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== History == {{Main|History of botany}} === Early botany === [[File:Cork Micrographia Hooke.png|thumb|alt=engraving of cork cells from Hooke's Micrographia, 1665|An engraving of the cells of [[Cork oak|cork]], from [[Robert Hooke]]'s ''[[Micrographia]]'', 1665]] {{multiple image|caption_align=center | total_width = 275 <!--image 1--> | image1 = Renaissance C14 Füllmaurer Leonhart Fuchs.jpg | width1 = 700 | height1 = 828 | alt1 = | link1 = | caption1 = [[Leonhart Fuchs]] <!--image 2--> | image2 = Brunfels-1.png | width2 = 700 | height2 = 828 | alt2 = | link2 = | caption2 = [[Otto Brunfels]] <!--image 3--> | image3 = Hieronymus Bock (1546).jpg | width3 = 700 | height3 = 828 | alt3 = | link3 = | caption3 = [[Hieronymus Bock]] }} Botany originated as [[herbalism]], the study and use of plants for their [[Medicinal plants|possible medicinal properties]].{{sfn|Sumner|2000|p = 16}} The early recorded history of botany includes many ancient writings and plant classifications. Examples of early botanical works have been found in ancient texts from India dating back to before 1100 BCE,{{sfn|Reed|1942|pp = 7–29}}{{sfn|Oberlies|1998|p = 155}} [[Ancient Egypt]],{{sfn|Manniche|2006}} in archaic [[Avestan language|Avestan]] writings, and in works from China purportedly from before 221 BCE.{{sfn|Reed|1942|pp = 7–29}}{{sfn|Needham|Lu|Huang|1986}} Modern botany traces its roots back to [[Ancient Greece]] specifically to [[Theophrastus]] ({{circa|371}}–287 BCE), a student of [[Aristotle]] who invented and described many of its principles and is widely regarded in the [[scientific community]] as the "Father of Botany".{{sfn|Greene|1909|pp = 140–142}} His major works, ''[[Historia Plantarum (Theophrastus)|Enquiry into Plants]]'' and ''On the Causes of Plants'', constitute the most important contributions to botanical science until the [[Middle Ages]], almost seventeen centuries later.{{sfn|Greene|1909|pp = 140–142}}{{sfn|Bennett|Hammond|1902|p = 30}} Another work from Ancient Greece that made an early impact on botany is {{Lang|la|[[De materia medica]]}}, a five-volume encyclopedia about [[Herbalism|preliminary herbal medicine]] written in the middle of the first century by Greek physician and pharmacologist [[Pedanius Dioscorides]]. {{Lang|la|De materia medica}} was widely read for more than 1,500 years.{{sfn|Mauseth|2003|p = 532}} Important contributions from the [[Islamic Golden Age|medieval Muslim world]] include [[Ibn Wahshiyya]]'s ''[[Nabatean Agriculture]]'', [[Abū Ḥanīfa Dīnawarī]]'s (828–896) the ''Book of Plants'', and [[Ibn Bassal]]'s ''The Classification of Soils''. In the early 13th century, [[Abu al-Abbas al-Nabati]], and [[Ibn al-Baitar]] (d. 1248) wrote on botany in a systematic and scientific manner.{{sfn|Dallal|2010|p = 197}}{{sfn|Panaino|2002|p = 93}}{{sfn|Levey|1973|p = 116}} In the mid-16th century, [[botanical garden]]s were founded in a number of Italian universities. The [[Orto botanico di Padova|Padua botanical garden]] in 1545 is usually considered to be the first which is still in its original location. These gardens continued the practical value of earlier "physic gardens", often associated with monasteries, in which plants were cultivated for suspected medicinal uses. They supported the growth of botany as an academic subject. Lectures were given about the plants grown in the gardens. Botanical gardens came much later to northern Europe; the first in England was the [[University of Oxford Botanic Garden]] in 1621.{{sfn|Hill|1915}} German physician [[Leonhart Fuchs]] (1501–1566) was one of "the three German fathers of botany", along with theologian [[Otto Brunfels]] (1489–1534) and physician [[Hieronymus Bock]] (1498–1554) (also called Hieronymus Tragus).{{sfn|National Museum of Wales|2007}}{{sfn|Yaniv|Bachrach|2005|p = 157}} Fuchs and Brunfels broke away from the tradition of copying earlier works to make original observations of their own. Bock created his own system of plant classification. Physician [[Valerius Cordus]] (1515–1544) authored a botanically and pharmacologically important herbal ''Historia Plantarum'' in 1544 and a [[pharmacopoeia]] of lasting importance, the ''Dispensatorium'' in 1546.{{sfn|Sprague|Sprague|1939}} Naturalist [[Conrad von Gesner]] (1516–1565) and herbalist [[John Gerard]] (1545 – {{circa|1611|lk=no}}) published herbals covering the supposed medicinal uses of plants. Naturalist [[Ulisse Aldrovandi]] (1522–1605) was considered the ''father of natural history'', which included the study of plants. In 1665, using an early microscope, [[Polymath]] [[Robert Hooke]] discovered [[cell (biology)|cells]] (a term he coined) in [[cork (material)|cork]], and a short time later in living plant tissue.{{sfn|Waggoner|2001}} === Early modern botany === {{Further|Taxonomy (biology)#History of taxonomy}} [[File:CarlvonLinne Garden.jpg|thumb|left|alt=Photograph of a garden|The [[Linnaean Garden]] of Linnaeus' residence in Uppsala, Sweden, was planted according to his ''Systema sexuale''.]] During the 18th century, systems of [[plant identification]] were developed comparable to [[single access key|dichotomous keys]], where unidentified plants are placed into [[taxon]]omic groups (e.g. family, genus and species) by making a series of choices between pairs of [[Character (biology)|characters]]. The choice and sequence of the characters may be artificial in keys designed purely for identification ([[single access key#Diagnostic ('artificial') versus synoptic ('natural') keys|diagnostic keys]]) or more closely related to the natural or [[taxonomic order|phyletic order]] of the [[taxon|taxa]] in synoptic keys.{{sfn|Scharf|2009|pp = 73–117}} By the 18th century, new plants for study were arriving in Europe in increasing numbers from newly discovered countries and the European colonies worldwide. In 1753, [[Carl Linnaeus]] published his [[Species Plantarum]], a hierarchical classification of plant species that remains the reference point for [[International Code of Nomenclature for algae, fungi, and plants|modern botanical nomenclature]]. This established a standardised binomial or two-part naming scheme where the first name represented the [[genus]] and the second identified the [[species]] within the genus.{{sfn|Capon|2005|pp = 220–223}} For the purposes of identification, Linnaeus's ''Systema Sexuale'' [[Linnaean taxonomy#Classification of plants|classified]] plants into 24 groups according to the number of their male sexual organs. The 24th group, ''Cryptogamia'', included all plants with concealed reproductive parts, [[moss]]es, [[liverwort]]s, [[fern]]s, [[algae]] and [[Fungus|fungi]].{{sfn|Hoek|Mann|Jahns|2005|p = 9}} Increasing knowledge of [[plant anatomy]], [[plant morphology|morphology]] and life cycles led to the realisation that there were more natural affinities between plants than the artificial sexual system of Linnaeus. [[Michel Adanson|Adanson]] (1763), [[Antoine Laurent de Jussieu|de Jussieu]] (1789), and [[Augustin Pyramus de Candolle|Candolle]] (1819) all proposed various alternative natural systems of classification that grouped plants using a wider range of shared characters and were widely followed. The [[Candollean system]] reflected his ideas of the progression of morphological complexity and the later [[Bentham & Hooker system]], which was influential until the mid-19th century, was influenced by Candolle's approach. [[Charles Darwin|Darwin]]'s publication of the ''[[On the Origin of Species|Origin of Species]]'' in 1859 and his concept of common descent required modifications to the Candollean system to reflect evolutionary relationships as distinct from mere morphological similarity.{{sfn|Starr|2009|pp =299–}} In the 19th century botany was a socially acceptable hobby for upper-class women. These women would collect and paint flowers and plants from around the world with scientific accuracy. The paintings were used to record many species that could not be transported or maintained in other environments. [[Marianne North]] illustrated over 900 species in extreme detail with watercolor and oil paintings.<ref>{{Cite web |last=Ross |first=Ailsa |date=2015-04-22 |title=The Victorian Gentlewoman Who Documented 900 Plant Species |url=http://www.atlasobscura.com/articles/marianne-north-early-female-explorer |access-date=2024-06-05 |website=Atlas Obscura |language=en}}</ref> Her work and many other women's botany work was the beginning of popularizing botany to a wider audience. Botany was greatly stimulated by the appearance of the first "modern" textbook, [[Matthias Jakob Schleiden|Matthias Schleiden]]'s ''{{lang|de|Grundzüge der Wissenschaftlichen Botanik}}'', published in English in 1849 as ''Principles of Scientific Botany''.{{sfn|Morton|1981|p = 377}} Schleiden was a microscopist and an early plant anatomist who co-founded the [[cell theory]] with [[Theodor Schwann]] and [[Rudolf Virchow]] and was among the first to grasp the significance of the [[cell nucleus]] that had been described by [[Robert Brown (botanist, born 1773)|Robert Brown]] in 1831.{{sfn|Harris|2000|pp = 76–81}} In 1855, [[Adolf Fick]] formulated [[Fick's laws of diffusion|Fick's laws]] that enabled the calculation of the rates of [[molecular diffusion]] in biological systems.{{sfn|Small|2012|pp =118–}} [[File:Echeveria glauca II.jpg|thumb|''Echeveria glauca'' in a Connecticut greenhouse. Botany uses Latin names for identification; here, the specific name ''glauca'' means blue.]] === Late modern botany === Building upon the gene-chromosome theory of heredity that originated with [[Gregor Mendel]] (1822–1884), [[August Weismann]] (1834–1914) proved that inheritance only takes place through [[gamete]]s. No other cells can pass on inherited characters.{{sfn|Karp|2009|p = 382}} The work of [[Katherine Esau]] (1898–1997) on plant anatomy is still a major foundation of modern botany. Her books ''Plant Anatomy'' and ''Anatomy of Seed Plants'' have been key plant structural biology texts for more than half a century.{{sfn|National Science Foundation|1989}}{{sfn|Chaffey|2007|pp = 481–482}} [[File:ETH-BIB-Kurs in Alpenbotanik-Ans 08784-01-031.tif|left|thumb|upright|Class of alpine botany in Switzerland, 1936]] The discipline of [[plant ecology]] was pioneered in the late 19th century by botanists such as [[Eugenius Warming]], who produced the hypothesis that plants form [[plant community|communities]], and his mentor and successor [[Christen C. Raunkiær]] whose system for describing [[Raunkiær plant life-form|plant life forms]] is still in use today. The concept that the composition of plant communities such as [[Temperate broadleaf and mixed forests|temperate broadleaf forest]] changes by a process of [[ecological succession]] was developed by [[Henry Chandler Cowles]], [[Arthur Tansley]] and [[Frederic Clements]]. Clements is credited with the idea of [[climax vegetation]] as the most complex vegetation that an environment can support and Tansley introduced the concept of [[ecosystem]]s to biology.{{sfn|Tansley|1935|pp=299–302}}{{sfn|Willis|1997|pp=267–271}}{{sfn|Morton|1981|p = 457}} Building on the extensive earlier work of [[Alphonse Pyramus de Candolle|Alphonse de Candolle]], [[Nikolai Ivanovich Vavilov|Nikolai Vavilov]] (1887–1943) produced accounts of the [[biogeography]], [[Center of origin|centres of origin]], and evolutionary history of economic plants.{{sfn|de Candolle|2006|pp = 9–25, 450–465}} Particularly since the mid-1960s there have been advances in understanding of the physics of [[Plant physiology|plant physiological]] processes such as [[transpiration]] (the transport of water within plant tissues), the temperature dependence of rates of water [[evaporation]] from the leaf surface and the [[molecular diffusion]] of water vapour and carbon dioxide through [[stomatal]] apertures. These developments, coupled with new methods for measuring the size of stomatal apertures, and the rate of [[photosynthesis]] have enabled precise description of the rates of [[gas exchange]] between plants and the atmosphere.{{sfn|Jasechko|Sharp|Gibson|Birks|2013|pp = 347–350}}{{sfn|Nobel|1983|p = 608}} Innovations in [[Statistics|statistical analysis]] by [[Ronald Fisher]],{{sfn|Yates|Mather|1963|pp = 91–129}} [[Frank Yates]] and others at [[Rothamsted Research#Statistical science|Rothamsted Experimental Station]] facilitated rational experimental design and data analysis in botanical research.{{sfn|Finney|1995|pp = 554–573}} The discovery and identification of the [[auxin]] plant hormones by [[Kenneth V. Thimann]] in 1948 enabled regulation of plant growth by externally applied chemicals. [[Frederick Campion Steward]] pioneered techniques of [[micropropagation]] and [[plant tissue culture]] controlled by [[Plant physiology#Plant hormones|plant hormones]].{{sfn|Cocking|1993}} The synthetic auxin [[2,4-Dichlorophenoxyacetic acid|2,4-dichlorophenoxyacetic acid]] or 2,4-D was one of the first commercial synthetic [[herbicide]]s.{{sfn|Cousens|Mortimer|1995}} [[File:Apfe-auf-Naehrboden.jpg|thumb|upright|alt=Micropropagation of transgenic plants|Micropropagation of transgenic plants]] 20th century developments in plant biochemistry have been driven by modern techniques of [[organic chemistry|organic chemical analysis]], such as [[spectroscopy]], [[chromatography]] and [[electrophoresis]]. With the rise of the related molecular-scale biological approaches of [[molecular biology]], [[genomics]], [[proteomics]] and [[metabolomics]], the relationship between the plant [[genome]] and most aspects of the biochemistry, physiology, morphology and behaviour of plants can be subjected to detailed experimental analysis.{{sfn|Ehrhardt|Frommer|2012|pp = 1–21}} The concept originally stated by [[Gottlieb Haberlandt]] in 1902{{sfn|Haberlandt|1902|pages=69–92}} that all plant cells are [[Cell potency#Totipotency|totipotent]] and can be grown ''in vitro'' ultimately enabled the use of [[genetic engineering]] experimentally to knock out a gene or genes responsible for a specific trait, or to add genes such as [[Green fluorescent protein|GFP]] that [[reporter gene|report]] when a gene of interest is being expressed. These technologies enable the biotechnological use of whole plants or plant cell cultures grown in [[bioreactors]] to synthesise [[Bt corn|pesticides]], [[Biopharmaceutics|antibiotics]] or other [[pharming (genetics)|pharmaceuticals]], as well as the practical application of [[genetically modified crops]] designed for traits such as improved yield.{{sfn|Leonelli|Charnley|Webb|Bastow|2012}} Modern morphology recognises a continuum between the major morphological categories of root, stem (caulome), leaf (phyllome) and [[trichome]].{{sfn|Sattler|Jeune|1992|pp = 249–262}} Furthermore, it emphasises structural dynamics.{{sfn|Sattler|1992|pp = 708–714}} Modern systematics aims to reflect and discover [[Phylogenetic nomenclature|phylogenetic relationships]] between plants.{{sfn|Ereshefsky|1997|pp = 493–519}}{{sfn|Gray|Sargent|1889|pp = 292–293}}{{sfn|Medbury|1993|pp = 14–16}}{{sfn|Judd|Campbell|Kellogg|Stevens|2002|pp = 347–350}} Modern [[molecular phylogenetics]] largely ignores morphological characters, relying on DNA sequences as data. Molecular analysis of [[nucleic acid sequence|DNA sequences]] from most families of flowering plants enabled the [[Angiosperm Phylogeny Group]] to publish in 1998 a [[phylogenetics|phylogeny]] of flowering plants, answering many of the questions about relationships among [[angiosperm]] families and species.{{sfn|Burger|2013}} The theoretical possibility of a practical method for identification of plant species and commercial varieties by [[DNA barcoding]] is the subject of active current research.{{sfn|Kress|Wurdack|Zimmer|Weigt|2005|pp = 8369–8374}}{{sfn|Janzen|Forrest|Spouge|Hajibabaei|2009|pp = 12794–12797}}
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