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The history of botany examines the human effort to understand life on Earth by tracing the historical development of the discipline of
botany Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek w ...
—that part of natural science dealing with organisms traditionally treated as plants. Rudimentary botanical science began with empirically based plant lore passed from generation to generation in the oral traditions of paleolithic hunter-gatherers. The first written records of plants were made in the Neolithic Revolution about 10,000 years ago as writing was developed in the settled agricultural communities where plants and animals were first domesticated. The first writings that show human curiosity about plants themselves, rather than the uses that could be made of them, appear in ancient Greece and ancient India. In Ancient Greece, the teachings of
Aristotle's Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatetic school of phil ...
student Theophrastus at the
Lyceum The lyceum is a category of educational institution defined within the education system of many countries, mainly in Europe. The definition varies among countries; usually it is a type of secondary school. Generally in that type of school the ...
in ancient Athens in about 350 BC are considered the starting point for Western botany. In ancient India, the Vŗkşăyurvĕda, attributed to Parăśara, is also considered one of the earliest texts to describe various branches of botany. In Europe, botanical science was soon overshadowed by a medieval preoccupation with the medicinal properties of plants that lasted more than 1000 years. During this time, the medicinal works of classical antiquity were reproduced in manuscripts and books called herbals. In China and the Arab world, the Greco-Roman work on medicinal plants was preserved and extended. In Europe the Renaissance of the 14th–17th centuries heralded a scientific revival during which botany gradually emerged from natural history as an independent science, distinct from medicine and agriculture. Herbals were replaced by
floras Flora is all the plant life present in a particular region or time, generally the naturally occurring ( indigenous) native plants. Sometimes bacteria and fungi are also referred to as flora, as in the terms ''gut flora'' or ''skin flora''. Et ...
: books that described the native plants of local regions. The invention of the microscope stimulated the study of
plant anatomy Plant anatomy or phytotomy is the general term for the study of the internal structure of plants. Originally it included plant morphology, the description of the physical form and external structure of plants, but since the mid-20th century plant ...
, and the first carefully designed experiments in plant physiology were performed. With the expansion of trade and exploration beyond Europe, the many new plants being discovered were subjected to an increasingly rigorous process of
naming Naming is assigning a name to something. Naming may refer to: * Naming (parliamentary procedure), a procedure in certain parliamentary bodies * Naming ceremony, an event at which an infant is named * Product naming, the discipline of deciding wh ...
, description, and
classification Classification is a process related to categorization, the process in which ideas and objects are recognized, differentiated and understood. Classification is the grouping of related facts into classes. It may also refer to: Business, organizat ...
. Progressively more sophisticated scientific technology has aided the development of contemporary botanical offshoots in the plant sciences, ranging from the applied fields of
economic botany ''Economic Botany'' is a quarterly peer-reviewed academic journal that covers all aspects of economic botany. The editor-in-chief is Robert A. Voeks (California State University, Fullerton). The journal was established in 1947 and is published by ...
(notably agriculture, horticulture and forestry), to the detailed examination of the structure and function of plants and their interaction with the environment over many scales from the large-scale global significance of vegetation and plant communities (
biogeography Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, i ...
and ecology) through to the small scale of subjects like
cell theory In biology, cell theory is a scientific theory first formulated in the mid-nineteenth century, that living organisms are made up of cells, that they are the basic structural/organizational unit of all organisms, and that all cells come from pre ...
, molecular biology and plant
biochemistry Biochemistry or biological chemistry is the study of chemical processes within and relating to living organisms. A sub-discipline of both chemistry and biology, biochemistry may be divided into three fields: structural biology, enzymology and ...
.


Introduction

Botany Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek w ...
( Greek - grass, fodder; Medieval Latin ' – herb, plant) and zoology are, historically, the core disciplines of
biology Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditary ...
whose history is closely associated with the natural sciences chemistry, physics and geology. A distinction can be made between botanical science in a pure sense, as the study of plants themselves, and botany as applied science, which studies the human use of plants. Early natural history divided pure botany into three main streams
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
-
classification Classification is a process related to categorization, the process in which ideas and objects are recognized, differentiated and understood. Classification is the grouping of related facts into classes. It may also refer to: Business, organizat ...
,
anatomy Anatomy () is the branch of biology concerned with the study of the structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old science, having it ...
and physiology – that is, external form, internal structure, and functional operation. The most obvious topics in applied botany are horticulture, forestry and
agriculture Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people to ...
although there are many others like
weed science A weed is a plant considered undesirable in a particular situation, "a plant in the wrong place", or a plant growing where it is not wanted.Harlan, J. R., & deWet, J. M. (1965). Some thoughts about weeds. ''Economic botany'', ''19''(1), 16-24. ...
,
plant pathology Plant pathology (also phytopathology) is the scientific study of diseases in plants caused by pathogens (infectious organisms) and environmental conditions (physiological factors). Organisms that cause infectious disease include fungi, oom ...
, floristry, pharmacognosy,
economic botany ''Economic Botany'' is a quarterly peer-reviewed academic journal that covers all aspects of economic botany. The editor-in-chief is Robert A. Voeks (California State University, Fullerton). The journal was established in 1947 and is published by ...
and ethnobotany which lie outside modern courses in botany. Since the origin of botanical science there has been a progressive increase in the scope of the subject as technology has opened up new techniques and areas of study. Modern molecular systematics, for example, entails the principles and techniques of taxonomy, molecular biology, computer science and more. Within botany there are a number of sub-disciplines that focus on particular plant groups, each with their own range of related studies (anatomy, morphology etc.). Included here are: phycology (
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthesis, photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from u ...
),
pteridology A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except th ...
( ferns),
bryology Bryology (from Greek , a moss, a liverwort) is the branch of botany concerned with the scientific study of bryophytes ( mosses, liverworts, and hornworts). Bryologists are people who have an active interest in observing, recording, classifying ...
( mosses and liverworts) and
palaeobotany Paleobotany, which is also spelled as palaeobotany, is the branch of botany dealing with the recovery and identification of plant remains from geological contexts, and their use for the biological reconstruction of past environments (paleogeogr ...
(fossil plants) and their histories are treated elsewhere (see side bar). To this list can be added mycology, the study of fungi, which were once treated as plants, but are now ranked as a unique kingdom.


Ancient knowledge

Nomadic hunter-gatherer societies passed on, by oral tradition, what they knew (their empirical observations) about the different kinds of plants that they used for food, shelter, poisons, medicines, for ceremonies and rituals etc. The uses of plants by these pre-literate societies influenced the way the plants were named and classified—their uses were embedded in folk-taxonomies, the way they were grouped according to use in everyday communication. The nomadic life-style was drastically changed when settled communities were established in about twelve centres around the world during the Neolithic Revolution which extended from about 10,000 to 2500 years ago depending on the region. With these communities came the development of the technology and skills needed for the
domestication of plants Domestication is a sustained multi-generational relationship in which humans assume a significant degree of control over the reproduction and care of another group of organisms to secure a more predictable supply of resources from that group. A ...
and animals and the emergence of the written word provided evidence for the passing of systematic knowledge and culture from one generation to the next.


Plant lore and plant selection

During the Neolithic Revolution plant knowledge increased most obviously through the use of plants for food and medicine. All of today's
staple food A staple food, food staple, or simply a staple, is a food that is eaten often and in such quantities that it constitutes a dominant portion of a standard Diet (nutrition), diet for a given person or group of people, supplying a large fraction of ...
s were domesticated in
prehistoric Prehistory, also known as pre-literary history, is the period of human history between the use of the first stone tools by hominins 3.3 million years ago and the beginning of recorded history with the invention of writing systems. The use o ...
times as a gradual process of selection of higher-yielding varieties took place, possibly unknowingly, over hundreds to thousands of years. Legumes were cultivated on all continents but cereals made up most of the regular diet: rice in East Asia, wheat and barley in the Middle east, and maize in Central and South America. By Greco-Roman times popular food plants of today, including grapes, apples,
fig The fig is the edible fruit of ''Ficus carica'', a species of small tree in the flowering plant family Moraceae. Native to the Mediterranean and western Asia, it has been cultivated since ancient times and is now widely grown throughout the world ...
s, and olives, were being listed as named varieties in early manuscripts. Botanical authority
William Stearn William Thomas Stearn (16 April 1911 – 9 May 2001) was a British botanist. Born in Cambridge in 1911, he was largely self-educated, and developed an early interest in books and natural history. His initial work experience was at a ...
has observed that "''cultivated plants are mankind's most vital and precious heritage from remote antiquity''". It is also from the Neolithic, in about 3000 BC, that we glimpse the first known illustrations of plants and read descriptions of impressive gardens in Egypt. However protobotany, the first pre-scientific written record of plants, did not begin with food; it was born out of the medicinal literature of Egypt, China, Mesopotamia and India. Botanical historian Alan Morton notes that agriculture was the occupation of the poor and uneducated, while medicine was the realm of socially influential
shaman Shamanism is a religious practice that involves a practitioner (shaman) interacting with what they believe to be a spirit world through altered states of consciousness, such as trance. The goal of this is usually to direct spirits or spiri ...
s, priests,
apothecaries ''Apothecary'' () is a mostly archaic term for a medical professional who formulates and dispenses '' materia medica'' (medicine) to physicians, surgeons, and patients. The modern chemist (British English) or pharmacist (British and North Ameri ...
,
magician Magician or The Magician may refer to: Performers * A practitioner of magic (supernatural) * A practitioner of magic (illusion) * Magician (fantasy), a character in a fictional fantasy context Entertainment Books * ''The Magician'', an 18th-ce ...
s and physicians, who were more likely to record their knowledge for posterity.


Early botany

;Ancient India An early example of ancient Indian plant classification is found in the Rigveda, a collection of Vedic Sanskrit hymns from about 3700–3100 BP. Plants are divided into ' (trees), ' (herbs useful to humans) and ' (creepers), with further subdivisions. The sacred Hindu text
Atharvaveda The Atharva Veda (, ' from ' and ''veda'', meaning "knowledge") is the "knowledge storehouse of ''atharvāṇas'', the procedures for everyday life".Laurie Patton (2004), Veda and Upanishad, in ''The Hindu World'' (Editors: Sushil Mittal and G ...
divides plants into eight classes: ' (spreading branches), ' (leaves with long clusters), ' (bushy plants), ' (which expands); ' (those with
monopodial Vascular plants with monopodial growth habits grow upward from a single point. They add leaves to the apex each year and the stem grows longer accordingly. The word ''Monopodial'' is derived from Greek "mono-", ''one'' and "podial", "foot", in ref ...
growth), ' (creeping plants), ' (with many stalks), and ' (plants with knotty joints). The Taittiriya Samhita classifies the plant kingdom into ', ' and ' (trees), ' (shrubs with spreading branches), ' (herbs), ' (spreading plant), ' (climber), ' (bushy plant), ' (creeper), and ' (spreading on the ground). Other examples of early Indian taxonomy include
Manusmriti The ''Manusmṛiti'' ( sa, मनुस्मृति), also known as the ''Mānava-Dharmaśāstra'' or Laws of Manu, is one of the many legal texts and constitution among the many ' of Hinduism. In ancient India, the sages often wrote the ...
, the Law book of Hindus, which classifies plants into eight major categories. Elaborate taxonomies also occur in the
Charaka Samhitā The ''Charaka Samhita'' (, “Compendium of ''Charaka''”) is a Sanskrit text on Ayurveda (Indian traditional medicine). Along with the ''Sushruta Samhita'', it is one of the two foundational texts of this field that have survived from ancien ...
, Sushruta Samhita and
Vaisesika Vaisheshika or Vaiśeṣika ( sa, वैशेषिक) is one of the six schools of Indian philosophy (Vedic systems) from ancient India. In its early stages, the Vaiśeṣika was an independent philosophy with its own metaphysics, epistemolog ...
. ;Ancient China In
ancient China The earliest known written records of the history of China date from as early as 1250 BC, from the Shang dynasty (c. 1600–1046 BC), during the reign of king Wu Ding. Ancient historical texts such as the ''Book of Documents'' (early chapter ...
lists of different plants and herb concoctions for pharmaceutical purposes date back to at least the time of the Warring States (481 BC-221 BC). Many Chinese writers over the centuries contributed to the written knowledge of herbal pharmaceutics. The Han Dynasty (202 BC-220 AD) includes the notable work of the Huangdi Neijing and the famous pharmacologist
Zhang Zhongjing Zhang Zhongjing (; 150–219), formal name Zhang Ji (), was a Chinese pharmacologist, physician, inventor, and writer of the Eastern Han dynasty and one of the most eminent Chinese physicians during the later years of the Han dynasty. He estab ...
. There were also the 11th century scientists and statesmen Su Song and Shen Kuo who compiled learned treatises on natural history, emphasising herbal medicine.


Theophrastus and the origin of botanical science

Ancient Athens, of the 6th century BC, was the busy trade centre at the confluence of Egyptian, Mesopotamian and
Minoa Minoa was the name of several Bronze-Age port cities on the coasts of the Aegean islands Crete, Paros, Siphnos, Amorgos and Corfu in Greece, as well as the Italian island of Sicily.F. Schachermeyer (1964). ''Die Minoische Kultur des alten Kr ...
n cultures at the height of Greek colonisation of the Mediterranean. The philosophical thought of this period ranged freely through many subjects. Empedocles (490–430 BC) foreshadowed Darwinian evolutionary theory in a crude formulation of the mutability of species and natural selection. The physician Hippocrates (460–370 BC) avoided the prevailing superstition of his day and approached healing by close observation and the test of experience. At this time a genuine non-
anthropocentric Anthropocentrism (; ) is the belief that human beings are the central or most important entity in the universe. The term can be used interchangeably with humanocentrism, and some refer to the concept as human supremacy or human exceptionalism. F ...
curiosity about plants emerged. The major works written about plants extended beyond the description of their medicinal uses to the topics of plant geography, morphology, physiology, nutrition, growth and reproduction. Foremost among the scholars studying botany was Theophrastus of Eressus ( Greek: ; c. 371–287 BC) who has been frequently referred to as the "Father of Botany". He was a student and close friend of Aristotle (384–322 BC) and succeeded him as head of the
Lyceum The lyceum is a category of educational institution defined within the education system of many countries, mainly in Europe. The definition varies among countries; usually it is a type of secondary school. Generally in that type of school the ...
(an educational establishment like a modern university) in Athens with its tradition of
peripatetic Peripatetic may refer to: *Peripatetic school The Peripatetic school was a school of philosophy in Ancient Greece. Its teachings derived from its founder, Aristotle (384–322 BC), and ''peripatetic'' is an adjective ascribed to his followers. ...
philosophy. Aristotle's special treatise on plants — — is now lost, although there are many botanical observations scattered throughout his other writings (these have been assembled by Christian Wimmer in ', 1836) but they give little insight into his botanical thinking. The Lyceum prided itself in a tradition of systematic observation of causal connections, critical experiment and rational theorizing. Theophrastus challenged the superstitious medicine employed by the physicians of his day, called rhizotomi, and also the control over medicine exerted by priestly authority and tradition. Together with Aristotle he had tutored
Alexander the Great Alexander III of Macedon ( grc, Ἀλέξανδρος, Alexandros; 20/21 July 356 BC – 10/11 June 323 BC), commonly known as Alexander the Great, was a king of the ancient Greek kingdom of Macedon. He succeeded his father Philip II to ...
whose military conquests were carried out with all the scientific resources of the day, the Lyceum garden probably containing many botanical trophies collected during his campaigns as well as other explorations in distant lands. It was in this garden where he gained much of his plant knowledge. Theophrastus's major botanical works were the ''
Enquiry into Plants Theophrastus's ''Enquiry into Plants'' or ''Historia Plantarum'' ( grc-gre, Περὶ φυτῶν ἱστορία, ''Peri phyton historia'') was, along with his mentor Aristotle's ''History of Animals'', Pliny the Elder's '' Natural History'' a ...
'' (''Historia Plantarum'') and ''Causes of Plants'' (''Causae Plantarum'') which were his lecture notes for the Lyceum. The opening sentence of the ''Enquiry'' reads like a botanical manifesto: "''We must consider the distinctive characters and the general nature of plants from the point of view of their
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
, their behaviour under external conditions, their mode of generation and the whole course of their life''". The ''Enquiry'' is 9 books of "applied" botany dealing with the forms and
classification Classification is a process related to categorization, the process in which ideas and objects are recognized, differentiated and understood. Classification is the grouping of related facts into classes. It may also refer to: Business, organizat ...
of plants and
economic botany ''Economic Botany'' is a quarterly peer-reviewed academic journal that covers all aspects of economic botany. The editor-in-chief is Robert A. Voeks (California State University, Fullerton). The journal was established in 1947 and is published by ...
, examining the techniques of
agriculture Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled people to ...
(relationship of crops to soil, climate, water and habitat) and horticulture. He described some 500 plants in detail, often including descriptions of habitat and geographic distribution, and he recognised some plant groups that can be recognised as modern-day plant families. Some names he used, like '' Crataegus'', '' Daucus'' and '' Asparagus'' have persisted until today. His second book ''Causes of Plants'' covers plant growth and reproduction (akin to modern physiology). Like Aristotle he grouped plants into "trees", "undershrubs", "shrubs" and "herbs" but he also made several other important botanical distinctions and observations. He noted that plants could be
annual Annual may refer to: *Annual publication, periodical publications appearing regularly once per year **Yearbook **Literary annual *Annual plant *Annual report *Annual giving *Annual, Morocco, a settlement in northeastern Morocco *Annuals (band), a ...
s,
perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...
s and
biennials A biennial plant is a flowering plant that, generally in a temperate climate, takes two years to complete its biological life cycle. Life cycle In its first year, the biennal plant undergoes primary growth, during which its vegetative structures ...
, they were also either
monocotyledon Monocotyledons (), commonly referred to as monocots, ( Lilianae ''sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of ...
s or
dicotyledon The dicotyledons, also known as dicots (or, more rarely, dicotyls), are one of the two groups into which all the flowering plants (angiosperms) were formerly divided. The name refers to one of the typical characteristics of the group: namely, t ...
s and he also noticed the difference between
determinate In logic and philosophy (especially metaphysics), a property is a characteristic of an object; a red object is said to have the property of redness. The property may be considered a form of object in its own right, able to possess other propertie ...
and indeterminate growth and details of floral structure including the degree of fusion of the petals, position of the ovary and more. These lecture notes of Theophrastus comprise the first clear exposition of the rudiments of plant anatomy, physiology, morphology and ecology — presented in a way that would not be matched for another eighteen centuries. Meanwhile, the study of medicinal plants was not being neglected and a full synthesis of ancient Greek pharmacology was compiled in ''Materia Medica'' c. 60 AD by
Pedanius Dioscorides Pedanius Dioscorides ( grc-gre, Πεδάνιος Διοσκουρίδης, ; 40–90 AD), “the father of pharmacognosy”, was a Greek physician, pharmacologist, botanist, and author of '' De materia medica'' (, On Medical Material) —a 5-vo ...
(c. 40-90 AD) who was a Greek physician with the Roman army. This work proved to be the definitive text on medicinal herbs, both oriental and occidental, for fifteen hundred years until the dawn of the European Renaissance being slavishly copied again and again throughout this period. Though rich in medicinal information with descriptions of about 600 medicinal herbs, the botanical content of the work was extremely limited.


Ancient Rome

The Romans contributed little to the foundations of botanical science laid by the ancient Greeks, but made a sound contribution to our knowledge of applied botany as agriculture. In works titled ' four Roman writers contributed to a compendium ''Scriptores Rei Rusticae'', published from the Renaissance on, which set out the principles and practice of agriculture. These authors were Cato (234–149 BC), Varro (116–27 BC) and, in particular,
Columella Lucius Junius Moderatus Columella (; Arabic: , 4 – ) was a prominent writer on agriculture in the Roman Empire. His ' in twelve volumes has been completely preserved and forms an important source on Roman agriculture, together with the wo ...
(4–70 AD) and Palladius (4th century AD). Roman encyclopaedist Pliny the Elder (23–79 AD) deals with plants in Books 12 to 26 of his 37-volume highly influential work ' in which he frequently quotes Theophrastus but with a lack of botanical insight although he does, nevertheless, draw a distinction between true botany on the one hand, and farming and medicine on the other. It is estimated that at the time of the Roman Empire between 1300 and 1400 plants had been recorded in the West.


Medieval knowledge


Medicinal plants of the early Middle Ages

In Western Europe, after Theophrastus, botany passed through a bleak period of 1800 years when little progress was made and, indeed, many of the early insights were lost. As Europe entered the Middle Ages (5th to 15th centuries), China, India and the Arab world enjoyed a golden age. Chinese philosophy had followed a similar path to that of the ancient Greeks. The Chinese dictionary-encyclopaedia
Erh Ya The ''Erya'' or ''Erh-ya'' is the first surviving Chinese dictionary. Bernhard Karlgren (1931:49) concluded that "the major part of its glosses must reasonably date from the 3rd century BC." Title Chinese scholars interpret the first title cha ...
probably dates from about 300 BC and describes about 334 plants classed as trees or shrubs, each with a common name and illustration. Between 100 and 1700 AD many new works on pharmaceutical botany were produced including encyclopaedic accounts and treatises compiled for the Chinese imperial court. These were free of superstition and myth with carefully researched descriptions and nomenclature; they included cultivation information and notes on economic and medicinal uses — and even elaborate monographs on ornamental plants. But there was no experimental method and no analysis of the plant sexual system, nutrition, or anatomy. The 400-year period from the 9th to 13th centuries AD was the
Islamic Renaissance The Islamic Golden Age was a period of cultural, economic, and scientific flourishing in the history of Islam, traditionally dated from the 8th century to the 14th century. This period is traditionally understood to have begun during the reign ...
, a time when Islamic culture and science thrived. Greco-Roman texts were preserved, copied and extended although new texts always emphasised the medicinal aspects of plants. Kurdish biologist Ābu Ḥanīfah Āḥmad ibn Dawūd Dīnawarī (828–896 AD) is known as the founder of Arabic botany; his ''Kitâb al-nabât'' ('Book of Plants') describes 637 species, discussing plant development from germination to senescence and including details of flowers and fruits. The
Mutazilite Muʿtazila ( ar, المعتزلة ', English: "Those Who Withdraw, or Stand Apart", and who called themselves ''Ahl al-ʿAdl wa al-Tawḥīd'', English: "Party of ivineJustice and Oneness f God); was an Islamic group that appeared in early Islami ...
philosopher and physician Ibn Sina (
Avicenna Ibn Sina ( fa, ابن سینا; 980 – June 1037 CE), commonly known in the West as Avicenna (), was a Persian polymath who is regarded as one of the most significant physicians, astronomers, philosophers, and writers of the Islamic ...
) (c. 980–1037 AD) was another influential figure, his '' The Canon of Medicine'' being a landmark in the history of medicine treasured until the Enlightenment. In India simple artificial plant classification systems of the Rigveda,
Atharvaveda The Atharva Veda (, ' from ' and ''veda'', meaning "knowledge") is the "knowledge storehouse of ''atharvāṇas'', the procedures for everyday life".Laurie Patton (2004), Veda and Upanishad, in ''The Hindu World'' (Editors: Sushil Mittal and G ...
and Taittiriya Samhita became more botanical with the work of Parashara (c. 400 – c. 500 AD), the author of ''Vṛksayurveda'' (the science of life of trees). He made close observations of cells and leaves and divided plants into Dvimatrka (
Dicotyledon The dicotyledons, also known as dicots (or, more rarely, dicotyls), are one of the two groups into which all the flowering plants (angiosperms) were formerly divided. The name refers to one of the typical characteristics of the group: namely, t ...
s) and Ekamatrka (
Monocotyledon Monocotyledons (), commonly referred to as monocots, ( Lilianae ''sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of ...
s). The dicotyledons were further classified into groupings (ganas) akin to modern floral families: ''Samiganiya'' ( Fabaceae), ''Puplikagalniya'' ( Rutaceae), ''Svastikaganiya'' (
Cruciferae Brassicaceae () or (the older) Cruciferae () is a medium-sized and economically important family of flowering plants commonly known as the mustards, the crucifers, or the cabbage family. Most are herbaceous plants, while some are shrubs. The leav ...
), ''Tripuspaganiya'' (
Cucurbitaceae The Cucurbitaceae, also called cucurbits or the gourd family, are a plant family consisting of about 965 species in around 95 genera, of which the most important to humans are: *''Cucurbita'' – squash, pumpkin, zucchini, some gourds *'' Lagen ...
), ''Mallikaganiya'' (
Apocynaceae Apocynaceae (from ''Apocynum'', Greek for "dog-away") is a family of flowering plants that includes trees, shrubs, herbs, stem succulents, and vines, commonly known as the dogbane family, because some taxa were used as dog poison Members of the ...
), and ''Kurcapuspaganiya'' (
Asteraceae The family Asteraceae, alternatively Compositae, consists of over 32,000 known species of flowering plants in over 1,900 genera within the order Asterales. Commonly referred to as the aster, daisy, composite, or sunflower family, Compositae w ...
). Important medieval Indian works of plant physiology include the ''Prthviniraparyam'' of
Udayana Udayana, (Devanagari: उदयन) also known as Udayanācārya (Udyanacharya, or Master Udayana), (circa 975 - 1050 CE) was an Indian philosopher and logician of the tenth century of the Nyaya school who attempted to devise a rational theol ...
, ''Nyayavindutika'' of Dharmottara, ''Saddarsana-samuccaya'' of Gunaratna, and ''Upaskara'' of Sankaramisra.


The Silk Road

Following the fall of Constantinople (1453), the newly expanded Ottoman Empire welcomed European embassies in its capital, which in turn became the sources of plants from those regions to the east which traded with the empire. In the following century twenty times as many plants entered Europe along the Silk Road as had been transported in the previous two thousand years, mainly as bulbs. Others were acquired primarily for their alleged medicinal value. Initially Italy benefited from this new knowledge, especially Venice, which traded extensively with the East. From there these new plants rapidly spread to the rest of Western Europe. By the middle of the sixteenth century there was already a flourishing export trade of various bulbs from Turkey to Europe.


The Age of Herbals

In the European Middle Ages of the 15th and 16th centuries the lives of European citizens were based around agriculture but when printing arrived, with movable type and woodcut illustrations, it was not treatises on agriculture that were published, but lists of medicinal plants with descriptions of their properties or "virtues". These first plant books, known as herbals showed that botany was still a part of medicine, as it had been for most of ancient history. Authors of herbals were often curators of university gardens, and most herbals were derivative compilations of classic texts, especially ''
De Materia Medica (Latin name for the Greek work , , both meaning "On Medical Material") is a pharmacopoeia of medicinal plants and the medicines that can be obtained from them. The five-volume work was written between 50 and 70 CE by Pedanius Dioscorides, a ...
''. However, the need for accurate and detailed plant descriptions meant that some herbals were more botanical than medicinal. German Otto Brunfels's (1464–1534) ''Herbarum Vivae Icones'' (1530) contained descriptions of about 47 species new to science combined with accurate illustrations. His fellow countryman
Hieronymus Bock Hieronymus Bock ( Latinised Hieronymus Tragus; c. 1498 – 21 February 1554) was a German botanist, physician, and Lutheran minister who began the transition from medieval botany to the modern scientific worldview by arranging plants by their re ...
's (1498–1554) ''Kreutterbuch'' of 1539 described plants he found in nearby woods and fields and these were illustrated in the 1546 edition. However, it was
Valerius Cordus Valerius Cordus (18 February 1515 – 25 September 1544) was a German physician, botanist and pharmacologist who authored the first pharmacopoeia North of the Alps and one of the most celebrated herbals in history. He is also widely credited ...
(1515–1544) who pioneered the formal botanical description that detailed both flowers and fruits, some anatomy including the number of chambers in the
ovary The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. ...
, and the type of ovule placentation. He also made observations on pollen and distinguished between inflorescence types. His five-volume ''Historia Plantarum'' was published about 18 years after his early death aged 29 in 1561–1563. In Holland Rembert Dodoens (1517–1585), in ''Stirpium Historiae'' (1583), included descriptions of many new species from the Netherlands in a scientific arrangement and in England William Turner (1515–1568) in his ''Libellus De Re Herbaria Novus'' (1538) published names, descriptions and localities of many native British plants. Herbals contributed to botany by setting in train the science of plant description, classification, and botanical illustration. Up to the 17th century botany and medicine were one and the same but those books emphasising medicinal aspects eventually omitted the plant lore to become modern pharmacopoeias; those that omitted the medicine became more botanical and evolved into the modern compilations of plant descriptions we call
Flora Flora is all the plant life present in a particular region or time, generally the naturally occurring ( indigenous) native plants. Sometimes bacteria and fungi are also referred to as flora, as in the terms '' gut flora'' or ''skin flora''. ...
s. These were often backed by specimens deposited in a herbarium which was a collection of dried plants that verified the plant descriptions given in the Floras. The transition from herbal to Flora marked the final separation of botany from medicine.


The Renaissance and Age of Enlightenment (1550–1800)

The revival of learning during the European Renaissance renewed interest in plants. The church, feudal aristocracy and an increasingly influential merchant class that supported science and the arts, now jostled in a world of increasing trade. Sea voyages of exploration returned botanical treasures to the large public, private, and newly established botanic gardens, and introduced an eager population to novel crops, drugs and spices from Asia, the
East Indies The East Indies (or simply the Indies), is a term used in historical narratives of the Age of Discovery. The Indies refers to various lands in the East or the Eastern hemisphere, particularly the islands and mainlands found in and around t ...
and the New World. The number of scientific publications increased. In England, for example, scientific communication and causes were facilitated by learned societies like Royal Society (founded in 1660) and the Linnaean Society (founded in 1788): there was also the support and activities of botanical institutions like the Jardin du Roi in Paris,
Chelsea Physic Garden The Chelsea Physic Garden was established as the Apothecaries' Garden in London, England, in 1673 by the Worshipful Society of Apothecaries to grow plants to be used as medicines. This four acre physic garden, the term here referring to the sc ...
,
Royal Botanic Gardens Kew Royal Botanic Gardens, Kew is a non-departmental public body in the United Kingdom sponsored by the Department for Environment, Food and Rural Affairs. An internationally important botanical research and education institution, it employs 1,100 ...
, and the Oxford and
Cambridge Botanic Garden The Cambridge University Botanic Garden is a botanical garden located in Cambridge, England, associated with the university Department of Plant Sciences (formerly Botany School). It lies between Trumpington Road to the west, Bateman Street to ...
s, as well as the influence of renowned private gardens and wealthy entrepreneurial nurserymen. By the early 17th century the number of plants described in Europe had risen to about 6000. The 18th century Enlightenment values of reason and science coupled with new voyages to distant lands instigating another phase of encyclopaedic plant identification, nomenclature, description and illustration, "flower painting" possibly at its best in this period of history. Plant trophies from distant lands decorated the gardens of Europe's powerful and wealthy in a period of enthusiasm for natural history, especially botany (a preoccupation sometimes referred to as "botanophilia") that is never likely to recur. Often such exotic new plant imports (primarily from Turkey), when they first appeared in print in English, lacked common names in the language. During the 18th century botany was one of the few sciences considered appropriate for genteel educated women. Around 1760, with the popularization of the Linnaean system, botany became much more widespread among educated women who painted plants, attended classes on plant classification, and collected herbarium specimens although emphasis was on the healing properties of plants rather than plant reproduction which had overtones of sexuality. Women began publishing on botanical topics and children's books on botany appeared by authors like
Charlotte Turner Smith Charlotte Smith (née Turner; – ) was an English novelist and poet of the School of Sensibility whose '' Elegiac Sonnets'' (1784) contributed to the revival of the form in England. She also helped to set conventions for Gothic fiction and wr ...
. Cultural authorities argued that education through botany created culturally and scientifically aware citizens, part of the thrust for 'improvement' that characterised the Enlightenment. However, in the early 19th century with the recognition of botany as an official science, women were again excluded from the discipline. Compared to other sciences, however, in botany the number of female researchers, collectors, or illustrators has always been remarkably high.


Botanical gardens and herbaria

Public and private gardens have always been strongly associated with the historical unfolding of botanical science. Early botanical gardens were physic gardens, repositories for the medicinal plants described in the herbals. As they were generally associated with universities or other academic institutions the plants were also used for study. The directors of these gardens were eminent physicians with an educational role as "scientific gardeners" and it was staff of these institutions that produced many of the published herbals. The botanical gardens of the modern tradition were established in northern Italy, the first being at Pisa (1544), founded by
Luca Ghini Luca Ghini (Casalfiumanese, 1490 – Bologna, 4 May 1556) was an Italian physician and botanist, notable as the creator of the first recorded herbarium, as well as the first botanical garden in Europe. Biography Ghini was born in Casalfiumanese, ...
(1490–1556). Although part of a medical faculty, the first chair of ', essentially a chair in botany, was established in Padua in 1533. Then in 1534, Ghini became Reader in ' at Bologna University, where
Ulisse Aldrovandi Ulisse Aldrovandi (11 September 1522 – 4 May 1605) was an Italian naturalist, the moving force behind Bologna's botanical garden, one of the first in Europe. Carl Linnaeus and the comte de Buffon reckoned him the father of natural history ...
established a similar garden in 1568 (see below). Collections of pressed and dried specimens were called a ' (garden of dry plants) and the first accumulation of plants in this way (including the use of a plant press) is attributed to Ghini. Buildings called
herbaria A herbarium (plural: herbaria) is a collection of preserved plant specimens and associated data used for scientific study. The specimens may be whole plants or plant parts; these will usually be in dried form mounted on a sheet of paper (called ...
housed these specimens mounted on card with descriptive labels. Stored in cupboards in systematic order they could be preserved in perpetuity and easily transferred or exchanged with other institutions, a taxonomic procedure that is still used today. By the 18th century the physic gardens had been transformed into "order beds" that demonstrated the classification systems that were being devised by botanists of the day — but they also had to accommodate the influx of curious, beautiful and new plants pouring in from voyages of exploration that were associated with European colonial expansion.


From Herbal to Flora

Plant classification systems of the 17th and 18th centuries now related plants to one another and not to man, marking a return to the non-anthropocentric botanical science promoted by Theophrastus over 1500 years before. In England, various herbals in either Latin or English were mainly compilations and translations of continental European works, of limited relevance to the British Isles. This included the rather unreliable work of Gerard (1597). The first systematic attempt to collect information on British plants was that of Thomas Johnson (1629), who was later to issue his own revision of Gerard's work (1633–1636). However Johnson was not the first apothecary or physician to organise botanical expeditions to systematise their local flora. In Italy
Ulisse Aldrovandi Ulisse Aldrovandi (11 September 1522 – 4 May 1605) was an Italian naturalist, the moving force behind Bologna's botanical garden, one of the first in Europe. Carl Linnaeus and the comte de Buffon reckoned him the father of natural history ...
(1522 – 1605) organised an expedition to the Sibylline mountains in
Umbria it, Umbro (man) it, Umbra (woman) , population_note = , population_blank1_title = , population_blank1 = , demographics_type1 = , demographics1_footnotes = , demographics1_title1 = , demographics1_info1 = , ...
in 1557, and compiled a local
Flora Flora is all the plant life present in a particular region or time, generally the naturally occurring ( indigenous) native plants. Sometimes bacteria and fungi are also referred to as flora, as in the terms '' gut flora'' or ''skin flora''. ...
. He then began to disseminate his findings amongst other European scholars, forming an early network of
knowledge sharing Knowledge sharing is an activity through which knowledge (namely, information, skills, or expertise) is exchanged among people, friends, peers, families, communities (for example, Wikipedia), or within or between organizations. It bridges the in ...
"''molti amici in molti luoghi''" (many friends in many places), including Charles de l'Écluse ( Clusius) (1526 – 1609) at
Montpellier Montpellier (, , ; oc, Montpelhièr ) is a city in southern France near the Mediterranean Sea. One of the largest urban centres in the region of Occitania, Montpellier is the prefecture of the department of Hérault. In 2018, 290,053 people l ...
and Jean de Brancion at
Malines Mechelen (; french: Malines ; traditional English name: MechlinMechelen has been known in English as ''Mechlin'', from where the adjective ''Mechlinian'' is derived. This name may still be used, especially in a traditional or historical contex ...
. Between them they started developing Latin names for plants, in addition to their common names. The exchange of information and specimens between scholars was often associated with the founding of
botanical gardens A botanical garden or botanic gardenThe terms ''botanic'' and ''botanical'' and ''garden'' or ''gardens'' are used more-or-less interchangeably, although the word ''botanic'' is generally reserved for the earlier, more traditional gardens, an ...
(above), and to this end Aldrovandi founded one of the earliest at his university in Bologna, the
Orto Botanico di Bologna Orto may refer to: * Orto, Corse-du-Sud Orto (; co, Ortu) is a commune in the Corse-du-Sud department of France on the island of Corsica. Population See also *Communes of the Corse-du-Sud department An intentional community is a v ...
in 1568. In France, Clusius journeyed throughout most of Western Europe, making discoveries in the vegetable kingdom along the way. He compiled Flora of Spain (1576), and Austria and Hungary (1583). He was the first to propose dividing plants into classes. Meanwhile, in Switzerland, from 1554, Conrad Gessner (1516 – 1565) made regular explorations of the
Swiss Alps The Alpine region of Switzerland, conventionally referred to as the Swiss Alps (german: Schweizer Alpen, french: Alpes suisses, it, Alpi svizzere, rm, Alps svizras), represents a major natural feature of the country and is, along with the Swis ...
from his native Zurich and discovered many new plants. He proposed that there were groups or genera of plants. He said that each genus was composed of many species and that these were defined by similar flowers and fruits. This principle of organization laid the groundwork for future botanists. He wrote his important '' Historia Plantarum'' shortly before his death. At Malines, in Flanders he established and maintained the botanical gardens of Jean de Brancion from 1568 to 1573, and first encountered
tulips Tulips (''Tulipa'') are a genus of spring-blooming perennial herbaceous bulbiferous geophytes (having bulbs as storage organs). The flowers are usually large, showy and brightly coloured, generally red, pink, yellow, or white (usually in warm ...
. This approach coupled with the new Linnaean system of
binomial nomenclature In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
resulted in plant encyclopaedias without medicinal information called ''Floras'' that meticulously described and illustrated the plants growing in particular regions. The 17th century also marked the beginning of experimental botany and application of a rigorous scientific method, while improvements in the microscope launched the new discipline of plant anatomy whose foundations, laid by the careful observations of Englishman
Nehemiah Grew Nehemiah Grew (26 September 164125 March 1712) was an English plant anatomist and physiologist, known as the "Father of Plant Anatomy". Biography Grew was the only son of Obadiah Grew (1607–1688), Nonconformist divine and vicar of St Mic ...
and Italian
Marcello Malpighi Marcello Malpighi (10 March 1628 – 30 November 1694) was an Italian biologist and physician, who is referred to as the "Founder of microscopical anatomy, histology & Father of physiology and embryology". Malpighi's name is borne by several ph ...
, would last for 150 years.


Botanical exploration

More new lands were opening up to European colonial powers, the botanical riches being returned to European botanists for description. This was a romantic era of botanical explorers, intrepid plant hunters and gardener-botanists. Significant botanical collections came from: the West Indies (
Hans Sloane Sir Hans Sloane, 1st Baronet (16 April 1660 – 11 January 1753), was an Irish physician, naturalist, and collector, with a collection of 71,000 items which he bequeathed to the British nation, thus providing the foundation of the British Mu ...
(1660–1753)); China (James Cunningham); the spice islands of the East Indies (Moluccas, George Rumphius (1627–1702)); China and Mozambique ( João de Loureiro (1717–1791)); West Africa ( Michel Adanson (1727–1806)) who devised his own classification scheme and forwarded a crude theory of the mutability of species; Canada, Hebrides, Iceland, New Zealand by
Captain James Cook James Cook (7 November 1728 Old Style date: 27 October – 14 February 1779) was a British explorer, navigator, cartographer, and captain in the British Royal Navy, famous for his three voyages between 1768 and 1779 in the Pacific Ocean and ...
's chief botanist
Joseph Banks Sir Joseph Banks, 1st Baronet, (19 June 1820) was an English naturalist, botanist, and patron of the natural sciences. Banks made his name on the 1766 natural-history expedition to Newfoundland and Labrador. He took part in Captain James C ...
(1743–1820).


Classification and morphology

By the middle of the 18th century the botanical booty resulting from the era of exploration was accumulating in gardens and herbaria – and it needed to be systematically catalogued. This was the task of the taxonomists, the plant classifiers. Plant classifications have changed over time from "artificial" systems based on general habit and form, to pre-evolutionary "natural" systems expressing similarity using one to many characters, leading to post-evolutionary "natural" systems that use characters to infer
evolutionary relationships Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
. Italian physician Andrea Caesalpino (1519–1603) studied medicine and taught botany at the University of Pisa for about 40 years eventually becoming Director of the Botanic Garden of Pisa from 1554 to 1558. His sixteen-volume ''De Plantis'' (1583) described 1500 plants and his herbarium of 260 pages and 768 mounted specimens still remains. Caesalpino proposed classes based largely on the detailed structure of the flowers and fruit; he also applied the concept of the genus. He was the first to try and derive principles of natural classification reflecting the overall similarities between plants and he produced a classification scheme well in advance of its day. Gaspard Bauhin (1560–1624) produced two influential publications ' (1620) and ' (1623). These brought order to the 6000 species now described and in the latter he used binomials and synonyms that may well have influenced Linnaeus's thinking. He also insisted that taxonomy should be based on natural affinities. To sharpen the precision of description and classification
Joachim Jung Joachim Jungius (born Joachim Junge; 22 October 1587 – 23 September 1657) was a German mathematician, logician and philosopher of science. Life Jungius was a native of Lübeck. He studied metaphysics at the Universities of Rostock and Giess ...
(1587–1657) compiled a much-needed botanical terminology which has stood the test of time. English botanist John Ray (1623–1705) built on Jung's work to establish the most elaborate and insightful classification system of the day. His observations started with the local plants of Cambridge where he lived, with the ' (1860) which later expanded to his ', essentially the first British Flora. Although his ' (1682, 1688, 1704) provided a step towards a world Flora as he included more and more plants from his travels, first on the continent and then beyond. He extended Caesalpino's natural system with a more precise definition of the higher classification levels, deriving many modern families in the process, and asserted that all parts of plants were important in classification. He recognised that variation arises from both internal (genotypic) and external environmental (phenotypic) causes and that only the former was of taxonomic significance. He was also among the first experimental physiologists. The ' can be regarded as the first botanical synthesis and textbook for modern botany. According to botanical historian Alan Morton, Ray "influenced both the theory and the practice of botany more decisively than any other single person in the latter half of the seventeenth century". Ray's family system was later extended by
Pierre Magnol Pierre Magnol (8 June 1638 – 21 May 1715) was a French botanist. He was born in the city of Montpellier, where he lived and worked for most of his life. He became Professor of Botany and Director of the Royal Botanic Garden of Montpellier and he ...
(1638–1715) and Joseph de Tournefort (1656–1708), a student of Magnol, achieved notoriety for his botanical expeditions, his emphasis on floral characters in classification, and for reviving the idea of the genus as the basic unit of classification. Above all it was Swedish Carl Linnaeus (1707–1778) who eased the task of plant cataloguing. He adopted a sexual system of classification using stamens and pistils as important characters. Among his most important publications were Systema Naturae (1735), Genera Plantarum (1737), and Philosophia Botanica (1751) but it was in his Species Plantarum (1753) that he gave every species a
binomial Binomial may refer to: In mathematics *Binomial (polynomial), a polynomial with two terms *Binomial coefficient, numbers appearing in the expansions of powers of binomials *Binomial QMF, a perfect-reconstruction orthogonal wavelet decomposition * ...
thus setting the path for the future accepted method of designating the names of all organisms. Linnaean thought and books dominated the world of taxonomy for nearly a century. His sexual system was later elaborated by
Bernard de Jussieu Bernard de Jussieu (; 17 August 1699 – 6 November 1777) was a French naturalist, younger brother of Antoine de Jussieu. Bernard de Jussieu was born in Lyon. He took a medical degree at Montpellier and began practice in 1720, but finding the w ...
(1699–1777) whose nephew
Antoine-Laurent de Jussieu Antoine-Laurent de Lavoisier ( , ; ; 26 August 17438 May 1794),
CNRS (
Michel Adanson (1727–1806) in his ' (1763, 1764), apart from extending the current system of family names, emphasized that a natural classification must be based on a consideration of all characters, even though these may later be given different emphasis according to their diagnostic value for the particular plant group. Adanson's method has, in essence, been followed to this day. 18th century plant taxonomy bequeathed to the 19th century a precise binomial nomenclature and botanical terminology, a system of classification based on natural affinities, and a clear idea of the ranks of family, genus and species — although the taxa to be placed within these ranks remains, as always, the subject of taxonomic research.


Anatomy

In the first half of the 18th century botany was beginning to move beyond descriptive science into experimental science. Although the microscope was invented in 1590 it was only in the late 17th century that lens grinding provided the resolution needed to make major discoveries. Antony van Leeuwenhoek is a notable example of an early lens grinder who achieved remarkable resolution with his single-lens microscopes. Important general biological observations were made by Robert Hooke (1635–1703) but the foundations of plant anatomy were laid by Italian
Marcello Malpighi Marcello Malpighi (10 March 1628 – 30 November 1694) was an Italian biologist and physician, who is referred to as the "Founder of microscopical anatomy, histology & Father of physiology and embryology". Malpighi's name is borne by several ph ...
(1628–1694) of the University of Bologna in his ' (1675) and Royal Society Englishman
Nehemiah Grew Nehemiah Grew (26 September 164125 March 1712) was an English plant anatomist and physiologist, known as the "Father of Plant Anatomy". Biography Grew was the only son of Obadiah Grew (1607–1688), Nonconformist divine and vicar of St Mic ...
(1628–1711) in his ''The Anatomy of Plants Begun'' (1671) and ''Anatomy of Plants'' (1682). These botanists explored what is now called developmental anatomy and morphology by carefully observing, describing and drawing the developmental transition from seed to mature plant, recording stem and wood formation. This work included the discovery and naming of
parenchyma Parenchyma () is the bulk of functional substance in an animal organ or structure such as a tumour. In zoology it is the name for the tissue that fills the interior of flatworms. Etymology The term ''parenchyma'' is New Latin from the word � ...
and stomata.


Physiology

In plant physiology research interest was focused on the movement of sap and the absorption of substances through the roots. Jan Helmont (1577–1644) by experimental observation and calculation, noted that the increase in weight of a growing plant cannot be derived purely from the soil, and concluded it must relate to water uptake. Englishman
Stephen Hales Stephen Hales (17 September 16774 January 1761) was an English clergyman who made major contributions to a range of scientific fields including botany, pneumatic chemistry and physiology. He was the first person to measure blood pressure. He al ...
(1677–1761) established by quantitative experiment that there is uptake of water by plants and a loss of water by transpiration and that this is influenced by environmental conditions: he distinguished "root pressure", "leaf suction" and "imbibition" and also noted that the major direction of sap flow in woody tissue is upward. His results were published in ''Vegetable Staticks'' (1727) He also noted that "air makes a very considerable part of the substance of vegetables". English chemist
Joseph Priestley Joseph Priestley (; 24 March 1733 – 6 February 1804) was an English chemist, natural philosopher, separatist theologian, grammarian, multi-subject educator, and liberal political theorist. He published over 150 works, and conducted exp ...
(1733–1804) is noted for his discovery of oxygen (as now called) and its production by plants. Later
Jan Ingenhousz Jan (or John) Ingenhousz or Ingen-Housz FRS (8 December 1730 – 7 September 1799) was a Dutch-born British physiologist, biologist and chemist. He is best known for discovering photosynthesis by showing that light is essential to the process ...
(1730–1799) observed that only in sunlight do the green parts of plants absorb air and release oxygen, this being more rapid in bright sunlight while, at night, the air (CO2) is released from all parts. His results were published in ''Experiments upon vegetables'' (1779) and with this the foundations for 20th century studies of carbon fixation were laid. From his observations he sketched the cycle of carbon in nature even though the composition of carbon dioxide was yet to be resolved. Studies in plant nutrition had also progressed. In 1804 Nicolas-Théodore de Saussure's (1767–1845) ' was an exemplary study of scientific exactitude that demonstrated the similarity of respiration in both plants and animals, that the fixation of carbon dioxide includes water, and that just minute amounts of salts and nutrients (which he analyzed in chemical detail from plant ash) have a powerful influence on plant growth.


Plant sexuality

It was Rudolf Camerarius (1665–1721) who was the first to establish plant sexuality conclusively by experiment. He declared in a letter to a colleague dated 1694 and titled ' that "no ovules of plants could ever develop into seeds from the female style and ovary without first being prepared by the pollen from the stamens, the male sexual organs of the plant". Much was learned about plant sexuality by unravelling the reproductive mechanisms of mosses, liverworts and algae. In his ' of 1851
Wilhelm Hofmeister Wilhelm Friedrich Benedikt Hofmeister (18 May 1824 – 12 January 1877) was a German biologist and botanist. He "stands as one of the true giants in the history of biology and belongs in the same pantheon as Darwin and Mendel." He was lar ...
(1824–1877) starting with the ferns and bryophytes demonstrated that the process of sexual reproduction in plants entails an "alternation of generations" between
sporophyte A sporophyte () is the diploid multicellular stage in the life cycle of a plant or alga which produces asexual spores. This stage alternates with a multicellular haploid gametophyte phase. Life cycle The sporophyte develops from the zygote p ...
s and gametophytes. This initiated the new field of
comparative morphology Comparative anatomy is the study of similarities and differences in the anatomy of different species. It is closely related to evolutionary biology and phylogeny (the evolution of species). The science began in the classical era, continuing ...
which, largely through the combined work of William Farlow (1844–1919),
Nathanael Pringsheim Nathanael Pringsheim (30 November 1823 – 6 October 1894) was a German botanist. Biography Nathanael Pringsheim was born at Landsberg, Prussian Silesia, and studied at the universities of Breslau, Leipzig, and Berlin successively. He graduat ...
(1823–1894), Frederick Bower,
Eduard Strasburger Eduard Adolf Strasburger (1 February 1844 – 18 May 1912) was a Polish- German professor and one of the most famous botanists of the 19th century. He discovered mitosis in plants. Life Eduard Strasburger was born in Warsaw, Congress Poland, th ...
and others, established that an "alternation of generations" occurs throughout the plant kingdom. Some time later the German academic and natural historian Joseph Kölreuter (1733–1806) extended this work by noting the function of nectar in attracting pollinators and the role of wind and insects in pollination. He also produced deliberate hybrids, observed the microscopic structure of pollen grains and how the transfer of matter from the pollen to the ovary inducing the formation of the embryo. One hundred years after Camerarius, in 1793, Christian Sprengel (1750–1816) broadened the understanding of flowers by describing the role of nectar guides in pollination, the adaptive floral mechanisms used for pollination, and the prevalence of cross-pollination, even though male and female parts are usually together on the same flower.


Nineteenth-century foundations of modern botany

In about the mid-19th century scientific communication changed. Until this time ideas were largely exchanged by reading the works of authoritative individuals who dominated in their field: these were often wealthy and influential "gentlemen scientists". Now research was reported by the publication of "papers" that emanated from research "schools" that promoted the questioning of conventional wisdom. This process had started in the late 18th century when specialist journals began to appear. Even so, botany was greatly stimulated by the appearance of the first "modern" textbook,
Matthias Schleiden Matthias Jakob Schleiden (; 5 April 1804 – 23 June 1881) was a German botanist and co-founder of cell theory, along with Theodor Schwann and Rudolf Virchow. Career Matthias Jakob Schleiden was born in Hamburg. on 5 April 1804. His father was ...
's (1804–1881) ', published in English in 1849 as ''Principles of Scientific Botany''. By 1850 an invigorated organic chemistry had revealed the structure of many plant constituents. Although the great era of plant classification had now passed the work of description continued.
Augustin de Candolle Augustin Pyramus (or Pyrame) de Candolle (, , ; 4 February 17789 September 1841) was a Swiss botanist. René Louiche Desfontaines launched de Candolle's botanical career by recommending him at a herbarium. Within a couple of years de Candolle ...
(1778–1841) succeeded
Antoine-Laurent de Jussieu Antoine-Laurent de Lavoisier ( , ; ; 26 August 17438 May 1794),
CNRS (
Alphonse Alphonse may refer to: * Alphonse (given name) * Alphonse (surname) * Alphonse Atoll, one of two atolls in the Seychelles' Alphonse Group See also *Alphons Alphons (Latinized ''Alphonsus'', ''Adelphonsus'', or ''Adefonsus'') is a male given n ...
(1806–1893) in the years from 1841 to 1873.


Plant geography and ecology

The opening of the 19th century was marked by an increase in interest in the connection between climate and plant distribution. Carl Willdenow (1765–1812) examined the connection between seed dispersal and distribution, the nature of plant associations and the impact of geological history. He noticed the similarities between the floras of N America and N Asia, the Cape and Australia, and he explored the ideas of "
centre of diversity A center of origin is a geographical area where a group of organisms, either domesticated or wild, first developed its distinctive properties. They are also considered centers of diversity. Centers of origin were first identified in 1924 by Ni ...
" and " centre of origin". German
Alexander von Humboldt Friedrich Wilhelm Heinrich Alexander von Humboldt (14 September 17696 May 1859) was a German polymath, geographer, naturalist, explorer, and proponent of Romantic philosophy and science. He was the younger brother of the Prussian minister, ...
(1769–1859) and Frenchman Aime Bonpland (1773–1858) published a massive and highly influential 30 volume work on their travels; Robert Brown (1773–1852) noted the similarities between the floras of S Africa, Australia and India, while Joakim Schouw (1789–1852) explored more deeply than anyone else the influence on plant distribution of temperature, soil factors, especially soil water, and light, work that was continued by
Alphonse de Candolle Alphonse Louis Pierre Pyramus (or Pyrame) de Candolle (28 October 18064 April 1893) was a French-Swiss botanist, the son of the Swiss botanist Augustin Pyramus de Candolle. Biography De Candolle, son of Augustin Pyramus de Candolle, first devot ...
(1806–1893). Joseph Hooker (1817–1911) pushed the boundaries of floristic studies with his work on Antarctica, India and the Middle East with special attention to endemism.
August Grisebach August Heinrich Rudolf Grisebach () was a German botanist and phytogeographer. He was born in Hannover on 17 April 1814 and died in Göttingen on 9 May 1879. Biography Grisebach studied at the Lyceum in Hanover, the cloister-school at Ilfeld, ...
(1814–1879) in ' (1872) examined physiognomy in relation to climate and in America geographic studies were pioneered by
Asa Gray Asa Gray (November 18, 1810 – January 30, 1888) is considered the most important American botanist of the 19th century. His ''Darwiniana'' was considered an important explanation of how religion and science were not necessarily mutually excl ...
(1810–1888). Physiological plant geography, or ecology, emerged from floristic biogeography in the late 19th century as environmental influences on plants received greater recognition. Early work in this area was synthesised by Danish professor Eugenius Warming (1841–1924) in his book ' (Ecology of Plants, generally taken to mark the beginning of modern ecology) including new ideas on plant communities, their adaptations and environmental influences. This was followed by another grand synthesis, the ' of
Andreas Schimper Andreas Franz Wilhelm Schimper (12 May 1856 – 9 September 1901) was a German botanist and phytogeographer who made major contributions in the fields of histology, ecology and plant geography. He travelled to South East Asia and the Caribbea ...
(1856–1901) in 1898 (published in English in 1903 as Plant-geography upon a physiological basis translated by W. R. Fischer, Oxford: Clarendon press, 839 pp.)


Anatomy

During the 19th century German scientists led the way towards a unitary theory of the structure and life-cycle of plants. Following improvements in the microscope at the end of the 18th century, Charles Mirbel (1776–1854) in 1802 published his ' and Johann Moldenhawer (1766–1827) published ' (1812) in which he describes techniques for separating cells from the middle lamella. He identified vascular and
parenchyma Parenchyma () is the bulk of functional substance in an animal organ or structure such as a tumour. In zoology it is the name for the tissue that fills the interior of flatworms. Etymology The term ''parenchyma'' is New Latin from the word � ...
tous tissues, described vascular bundles, observed the cells in the
cambium A cambium (plural cambia or cambiums), in plants, is a tissue layer that provides partially undifferentiated cells for plant growth. It is found in the area between xylem and phloem. A cambium can also be defined as a cellular plant tissue from ...
, and interpreted tree rings. He found that
stoma In botany, a stoma (from Greek ''στόμα'', "mouth", plural "stomata"), also called a stomate (plural "stomates"), is a pore found in the epidermis of leaves, stems, and other organs, that controls the rate of gas exchange. The pore is bor ...
ta were composed of pairs of cells, rather than a single cell with a hole. Anatomical studies on the stele were consolidated by Carl Sanio (1832–1891) who described the secondary tissues and meristem including
cambium A cambium (plural cambia or cambiums), in plants, is a tissue layer that provides partially undifferentiated cells for plant growth. It is found in the area between xylem and phloem. A cambium can also be defined as a cellular plant tissue from ...
and its action.
Hugo von Mohl Hugo von Mohl FFRS HFRSE (8 April 1805 – 1 April 1872) was a German botanist from Stuttgart. He was the first person to use the word "protoplasm". Life He was a son of the Württemberg statesman Benjamin Ferdinand von Mohl (1766–1845), ...
(1805–1872) summarized work in anatomy leading up to 1850 in ' (1851) but this work was later eclipsed by the encyclopaedic comparative anatomy of
Heinrich Anton de Bary Heinrich Anton de Bary (26 January 183119 January 1888) was a German surgeon, botanist, microbiologist, and mycologist (fungal systematics and physiology). He is considered a founding father of plant pathology (phytopathology) as well as the fou ...
in 1877. An overview of knowledge of the stele in root and stem was completed by Van Tieghem (1839–1914) and of the meristem by
Carl Nägeli Carl Wilhelm von Nägeli (26 or 27 March 1817 – 10 May 1891) was a Swiss botanist. He studied cell division and pollination but became known as the man who discouraged Gregor Mendel from further work on genetics. He rejected natural selection ...
(1817–1891). Studies had also begun on the origins of the
carpel Gynoecium (; ) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) ''pistils' ...
and flower that continue to the present day.


Water relations

The riddle of water and nutrient transport through the plant remained. Physiologist Von Mohl explored solute transport and the theory of water uptake by the roots using the concepts of cohesion, transpirational pull, capillarity and root pressure. German dominance in the field of physiology was underlined by the publication of the definitive textbook on plant physiology synthesising the work of this period, Sach's ''e'' of 1882. There were, however, some advances elsewhere such as the early exploration of geotropism (the effect of gravity on growth) by Englishman Thomas Knight, and the discovery and naming of osmosis by Frenchman
Henri Dutrochet René Joachim Henri Dutrochet (14 November 1776 – 4 February 1847) was a French physician, botanist and physiologist. He is best known for his investigation into osmosis. Early career Dutrochet was born on Néons to a noble family, soon ruine ...
(1776–1847).


Cytology

The cell nucleus was discovered by Robert Brown in 1831. Demonstration of the cellular composition of all organisms, with each cell possessing all the characteristics of life, is attributed to the combined efforts of botanist Matthias Schleiden and zoologist Theodor Schwann (1810–1882) in the early 19th century although Moldenhawer had already shown that plants were wholly cellular with each cell having its own wall and
Julius von Sachs Julius von Sachs (; 2 October 1832 – 29 May 1897) was a German botanist from Breslau, Prussian Silesia. He is considered the founder of experimental plant physiology and co-founder of modern water culture. Julius von Sachs and Wilhelm Knop a ...
had shown the continuity protoplasm between
cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It can be tough, flexible, and sometimes rigid. It provides the cell with both structural support and protection, and also acts as a filtering me ...
s. From 1870 to 1880 it became clear that cell nuclei are never formed anew but always derived from the substance of another nucleus. In 1882 Flemming observed the longitudinal splitting of chromosomes in the dividing nucleus and concluded that each daughter nucleus received half of each of the chromosomes of the mother nucleus: then by the early 20th century it was found that the number of chromosomes in a given species is constant. With genetic continuity confirmed and the finding by
Eduard Strasburger Eduard Adolf Strasburger (1 February 1844 – 18 May 1912) was a Polish- German professor and one of the most famous botanists of the 19th century. He discovered mitosis in plants. Life Eduard Strasburger was born in Warsaw, Congress Poland, th ...
that the nuclei of reproductive cells (in pollen and embryo) have a reducing division (halving of chromosomes, now known as meiosis) the field of heredity was opened up. By 1926 Thomas Morgan was able to outline a theory of the gene and its structure and function. The form and function of plastids received similar attention, the association with starch being noted at an early date. With observation of the cellular structure of all organisms and the process of cell division and continuity of genetic material, the analysis of the structure of protoplasm and the cell wall as well as that of plastids and
vacuole A vacuole () is a membrane-bound organelle which is present in plant and fungal cells and some protist, animal, and bacterial cells. Vacuoles are essentially enclosed compartments which are filled with water containing inorganic and organic mo ...
s – what is now known as
cytology Cell biology (also cellular biology or cytology) is a branch of biology that studies the structure, function, and behavior of cells. All living organisms are made of cells. A cell is the basic unit of life that is responsible for the living and ...
, or
cell theory In biology, cell theory is a scientific theory first formulated in the mid-nineteenth century, that living organisms are made up of cells, that they are the basic structural/organizational unit of all organisms, and that all cells come from pre ...
became firmly established. Later, the cytological basis of the gene-chromosome theory of heredity extended from about 1900–1944 and was initiated by the rediscovery of Gregor Mendel's (1822–1884) laws of plant heredity first published in 1866 in ''Experiments on Plant Hybridization'' and based on cultivated pea, ''Pisum sativum'': this heralded the opening up of plant genetics. The cytological basis for gene-chromosome theory was explored through the role of polyploidy and
hybridization Hybridization (or hybridisation) may refer to: *Hybridization (biology), the process of combining different varieties of organisms to create a hybrid *Orbital hybridization, in chemistry, the mixing of atomic orbitals into new hybrid orbitals *Nu ...
in
speciation Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within ...
and it was becoming better understood that interbreeding populations were the unit of adaptive change in biology.


Developmental morphology and evolution

Until the 1860s it was believed that species had remained unchanged through time: each biological form was the result of an independent act of creation and therefore absolutely distinct and immutable. But the hard reality of geological formations and strange fossils needed scientific explanation.
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended fr ...
's '' Origin of Species'' (1859) replaced the assumption of constancy with the theory of descent with modification. Phylogeny became a new principle as "natural" classifications became classifications reflecting, not just similarities, but evolutionary relationships.
Wilhelm Hofmeister Wilhelm Friedrich Benedikt Hofmeister (18 May 1824 – 12 January 1877) was a German biologist and botanist. He "stands as one of the true giants in the history of biology and belongs in the same pantheon as Darwin and Mendel." He was lar ...
established that there was a similar pattern of organization in all plants expressed through the
alternation of generations Alternation of generations (also known as metagenesis or heterogenesis) is the predominant type of life cycle in plants and algae. It consists of a multicellular haploid sexual phase, the gametophyte, which has a single set of chromosomes alte ...
and extensive homology of structures. Polymath German intellect
Johann Goethe Johann Wolfgang von Goethe (28 August 1749 – 22 March 1832) was a German poet, playwright, novelist, scientist, statesman, theatre director, and critic. His works include plays, poetry, literature, and aesthetic criticism, as well as treat ...
(1749–1832) had interests and influence that extended into botany. In ' (1790) he provided a theory of plant morphology (he coined the word "morphology") and he included within his concept of "metamorphosis" modification during evolution, thus linking comparative morphology with phylogeny. Though the botanical basis of his work has been challenged there is no doubt that he prompted discussion and research on the origin and function of floral parts. His theory probably stimulated the opposing views of German botanists
Alexander Braun Alexander Carl Heinrich Braun (10 May 1805 – 29 March 1877) was a German botanist from Regensburg, Bavaria. His research centered on the morphology of plants. Biography He studied botany in Heidelberg, Paris and Munich. In 1833 he began teachi ...
(1805–1877) and Matthias Schleiden who applied the experimental method to the principles of growth and form that were later extended by
Augustin de Candolle Augustin Pyramus (or Pyrame) de Candolle (, , ; 4 February 17789 September 1841) was a Swiss botanist. René Louiche Desfontaines launched de Candolle's botanical career by recommending him at a herbarium. Within a couple of years de Candolle ...
(1778–1841).


Carbon fixation (photosynthesis)

At the start of the 19th century the idea that plants could synthesize almost all their tissues from atmospheric gases had not yet emerged. The energy component of photosynthesis, the capture and storage of the Sun's radiant energy in carbon bonds (a process on which all life depends) was first elucidated in 1847 by Mayer, but the details of how this was done would take many more years. Chlorophyll was named in 1818 and its chemistry gradually determined, to be finally resolved in the early 20th century. The mechanism of photosynthesis remained a mystery until the mid-19th century when Sachs, in 1862, noted that starch was formed in green cells only in the presence of light and in 1882 he confirmed carbohydrates as the starting point for all other organic compounds in plants. The connection between the pigment chlorophyll and starch production was finally made in 1864 but tracing the precise biochemical pathway of starch formation did not begin until about 1915.


Nitrogen fixation

Significant discoveries relating to nitrogen assimilation and metabolism, including
ammonification The nitrogen cycle is the biogeochemical cycle by which nitrogen is converted into multiple chemical forms as it circulates among atmospheric, terrestrial, and marine ecosystems. The conversion of nitrogen can be carried out through both biologi ...
, nitrification and
nitrogen fixation Nitrogen fixation is a chemical process by which molecular nitrogen (), with a strong triple covalent bond, in the air is converted into ammonia () or related nitrogenous compounds, typically in soil or aquatic systems but also in industry. A ...
(the uptake of atmospheric nitrogen by
symbiotic Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasi ...
soil microorganisms) had to wait for advances in chemistry and bacteriology in the late 19th century and this was followed in the early 20th century by the elucidation of protein and
amino-acid Amino acids are organic compounds that contain both amino and carboxylic acid functional groups. Although hundreds of amino acids exist in nature, by far the most important are the alpha-amino acids, which comprise proteins. Only 22 alpha am ...
synthesis and their role in plant metabolism. With this knowledge it was then possible to outline the global nitrogen cycle.


Twentieth century

20th century science grew out of the solid foundations laid by the breadth of vision and detailed experimental observations of the 19th century. A vastly increased research force was now rapidly extending the horizons of botanical knowledge at all levels of plant organization from molecules to global plant ecology. There was now an awareness of the unity of biological structure and function at the cellular and biochemical levels of organisation. Botanical advance was closely associated with advances in physics and chemistry with the greatest advances in the 20th century mainly relating to the penetration of molecular organization. However, at the level of plant communities it would take until mid century to consolidate work on ecology and
population genetics Population genetics is a subfield of genetics that deals with genetic differences within and between populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as adaptation, speciation, and pop ...
. By 1910 experiments using labelled isotopes were being used to elucidate plant biochemical pathways, to open the line of research leading to gene technology. On a more practical level research funding was now becoming available from agriculture and industry.


Molecules

In 1903
Chlorophyll Chlorophyll (also chlorophyl) is any of several related green pigments found in cyanobacteria and in the chloroplasts of algae and plants. Its name is derived from the Greek words , ("pale green") and , ("leaf"). Chlorophyll allow plants to ...
s a and b were separated by thin layer chromatography then, through the 1920s and 1930s, biochemists, notably Hans Krebs (1900–1981) and
Carl Carl may refer to: * Carl, Georgia, city in USA * Carl, West Virginia, an unincorporated community *Carl (name), includes info about the name, variations of the name, and a list of people with the name * Carl², a TV series * "Carl", an episode of ...
(1896–1984) and
Gerty Cori Gerty Theresa Cori (; August 15, 1896 – October 26, 1957) was an Austro-Hungarian and American biochemist who in 1947 was the third woman to win a Nobel Prize in science, and the first woman to be awarded the Nobel Prize in Physiology or Me ...
(1896–1957) began tracing out the central metabolic pathways of life. Between the 1930s and 1950s it was determined that ATP, located in mitochondria, was the source of cellular chemical energy and the constituent reactions of photosynthesis were progressively revealed. Then, in 1944 DNA was extracted for the first time. Along with these revelations there was the discovery of plant hormones or "growth substances", notably
auxin Auxins (plural of auxin ) are a class of plant hormones (or plant-growth regulators) with some morphogen-like characteristics. Auxins play a cardinal role in coordination of many growth and behavioral processes in plant life cycles and are essenti ...
s, (1934)
gibberellin Gibberellins (GAs) are plant hormones that regulate various developmental processes, including stem elongation, germination, dormancy, flowering, flower development, and leaf and fruit senescence. GAs are one of the longest-known classes of plan ...
s (1934) and
cytokinin Cytokinins (CK) are a class of plant hormones that promote cell division, or cytokinesis, in plant roots and shoots. They are involved primarily in cell growth and differentiation, but also affect apical dominance, axillary bud growth, and lea ...
s (1964) and the effects of
photoperiodism Photoperiodism is the physiological reaction of organisms to the length of night or a dark period. It occurs in plants and animals. Plant photoperiodism can also be defined as the developmental responses of plants to the relative lengths of light ...
, the control of plant processes, especially flowering, by the relative lengths of day and night. Following the establishment of Mendel's laws, the gene-chromosome theory of heredity was confirmed by the work of
August Weismann August Friedrich Leopold Weismann FRS (For), HonFRSE, LLD (17 January 18345 November 1914) was a German evolutionary biologist. Fellow German Ernst Mayr ranked him as the second most notable evolutionary theorist of the 19th century, after Cha ...
who identified chromosomes as the hereditary material. Also, in observing the halving of the chromosome number in germ cells he anticipated work to follow on the details of meiosis, the complex process of redistribution of hereditary material that occurs in the germ cells. In the 1920s and 1930s
population genetics Population genetics is a subfield of genetics that deals with genetic differences within and between populations, and is a part of evolutionary biology. Studies in this branch of biology examine such phenomena as adaptation, speciation, and pop ...
combined the theory of evolution with
Mendelian genetics Mendelian inheritance (also known as Mendelism) is a type of biological inheritance following the principles originally proposed by Gregor Mendel in 1865 and 1866, re-discovered in 1900 by Hugo de Vries and Carl Correns, and later popularized ...
to produce the modern synthesis. By the mid-1960s the molecular basis of metabolism and reproduction was firmly established through the new discipline of molecular biology. Genetic engineering, the insertion of genes into a host cell for cloning, began in the 1970s with the invention of
recombinant DNA Recombinant DNA (rDNA) molecules are DNA molecules formed by laboratory methods of genetic recombination (such as molecular cloning) that bring together genetic material from multiple sources, creating sequences that would not otherwise be fou ...
techniques and its commercial applications applied to agricultural crops followed in the 1990s. There was now the potential to identify organisms by molecular "
fingerprinting A fingerprint is an impression left by the friction ridges of a human finger. The recovery of partial fingerprints from a crime scene is an important method of forensic science. Moisture and grease on a finger result in fingerprints on surfac ...
" and to estimate the times in the past when critical evolutionary changes had occurred through the use of "
molecular clock The molecular clock is a figurative term for a technique that uses the mutation rate of biomolecules to deduce the time in prehistory when two or more life forms diverged. The biomolecular data used for such calculations are usually nucleotid ...
s".


Computers, electron microscopes and evolution

Increased experimental precision combined with vastly improved scientific instrumentation was opening up exciting new fields. In 1936
Alexander Oparin Alexander Ivanovich Oparin (russian: Александр Иванович Опарин; – April 21, 1980) was a Soviet biochemist notable for his theories about the origin of life, and for his book ''The Origin of Life''. He also studied the bi ...
(1894–1980) demonstrated a possible mechanism for the synthesis of organic matter from inorganic molecules. In the 1960s it was determined that the Earth's earliest life-forms treated as plants, the cyanobacteria known as
stromatolite Stromatolites () or stromatoliths () are layered sedimentary formations (microbialite) that are created mainly by photosynthetic microorganisms such as cyanobacteria, sulfate-reducing bacteria, and Pseudomonadota (formerly proteobacteria). The ...
s, dated back some 3.5 billion years. Mid-century transmission and scanning electron microscopy presented another level of resolution to the structure of matter, taking anatomy into the new world of " ultrastructure". New and revised "phylogenetic" classification systems of the plant kingdom were produced by several botanists, including
August Eichler August Wilhelm Eichler, also known under his Latinized name, Augustus Guilielmus Eichler (22 April 1839 – 2 March 1887), was a German botanist who developed a new system of classification of plants to reflect the concept of evolution. His aut ...
. A massive 23 volume ' was published by
Adolf Engler Heinrich Gustav Adolf Engler (25 March 1844 – 10 October 1930) was a German botanist. He is notable for his work on plant taxonomy and phytogeography, such as ''Die natürlichen Pflanzenfamilien'' (''The Natural Plant Families''), edited with ...
& Karl Prantl over the period 1887 to 1915. Taxonomy based on gross morphology was now being supplemented by using characters revealed by pollen morphology,
embryology Embryology (from Greek ἔμβρυον, ''embryon'', "the unborn, embryo"; and -λογία, ''-logia'') is the branch of animal biology that studies the prenatal development of gametes (sex cells), fertilization, and development of embryos an ...
,
anatomy Anatomy () is the branch of biology concerned with the study of the structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old science, having it ...
,
cytology Cell biology (also cellular biology or cytology) is a branch of biology that studies the structure, function, and behavior of cells. All living organisms are made of cells. A cell is the basic unit of life that is responsible for the living and ...
,
serology Serology is the scientific study of serum and other body fluids. In practice, the term usually refers to the diagnostic identification of antibodies in the serum. Such antibodies are typically formed in response to an infection (against a given mi ...
, macromolecules and more. The introduction of computers facilitated the rapid analysis of large data sets used for
numerical taxonomy Numerical taxonomy is a classification system in biological systematics which deals with the grouping by numerical methods of taxonomic units based on their character states. It aims to create a taxonomy using numeric algorithms like cluster ...
(also called taximetrics or phenetics). The emphasis on truly natural phylogenies spawned the disciplines of
cladistics Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived cha ...
and
phylogenetic systematics Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived cha ...
. The grand taxonomic synthesis ''An Integrated System of Classification of Flowering Plants'' (1981) of American
Arthur Cronquist Arthur John Cronquist (March 19, 1919 – March 22, 1992) was an American biologist, botanist and a specialist on Compositae. He is considered one of the most influential botanists of the 20th century, largely due to his formulation of the Cr ...
(1919–1992) was superseded when, in 1998, the
Angiosperm Phylogeny Group The Angiosperm Phylogeny Group (APG) is an informal international group of systematic botanists who collaborate to establish a consensus on the taxonomy of flowering plants (angiosperms) that reflects new knowledge about plant relationships disco ...
published a phylogeny of flowering plants based on the analysis of DNA sequences using the techniques of the new molecular systematics which was resolving questions concerning the earliest evolutionary branches of the
angiosperms Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
(flowering plants). The exact relationship of fungi to plants had for some time been uncertain. Several lines of evidence pointed to fungi being different from plants, animals and bacteria – indeed, more closely related to animals than plants. In the 1980s-90s molecular analysis revealed an evolutionary divergence of fungi from other organisms about 1 billion years ago – sufficient reason to erect a unique kingdom separate from plants.


Biogeography and ecology

The publication of
Alfred Wegener Alfred Lothar Wegener (; ; 1 November 1880 – November 1930) was a German climatologist, geologist, geophysicist, meteorologist, and polar researcher. During his lifetime he was primarily known for his achievements in meteorology and ...
's (1880–1930) theory of
continental drift Continental drift is the hypothesis that the Earth's continents have moved over geologic time relative to each other, thus appearing to have "drifted" across the ocean bed. The idea of continental drift has been subsumed into the science of pl ...
1912 gave additional impetus to comparative physiology and the study of
biogeography Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, i ...
while ecology in the 1930s contributed the important ideas of plant community,
succession Succession is the act or process of following in order or sequence. Governance and politics *Order of succession, in politics, the ascension to power by one ruler, official, or monarch after the death, resignation, or removal from office of ...
, community change, and energy flows. From 1940 to 1950 ecology matured to become an independent discipline as
Eugene Odum Eugene Pleasants Odum (September 17, 1913 – August 10, 2002) was an American biologist at the University of Georgia known for his pioneering work on ecosystem ecology. He and his brother Howard T. Odum wrote the popular ecology textbook, ''Funda ...
(1913–2002) formulated many of the concepts of
ecosystem ecology Ecosystem ecology is the integrated study of living ( biotic) and non-living (abiotic) components of ecosystems and their interactions within an ecosystem framework. This science examines how ecosystems work and relates this to their components ...
, emphasising relationships between groups of organisms (especially material and energy relationships) as key factors in the field. Building on the extensive earlier work of Alphonse de Candolle, Nikolai Vavilov (1887–1943) from 1914 to 1940 produced accounts of the geography, centres of origin, and evolutionary history of economic plants.


Twenty-first century

In reviewing the sweep of botanical history it is evident that, through the power of the scientific method, most of the basic questions concerning the structure and function of plants have, in principle, been resolved. Now the distinction between pure and applied botany becomes blurred as our historically accumulated botanical wisdom at all levels of plant organisation is needed (but especially at the molecular and global levels) to improve human custodianship of planet earth. The most urgent unanswered botanical questions now relate to the role of plants as primary producers in the global cycling of life's basic ingredients: energy, carbon, hydrogen, oxygen, and nitrogen, and ways that our plant stewardship can help address the global environmental issues of
resource management In organizational studies, resource management is the efficient and effective development of an organization's resources when they are needed. Such resources may include the financial resources, inventory, human skills, production resources, or i ...
, conservation, human food security, biologically invasive organisms,
carbon sequestration Carbon sequestration is the process of storing carbon in a carbon pool. Carbon dioxide () is naturally captured from the atmosphere through biological, chemical, and physical processes. These changes can be accelerated through changes in lan ...
, climate change, and sustainability.


See also

*
International Botanical Congress International Botanical Congress (IBC) is an international meeting of botanists in all scientific fields, authorized by the International Association of Botanical and Mycological Societies (IABMS) and held every six years, with the location rotati ...
*
History of plant systematics The history of plant systematics—the biological classification of plants—stretches from the work of ancient Greek to modern evolutionary biologists. As a field of science, plant systematics came into being only slowly, early plant lo ...
*
Botanical illustration Botanical illustration is the art of depicting the form, color, and details of plant species, frequently in watercolor paintings. They must be scientifically accurate but often also have an artistic component and may be printed with a botanical ...
*
History of phycology The history of phycology is the history of the scientific study of algae. Human interest in plants as food goes back into the origins of the species, and knowledge of algae can be traced back more than two thousand years. However, only in the last ...
*
List of botanists This is a list of botanists who have Wikipedia articles, in alphabetical order by surname. The List of botanists by author abbreviation is mostly a list of plant taxonomists because an author receives a standard abbreviation only when that aut ...
* List of botanists by author abbreviation


References


Bibliography


Books


History of science

* (see also The Jewel House) * * * * *


History of botany, agriculture and horticulture

* * * * * * * Fries, Robert Elias (1950). ''A short history of botany in Sweden''. Uppsala: Almqvist & Wiksells boktr. OCL
3954193
* ; originally published as * *
1999 reprint Google Books
* * * * * * * * * * * * * , see also * * * * * *


Antiquity

* * * *


British botany

* * * * * * * * * *


Cultural studies

* * * * * * * * * *


Botanical art and illustration

* * *


Historical sources

* * * * * * ** , i
Botany pp. 243–254
*


Bibliographic sources

* *


Articles

* * * * * * * * * * * * *


Websites

* * * * * ;National Library of Medicine * ** ** ** {{DEFAULTSORT:History Of Botany
Botany Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek w ...
Botany Botany, also called , plant biology or phytology, is the science of plant life and a branch of biology. A botanist, plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek w ...