Hugo de Vries's theory of pangenes. Weismannism was extremely influential, especially in the new field of experimental embryology.

By the mid 1850s the miasma theory of disease was largely superseded by the germ theory of disease, creating extensive interest in microorganisms and their interactions with other forms of life. By the 1880s, bacteriology was becoming a coherent discipline, especially through the work of Robert Koch, who introduced methods for growing pure cultures on agar gels containing specific nutrients in Petri dishes. The long-held idea that living organisms could easily originate from nonliving matter (spontaneous generation) was attacked in a series of experiments carried out by Louis Pasteur, while debates over vitalism vs. mechanism (a perennial issue since the time of Aristotle and the Greek atomists) continued apace.

Klasszikus és modern genetika, evolúciós elmélet
<!--  1900 marked the so-called rediscovery of Mendel: Hugo de Vries, Carl Correns, and Erich von Tschermak independently arrived at Mendel's laws (which were not actually present in Mendel's work). Soon after, cytologists (cell biologists) proposed that chromosomes were the hereditary material. Between 1910 and 1915, Thomas Hunt Morgan and the "Drosophilists" in his fly lab forged these two ideas&mdash;both controversial&mdash;into the "Mendelian-chromosome theory" of heredity. They quantified the phenomenon of genetic linkage and postulated that genes reside on chromosomes like beads on string; they hypothesized crossing over to explain linkage and constructed genetic maps of the fruit fly Drosophila melanogaster, which became a widely used model organism.

Hugo de Vries tried to link link the new genetics with evolution; building on his work with heredity and hybridization, he proposed a theory of mutationism, which was widely accepted in the early 20th century. Lamarckism also had many adherents. Darwinism was seen as incompatible with the continuously variable traits studied by biometricians, which seemed only partially heritable. In the 1920s and 1930s&mdash;following the acceptance of the Mendelian-chromosome theory&mdash; the emergence of the discipline of population genetics, with the work of R.A. Fisher, J.B.S. Haldane and Sewall Wright, unified the idea of evolution by natural selection with Mendelian genetics producing the modern synthesis. The inheritance of acquired characters was rejected, while mutationism gave way as genetic theories matured.

In the second half of the century the ideas of population genetics began to be applied in the new discipline of the genetics of behavior, sociobiology, and, especially in humans, evolutionary psychology. In the 1960s W.D. Hamilton and others developed game theory approaches to explain altruism from an evolutionary perspective through kin selection. The possible origin of higher organisms through endosymbiosis, and contrasting approaches to molecular evolution in the gene-centered view (which held selection as the predominant cause of evolution) and the neutral theory (which made genetic drift a key factor) spawned perennial debates over the proper balance of adaptationism and contingency in evolutionary theory.

In the 1970s Stephen Jay Gould and Niles Eldredge proposed the theory of punctuated equilibrium which holds that stasis is the most prominent feature of the fossil record, and that most evolutionary changes occur rapidly over relatively short periods of time. In 1980 Luis Alvarez and Walter Alvarez proposed the hypothesis that an impact event was responsible for the Cretaceous-Tertiary extinction event. Also in the early 1980s, statistical analysis of the fossil record of marine organisms published by Jack Sepkoski and David M. Raup lead to a better appreciation of the importance of mass extinction events to the history of life on earth.

A molekuláris biológia eredete
<!--  Following the rise of classical genetics, many biologists&mdash;including a new wave of physical scientists in biology&mdash;pursued the question of the gene and its physical nature. Warren Weaver&mdash;head of the science division of the Rockefeller Foundation&mdash;issued grants to promote research that applied the methods of physics and chemistry to basic biological problems, coining the term molecular biology for this approach in 1938; many of the significant biological breakthroughs of the 1930s and 1940s were funded by the Rockefeller Foundation.



Like biochemistry, the overlapping disciplines of bacteriology and virology (later combined as microbiology), situated between science and medicine, developed rapidly in the early 20th century. Félix d'Herelle's isolation of bacteriophage during World War I initiated a long line of research focused of phage viruses and the bacteria they infect.

The development of standard, genetically uniform organisms that could produce repeatable experimental results was essential for the development of molecular genetics. After early work with Drosophila and maize, the adoption of simpler model systems like the bread mold Neurospora crassa made it possible to connect genetics to biochemistry, most importantly with Beadle and Tatum's "one gene, one enzyme" hypothesis in 1941. Genetics experiments on even simpler systems like tobacco mosaic virus and bacteriophage, aided by the new technologies of electron microscopy and ultracentrifugation, forced scientists to re-evaluate the literal meaning of life; virus heredity and reproducing nucleoprotein cell structures outside the nucleus ("plasmagenes") complicated the accepted Mendelian-chromosome theory.



Oswald Avery showed in 1943 that DNA was likely the genetic material of the chromosome, not its protein; the issue was settled decisively with the 1952 Hershey-Chase experiment&mdash;one of many contribution from the so-called phage group centered around physicist-turned-biologist Max Delbrück. In 1953 James D. Watson and Francis Crick, building on the work of Maurice Wilkins and Rosalind Franklin, suggested that the structure of DNA was a double helix. In their famous paper "Molecular structure of Nucleic Acids", Watson and Crick noted coyly, "It has not escaped our notice that the specific pairing we have postulated immediately suggests a possible copying mechanism for the genetic material." After the 1958 Meselson-Stahl experiment confirmed the semiconservative replication of DNA, it was clear to most biologists that nucleic acid sequence must somehow determine amino acid sequence in proteins; physicist George Gamow proposed that a fixed genetic code connected proteins and DNA. Between 1953 and 1961, there were few known biological sequences&mdash;either DNA or protein&mdash;but an abundance of proposed code systems, a situation made even more complicated by expanding knowledge of the intermediate role of RNA. To actually decipher the code, it took an extensive series of experiments in biochemistry and bacterial genetics, between 1961 and 1966&mdash;most importantly the work of Nirenberg and Khorana.



A molekuláris biológia elterjedése
<!--  In addition to the Division of Biology at Caltech, the Laboratory of Molecular Biology (and its precursors) at Cambridge, and a handful of other institutions, the Pasteur Institute became a major center for molecular biology research in the late 1950s. Scientists at Cambridge, led by Max Perutz and John Kendrew, focused on the rapidly developing field of structural biology, combining X-ray crystallography with molecular modelling and the new computational possibilities of digital computing (benefiting both directly and indirectly from the military funding of science). A number of biochemists led by Fred Sanger later joined the Cambridge lab, bringing together the study of macromolecular structure and function. At the Pasteur Institute, François Jacob and Jacques Monod followed the 1959 PaJaMo experiment with a series of publications regarding the lac operon that established the concept of gene regulation and identified what came to be known as messenger RNA. By the mid-1960s, the intellectual core of molecular biology&mdash;a model for the molecular basis of metabolism and reproduction&mdash; was largely complete.

The late 1950s to the early 1970s was a period of intense research and institutional expansion for molecular biology, which had only recently become a somewhat coherent discipline. In what organismic biologist E. O. Wilson called "The Molecular Wars", the methods and practitioners of molecular biology spread rapidly, often coming to dominate departments and even entire disciplines. Molecularization was particularly important in genetics, immunology, embryology, and neurobiology, while the idea that life is controlled by a "genetic program"&mdash;a metaphor Jacob and Monod introduced from the emerging fields of cybernetics and computer science&mdash;became an influential perspective throughout biology. Immunology in particular became linked with molecular biology, with innovation flowing both ways: the clonal selection theory developed by Niels Jerne and Frank Macfarlane Burnet in the mid 1950s helped shed light on the general mechanisms of protein synthesis.

Resistance to the growing influence molecular biology was especially evident in evolutionary biology. Protein sequencing had great potential for the quantitative study of evolution (through the molecular clock hypothesis), but leading evolutionary biologists questioned the relevance of molecular biology for answering the big questions of evolutionary causation. Departments and disciplines fractured as organismic biologists asserted their importance and independence: Theodosius Dobzhansky made the famous statement that "nothing in biology makes sense except in the light of evolution" as a response to the molecular challenge. The issue became even more critical after 1968; Motoo Kimura's neutral theory of molecular evolution suggested natural selection was not the ubiquitous cause of evolution, at least at the molecular level, and that molecular evolution might be a fundamentally different process from morphological evolution. (Resolving this "molecular/morphological paradox" has been a central focus of molecular evolution research since the 1960s.)

Biotechnológia, génmanipuláció és genomika
<!--

Biotechnology in the general sense has been an important part of biology since the late 19th century. With the industrialization of brewing and agriculture, chemists and biologists became aware of the great potential of human-controlled biological processes. In particular, fermentation proved a great boon to chemical industries. By the early 1970s, a wide range of biotechnologies were being developed, from drugs like penicillin and steroids to foods like Chlorella and single-cell protein to gasohol&mdash;as well as a wide range of hybrid high-yield crops and agricultural technologies, the basis for the Green Revolution.



A rekombináns DNS
Biotechnology in the modern sense of genetic engineering began in the 1970s, with the invention of recombinant DNA techniques. Restriction enzymes were discovered and characterized in the late 1960s, following on the heels of the isolation, then duplication, then synthesis of viral genes. Beginning with the lab of Paul Berg in 1972 (aided by EcoRI from Herbert Boyer's lab, building on work with ligase by Arthur Kornberg's lab), molecular biologists put these pieces together to produce the first transgenic organisms. Soon after, others began using plasmid vectors and adding genes for antibiotic resistance, greatly increasing the reach of the recombinant techniques.

Wary of the potential dangers (particularly the possibility of a prolific bacteria with a viral cancer-causing gene), the scientific community as well as a wide range of scientific outsiders reacted to these developments with both enthusiasm and fearful restraint. Prominent molecular biologists led by Berg suggested a temporary moratorium on recombinant DNA research until the dangers could be assessed and policies could be created. This moratorium was largely respected, until the participants in the 1975 Asilomar Conference on Recombinant DNA created policy recommendations and concluded that the technology could be used safely.

Following Asilomar, new genetic engineering techniques and applications developed rapidly. DNA sequencing methods improved greatly (pioneered by Fred Sanger and Walter Gilbert), as did oligonucleotide synthesis and transfection techniques. Researchers learned to control the expression of transgenes, and were soon racing&mdash;in both academic and industrial contexts&mdash;to create organisms capable of expressing human genes for the production of human hormones. However, this was a more daunting task than molecular biologists had expected; developments between 1977 and 1980 showed that, due to the phenomena of split genes and splicing, higher organisms had a much more complex system of gene expression than the bacteria models of earlier studies. The first such race, for synthesizing human insulin, was won by Genentech. This marked the beginning of the biotech boom (and with it, the era of gene patents), with an unprecedented level of overlap between biology, industry, and law.

Irodalom

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 * Sturtevant, A. H. A History of Genetics.  Cold Spring Harbor Laboratory Press: Cold Spring Harbor, 2001.  ISBN 0879696079
 * Thackray, Arnold, ed. Private Science: Biotechnology and the Rise of the Molecular Sciences. University of Pennsylvania Press: Philadelphia, 1998. ISBN 0812234286
 * Wilson, Edward O. Naturalist.  Island Press, 1994.
 * Zimmer, Carl. Evolution: the triumph of an idea. HarperCollins: New York, 2001. ISBN 0-06-113840-1
 * Zimmer, Carl. Evolution: the triumph of an idea. HarperCollins: New York, 2001. ISBN 0-06-113840-1

Külső hivatkozások

 * International Society for History, Philosophy, and Social Studies of Biology - professional history of biology organization
 * History of Biology - Historyworld article
 * History of Biology at Bioexplorer.Net - a collection of history of biology links
 * Biology - historically-oriented article on Citizendium
 * Miall, L. C. (1911) History of biology. Watts & Co. London

History of biology Geschichte der Biologie Historia de la biología Histoire de la biologie היסטוריה של הביולוגיה 生物学史 Historia biologii História da biologia Dejiny biológie