History of botany

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Some traditional tools of botanical science

The history of botany examines the human effort to understand life on Earth by tracing the historical development of the discipline of botany—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

ancient Athens in about 350 BC are considered the starting point for Western botany. In ancient India, the Vṛkṣāyurveda, attributed to Parashara, is also considered one of the earliest texts to describe various branches of botany.[1]

In Europe, botanical science was soon overshadowed by a

Greco-Roman
work on medicinal plants was preserved and extended.

In Europe, the

classification
.

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 (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 and ecology) through to the small scale of subjects like cell theory, molecular biology and plant biochemistry.

Introduction

molecular systematics, for example, entails the principles and techniques of taxonomy, molecular biology, computer science
and more.

Within botany, there are a number of sub-disciplines that focus on particular plant groups, each with their own range of related studies (anatomy, morphology etc.). Included here are:

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

Ancient knowledge

Nomadic hunter-gatherer societies passed on, by oral tradition, what they knew (their empirical observations) about the different kinds of plants that they used for food, shelter, poisons, medicines, for ceremonies and rituals etc. The uses of plants by these pre-literate societies influenced the way the plants were named and classified—their uses were embedded in folk-taxonomies, the way they were grouped according to use in everyday communication.[4] The nomadic life-style was drastically changed when settled communities were established in about twelve centres around the world during the Neolithic Revolution which extended from about 10,000 to 2500 years ago depending on the region. With these communities came the development of the technology and skills needed for the domestication of plants 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.[5]

Plant lore and plant selection

A Sumerian harvester's sickle dated to 3000 BC

During the Neolithic Revolution, plant knowledge increased most obviously through the use of plants for food and medicine. All of today's

William Stearn has observed that "cultivated plants are mankind's most vital and precious heritage from remote antiquity".[7]

It is also from the Neolithic, in about 3000 BC, that we glimpse the first known illustrations of plants

magicians and physicians, who were more likely to record their knowledge for posterity.[11]

Early botany

Ancient India

An early example of ancient Indian plant classification is found in the

Classical antiquity

Classical Greece

1509–1510

Ancient Athens, of the 6th century BC, was the busy trade centre at the confluence of

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.[14]

Theophrastus and the origin of botanical science

Statue of Theophrastus 371–287 BC
"Father of Botany"
Palermo Botanic Gardens

Foremost among the scholars studying botany was Theophrastus of Eressus (Greek: Θεόφραστος; c. 371–287 BC) who has been frequently referred to as the "Father of Botany". He was a student and close friend of Aristotle (384–322 BC) and succeeded him as head of the Lyceum (an educational establishment like a modern university) in Athens with its tradition of peripatetic philosophy. Aristotle's special treatise on plants — θεωρία περὶ φυτῶν — is now lost, although there are many botanical observations scattered throughout his other writings (these have been assembled by Christian Wimmer in Phytologiae Aristotelicae Fragmenta, 1836) but they give little insight into his botanical thinking.[15] 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.[16] Together with Aristotle, he had tutored Alexander the Great 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.[17] It was in this garden where he gained much of his plant knowledge.[18]

Enquiry into Plants and Causes of Plants
The frontispiece to an illustrated 1644 edition of Historia Plantarum
Wild asparagus (Asparagus aphyllus) native to the Levant

Theophrastus's major botanical works were the

Enquiry into Plants (Historia Plantarum) and Causes of Plants (Causae Plantarum) which were his lecture notes for the Lyceum.[19] The opening sentence of the Enquiry reads like a botanical manifesto
:

We must consider the distinctive characters and the general nature of plants from the point of view of their morphology, their behaviour under external conditions, their mode of generation and the whole course of their life.

— Theophrastus, Enquiry into Plants

The Enquiry is 9 books of "applied" botany dealing with the forms and

determinate and indeterminate growth and details of floral structure including the degree of fusion of the petals, position of the ovary and more.[21][22] 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.[23]

Pedanius Dioscorides

Dioscorides and Heuresis

A full synthesis of ancient Greek pharmacology was compiled in De Materia Medica c. 60 AD by Pedanius Dioscorides (c. 40-90 AD) who was a Greek physician with the Roman army. This work proved to be the definitive text on medicinal herbs, both oriental and occidental, for fifteen hundred years until the dawn of the European Renaissance being slavishly copied again and again throughout this period.[24] Though rich in medicinal information with descriptions of about 600 medicinal herbs, the botanical content of the work was extremely limited.[25]

Ancient Rome

Gallic-Roman harvester. Relief from Trier

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 De Re Rustica, four Roman writers contributed to a compendium Scriptores Rei Rusticae, published from the Renaissance on, which set out the principles and practice of agriculture. These authors were Cato (234–149 BC), Varro (116–27 BC) and, in particular, Columella (4–70 AD) and Palladius (4th century AD).[26]

Pliny the Elder

Roman encyclopaedist

Naturalis Historia 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.[27] It is estimated that at the time of the Roman Empire between 1300 and 1400 plants had been recorded in the West.[28]

Ancient China

In

Han Dynasty (202 BC-220 AD) includes the notable work of the Huangdi Neijing and the famous pharmacologist Zhang Zhongjing
.

Medieval knowledge

Medicinal plants of the early Middle Ages

's Canon of Medicine dated 1593

In Western Europe, after Theophrastus, botany passed through a bleak period of 1800 years when little progress was made and, indeed, many of the early insights were lost. As Europe entered the Middle Ages (5th to 15th centuries), China, India and the Arab world enjoyed a golden age.

Medieval China

Chinese philosophy had followed a similar path to that of the ancient Greeks. Between 100 and 1700 AD, many new works on pharmaceutical botany were produced. The 11th century scientists and statesmen Su Song and Shen Kuo compiled learned treatises on natural history, emphasising herbal medicine.[29] Among the pharmaceutical botany works were 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.[30]

Medieval India

In India, simple artificial plant classification systems of the

Cruciferae), Tripuspaganiya (Cucurbitaceae), Mallikaganiya (Apocynaceae), and Kurcapuspaganiya (Asteraceae).[31][32] Important medieval Indian works of plant physiology include the Prthviniraparyam of Udayana
, Nyayavindutika of Dharmottara, Saddarsana-samuccaya of Gunaratna, and Upaskara of Sankaramisra.

Islamic Golden Age

Physician preparing an elixir, from an Arabic version of the De Materia Medica by Dioscorides

The 400-year period from the 9th to 13th centuries AD was the

Ibn Sina (Avicenna) (c. 980–1037 AD) was another influential figure, his The Canon of Medicine being a landmark in the history of medicine treasured until the Enlightenment.[34]

The Silk Road

Following the fall of Constantinople (1453), the newly expanded Ottoman Empire welcomed European embassies in its capital, which in turn became the sources of plants from those regions to the east which traded with the empire. In the following century, twenty times as many plants entered Europe along the Silk Road as had been transported in the previous two thousand years, mainly as bulbs. Others were acquired primarily for their alleged medicinal value. Initially, Italy benefited from this new knowledge, especially Venice, which traded extensively with the East. From there, these new plants rapidly spread to the rest of Western Europe.[35] By the middle of the sixteenth century, there was already a flourishing export trade of various bulbs from Turkey to Europe.[36]

The Age of Herbals

De Materia Medica, Byzantium
, 15th century.

In the European

De Materia Medica
.

European white waterlily Nymphaea alba, from Herbarium Vivae Eicones

The authors of the oldest herbals of the 16th century, Brunfels, Fuchs, Bock, Mattioli and others, regarded plants mainly as the vehicles of medicinal virtues. ... Their chief object was to discover the plants employed by the physicians of antiquity, the knowledge of which had been lost in later times. The corrupt texts of Theophrastus, Dioscorides, Pliny and Galen had been in many respects improved and illustrated by ... Italian commentators of the 15th and ... early part of the 16th century; but there was one imperfection which no criticism could remove,—the highly unsatisfactory descriptions of the old authors or the entire absence of descriptions.[38]

It was moreover at first assumed that the plants described by the Greek physicians must grow wild in Germany also, and generally in the rest of Europe; each author identified a different native plant with some one mentioned by Dioscorides or Theophrastus or others, and thus there arose [in] the 16th century a confusion of nomenclature.[38]

However, the need for accurate and detailed plant descriptions meant that some herbals were more botanical than medicinal.

Two Lavandula species. Woodcut from Hieronymus Bock's Kreütterbuch (2nd ed.) 1546

A great advance was made by the first German composers of herbals, who went straight to nature, described the wild plants growing around them and had figures of them carefully executed in wood. Thus was made the first beginning of a really scientific examination of plants, though the aims pursued were not yet truly scientific, for no questions were proposed as to the nature of plants, their organisation or mutual relations; the only point of interest was the knowledge of individual forms and of their medicinal virtues.[39]

— Julius von Sachs, History of Botany

German Otto Brunfels's (1464–1534) Herbarum Vivae Icones (1530) contained descriptions of about 47 species new to science combined with accurate illustrations. His fellow countryman Hieronymus Bock's (1498–1554) Kreutterbuch of 1539 described plants he found in nearby woods and fields and these were illustrated in the 1546 edition.[40] However, it was Valerius Cordus (1515–1544) who pioneered the formal botanical description that detailed both flowers and fruits, some anatomy including the number of chambers in the ovary, and the type of ovule placentation. He also made observations on pollen and distinguished between inflorescence types.[40] His five-volume Historia Plantarum was published about 18 years after his early death aged 29 in 1561–1563. In England, William Turner (1515–1568) in his Libellus De Re Herbaria Novus (1538) published names, descriptions and localities of many native British plants[41] and in Holland Rembert Dodoens (1517–1585), in Stirpium Historiae (1583), included descriptions of many new species from the Netherlands in a scientific arrangement.[42]

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 Floras. These were often backed by specimens deposited in a herbarium which was a collection of dried plants that verified the plant descriptions given in the Floras. The transition from herbal to Flora marked the final separation of botany from medicine.[43]

The Renaissance and Age of Enlightenment (1550–1800)

A 1647 portrait of a scholar holding a book of plant diagrams.

The revival of learning during the European Renaissance renewed interest in plants. The church, feudal aristocracy and an increasingly influential merchant class that supported science and the arts, now jostled in a world of increasing trade. Sea voyages of exploration returned botanical treasures to the large public, private, and newly established botanic gardens, and introduced an eager population to novel crops, drugs and spices from Asia, the East Indies and the New World.

The number of scientific publications increased. In England, for example, scientific communication and causes were facilitated by learned societies like Royal Society (founded in 1660) and the

Cambridge Botanic Gardens, as well as the influence of renowned private gardens and wealthy entrepreneurial nurserymen.[44] By the early 17th century the number of plants described in Europe had risen to about 6000.[45] The 18th century Enlightenment values of reason and science coupled with new voyages to distant lands instigating another phase of encyclopaedic plant identification, nomenclature, description and illustration, "flower painting" possibly at its best in this period of history.[46][47] 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.[48] Often such exotic new plant imports (primarily from Turkey), when they first appeared in print in English, lacked common names in the language.[47]

During the 18th century, botany was one of the few sciences considered appropriate for genteel educated women. Around 1760, with the popularization of the Linnaean system, botany became much more widespread among educated women who painted plants, attended classes on plant classification, and collected herbarium specimens although emphasis was on the healing properties of plants rather than plant reproduction which had overtones of sexuality. Women began publishing on botanical topics and children's books on botany appeared by authors like

Charlotte Turner Smith. Cultural authorities argued that education through botany created culturally and scientifically aware citizens, part of the thrust for 'improvement' that characterised the Enlightenment. However, in the early 19th century with the recognition of botany as an official science, women were again excluded from the discipline.[49] Compared to other sciences, however, in botany the number of female researchers, collectors, or illustrators has always been remarkably high.[50]

Botanical gardens and herbaria

A 16th century print of the Botanical Garden of Padova (Garden of the Simples) — the oldest academic botanic garden that is still in its original location
Preparing a herbarium specimen

Public and private gardens have always been strongly associated with the historical unfolding of botanical science.[51] Early botanical gardens were physic gardens, repositories for the medicinal plants described in the herbals. As they were generally associated with universities or other academic institutions, the plants were also used for study. The directors of these gardens were eminent physicians with an educational role as "scientific gardeners" and it was staff of these institutions that produced many of the published herbals.

The botanical gardens of the modern tradition were established in northern Italy, the first being at Pisa (1544), founded by Luca Ghini (1490–1556). Although part of a medical faculty, the first chair of materia medica, essentially a chair in botany, was established in Padua in 1533. Then in 1534, Ghini became Reader in materia medica at Bologna University, where Ulisse Aldrovandi established a similar garden in 1568 (see below).[52] Collections of pressed and dried specimens were called a hortus siccus (garden of dry plants) and the first accumulation of plants in this way (including the use of a plant press) is attributed to Ghini.[53][54] Buildings called herbaria housed these specimens mounted on card with descriptive labels. Stored in cupboards in systematic order, they could be preserved in perpetuity and easily transferred or exchanged with other institutions, a taxonomic procedure that is still used today.

By the 18th century, the physic gardens had been transformed into "order beds" that demonstrated the classification systems that were being devised by botanists of the day — but they also had to accommodate the influx of curious, beautiful and new plants pouring in from voyages of exploration that were associated with European colonial expansion.

From Herbal to Flora

Plant classification systems of the 17th and 18th centuries now related plants to one another and not to man, marking a return to the non-anthropocentric botanical science promoted by Theophrastus over 1500 years before. In England, various herbals in either Latin or English were mainly compilations and translations of continental European works, of limited relevance to the British Isles. This included the rather unreliable work of Gerard (1597).[55] The first systematic attempt to collect information on British plants was that of Thomas Johnson (1629),[56][57] who was later to issue his own revision of Gerard's work (1633–1636).[58]

However, Johnson was not the first apothecary or physician to organise botanical expeditions to

Orto Botanico di Bologna in 1568.[52]

In France, Clusius journeyed throughout most of

This approach coupled with the new Linnaean system of binomial nomenclature resulted in plant encyclopaedias without medicinal information called Floras that meticulously described and illustrated the plants growing in particular regions.[66] 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[67] and Italian Marcello Malpighi, would last for 150 years.[68]

Botanical exploration

More new lands were opening up to European colonial powers, the botanical riches being returned to European botanists for description. This was a romantic era of botanical explorers, intrepid

Captain James Cook's chief botanist Joseph Banks (1743–1820).[69]

Classification and morphology

Portrait of Carl Linnaeus by Alexander Roslin, 1775

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.[70]

Italian physician

Botanic Garden of Pisa from 1554 to 1558. His sixteen-volume De Plantis (1583) described 1500 plants and his herbarium of 260 pages and 768 mounted specimens still remains. Caesalpino proposed classes based largely on the detailed structure of the flowers and fruit;[63] he also applied the concept of the genus.[71] 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.[72] Gaspard Bauhin (1560–1624) produced two influential publications Prodromus Theatrici Botanici (1620) and Pinax (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.[73]

Cover page of Species Plantarum of Carl Linnaeus published in 1753

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 (1623–1705) built on Jung's work to establish the most elaborate and insightful classification system of the day.[74] His observations started with the local plants of Cambridge where he lived, with the Catalogus Stirpium circa Cantabrigiam Nascentium (1860) which later expanded to his Synopsis Methodica Stirpium Britannicarum, essentially the first British Flora. Although his Historia Plantarum (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 Historia Plantarum can be regarded as the first botanical synthesis and textbook for modern botany. According to botanical historian Alan Morton, Ray "influenced both the theory and the practice of botany more decisively than any other single person in the latter half of the seventeenth century".[75] Ray's family system was later extended by Pierre Magnol (1638–1715) and Joseph de Tournefort (1656–1708), a student of Magnol, achieved notoriety for his botanical expeditions, his emphasis on floral characters in classification, and for reviving the idea of the genus as the basic unit of classification.[76]

Above all it was Swedish

Antoine-Laurent de Jussieu (1748–1836) extended it yet again to include about 100 orders (present-day families).[78] Frenchman Michel Adanson (1727–1806) in his Familles des Plantes (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.[79]

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

Robert Hooke's microscope which he described in the 1665 Micrographia: he coined the biological use of the term cell

In the first half of the 18th century, botany was beginning to move beyond descriptive science into experimental science. Although the

stomata.[80]

Physiology

In plant physiology, research interest was focused on the movement of sap and the absorption of substances through the roots.

Nicolas-Théodore de Saussure's (1767–1845) Recherches Chimiques sur la Végétation was an exemplary study of scientific exactitude that demonstrated the similarity of respiration in both plants and animals, that the fixation of carbon dioxide includes water, and that just minute amounts of salts and nutrients (which he analyzed in chemical detail from plant ash) have a powerful influence on plant growth.[85]

Plant sexuality

Diagram showing the sexual parts of a mature flower

It was Rudolf Camerarius (1665–1721) who was the first to establish plant sexuality conclusively by experiment. He declared in a letter to a colleague, dated 1694 and titled De Sexu Plantarum Epistola, 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".[86]

Some time later, the German academic and natural historian

Angiosperm
(flowering plant) life cycle showing alternation of generations