Mycology is the branch of biology concerned with the study of fungi , including their taxonomy , genetics , biochemical properties, and use by humans . Fungi can be a source of tinder , food , traditional medicine , as well as entheogens , poison , and infection . Mycology branches into the field of phytopathology , the study of plant diseases. The two disciplines are closely related, because the vast majority of plant pathogens are fungi. A biologist specializing in mycology is called a mycologist .
191-470: Although mycology was historically considered a branch of botany , the 1969 discovery of fungi's close evolutionary relationship to animals resulted in the study's reclassification as an independent field. Pioneer mycologists included Elias Magnus Fries , Christiaan Hendrik Persoon , Heinrich Anton de Bary , Elizabeth Eaton Morse , and Lewis David de Schweinitz . Beatrix Potter , author of The Tale of Peter Rabbit , also made significant contributions to
382-794: A folk medicine in China , Japan , and Russia . Although the use of mushrooms in folk medicine is centered largely on the Asian continent, people in other parts of the world like the Middle East , Poland , and Belarus have been documented using mushrooms for medicinal purposes. Mushrooms produce large amounts of vitamin D when exposed to ultraviolet (UV) light . Penicillin , ciclosporin , griseofulvin , cephalosporin and psilocybin are examples of drugs that have been isolated from molds or other fungi. Botany Botany , also called plant science (or plant sciences ), plant biology or phytology ,
573-483: A botanically and pharmacologically important herbal Historia Plantarum in 1544 and a pharmacopoeia of lasting importance, the Dispensatorium in 1546. Naturalist Conrad von Gesner (1516–1565) and herbalist John Gerard (1545– c. 1611 ) published herbals covering the supposed medicinal uses of plants. Naturalist Ulisse Aldrovandi (1522–1605) was considered the father of natural history , which included
764-414: A considerable problem in agriculture, and the biology and control of plant pathogens in agriculture and natural ecosystems . Ethnobotany is the study of the relationships between plants and people. When applied to the investigation of historical plant–people relationships ethnobotany may be referred to as archaeobotany or palaeoethnobotany . Some of the earliest plant-people relationships arose between
955-452: A continuum between the major morphological categories of root, stem (caulome), leaf (phyllome) and trichome . Furthermore, it emphasises structural dynamics. Modern systematics aims to reflect and discover phylogenetic relationships between plants. Modern Molecular phylogenetics largely ignores morphological characters, relying on DNA sequences as data. Molecular analysis of DNA sequences from most families of flowering plants enabled
1146-421: A few families and still others lived on isolated farms spread over the countryside. There were also areas where the pattern was a mix of two or more of those systems. Unlike in the late Roman period, there was no sharp break between the legal status of the free peasant and the aristocrat, and it was possible for a free peasant's family to rise into the aristocracy over several generations through military service to
1337-400: A hierarchical classification of plant species that remains the reference point for modern botanical nomenclature . This established a standardised binomial or two-part naming scheme where the first name represented the genus and the second identified the species within the genus. For the purposes of identification, Linnaeus's Systema Sexuale classified plants into 24 groups according to
1528-586: A king to rule over them all. By the late sixth century, this arrangement had been replaced by a permanent monarchy, the Kingdom of the Lombards . The invasions brought new ethnic groups to Europe, although some regions received a larger influx of new peoples than others. In Gaul for instance, the invaders settled much more extensively in the north-east than in the south-west. Slavs settled in Central and Eastern Europe and
1719-404: A plant sucks water through them under water stress. Lignin is also used in other cell types like sclerenchyma fibres that provide structural support for a plant and is a major constituent of wood. Sporopollenin is a chemically resistant polymer found in the outer cell walls of spores and pollen of land plants responsible for the survival of early land plant spores and the pollen of seed plants in
1910-552: A polymer of fructose is used for the same purpose in the sunflower family Asteraceae . Some of the glucose is converted to sucrose (common table sugar) for export to the rest of the plant. Unlike in animals (which lack chloroplasts), plants and their eukaryote relatives have delegated many biochemical roles to their chloroplasts , including synthesising all their fatty acids , and most amino acids . The fatty acids that chloroplasts make are used for many things, such as providing material to build cell membranes out of and making
2101-567: A powerful lord. Roman city life and culture changed greatly in the early Middle Ages. Although Italian cities remained inhabited, they contracted significantly in size. Rome, for instance, shrank from a population of hundreds of thousands to around 30,000 by the end of the 6th century. Roman temples were converted into Christian churches and city walls remained in use. In Northern Europe, cities also shrank, while civic monuments and other public buildings were raided for building materials. The establishment of new kingdoms often meant some growth for
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#17328722149002292-462: A practical skill rather than a sign of elite status. In the 4th century, Jerome (d. 420) dreamed that God rebuked him for spending more time reading Cicero than the Bible . By the 6th century, Gregory of Tours (d. 594) had a similar dream, but instead of being chastised for reading Cicero, he was chastised for learning shorthand . By the late 6th century, the principal means of religious instruction in
2483-460: A process that generates molecular oxygen (O 2 ) as a by-product. The light energy captured by chlorophyll a is initially in the form of electrons (and later a proton gradient ) that's used to make molecules of ATP and NADPH which temporarily store and transport energy. Their energy is used in the light-independent reactions of the Calvin cycle by the enzyme rubisco to produce molecules of
2674-505: A pure form of carbon made by pyrolysis of wood, has a long history as a metal- smelting fuel, as a filter material and adsorbent and as an artist's material and is one of the three ingredients of gunpowder . Cellulose , the world's most abundant organic polymer, can be converted into energy, fuels, materials and chemical feedstock. Products made from cellulose include rayon and cellophane , wallpaper paste , biobutanol and gun cotton . Sugarcane , rapeseed and soy are some of
2865-467: A small foothold in southern Spain. Justinian's reconquests have been criticised by historians for overextending his realm and setting the stage for the early Muslim conquests , but many of the difficulties faced by Justinian's successors were due not just to over-taxation to pay for his wars but to the essentially civilian nature of the empire, which made raising troops difficult. In the Eastern Empire
3056-526: A student of Aristotle who invented and described many of its principles and is widely regarded in the scientific community as the "Father of Botany". His major works, Enquiry into Plants and On the Causes of Plants , constitute the most important contributions to botanical science until the Middle Ages , almost seventeen centuries later. Another work from Ancient Greece that made an early impact on botany
3247-454: A useful proxy for temperature in historical climatology , and the biological impact of climate change and global warming . Palynology , the analysis of fossil pollen deposits in sediments from thousands or millions of years ago allows the reconstruction of past climates. Estimates of atmospheric CO 2 concentrations since the Palaeozoic have been obtained from stomatal densities and
3438-1010: A variety of spatial scales in groups, populations and communities that collectively constitute vegetation. Regions with characteristic vegetation types and dominant plants as well as similar abiotic and biotic factors, climate , and geography make up biomes like tundra or tropical rainforest . Herbivores eat plants, but plants can defend themselves and some species are parasitic or even carnivorous . Other organisms form mutually beneficial relationships with plants. For example, mycorrhizal fungi and rhizobia provide plants with nutrients in exchange for food, ants are recruited by ant plants to provide protection, honey bees , bats and other animals pollinate flowers and humans and other animals act as dispersal vectors to spread spores and seeds . Plant responses to climate and other environmental changes can inform our understanding of how these changes affect ecosystem function and productivity. For example, plant phenology can be
3629-628: A way of drug discovery . Plants can synthesise coloured dyes and pigments such as the anthocyanins responsible for the red colour of red wine , yellow weld and blue woad used together to produce Lincoln green , indoxyl , source of the blue dye indigo traditionally used to dye denim and the artist's pigments gamboge and rose madder . Sugar, starch , cotton, linen , hemp , some types of rope , wood and particle boards , papyrus and paper, vegetable oils , wax , and natural rubber are examples of commercially important materials made from plant tissues or their secondary products. Charcoal ,
3820-452: A wider range of shared characters and were widely followed. The Candollean system reflected his ideas of the progression of morphological complexity and the later Bentham & Hooker system , which was influential until the mid-19th century, was influenced by Candolle's approach. Darwin 's publication of the Origin of Species in 1859 and his concept of common descent required modifications to
4011-406: Is De materia medica , a five-volume encyclopedia about preliminary herbal medicine written in the middle of the first century by Greek physician and pharmacologist Pedanius Dioscorides . De materia medica was widely read for more than 1,500 years. Important contributions from the medieval Muslim world include Ibn Wahshiyya 's Nabatean Agriculture , Abū Ḥanīfa Dīnawarī 's (828–896)
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#17328722149004202-405: Is peppermint , Mentha × piperita , a sterile hybrid between Mentha aquatica and spearmint, Mentha spicata . The many cultivated varieties of wheat are the result of multiple inter- and intra- specific crosses between wild species and their hybrids. Angiosperms with monoecious flowers often have self-incompatibility mechanisms that operate between the pollen and stigma so that
4393-476: Is a broad, multidisciplinary subject with contributions and insights from most other areas of science and technology. Research topics include the study of plant structure , growth and differentiation, reproduction , biochemistry and primary metabolism , chemical products, development , diseases , evolutionary relationships , systematics , and plant taxonomy . Dominant themes in 21st-century plant science are molecular genetics and epigenetics , which study
4584-762: Is gathered by ethnobotanists. This information can relay a great deal of information on how the land once was thousands of years ago and how it has changed over that time. The goals of plant ecology are to understand the causes of their distribution patterns, productivity, environmental impact, evolution, and responses to environmental change. Plants depend on certain edaphic (soil) and climatic factors in their environment but can modify these factors too. For example, they can change their environment's albedo , increase runoff interception, stabilise mineral soils and develop their organic content, and affect local temperature. Plants compete with other organisms in their ecosystem for resources. They interact with their neighbours at
4775-627: Is itself subdivided into the Early , High , and Late Middle Ages . Population decline , counterurbanisation , the collapse of centralized authority, invasions, and mass migrations of tribes , which had begun in Late Antiquity , continued into the Early Middle Ages. The large-scale movements of the Migration Period , including various Germanic peoples , formed new kingdoms in what remained of
4966-649: Is no universally agreed upon end date. Depending on the context, events such as the conquest of Constantinople by the Turks in 1453, Christopher Columbus 's first voyage to the Americas in 1492, or the Protestant Reformation in 1517 are sometimes used. English historians often use the Battle of Bosworth Field in 1485 to mark the end of the period. For Spain, dates commonly used are the death of King Ferdinand II in 1516,
5157-451: Is still a major foundation of modern botany. Her books Plant Anatomy and Anatomy of Seed Plants have been key plant structural biology texts for more than half a century. The discipline of plant ecology was pioneered in the late 19th century by botanists such as Eugenius Warming , who produced the hypothesis that plants form communities , and his mentor and successor Christen C. Raunkiær whose system for describing plant life forms
5348-400: Is still given to these groups by botanists, and fungi (including lichens) and photosynthetic protists are usually covered in introductory botany courses. Palaeobotanists study ancient plants in the fossil record to provide information about the evolutionary history of plants . Cyanobacteria , the first oxygen-releasing photosynthetic organisms on Earth, are thought to have given rise to
5539-444: Is still in use today. The concept that the composition of plant communities such as temperate broadleaf forest changes by a process of ecological succession was developed by Henry Chandler Cowles , Arthur Tansley and Frederic Clements . Clements is credited with the idea of climax vegetation as the most complex vegetation that an environment can support and Tansley introduced the concept of ecosystems to biology. Building on
5730-495: Is the science of plant life and a branch of biology . A botanist , plant scientist or phytologist is a scientist who specialises in this field. The term "botany" comes from the Ancient Greek word botanē ( βοτάνη ) meaning " pasture ", " herbs " " grass ", or " fodder "; Botanē is in turn derived from boskein ( Greek : βόσκειν ), "to feed" or "to graze ". Traditionally, botany has also included
5921-498: Is the study of ferns and allied plants. A number of other taxa of ranks varying from family to subgenus have terms for their study, including agrostology (or graminology) for the study of grasses, synantherology for the study of composites, and batology for the study of brambles. Study can also be divided by guild rather than clade or grade . Dendrology is the study of woody plants. Many divisions of biology have botanical subfields. These are commonly denoted by prefixing
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6112-467: The fyrd , which were led by the local elites. In military technology, one of the main changes was the return of the crossbow , which had been known in Roman times and reappeared as a military weapon during the last part of the Early Middle Ages. Another change was the introduction of the stirrup, which increased the effectiveness of cavalry as shock troops. A technological advance that had implications beyond
6303-690: The Alans , Vandals , and Suevi crossed into Gaul ; over the next three years they spread across Gaul and in 409 crossed the Pyrenees Mountains into modern-day Spain. The Migration Period began, when various peoples, initially largely Germanic peoples , moved across Europe. The Franks , Alemanni , and the Burgundians all ended up in northern Gaul while the Angles , Saxons , and Jutes settled in Britain , and
6494-553: The Angiosperm Phylogeny Group to publish in 1998 a phylogeny of flowering plants, answering many of the questions about relationships among angiosperm families and species. The theoretical possibility of a practical method for identification of plant species and commercial varieties by DNA barcoding is the subject of active current research. Botany is divided along several axes. Some subfields of botany relate to particular groups of organisms. Divisions related to
6685-521: The Benedictine Rule for Western monasticism during the 6th century, detailing the administrative and spiritual responsibilities of a community of monks led by an abbot . Monks and monasteries had a deep effect on the religious and political life of the Early Middle Ages, in various cases acting as land trusts for powerful families, centres of propaganda and royal support in newly conquered regions, and bases for missions and proselytisation. They were
6876-495: The Book of Plants , and Ibn Bassal 's The Classification of Soils . In the early 13th century, Abu al-Abbas al-Nabati , and Ibn al-Baitar (d. 1248) wrote on botany in a systematic and scientific manner. In the mid-16th century, botanical gardens were founded in a number of Italian universities. The Padua botanical garden in 1545 is usually considered to be the first which is still in its original location. These gardens continued
7067-417: The C 4 carbon fixation pathway for photosynthesis which avoid the losses resulting from photorespiration in the more common C 3 carbon fixation pathway. These biochemical strategies are unique to land plants. Phytochemistry is a branch of plant biochemistry primarily concerned with the chemical substances produced by plants during secondary metabolism . Some of these compounds are toxins such as
7258-575: The Carolingian Empire during the later 8th and early 9th centuries. It covered much of Western Europe but later succumbed to the pressures of internal civil wars combined with external invasions: Vikings from the north, Magyars from the east, and Saracens from the south. During the High Middle Ages, which began after 1000, the population of Europe increased greatly as technological and agricultural innovations allowed trade to flourish and
7449-520: The Desert Fathers of Egypt and Syria . Most European monasteries were of the type that focuses on community experience of the spiritual life, called cenobitism , which was pioneered by Pachomius (d. 348) in the 4th century. Monastic ideals spread from Egypt to Western Europe in the 5th and 6th centuries through hagiographical literature such as the Life of Anthony . Benedict of Nursia (d. 547) wrote
7640-568: The Gregorian chant in liturgical music for the churches. An important activity for scholars during this period was the copying, correcting, and dissemination of basic works on religious and secular topics, with the aim of encouraging learning. New works on religious topics and schoolbooks were also produced. Grammarians of the period modified the Latin language, changing it from the Classical Latin of
7831-698: The Macedonian dynasty . Commerce revived and the emperors oversaw the extension of a uniform administration to all the provinces. The military was reorganised, which allowed the emperors John I (r. 969–976) and Basil II (r. 976–1025) to expand the frontiers of the empire on all fronts. The imperial court was the centre of a revival of classical learning, a process known as the Macedonian Renaissance . Writers such as John Geometres ( fl. early 10th century) composed new hymns, poems, and other works. Missionary efforts by both Eastern and Western clergy resulted in
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8022-477: The Medieval Warm Period climate change allowed crop yields to increase. Manorialism , the organisation of peasants into villages that owed rent and labour services to the nobles , and feudalism , the political structure whereby knights and lower-status nobles owed military service to their overlords in return for the right to rent from lands and manors , were two of the ways society was organized in
8213-504: The alkaloid coniine from hemlock . Others, such as the essential oils peppermint oil and lemon oil are useful for their aroma, as flavourings and spices (e.g., capsaicin ), and in medicine as pharmaceuticals as in opium from opium poppies . Many medicinal and recreational drugs , such as tetrahydrocannabinol (active ingredient in cannabis ), caffeine , morphine and nicotine come directly from plants. Others are simple derivatives of botanical natural products. For example,
8404-565: The auxin plant hormones by Kenneth V. Thimann in 1948 enabled regulation of plant growth by externally applied chemicals. Frederick Campion Steward pioneered techniques of micropropagation and plant tissue culture controlled by plant hormones . The synthetic auxin 2,4-dichlorophenoxyacetic acid or 2,4-D was one of the first commercial synthetic herbicides . 20th century developments in plant biochemistry have been driven by modern techniques of organic chemical analysis , such as spectroscopy , chromatography and electrophoresis . With
8595-402: The botanical kingdom Fungi and reasonably modern. Many fungi produce toxins , antibiotics , and other secondary metabolites . For example, the cosmopolitan genus Fusarium and their toxins associated with fatal outbreaks of alimentary toxic aleukia in humans were extensively studied by Abraham Z. Joffe . Fungi are fundamental for life on earth in their roles as symbionts , e.g. in
8786-626: The cell theory with Theodor Schwann and Rudolf Virchow and was among the first to grasp the significance of the cell nucleus that had been described by Robert Brown in 1831. In 1855, Adolf Fick formulated Fick's laws that enabled the calculation of the rates of molecular diffusion in biological systems. Building upon the gene-chromosome theory of heredity that originated with Gregor Mendel (1822–1884), August Weismann (1834–1914) proved that inheritance only takes place through gametes . No other cells can pass on inherited characters. The work of Katherine Esau (1898–1997) on plant anatomy
8977-444: The cellulose and lignin used to build their bodies, and secondary products like resins and aroma compounds . Plants and various other groups of photosynthetic eukaryotes collectively known as " algae " have unique organelles known as chloroplasts . Chloroplasts are thought to be descended from cyanobacteria that formed endosymbiotic relationships with ancient plant and algal ancestors. Chloroplasts and cyanobacteria contain
9168-427: The indigenous people of Canada in identifying edible plants from inedible plants. This relationship the indigenous people had with plants was recorded by ethnobotanists. Plant biochemistry is the study of the chemical processes used by plants. Some of these processes are used in their primary metabolism like the photosynthetic Calvin cycle and crassulacean acid metabolism . Others make specialised materials like
9359-426: The pines , and flowering plants ) and the free-sporing cryptogams including ferns , clubmosses , liverworts , hornworts and mosses . Embryophytes are multicellular eukaryotes descended from an ancestor that obtained its energy from sunlight by photosynthesis . They have life cycles with alternating haploid and diploid phases. The sexual haploid phase of embryophytes, known as the gametophyte , nurtures
9550-462: The 17th-century German historian Christoph Cellarius divided history into three periods: ancient, medieval, and modern. The most commonly given starting point for the Middle Ages is around 500, with the date of 476 first used by Bruni. Later starting dates are sometimes used in the outer parts of Europe. For Europe as a whole, 1500 is often considered to be the end of the Middle Ages, but there
9741-408: The 3-carbon sugar glyceraldehyde 3-phosphate (G3P). Glyceraldehyde 3-phosphate is the first product of photosynthesis and the raw material from which glucose and almost all other organic molecules of biological origin are synthesised. Some of the glucose is converted to starch which is stored in the chloroplast. Starch is the characteristic energy store of most land plants and algae, while inulin ,
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#17328722149009932-474: The 5th and the 7th centuries, going first to England and Scotland and then on to the continent. Under such monks as Columba (d. 597) and Columbanus (d. 615), they founded monasteries, taught in Latin and Greek, and authored secular and religious works. The Early Middle Ages witnessed the rise of monasticism in the West. The shape of European monasticism was determined by traditions and ideas that originated with
10123-702: The 5th century. The Eastern Empire was marked by closer relations between the political state and Christian Church, with doctrinal matters assuming an importance in Eastern politics that they did not have in Western Europe. Legal developments included the codification of Roman law ; the first effort—the Codex Theodosianus —was completed in 438. Under Emperor Justinian (r. 527–565), another compilation took place—the Corpus Juris Civilis . Justinian also oversaw
10314-616: The 5th to the late 15th centuries, similarly to the Post-classical period of global history . It began with the fall of the Western Roman Empire and transitioned into the Renaissance and the Age of Discovery . The Middle Ages is the middle period of the three traditional divisions of Western history: classical antiquity , the medieval period, and the modern period . The medieval period
10505-665: The 6th and 7th centuries, all of them ruled by the Merovingian dynasty, who were descended from Clovis. The 7th century was a tumultuous period of wars between Austrasia and Neustria. Such warfare was exploited by Pippin (d. 640), the Mayor of the Palace for Austrasia who became the power behind the Austrasian throne. Later members of his family inherited the office, acting as advisers and regents. One of his descendants, Charles Martel (d. 741), won
10696-636: The Arabs. The migrations and invasions of the 4th and 5th centuries disrupted trade networks around the Mediterranean. African goods stopped being imported into Europe, first disappearing from the interior and by the 7th century found only in a few cities such as Rome or Naples . By the end of the 7th century, under the impact of the Muslim conquests , African products were no longer found in Western Europe. The replacement of goods from long-range trade with local products
10887-526: The Bald received the western Frankish lands, comprising most of modern-day France. Charlemagne's grandsons and great-grandsons divided their kingdoms between their descendants, eventually causing all internal cohesion to be lost. In 987 the Carolingian dynasty was replaced in the western lands, with the crowning of Hugh Capet (r. 987–996) as king. In the eastern lands the dynasty had died out earlier, in 911, with
11078-535: The Balkan Peninsula. The settlement of peoples was accompanied by changes in languages. Latin , the literary language of the Western Roman Empire, was gradually replaced by vernacular languages which evolved from Latin, but were distinct from it, collectively known as Romance languages . These changes from Latin to the new languages took many centuries. Greek remained the language of the Byzantine Empire, but
11269-628: The Battle of Poitiers in 732, halting the advance of Muslim armies across the Pyrenees. Great Britain was divided into small states dominated by the kingdoms of Northumbria , Mercia , Wessex , and East Anglia which descended from the Anglo-Saxon invaders. Smaller kingdoms in present-day Wales and Scotland were still under the control of the native Britons and Picts . Ireland was divided into even smaller political units, usually known as tribal kingdoms, under
11460-566: The Byzantine Empire, as the assumption of the imperial title by the Carolingians asserted their equivalence to the Byzantine state. There were several differences between the newly established Carolingian Empire and both the older Western Roman Empire and the concurrent Byzantine Empire. The Frankish lands were rural in character, with only a few small cities. Most of the people were peasants settled on small farms. Little trade existed and much of that
11651-518: The Byzantine Empire, which he sealed with the marriage of his son Otto II (r. 967–983) to Theophanu (d. 991), daughter of an earlier Byzantine Emperor Romanos II (r. 959–963). By the late 10th century Italy had been drawn into the Ottonian sphere after a period of instability; Otto III (r. 996–1002) spent much of his later reign in the kingdom. The western Frankish kingdom was more fragmented, and although kings remained nominally in charge, much of
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#173287221490011842-506: The Candollean system to reflect evolutionary relationships as distinct from mere morphological similarity. Botany was greatly stimulated by the appearance of the first "modern" textbook, Matthias Schleiden 's Grundzüge der Wissenschaftlichen Botanik , published in English in 1849 as Principles of Scientific Botany . Schleiden was a microscopist and an early plant anatomist who co-founded
12033-629: The Christian period as nova (or "new"). Petrarch regarded the post-Roman centuries as " dark " compared to the "light" of classical antiquity . Leonardo Bruni was the first historian to use tripartite periodisation in his History of the Florentine People (1442), with a middle period "between the fall of the Roman Empire and the revival of city life sometime in late eleventh and twelfth centuries". Tripartite periodisation became standard after
12224-642: The Church had widened to the extent that the cultural and religious differences were greater than the similarities. The formal break, known as the East–West Schism , came in 1054, when the papacy and the patriarchy of Constantinople clashed over papal supremacy and excommunicated each other, which led to the division of Christianity into two Churches—the Western branch became the Roman Catholic Church and
12415-499: The Church had become music and art rather than the book. Most intellectual efforts went towards imitating classical scholarship, but some original works were created, along with now-lost oral compositions. The writings of Sidonius Apollinaris (d. 489), Cassiodorus (d. c. 585 ), and Boethius (d. c. 525) were typical of the age. Changes also took place among laymen, as aristocratic culture focused on great feasts held in halls rather than on literary pursuits. Clothing for
12606-676: The Dnieper River in modern Ukraine to the Adriatic Sea. By 1018, the last Bulgarian nobles had surrendered to the Byzantine Empire. Few large stone buildings were constructed between the Constantinian basilicas of the 4th century and the 8th century, although many smaller ones were built during the 6th and 7th centuries. By the beginning of the 8th century, the Carolingian Empire revived the basilica form of architecture. One feature of
12797-581: The Early Middle Ages are mostly illuminated manuscripts and carved ivories , originally made for metalwork that has since been melted down. Objects in precious metals were the most prestigious form of art, but almost all are lost except for a few crosses such as the Cross of Lothair , several reliquaries , and finds such as the Anglo-Saxon burial at Sutton Hoo and the hoards of Gourdon from Merovingian France, Guarrazar from Visigothic Spain and Nagyszentmiklós near Byzantine territory. There are survivals from
12988-584: The Eastern Mediterranean and remained a major power. The empire's law code, the Corpus Juris Civilis or "Code of Justinian", was rediscovered in Northern Italy in the 11th century. In the West, most kingdoms incorporated the few extant Roman institutions. Monasteries were founded as campaigns to Christianise pagan Europe continued. The Franks , under the Carolingian dynasty , briefly established
13179-520: The Eastern branch the Eastern Orthodox Church . The ecclesiastical structure of the Roman Empire survived the movements and invasions in the west mostly intact, but the papacy was little regarded, and few of the Western bishops looked to the bishop of Rome for religious or political leadership. Many of the popes prior to 750 were more concerned with Byzantine affairs and Eastern theological controversies. The register, or archived copies of
13370-528: The Eastern emperors to pay tribute. They remained a strong power until 796. An additional problem to face the empire came as a result of the involvement of Emperor Maurice (r. 582–602) in Persian politics when he intervened in a succession dispute . This led to a period of peace, but when Maurice was overthrown, the Persians invaded and during the reign of Emperor Heraclius (r. 610–641) controlled large chunks of
13561-487: The European population remained rural peasants. Many were no longer settled in isolated farms but had gathered into small communities, usually known as manors or villages. These peasants were often subject to noble overlords and owed them rents and other services, in a system known as manorialism . There remained a few free peasants throughout this period and beyond, with more of them in the regions of Southern Europe than in
13752-705: The Frankish King Charles the Simple (r. 898–922) to settle in what became Normandy . The eastern parts of the Frankish kingdoms, especially Germany and Italy, were under continual Magyar assault until the invader's defeat at the Battle of Lechfeld in 955. The breakup of the Abbasid dynasty meant that the Islamic world fragmented into smaller political states, some of which began expanding into Italy and Sicily, as well as over
13943-568: The Franks and Celtic Britons set up small polities. Francia was centred in northern Gaul, and the first king of whom much is known is Childeric I (d. 481). His grave was discovered in 1653 and is remarkable for its grave goods , which included weapons and a large quantity of gold. Under Childeric's son Clovis I (r. 509–511), the founder of the Merovingian dynasty , the Frankish kingdom expanded and converted to Christianity. The Britons, related to
14134-665: The German tried to annex all of East Francia. Louis the Pious died in 840, with the empire still in chaos. A three-year civil war followed his death. By the Treaty of Verdun (843), a kingdom between the Rhine and Rhone rivers was created for Lothair to go with his lands in Italy, and his imperial title was recognised. Louis the German was in control of Bavaria and the eastern lands in modern-day Germany. Charles
14325-677: The Great (d. 526) and set up a kingdom marked by its co-operation between the Italians and the Ostrogoths, at least until the last years of Theodoric's reign. The Burgundians settled in Gaul, and after an earlier realm was destroyed by the Huns in 436, formed a new kingdom in the 440s. Between today's Geneva and Lyon , it grew to become the realm of Burgundy in the late 5th and early 6th centuries. Elsewhere in Gaul,
14516-547: The High Middle Ages. This period also saw the collapse of the unified Christian church, with the East-West Schism of 1054 . The Crusades , first preached in 1095, were military attempts by Western European Christians to regain control of the Holy Land from Muslims . Kings became the heads of centralised nation-states , reducing crime and violence but making the ideal of a unified Christendom more distant. Intellectual life
14707-505: The Italian peninsula was conquered by the Ostrogoths . The Eastern Roman Empire, often referred to as the Byzantine Empire after the fall of its western counterpart, had little ability to assert control over the lost western territories. The Byzantine emperors maintained a claim over the territory, but while none of the new kings in the west dared to elevate himself to the position of emperor of
14898-733: The Late Middle Ages and beginning the early modern period . The Middle Ages is one of the three major periods in the most enduring scheme for analysing European history : classical civilisation or Antiquity , the Middle Ages and the Modern Period . The "Middle Ages" first appears in Latin in 1469 as media tempestas or "middle season". In early usage, there were many variants, including medium aevum , or "middle age", first recorded in 1604, and media saecula , or "middle centuries", first recorded in 1625. The adjective "medieval" (or sometimes "mediaeval" or "mediæval"), meaning pertaining to
15089-547: The Lombards, which freed the papacy from the fear of Lombard conquest and marked the beginnings of the Papal States . The coronation of Charlemagne as emperor on Christmas Day 800 is regarded as a turning point in medieval history, marking a return of the Western Roman Empire, since the new emperor ruled over much of the area previously controlled by the Western emperors. It also marks a change in Charlemagne's relationship with
15280-405: The Mediterranean, pottery remained prevalent and appears to have been traded over medium-range networks, not just produced locally. The various Germanic states in the west all had coinages that imitated existing Roman and Byzantine forms. Gold continued to be minted until the end of the 7th century in 693-94 when it was replaced by silver in the Merovingian kingdom. The basic Frankish silver coin
15471-411: The Middle Ages, derives from medium aevum . Medieval writers divided history into periods such as the " Six Ages " or the " Four Empires ", and considered their time to be the last before the end of the world. When referring to their own times, they spoke of them as being "modern". In the 1330s, the Italian humanist and poet Petrarch referred to pre-Christian times as antiqua (or "ancient") and to
15662-760: The Middle East than Europe, losing control of sections of the Muslim lands. Umayyad descendants took over the Iberian Peninsula, the Aghlabids controlled North Africa, and the Tulunids became rulers of Egypt. By the middle of the 8th century, new trading patterns were emerging in the Mediterranean; trade between the Franks and the Arabs replaced the old Roman economy . Franks traded timber, furs, swords and slaves in return for silks and other fabrics, spices, and precious metals from
15853-652: The Pyrenees into the southern parts of the Frankish kingdoms. Efforts by local kings to fight the invaders led to the formation of new political entities. In Anglo-Saxon England , King Alfred the Great (r. 871–899) came to an agreement with the Viking invaders in the late 9th century, resulting in Danish settlements in Northumbria, Mercia, and parts of East Anglia. By the middle of the 10th century, Alfred's successors had conquered Northumbria, and restored English control over most of
16044-607: The Rhine and eastwards, leaving Charles West Francia with the empire to the west of the Rhineland and the Alps. Louis the German (d. 876), the middle child, who had been rebellious to the last, was allowed to keep Bavaria under the suzerainty of his elder brother. The division was disputed. Pepin II of Aquitaine (d. after 864), the emperor's grandson, rebelled in a contest for Aquitaine , while Louis
16235-515: The Roman Empire into a more flexible form to fit the needs of the Church and government. By the reign of Charlemagne, the language had so diverged from the classical Latin that it was later called Medieval Latin . Charlemagne planned to continue the Frankish tradition of dividing his kingdom between all his heirs, but was unable to do so as only one son, Louis the Pious (r. 814–840), was still alive by 813. Just before Charlemagne died in 814, he crowned Louis as his successor. Louis's reign of 26 years
16426-492: The Romans and the invaders are often similar, and tribal items were often modelled on Roman objects. Much of the scholarly and written culture of the new kingdoms was also based on Roman intellectual traditions. An important difference was the gradual loss of tax revenue by the new polities. Many of the new political entities no longer supported their armies through taxes, instead relying on granting them land or rents. This meant there
16617-571: The Vandals went on to cross the strait of Gibraltar after which they conquered the province of Africa . In the 430s the Huns began invading the empire; their king Attila (r. 434–453) led invasions into the Balkans in 442 and 447, Gaul in 451, and Italy in 452. The Hunnic threat remained until Attila's death in 453, when the Hunnic confederation he led fell apart. These invasions by the tribes completely changed
16808-474: The Vegetable Kingdom at the start of chapter XII noted "The first and most important of the conclusions which may be drawn from the observations given in this volume, is that generally cross-fertilisation is beneficial and self-fertilisation often injurious, at least with the plants on which I experimented." An important adaptive benefit of outcrossing is that it allows the masking of deleterious mutations in
16999-455: The West were not uniform; some areas had greatly fragmented landholding patterns, but in other areas large contiguous blocks of land were the norm. These differences allowed for a wide variety of peasant societies, some dominated by aristocratic landholders and others having a great deal of autonomy. Land settlement also varied greatly. Some peasants lived in large settlements that numbered as many as 700 inhabitants. Others lived in small groups of
17190-611: The Western Roman Empire. In the 7th century, North Africa and the Middle East—once part of the Byzantine Empire —came under the rule of the Umayyad Caliphate , an Islamic empire, after conquest by Muhammad's successors . Although there were substantial changes in society and political structures, the break with classical antiquity was not complete. The still-sizeable Byzantine Empire, Rome's direct continuation, survived in
17381-412: The above pair of categories gives rise to fields such as bryogeography (the study of the distribution of mosses). Different parts of plants also give rise to their own subfields, including xylology , carpology (or fructology) and palynology , these been the study of wood, fruit and pollen/spores respectively. Botany also overlaps on the one hand with agriculture, horticulture and silviculture, and on
17572-556: The acceptance of figurative monumental sculpture in Christian art , and by the end of the period near life-sized figures such as the Gero Cross were common in important churches. During the later Roman Empire, the principal military developments were attempts to create an effective cavalry force as well as the continued development of highly specialised types of troops. The creation of heavily armoured cataphract -type soldiers as cavalry
17763-439: The ancestor of plants by entering into an endosymbiotic relationship with an early eukaryote, ultimately becoming the chloroplasts in plant cells. The new photosynthetic plants (along with their algal relatives) accelerated the rise in atmospheric oxygen started by the cyanobacteria , changing the ancient oxygen-free, reducing , atmosphere to one in which free oxygen has been abundant for more than 2 billion years. Among
17954-447: The base of most food chains because they use the energy from the sun and nutrients from the soil and atmosphere, converting them into a form that can be used by animals. This is what ecologists call the first trophic level . The modern forms of the major staple foods , such as hemp , teff , maize, rice, wheat and other cereal grasses, pulses , bananas and plantains, as well as hemp , flax and cotton grown for their fibres, are
18145-417: The basilica is the use of a transept , or the "arms" of a cross-shaped building that are perpendicular to the long nave . Other new features of religious architecture include the crossing tower and a monumental entrance to the church , usually at the west end of the building. Carolingian art was produced for a small group of figures around the court, and the monasteries and churches they supported. It
18336-500: The blue-green pigment chlorophyll a . Chlorophyll a (as well as its plant and green algal-specific cousin chlorophyll b ) absorbs light in the blue-violet and orange/red parts of the spectrum while reflecting and transmitting the green light that we see as the characteristic colour of these organisms. The energy in the red and blue light that these pigments absorb is used by chloroplasts to make energy-rich carbon compounds from carbon dioxide and water by oxygenic photosynthesis ,
18527-446: The broader historical sense of botany include bacteriology , mycology (or fungology) and phycology - the study of bacteria, fungi and algae respectively - with lichenology as a subfield of mycology. The narrower sense of botany in the sense of the study of embryophytes (land plants) is disambiguated as phytology. Bryology is the study of mosses (and in the broader sense also liverworts and hornworts). Pteridology (or filicology)
18718-453: The characters may be artificial in keys designed purely for identification ( diagnostic keys ) or more closely related to the natural or phyletic order of the taxa in synoptic keys. By the 18th century, new plants for study were arriving in Europe in increasing numbers from newly discovered countries and the European colonies worldwide. In 1753, Carl Linnaeus published his Species Plantarum ,
18909-407: The city of Byzantium as the newly renamed eastern capital, Constantinople . Diocletian's reforms strengthened the governmental bureaucracy, reformed taxation, and strengthened the army, which bought the empire time but did not resolve the problems it was facing: excessive taxation, a declining birthrate, and pressures on its frontiers, among others. Civil war between rival emperors became common in
19100-577: The conquest of North Africa sundered maritime connections between those areas. Increasingly, the Byzantine Church differed in language, practices, and liturgy from the Western Church. The Eastern Church used Greek instead of the Western Latin. Theological and political differences emerged, and by the early and middle 8th century issues such as iconoclasm , clerical marriage , and state control of
19291-660: The construction of the Hagia Sophia in Constantinople and the reconquest of North Africa from the Vandals and Italy from the Ostrogoths, under Belisarius (d. 565). The conquest of Italy was not complete, as a deadly outbreak of plague in 542 led to the rest of Justinian's reign concentrating on defensive measures rather than further conquests. At the Emperor's death, the Byzantines had control of most of Italy , North Africa, and
19482-543: The control of kings. There were perhaps as many as 150 local kings in Ireland, of varying importance. The Carolingian dynasty , as the successors to Charles Martel are known, officially took control of the kingdoms of Austrasia and Neustria in a coup of 753 led by Pippin III (r. 752–768). A contemporary chronicle claims that Pippin sought, and gained, authority for this coup from Pope Stephen II (pope 752–757). Pippin's takeover
19673-616: The conversion of the Moravians , Bulgars , Bohemians , Poles , Magyars, and Slavic inhabitants of the Kievan Rus' . These conversions contributed to the founding of political states in the lands of those peoples—the states of Moravia , Bulgaria , Bohemia , Poland , Hungary, and the Kievan Rus'. Bulgaria, which was founded around 680, at its height reached from Budapest to the Black Sea and from
19864-675: The death of Louis the Child , and the selection of the unrelated Conrad I (r. 911–918) as king. The breakup of the Carolingian Empire was accompanied by invasions, migrations, and raids by external foes. The Atlantic and northern shores were harassed by the Vikings , who also raided the British Isles and settled there as well as in Iceland. In 911, the Viking chieftain Rollo (d. c. 931) received permission from
20055-401: The death of Queen Isabella I of Castile in 1504, or the conquest of Granada in 1492. Historians from Romance-speaking countries tend to divide the Middle Ages into two parts: an earlier "High" and later "Low" period. English-speaking historians, following their German counterparts, generally subdivide the Middle Ages into three intervals: "Early", "High", and "Late". In the 19th century,
20246-553: The descriptions were later amended as invalid by modern rules). The founding nomenclaturist Carl Linnaeus included fungi in his binomial naming system in 1753, where each type of organism has a two-word name consisting of a genus and species (whereas up to then organisms were often designated with Latin phrases containing many words). He originated the scientific names of numerous well-known mushroom taxa , such as Boletus and Agaricus , which are still in use today. During this period, fungi were still considered to belong to
20437-461: The developing diploid embryo sporophyte within its tissues for at least part of its life, even in the seed plants, where the gametophyte itself is nurtured by its parent sporophyte. Other groups of organisms that were previously studied by botanists include bacteria (now studied in bacteriology ), fungi ( mycology ) – including lichen -forming fungi ( lichenology ), non- chlorophyte algae ( phycology ), and viruses ( virology ). However, attention
20628-561: The disorder, was killed fighting the Goths at the Battle of Adrianople on 9 August 378. In addition to the threat from such tribal confederacies in the north, internal divisions within the empire, especially within the Christian Church, caused problems. In 400, the Visigoths invaded the Western Roman Empire and, although briefly forced back from Italy, in 410 sacked the city of Rome . In 406
20819-459: The earliest was the Padua botanical garden . These gardens facilitated the academic study of plants. Efforts to catalogue and describe their collections were the beginnings of plant taxonomy and led in 1753 to the binomial system of nomenclature of Carl Linnaeus that remains in use to this day for the naming of all biological species. In the 19th and 20th centuries, new techniques were developed for
21010-442: The early Carolingian period, with a growing dominance of elite heavy cavalry. The use of militia-type levies of the free population declined over the Carolingian period. Although much of the Carolingian armies were mounted, a large proportion during the early period appear to have been mounted infantry , rather than true cavalry. One exception was Anglo-Saxon England, where the armies were still composed of regional levies, known as
21201-419: The efforts of early humans to identify – and later cultivate – plants that were edible, poisonous, and possibly medicinal, making it one of the first endeavours of human investigation. Medieval physic gardens , often attached to monasteries , contained plants possibly having medicinal benefit. They were forerunners of the first botanical gardens attached to universities , founded from the 1540s onwards. One of
21392-523: The elites was richly embellished with jewels and gold. Lords and kings supported entourages of fighters who formed the backbone of the military forces. Family ties within the elites were important, as were the virtues of loyalty, courage, and honour. These ties led to the prevalence of the feud in aristocratic society, examples of which included those related by Gregory of Tours that took place in Merovingian Gaul. Most feuds seem to have ended quickly with
21583-570: The emergence of Islam in Arabia during the lifetime of Muhammad (d. 632). After his death, Islamic forces conquered much of the Eastern Roman Empire and Persia, starting with Syria in 634–635, continuing with Persia between 637 and 642, reaching Egypt in 640–641, North Africa in the later seventh century, and the Iberian Peninsula in 711. By 714, Islamic forces controlled much of
21774-403: The empire than tax-payers. The Emperor Diocletian (r. 284–305) split the empire into separately administered eastern and western halves in 286; the empire was not considered divided by its inhabitants or rulers, as legal and administrative promulgations in one division were considered valid in the other. In 330, after a period of civil war, Constantine the Great (r. 306–337) refounded
21965-560: The empire to Christianity , a gradual process that lasted from the 2nd to the 5th centuries. In 376, the Goths , fleeing from the Huns , received permission from Emperor Valens (r. 364–378) to settle in the Roman province of Thracia in the Balkans . The settlement did not go smoothly, and when Roman officials mishandled the situation, the Goths began to raid and plunder. Valens, attempting to put down
22156-447: The empire, including Egypt, Syria, and Anatolia until Heraclius' successful counterattack. In 628 the empire secured a peace treaty and recovered all of its lost territories. In Western Europe, some of the older Roman elite families died out while others became more involved with ecclesiastical than secular affairs. Values attached to Latin scholarship and education mostly disappeared, and while literacy remained important, it became
22347-479: The empire. Such movements were aided by the refusal of the Western Roman elites to support the army or pay the taxes that would have allowed the military to suppress the migration. The emperors of the 5th century were often controlled by military strongmen such as Stilicho (d. 408), Aetius (d. 454), Aspar (d. 471), Ricimer (d. 472), or Gundobad (d. 516), who were partly or fully of non-Roman background. When
22538-454: The entire Middle Ages were often referred to as the " Dark Ages ", but with the adoption of these subdivisions, use of this term was restricted to the Early Middle Ages, at least among historians. The Roman Empire reached its greatest territorial extent during the 2nd century AD; the following two centuries witnessed the slow decline of Roman control over its outlying territories. Economic issues, including inflation, and external pressure on
22729-453: The extensive earlier work of Alphonse de Candolle , Nikolai Vavilov (1887–1943) produced accounts of the biogeography , centres of origin , and evolutionary history of economic plants. Particularly since the mid-1960s there have been advances in understanding of the physics of plant physiological processes such as transpiration (the transport of water within plant tissues), the temperature dependence of rates of water evaporation from
22920-407: The field. Pier Andrea Saccardo developed a system for classifying the imperfect fungi by spore color and form, which became the primary system used before classification by DNA analysis . He is most famous for his Sylloge Fungorum , which was a comprehensive list of all of the names that had been used for mushrooms . Sylloge is still the only work of this kind that was both comprehensive for
23111-682: The form of mycorrhizae , insect symbionts, and lichens . Many fungi are able to break down complex organic biomolecules such as lignin , the more durable component of wood , and pollutants such as xenobiotics , petroleum , and polycyclic aromatic hydrocarbons . By decomposing these molecules, fungi play a critical role in the global carbon cycle . Fungi and other organisms traditionally recognized as fungi, such as oomycetes and myxomycetes ( slime molds ), often are economically and socially important, as some cause diseases of animals (including humans) and of plants. Apart from pathogenic fungi, many fungal species are very important in controlling
23302-563: The fossil record. It is widely regarded as a marker for the start of land plant evolution during the Ordovician period. The concentration of carbon dioxide in the atmosphere today is much lower than it was when plants emerged onto land during the Ordovician and Silurian periods. Many monocots like maize and the pineapple and some dicots like the Asteraceae have since independently evolved pathways like Crassulacean acid metabolism and
23493-613: The frontiers combined to create the Crisis of the Third Century , with emperors coming to the throne only to be rapidly replaced by new usurpers. Military expenses increased steadily during the 3rd century, mainly in response to the war with the Sasanian Empire , which revived in the middle of the 3rd century. The army doubled in size, and cavalry and smaller units replaced the Roman legion as
23684-451: The functional relationships between plants and their habitats – the environments where they complete their life cycles . Plant ecologists study the composition of local and regional floras , their biodiversity , genetic diversity and fitness , the adaptation of plants to their environment, and their competitive or mutualistic interactions with other species. Some ecologists even rely on empirical data from indigenous people that
23875-462: The fungi that had been perpetuated by the classical authors. The start of the modern age of mycology begins with Pier Antonio Micheli 's 1737 publication of Nova plantarum genera . Published in Florence , this seminal work laid the foundations for the systematic classification of grasses, mosses and fungi. He originated the still current genus names Polyporus and Tuber , both dated 1729 (though
24066-417: The funguses [ sic ]". Some fungi can cause disease in humans and other animals; the study of pathogenic fungi that infect animals is referred to as medical mycology . It is believed that humans started collecting mushrooms as food in prehistoric times. Mushrooms were first written about in the works of Euripides (480–406 BC). The Greek philosopher Theophrastos of Eresos (371–288 BC)
24257-436: The genome of progeny. This beneficial effect is also known as hybrid vigor or heterosis. Once outcrossing is established, subsequent switching to inbreeding becomes disadvantageous since it allows expression of the previously masked deleterious recessive mutations, commonly referred to as inbreeding depression. Middle Ages In the history of Europe , the Middle Ages or medieval period lasted approximately from
24448-444: The global carbon and water cycles and plant roots bind and stabilise soils, preventing soil erosion . Plants are crucial to the future of human society as they provide food, oxygen, biochemicals , and products for people, as well as creating and preserving soil. Historically, all living things were classified as either animals or plants and botany covered the study of all organisms not considered animals. Botanists examine both
24639-447: The imperial officials called missi dominici , who served as roving inspectors and troubleshooters. Charlemagne's court in Aachen was the centre of the cultural revival sometimes referred to as the " Carolingian Renaissance ". Literacy increased, as did development in the arts, architecture and jurisprudence, as well as liturgical and scriptural studies. The English monk Alcuin (d. 804)
24830-621: The important botanical questions of the 21st century are the role of plants as primary producers in the global cycling of life's basic ingredients: energy, carbon, oxygen, nitrogen and water, and ways that our plant stewardship can help address the global environmental issues of resource management , conservation , human food security , biologically invasive organisms , carbon sequestration , climate change , and sustainability . Virtually all staple foods come either directly from primary production by plants, or indirectly from animals that eat them. Plants and other photosynthetic organisms are at
25021-478: The internal functions and processes within plant organelles , cells, tissues, whole plants, plant populations and plant communities. At each of these levels, a botanist may be concerned with the classification ( taxonomy ), phylogeny and evolution , structure ( anatomy and morphology ), or function ( physiology ) of plant life. The strictest definition of "plant" includes only the "land plants" or embryophytes , which include seed plants (gymnosperms, including
25212-415: The king of the united Austrasia and Neustria. Charles, more often known as Charles the Great or Charlemagne , embarked upon a programme of systematic expansion in 774 that unified a large portion of Europe, eventually controlling modern-day France, northern Italy, and Saxony . In the wars that lasted beyond 800, he rewarded allies with war booty and command over parcels of land. In 774, Charlemagne conquered
25403-615: The large brooches in fibula or penannular form that were a key piece of personal adornment for elites, including the Irish Tara Brooch . Highly decorated books were mostly Gospel Books and these have survived in larger numbers , including the Insular Book of Kells , the Book of Lindisfarne , and the imperial Codex Aureus of St. Emmeram , which is one of the few to retain its " treasure binding " of gold encrusted with jewels. Charlemagne's court seems to have been responsible for
25594-403: The leaf shapes and sizes of ancient land plants . Ozone depletion can expose plants to higher levels of ultraviolet radiation-B (UV-B), resulting in lower growth rates. Moreover, information from studies of community ecology , plant systematics , and taxonomy is essential to understanding vegetation change , habitat destruction and species extinction . Inheritance in plants follows
25785-576: The leaf surface and the molecular diffusion of water vapour and carbon dioxide through stomatal apertures. These developments, coupled with new methods for measuring the size of stomatal apertures, and the rate of photosynthesis have enabled precise description of the rates of gas exchange between plants and the atmosphere. Innovations in statistical analysis by Ronald Fisher , Frank Yates and others at Rothamsted Experimental Station facilitated rational experimental design and data analysis in botanical research. The discovery and identification of
25976-470: The letters, of Pope Gregory the Great (pope 590–604) survived, and of those more than 850 letters, the vast majority were concerned with affairs in Italy or Constantinople. The only part of Western Europe where the papacy had influence was Britain, where Gregory had sent the Gregorian mission in 597 to convert the Anglo-Saxons to Christianity. Irish missionaries were most active in Western Europe between
26167-417: The line of Western emperors ceased, many of the kings who replaced them were from the same background. Intermarriage between the new kings and the Roman elites was common. This led to a fusion of Roman culture with the customs of the invading tribes, including the popular assemblies that allowed free male tribal members more say in political matters than was common in the Roman state. Material artefacts left by
26358-517: The main and sometimes only outposts of education and literacy in a region. Many of the surviving manuscripts of the Latin classics were copied in monasteries in the Early Middle Ages. Monks were also the authors of new works, including history, theology, and other subjects, written by authors such as Bede (d. 735), a native of northern England who wrote in the late 7th and early 8th centuries. The Frankish kingdom in northern Gaul split into kingdoms called Austrasia , Neustria , and Burgundy during
26549-410: The main tactical unit. The need for revenue led to increased taxes and a decline in numbers of the curial , or landowning, class, and decreasing numbers of them willing to shoulder the burdens of holding office in their native towns. More bureaucrats were needed in the central administration to deal with the needs of the army, which led to complaints from civilians that there were more tax-collectors in
26740-576: The maintenance of biodiversity . Botany originated as herbalism , the study and use of plants for their possible medicinal properties . The early recorded history of botany includes many ancient writings and plant classifications. Examples of early botanical works have been found in ancient texts from India dating back to before 1100 BCE, Ancient Egypt , in archaic Avestan writings, and in works from China purportedly from before 221 BCE. Modern botany traces its roots back to Ancient Greece specifically to Theophrastus ( c. 371 –287 BCE),
26931-493: The major groups of organisms that carry out photosynthesis , a process that uses the energy of sunlight to convert water and carbon dioxide into sugars that can be used both as a source of chemical energy and of organic molecules that are used in the structural components of cells. As a by-product of photosynthesis, plants release oxygen into the atmosphere, a gas that is required by nearly all living things to carry out cellular respiration. In addition, they are influential in
27122-474: The mechanisms and control of gene expression during differentiation of plant cells and tissues . Botanical research has diverse applications in providing staple foods , materials such as timber , oil , rubber, fibre and drugs, in modern horticulture , agriculture and forestry , plant propagation , breeding and genetic modification , in the synthesis of chemicals and raw materials for construction and energy production, in environmental management , and
27313-459: The middle of the 4th century, diverting soldiers from the empire's frontier forces and allowing invaders to encroach. For much of the 4th century, Roman society stabilised in a new form that differed from the earlier classical period , with a widening gulf between the rich and poor, and a decline in the vitality of the smaller towns. Another change was the Christianisation , or conversion of
27504-484: The migrations of the Slavs added Slavic languages to Eastern Europe. As Western Europe witnessed the formation of new kingdoms, the Eastern Roman Empire remained intact and experienced an economic revival that lasted into the early 7th century. There were fewer invasions of the eastern section of the empire; most occurred in the Balkans. Peace with the Sasanian Empire , the traditional enemy of Rome, lasted throughout most of
27695-464: The military was the horseshoe , which allowed horses to be used in rocky terrain. The High Middle Ages was a period of tremendous expansion of population . The estimated population of Europe grew from 35 to 80 million between 1000 and 1347, although the exact causes remain unclear: improved agricultural techniques, the decline of slaveholding, a more clement climate and the lack of invasion have all been suggested. As much as 90 per cent of
27886-652: The name Agaricus deliciosus to the saffron milk-cap, but its current name is Lactarius deliciosus . On the other hand, the field mushroom Agaricus campestris has kept the same name ever since Linnaeus's publication. The English word " agaric " is still used for any gilled mushroom, which corresponds to Linnaeus's use of the word. The term mycology and the complementary term mycologist are traditionally attributed to M.J. Berkeley in 1836. However, mycologist appeared in writings by English botanist Robert Kaye Greville as early as 1823 in reference to Schweinitz . For centuries, certain mushrooms have been documented as
28077-750: The natives of Britannia – modern-day Great Britain – settled in what is now Brittany . Other monarchies were established by the Visigothic Kingdom in the Iberian Peninsula , the Suebi in northwestern Iberia, and the Vandal Kingdom in North Africa . In the sixth century, the Lombards settled in Northern Italy , replacing the Ostrogothic kingdom with a grouping of duchies that occasionally selected
28268-596: The number of their male sexual organs. The 24th group, Cryptogamia , included all plants with concealed reproductive parts, mosses , liverworts , ferns , algae and fungi . Botany was originally a hobby for upper-class women. These women would collect and paint flowers and plants from around the world with scientific accuracy. The paintings were used to record many species that could not be transported or maintained in other environments. Marianne North illustrated over 900 species in extreme detail with watercolor and oil paintings. Her work and many other women's botany work
28459-447: The other hand with medicine and pharmacology, giving rise to fields such as agronomy , horticultural botany, phytopathology and phytopharmacology . The study of plants is vital because they underpin almost all animal life on Earth by generating a large proportion of the oxygen and food that provide humans and other organisms with aerobic respiration with the chemical energy they need to exist. Plants, algae and cyanobacteria are
28650-404: The outcome of prehistoric selection over thousands of years from among wild ancestral plants with the most desirable characteristics. Botanists study how plants produce food and how to increase yields, for example through plant breeding , making their work important to humanity's ability to feed the world and provide food security for future generations. Botanists also study weeds, which are
28841-791: The pain killer aspirin is the acetyl ester of salicylic acid , originally isolated from the bark of willow trees, and a wide range of opiate painkillers like heroin are obtained by chemical modification of morphine obtained from the opium poppy . Popular stimulants come from plants, such as caffeine from coffee, tea and chocolate, and nicotine from tobacco. Most alcoholic beverages come from fermentation of carbohydrate -rich plant products such as barley (beer), rice ( sake ) and grapes (wine). Native Americans have used various plants as ways of treating illness or disease for thousands of years. This knowledge Native Americans have on plants has been recorded by enthnobotanists and then in turn has been used by pharmaceutical companies as
29032-537: The payment of some sort of compensation . Women took part in aristocratic society mainly in their roles as wives and mothers of men, with the role of mother of a ruler being especially prominent in Merovingian Gaul. In Anglo-Saxon society the lack of many child rulers meant a lesser role for women as queen mothers, but this was compensated for by the increased role played by abbesses of monasteries. Only in Italy does it appear that women were always considered under
29223-526: The peninsula in a region they called Al-Andalus . The Islamic conquests reached their peak in the mid-eighth century. The defeat of Muslim forces at the Battle of Tours in 732 led to the reconquest of southern France by the Franks, but the main reason for the halt of Islamic growth in Europe was the overthrow of the Umayyad Caliphate and its replacement by the Abbasid Caliphate . The Abbasids moved their capital to Baghdad and were more concerned with
29414-526: The plant diseases caused by different pathogens. For example, species of the filamentous fungal genus Trichoderma are considered one of the most important biological control agents as an alternative to chemical-based products for effective crop diseases management. Field meetings to find interesting species of fungi are known as 'forays', after the first such meeting organized by the Woolhope Naturalists' Field Club in 1868 and entitled "A foray among
29605-471: The plant kingdom, so they were categorized in his Species Plantarum . Linnaeus' fungal taxa were not nearly as comprehensive as his plant taxa, however, grouping together all gilled mushrooms with a stem in genus Agaricus . Thousands of gilled species exist, which were later divided into dozens of diverse genera; in its modern usage, Agaricus only refers to mushrooms closely related to the common shop mushroom, Agaricus bisporus . For example, Linnaeus gave
29796-412: The plants with a highly fermentable sugar or oil content that are used as sources of biofuels , important alternatives to fossil fuels , such as biodiesel . Sweetgrass was used by Native Americans to ward off bugs like mosquitoes . These bug repelling properties of sweetgrass were later found by the American Chemical Society in the molecules phytol and coumarin . Plant ecology is the science of
29987-408: The political and demographic nature of what had been the Western Roman Empire. By the end of the 5th century the western section of the empire was divided into smaller political units, ruled by the tribes that had invaded in the early part of the century. The deposition of the last emperor of the west, Romulus Augustulus , in 476 has traditionally marked the end of the Western Roman Empire. By 493
30178-532: The political power devolved to the local lords. Missionary efforts to Scandinavia during the 9th and 10th centuries helped strengthen the growth of kingdoms such as Sweden , Denmark , and Norway , which gained power and territory. Some kings converted to Christianity, although not all by 1000. Scandinavians also expanded and colonised throughout Europe. Besides the settlements in Ireland, England, and Normandy, further settlement took place in what became Russia and Iceland . Swedish traders and raiders ranged down
30369-486: The pollen either fails to reach the stigma or fails to germinate and produce male gametes . This is one of several methods used by plants to promote outcrossing . In many land plants the male and female gametes are produced by separate individuals. These species are said to be dioecious when referring to vascular plant sporophytes and dioicous when referring to bryophyte gametophytes . Charles Darwin in his 1878 book The Effects of Cross and Self-Fertilization in
30560-442: The polymer cutin which is found in the plant cuticle that protects land plants from drying out. Plants synthesise a number of unique polymers like the polysaccharide molecules cellulose , pectin and xyloglucan from which the land plant cell wall is constructed. Vascular land plants make lignin , a polymer used to strengthen the secondary cell walls of xylem tracheids and vessels to keep them from collapsing when
30751-443: The practical value of earlier "physic gardens", often associated with monasteries, in which plants were cultivated for suspected medicinal uses. They supported the growth of botany as an academic subject. Lectures were given about the plants grown in the gardens. Botanical gardens came much later to northern Europe; the first in England was the University of Oxford Botanic Garden in 1621. German physician Leonhart Fuchs (1501–1566)
30942-419: The protection and control of a male relative. Peasant society is much less documented than the nobility. Most of the surviving information available to historians comes from archaeology ; few detailed written records documenting peasant life remain from before the 9th century. Most of the descriptions of the lower classes come from either law codes or writers from the upper classes. Landholding patterns in
31133-461: The rise of the related molecular-scale biological approaches of molecular biology , genomics , proteomics and metabolomics , the relationship between the plant genome and most aspects of the biochemistry, physiology, morphology and behaviour of plants can be subjected to detailed experimental analysis. The concept originally stated by Gottlieb Haberlandt in 1902 that all plant cells are totipotent and can be grown in vitro ultimately enabled
31324-470: The rivers of the Russian steppe, and even attempted to seize Constantinople in 860 and 907 . Christian Spain, initially driven into a small section of the peninsula in the north, expanded slowly south during the 9th and 10th centuries, establishing the kingdoms of Asturias and León . In Eastern Europe, Byzantium revived its fortunes under Emperor Basil I (r. 867–886) and his successors Leo VI (r. 886–912) and Constantine VII (r. 913–959), members of
31515-629: The same fundamental principles of genetics as in other multicellular organisms. Gregor Mendel discovered the genetic laws of inheritance by studying inherited traits such as shape in Pisum sativum ( peas ). What Mendel learned from studying plants has had far-reaching benefits outside of botany. Similarly, " jumping genes " were discovered by Barbara McClintock while she was studying maize. Nevertheless, there are some distinctive genetic differences between plants and other organisms. Species boundaries in plants may be weaker than in animals, and cross species hybrids are often possible. A familiar example
31706-449: The slow infiltration of the Balkans by the Slavs added a further difficulty for Justinian's successors. It began gradually, but by the late 540s Slavic tribes were in Thrace and Illyrium , and had defeated an imperial army near Adrianople in 551. In the 560s the Avars began to expand from their base on the north bank of the Danube ; by the end of the 6th-century, they were the dominant power in Central Europe and routinely able to force
31897-430: The southern part of Great Britain. In northern Britain, Kenneth MacAlpin (d. c. 860) united the Picts and the Scots into the Kingdom of Alba . In the early 10th century, the Ottonian dynasty had established itself in Germany , and was engaged in driving back the Magyars. Its efforts culminated in the coronation in 962 of Otto I (r. 936–973) as Holy Roman Emperor . In 972, he secured recognition of his title by
32088-516: The study of fungi and algae by mycologists and phycologists respectively, with the study of these three groups of organisms remaining within the sphere of interest of the International Botanical Congress . Nowadays, botanists (in the strict sense) study approximately 410,000 species of land plants , including some 391,000 species of vascular plants (of which approximately 369,000 are flowering plants ) and approximately 20,000 bryophytes . Botany originated in prehistory as herbalism with
32279-450: The study of plants, including methods of optical microscopy and live cell imaging , electron microscopy , analysis of chromosome number , plant chemistry and the structure and function of enzymes and other proteins . In the last two decades of the 20th century, botanists exploited the techniques of molecular genetic analysis , including genomics and proteomics and DNA sequences to classify plants more accurately. Modern botany
32470-471: The study of plants. In 1665, using an early microscope, Polymath Robert Hooke discovered cells (a term he coined) in cork , and a short time later in living plant tissue. During the 18th century, systems of plant identification were developed comparable to dichotomous keys , where unidentified plants are placed into taxonomic groups (e.g. family, genus and species) by making a series of choices between pairs of characters . The choice and sequence of
32661-427: The superfluous moisture or earth, of trees, or rotten wood, and of other rotting things. This is plain from the fact that all fungi and truffles, especially those that are used for eating, grow most commonly in thundery and wet weather. The Middle Ages saw little advancement in the body of knowledge about fungi. However, the invention of the printing press allowed authors to dispel superstitions and misconceptions about
32852-447: The towns chosen as capitals. Although there had been Jewish communities in many Roman cities , the Jews suffered periods of persecution after the conversion of the empire to Christianity. Officially they were tolerated, if subject to conversion efforts, and at times were even encouraged to settle in new areas. Religious beliefs in the Eastern Roman Empire and Iran were in flux during the late sixth and early seventh centuries. Judaism
33043-526: The use of genetic engineering experimentally to knock out a gene or genes responsible for a specific trait, or to add genes such as GFP that report when a gene of interest is being expressed. These technologies enable the biotechnological use of whole plants or plant cell cultures grown in bioreactors to synthesise pesticides , antibiotics or other pharmaceuticals , as well as the practical application of genetically modified crops designed for traits such as improved yield. Modern morphology recognises
33234-514: The west, Byzantine control of most of the Western Empire could not be sustained; the reconquest of the Mediterranean periphery and the Italian Peninsula ( Gothic War ) in the reign of Justinian (r. 527–565) was the sole, and temporary, exception. The political structure of Western Europe changed with the end of the united Roman Empire. Although the movements of peoples during this period are usually described as "invasions", they were not just military expeditions but migrations of entire peoples into
33425-485: The word plant (e.g. plant taxonomy, plant ecology, plant anatomy, plant morphology, plant systematics, plant ecology), or prefixing or substituting the prefix phyto- (e.g. phytochemistry , phytogeography ). The study of fossil plants is palaeobotany . Other fields are denoted by adding or substituting the word botany (e.g. systematic botany ). Phytosociology is a subfield of plant ecology that classifies and studies communities of plants. The intersection of fields from
33616-438: Was a trend throughout the old Roman lands that happened in the Early Middle Ages. This was especially marked in the lands that did not lie on the Mediterranean, such as northern Gaul or Britain. Non-local goods appearing in the archaeological record are usually luxury goods. In the northern parts of Europe, not only were the trade networks local, but the goods carried were simple, with little pottery or other complex products. Around
33807-402: Was an active proselytising faith, and at least one Arab political leader converted to it. In addition Jewish theologians wrote polemics defending their religion against Christian and Islamic influences. Christianity had active missions competing with the Persians' Zoroastrianism in seeking converts, especially among residents of the Arabian Peninsula . All these strands came together with
33998-438: Was an important feature of the 5th-century Roman military. The various invading tribes had differing emphases on types of soldiers—ranging from the primarily infantry Anglo-Saxon invaders of Britain to the Vandals and Visigoths who had a high proportion of cavalry in their armies. During the early invasion period, the stirrup had not been introduced into warfare, which limited the usefulness of cavalry as shock troops because it
34189-420: Was dominated by efforts to regain the dignity and classicism of imperial Roman and Byzantine art , but was also influenced by the Insular art of the British Isles. Insular art integrated the energy of Irish Celtic and Anglo-Saxon Germanic styles of ornament with Mediterranean forms such as the book, and established many characteristics of art for the rest of the medieval period. Surviving religious works from
34380-408: Was invited to Aachen and brought the education available in the monasteries of Northumbria. Charlemagne's chancery —or writing office—made use of a new script today known as Carolingian minuscule , allowing a common writing style that advanced communication across much of Europe. Charlemagne sponsored changes in church liturgy , imposing the Roman form of church service on his domains, as well as
34571-434: Was less need for large tax revenues and so the taxation systems decayed. Warfare was common between and within the kingdoms. Slavery declined as the supply weakened, and society became more rural. Between the 5th and 8th centuries, new peoples and individuals filled the political void left by Roman centralised government. The Ostrogoths , a Gothic tribe, settled in Roman Italy in the late fifth century under Theoderic
34762-436: Was marked by scholasticism , a philosophy that emphasised joining faith to reason, and by the founding of universities . The theology of Thomas Aquinas , the paintings of Giotto , the poetry of Dante and Chaucer , the travels of Marco Polo , and the Gothic architecture of cathedrals such as Chartres are among the outstanding achievements toward the end of this period and into the Late Middle Ages. The Late Middle Ages
34953-470: Was marked by difficulties and calamities including famine, plague, and war, which significantly diminished the population of Europe; between 1347 and 1350, the Black Death killed about a third of Europeans. Controversy, heresy , and the Western Schism within the Catholic Church paralleled the interstate conflict, civil strife, and peasant revolts that occurred in the kingdoms. Cultural and technological developments transformed European society, concluding
35144-439: Was marked by numerous divisions of the empire among his sons and, after 829, civil wars between various alliances of father and sons over the control of various parts of the empire. Eventually, Louis recognised his eldest son Lothair I (d. 855) as emperor and gave him Italy. Louis divided the rest of the empire between Lothair and Charles the Bald (d. 877), his youngest son. Lothair took East Francia , comprising both banks of
35335-496: Was not possible to put the full force of the horse and rider behind blows struck by the rider. The greatest change in military affairs during the invasion period was the adoption of the Hunnic composite bow in place of the earlier, and weaker, Scythian composite bow. Another development was the increasing use of longswords and the progressive replacement of scale armour by mail armour and lamellar armour . The importance of infantry and light cavalry began to decline during
35526-399: Was one of "the three German fathers of botany", along with theologian Otto Brunfels (1489–1534) and physician Hieronymus Bock (1498–1554) (also called Hieronymus Tragus). Fuchs and Brunfels broke away from the tradition of copying earlier works to make original observations of their own. Bock created his own system of plant classification. Physician Valerius Cordus (1515–1544) authored
35717-467: Was perhaps the first to try to systematically classify plants; mushrooms were considered to be plants missing certain organs. It was later Pliny the Elder (23–79 AD), who wrote about truffles in his encyclopedia Natural History . The word mycology comes from the Ancient Greek : μύκης ( mukēs ), meaning "fungus" and the suffix -λογία ( -logia ), meaning "study". Fungi and truffles are neither herbs, nor roots, nor flowers, nor seeds, but merely
35908-442: Was reinforced with propaganda that portrayed the Merovingians as inept or cruel rulers, exalted the accomplishments of Charles Martel, and circulated stories of the family's great piety. At the time of his death in 768, Pippin left his kingdom in the hands of his two sons, Charles (r. 768–814) and Carloman (r. 768–771). When Carloman died of natural causes, Charles blocked the succession of Carloman's young son and installed himself as
36099-417: Was the denarius or denier , while the Anglo-Saxon version was called a penny . From these areas, the denier or penny spread throughout Europe from 700 to 1000 AD. Copper or bronze coins were not struck, nor were gold except in Southern Europe. No silver coins denominated in multiple units were minted. Christianity was a major unifying factor between Eastern and Western Europe before the Arab conquests, but
36290-412: Was the beginning of popularizing botany to a wider audience. Increasing knowledge of plant anatomy , morphology and life cycles led to the realisation that there were more natural affinities between plants than the artificial sexual system of Linnaeus. Adanson (1763), de Jussieu (1789), and Candolle (1819) all proposed various alternative natural systems of classification that grouped plants using
36481-409: Was with the British Isles and Scandinavia, in contrast to the older Roman Empire with its trading networks centred on the Mediterranean. The empire was administered by an itinerant court that travelled with the emperor, as well as approximately 300 imperial officials called counts , who administered the counties the empire had been divided into. Clergy and local bishops served as officials, as well as
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