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/study/ and is defined as the science of life and living organisms. An organism is a living entity consisting of one cell e.g. bacteria, or several cells e.g. animals, plants and fungi. Aspects of biological science range from the study of molecular mechanisms in cells, to the classification and behaviour of organisms, how species evolve and interaction between ecosystems. The study of biology can be divided into different disciplines Ethology Evolutionary Biology Physiology enetics !olecular Biology !orphology "ystematics Ecology
Biology often overlaps with other sciences# for e$ample, biochemistry and to$icology with biology, chemistry, and medicine# biophysics with biology and physics# stratigraphy with biology and geography# astrobiology with biology and astronomy. "ocial sciences such as geography, philosophy, psychology and sociology can also interact with biology, for e$ample, in administration of biological resources, developmental biology, biogeography, evolutionary psychology and ethics. Biology is a natural science concerned with the study of life and living organisms, including their structure, function, growth, origin, evolution, distribution, and ta$onomy. "ubdisciplines of biology are recogni%ed on the basis of the scale at which organisms are studied and the methods used to study them& biochemistry e$amines the rudimentary chemistry of life# molecular biology studies the comple$ interactions of systems of biological molecules# cellular biology e$amines the basic building block of all life, the cell# physiology e$amines the physical and chemical functions of the tissues, organs, and organ systems of an organism# and ecology e$amines how various organisms interact and associate with their environment. These are the main branches of biology&'()*'(+* Aerobiology , the study of airborne organic particles Agriculture , the study of producing crops from the land, with an emphasis on practical applications Anatomy , the study of form and function, in plants, animals, and other organisms, or specifically in humans Bioengineering , the study of biology through the means of engineering with an emphasis on applied knowledge and especially related to biotechnology Biomathematics or Mathematical Biology - the .uantitative or mathematical study of biological processes, with an emphasis on modeling Biotechnology , a new and sometimes controversial branch of biology that studies the manipulation of living matter, including genetic modification and synthetic biology Botany , the study of plants
ell biology , the study of the cell as a complete unit, and the molecular and chemical interactions that occur within a living cell !cology , the study of the interactions of living organisms with one another and with the nonliving elements of their environment !pidemiology , a ma/or component of public health research, studying factors affecting the health of populations !pigenetics , the study of heritable changes in gene e$pression or cellular phenotype caused by mechanisms other than changes in the underlying 01A se.uence !thology , the study of animal behavior !volutionary Biology , the study of the origin and descent of species over time Genetics , the study of genes and heredity "erpetology , the study of reptiles and amphibians "istology , the study of cells and tissues, a microscopic branch of anatomy Ichthyology , the study of fish Marine Biology , the study of ocean ecosystems, plants, animals, and other living beings Microbiology , the study of microscopic organisms 2microorganisms3 and their interactions with other living things Molecular Biology , the study of biology and biological functions at the molecular level, some cross over with biochemistry Mycology , the study of fungi Oceanography , the study of the ocean, including ocean life, environment, geography, weather, and other aspects influencing the ocean Oncology , the study of cancer processes, including virus or mutation oncogenesis, angiogenesis and tissues remoldings #opulation genetics , the study of changes in gene fre.uencies in populations of organisms #aleontology , the study of fossils and sometimes geographic evidence of prehistoric life #athobiology or pathology , the study of diseases, and the causes, Parasitology , the study of parasites and parasitism #harmacology , the study and practical application of preparation, use, and effects of drugs and synthetic medicines Physiology , the study of the functioning of living organisms and the organs and parts of living organisms #hytopathology , the study of plant diseases 2also called Plant Pathology3 #sychobiology , the study of the biological bases of psychology $irology , the study of viruses and some other virus-like agents %oology , the study of animals, including classification, physiology, development, and behavior 2"ee also Entomology, Ethology, 4erpetology, 5chthyology, !ammalogy, and 6rnithology3
Basic #rinciples of Biology The foundation of biology as it e$ists today is based on five basic principles. They are the cell theory, gene theory, evolution, homeostasis, and laws of thermodynamics. 7ell Theory& all living organisms are composed of cells. The cell is the basic unit of life. ene Theory& traits are inherited through gene transmission. onchromosomes and consist of 01A. enes are located
Evolution& any genetic change in a population that is inherited over several generations. These changes may be small or large, noticeable or not so noticeable.
4omeostasis& ability to maintain a constant internal environment in response to environmental changes. Thermodynamics& energy is constant and energy transformation is not completely efficient.
&ubdiciplines of Biology The field of biology is very broad in scope and can be divided into several disciplines. 5n the most general sense, these disciplines are categori%ed based on the type of organism studied. 8or e$ample, %oology deals with animal studies, botany deals with plant studies, and microbiology is the study of microorganisms. These fields of study can be broken down further into several speciali%ed sub-disciplines. "ome of which include anatomy, cell biology, genetics, and physiology. "I&TO'I AL BA (G'O)*+ O, BIOLOGY The history of biology traces the study of the living world from ancient to modern times. Although the concept of biology as a single coherent field arose in the 9:th century, the biological sciences emerged from traditions of medicine andnatural history reaching back to ayurveda, ancient Egyptian medicine and the works of Aristotle and alen in the ancient reco-;oman world. This ancient work was further developed in the !iddle Ages by !uslim physicians and scholars such as Avicenna. 0uring the European ;enaissance and early modern period, biological thought was revolutioni%ed in Europe by a renewed interest in empiricism and the discovery of many novel organisms. Prominent in this movement were <esalius and 4arvey, who used e$perimentation and careful observation in physiology, and naturalists such as =innaeus and Buffon who began to classify the diversity of life and the fossil record, as well as the development and behavior of organisms. !icroscopy revealed the previously unknown world of microorganisms, laying the groundwork for cell theory. The growing importance of natural theology, partly a response to the rise of mechanical philosophy, encouraged the growth of natural history 2although it entrenched the argument from design3. 6ver the 9>th and 9:th centuries, biological sciences such as botany and %oology became increasingly professional scientific disciplines. =avoisier and other physical scientists began to connect the animate and inanimate worlds through physics and chemistry. E$plorer-naturalists such as Ale$ander von 4umboldt investigated the interaction between organisms and their environment, and the ways this relationship depends on geography,laying the foundations forbiogeography, ecology and ethology. 1aturalists began to re/ect essentialism and consider the importance of e$tinction and the mutability of species. 7ell theory provided a new perspective on the fundamental basis of life. These developments, as well as the results from embryology and paleontology, were synthesi%ed in 7harles 0arwin?s theory of evolution by natural selection. The end of the 9:th century saw the fall of spontaneous generation and the rise of thegerm theory of disease, though the mechanism of inheritance remained a mystery.
5n the early @Ath century, the rediscovery of !endel?s work led to the rapid development of genetics by Thomas 4unt !organ and his students, and by the 9:BAs the combination of population genetics and natural selection in the Cneo-0arwinian synthesisC. 1ew disciplines developed rapidly, especially after Datson and 7rick proposed the structure of 01A. 8ollowing the establishment of the 7entral 0ogma and the cracking of the genetic code, biology was largely split between organismal biology,the fields that deal with whole organisms and groups of organisms,and the fields related to cellular and molecular biology. By the late @Ath century, new fields like genomics and proteomics were reversing this trend, with organismal biologists using molecular techni.ues, and molecular and cell biologists investigating the interplay between genes and the environment, as well as the genetics of natural populations of organisms.
Etymology of "biology"[edit]
The word biology is formed by combining the reek EFGH 2bios3, meaning ClifeC, and the suffi$ ?logy?, meaning Cscience ofC, Cknowledge ofC, Cstudy ofC, based on the reek verb IJKLMN, ?legein? Cto selectC, Cto gatherC 2cf. the noun IOKGH, ?logos? CwordC3. The term biology in its modern sense appears to have been introduced independently by Thomas Beddoes 2in 9P::3,'9* Qarl 8riedrich Burdach 2in 9>AA3, ottfried ;einhold Treviranus 2Biologie oder Philosophie der lebenden Natur , 9>A@3 and Rean-Baptiste =amarck 2Hydrogologie, 9>A@3.'@*'B* The word itself appears in the title of <olume B of !ichael 7hristoph 4anow?s Philosophiae naturalis sive physicae dogmaticae: Geologia, biologia, phytologia generalis et dendrologia , published in 9P((. Before biology, there were several terms used for the study of animals and plants. Natural history referred to the descriptive aspects of biology, though it also included mineralogy and other non-biological fields# from the !iddle Ages through the ;enaissance, the unifying framework of natural history was the scala naturae or reat 7hain of Being. Natural philosophy and natural theology encompassed the conceptual and metaphysical basis of plant and animal life, dealing with problems of why organisms e$ist and behave the way they do, though these sub/ects also included what is now geology, physics, chemistry, and astronomy. Physiology and 2botanical3 pharmacology were the province of medicine. Botany, zoology, and 2in the case of fossils3 geology replaced natural history and natural philosophy in the 9>th and 9:th centuries before biology was widely adopted.')*'+* To this day, CbotanyC and C%oologyC are widely used, although they have been /oined by other sub-disciplines of biology, such as mycology and molecular biology.
4umans first domesticated plants for farming, then livestock animals to accompany the resulting sedentary societies.'(* The ancient cultures of !esopotamia, Egypt, the 5ndian subcontinent, and 7hina, among others, produced renowned surgeons and students of the natural sciences such as "usruta and Shang Shong/ing, reflecting independent sophisticated systems of natural philosophy. 4owever, the roots of modern biology are usually traced back to the secular tradition of ancient reek philosophy.'P*
Ancient
hinese traditions-edit.
5n ancient 7hina, biological topics can be found dispersed across several different disciplines, including the work of herbologists, physicians, alchemists, and philosophers. The Taoist tradition of 7hinese alchemy, for e$ample, can be considered part of the life sciences due to its emphasis on health 2with the ultimate goal being the eli$ir of life3. The system of classical 7hinese medicine usually revolved around the theory of yin and yang, and the five phases.'>* Taoist philosophers, such as Shuang%i in the )th century B7E, also e$pressed ideas related to evolution, such as denying the fi$ity of biological species and speculating that species had developed differing attributes in response to differing environments. ':*
8rontispiece to a 9()) version of the e$panded and illustrated edition of Historia Plantarum 2ca. 9@AA3, which was originally written around BAA B7
The pre-"ocratic philosophers asked many .uestions about life but produced little systematic knowledge of specifically biological interest,though the attempts of the atomists to e$plain life in purely physical terms would recur periodically through the history of biology. 4owever, the medical theories of 4ippocrates and his followers, especially humorism, had a lasting impact.'9B* The philosopher Aristotle was the most influential scholar of the living world from classical anti.uity. 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. 4e 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 categori%ing. 5n all, Aristotle classified +)A animal species, and dissected at least +A. 4e believed that intellectual purposes, formal causes, guided all natural processes.'9)* Aristotle, and nearly all Destern scholars after him until the 9>th century, believed that creatures were arranged in a graded scale of perfection rising from plants on up to humans& the scala naturae or reat 7hain of Being.'9+* Aristotle?s successor at the=yceum, Theophrastus, wrote a series of books on botany,the History of Plants,which survived as the most important contribution of anti.uity to botany, even into the !iddle Ages. !any of Theophrastus? names survive into modern times, such as carposfor 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 %oological descriptions.'9(* A few scholars in the 4ellenistic period under the Ptolemies,particularly 4erophilus of 7halcedon and Erasistratus of 7hios,amended Aristotle?s physiological work, even performing dissections and vivisections.'9P* 7laudius alen became the most important authority on medicine and anatomy. Though a few ancient atomists such as =ucretius challenged the teleological Aristotelian viewpoint that all aspects of life are the result of design or purpose,
teleology 2and after the rise of 7hristianity, natural theology3 would remain central to biological thought essentially until the 9>th and 9:th centuries. Ernst D. !ayr argued that C1othing of any real conse.uence happened in biology after =ucretius and alen until the ;enaissance.C '9>* The ideas of the reek traditions of natural history and medicine survived, but they were generally taken un.uestioningly in medieval Europe.'9:*
A biomedical work by 5bn al-1afis, an early adherent of e$perimental dissection who discovered the pulmonary circulation and coronary circulation.
The decline of the ;oman Empire led to the disappearance or destruction of much knowledge, though physicians still incorporated many aspects of the reek tradition into training and practice. 5n By%antium and the5slamic world, many of the reek works were translated into Arabic and many of the works of Aristotle were preserved.'@A* !edieval !uslim physicians, scientists and philosophers made significant contributions to biological knowledge between the >th and 9Bth centuries during what is known as the C5slamic olden AgeC or C!uslim Agricultural ;evolutionC. 5n %oology, for e$ample, the AfroArab scholar al-Rahi% 2P>9U>(:3 described early evolutionary ideas'@9*'@@* such as the struggle for e$istence.'@B* 4e also introduced the idea of a food chain,'@)* and was an early adherent ofenvironmental determinism.'@+* The Persian biologist Al-0inawari 2>@>U>:(3 authored the Boo" of Plants, in which he described at least (BP species and discussed plant development, plant growth and the production of flowers and fruit.'@(* Persian polymath Abu ;ayhan Biruni described the idea of artificial selection and argued that nature works in much the same way, an idea that has been compared to natural selection.'@P* 5n e$perimental medicine, the Persian physician Avicenna 2:>AU9ABP3 introduced clinical trials and clinical pharmacology in %he &anon of $edicine,'@>* which remained an authoritative te$t in European medical education up until the 9Pth century. '@:*'BA* TheAndalusian-
Arabian physician Aven%oar 29A:9U99(93 was an early adherent of e$perimental dissection and autopsy, which he carried out to prove that the skin disease scabies was caused by a parasite, a discovery which upset the theory of humorism.'B9*4e also introduced e$perimental surgery,'B@* where animal testing is used to e$periment with surgical techni.ues prior to using them on humans.'BB* 0uring a famine in Egypt in 9@AA, Abd-ellatif observed and e$amined a large number of skeletons, and he discovered that alen was incorrect regarding the formation of the bones of the lower /aw and sacrum.'B)* 5n the early 9Bth century, the Andalusian-Arabian biologist Abu al-Abbas al-1abati developed an early scientific method for botany, introducing empirical and e$perimental techni.ues in the testing, description and identification of numerous materia medica, and separating unverified reports from those supported by actual tests and observations.'B+* 4is student 5bn al-Baitar 2d. 9@)>3 wrote a pharmaceutical encyclopedia describing 9,)AA plants, foods, and drugs, BAA of which were his own original discoveries. A =atin translation of his work was useful to European biologists and pharmacists in the 9>th and 9:th centuries. 'B(* The Arabian physician 5bn al-1afis 29@9BU9@>>3 was another early adherent of e$perimental dissection and autopsy,'BP*'B>* who in 9@)@ discovered pulmonary circulation'B:* and coronary circulation,')A*')9* which form the basis of the circulatory system.')@*4e also described the concept of metabolism,')B* and discredited the incorrect alenic and Avicennian theories on the four humours, pulsation,'))* bones, muscles, intestines, sensory organs, bilious canals, esophagus and stomach.'BP*
'e arte venandi, by 8rederick 55, 4oly ;oman Emperor, was an influential medieval natural history te$t that e$plored birdmorphology.
0uring the 4igh !iddle Ages, a few European scholars such as 4ildegard of Bingen, Albertus !agnus and 8rederick 55 e$panded the natural history canon. The rise of European universities, though important for the development of physics and philosophy, had little impact on biological scholarship.')+*
7abinets of curiosities, such as that of6le Dorm, were centers of biological knowledge in the early modern period, bringing organisms from across the world together in one place. Before the Age of E$ploration, naturalists had little idea of the sheer scale of biological diversity.
E$tending the work of <esalius into e$periments on still living bodies 2of both humans and animals3, Dilliam 4arvey and other natural philosophers investigated the roles of blood, veins and arteries. 4arvey?s'e motu cordis in 9(@> was the beginning of the end for alenic theory, and alongside "antorio "antorio?s studies of metabolism, it served as an influential model of .uantitative approaches to physiology. '+)* 5n the early 9Pth century, the micro-world of biology was /ust beginning to open up. A few lensmakers and natural philosophers had been creating crude microscopes since the late 9(th century, and ;obert 4ooke published the seminal $icrographiabased on observations with his own compound microscope in 9((+. But it was not until Antony van =eeuwenhoek?s dramatic improvements in lensmaking beginning in the 9(PAs,ultimately producing up to @AA-fold magnification with a single lens,that scholars discovered spermato%oa, bacteria, infusoria and the sheer strangeness and diversity of microscopic life. "imilar investigations by Ran "wammerdam led to new interest in entomology and built the basic techni.ues of microscopic dissection and staining.'++*
5n $icrographia, ;obert 4ooke had applied the word cell to biological structures such as this piece of cork, but it was not until the 9:th century that scientists considered cells the universal basis of life.
As the microscopic world was e$panding, the macroscopic world was shrinking. Botanists such as Rohn ;ay worked to incorporate the flood of newly discovered organisms shipped from across the globe into a coherent ta$onomy, and a coherent theology 2natural theology3.'+(* 0ebate over another flood, the 1oachian, cataly%ed the development of paleontology# in 9((: 1icholas "teno published an essay on how the remains of living organisms could be trapped in layers of sediment and minerali%ed to produce fossils. Although "teno?s ideas about fossili%ation were well known and much debated among natural philosophers, an organic origin for all fossils would not be accepted by all naturalists until the end of the 9>th century due to philosophical and theological debate about issues such as the age of the earth and e$tinction.'+P*
5n the course of his travels, Ale$ander von 4umboldt mapped the distribution of plants across landscapes and recorded a variety of physical conditions such as pressure and temperature.
7harles 0arwin?s first sketch of an evolutionary tree from his )irst Noteboo" on %ransmutation of Species 29>BP3
See also: History of evolutionary thought The most significant evolutionary theory before 0arwin?s was that of Rean-Baptiste =amarck# based on the inheritance of ac.uired characteristics 2an inheritance mechanism that was widely accepted until the @Ath century3, it described a chain of development stretching from the lowliest microbe to humans.'(@* The British naturalist 7harles 0arwin, combining the biogeographical approach of 4umboldt, the uniformitarian geology of =yell, Thomas !althus?s writings on population growth, and his own morphological e$pertise, created a more successful evolutionary theory based on natural selection# similar evidence led Alfred ;ussel Dallace to independently reach the same conclusions.'(B* The 9>+: publication of 0arwin?s theory in *n the *rigin of Species by $eans of Natural Selection, or the Preservation of )avoured (aces in the Struggle for +ife is often considered the central event in the history of modern biology. 0arwin?s established credibility as a naturalist, the sober tone of the work, and most of all the sheer strength and volume of evidence presented, allowed *rigin to succeed where previous evolutionary works such as the anonymous ,estiges of &reation had failed. !ost scientists were convinced of evolution and common descent by the end of the 9:th century. 4owever, natural selection would not be accepted as the primary mechanism of evolution until well into the @Ath century, as most contemporary theories of heredity seemed incompatible with the inheritance of random variation. '()* Dallace, following on earlier work by de 7andolle, 4umboldt and 0arwin, made ma/or contributions to %oogeography. Because of his interest in the transmutation hypothesis, he paid particular attention to the geographical distribution of closely allied species during his field work first in "outh America and then in the !alay archipelago. Dhile in the archipelago he identified the Dallace line, which runs through the "pice 5slands dividing the fauna of the archipelago between an Asian %one and a1ew uinea/Australian %one. 4is key .uestion, as to why the fauna of islands with such similar climates should be so different, could only be answered by considering their origin. 5n 9>P( he wrote %he Geographical 'istribution of !nimals, which was the standard reference work for over half a century, and a se.uel, #sland +ife, in 9>>A that focused on island biogeography. 4e e$tended the si$-%one system developed by Philip
"clater for describing the geographical distribution of birds to animals of all kinds. 4is method of tabulating data on animal groups in geographic %ones highlighted the discontinuities# and his appreciation of evolution allowed him to propose rational e$planations, which had not been done before.'(+*'((* The scientific study of heredity grew rapidly in the wake of 0arwin?s *rigin of Species with the work of 8rancis alton and the biometricians. The origin of genetics is usually traced to the 9>(( work of the monk regor !endel, who would later be credited with the laws of inheritance. 4owever, his work was not recogni%ed as significant until B+ years afterward. 5n the meantime, a variety of theories of inheritance 2based on pangenesis, orthogenesis, or other mechanisms3 were debated and investigated vigorously. '(P* Embryology and ecology also became central biological fields, especially as linked to evolution and populari%ed in the work of Ernst 4aeckel. !ost of the 9:th century work on heredity, however, was not in the realm of natural history, but that of e$perimental physiology.
#hysiology-edit.
6ver the course of the 9:th century, the scope of physiology e$panded greatly, from a primarily medically oriented field to a wide-ranging investigation of the physical and chemical processes of life,including plants, animals, and even microorganisms in addition to man. +iving things as machines became a dominant metaphor in biological 2and social3 thinking. '(>*
5nnovative laboratory glassware and e$perimental methods developed by =ouis Pasteur and other biologists contributed to the young field of bacteriology in the late 9:th century.
;emak and ;udolf <irchow, however, by the 9>(As most biologists accepted all three tenets of what came to be known as cell theory.'(:* 7ell theory led biologists to re-envision individual organisms as interdependent assemblages of individual cells. "cientists in the rising field of cytology, armed with increasingly powerful microscopes and new staining methods, soon found that even single cells were far more comple$ than the homogeneous fluid-filled chambers described by earlier microscopists. ;obert Brown had described the nucleus in 9>B9, and by the end of the 9:th century cytologists identified many of the key cell components& chromosomes, centrosomes mitochondria, chloroplasts, and other structures made visible through staining. Between 9>P) and 9>>) Dalther 8lemming 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 /ust before the cell divided and a daughter cell was produced. !uch of the research on cell reproduction came together in August Deismann?s theory of heredity& he identified the nucleus 2in particular chromosomes3 as the hereditary material, proposed the distinction between somatic cells and germ cells 2arguing that chromosome number must be halved for germ cells, a precursor to the concept ofmeiosis3, and adopted 4ugo de <ries?s theory of pangenes. Deismannism was e$tremely influential, especially in the new field of e$perimental embryology.'PA* By the mid-9>+As the miasma theory of disease was largely superseded by the germ theory of disease, creating e$tensive interest in microorganisms and their interactions with other forms of life. By the 9>>As, bacteriology was becoming a coherent discipline, especially through the work of ;obert Qoch, who introduced methods for growing pure cultures on agar gels containing specific nutrients in Petri dishes. The long-held idea that living organisms could easily originate from nonliving matter 2spontaneous generation3 was attacked in a series of e$periments carried out by =ouis Pasteur, while debates over vitalism vs. mechanism 2a perennial issue since the time of Aristotle and the reek atomists3 continued apace. 'P9*
importance and diversity of e$perimental physiology methods, within both medicine and biology, grew dramatically over the second half of the 9:th century. The control and manipulation of life processes became a central concern, and e$periment was placed at the center of biological education.'PB*
issues, while smaller-scale independent efforts such as island biogeography and the 4ubbard Brook E$perimental 8orest helped redefine the scope of an increasingly diverse discipline. 'P>*
Thomas 4unt !organ?s illustration ofcrossing over, part of the !endelian-chromosome theory of heredity
9:AA marked the so-called rediscovery of $endel& 4ugo de <ries, 7arl 7orrens, and Erich von Tschermak independently arrived at !endel?s laws 2which were not actually present in !endel?s work3.'P:* "oon after, cytologists 2cell biologists3 proposed that chromosomes were the hereditary material. Between 9:9A and 9:9+, Thomas 4unt !organ and the C0rosophilistsC in his fly lab forged these two ideas,both controversial,into the C!endelian-chromosome theoryC of heredity.'>A* They .uantified the phenomenon of genetic linkage and postulated that genes reside on chromosomes like beads on string# they hypothesi%ed crossing over to e$plain linkage and constructed genetic maps of the fruit fly 'rosophila melanogaster, which became a widely used model organism.'>9* 4ugo de <ries tried to link the new genetics with evolution# building on his work with heredity and hybridi%ation, he proposed a theory of mutationism, which was widely accepted in the early @Ath century. =amarckism also had many adherents.0arwinism was seen as incompatible with the continuously variable traits studied by biometricians, which seemed only partially heritable. 5n the 9:@As and 9:BAs,following the acceptance of the !endelian-chromosome theory, the emergence of the discipline of population genetics, with the work of ;.A. 8isher, R.B.". 4aldane and "ewall Dright, unified the idea of evolution by natural selection with !endelian genetics, producing the modern synthesis. The inheritance of ac.uired characterswas re/ected, while mutationism gave way as genetic theories matured. '>@* 5n 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. 5n the 9:(As D.0. 4amilton and others developed game theory approaches to e$plain altruism from an evolutionary perspective through kin selection. The possible origin of
higher organisms through endosymbiosis, and contrasting approaches to molecular evolution in the gene-centered view 2which held selection as the predominant cause of evolution3 and the neutral theory 2which made genetic drift a key factor3 spawned perennial debates over the proper balance of adaptationism and contingency in evolutionary theory. '>B* 5n the 9:PAs "tephen Ray ould and 1iles Eldredge proposed the theory of punctuated e.uilibrium which holds that stasis is the most prominent feature of the fossil record, and that most evolutionary changes occur rapidly over relatively short periods of time. '>)* 5n 9:>A =uis Alvare% and Dalter Alvare% proposed the hypothesis that an impact event was responsible for the 7retaceousUPaleogene e$tinction event.'>+* Also in the early 9:>As, statistical analysis of the fossil record of marine organisms published by Rack "epkoski and 0avid !. ;aup led to a better appreciation of the importance of mass e$tinction events to the history of life on earth.'>(*
Dendell "tanley?s crystalli%ation oftobacco mosaic virus as a purenucleoprotein in 9:B+ convinced many scientists that heredity might be e$plained purely through physics and chemistry.
=ike biochemistry, the overlapping disciplines of bacteriology and virology 2later combined as microbiology3, situated between science and medicine, developed rapidly in the early @Ath century. 8Yli$ d?4erelle?s isolation of bacteriophage during Dorld Dar 5 initiated a long line of research focused on phage viruses and the bacteria they infect. ':A* The development of standard, genetically uniform organisms that could produce repeatable e$perimental results was essential for the development of molecular genetics. After early work with 'rosophila and mai%e, the adoption of simpler model systems like the bread mold Neurospora crassa made it possible to connect genetics to biochemistry, most importantly with Beadle and Tatum?s Cone gene, one en%ymeC hypothesis in 9:)9. enetics e$periments on even simpler systems liketobacco 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 2CplasmagenesC3 complicated the accepted !endelian-chromosome theory. ':9*
The Ccentral dogma of molecular biologyC 2originally a CdogmaC only in /est3 was proposed by 8rancis 7rick in 9:+>.
':@*
This is 7rick?s reconstruction of how he conceived of the central dogma at the time. The solid lines represent 2as
it seemed in 9:+>3 known modes of information transfer, and the dashed lines represent postulated ones.
6swald Avery showed in 9:)B that 01A was likely the genetic material of the chromosome, not its protein# the issue was settled decisively with the 9:+@ 4ershey-7hase e$periment,one of many contributions from the so-called phage group centered around physicist-turnedbiologist !a$ 0elbrWck. 5n 9:+B Rames 0. Datson and 8rancis 7rick, building on the work of !aurice Dilkins and ;osalind 8ranklin, suggested that the structure of 01A was a double heli$. 5n their famous paper C$olecular structure of Nucleic !cidsC, Datson and 7rick noted coyly, C5t has not escaped our notice that the specific pairing we have postulated immediately suggests a possible copying mechanism for the genetic material.C ':B* After the 9:+> !eselson-"tahl e$periment confirmed the semiconservative replicationof 01A, it was clear to most biologists that nucleic acid se.uence must somehow determine amino acid se.uence in proteins# physicist eorge amow proposed that a fi$ed genetic code connected proteins and 01A. Between 9:+B and 9:(9, there were few known biological se.uences,either 01A or protein, but an abundance of proposed code systems, a situation made even more complicated by e$panding knowledge of the intermediate role of ;1A. To actually decipher the code, it took an e$tensive series of e$periments in biochemistry and bacterial genetics, between 9:(9 and 9:(( ,most importantly the work of 1irenberg and Qhorana.':)*
molecular biology for answering the big .uestions of evolutionary causation. 0epartments and disciplines fractured as organismic biologists asserted their importance and independence& Theodosius 0ob%hansky made the famous statement that Cnothing in biology makes sense e$cept in the light of evolutionC as a response to the molecular challenge. The issue became even more critical after 9:(># !otoo Qimura?s neutral theory of molecular evolution suggested that natural selection was not the ubi.uitous cause of evolution, at least at the molecular level, and that molecular evolution might be a fundamentally different process from morphological evolution. 2;esolving this Cmolecular/morphological parado$C has been a central focus of molecular evolution research since the 9:(As.3'9A@*
7arefully engineered strains of the bacterium -scherichia coli are crucial tools in biotechnology as well as many other biological fields.
'ecombinant +*A-edit.
Biotechnology in the modern sense of genetic engineering began in the 9:PAs, with the invention of recombinant 01A techni.ues.'9A)* ;estriction en%ymes were discovered and characteri%ed in the late 9:(As, following on the heels of the isolation, then duplication, then synthesis of viral genes. Beginning with the lab of Paul Berg in 9:P@ 2aided by -co(# from 4erbert Boyer?s lab, building on work with ligase by Arthur Qornberg?s lab3, molecular biologists put these pieces together to produce the first transgenic organisms. "oon after, others began using plasmid vectors and adding genes for antibiotic resistance, greatly increasing the reach of the recombinant techni.ues.'9A+* Dary of the potential dangers 2particularly the possibility of a prolific bacteria with a viral cancercausing gene3, the scientific community as well as a wide range of scientific outsiders reacted to these developments with both enthusiasm and fearful restraint. Prominent molecular biologists
led by Berg suggested a temporary moratorium on recombinant 01A research until the dangers could be assessed and policies could be created. This moratorium was largely respected, until the participants in the 9:P+ Asilomar 7onference on ;ecombinant 01A created policy recommendations and concluded that the technology could be used safely. '9A(* 8ollowing Asilomar, new genetic engineering techni.ues and applications developed rapidly. 01A se.uencing methods improved greatly 2pioneered by 8rederick "anger and Dalter ilbert3, as did oligonucleotide synthesis and transfectiontechni.ues.'9AP* ;esearchers learned to control the e$pression of transgenes, and were soon racing,in both academic and industrial conte$ts,to create organisms capable of e$pressing human genes for the production of human hormones. 4owever, this was a more daunting task than molecular biologists had e$pected# developments between 9:PP and 9:>A showed that, due to the phenomena of split genes and splicing, higher organisms had a much more comple$ system of gene e$pression than the bacteria models of earlier studies.'9A>* The first such race, for synthesi%ing human insulin, was won by enentech. This marked the beginning of the biotech boom 2and with it, the era of gene patents3, with an unprecedented level of overlap between biology, industry, and law. '9A:*
5nside of a )>-well thermal cycler, a device used to performpolymerase chain reaction on many samples at once
By the 9:>As, protein se.uencing had already transformed methods of scientific classification of organisms 2especially cladistics3 but biologists soon began to use ;1A and 01A se.uences as characters# this e$panded the significance of molecular evolution within evolutionary biology, as the results of molecular systematics could be compared with traditional evolutionary trees based on morphology. 8ollowing the pioneering ideas of =ynn !argulis on endosymbiotic theory, which holds that some of theorganelles of eukaryotic cells originated from free living prokaryotic organisms through symbiotic relationships, even the overall division of the tree of life was revised. 5nto the 9::As, the five domains 2Plants, Animals, 8ungi, Protists, and !onerans3 became three 2the Archaea, the Bacteria, and the Eukarya3 based on 7arl Doese?s pioneering molecular systematics work with 9(" r;1A se.uencing.'99A* The development and populari%ation of the polymerase chain reaction 2P7;3 in mid-9:>As 2by Qary !ullis and others at 7etus 7orp.3 marked another watershed in the history of modern
biotechnology, greatly increasing the ease and speed of genetic analysis. '999* 7oupled with the use of e$pressed se.uence tags, P7; led to the discovery of many more genes than could be found through traditional biochemical or genetic methods and opened the possibility of se.uencing entire genomes.'99@* The unity of much of the morphogenesis of organisms from fertili%ed egg to adult began to be unraveled after the discovery of the homeobo$ genes, first in fruit flies, then in other insects and animals, including humans. These developments led to advances in the field of evolutionary developmental biology towards understanding how the various body plans of the animal phyla have evolved and how they are related to one another. '99B* The 4uman enome Pro/ect,the largest, most costly single biological study ever undertaken, began in 9:>> under the leadership of Rames 0. Datson, after preliminary work with genetically simpler model organisms such as -. coli, S. cerevisiae and &. elegans. "hotgun se.uencing and gene discovery methods pioneered by 7raig <enter,and fueled by the financial promise of gene patents with 7elera enomics, led to a publicUprivate se.uencing competition that ended in compromise with the first draft of the human 01A se.uence announced in @AAA. '99)*