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{{For|the video game|History of Biology (video game)}}
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[[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===
{{
Widespread travel by naturalists in the early-
====Geology and paleontology====
{{
The emerging discipline of geology also brought natural history and natural philosophy closer together; the establishment of the [[stratigraphy|stratigraphic column]] linked the
====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
[[
[[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>{{Cite web|url=https://history.info/on-this-day/1843-robert-koch-man-saved-millions-lives/|title=1843: Robert Koch: The Man who Saved Millions of Lives | History.info|date=10 December 2019 |accessdate=26 June 2025}}</ref> For his life's work Koch is seen as one of the founders of modern medicine.<ref>{{Cite web|url=https://www.facebook.com/IndianHumanists/videos/e181-founders-of-modern-medicine-robert-koch-father-of-medical-bacteriology/245261433654285/|title=E181: Founders of Modern Medicine: Robert Koch (Father of Medical Bacteriology) | Episode 181: 26 June 2021- Science Hour Discussion topic: Founders of Modern Medicine: Robert Koch (Father of Medical Bacteriology) రాబర్ట్ కోచ్... | By Indian Humanists | Facebook|accessdate=26 June 2025|via=www.facebook.com}}</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.
By the mid
====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]].
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]].
==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===
{{
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.
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
In the 1960s, as evolutionary theorists explored the possibility of multiple [[units of selection]], ecologists turned to evolutionary approaches.
===Classical genetics, the modern synthesis, and evolutionary theory===
{{
[[
1900 marked the so-called ''rediscovery of Mendel''
Hugo de Vries tried to
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]].
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>
===Biochemistry, microbiology, and molecular biology===
{{
{{multiple image
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| image1 = Portrait of Fritz Albert Lipmann (1899-1986), Biochemist (2551001689).jpg
| caption1 = [[Fritz Albert Lipmann]]
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| caption2 = [[Hans Krebs (biochemist)|Hans Krebs]]
}}
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
====Origins of molecular biology====
Following the rise of classical genetics, many
[[
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.
The development of standard, genetically uniform organisms that could produce repeatable experimental results was essential for the development of [[molecular genetics]].
[[
[[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 [[
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 | last1=Epstein | first1=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 | 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>
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.
Resistance to the growing influence of molecular biology was especially evident in [[evolutionary biology]].
===Biotechnology, genetic engineering, and genomics===
{{
[[Biotechnology]] in the general sense has been an important part of biology since the late 19th century.
[[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]].
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.
Following Asilomar, new genetic engineering techniques and applications developed rapidly.
====Molecular systematics and genomics====
{{
[[
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]].
The development and popularization of the [[polymerase chain reaction]] (PCR) in mid
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]]
==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]].
==
* [[History of botany]]
* [[Outline of biology]]
* [[Timeline of biology and organic chemistry]]
== References ==
=== Citations ===
{{Reflist}}
=== Sources ===
{{refbegin|60em}}
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* Raby, Peter. ''Bright Paradise: Victorian Scientific Travellers''. Princeton University Press: Princeton, 1997. {{ISBN|0-691-04843-6}}
* Rothman, Sheila M. and David J. Rothman. ''The Pursuit of Perfection: The Promise and Perils of Medical Enhancement''. Vintage Books: New York, 2003. {{ISBN|0-679-75835-6}}
* [[Jan Sapp|Sapp, Jan]]. ''Genesis: The Evolution of Biology''. Oxford University Press: New York, 2003. {{ISBN|0-19-515618-8}}
* [[James A. Secord|Secord, James A.]] ''Victorian Sensation: The Extraordinary Publication, Reception, and Secret Authorship of ''Vestiges of the Natural History of Creation. University of Chicago Press: Chicago, 2000. {{ISBN|0-226-74410-8}}
* Serafini, Anthony ''The Epic History of Biology'', Perseus Publishing, 1993.
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* Smocovitis, Vassiliki Betty. ''Unifying Biology: The Evolutionary Synthesis and Evolutionary Biology''. Princeton University Press: Princeton, 1996. {{ISBN|0-691-03343-9}}
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* Thackray, Arnold, ed. ''Private Science: Biotechnology and the Rise of the Molecular Sciences''. University of Pennsylvania Press: Philadelphia, 1998. {{ISBN|0-8122-3428-6}}
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{{refend}}
==External links==
{{Library resources box
|onlinebooks=yes
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|label=History of biology
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* [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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