History of botany
Encyclopedia
The history of botany examines the human effort to understand life on Earth by tracing the historical development of the discipline of botany
Botany
Botany, plant science, or plant biology is a branch of biology that involves the scientific study of plant life. Traditionally, botany also included the study of fungi, algae and viruses...

—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
Paleolithic
The Paleolithic Age, Era or Period, is a prehistoric period of human history distinguished by the development of the most primitive stone tools discovered , and covers roughly 99% of human technological prehistory...

 hunter-gatherer
Hunter-gatherer
A hunter-gatherer or forage society is one in which most or all food is obtained from wild plants and animals, in contrast to agricultural societies which rely mainly on domesticated species. Hunting and gathering was the ancestral subsistence mode of Homo, and all modern humans were...

s. The first written records of plants were made in the Neolithic Revolution
Neolithic Revolution
The Neolithic Revolution was the first agricultural revolution. It was the transition from hunting and gathering to agriculture and settlement. Archaeological data indicates that various forms of plants and animal domestication evolved independently in 6 separate locations worldwide circa...

 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, appears in the teachings of Aristotle's student Theophrastus
Theophrastus
Theophrastus , a Greek native of Eresos in Lesbos, was the successor to Aristotle in the Peripatetic school. He came to Athens at a young age, and initially studied in Plato's school. After Plato's death he attached himself to Aristotle. Aristotle bequeathed to Theophrastus his writings, and...

 at the Lyceum
Lyceum
The lyceum is a category of educational institution defined within the education system of many countries, mainly in Europe. The definition varies between countries; usually it is a type of secondary school.-History:...

 in ancient Athens in about 350 BC; this is considered the starting point for modern botany. In Europe, this early 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 herbal
Herbal
AThe use of a or an depends on whether or not herbal is pronounced with a silent h. herbal is "a collection of descriptions of plants put together for medicinal purposes." Expressed more elaborately — it is a book containing the names and descriptions of plants, usually with information on their...

s. In China and the Arab world, the Greco-Roman work on medicinal plants was preserved and extended.

In Europe the Renaissance
Renaissance
The Renaissance was a cultural movement that spanned roughly the 14th to the 17th century, beginning in Italy in the Late Middle Ages and later spreading to the rest of Europe. The term is also used more loosely to refer to the historical era, but since the changes of the Renaissance were not...

 of the 14th–17th centuries heralded a scientific revival during which botany gradually emerged from natural history
Natural history
Natural history is the scientific research of plants or animals, leaning more towards observational rather than experimental methods of study, and encompasses more research published in magazines than in academic journals. Grouped among the natural sciences, natural history is the systematic study...

 as an independent science, distinct from medicine and agriculture. Herbal
Herbal
AThe use of a or an depends on whether or not herbal is pronounced with a silent h. herbal is "a collection of descriptions of plants put together for medicinal purposes." Expressed more elaborately — it is a book containing the names and descriptions of plants, usually with information on their...

s were replaced by Flora
Flora
Flora is the plant life occurring in a particular region or time, generally the naturally occurring or indigenous—native plant life. The corresponding term for animals is fauna.-Etymology:...

s: books that described the native plants of local regions. The invention of the microscope
Microscope
A microscope is an instrument used to see objects that are too small for the naked eye. The science of investigating small objects using such an instrument is called microscopy...

 stimulated the study of plant anatomy
Plant anatomy
Plant anatomy or phytotomy is the general term for the study of the internal structure of plants. While originally it included plant morphology, which is the description of the physical form and external structure of plants, since the mid-20th century the investigations of plant anatomy are...

, and the first carefully designed experiments in plant physiology
Plant physiology
Plant physiology is a subdiscipline of botany concerned with the functioning, or physiology, of plants. Closely related fields include plant morphology , plant ecology , phytochemistry , cell biology, and molecular biology.Fundamental processes such as photosynthesis, respiration, plant nutrition,...

 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
Nomenclature
Nomenclature is a term that applies to either a list of names or terms, or to the system of principles, procedures and terms related to naming - which is the assigning of a word or phrase to a particular object or property...

, description, and classification
Biological classification
Biological classification, or scientific classification in biology, is a method to group and categorize organisms by biological type, such as genus or species. Biological classification is part of scientific taxonomy....

.

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
Economic botany can be very broadly defined as a study of relationships between plants and people. Economic botany contributes significantly to anthropology, biology, conservation, botany, and other fields of science...

 (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
Biogeography is the study of the distribution of species , organisms, and ecosystems in space and through geological time. Organisms and biological communities vary in a highly regular fashion along geographic gradients of latitude, elevation, isolation and habitat area...

 and ecology
Ecology
Ecology is the scientific study of the relations that living organisms have with respect to each other and their natural environment. Variables of interest to ecologists include the composition, distribution, amount , number, and changing states of organisms within and among ecosystems...

) through to the small scale of subjects like cell theory
Cell theory
Cell theory refers to the idea that cells are the basic unit of structure in every living thing. Development of this theory during the mid 17th century was made possible by advances in microscopy. This theory is one of the foundations of biology...

, molecular biology
Molecular biology
Molecular biology is the branch of biology that deals with the molecular basis of biological activity. This field overlaps with other areas of biology and chemistry, particularly genetics and biochemistry...

 and plant biochemistry
Biochemistry
Biochemistry, sometimes called biological chemistry, is the study of chemical processes in living organisms, including, but not limited to, living matter. Biochemistry governs all living organisms and living processes...

.

Introduction

Botany
Botany
Botany, plant science, or plant biology is a branch of biology that involves the scientific study of plant life. Traditionally, botany also included the study of fungi, algae and viruses...

 (Greek
Greek language
Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...

  - grass, fodder; Medieval Latin
Medieval Latin
Medieval Latin was the form of Latin used in the Middle Ages, primarily as a medium of scholarly exchange and as the liturgical language of the medieval Roman Catholic Church, but also as a language of science, literature, law, and administration. Despite the clerical origin of many of its authors,...

  – herb, plant) and zoology
Zoology
Zoology |zoölogy]]), is the branch of biology that relates to the animal kingdom, including the structure, embryology, evolution, classification, habits, and distribution of all animals, both living and extinct...

 are, historically, the core disciplines of biology
Biology
Biology is a natural science concerned with the study of life and living organisms, including their structure, function, growth, origin, evolution, distribution, and taxonomy. Biology is a vast subject containing many subdivisions, topics, and disciplines...

 whose history is closely associated with the natural sciences chemistry
Chemistry
Chemistry is the science of matter, especially its chemical reactions, but also its composition, structure and properties. Chemistry is concerned with atoms and their interactions with other atoms, and particularly with the properties of chemical bonds....

, physics
Physics
Physics is a natural science that involves the study of matter and its motion through spacetime, along with related concepts such as energy and force. More broadly, it is the general analysis of nature, conducted in order to understand how the universe behaves.Physics is one of the oldest academic...

 and geology
Geology
Geology is the science comprising the study of solid Earth, the rocks of which it is composed, and the processes by which it evolves. Geology gives insight into the history of the Earth, as it provides the primary evidence for plate tectonics, the evolutionary history of life, and past climates...

. 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
Natural history
Natural history is the scientific research of plants or animals, leaning more towards observational rather than experimental methods of study, and encompasses more research published in magazines than in academic journals. Grouped among the natural sciences, natural history is the systematic study...

 divided pure botany into three main streams morphology
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....

-classification
Biological classification
Biological classification, or scientific classification in biology, is a method to group and categorize organisms by biological type, such as genus or species. Biological classification is part of scientific taxonomy....

, anatomy
Anatomy
Anatomy is a branch of biology and medicine that is the consideration of the structure of living things. It is a general term that includes human anatomy, animal anatomy , and plant anatomy...

 and physiology
Physiology
Physiology is the science of the function of living systems. This includes how organisms, organ systems, organs, cells, and bio-molecules carry out the chemical or physical functions that exist in a living system. The highest honor awarded in physiology is the Nobel Prize in Physiology or...

 – that is, external form, internal structure, and functional operation. The most obvious topics in applied botany are horticulture
Horticulture
Horticulture is the industry and science of plant cultivation including the process of preparing soil for the planting of seeds, tubers, or cuttings. Horticulturists work and conduct research in the disciplines of plant propagation and cultivation, crop production, plant breeding and genetic...

, forestry
Forestry
Forestry is the interdisciplinary profession embracing the science, art, and craft of creating, managing, using, and conserving forests and associated resources in a sustainable manner to meet desired goals, needs, and values for human benefit. Forestry is practiced in plantations and natural stands...

 and agriculture
Agriculture
Agriculture is the cultivation of animals, plants, fungi and other life forms for food, fiber, and other products used to sustain life. Agriculture was the key implement in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that nurtured the...

 although there are many others like weed science
Invasive species
"Invasive species", or invasive exotics, is a nomenclature term and categorization phrase used for flora and fauna, and for specific restoration-preservation processes in native habitats, with several definitions....

, plant pathology, floristry
Floristry
Floristry is the general term used to describe production, commerce and trade in flowers. It encompasses flower care and handling, floral design or flower arranging, merchandising, and display and flower delivery. Wholesale florists sell bulk flowers and related supplies to professionals in the trade...

, pharmacognosy
Pharmacognosy
Pharmacognosy is the study of medicines derived from natural sources. The American Society of Pharmacognosy defines pharmacognosy as "the study of the physical, chemical, biochemical and biological properties of drugs, drug substances or potential drugs or drug substances of natural origin as well...

, economic botany
Economic botany
Economic botany can be very broadly defined as a study of relationships between plants and people. Economic botany contributes significantly to anthropology, biology, conservation, botany, and other fields of science...

 and ethnobotany
Ethnobotany
Ethnobotany is the scientific study of the relationships that exist between people and plants....

 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
Taxonomy
Taxonomy is the science of identifying and naming species, and arranging them into a classification. The field of taxonomy, sometimes referred to as "biological taxonomy", revolves around the description and use of taxonomic units, known as taxa...

, molecular biology
Molecular biology
Molecular biology is the branch of biology that deals with the molecular basis of biological activity. This field overlaps with other areas of biology and chemistry, particularly genetics and biochemistry...

, computer science
Computer science
Computer science or computing science is the study of the theoretical foundations of information and computation and of practical techniques for their implementation and application in computer systems...

 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
Phycology
Phycology is the scientific study of algae. Phycology is a branch of life science and often is regarded as a subdiscipline of botany....

 (algae
Algae
Algae are a large and diverse group of simple, typically autotrophic organisms, ranging from unicellular to multicellular forms, such as the giant kelps that grow to 65 meters in length. They are photosynthetic like plants, and "simple" because their tissues are not organized into the many...

), pteridology (fern
Fern
A fern is any one of a group of about 12,000 species of plants belonging to the botanical group known as Pteridophyta. Unlike mosses, they have xylem and phloem . They have stems, leaves, and roots like other vascular plants...

s), bryology
Bryology
Bryology is the branch of botany concerned with the scientific study of bryophytes . Bryophytes were first studied in detail in the 18th century...

 (moss
Moss
Mosses are small, soft plants that are typically 1–10 cm tall, though some species are much larger. They commonly grow close together in clumps or mats in damp or shady locations. They do not have flowers or seeds, and their simple leaves cover the thin wiry stems...

es and liverworts
Marchantiophyta
The Marchantiophyta are a division of bryophyte plants commonly referred to as hepatics or liverworts. Like other bryophytes, they have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of genetic information....

) and palaeobotany (fossil plants) and their histories are treated elsewhere (see side bar). To this list can be added mycology
Mycology
Mycology is the branch of biology concerned with the study of fungi, including their genetic and biochemical properties, their taxonomy and their use to humans as a source for tinder, medicinals , food and entheogens, as well as their dangers, such as poisoning or...

, the study of fungi, which were once treated as plants, but are now ranked as a unique kingdom.

Ancient and medieval knowledge

Nomad
Nomad
Nomadic people , commonly known as itinerants in modern-day contexts, are communities of people who move from one place to another, rather than settling permanently in one location. There are an estimated 30-40 million nomads in the world. Many cultures have traditionally been nomadic, but...

ic hunter-gatherer
Hunter-gatherer
A hunter-gatherer or forage society is one in which most or all food is obtained from wild plants and animals, in contrast to agricultural societies which rely mainly on domesticated species. Hunting and gathering was the ancestral subsistence mode of Homo, and all modern humans were...

 societies passed on, by oral tradition
Oral tradition
Oral tradition and oral lore is cultural material and traditions transmitted orally from one generation to another. The messages or testimony are verbally transmitted in speech or song and may take the form, for example, of folktales, sayings, ballads, songs, or chants...

, 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
Folk taxonomy
A folk taxonomy is a vernacular naming system, and can be contrasted with scientific taxonomy. Folk biological classification is the way peoples describe and organize their natural surroundings/the world around them, typically making generous use of form taxa like "shrubs", "bugs", "ducks",...

, 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
Neolithic Revolution
The Neolithic Revolution was the first agricultural revolution. It was the transition from hunting and gathering to agriculture and settlement. Archaeological data indicates that various forms of plants and animal domestication evolved independently in 6 separate locations worldwide circa...

 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
Domestication or taming is the process whereby a population of animals or plants, through a process of selection, becomes accustomed to human provision and control. In the Convention on Biological Diversity a domesticated species is defined as a 'species in which the evolutionary process has been...

 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
Staple food
A staple food is one that is eaten regularly and in such quantities that it constitutes a dominant portion of a diet, and that supplies a high proportion of energy and nutrient needs. Most people live on a diet based on one or more staples...

s were domesticated in prehistoric 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
Rice
Rice is the seed of the monocot plants Oryza sativa or Oryza glaberrima . As a cereal grain, it is the most important staple food for a large part of the world's human population, especially in East Asia, Southeast Asia, South Asia, the Middle East, and the West Indies...

 in East Asia, wheat
Wheat
Wheat is a cereal grain, originally from the Levant region of the Near East, but now cultivated worldwide. In 2007 world production of wheat was 607 million tons, making it the third most-produced cereal after maize and rice...

 and barley
Barley
Barley is a major cereal grain, a member of the grass family. It serves as a major animal fodder, as a base malt for beer and certain distilled beverages, and as a component of various health foods...

 in the Middle east, and maize
Maize
Maize known in many English-speaking countries as corn or mielie/mealie, is a grain domesticated by indigenous peoples in Mesoamerica in prehistoric times. The leafy stalk produces ears which contain seeds called kernels. Though technically a grain, maize kernels are used in cooking as a vegetable...

 in Central and South America. By Greco-Roman times popular food plants of today, including grape
Grape
A grape is a non-climacteric fruit, specifically a berry, that grows on the perennial and deciduous woody vines of the genus Vitis. Grapes can be eaten raw or they can be used for making jam, juice, jelly, vinegar, wine, grape seed extracts, raisins, molasses and grape seed oil. Grapes are also...

s, apple
Apple
The apple is the pomaceous fruit of the apple tree, species Malus domestica in the rose family . It is one of the most widely cultivated tree fruits, and the most widely known of the many members of genus Malus that are used by humans. Apple grow on small, deciduous trees that blossom in the spring...

s, fig
Ficus
Ficus is a genus of about 850 species of woody trees, shrubs, vines, epiphytes, and hemiepiphyte in the family Moraceae. Collectively known as fig trees or figs, they are native throughout the tropics with a few species extending into the semi-warm temperate zone. The Common Fig Ficus is a genus of...

s, and olive
Olive
The olive , Olea europaea), is a species of a small tree in the family Oleaceae, native to the coastal areas of the eastern Mediterranean Basin as well as northern Iran at the south end of the Caspian Sea.Its fruit, also called the olive, is of major agricultural importance in the...

s, were being listed as named varieties in early manuscripts. Botanical authority William Stearn 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 BCE, 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
Egypt
Egypt , officially the Arab Republic of Egypt, Arabic: , is a country mainly in North Africa, with the Sinai Peninsula forming a land bridge in Southwest Asia. Egypt is thus a transcontinental country, and a major power in Africa, the Mediterranean Basin, the Middle East and the Muslim world...

, China
China
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...

, Mesopotamia
Mesopotamia
Mesopotamia is a toponym for the area of the Tigris–Euphrates river system, largely corresponding to modern-day Iraq, northeastern Syria, southeastern Turkey and southwestern Iran.Widely considered to be the cradle of civilization, Bronze Age Mesopotamia included Sumer and the...

 and India
India
India , officially the Republic of India , is a country in South Asia. It is the seventh-largest country by geographical area, the second-most populous country with over 1.2 billion people, and the most populous democracy in the world...

. Botanical historian Alan Morton notes that agriculture was the occupation of the poor and uneducated, while medicine was the realm of socially influential shamans, priest
Priest
A priest is a person authorized to perform the sacred rites of a religion, especially as a mediatory agent between humans and deities. They also have the authority or power to administer religious rites; in particular, rites of sacrifice to, and propitiation of, a deity or deities...

s, apothecaries
Apothecary
Apothecary is a historical name for a medical professional who formulates and dispenses materia medica to physicians, surgeons and patients — a role now served by a pharmacist and some caregivers....

, magicians and physician
Physician
A physician is a health care provider who practices the profession of medicine, which is concerned with promoting, maintaining or restoring human health through the study, diagnosis, and treatment of disease, injury and other physical and mental impairments...

s, 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
Rigveda
The Rigveda is an ancient Indian sacred collection of Vedic Sanskrit hymns...

, a collection of Vedic Sanskrit
Vedic Sanskrit
Vedic Sanskrit is an old Indo-Aryan language. It is an archaic form of Sanskrit, an early descendant of Proto-Indo-Iranian. It is closely related to Avestan, the oldest preserved Iranian language...

 hymns from about 3700–3100 BP. Plants are divided into (trees), (herbs useful to humans) and (creepers), with further subdivisions. The sacred Hindu
Hindu
Hindu refers to an identity associated with the philosophical, religious and cultural systems that are indigenous to the Indian subcontinent. As used in the Constitution of India, the word "Hindu" is also attributed to all persons professing any Indian religion...

 text Atharvaveda
Atharvaveda
The Atharvaveda is a sacred text of Hinduism and one of the four Vedas, often called the "fourth Veda"....

 divides plants into eight classes: (spreading branches), (leaves with long clusters), (bushy plants), (which expands); (those with monopodial
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...

 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 Law book of Hindu
Hindu
Hindu refers to an identity associated with the philosophical, religious and cultural systems that are indigenous to the Indian subcontinent. As used in the Constitution of India, the word "Hindu" is also attributed to all persons professing any Indian religion...

s, which classifies plants into eight major categories. Elaborate taxonomies also occur in the Charaka Samhitā
Charaka Samhita
The ' is an early Ayurvedic text on internal medicine. It is believed to be the oldest of the three ancient treatises of Ayurveda...

, Sushruta Samhita
Sushruta Samhita
The Sushruta Samhita is a Sanskrit text, attributed to one Sushruta, foundational to Ayurvedic medicine , with innovative chapters on surgery....

 and Vaisesika.

Ancient China
In ancient China 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
Han Dynasty
The Han Dynasty was the second imperial dynasty of China, preceded by the Qin Dynasty and succeeded by the Three Kingdoms . It was founded by the rebel leader Liu Bang, known posthumously as Emperor Gaozu of Han. It was briefly interrupted by the Xin Dynasty of the former regent Wang Mang...

 (202 BC-220 AD) includes the notable work of the Huangdi Neijing and the famous pharmacologist Zhang Zhongjing
Zhang Zhongjing
Zhang Zhongjing , formal name Zhang Ji, was a Han Dynasty physician and one of the most eminent Chinese physicians during the later years of the Han Dynasty...

. There were also the 11th century scientists and statesmen Su Song
Su Song
Su Song was a renowned Chinese polymath who specialized himself as a statesman, astronomer, cartographer, horologist, pharmacologist, mineralogist, zoologist, botanist, mechanical and architectural engineer, poet, antiquarian, and ambassador of the Song Dynasty .Su Song was the engineer of a...

 and Shen Kuo
Shen Kuo
Shen Kuo or Shen Gua , style name Cunzhong and pseudonym Mengqi Weng , was a polymathic Chinese scientist and statesman of the Song Dynasty...

 who compiled learned treatises on natural history, emphasising herbal medicine.

Theophrastus and the origin of botanical science

Ancient Athens, of the 6th century BCE, was the busy trade centre at the confluence of Egypt
Egypt
Egypt , officially the Arab Republic of Egypt, Arabic: , is a country mainly in North Africa, with the Sinai Peninsula forming a land bridge in Southwest Asia. Egypt is thus a transcontinental country, and a major power in Africa, the Mediterranean Basin, the Middle East and the Muslim world...

ian, Mesopotamia
Mesopotamia
Mesopotamia is a toponym for the area of the Tigris–Euphrates river system, largely corresponding to modern-day Iraq, northeastern Syria, southeastern Turkey and southwestern Iran.Widely considered to be the cradle of civilization, Bronze Age Mesopotamia included Sumer and the...

n and Minoa
Minoa
Minoa is the name of several bronze-age settlements in the Aegean coasts, in Corfu and in Sicily. The original meaning of the word remains unknown, but it seems that there is a strong connection with the mythic king of Crete Minos, during the bronze-age Minoan civilization which flourished in Crete...

n cultures at the height of Greek colonisation of the Mediterranean. The philosophical thought of this period ranged freely through many subjects. Empedocles
Empedocles
Empedocles was a Greek pre-Socratic philosopher and a citizen of Agrigentum, a Greek city in Sicily. Empedocles' philosophy is best known for being the originator of the cosmogenic theory of the four Classical elements...

 (490–430 BCE) foreshadowed Darwinian evolutionary theory in a crude formulation of the mutability of species and natural selection
Natural selection
Natural selection is the nonrandom process by which biologic traits become either more or less common in a population as a function of differential reproduction of their bearers. It is a key mechanism of evolution....

. The physician Hippocrates
Hippocrates
Hippocrates of Cos or Hippokrates of Kos was an ancient Greek physician of the Age of Pericles , and is considered one of the most outstanding figures in the history of medicine...

 (460–370 BCE) 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 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
Theophrastus
Theophrastus , a Greek native of Eresos in Lesbos, was the successor to Aristotle in the Peripatetic school. He came to Athens at a young age, and initially studied in Plato's school. After Plato's death he attached himself to Aristotle. Aristotle bequeathed to Theophrastus his writings, and...

 of Eressus (Greek
Greek language
Greek is an independent branch of the Indo-European family of languages. Native to the southern Balkans, it has the longest documented history of any Indo-European language, spanning 34 centuries of written records. Its writing system has been the Greek alphabet for the majority of its history;...

: ; c. 371–287 BCE) who has been frequently referred to as the ”Father of Botany”. He was a student and close friend of Aristotle
Aristotle
Aristotle was a Greek philosopher and polymath, a student of Plato and teacher of Alexander the Great. His writings cover many subjects, including physics, metaphysics, poetry, theater, music, logic, rhetoric, linguistics, politics, government, ethics, biology, and zoology...

 (384–322 BCE) and succeeded him as head of the Lyceum
Lyceum
The lyceum is a category of educational institution defined within the education system of many countries, mainly in Europe. The definition varies between countries; usually it is a type of secondary school.-History:...

 (an educational establishment like a modern university) in Athens with its tradition of peripatetic 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
Christian Friedrich Heinrich Wimmer
Christian Friedrich Heinrich Wimmer was a German botanist and educator who was a native of Breslau.He worked as a Schulrat in Breslau, and was the author of several publications of Silesian flora. He specialized in the study of the willow genus Salix...

 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 I of Macedon
- Biography :Alexander was the son of Amyntas I and Queen Eurydice.According to Herodotus, he was unfriendly to Persia, and had the envoys of Darius I killed when they arrived at the court of his father during the Ionian Revolt...

 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
Historia Plantarum
Historia Plantarum is Latin and literally means History of Plants, although in reality it means something closer to "on plants" or "treatise on plants". There has been more than one book by this title....

(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
Manifesto
A manifesto is a public declaration of principles and intentions, often political in nature. Manifestos relating to religious belief are generally referred to as creeds. Manifestos may also be life stance-related.-Etymology:...

: “We must consider the distinctive characters and the general nature of plants from the point of view of their morphology
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....

, 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
Biological classification
Biological classification, or scientific classification in biology, is a method to group and categorize organisms by biological type, such as genus or species. Biological classification is part of scientific taxonomy....

 of plants and economic botany
Economic botany
Economic botany can be very broadly defined as a study of relationships between plants and people. Economic botany contributes significantly to anthropology, biology, conservation, botany, and other fields of science...

, examining the techniques of agriculture
Agriculture
Agriculture is the cultivation of animals, plants, fungi and other life forms for food, fiber, and other products used to sustain life. Agriculture was the key implement in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that nurtured the...

 (relationship of crops to soil, climate, water and habitat) and horticulture
Horticulture
Horticulture is the industry and science of plant cultivation including the process of preparing soil for the planting of seeds, tubers, or cuttings. Horticulturists work and conduct research in the disciplines of plant propagation and cultivation, crop production, plant breeding and genetic...

. 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
Crataegus
Crataegus , commonly called hawthorn or thornapple, is a large genus of shrubs and trees in the rose family, Rosaceae, native to temperate regions of the Northern Hemisphere in Europe, Asia and North America. The name hawthorn was originally applied to the species native to northern Europe,...

, Daucus
Daucus
Daucus is a worldwide genus of herbaceous plants of the family Apiaceae of which the best-known species is the cultivated carrot. Daucus genus of Umbelliferae Apiaceae, has about 25 species.- Features :...

and Asparagus
Asparagus
Asparagus officinalis is a spring vegetable, a flowering perennialplant species in the genus Asparagus. It was once classified in the lily family, like its Allium cousins, onions and garlic, but the Liliaceae have been split and the onion-like plants are now in the family Amaryllidaceae and...

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 plant
An annual plant is a plant that usually germinates, flowers, and dies in a year or season. True annuals will only live longer than a year if they are prevented from setting seed...

s, perennials and biennials
Biennial plant
A biennial plant is a flowering plant that takes two years to complete its biological lifecycle. In the first year the plant grows leaves, stems, and roots , then it enters a period of dormancy over the colder months. Usually the stem remains very short and the leaves are low to the ground, forming...

, they were also either monocotyledon
Monocotyledon
Monocotyledons, also known as monocots, are one of two major groups of flowering plants that are traditionally recognized, the other being dicotyledons, or dicots. Monocot seedlings typically have one cotyledon , in contrast to the two cotyledons typical of dicots...

s or dicotyledon
Dicotyledon
The dicotyledons, also known as dicots, are a group of flowering plants whose seed typically has two embryonic leaves or cotyledons. There are around 199,350 species within this group...

s and he also noticed the difference between determinate 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 CE by Pedanius Dioscorides
Pedanius Dioscorides
Pedanius Dioscorides was a Greek physician, pharmacologist and botanist, the author of a 5-volume encyclopedia about herbal medicine and related medicinal substances , that was widely read for more than 1,500 years.-Life:...

 (c. 40-90 CE) 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
Renaissance
The Renaissance was a cultural movement that spanned roughly the 14th to the 17th century, beginning in Italy in the Late Middle Ages and later spreading to the rest of Europe. The term is also used more loosely to refer to the historical era, but since the changes of the Renaissance were not...

 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
Cato the Elder
Marcus Porcius Cato was a Roman statesman, commonly referred to as Censorius , Sapiens , Priscus , or Major, Cato the Elder, or Cato the Censor, to distinguish him from his great-grandson, Cato the Younger.He came of an ancient Plebeian family who all were noted for some...

 (234–149 BCE), Varro
Marcus Terentius Varro
Marcus Terentius Varro was an ancient Roman scholar and writer. He is sometimes called Varro Reatinus to distinguish him from his younger contemporary Varro Atacinus.-Biography:...

 (116–27 BCE) and, in particular, Columella
Columella
Lucius Junius Moderatus Columella is the most important writer on agriculture of the Roman empire. Little is known of his life. He was probably born in Gades , possibly of Roman parents. After a career in the army , he took up farming...

 (4–70 CE) and Palladius
Rutilius Taurus Aemilianus Palladius
Rutilius Taurus Aemilianus Palladius, also known as Palladius Rutilius Taurus Aemilianus or just Palladius, was a Roman writer of the 4th century AD. He is principally known for his book on agriculture, Opus agriculturae, sometimes known as De re rustica.-Opus agriculturae:The Opus agriculturae is...

 (4th century CE). Roman encyclopaedist Pliny the Elder (23–79 CE) 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.

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
Middle Ages
The Middle Ages is a periodization of European history from the 5th century to the 15th century. The Middle Ages follows the fall of the Western Roman Empire in 476 and precedes the Early Modern Era. It is the middle period of a three-period division of Western history: Classic, Medieval and Modern...

, a period of disorganised feudalism and indifference to learning, 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 probably dates from about 300 BCE and describes about 334 plants classed as trees or shrubs, each with a common name and illustration. Between 100 and 1700 CE 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 CE was the Islamic Renaissance, 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. Kurd
Kürd
Kürd or Kyurd or Kyurt may refer to:*Kürd Eldarbəyli, Azerbaijan*Kürd Mahrızlı, Azerbaijan*Kürd, Goychay, Azerbaijan*Kürd, Jalilabad, Azerbaijan*Kürd, Qabala, Azerbaijan*Qurdbayram, Azerbaijan...

ish biologist Ābu Ḥanīfah Āḥmad ibn Dawūd Dīnawarī (828–896 CE) 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 philosopher and physician Ibn Sina (Avicenna
Avicenna
Abū ʿAlī al-Ḥusayn ibn ʿAbd Allāh ibn Sīnā , commonly known as Ibn Sīnā or by his Latinized name Avicenna, was a Persian polymath, who wrote almost 450 treatises on a wide range of subjects, of which around 240 have survived...

) (c. 980–1037 CE) was another influential figure, his The Canon of Medicine
The Canon of Medicine
The Canon of Medicine is an encyclopedia of Galenic medicine in five books compiled by Ibn Sīnā and completed in 1025. It presents a clear and organized summary of all the medical knowledge of the time...

being a landmark in the history of medicine treasured until the Enlightenment
Age of Enlightenment
The Age of Enlightenment was an elite cultural movement of intellectuals in 18th century Europe that sought to mobilize the power of reason in order to reform society and advance knowledge. It promoted intellectual interchange and opposed intolerance and abuses in church and state...

. In the early 13th century, the Andalusian
Al-Andalus
Al-Andalus was the Arabic name given to a nation and territorial region also commonly referred to as Moorish Iberia. The name describes parts of the Iberian Peninsula and Septimania governed by Muslims , at various times in the period between 711 and 1492, although the territorial boundaries...

-Arab
Arab
Arab people, also known as Arabs , are a panethnicity primarily living in the Arab world, which is located in Western Asia and North Africa. They are identified as such on one or more of genealogical, linguistic, or cultural grounds, with tribal affiliations, and intra-tribal relationships playing...

ian biologist Abu al-Abbas al-Nabati developed an early scientific method
Scientific method
Scientific method refers to a body of techniques for investigating phenomena, acquiring new knowledge, or correcting and integrating previous knowledge. To be termed scientific, a method of inquiry must be based on gathering empirical and measurable evidence subject to specific principles of...

 for botany, introducing empirical
Empirical
The word empirical denotes information gained by means of observation or experimentation. Empirical data are data produced by an experiment or observation....

 and experiment
Experiment
An experiment is a methodical procedure carried out with the goal of verifying, falsifying, or establishing the validity of a hypothesis. Experiments vary greatly in their goal and scale, but always rely on repeatable procedure and logical analysis of the results...

al techniques in the testing, description and identification of numerous materia medica
Materia medica
Materia medica is a Latin medical term for the body of collected knowledge about the therapeutic properties of any substance used for healing . The term 'materia medica' derived from the title of a work by the Ancient Greek physician Pedanius Dioscorides in the 1st century AD, De materia medica libre...

, and separating unverified reports from those supported by actual tests and observation
Observation
Observation is either an activity of a living being, such as a human, consisting of receiving knowledge of the outside world through the senses, or the recording of data using scientific instruments. The term may also refer to any data collected during this activity...

s. His student Ibn al-Baitar (circa, 1188–1248) was an eminent Arab
Arab
Arab people, also known as Arabs , are a panethnicity primarily living in the Arab world, which is located in Western Asia and North Africa. They are identified as such on one or more of genealogical, linguistic, or cultural grounds, with tribal affiliations, and intra-tribal relationships playing...

 scientist
Islamic science
Science in the medieval Islamic world, also known as Islamic science or Arabic science, is the science developed and practised in the Islamic world during the Islamic Golden Age . During this time, Indian, Iranian and especially Greek knowledge was translated into Arabic...

. His Kitab al-Jami fi al-Adwiya al-Mufrada was a pharmacopoeia
Pharmacopoeia
Pharmacopoeia, pharmacopeia, or pharmacopoea, , in its modern technical sense, is a book containing directions for the identification of samples and the preparation of compound medicines, and published by the authority of a government or a medical or pharmaceutical society.In a broader sense it is...

 describing 1400 species, 300 discovered by himself. Translated into Latin
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...

 in 1758 this was used in Europe until the early 19th century as a critical summing up of centuries of Arab pharmacology.

In India simple artificial plant classification systems of the Rigveda
Rigveda
The Rigveda is an ancient Indian sacred collection of Vedic Sanskrit hymns...

, Atharvaveda
Atharvaveda
The Atharvaveda is a sacred text of Hinduism and one of the four Vedas, often called the "fourth Veda"....

 and Taittiriya Samhita became more botanical with the work of Parashara
Parashara
' is a Rigvedic Maharishi and author of many ancient Indian texts. Parāśara was the grandson of Vashista, the son of Śakti Maharṣi, and the father of Vyasa. There are several texts which give reference to Parāśara as an author/speaker...

 (c.400–500 CE), the author of Vṛksayurveda (the science of life of trees). He made close observations of cells and leaves and divided plants into Dvimatrka (Dicotyledon
Dicotyledon
The dicotyledons, also known as dicots, are a group of flowering plants whose seed typically has two embryonic leaves or cotyledons. There are around 199,350 species within this group...

s) and Ekamatrka (Monocotyledon
Monocotyledon
Monocotyledons, also known as monocots, are one of two major groups of flowering plants that are traditionally recognized, the other being dicotyledons, or dicots. Monocot seedlings typically have one cotyledon , in contrast to the two cotyledons typical of dicots...

s). The dicotyledons were further classified into groupings (ganas) akin to modern floral families: Samiganiya (Fabaceae
Fabaceae
The Fabaceae or Leguminosae, commonly known as the legume, pea, or bean family, is a large and economically important family of flowering plants. The group is the third largest land plant family, behind only the Orchidaceae and Asteraceae, with 730 genera and over 19,400 species...

), Puplikagalniya (Rutaceae
Rutaceae
Rutaceae, commonly known as the rue or citrus family, is a family of flowering plants, usually placed in the order Sapindales.Species of the family generally have flowers that divide into four or five parts, usually with strong scents...

), Svastikaganiya (Cruciferae), Tripuspaganiya (Cucurbitaceae
Cucurbitaceae
The plant family Cucurbitaceae consists of various squashes, melons, and gourds, including crops such as cucumber, pumpkins, luffas, and watermelons...

), Mallikaganiya (Apocynaceae
Apocynaceae
The Apocynaceae or dogbane family is a family of flowering plants that includes trees, shrubs, herbs, and lianas.Many species are tall trees found in tropical rainforests, and most are from the tropics and subtropics, but some grow in tropical dry, xeric environments. There are also perennial herbs...

), and Kurcapuspaganiya (Asteraceae
Asteraceae
The Asteraceae or Compositae , is an exceedingly large and widespread family of vascular plants. The group has more than 22,750 currently accepted species, spread across 1620 genera and 12 subfamilies...

). Important medieval Indian works of plant physiology include the Prthviniraparyam of Udayana
Udayana
- Introduction :Udayana also known as Udayanācārya lived in the 10th century, in Kariyan village in Mithila, near present day Darbhanga, Bihar state, India. Udayana was a very important Hindu logician who attempted to reconcile the views held by the two major schools of logic...

, Nyayavindutika of Dharmottara, Saddarsana-samuccaya of Gunaratna, and Upaskara of Sankaramisra.

The Age of Herbals

In the European Middle Ages
Middle Ages
The Middle Ages is a periodization of European history from the 5th century to the 15th century. The Middle Ages follows the fall of the Western Roman Empire in 476 and precedes the Early Modern Era. It is the middle period of a three-period division of Western history: Classic, Medieval and Modern...

 of the 15th and 16th centuries the lives of European citizens were based around agriculture but when printing arrived, with movable type and woodcut
Woodcut
Woodcut—occasionally known as xylography—is a relief printing artistic technique in printmaking in which an image is carved into the surface of a block of wood, with the printing parts remaining level with the surface while the non-printing parts are removed, typically with gouges...

 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 herbal
Herbal
AThe use of a or an depends on whether or not herbal is pronounced with a silent h. herbal is "a collection of descriptions of plants put together for medicinal purposes." Expressed more elaborately — it is a book containing the names and descriptions of plants, usually with information on their...

s 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. However, the need for accurate and detailed plant descriptions meant that some herbals were more botanical than medicinal. German Otto Brunfels
Otto Brunfels
Otto Brunfels was a German theologian and botanist...

'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
Hieronymus Bock 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 relation or resemblance....

'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
Valerius Cordus was a German physician and botanist who authored one of the greatest pharmacopoeias and one of the most celebrated herbals in history...

 (1515–1544) who pioneered the formal botanical description that detailed both flowers and fruits, some anatomy including the number of chambers in the ovary
Ovary
The ovary is an ovum-producing reproductive organ, often found in pairs as part of the vertebrate female reproductive system. Ovaries in anatomically female individuals are analogous to testes in anatomically male individuals, in that they are both gonads and endocrine glands.-Human anatomy:Ovaries...

, and the type of ovule
Ovule
Ovule means "small egg". In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: The integument forming its outer layer, the nucellus , and the megaspore-derived female gametophyte in its center...

 placentation
Placentation
In biology, placentation refers to the formation, type and structure, or arrangement of placentas. The function of placentation is to transfer nutrients from maternal tissue to a growing embryo...

. He also made observations on pollen and distinguished between inflorescence
Inflorescence
An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Strictly, it is the part of the shoot of seed plants where flowers are formed and which is accordingly modified...

 types. His five-volume Historia Plantarum was published about 18 years after his early death aged 29 in 1561-1563. In Holland Rembert Dodoens
Rembert Dodoens
Rembert Dodoens was a Flemish physician and botanist, also known under his Latinized name Rembertus Dodonaeus.-Biography:...

 (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
Flora is the plant life occurring in a particular region or time, generally the naturally occurring or indigenous—native plant life. The corresponding term for animals is fauna.-Etymology:...

s. These were often backed by specimens deposited in a herbarium
Herbarium
In botany, a herbarium – sometimes known by the Anglicized term herbar – is a collection of preserved plant specimens. These specimens may be whole plants or plant parts: these will usually be in a dried form, mounted on a sheet, but depending upon the material may also be kept in...

 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.

1550–1800 The Renaissance and Enlightenment

The revival of learning during the European Renaissance
Renaissance
The Renaissance was a cultural movement that spanned roughly the 14th to the 17th century, beginning in Italy in the Late Middle Ages and later spreading to the rest of Europe. The term is also used more loosely to refer to the historical era, but since the changes of the Renaissance were not...

 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
East Indies
East Indies is a term used by Europeans from the 16th century onwards to identify what is now known as Indian subcontinent or South Asia, Southeastern Asia, and the islands of Oceania, including the Malay Archipelago and the Philippines...

 and the New World
New World
The New World is one of the names used for the Western Hemisphere, specifically America and sometimes Oceania . The term originated in the late 15th century, when America had been recently discovered by European explorers, expanding the geographical horizon of the people of the European middle...

.

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
Chelsea Physic Garden
The Chelsea Physic Garden was established as the Apothecaries’ Garden in London, England in 1673. It is the second oldest botanical garden in Britain, after the University of Oxford Botanic Garden, which was founded in 1621.Its rock garden is the oldest English garden devoted to alpine plants...

, Royal Botanic Gardens Kew, and the Oxford and Cambridge Botanic Gardens, 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
Age of Enlightenment
The Age of Enlightenment was an elite cultural movement of intellectuals in 18th century Europe that sought to mobilize the power of reason in order to reform society and advance knowledge. It promoted intellectual interchange and opposed intolerance and abuses in church and state...

 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.

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
Orto botanico di Pisa
The Orto botanico di Pisa, also known as the Orto Botanico dell'Università di Pisa, is a botanical garden operated by the University of Pisa, and located at via Luca Ghini 5, Pisa, Italy...

 (1544), founded by Luca Ghini
Luca Ghini
Luca Ghini was an Italian physician and botanist, notable as the creator of the first recorded herbarium, as well as the first botanical garden in Europe....

 (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. 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
Herbarium
In botany, a herbarium – sometimes known by the Anglicized term herbar – is a collection of preserved plant specimens. These specimens may be whole plants or plant parts: these will usually be in a dried form, mounted on a sheet, but depending upon the material may also be kept in...

 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. This approach coupled with the new Linnaean system of binomial nomenclature
Binomial nomenclature
Binomial nomenclature is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin grammatical forms, although they can be based on words from other languages...

 resulted in plant encyclopaedias without medicinal information called Flora
Flora
Flora is the plant life occurring in a particular region or time, generally the naturally occurring or indigenous—native plant life. The corresponding term for animals is fauna.-Etymology:...

s
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
Nehemiah Grew was an English plant anatomist and physiologist, very famously known as the "Father of Plant Physiology"...

 and Italian Marcello Malpighi
Marcello Malpighi
Marcello Malpighi was an Italian doctor, who gave his name to several physiological features, like the Malpighian tubule system.-Early years:...

, would last for 150 years.

Botanical exploration

Konrad Gessner discovered many new plants while climbing the Swiss Alps
Swiss Alps
The Swiss Alps are the portion of the Alps mountain range that lies within Switzerland. Because of their central position within the entire Alpine range, they are also known as the Central Alps....

. He proposed that there were groups or genera of plants. He said that each genera 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. Clusius
Clusius
Clusius may refer to:* Carolus Clusius, a Flemish doctor and botanist* Clusius , a hybrid elm cultivar...

 journeyed throughout most of Western Europe
Western Europe
Western Europe is a loose term for the collection of countries in the western most region of the European continents, though this definition is context-dependent and carries cultural and political connotations. One definition describes Western Europe as a geographic entity—the region lying in the...

, making discoveries in the vegetable kingdom along the way. He was the first to propose dividing plants into classes.

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
Plant collecting
Plant collecting involves procuring live or dried plant specimens, for the purposes of research, cultivation or as a hobby.-Collection of live specimens:...

 and gardener-botanists. Significant botanical collections came from: the West Indies (Hans Sloane
Hans Sloane
Sir Hans Sloane, 1st Baronet, PRS was an Ulster-Scot physician and collector, notable for bequeathing his collection to the British nation which became the foundation of the British Museum...

 (1660–1753)); China (James Cunningham); the spice islands of the East Indies (Moluccas, George Rumphius
Georg Eberhard Rumphius
Georg Eberhard Rumphius or originally Rumpf was a German-born botanist employed by the Dutch East India Company in what is now eastern Indonesia, and is best known for his work, Herbarium Amboinense....

 (1627–1702)); China and Mozambique (João de Loureiro
João de Loureiro
João de Loureiro was a Portuguese Jesuit missionary, paleontologist, physician and botanist.In 1742 he travelled to Cochinchina, remaining there for 30 years. He became a specialist in Asian flora and on his return published Flora Cochinchinensis .- References :...

 (1717–1791)); West Africa (Michel Adanson
Michel Adanson
Michel Adanson was a French naturalist of Scottish descent.Adanson was born at Aix-en-Provence. His family moved to Paris on 1730. After leaving the College Sainte Barbe he was employed in the cabinets of R. A. F. Reaumur and Bernard de Jussieu, as well as in the Jardin des Plantes. At the end of...

 (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's chief botanist Joseph Banks
Joseph Banks
Sir Joseph Banks, 1st Baronet, GCB, PRS was an English naturalist, botanist and patron of the natural sciences. He took part in Captain James Cook's first great voyage . Banks is credited with the introduction to the Western world of eucalyptus, acacia, mimosa and the genus named after him,...

 (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
Phylogenetics
In biology, phylogenetics is the study of evolutionary relatedness among groups of organisms , which is discovered through molecular sequencing data and morphological data matrices...

.

Italian physician Andrea Caesalpino (1519–1603) studied medicine and taught botany at the University of Pisa
University of Pisa
The University of Pisa , located in Pisa, Tuscany, is one of the oldest universities in Italy. It was formally founded on September 3, 1343 by an edict of Pope Clement VI, although there had been lectures on law in Pisa since the 11th century...

 for about 40 years eventually becoming Director of the Botanic Garden of Pisa
Orto botanico di Pisa
The Orto botanico di Pisa, also known as the Orto Botanico dell'Università di Pisa, is a botanical garden operated by the University of Pisa, and located at via Luca Ghini 5, Pisa, Italy...

 from 1554 to 1558. His sixteen-volume De Plantis (1583) described 1500 plants and his herbarium
Herbarium
In botany, a herbarium – sometimes known by the Anglicized term herbar – is a collection of preserved plant specimens. These specimens may be whole plants or plant parts: these will usually be in a dried form, mounted on a sheet, but depending upon the material may also be kept in...

 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
Gaspard Bauhin
Gaspard Bauhin, or Caspar Bauhin , was a Swiss botanist who wrote Pinax theatri botanici , which described thousands of plants and classified them in a manner that draws comparisons to the later binomial nomenclature of Linnaeus...

 (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 (1587–1657) compiled a much-needed botanical terminology which has stood the test of time. English botanist John Ray
John Ray
John Ray was an English naturalist, sometimes referred to as the father of English natural history. Until 1670, he wrote his name as John Wray. From then on, he used 'Ray', after "having ascertained that such had been the practice of his family before him".He published important works on botany,...

 (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 text book 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
Pierre Magnol was a French botanist. He was born in the city of Montpellier, where he lived and worked for the biggest part of his life. He eventually became Professor of Botany and Director of the Royal Botanic Garden of Montpellier and even held a seat in the Académie Royale des Sciences de...

 (1638–1715) and Joseph de Tournefort
Joseph Pitton de Tournefort
Joseph Pitton de Tournefort was a French botanist, notable as the first to make a clear definition of the concept of genus for plants.- Biography :...

 (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
Systema Naturae
The book was one of the major works of the Swedish botanist, zoologist and physician Carolus Linnaeus. The first edition was published in 1735...

 (1735), Genera Plantarum
Genera Plantarum
Genera Plantarum is a publication of Swedish naturalist Carl Linnaeus . The fifth edition served as a complementary volume to Species Plantarum . Article 13 of the International Code of Botanical Nomenclature states that "It is agreed to associate generic names which first appear in Linnaeus'...

 (1737), and Philosophia Botanica
Philosophia Botanica
Philosophia Botanica was published by the Swedish naturalist and physician Carolus Linnaeus who greatly influenced the development of botanical taxonomy and systematics in the 18th and 19th centuries. It is "the first textbook of descriptive systematic botany and botanical Latin"...

 (1751) but it was in his Species Plantarum
Species Plantarum
Species Plantarum was first published in 1753, as a two-volume work by Carl Linnaeus. Its prime importance is perhaps that it is the primary starting point of plant nomenclature as it exists today. This means that the first names to be considered validly published in botany are those that appear...

 (1753) that he gave every species a binomial
Binomial nomenclature
Binomial nomenclature is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin grammatical forms, although they can be based on words from other languages...

 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
Bernard de Jussieu was a French naturalist, younger brother of Antoine de Jussieu.Bernard de Jussieu was born in Lyon...

 (1699–1777) whose nephew Antoine-Laurent de Jussieu (1748–1836) extended it yet again to include about 100 orders (present-day families). Frenchman Michel Adanson
Michel Adanson
Michel Adanson was a French naturalist of Scottish descent.Adanson was born at Aix-en-Provence. His family moved to Paris on 1730. After leaving the College Sainte Barbe he was employed in the cabinets of R. A. F. Reaumur and Bernard de Jussieu, as well as in the Jardin des Plantes. At the end of...

 (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
Microscope
A microscope is an instrument used to see objects that are too small for the naked eye. The science of investigating small objects using such an instrument is called microscopy...

 was invented in 1590 it was only in the late 17th century that lens grinding by Antony van Leeuwenhoek provided the resolution needed to make major discoveries. Important general biological observations were made by Robert Hooke
Robert Hooke
Robert Hooke FRS was an English natural philosopher, architect and polymath.His adult life comprised three distinct periods: as a scientific inquirer lacking money; achieving great wealth and standing through his reputation for hard work and scrupulous honesty following the great fire of 1666, but...

 (1635–1703) but the foundations of plant anatomy were laid by Italian Marcello Malpighi
Marcello Malpighi
Marcello Malpighi was an Italian doctor, who gave his name to several physiological features, like the Malpighian tubule system.-Early years:...

 (1628–1694) of the University of Bologna in his (1675) and Royal Society Englishman Nehemiah Grew
Nehemiah Grew
Nehemiah Grew was an English plant anatomist and physiologist, very famously known as the "Father of Plant Physiology"...

 (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
Parenchyma is a term used to describe a bulk of a substance. It is used in different ways in animals and in plants.The term is New Latin, f. Greek παρέγχυμα - parenkhuma, "visceral flesh", f. παρεγχεῖν - parenkhein, "to pour in" f. para-, "beside" + en-, "in" + khein, "to pour"...

 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
Jan Baptist van Helmont
Jan Baptist van Helmont was an early modern period Flemish chemist, physiologist, and physician. He worked during the years just after Paracelsus and iatrochemistry, and is sometimes considered to be "the founder of pneumatic chemistry"...

 (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
Stephen Hales, FRS was an English physiologist, chemist and inventor.Hales studied the role of air and water in the maintenance of both plant and animal life. He gave accurate accounts of the movements of water in plants, and demonstrated that plants absorb air...

 (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 Priestly (1733–1804) is noted for his discovery of oxygen (as now called) and its production by plants. Later Jan Ingenhousz
Jan Ingenhousz
Jan Ingenhousz or Ingen-Housz FRS was a Dutch physiologist, biologist and chemist. He is best known for showing that light is essential to photosynthesis and thus having discovered photosynthesis. He also discovered that plants, like animals, have cellular respiration...

 (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
Nicolas-Théodore de Saussure
Nicolas-Théodore de Saussure was a Swiss chemist and student of plant physiology who made seminal advances in phytochemistry....

'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 analysed in chemical detail from plant ash) have a powerful influence on plant growth.

Plant sexuality

It was Rudolf Camerarius
Rudolf Jakob Camerarius
Rudolf Jakob Camerarius or Camerer was a German botanist and physician.Camerarius was born at Tübingen, and became professor of medicine and director of the botanical gardens at Tübingen in 1687...

 (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 Hofmeister
Wilhelm Friedrich Benedikt Hofmeister 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 largely self-taught....

 (1824–1877) starting with the ferns and bryophytes demonstrated that the process of sexual reproduction in plants entails an “alternation of generations” between sporophyte
Sporophyte
All land plants, and some algae, have life cycles in which a haploid gametophyte generation alternates with a diploid sporophyte, the generation of a plant or algae that has a double set of chromosomes. A multicellular sporophyte generation or phase is present in the life cycle of all land plants...

s and gametophyte
Gametophyte
A gametophyte is the haploid, multicellular phase of plants and algae that undergo alternation of generations, with each of its cells containing only a single set of chromosomes....

s. This initiated the new field of comparative morphology which, largely through the combined work of William Farlow
William Gilson Farlow
William Gilson Farlow was an American botanist, born in Boston, Massachusetts, and educated at Harvard , where, after several years of European study, he became adjunct professor of botany in 1874 and professor of cryptogamic botany in 1879.In 1899 he was president of the American Society of...

 (1844–1919), Nathanael Pringsheim
Nathanael Pringsheim
Nathanael Pringsheim was a German botanist.-Biography:Nathanael Pringsheim was born at Landsberg, Prussian Silesia, and studied at the universities of Breslau, Leipzig, and Berlin successively...

 (1823–1894), Frederick Bower
Frederick Orpen Bower
Frederick Orpen Bower FRS was a British botanist. He was elected a Fellow of the Royal Society in 1891. He was awarded the Gold Medal of the Linnean Society in 1909 and the Darwin Medal of the Royal Society in 1938....

, Eduard Strasburger
Eduard Strasburger
Eduard Adolf Strasburger was a German professor who was one of the most famous botanists of the 19th century....

 and others, established that an "alternation of generations" occurs throughout the plant kingdom.

Some time later the German academic and natural historian Joseph Kölreuter
Joseph Gottlieb Kölreuter
Joseph Gottlieb Kölreuter , also spelled Koelreuter or Kohlreuter, was a German botanist.Kölreuter was born the son of a pharmacist in Karlsruhe, Germany, and grew up in Sulz. He studied medicine at the University of Tübingen under physician and botanist Johann Georg Gmelin, receiving his PhD in...

 (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
Christian Konrad Sprengel
Christian Konrad Sprengel was a German theologist, teacher and, most importantly, a naturalist. He is most famously known for his research into plant sexuality....

 (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” text book, Matthias Schleiden
Matthias Jakob Schleiden
Matthias Jakob Schleiden was a German botanist and co-founder of the cell theory, along with Theodor Schwann and Rudolf Virchow....

'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 (1778–1841) succeeded Antoine-Laurent de Jussieu
Antoine de Jussieu
Antoine de Jussieu was a French naturalist.Jussieu was born in Lyon, the son of Christophe de Jussieu , an apothecary of some repute, who published a Nouveau traité de la theriaque . Antoine studied at the university of Montpellier, and travelled with his brother Bernard through Spain, Portugal...

 in managing the botanical project (1824–1841) which involved 35 authors: it contained all the dicotyledons known in his day, some 58000 species in 161 families, and he doubled the number of recognized plant families, the work being completed by his son Alphonse (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
Carl Ludwig Willdenow
Carl Ludwig Willdenow was a German botanist, pharmacist, and plant taxonomist. He is considered one of the founders of phytogeography, the study of the geographic distribution of plants...

 (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" and "centre of origin”. German Alexander von Humbolt (1769–1859) and Frenchman Aime Bonpland
Aimé Bonpland
Aimé Jacques Alexandre Bonpland was a French explorer and botanist.Bonpland's real name was Goujaud, and he was born in La Rochelle, a coastal city in France. After serving as a surgeon in the French army, and studying under J. N...

 (1773–1858) published a massive and highly influential 30 volume work on their travels; Robert Brown
Robert Brown (botanist)
Robert Brown was a Scottish botanist and palaeobotanist who made important contributions to botany largely through his pioneering use of the microscope...

 (1773–1852) noted the similarities between the floras of S Africa, Australia and India, while Joakim Schouw
Joakim Frederik Schouw
Joakim Frederik Schouw was a Danish lawyer, botanist and politician. From 1821, professor in botany at the University of Copenhagen - first extraordinary professor, but after the death of J.W. Hornemann in 1841 ordinary...

 (1789–1852) explored more deeply than anyone else the influence on plant distribution of temperature, soil
Edaphic
Edaphic is a nature related to soil. Edaphic qualities may characterize the soil itself, including drainage, texture, or chemical properties such as pH. Edaphic may also characterize organisms, such as plant communities, where it specifies their relationships with soil...

 factors, especially soil water, and light, work that was continued by Alphonse de Candolle (1806–1893). Joseph Hooker
Joseph Dalton Hooker
Sir Joseph Dalton Hooker OM, GCSI, CB, MD, FRS was one of the greatest British botanists and explorers of the 19th century. Hooker was a founder of geographical botany, and Charles Darwin's closest friend...

 (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 Grisebach
August Heinrich Rudolf Grisebach was a German botanist and phytogeographer. Born in Hannover on April 17, 1814, he died at Göttingen on May 9, 1879.- Biography :...

 (1814–1879) in (1872) examined physiognomy
Physiognomy
Physiognomy is the assessment of a person's character or personality from their outer appearance, especially the face...

 in relation to climate and in America geographic studies were pioneered by Asa Gray
Asa Gray
-References:*Asa Gray. Dictionary of American Biography. American Council of Learned Societies, 1928–1936.*Asa Gray. Encyclopedia of World Biography, 2nd ed. 17 Vols. Gale Research, 1998.*Asa Gray. Plant Sciences. 4 vols. Macmillan Reference USA, 2001....

 (1810–1888).

Physiological plant geography, perhaps more familiarly termed ecology
Ecology
Ecology is the scientific study of the relations that living organisms have with respect to each other and their natural environment. Variables of interest to ecologists include the composition, distribution, amount , number, and changing states of organisms within and among ecosystems...

, emerged out of 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
Eugenius Warming
Johannes Eugenius Bülow Warming , known as Eugen Warming, was a Danish botanist and a main founding figure of the scientific discipline of ecology...

 (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
Andreas Franz Wilhelm Schimper was a botanist and phytogeographer who made major contributions in the fields of histology, ecology and plant geography.-Biography:...

 (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
Johann Jacob Paul Moldenhawer
Johann Jacob Paul Moldenhawer was a German botanist who made a number of important discoveries in plant anatomy.He was born in Hamburg, the son of a minister, and started out studying theology and the classics...

 (1766–1827) published (1812) in which he describes techniques for separating cells from the middle lamella
Middle lamella
The middle lamella is a pectin layer which cements the cell walls of two adjoining cells together. Plants need this to give them stability and so that they can form plasmodesmata between the cells. It is the first formed layer which is deposited at the time of cytokinesis. The cell plate that is...

. He identified vascular
Vascular tissue
Vascular tissue is a complex conducting tissue, formed of more than one cell type, found in vascular plants. The primary components of vascular tissue are the xylem and phloem. These two tissues transport fluid and nutrients internally. There are also two meristems associated with vascular tissue:...

 and parenchyma
Parenchyma
Parenchyma is a term used to describe a bulk of a substance. It is used in different ways in animals and in plants.The term is New Latin, f. Greek παρέγχυμα - parenkhuma, "visceral flesh", f. παρεγχεῖν - parenkhein, "to pour in" f. para-, "beside" + en-, "in" + khein, "to pour"...

tous tissues, described vascular bundles, observed the cells in the cambium
Vascular cambium
The vascular cambium is a part of the morphology of plants. It consists of cells that are partly specialized, for the tissues that transport water solutions, but have not reached any of the final forms that occur in their branch of the specialization graph...

, and interpreted tree rings. He found that stoma
Stoma
In botany, a stoma is a pore, found in the leaf and stem epidermis that is used forgas exchange. The pore is bordered by a pair of specialized parenchyma cells known as guard cells that are responsible for regulating the size of the opening...

ta were composed of pairs of cells, rather than a single cell with a hole.

Anatomical studies on the stele
Stele
A stele , also stela , is a stone or wooden slab, generally taller than it is wide, erected for funerals or commemorative purposes, most usually decorated with the names and titles of the deceased or living — inscribed, carved in relief , or painted onto the slab...

 were consolidated by Carl Sanio (1832–1891) who described the secondary tissues and meristem
Meristem
A meristem is the tissue in most plants consisting of undifferentiated cells , found in zones of the plant where growth can take place....

 including cambium
Meristem
A meristem is the tissue in most plants consisting of undifferentiated cells , found in zones of the plant where growth can take place....

 and its action. Hugo von Mohl
Hugo von Mohl
Hugo von Mohl was a German botanist from Stuttgart.He was a son of the Württemberg statesman Benjamin Ferdinand von Mohl , the family being connected on both sides with the higher class of state officials of Württemberg...

 (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 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 Karl Nägeli
Karl Wilhelm von Nägeli
Karl Wilhelm von Nägeli 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.-Birth and education:...

 (1817–1891). Studies had also begun on the origins of the carpel and flower
Flower
A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants . The biological function of a flower is to effect reproduction, usually by providing a mechanism for the union of sperm with eggs...

 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
Osmosis
Osmosis is the movement of solvent molecules through a selectively permeable membrane into a region of higher solute concentration, aiming to equalize the solute concentrations on the two sides...

 by Frenchman Henri Dutrochet
Henri Dutrochet
René Joachim Henri Dutrochet was a French physician, botanist and physiologist. He is best known for his investigation into osmosis.-Early Career:...

 (1776–1847).

Cytology

The cell nucleus was discovered by Robert Brown
Robert Brown (botanist)
Robert Brown was a Scottish botanist and palaeobotanist who made important contributions to botany largely through his pioneering use of the microscope...

 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
Theodor Schwann
Theodor Schwann was a German physiologist. His many contributions to biology include the development of cell theory, the discovery of Schwann cells in the peripheral nervous system, the discovery and study of pepsin, the discovery of the organic nature of yeast, and the invention of the term...

 (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
Julius von Sachs was a German botanist from Breslau, Prussian Silesia.At an early age he showed a taste for natural history, becoming acquainted with the Breslau physiologist Jan Evangelista Purkyně. In 1851 he began studying at Charles University in Prague...

 had shown the continuity protoplasm
Protoplasm
Protoplasm is the living contents of a cell that is surrounded by a plasma membrane. It is a general term of the Cytoplasm . Protoplasm is composed of a mixture of small molecules such as ions, amino acids, monosaccharides and water, and macromolecules such as nucleic acids, proteins, lipids and...

 between cell wall
Cell wall
The cell wall is the tough, usually flexible but sometimes fairly rigid layer that surrounds some types of cells. It is located outside the cell membrane and provides these cells with structural support and protection, and also acts as a filtering mechanism. A major function of the cell wall is to...

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 chromosome
Chromosome
A chromosome is an organized structure of DNA and protein found in cells. It is a single piece of coiled DNA containing many genes, regulatory elements and other nucleotide sequences. Chromosomes also contain DNA-bound proteins, which serve to package the DNA and control its functions.Chromosomes...

s 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 that the nuclei of reproductive cells (in pollen and embryo) have a reducing division (halving of chromosomes, now known as meiosis
Meiosis
Meiosis is a special type of cell division necessary for sexual reproduction. The cells produced by meiosis are gametes or spores. The animals' gametes are called sperm and egg cells....

) the field of heredity was opened up. By 1926 Thomas Morgan
Thomas Hunt Morgan
Thomas Hunt Morgan was an American evolutionary biologist, geneticist and embryologist and science author who won the Nobel Prize in Physiology or Medicine in 1933 for discoveries relating the role the chromosome plays in heredity.Morgan received his PhD from Johns Hopkins University in zoology...

 was able to outline a theory of the gene
Gene
A gene is a molecular unit of heredity of a living organism. It is a name given to some stretches of DNA and RNA that code for a type of protein or for an RNA chain that has a function in the organism. Living beings depend on genes, as they specify all proteins and functional RNA chains...

 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 plastid
Plastid
Plastids are major organelles found in the cells of plants and algae. Plastids are the site of manufacture and storage of important chemical compounds used by the cell...

s and vacuole
Vacuole
A vacuole is a membrane-bound organelle which is present in all 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 molecules including enzymes in solution, though in certain...

s – what is now known as cytology
Cell biology
Cell biology is a scientific discipline that studies cells – their physiological properties, their structure, the organelles they contain, interactions with their environment, their life cycle, division and death. This is done both on a microscopic and molecular level...

, or cell theory
Cell theory
Cell theory refers to the idea that cells are the basic unit of structure in every living thing. Development of this theory during the mid 17th century was made possible by advances in microscopy. This theory is one of the foundations of biology...

 became firmly established.

Later, the cytological basis of the gene-chromosome theory of heredity
Heredity
Heredity is the passing of traits to offspring . This is the process by which an offspring cell or organism acquires or becomes predisposed to the characteristics of its parent cell or organism. Through heredity, variations exhibited by individuals can accumulate and cause some species to evolve...

 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
Polyploidy
Polyploid is a term used to describe cells and organisms containing more than two paired sets of chromosomes. Most eukaryotic species are diploid, meaning they have two sets of chromosomes — one set inherited from each parent. However polyploidy is found in some organisms and is especially common...

 and hybridization in speciation
Speciation
Speciation is the evolutionary process by which new biological species arise. The biologist Orator F. Cook seems to have been the first to coin the term 'speciation' for the splitting of lineages or 'cladogenesis,' as opposed to 'anagenesis' or 'phyletic evolution' occurring within lineages...

 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 Darwin
Charles Robert Darwin FRS was an English naturalist. He established that all species of life have descended over time from common ancestry, and proposed the scientific theory that this branching pattern of evolution resulted from a process that he called natural selection.He published his theory...

’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 Hofmeister
Wilhelm Friedrich Benedikt Hofmeister 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 largely self-taught....

 established that there was a similar pattern of organization in all plants expressed through the alternation of generations
Alternation of generations
Alternation of generations is a term primarily used in describing the life cycle of plants . A multicellular sporophyte, which is diploid with 2N paired chromosomes , alternates with a multicellular gametophyte, which is haploid with N unpaired chromosomes...

 and extensive homology
Homology (biology)
Homology forms the basis of organization for comparative biology. In 1843, Richard Owen defined homology as "the same organ in different animals under every variety of form and function". Organs as different as a bat's wing, a seal's flipper, a cat's paw and a human hand have a common underlying...

 of structures.

Polymath
Polymath
A polymath is a person whose expertise spans a significant number of different subject areas. In less formal terms, a polymath may simply be someone who is very knowledgeable...

 German intellect Johann Goethe
Johann Wolfgang von Goethe
Johann Wolfgang von Goethe was a German writer, pictorial artist, biologist, theoretical physicist, and polymath. He is considered the supreme genius of modern German literature. His works span the fields of poetry, drama, prose, philosophy, and science. His Faust has been called the greatest long...

 (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 Braun
Alexander Carl Heinrich Braun was a German botanist from Regensburg, Bavaria.He studied botany in Heidelberg, Paris and Munich. In 1833 he began teaching botany at the Polytechnic School of Karlsruhe, staying there until 1846...

 (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 (1778–1841).

Carbon fixation (photosynthesis)

At the start of the 19th century the idea that plants could synthesise 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
Julius Robert von Mayer
Julius Robert von Mayer was a German physician and physicist and one of the founders of thermodynamics...

, 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, nitrification
Nitrification
Nitrification is the biological oxidation of ammonia with oxygen into nitrite followed by the oxidation of these nitrites into nitrates. Degradation of ammonia to nitrite is usually the rate limiting step of nitrification. Nitrification is an important step in the nitrogen cycle in soil...

 and nitrogen fixation
Nitrogen fixation
Nitrogen fixation is the natural process, either biological or abiotic, by which nitrogen in the atmosphere is converted into ammonia . This process is essential for life because fixed nitrogen is required to biosynthesize the basic building blocks of life, e.g., nucleotides for DNA and RNA and...

 (the uptake of atmospheric nitrogen by symbiotic 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
Protein
Proteins are biochemical compounds consisting of one or more polypeptides typically folded into a globular or fibrous form, facilitating a biological function. A polypeptide is a single linear polymer chain of amino acids bonded together by peptide bonds between the carboxyl and amino groups of...

 and amino-acid synthesis and their role in plant metabolism. With this knowledge it was then possible to outline the global nitrogen cycle
Nitrogen cycle
The nitrogen cycle is the process by which nitrogen is converted between its various chemical forms. This transformation can be carried out by both biological and non-biological processes. Important processes in the nitrogen cycle include fixation, mineralization, nitrification, and denitrification...

.

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
Population genetics is the study of allele frequency distribution and change under the influence of the four main evolutionary processes: natural selection, genetic drift, mutation and gene flow. It also takes into account the factors of recombination, population subdivision and population...

.
By 1910 experiments using labelled isotope
Isotope
Isotopes are variants of atoms of a particular chemical element, which have differing numbers of neutrons. Atoms of a particular element by definition must contain the same number of protons but may have a distinct number of neutrons which differs from atom to atom, without changing the designation...

s 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
Chlorophyll is a green pigment found in almost all plants, algae, and cyanobacteria. Its name is derived from the Greek words χλωρος, chloros and φύλλον, phyllon . Chlorophyll is an extremely important biomolecule, critical in photosynthesis, which allows plants to obtain energy from light...

s a and b were separated by thin layer chromatography
Chromatography
Chromatography is the collective term for a set of laboratory techniques for the separation of mixtures....

 then, through the 1920s and 1930s, biochemists, notably Hans Krebs
Hans Adolf Krebs
Sir Hans Adolf Krebs was a German-born British physician and biochemist. Krebs is best known for his identification of two important metabolic cycles: the urea cycle and the citric acid cycle...

 (1900–1981) and Carl (1896–1984) and Gerty Cori
Gerty Cori
Gerty Theresa Cori was an American biochemist who became the third woman—and first American woman—to win a Nobel Prize in science, and the first woman to be awarded the Nobel Prize in Physiology or Medicine.Cori was born in Prague...

 (1896–1957) began tracing out the central metabolic pathways of life. Between the 1930s and 1950s it was determined that ATP
Adenosine triphosphate
Adenosine-5'-triphosphate is a multifunctional nucleoside triphosphate used in cells as a coenzyme. It is often called the "molecular unit of currency" of intracellular energy transfer. ATP transports chemical energy within cells for metabolism...

, located in mitochondria, was the source of cellular chemical energy and the constituent reactions of photosynthesis
Photosynthesis
Photosynthesis is a chemical process that converts carbon dioxide into organic compounds, especially sugars, using the energy from sunlight. Photosynthesis occurs in plants, algae, and many species of bacteria, but not in archaea. Photosynthetic organisms are called photoautotrophs, since they can...

 were progressively revealed. Then, in 1944 DNA
DNA
Deoxyribonucleic acid is a nucleic acid that contains the genetic instructions used in the development and functioning of all known living organisms . The DNA segments that carry this genetic information are called genes, but other DNA sequences have structural purposes, or are involved in...

 was extracted for the first time. Along with these revelations there was the discovery of plant hormones or “growth substances”, notably auxin
Auxin
Auxins are a class of plant hormones with some morphogen-like characteristics. Auxins have a cardinal role in coordination of many growth and behavioral processes in the plant's life cycle and are essential for plant body development. Auxins and their role in plant growth were first described by...

s, (1934) gibberellin
Gibberellin
Gibberellins are plant hormones that regulate growth and influence various developmental processes, including stem elongation, germination, dormancy, flowering, sex expression, enzyme induction, and leaf and fruit senescence....

s (1934) and cytokinin
Cytokinin
Cytokinins are a class of plant growth substances 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 leaf senescence...

s (1964) and the effects of photoperiodism
Photoperiodism
Photoperiodism is the physiological reaction of organisms to the length of day or night. It occurs in plants and animals.Photoperiodism can also be defined as the developmental responses of plants to the relative lengths of the light and dark periods...

, 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 Weismann
Friedrich Leopold August Weismann was a German evolutionary biologist. Ernst Mayr ranked him the second most notable evolutionary theorist of the 19th century, after Charles Darwin...

 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
Meiosis
Meiosis is a special type of cell division necessary for sexual reproduction. The cells produced by meiosis are gametes or spores. The animals' gametes are called sperm and egg cells....

, the complex process of redistribution of hereditary material that occurs in the germ cells. In the 1920s and 1930s population genetics
Population genetics
Population genetics is the study of allele frequency distribution and change under the influence of the four main evolutionary processes: natural selection, genetic drift, mutation and gene flow. It also takes into account the factors of recombination, population subdivision and population...

 combined the theory of evolution with Mendelian genetics 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
Molecular biology
Molecular biology is the branch of biology that deals with the molecular basis of biological activity. This field overlaps with other areas of biology and chemistry, particularly genetics and biochemistry...

. Genetic engineering
Genetic engineering
Genetic engineering, also called genetic modification, is the direct human manipulation of an organism's genome using modern DNA technology. It involves the introduction of foreign DNA or synthetic genes into the organism of interest...

, the insertion of genes into a host cell for cloning, began in the 1970s with the invention of recombinant DNA
Recombinant DNA
Recombinant DNA molecules are DNA sequences that result from the use of laboratory methods to bring together genetic material from multiple sources, creating sequences that would not otherwise be found in biological organisms...

 techniques and its commercial applications applied to agricultural crops followed in the 1990s. There was now the potential to identify organisms by molecular “fingerprinting” and to estimate the times in the past when critical evolutionary changes had occurred through the use of “molecular clock
Molecular clock
The molecular clock is a technique in molecular evolution that uses fossil constraints and rates of molecular change to deduce the time in geologic history when two species or other taxa diverged. It is used to estimate the time of occurrence of events called speciation or radiation...

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 (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
Stromatolite
Stromatolites or stromatoliths are layered accretionary structures formed in shallow water by the trapping, binding and cementation of sedimentary grains by biofilms of microorganisms, especially cyanobacteria ....

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
Ultrastructure
Ultrastructure is the detailed structure of a biological specimen, such as a cell, tissue, or organ, that can be observed by electron microscopy...

”.

New and revised “phylogenetic” classification systems of the plant kingdom were produced, perhaps the most notable being that of August Eichler
August W. Eichler
August Wilhelm Eichler, also known under his Latinized name, Augustus Guilielmus Eichler , was a German botanist who modified the classification system to better reflect the relationships between plants...

 (1839–1887), and the massive 23 volume of Adolf Engler
Adolf Engler
Heinrich Gustav Adolf Engler was a German botanist. He is notable for his work on plant taxonomy and phytogeography, like Die Natürlichen Pflanzenfamilien , edited with Karl A. E...

 (1844–1930) & Karl Prantl
Karl Anton Eugen Prantl
Karl Anton Eugen Prantl , also known as Carl Anton Eugen Prantl, was a German botanist.Prantl was born in Munich, Kingdom of Bavaria, and studied in Munich. In 1870 he graduated with the dissertation Das Inulin. Ein Beitrag zur Pflanzenphysiologie...

 (1849–1893) published over the period 1887 and 1915. Taxonomy
Taxonomy
Taxonomy is the science of identifying and naming species, and arranging them into a classification. The field of taxonomy, sometimes referred to as "biological taxonomy", revolves around the description and use of taxonomic units, known as taxa...

 based on gross morphology was now being supplemented by using characters revealed by pollen morphology
Palynology
Palynology is the science that studies contemporary and fossil palynomorphs, including pollen, spores, orbicules, dinoflagellate cysts, acritarchs, chitinozoans and scolecodonts, together with particulate organic matter and kerogen found in sedimentary rocks and sediments...

, embryology
Embryology
Embryology is a science which is about the development of an embryo from the fertilization of the ovum to the fetus stage...

, anatomy
Anatomy
Anatomy is a branch of biology and medicine that is the consideration of the structure of living things. It is a general term that includes human anatomy, animal anatomy , and plant anatomy...

, cytology
Cell biology
Cell biology is a scientific discipline that studies cells – their physiological properties, their structure, the organelles they contain, interactions with their environment, their life cycle, division and death. This is done both on a microscopic and molecular level...

, serology
Serology
Serology is the scientific study of blood serum and other bodily fluids. In practice, the term usually refers to the diagnostic identification of antibodies in the serum...

, macromolecule
Macromolecule
A macromolecule is a very large molecule commonly created by some form of polymerization. In biochemistry, the term is applied to the four conventional biopolymers , as well as non-polymeric molecules with large molecular mass such as macrocycles...

s and more. The introduction of computers facilitated the rapid analysis of large data sets used for numerical taxonomy
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 analysis rather than using subjective evaluation of...

 (also called taximetrics or phenetics
Phenetics
In biology, phenetics, also known as taximetrics, is an attempt to classify organisms based on overall similarity, usually in morphology or other observable traits, regardless of their phylogeny or evolutionary relation. It is closely related to numerical taxonomy which is concerned with the use of...

). The emphasis on truly natural phylogenies spawned the disciplines of cladistics
Cladistics
Cladistics is a method of classifying species of organisms into groups called clades, which consist of an ancestor organism and all its descendants . For example, birds, dinosaurs, crocodiles, and all descendants of their most recent common ancestor form a clade...

 and phylogenetic systematics. The grand taxonomic synthesis An Integrated System of Classification of Flowering Plants (1981) of American Arthur Cronquist
Arthur Cronquist
Arthur John Cronquist was a North American 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 Cronquist system. Two plant genera in the aster family have been named in his honor...

 (1919–1992) was superseded when, in 1998, the Angiosperm Phylogeny Group
Angiosperm Phylogeny Group
The Angiosperm Phylogeny Group, or APG, refers to an informal international group of systematic botanists who came together to try to establish a consensus on the taxonomy of flowering plants that would reflect new knowledge about plant relationships discovered through phylogenetic studies., three...

 published a phylogeny of flowering plants based on the analysis of DNA
DNA
Deoxyribonucleic acid is a nucleic acid that contains the genetic instructions used in the development and functioning of all known living organisms . The DNA segments that carry this genetic information are called genes, but other DNA sequences have structural purposes, or are involved in...

 sequences using the techniques of the new molecular systematics which was resolving questions concerning the earliest evolutionary branches of the angiosperms (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 Wegener
Alfred Lothar Wegener was a German scientist, geophysicist, and meteorologist.He is most notable for his theory of continental drift , proposed in 1912, which hypothesized that the continents were slowly drifting around the Earth...

's (1880–1930) theory of continental drift
Continental drift
Continental drift is the movement of the Earth's continents relative to each other. The hypothesis that continents 'drift' was first put forward by Abraham Ortelius in 1596 and was fully developed by Alfred Wegener in 1912...

 1912 gave additional impetus to comparative physiology and the study of biogeography
Biogeography
Biogeography is the study of the distribution of species , organisms, and ecosystems in space and through geological time. Organisms and biological communities vary in a highly regular fashion along geographic gradients of latitude, elevation, isolation and habitat area...

 while ecology in the 1930s contributed the important ideas of plant community, succession
Ecological succession
Ecological succession, is the phenomenon or process by which a community progressively transforms itself until a stable community is formed. It is a fundamental concept in ecology, and refers to more or less predictable and orderly changes in the composition or structure of an ecological community...

, community change, and energy flows. From 1940 to 1950 ecology matured to become an independent discipline as Eugene Odum (1913–2002) formulated many of the concepts of ecosystem ecology
Ecosystem ecology
Ecosystem ecology is the integrated study of biotic and abiotic components of ecosystems and their interactions within an ecosystem framework. This science examines how ecosystems work and relates this to their components such as chemicals, bedrock, soil, plants, and animals.Ecosystem ecology...

, 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
Resource management
In organizational studies, resource management is the efficient and effective deployment of an organization's resources when they are needed. Such resources may include financial resources, inventory, human skills, production resources, or information technology...

, conservation, human food security
Food security
Food security refers to the availability of food and one's access to it. A household is considered food-secure when its occupants do not live in hunger or fear of starvation. According to the World Resources Institute, global per capita food production has been increasing substantially for the past...

, biologically invasive organisms
Introduced species
An introduced species — or neozoon, alien, exotic, non-indigenous, or non-native species, or simply an introduction, is a species living outside its indigenous or native distributional range, and has arrived in an ecosystem or plant community by human activity, either deliberate or accidental...

, carbon sequestration, climate change
Climate change
Climate change is a significant and lasting change in the statistical distribution of weather patterns over periods ranging from decades to millions of years. It may be a change in average weather conditions or the distribution of events around that average...

, and sustainability
Sustainability
Sustainability is the capacity to endure. For humans, sustainability is the long-term maintenance of well being, which has environmental, economic, and social dimensions, and encompasses the concept of union, an interdependent relationship and mutual responsible position with all living and non...

.

See also

  • History of plant systematics
    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 lore usually being treated as part of the study of...

  • History of phycology
    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...

  • List of botanists
  • List of botanists by author abbreviation

Further reading

The source of this article is wikipedia, the free encyclopedia.  The text of this article is licensed under the GFDL.
 
x
OK