Editing History of biology

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{{For|the video game|History of Biology (video game)}}
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{{Use dmy dates|date=April 2019}}
[[File:Erasmus Darwin Temple of Nature.jpg|thumb|right|180px|The frontispiece to [[Erasmus Darwin]]'s [[evolution]]-themed poem ''The Temple of Nature'' shows a goddess pulling back the veil from nature (in the person of [[Artemis]]). Allegory and metaphor have often played an important role in the history of biology.]]
{{TopicTOC-Biology}}
The '''history of biology''' traces the study of the [[life|living world]] from [[ancient]] to [[Modernity|modern]] times. Although the concept of ''[[biology]]'' as a single coherent field arose in the 19th century, the [[biological sciences]] emerged from [[history of medicine|traditions of medicine]] and [[natural history]] reaching back to [[Ayurveda]], [[ancient Egyptian medicine]] and the works of [[Aristotle]], [[Theophrastus]] and [[Galen]] in the ancient [[Greco-Roman world]]. This ancient work was further developed in the [[Middle Ages]] by [[Islamic medicine|Muslim physicians]] and scholars such as [[Avicenna]]. During the European [[Renaissance]] and early modern period, biological thought was revolutionized in [[Europe]] by a renewed interest in [[empiricism]] and the discovery of many novel organisms. Prominent in this movement were [[Vesalius]] and [[William Harvey|Harvey]], who used experimentation and careful observation in [[physiology]], and [[naturalists]] such as [[Carl Linnaeus|Linnaeus]] and [[Georges-Louis Leclerc, Comte de Buffon|Buffon]] who began to [[Scientific classification|classify the diversity of life]] and the [[fossil record]], as well as the development and behavior of organisms. [[Antonie van Leeuwenhoek]] revealed by means of [[microscopy]] 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 (although it entrenched the [[teleological argument|argument from design]]).

Over the 18th and 19th centuries, biological sciences such as [[botany]] and [[zoology]] became increasingly professional [[scientific discipline]]s. [[Lavoisier]] and other physical scientists began to connect the animate and inanimate worlds through physics and chemistry. Explorer-naturalists such as [[Alexander von Humboldt]] investigated the interaction between organisms and their environment, and the ways this relationship depends on geography—laying the foundations for [[biogeography]], [[ecology]] and [[ethology]]. Naturalists began to reject [[essentialism]] and consider the importance of [[extinction]] and the [[history of evolutionary thought|mutability of species]]. [[Cell theory]] provided a new perspective on the fundamental basis of life. These developments, as well as the results from [[embryology]] and [[paleontology]], were synthesized in [[Charles Darwin]]'s theory of [[evolution]] by [[natural selection]]. The end of the 19th century saw the fall of [[spontaneous generation]] and the rise of the [[germ theory of disease]], though the mechanism of [[biological inheritance|inheritance]] remained a mystery.

In the early 20th century, the rediscovery of [[Gregor Mendel|Mendel's]] work in botany by [[Carl Correns]] led to the rapid development of [[genetics]] applied to fruit flies by [[Thomas Hunt Morgan]] and his students, and by the 1930s the combination of [[population genetics]] and natural selection in the "[[Modern synthesis (20th century)|neo-Darwinian synthesis]]". New disciplines developed rapidly, especially after [[James D. Watson|Watson]] and [[Francis Crick|Crick]] proposed the structure of [[DNA]]. Following the establishment of the [[Central Dogma]] 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 ''[[cell biology|cellular]] and [[molecular biology]]''. By the late 20th century, new fields like [[genomics]] and [[proteomics]] were reversing this trend, with organismal biologists using molecular techniques, and molecular and cell biologists investigating the interplay between genes and the environment, as well as the genetics of natural populations of organisms.

==Prehistoric times==
{{Further|Human history|History of agriculture|History of medicine}}
[[File:Divinatory livers Louvre AO19837.jpg|thumb|left|upright=0.5|Clay models of animal livers dating between the nineteenth and eighteenth centuries BCE, found in the royal palace at [[Mari, Syria|Mari]]]]
The [[earliest humans]] must have had and passed on knowledge about [[plant]]s and [[animal]]s to increase their chances of survival. This may have included knowledge of human and animal anatomy and aspects of animal behavior (such as migration patterns). However, the first major turning point in biological knowledge came with the [[Neolithic Revolution]] about 10,000 years ago. Humans first domesticated plants for farming, then [[livestock]] animals to accompany the resulting [[Sedentary lifestyle|sedentary societies]].<ref name = "magner2002a">{{cite book | last = Magner | first = Louis N. | date = 2002 | chapter = The origins of the life sciences | title =  A History of the Life Sciences | edition = 3rd | pages = 1–40 | location = New York | publisher = CRC Press | isbn = 0824708245}}</ref>

==Earliest roots==
Between around 3000 and 1200 [[Common Era|BCE]], the [[Ancient Egypt]]ians and [[Mesopotamia]]ns made contributions to [[astronomy]], [[mathematics]], and [[medicine]],<ref name= "Lindberg1" >{{cite book | last= Lindberg | first= David C. | year = 2007 | chapter = Science before the Greeks | title= The beginnings of Western science: the European Scientific tradition in philosophical, religious, and institutional context | pages = 1–20 | edition = Second | location = Chicago, Illinois | publisher = University of Chicago Press | isbn= 978-0-226-48205-7}}</ref><ref name= "Grant2007a">{{cite book | last= Grant| first= Edward | year = 2007 | chapter = Ancient Egypt to Plato | title= A History of Natural Philosophy: From the Ancient World to the Nineteenth Century | url= https://archive.org/details/historynaturalph00gran| url-access= limited| pages = [https://archive.org/details/historynaturalph00gran/page/n16 1]–26 | edition = First | location = New York, New York | publisher = Cambridge University Press | isbn= 978-052-1-68957-1}}</ref> which later entered and shaped Greek [[natural philosophy]] of [[classical antiquity]], a period that profoundly influenced the development of what came to be known as biology.<ref name = "magner2002a"/>

===Ancient Egypt===
Over a dozen [[medical papyri]] have been preserved, most notably the [[Edwin Smith Papyrus]] (the oldest extant surgical handbook) and the [[Ebers Papyrus]] (a handbook of preparing and using materia medica for various diseases), both from around 1600 BCE.<ref name= "Lindberg1"/>

Ancient Egypt is also known for developing [[embalming]], which was used for [[Mummy|mummification]], in order to preserve human remains and forestall [[decomposition]].<ref name = "magner2002a"/>

===Mesopotamia===
{{Further|Babylonian medicine}}
The Mesopotamians seem to have had little interest in the natural world as such, preferring to study how the gods had ordered the universe. [[Animal physiology]] was studied for [[divination]], including especially the anatomy of the [[liver]], seen as an important organ in [[haruspicy]]. [[Animal behavior]] too was studied for divinatory purposes. Most information about the training and domestication of animals was probably transmitted orally, but one text dealing with the training of horses has survived.<ref name="McIntosh2005">{{cite book |last1=McIntosh |first1=Jane R. |title=Ancient Mesopotamia: New Perspectives |date=2005 |publisher=ABC-CLIO |location=Santa Barbara, California, Denver, Colorado, and Oxford, England |isbn=978-1-57607-966-9 |pages=273–276 |url=https://books.google.com/books?id=9veK7E2JwkUC&q=science+in+ancient+Mesopotamia }}</ref>

The ancient Mesopotamians had no distinction between "rational science" and [[Magic (paranormal)|magic]].<ref name="Farber1995">{{Cite book |last=Farber |first=Walter |date=1995 |title=Witchcraft, Magic, and Divination in Ancient Mesopotamia |url=https://archive.org/details/isbn_9780684192796/page/1891 |journal=Civilizations of the Ancient Near East |volume=3 |location=New York City, New York |publisher=Charles Schribner’s Sons, MacMillan Library Reference USA, Simon & Schuster MacMillan |pages=[https://archive.org/details/isbn_9780684192796/page/1891 1891–1908] |isbn=9780684192796 |access-date=12 May 2018 }}</ref><ref name="Abusch">{{cite book |last=Abusch |first=Tzvi |title=Mesopotamian Witchcraft: Towards a History and Understanding of Babylonian Witchcraft Beliefs and Literature |url=https://books.google.com/books?id=Slhv-0ewLHwC |location=Leiden, The Netherlands |publisher=Brill |year=2002 |isbn=9789004123878 |page=56}}</ref><ref name="Brown">{{cite book |last=Brown |first=Michael |date=1995 |title=Israel's Divine Healer |url=https://books.google.com/books?id=KCzmNKnLqMkC |location=Grand Rapids, Michigan |publisher=Zondervan |isbn=9780310200291 |page=42}}</ref> When a person became ill, doctors prescribed both magical formulas to be recited and medicinal treatments.<ref name="Farber1995"/><ref name="Abusch"/><ref name="Brown"/> The earliest medical prescriptions appear in [[Sumerian language|Sumerian]] during the [[Third Dynasty of Ur]] ({{circa|2112|2004 BCE}}).<ref>{{cite journal  |title=Medicine, Surgery, and Public Health in Ancient Mesopotamia  |author=R D. Biggs  |journal=Journal of Assyrian Academic Studies  |volume=19  |number=1  |year=2005  |pages=7–18}}</ref> The most extensive Babylonian medical text, however, is the ''Diagnostic Handbook'' written by the ''ummânū'', or chief scholar, [[Esagil-kin-apli]] of [[Borsippa]],<ref name="Stol-99">{{cite book |last=Heeßel |first=N. P. |date=2004 |chapter=Diagnosis, Divination, and Disease: Towards an Understanding of the ''Rationale'' Behind the Babylonian ''Diagnostic Handbook'' |chapter-url=https://books.google.com/books?id=p6rejN-iF0IC&q=Diagnostic+Handbook |title=Magic and Rationality in Ancient Near Eastern and Graeco-Roman Medicine |editor1-last=Horstmanshoff |editor1-first=H. F. J. |editor2-last=Stol |editor2-first=Marten |editor3-last=Tilburg |editor3-first=Cornelis |series=Studies in Ancient Medicine |volume=27 |location=Leiden, The Netherlands |publisher=Brill |isbn=978-90-04-13666-3 |pages=97–116 }}</ref> during the reign of the Babylonian king [[Adad-apla-iddina]] (1069 – 1046 BCE).<ref>Marten Stol (1993), ''Epilepsy in Babylonia'', p. 55, [[Brill Publishers]], {{ISBN |90-72371-63-1}}.</ref> In [[East Semitic]] cultures, the main medicinal authority was an exorcist-healer known as an ''[[Asipu|āšipu]]''.<ref name="Farber1995"/><ref name="Abusch"/><ref name="Brown"/> The profession was passed down from father to son and was held in high regard.<ref name="Farber1995"/> Of less frequent recourse was the ''asu'', a healer who treated physical symptoms using remedies composed of herbs, animal products, and minerals, as well as potions, enemas, and ointments or [[poultices]]. These physicians, who could be either male or female, also dressed wounds, set limbs, and performed simple surgeries. The ancient Mesopotamians also practiced [[prophylaxis]] and took measures to prevent the spread of disease.<ref name="McIntosh2005"/>

==Separate developments in China and India==
[[File:Huang-Quan-Xie-sheng-zhen-qin-tu.jpg|thumb|300px|''Description of rare animals'' (写生珍禽图), by Huang Quan (903–965) during the [[Song dynasty]]]]
Observations and theories regarding nature and human health, separate from [[Western culture#Scientific and technological inventions and discoveries|Western tradition]]s, had emerged independently in other civilizations such as those in [[History of China|China]] and the [[History of India|Indian subcontinent]].<ref name = "magner2002a"/> In ancient China, earlier conceptions can be found dispersed across several different disciplines, including the work of [[Chinese herbology|herbologists]], physicians, alchemists, and [[Chinese philosophy|philosophers]]. The [[Taoism|Taoist]] tradition of [[Chinese alchemy]], for example, emphasized health (with the ultimate goal being the [[elixir of life]]). The system of [[classical Chinese medicine]] usually revolved around the theory of [[yin and yang]], and the [[Wuxing (Chinese philosophy)|five phases]].<ref name = "magner2002a"/> Taoist philosophers, such as [[Zhuang Zhou|Zhuangzi]] in the 4th century BCE, also expressed ideas related to [[evolution]], such as denying the fixity of biological species and speculating that species had developed differing attributes in response to differing environments.<ref>{{cite book|last1=Needham|first1=Joseph|author-link=Joseph Needham|last2=Ronan|first2=Colin Alistair|title=The Shorter Science and Civilisation in China: An Abridgement of Joseph Needham's Original Text, Vol. 1|publisher=[[Cambridge University Press]]|year=1995|isbn=978-0-521-29286-3|page=101}}</ref>

One of the oldest organised systems of medicine is known from ancient India in the form of [[Ayurveda]], which originated around 1500 BCE from [[Atharvaveda]] (one of the four most ancient books of Indian knowledge, wisdom and culture).

The ancient Indian [[Ayurveda]] tradition independently developed the concept of three humours, resembling that of the [[Humorism|four humours]] of [[ancient Greek medicine]], though the Ayurvedic system included further complications, such as the body being composed of [[Classical element|five elements]] and seven basic [[Tissue (biology)|tissues]]. Ayurvedic writers also classified living things into four categories based on the method of birth (from the womb, eggs, heat & moisture, and seeds) and explained the conception of a [[fetus]] in detail. They also made considerable advances in the field of [[surgery]], often without the use of human [[dissection]] or animal [[vivisection]].<ref name = "magner2002a"/> One of the earliest Ayurvedic treatises was the ''[[Sushruta Samhita]]'', attributed to Sushruta in the 6th century BCE. It was also an early [[materia medica]], describing 700 medicinal plants, 64 preparations from mineral sources, and 57 preparations based on animal sources.<ref>{{Cite journal|last=Girish Dwivedi |first=Shridhar Dwivedi |year=2007 |title=History of Medicine: Sushruta – the Clinician – Teacher par Excellence |publisher=[[National Informatics Centre]] |url=http://medind.nic.in/iae/t07/i4/iaet07i4p243.pdf |access-date=8 October 2008 |url-status=dead |archive-url=https://web.archive.org/web/20081010045900/http://medind.nic.in/iae/t07/i4/iaet07i4p243.pdf |archive-date=10 October 2008 |journal=Indian J Chest Dis Allied Sci|volume=49|pages=243–244}}</ref>

==Classical antiquity==
{{Further|Ancient Greek medicine|Aristotle's biology}}
[[File:161Theophrastus 161 frontespizio.jpg|thumb|upright|left|Frontispiece to a 1644 version of the expanded and illustrated edition of ''[[Historia Plantarum (Theophrastus)|Historia Plantarum]]'', originally written by [[Theophrastus]] around 300 BCE]]

The [[Pre-Socratic philosophy|pre-Socratic philosophers]] asked many questions about life but produced little systematic knowledge of specifically biological interest—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.<ref name = "magner2002a"/>

The philosopher [[Aristotle]] was the most influential scholar of the living world from [[classical antiquity]].<ref>Lennox, J.G. 2001. ''Aristotle's Philosophy of Biology: Studies in the Origins of Life Science''. Cambridge: Cambridge University Press.</ref> Though his early work in natural philosophy was speculative, [[Aristotle's biology|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 [[Abstraction|attributes]] of [[plant]]s and [[animal]]s in the world around him, which he devoted considerable attention to [[categorization|categorizing]]. In all, Aristotle classified 540 animal species, and dissected at least 50. He believed that intellectual purposes, [[formal cause]]s, guided all natural processes.<ref>Mayr, ''The Growth of Biological Thought'', pp 84–90, 135; Mason, ''A History of the Sciences'', p 41–44</ref>

Aristotle's successor at the [[Lyceum]], [[Theophrastus]], wrote a series of books on botany, the ''[[Historia Plantarum (Theophrastus)|History of Plants]]'', 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 ''karpós'' for fruit, and ''perikárpion'' for seed vessel. [[Dioscorides]] wrote a pioneering and [[Encyclopedia|encyclopedic]] [[pharmacopoeia]], ''[[De materia medica]]'', incorporating descriptions of some 600 plants and their uses in [[Roman medicine|medicine]]. [[Pliny the Elder]], in his ''[[Natural History (Pliny)|Natural History]]'', assembled a similarly encyclopaedic account of things in nature, including accounts of many plants and animals.<ref>Mayr, ''The Growth of Biological Thought'', pp 90–91; Mason, ''A History of the Sciences'', p 46</ref> Aristotle, and nearly all Western 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]].<ref>Mayr, ''The Growth of Biological Thought'', pp 201–202; see also: Lovejoy, ''The Great Chain of Being''</ref>

A few scholars in the [[Hellenistic period]] under the [[Ptolemaic dynasty|Ptolemies]]—particularly [[Herophilos|Herophilus of Chalcedon]] and [[Erasistratus|Erasistratus of Chios]]—amended Aristotle's physiological work, even performing dissections and vivisections.<ref>Barnes, ''Hellenistic Philosophy and Science'', p 383–384</ref> [[Galen|Claudius Galen]] became the most important authority on medicine and anatomy. Though a few ancient [[atomism|atomists]] such as [[Lucretius]] challenged the [[teleology|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. [[Ernst W. Mayr]] argued that "Nothing of any real consequence happened in biology after Lucretius and Galen until the Renaissance."<ref>Mayr, ''The Growth of Biological Thought'', pp 90–94; quotation from p 91</ref> The ideas of the Greek traditions of natural history and medicine survived, but they were generally taken unquestioningly in [[Middle Ages|medieval Europe]].<ref>Annas, ''Classical Greek Philosophy'', p 252</ref>

==Middle Ages==
{{Further|Islamic medicine|Byzantine medicine|Medieval medicine of Western Europe}}
[[File:ibn al-nafis page.jpg|thumb|upright|A biomedical work by [[Ibn al-Nafis]], an early adherent of experimental dissection who discovered the [[pulmonary circulation|pulmonary]] and [[coronary circulation]]]]

The decline of the [[Roman Empire]] led to the disappearance or destruction of much knowledge, though physicians still incorporated many aspects of the Greek tradition into training and practice. In [[Byzantium]] and the [[Islamic]] world, many of the Greek works were translated into [[Arabic]] and many of the works of Aristotle were preserved.<ref name=Mayr-91-94>Mayr, ''The Growth of Biological Thought'', pp 91–94</ref>

[[File:Frederick II and eagle.jpg|thumb|upright|''[[De arte venandi]]'', by [[Frederick II, Holy Roman Emperor]], was an influential medieval natural history text that explored bird [[Morphology (biology)|morphology]].]]

During the [[High Middle Ages]], a few European scholars such as [[Hildegard of Bingen]], [[Albertus Magnus]] and [[Frederick II, Holy Roman Emperor|Frederick II]] wrote on natural history. The [[History of European research universities|rise of European universities]], though important for the development of physics and philosophy, had little impact on biological scholarship.<ref>Mayr, ''The Growth of Biological Thought'', pp 91–94: {{blockquote|"As far as biology as a whole is concerned, it was not until the late eighteenth and early nineteenth century that the universities became centers of biological research."}}</ref>

==Renaissance==
{{Further|History of anatomy|Scientific Revolution}}
The [[European Renaissance]] brought expanded interest in both empirical natural history and physiology. In 1543, [[Andreas Vesalius]] inaugurated the modern era of Western medicine with his seminal [[human anatomy]] treatise ''[[De humani corporis fabrica]]'', which was based on dissection of corpses. Vesalius was the first in a series of anatomists who gradually replaced [[scholasticism]] with [[empiricism]] in physiology and medicine, relying on first-hand experience rather than authority and abstract reasoning. Via [[herbalism]], medicine was also indirectly the source of renewed empiricism in the study of plants. [[Otto Brunfels]], [[Hieronymus Bock]] and [[Leonhart Fuchs]] wrote extensively on wild plants, the beginning of a nature-based approach to the full range of plant life.<ref>Mayr, ''The Growth of Biological Thought'', pp 94–95, 154–158</ref> [[Bestiaries]]—a genre that combines both the natural and figurative knowledge of animals—also became more sophisticated, especially with the work of [[William Turner (naturalist)|William Turner]], [[Pierre Belon]], [[Guillaume Rondelet]], [[Conrad Gessner]], and [[Ulisse Aldrovandi]].<ref>Mayr, ''The Growth of Biological Thought'', pp 166–171</ref>

Artists such as [[Albrecht Dürer]] and [[Leonardo da Vinci]], often working with naturalists, were also interested in the bodies of animals and humans, studying physiology in detail and contributing to the growth of anatomical knowledge.<ref>Magner, ''A History of the Life Sciences'', pp 80–83</ref> The traditions of [[alchemy]] and [[natural magic]], especially in the work of [[Paracelsus]], also laid claim to knowledge of the living world. Alchemists subjected organic matter to chemical analysis and experimented liberally with both biological and mineral [[pharmacology]].<ref>Magner, ''A History of the Life Sciences'', pp 90–97</ref> This was part of a larger transition in world views (the rise of the [[mechanical philosophy]]) that continued into the 17th century, as the traditional metaphor of ''nature as organism'' was replaced by the ''nature as machine'' metaphor.<ref>Merchant, ''The Death of Nature'', chapters 1, 4, and 8</ref>

==Age of Enlightenment==
{{Further|History of plant systematics}}
[[Scientific classification|Systematizing]], naming and classifying dominated natural history throughout much of the 17th and 18th centuries. [[Carl Linnaeus]] published a basic [[Taxonomy (biology)|taxonomy]] for the natural world in 1735 (variations of which have been in use ever since), and in the 1750s introduced [[Binomial nomenclature|scientific names]] for all his species.<ref>Mayr, ''The Growth of Biological Thought'', chapter 4</ref> 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—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 [[Charles Darwin|Darwin]].<ref>Mayr, ''The Growth of Biological Thought'', chapter 7</ref>

The discovery and description of new species and the [[collecting|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.<ref>See Raby, ''Bright Paradise''</ref>

[[File:1655 - Frontispiece of Museum Wormiani Historia.jpg|thumb|left|[[Cabinet of curiosities|Cabinets of curiosities]], such as that of [[Ole Worm]], were centers of biological knowledge in the early modern period, bringing organisms from across the world together in one place. Before the [[Age of Exploration]], naturalists had little idea of the sheer scale of biological diversity.]]

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.<ref>Magner, ''A History of the Life Sciences'', pp 103–113</ref>

In the early 17th century, the micro-world of biology was just beginning to open up. A few lensmakers and natural philosophers had been creating crude [[microscope]]s 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 [[Antonie van Leeuwenhoek]]'s dramatic improvements in lensmaking beginning in the 1670s—ultimately producing up to 200-fold magnification with a single lens—that scholars discovered [[spermatozoa]], [[bacteria]], [[infusoria]] and the sheer strangeness and diversity of microscopic life. Similar investigations by [[Jan Swammerdam]] led to a new interest in [[entomology]] and built the basic techniques of microscopic dissection and [[staining]].<ref>Magner, ''A History of the Life Sciences'', pp 133–144</ref>
[[File:Cork Micrographia Hooke.png|thumb|right|upright|In ''[[Micrographia]]'', Robert Hooke had applied the word ''cell'' to biological structures such as this piece of [[Cork cambium|cork]], but it was not until the 19th century that scientists considered cells the universal basis of life.]]

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]]).<ref>Mayr, ''The Growth of Biological Thought'', pp 162–166</ref> Debate over another flood, the [[Noachian flood|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 [[fossil]]s. 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]].<ref>Rudwick, ''The Meaning of Fossils'', pp 41–93</ref>

==19th century: the emergence of biological disciplines==<!--section title linked from [[On the Origin of Species]], please don't change-->
Up through the 19th century, the scope of biology was largely divided between medicine, which investigated questions of form and function (i.e., physiology), and natural history, which was concerned with the diversity of life and interactions among different forms of life and between life and non-life. By 1900, much of these domains overlapped, while natural history (and its counterpart [[natural philosophy]]) had largely given way to more specialized scientific disciplines—[[cell biology|cytology]], [[bacteriology]], [[morphology (biology)|morphology]], [[embryology]], [[geography]], and [[geology]].

[[File:Geographie der Pflanzen in den Tropen-Ländern.jpg|thumb|left|In the course of his travels, [[Alexander von Humboldt]] mapped the distribution of plants across landscapes and recorded a variety of physical conditions such as pressure and temperature.]]

===Use of the term ''biology''===
The term ''biology'' in its modern sense appears to have been introduced independently by [[Thomas Beddoes]] (in 1799),<ref>{{cite web|title=biology, ''n''.|work=[[Oxford English Dictionary]] online version|publisher=Oxford University Press|date=September 2011|url=http://www.oed.com/view/Entry/19228?redirectedFrom=Biology#eid|access-date=1 November 2011}} {{OEDsub}}</ref> [[Karl Friedrich Burdach]] (in 1800), [[Gottfried Reinhold Treviranus]] (''Biologie oder Philosophie der lebenden Natur'', 1802) and [[Jean-Baptiste Lamarck]] (''Hydrogéologie'', 1802).<ref>Junker ''Geschichte der Biologie'', p8.</ref><ref>Coleman, ''Biology in the Nineteenth Century'', pp 1–2.</ref> The word itself appears in the title of Volume 3 of [[Michael Christoph Hanow]]'s ''Philosophiae naturalis sive physicae dogmaticae: Geologia, biologia, phytologia generalis et dendrologia'', published in 1766. The term ''biology'' derives from the [[Greek language|Greek]] [[wikt:βίος|βίος]] (''bíos'') 'life', and [[wikt:λογία|λογία]] (''logia'') 'branch of study'.

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 Middle Ages through the Renaissance, the unifying framework of natural history was the ''scala naturae'' or [[Great Chain of Being]]. ''[[Natural philosophy]]'' and ''[[natural theology]]'' encompassed the conceptual and metaphysical basis of plant and animal life, dealing with problems of why organisms exist and behave the way they do, though these subjects also included what is now [[geology]], [[physics]], [[chemistry]], and [[astronomy]]. Physiology and (botanical) pharmacology were the province of medicine. ''Botany'', ''Zoology'', and (in the case of fossils) ''Geology'' replaced ''natural history'' and ''natural philosophy'' in the 18th and 19th centuries before ''biology'' was widely adopted.<ref>Mayr, ''The Growth of Biological Thought'', pp36–37</ref><ref>Coleman, ''Biology in the Nineteenth Century'', pp 1–3.</ref> To this day, "botany" and "zoology" are widely used, although they have been joined by other sub-disciplines of biology.

===Natural history and natural philosophy===
{{Further|Humboldtian science}}
Widespread travel by naturalists in the early-to-mid-19th century resulted in a wealth of new information about the diversity and distribution of living organisms. Of particular importance was the work of [[Alexander von Humboldt]], which analyzed the relationship between organisms and their environment (i.e., the domain of [[natural history]]) using the quantitative approaches of [[natural philosophy]] (i.e., [[physics]] and [[chemistry]]). Humboldt's work laid the foundations of [[biogeography]] and inspired several generations of scientists.<ref>Bowler, ''The Earth Encompassed'', pp 204–211</ref>

====Geology and paleontology====
{{Further|History of geology|History of paleontology}}
The emerging discipline of geology also brought natural history and natural philosophy closer together; the establishment of the [[stratigraphy|stratigraphic column]] linked the spatial distribution of organisms to their temporal distribution, a key precursor to concepts of evolution. [[Georges Cuvier]] and others made great strides in [[comparative anatomy]] and [[paleontology]] in the late 1790s and early 19th century. In a series of lectures and papers that made detailed comparisons between living mammals and [[fossil]] remains Cuvier was able to establish that the fossils were remains of species that had become [[extinct]]—rather than being remains of species still alive elsewhere in the world, as had been widely believed.<ref>Rudwick, ''The Meaning of Fossils'', pp 112–113</ref> Fossils discovered and described by [[Gideon Mantell]], [[William Buckland]], [[Mary Anning]], and [[Richard Owen]] among others helped establish that there had been an 'age of reptiles' that had preceded even the prehistoric mammals. These discoveries captured the public imagination and focused attention on the history of life on earth.<ref>Bowler, ''The Earth Encompassed'', pp 211–220</ref> Most of these geologists held to [[catastrophism]], but [[Charles Lyell]]'s influential ''Principles of Geology'' (1830) popularised [[James Hutton|Hutton's]] [[uniformitarianism (science)|uniformitarianism]], a theory that explained the geological past and present on equal terms.<ref>Bowler, ''The Earth Encompassed'', pp 237–247</ref>

====Evolution and biogeography====
{{Further|History of evolutionary thought|History of speciation}}

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.<ref>Mayr, ''The Growth of Biological Thought'', pp 343–357</ref> 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 led [[Alfred Russel Wallace]] to independently reach the same conclusions.<ref>Mayr, ''The Growth of Biological Thought'', chapter 10: "Darwin's evidence for evolution and common descent"; and chapter 11: "The causation of evolution: natural selection"; Larson, ''Evolution'', chapter 3</ref>

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.<ref>Larson, ''Evolution'', chapter 5: "Ascent of Evolutionism"; see also: Bowler, ''The Eclipse of Darwinism''; Secord, ''Victorian Sensation''</ref>
[[File:Darwins first tree.jpg|left|thumb|[[Charles Darwin]]'s first sketch of an evolutionary tree from his ''First Notebook on Transmutation of Species'' (1837)]]

Wallace, following on earlier work by [[A.P. de Candolle|de Candolle]], [[Alexander von Humboldt|Humboldt]] and Darwin, made major contributions to [[zoogeography]]. 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 [[South America]] and then in the [[Malay Archipelago]]. While in the archipelago he identified the [[Wallace line]], which runs through the [[Maluku Islands|Spice Islands]] dividing the fauna of the archipelago between an Asian zone and a [[New Guinea]]/Australian zone. His key question, as to why the fauna of islands with such similar climates should be so different, could only be answered by considering their origin. In 1876 he wrote ''The Geographical Distribution of Animals'', which was the standard reference work for over half a century, and a sequel, ''Island Life'', in 1880 that focused on island biogeography. He extended the six-zone system developed by [[Philip Sclater]] for describing the geographical distribution of birds to animals of all kinds. His method of tabulating data on animal groups in geographic zones highlighted the discontinuities; and his appreciation of evolution allowed him to propose rational explanations, which had not been done before.<ref>Larson, ''Evolution'', pp 72–73, 116–117; see also: Browne, ''The Secular Ark''.</ref><ref>Bowler ''Evolution: The History of an Idea'' p. 174</ref>

[[File:Gregor Mendel 2.jpg|thumb|[[Gregor Mendel]], "father of modern genetics"<ref>{{cite web | url=https://www.biography.com/scientists/gregor-mendel | title=Gregor Mendel - Life, Experiments & Facts | date=21 May 2021 }}</ref>]]
The scientific study of [[heredity]] grew rapidly in the wake of Darwin's ''Origin of Species'' with the work of [[Francis Galton]] and the [[biometry|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.<ref>Mayr, ''The Growth of Biological Thought'', pp 693–710</ref> [[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.

===Physiology===
Over the course of the 19th century, the scope of physiology expanded 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. ''Living things as machines'' became a dominant metaphor in biological (and social) thinking.<ref>Coleman, ''Biology in the Nineteenth Century'', chapter 6; on the machine metaphor, see also: Rabinbach, ''The Human Motor''</ref>

[[File:Albert Edelfelt - Louis Pasteur - 1885.jpg|thumb|Innovative [[laboratory glassware]] and experimental methods developed by [[Louis Pasteur]] and other biologists contributed to the young field of [[bacteriology]] in the late 19th century.]]
[[File:Statue of Robert Koch in Berlin.jpg|thumb|Statue of [[Robert Koch]] in Berlin. Koch directly provided proof for the [[germ theory of diseases]], therefore creating the scientific basis of [[public health]],<ref name=":16">{{Cite journal|last=Lakhtakia|first=Ritu|date=2014|title=The Legacy of Robert Koch: Surmise, search, substantiate|journal=Sultan Qaboos University Medical Journal|volume=14|issue=1|pages=e37–41|doi=10.12816/0003334|pmc=3916274|pmid=24516751}}</ref> saving millions of lives.<ref>https://history.info/on-this-day/1843-robert-koch-man-saved-millions-lives/ {{Bare URL inline|date=August 2024}}</ref> For his life's work Koch is seen as one of the founders of modern medicine.<ref>https://www.facebook.com/watch/?v=245261433654285 {{Bare URL inline|date=August 2024}}</ref><ref>{{cite web | url=https://www.youtube.com/watch?v=XCVnOb6VXmg | title=Louis Pasteur vs Robert Koch: The History of Germ Theory | website=[[YouTube]] | date=26 May 2023 }}</ref>]]

====Cell theory, embryology and germ theory====
Advances in [[microscopy]] also had a profound impact on biological thinking. In the early 19th century, a number of biologists pointed to the central importance of the [[cell (biology)|cell]]. In 1838 and 1839, [[Matthias Jakob Schleiden|Schleiden]] and [[Theodor Schwann|Schwann]] began promoting 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]].<ref>Sapp, ''Genesis'', chapter 7; Coleman, ''Biology in the Nineteenth Century'', chapters 2</ref>

Cell theory led biologists to re-envision individual organisms as interdependent assemblages of individual cells. Scientists in the rising field of [[cell biology|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 described by earlier microscopists. [[Robert Brown (Scottish botanist from Montrose)|Robert Brown]] had described the [[Cell nucleus|nucleus]] in 1831, and by the end of the 19th century cytologists identified many of the key cell components: [[chromosome]]s, [[centrosome]]s [[mitochondria]], [[chloroplast]]s, and other structures made visible through staining. Between 1874 and 1884 [[Walther Flemming]] described the discrete stages of mitosis, showing that they were not [[Artifact (observational)|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 chromosomes) as the hereditary material, proposed the distinction between [[somatic cell]]s and [[germ cell]]s (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 [[pangene]]s. Weismannism was extremely influential, especially in the new field of experimental [[embryology]].<ref>Sapp, ''Genesis'', chapter 8; Coleman, ''Biology in the Nineteenth Century'', chapter 3</ref>

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 plate|agar gels]] containing specific nutrients in [[Petri dish]]es. 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 (philosophy)|mechanism]] (a perennial issue since the time of Aristotle and the Greek atomists) continued apace.<ref>Magner, ''A History of the Life Sciences'', pp 254–276</ref>

====Rise of organic chemistry and experimental physiology====
In chemistry, one central issue was the distinction between organic and inorganic substances, especially in the context of organic transformations such as [[Fermentation (biochemistry)|fermentation]] and [[putrefaction]]. Since Aristotle these had been considered essentially biological (''[[vitalism|vital]]'') processes. However, [[Friedrich Wöhler]], [[Justus Liebig]] and other pioneers of the rising field of [[organic chemistry]]—building on the work of Lavoisier—showed that the organic world could often be analyzed by physical and chemical methods. In 1828 Wöhler showed that the organic substance [[urea]] could be created by chemical means that do not involve life, providing a powerful challenge to [[vitalism]]. Cell extracts ("ferments") that could effect chemical transformations were discovered, beginning with [[diastase]] in 1833. By the end of the 19th century the concept of [[enzymes]] was well established, though equations of [[chemical kinetics]] would not be applied to enzymatic reactions until the early 20th century.<ref>Fruton, ''Proteins, Enzymes, Genes'', chapter 4; Coleman, ''Biology in the Nineteenth Century'', chapter 6</ref>

Physiologists such as [[Claude Bernard]] explored (through vivisection and other experimental methods) the chemical and physical functions of living bodies to an unprecedented degree, laying the groundwork for [[endocrinology]] (a field that developed quickly after the discovery of the first [[hormone]], [[secretin]], in 1902), [[biomechanics]], and the study of [[nutrition]] and [[digestion]]. The importance and diversity of experimental physiology methods, within both medicine and biology, grew dramatically over the second half of the 19th century. The control and manipulation of life processes became a central concern, and experiment was placed at the center of biological education.<ref>Rothman and Rothman, ''The Pursuit of Perfection'', chapter 1; Coleman, ''Biology in the Nineteenth Century'', chapter 7</ref>

==Twentieth century biological sciences==
[[File:Embryonic development of a salamander, filmed in the 1920s.ogv|thumb|Embryonic development of a salamander, filmed in the 1920s]]
At the beginning of the 20th century, biological research was largely a professional endeavour. Most work was still done in the [[natural history]] mode, which emphasized morphological and phylogenetic analysis over experiment-based causal explanations. However, anti-[[vitalism|vitalist]] experimental physiologists and embryologists, especially in Europe, were increasingly influential. The tremendous success of experimental approaches to development, heredity, and metabolism in the 1900s and 1910s demonstrated the power of experimentation in biology. In the following decades, experimental work replaced natural history as the dominant mode of research.<ref>See: Coleman, ''Biology in the Nineteenth Century''; Kohler, ''Landscapes and Labscapes''; Allen, ''Life Science in the Twentieth Century''; Agar, ''Science in the Twentieth Century and Beyond''</ref>

===Ecology and environmental science===
{{Further|History of ecology}}

In the early 20th century, naturalists were faced with increasing pressure to add rigor and preferably experimentation to their methods, as the newly prominent laboratory-based biological disciplines had done. [[Ecology]] had emerged as a combination of biogeography with the [[biogeochemical cycle]] concept pioneered by chemists; field biologists developed quantitative methods such as the [[quadrat]] and adapted laboratory instruments and cameras for the field to further set their work apart from traditional natural history. Zoologists and botanists did what they could to mitigate the unpredictability of the living world, performing laboratory experiments and studying semi-controlled natural environments such as gardens; new institutions like the [[Carnegie Station for Experimental Evolution]] and the [[Marine Biological Laboratory]] provided more controlled environments for studying organisms through their entire life cycles.<ref>Kohler, ''Landscapes and Labscapes'', chapters 2, 3, 4</ref>

The [[ecological succession]] concept, pioneered in the 1900s and 1910s by [[Henry Chandler Cowles]] and [[Frederic Clements]], was important in early plant ecology.<ref>Agar, ''Science in the Twentieth Century and Beyond'', p. 145</ref> [[Alfred Lotka]]'s [[predator-prey equations]], [[G. Evelyn Hutchinson]]'s studies of the biogeography and biogeochemical structure of lakes and rivers ([[limnology]]) and [[Charles Sutherland Elton|Charles Elton's]] studies of animal [[food chain]]s were pioneers among the succession of quantitative methods that colonized the developing ecological specialties. Ecology became an independent discipline in the 1940s and 1950s after [[Eugene P. Odum]] synthesized many of the concepts of [[ecosystem ecology]], placing relationships between groups of organisms (especially material and energy relationships) at the center of the field.<ref>Hagen, ''An Entangled Bank'', chapters 2–5</ref>

In the 1960s, as evolutionary theorists explored the possibility of multiple [[units of selection]], ecologists turned to evolutionary approaches. In [[population ecology]], debate over [[group selection]] was brief but vigorous; by 1970, most biologists agreed that natural selection was rarely effective above the level of individual organisms. The evolution of ecosystems, however, became a lasting research focus. Ecology expanded rapidly with the rise of the environmental movement; the [[International Biological Program]] attempted to apply the methods of [[big science]] (which had been so successful in the physical sciences) to ecosystem ecology and pressing environmental issues, while smaller-scale independent efforts such as [[island biogeography]] and the [[Hubbard Brook Experimental Forest]] helped redefine the scope of an increasingly diverse discipline.<ref>Hagen, ''An Entangled Bank'', chapters 8–9</ref>

===Classical genetics, the modern synthesis, and evolutionary theory===
{{Further|History of genetics|History of model organisms|Modern synthesis (20th century)}}

[[File:Morgan crossover 1.jpg|thumb|right|[[Thomas Hunt Morgan]]'s illustration of [[Chromosomal crossover|crossing over]], part of the Mendelian-chromosome theory of heredity]]
1900 marked the so-called ''rediscovery of Mendel'' by [[Carl Correns]], who arrived at [[Mendel's laws]] (which were not actually present in Mendel's work).<ref>Randy Moore, "[http://papa.indstate.edu/amcbt/volume_27/v27-2p13-24.pdf The 'Rediscovery' of Mendel's Work] {{webarchive|url=https://web.archive.org/web/20120401145158/http://papa.indstate.edu/amcbt/volume_27/v27-2p13-24.pdf |date=2012-04-01 }}", ''Bioscene'', Volume 27(2) pp. 13–24, May 2001.</ref> Soon after, cytologists (cell biologists) proposed that [[chromosome]]s were the hereditary material. This was taken up by [[Carl Correns]] and others between 1910 and 1915 as the "Mendelian-chromosome theory" of heredity.  [[Thomas Hunt Morgan]] and the "[[Drosophilists]]" in his fly lab applied this to a new model organism.<ref>T. H. Morgan, A. H. Sturtevant, H. J. Muller, C. B. Bridges (1915) [http://www.esp.org/books/morgan/mechanism/facsimile/title3.html ''The Mechanism of Mendelian Heredity''] Henry Holt and Company.</ref> They hypothesized [[Chromosomal crossover|crossing over]] to explain linkage and constructed [[genetic map]]s of the fruit fly ''[[Drosophila melanogaster]]'', which became a widely used [[model organism]].<ref>Garland Allen, ''Thomas Hunt Morgan: The Man and His Science'' (1978), chapter 5; see also: Kohler, ''Lords of the Fly'' and Sturtevant, ''A History of Genetics''</ref>

Hugo de Vries tried to link the new genetics with evolution; building on his work with heredity and [[Hybrid (biology)|hybridization]], he proposed a theory of [[mutationism]], which was widely accepted in the early 20th century. [[Lamarckism]], or the theory of inheritance of acquired characteristics also had many adherents. [[Darwinism]] was seen as incompatible with the continuously variable traits studied by [[biometry|biometricians]], which seemed only partially heritable. In the 1920s and 1930s—following the acceptance of the Mendelian-chromosome theory— 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 inheritance|Mendelian genetics]], producing the [[Modern synthesis (20th century)|modern synthesis]]. The [[inheritance of acquired characters]] was rejected, while mutationism gave way as genetic theories matured.<ref>Smocovitis, ''Unifying Biology'', chapter 5; see also: Mayr and Provine (eds.), ''The Evolutionary Synthesis''</ref>

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 of evolution|gene-centered view]] (which held selection as the predominant cause of evolution) and the [[neutral theory of molecular evolution|neutral theory]] (which made [[genetic drift]] a key factor) spawned perennial debates over the proper balance of [[adaptationism]] and contingency in evolutionary theory.<ref>Gould, ''The Structure of Evolutionary Theory'', chapter 8; Larson, ''Evolution'', chapter 12</ref>

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.<ref>Larson, ''Evolution'', pp 271–283</ref> In 1980 [[Luis Walter Alvarez|Luis Alvarez]] and [[Walter Alvarez]] proposed the hypothesis that an [[impact event]] was responsible for the [[Cretaceous–Paleogene extinction event]].<ref>Zimmer, ''Evolution'', pp 188–195</ref> Also in the early 1980s, statistical analysis of the fossil record of marine organisms published by [[Jack Sepkoski]] and [[David M. Raup]] led to a better appreciation of the importance of [[mass extinction events]] to the history of life on earth.<ref>Zimmer, ''Evolution'', pp 169–172</ref>

===Biochemistry, microbiology, and molecular biology===
{{Further|History of biochemistry|History of molecular biology}}
{{multiple image
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By the end of the 19th century all of the major pathways of [[drug metabolism]] had been discovered, along with the outlines of protein and fatty acid metabolism and urea synthesis.<ref>Caldwell, "Drug metabolism and pharmacogenetics"; Fruton, ''Proteins, Enzymes, Genes'', chapter 7</ref> In the early decades of the 20th century, the minor components of foods in human nutrition, the [[vitamins]], began to be isolated and synthesized. Improved laboratory techniques such as [[chromatography]] and [[electrophoresis]] led to rapid advances in physiological chemistry, which—as ''biochemistry''—began to achieve independence from its medical origins. In the 1920s and 1930s, biochemists—led by [[Hans Adolf Krebs|Hans Krebs]] and [[Carl Ferdinand Cori|Carl]] and [[Gerty Cori]]—began to work out many of the central [[metabolic pathways]] of life: the [[citric acid cycle]], [[glycogenesis]] and [[glycolysis]], and the synthesis of [[steroid]]s and [[porphyrin]]s. Between the 1930s and 1950s, [[Fritz Lipmann]] and others established the role of [[Adenosine triphosphate|ATP]] as the universal carrier of energy in the cell, and [[mitochondria]] as the powerhouse of the cell. Such traditionally biochemical work continued to be very actively pursued throughout the 20th century and into the 21st.<ref>Fruton, ''Proteins, Enzymes, Genes'', chapters 6 and 7</ref>

====Origins of molecular biology====
Following the rise of classical genetics, many biologists—including a new wave of physical scientists in biology—pursued the question of the gene and its physical nature. [[Warren Weaver]]—head of the science division of the [[Rockefeller Foundation]]—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.<ref>Morange, ''A History of Molecular Biology'', chapter 8; Kay, ''The Molecular Vision of Life'', Introduction, Interlude I, and Interlude II</ref>

[[File:TMV virus under magnification.jpg|thumb|left|[[Wendell Meredith Stanley|Wendell Stanley]]'s crystallization of [[tobacco mosaic virus]] as a pure [[nucleoprotein]] in 1935 convinced many scientists that heredity might be explained purely through physics and chemistry.]]

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 on phage viruses and the bacteria they infect.<ref>See: Summers, ''Félix d'Herelle and the Origins of Molecular Biology''</ref>

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 [[Scientific modelling|model system]]s like the bread mold ''[[Neurospora crassa]]'' made it possible to connect genetics to biochemistry, most importantly with [[George Wells Beadle|Beadle]] and [[Edward Lawrie Tatum|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 microscope|electron microscopy]] and [[ultracentrifuge|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.<ref>Creager, ''The Life of a Virus'', chapters 3 and 6; Morange, ''A History of Molecular Biology'', chapter 2</ref>

[[File:Crick's 1958 central dogma.svg|thumb|The "[[central dogma of molecular biology]]" (originally a "dogma" only in jest) was proposed by Francis Crick in 1958.<ref>{{Cite journal| first1 = F. | title = Central Dogma of Molecular Biology | journal = Nature | volume = 227| last1 = Crick| issue = 5258 | pages = 561–563 | year = 1970| pmid = 4913914 | doi = 10.1038/227561a0|bibcode = 1970Natur.227..561C | s2cid = 4164029 }}</ref> This is Crick's reconstruction of how he conceived of the central dogma at the time. The solid lines represent (as it seemed in 1958) known modes of information transfer, and the dashed lines represent postulated ones.]]

[[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]]—one of many contributions from the so-called [[phage group]] centered around physicist-turned-biologist [[Max Delbrück]]. In 1953 [[James 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."<ref>Watson, James D. and Francis Crick. "[http://www.nature.com/nature/dna50/watsoncrick.pdf Molecular structure of Nucleic Acids: A Structure for Deoxyribose Nucleic Acid]", ''[[Nature (journal)|Nature]]'', vol. 171, no. 4356, pp 737–738</ref> 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 [[Peptide sequence|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—either DNA or protein—but an abundance of proposed code systems, a situation made even more complicated by expanding knowledge of the intermediate role of [[RNA]].  
In 1961, it was demonstrated that when a [[gene]] encodes a [[protein]], three sequential bases of a gene’s [[DNA]] specify each successive amino acid of the protein.<ref>Crick FH, Barnett L, Brenner S, Watts-Tobin RJ  (December 1961). "General nature of the genetic code for proteins". Nature. 192 (4809): 1227–32. Bibcode:1961Natur.192.1227C. doi:10.1038/1921227a0. PMID 13882203. S2CID 4276146</ref>  Thus the [[genetic code]] is a triplet code, where each triplet (called a codon) specifies a particular amino acid.  Furthermore, it was shown that the codons do not overlap with each other in the DNA sequence encoding a protein, and that each sequence is read from a fixed starting point.  
To actually decipher the code, it took an extensive series of experiments in biochemistry and bacterial genetics, between 1961 and 1966—most importantly the work of [[Marshall Warren Nirenberg|Nirenberg]] and [[Har Gobind Khorana|Khorana]].<ref>Morange, ''A History of Molecular Biology'', chapters 3, 4, 11, and 12; Fruton, ''Proteins, Enzymes, Genes'', chapter 8; on the Meselson-Stahl experiment, see: Holmes, ''Meselson, Stahl, and the Replication of DNA''</ref>
During 1962-1964, numerous conditional lethal mutants of a bacterial virus were isolated.<ref>{{cite journal | last=Epstein | first=R. H. | last2=Bolle | first2=A. | last3=Steinberg | first3=C. M. | last4=Kellenberger | first4=E. | last5=Boy de la Tour | first5=E. | last6=Chevalley | first6=R. | last7=Edgar | first7=R. S. | last8=Susman | first8=M. | last9=Denhardt | first9=G. H. | last10=Lielausis | first10=A. |display-authors=3| title=Physiological Studies of Conditional Lethal Mutants of Bacteriophage T4D | journal=Cold Spring Harbor Symposia on Quantitative Biology | volume=28 | issue=0 | date=1963-01-01 | issn=0091-7451 | doi=10.1101/SQB.1963.028.01.053 | pages=375–394}}</ref> These mutants were used in several different labs to  advance fundamental understanding of the functions and interactions of the proteins employed in the machinery of [[DNA replication]], [[DNA repair]],  [[genetic recombination|DNA recombination]], and in the assembly of molecular structures.

====Expansion of molecular biology====
In addition to the Division of Biology at [[Caltech]], the [[Laboratory of Molecular Biology]] (and its precursors) at [[Cambridge University|Cambridge]], and a handful of other institutions, the [[Pasteur Institute]] became a major center for molecular biology research in the late 1950s.<ref>On Caltech molecular biology, see Kay, ''The Molecular Vision of Life'', chapters 4–8; on the Cambridge lab, see de Chadarevian, ''Designs for Life''; on comparisons with the Pasteur Institute, see Creager, "Building Biology across the Atlantic"</ref> 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 [[History of computing hardware|digital computing]] (benefiting both directly and indirectly from the [[military funding of science]]). A number of biochemists led by [[Frederick Sanger]] later joined the Cambridge lab, bringing together the study of [[macromolecule|macromolecular]] structure and function.<ref>de Chadarevian, ''Designs for Life'', chapters 4 and 7</ref> At the Pasteur Institute, [[François Jacob]] and [[Jacques Monod]] followed the 1959 [[Arthur Pardee#The PaJaMo experiment|PaJaMo experiment]] with a series of publications regarding the [[lac operon|''lac'']] [[operon]] that established the concept of [[gene regulation]] and identified what came to be known as [[messenger RNA]].<ref>{{cite journal |author=Pardee A |title=PaJaMas in Paris |journal=Trends Genet. |volume=18 |issue=11 |pages=585–7 |year=2002 |pmid=12414189 |doi=10.1016/S0168-9525(02)02780-4}}</ref> By the mid-1960s, the intellectual core of molecular biology—a model for the molecular basis of metabolism and reproduction— was largely complete.<ref>Morange, ''A History of Molecular Biology'', chapter 14</ref>

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.<ref>Wilson, ''Naturalist'', chapter 12; Morange, ''A History of Molecular Biology'', chapter 15</ref> Molecularization was particularly important in [[genetics]], [[immunology]], [[embryology]], and [[neurobiology]], while the idea that life is controlled by a "[[genetic program]]"—a metaphor Jacob and Monod introduced from the emerging fields of [[cybernetics]] and [[computer science]]—became an influential perspective throughout biology.<ref>Morange, ''A History of Molecular Biology'', chapter 15; Keller, ''The Century of the Gene'', chapter 5</ref> 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.<ref>Morange, ''A History of Molecular Biology'', pp 126–132, 213–214</ref>

Resistance to the growing influence of 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 that [[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 [[Morphology (biology)|morphological]] evolution. (Resolving this "molecular/morphological paradox" has been a central focus of molecular evolution research since the 1960s.)<ref>Dietrich, "Paradox and Persuasion", pp 100–111</ref>

===Biotechnology, genetic engineering, and genomics===
{{Further|History of biotechnology}}

[[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, [[Industrial fermentation|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]]—as well as a wide range of [[Hybrid (biology)|hybrid]] high-yield crops and agricultural technologies, the basis for the [[Green Revolution]].<ref>Bud, ''The Uses of Life'', chapters 2 and 6</ref>

[[File:E coli at 10000x, original.jpg|thumb|left|Carefully engineered [[Strain (biology)|strains]] of the bacterium ''[[Escherichia coli]]'' are crucial tools in biotechnology as well as many other biological fields.]]

====Recombinant DNA====
Biotechnology in the modern sense of [[genetic engineering]] began in the 1970s, with the invention of [[recombinant DNA]] techniques.<ref>Agar, ''Science in the Twentieth Century and Beyond'', p. 436</ref> [[Restriction enzyme]]s 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]] [[Vector (molecular biology)|vectors]] and adding genes for [[antibiotic resistance]], greatly increasing the reach of the recombinant techniques.<ref>Morange, ''A History of Molecular Biology'', chapters 15 and 16</ref>

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.<ref>Bud, ''The Uses of Life'', chapter 8; Gottweis, ''Governing Molecules'', chapter 3; Morange, ''A History of Molecular Biology'', chapter 16</ref>

Following Asilomar, new genetic engineering techniques and applications developed rapidly. [[DNA sequencing]] methods improved greatly (pioneered by [[Frederick Sanger]] and [[Walter Gilbert]]), as did [[oligonucleotide]] synthesis and [[transfection]] techniques.<ref>Morange, ''A History of Molecular Biology'', chapter 16</ref> Researchers learned to control the expression of [[transgene]]s, and were soon racing—in both academic and industrial contexts—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 (genetics)|splicing]], higher organisms had a much more complex system of [[gene expression]] than the bacteria models of earlier studies.<ref>Morange, ''A History of Molecular Biology'', chapter 17</ref> 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 patent]]s), with an unprecedented level of overlap between biology, industry, and law.<ref>Krimsky, ''Biotechnics and Society'', chapter 2; on the race for insulin, see: Hall, ''Invisible Frontiers''; see also: Thackray (ed.), ''Private Science''</ref>

====Molecular systematics and genomics====
{{Further|History of molecular evolution}}
[[File:Cycler.jpg|thumb|upright|Inside of a 48-well [[thermal cycler]], a device used to perform [[polymerase chain reaction]] on many samples at once]]

By the 1980s, protein sequencing had already transformed methods of [[scientific classification]] of organisms (especially [[cladistics]]) but biologists soon began to use RNA and DNA sequences as [[Trait (biology)|characters]]; this expanded the significance of [[molecular evolution]] within evolutionary biology, as the results of [[molecular systematics]] could be compared with traditional evolutionary trees based on [[morphology (biology)|morphology]]. Following the pioneering ideas of [[Lynn Margulis]] on [[endosymbiotic theory]], which holds that some of the [[organelles]] of [[eukaryotic]] cells originated from free living [[prokaryotic]] organisms through [[symbiotic]] relationships, even the overall division of the tree of life was revised. Into the 1990s, the five domains (Plants, Animals, Fungi, Protists, and Monerans) became three (the [[Archaea]], the [[Bacteria]], and the [[Eukarya]]) based on [[Carl Woese]]'s pioneering [[molecular systematics]] work with [[16S ribosomal RNA|16S rRNA]] sequencing.<ref>Sapp, ''Genesis'', chapters 18 and 19</ref>

The development and popularization of the [[polymerase chain reaction]] (PCR) in mid-1980s (by [[Kary Mullis]] and others at [[Cetus Corp.]]) marked another watershed in the history of modern biotechnology, greatly increasing the ease and speed of genetic analysis.<ref>Agar, ''Science in the Twentieth Century and Beyond'', p. 456</ref> Coupled with the use of [[expressed sequence tags]], PCR led to the discovery of many more genes than could be found through traditional biochemical or genetic methods and opened the possibility of sequencing entire genomes.<ref>Morange, ''A History of Molecular Biology'', chapter 20; see also: Rabinow, ''Making PCR''</ref>

The unity of much of the [[morphogenesis]] of organisms from fertilized egg to adult began to be unraveled after the discovery of the [[homeobox]] 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 plan]]s of the animal phyla have evolved and how they are related to one another.<ref>Gould, ''The Structure of Evolutionary Theory'', chapter 10</ref>

The [[Human Genome Project]]—the largest, most costly single biological study ever undertaken—began in 1988 under the leadership of [[James D. Watson]], after preliminary work with genetically simpler model organisms such as ''[[E. coli]]'', ''[[S. cerevisiae]]'' and ''[[Caenorhabditis elegans|C. elegans]]''. [[Shotgun sequencing]] and gene discovery methods pioneered by [[Craig Venter]]—and fueled by the financial promise of gene patents with [[Celera Genomics]]— led to a public–private sequencing competition that ended in compromise with the first draft of the human DNA sequence announced in 2000.<ref>Davies, ''Cracking the Genome'', Introduction; see also: Sulston, ''The Common Thread''</ref>

==Twenty-first century biological sciences==
At the beginning of the 21st century, biological sciences converged with previously differentiated new and classic disciplines like [[physics]] into research fields like [[biophysics]]. Advances were made in [[analytical chemistry]] and physics instrumentation including improved sensors, [[optics]], tracers, instrumentation, signal processing, networks, [[Robot|robots]], satellites, and compute power for data collection, storage, analysis, modeling, visualization, and simulations. These technological advances allowed theoretical and experimental research including internet publication of molecular [[biochemistry]], [[biological system]]s, and ecosystems science. This enabled worldwide access to better measurements, theoretical models, complex simulations, theory predictive model experimentation, analysis, worldwide internet observational [[data reporting]], open peer-review, collaboration, and internet publication. New fields of biological sciences research emerged including [[bioinformatics]], [[neuroscience]], [[theoretical biology]], [[computational genomics]], [[astrobiology]] and [[synthetic biology]].

==See also==
* [[History of botany]]
* [[Outline of biology]]
* [[Timeline of biology and organic chemistry]]

== References ==

=== Citations ===
{{Reflist}}

=== Sources ===
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{{refend}}

==External links==
{{Library resources box
 |onlinebooks=yes
 |by=no
 |lcheading= Biology History
 |label=History of biology
 }}
* [http://www.ishpssb.org/ International Society for History, Philosophy, and Social Studies of Biology] – professional history of biology organization
* [http://www.historyworld.net/wrldhis/PlainTextHistories.asp?historyid=ac22 History of Biology] – Historyworld article
* [https://www.bioexplorer.net/History_of_Biology/ History of Biology] at Bioexplorer.Net – a collection of history of biology links
* [http://en.citizendium.org/wiki/Biology Biology] – historically oriented article on Citizendium
* [https://archive.org/details/historyofbiology00mialrich Miall, L. C. (1911) History of biology.] Watts & Co. London
* {{Cite Americana|wstitle=Biology|author=[[Ernest Ingersoll]] |short=x}}

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