Narcissus (plant)

General Narcissus is a genus of perennial herbaceous bulbiferous geophytes, which die back after flowering to an underground storage bulb. They regrow in the following year from brown-skinned ovoid bulbs with pronounced necks, and reach heights of depending on the species. Dwarf species such as N. asturiensis have a maximum height of , while Narcissus tazetta may grow as tall as . Reproductive ; Inflorescence : The inflorescence is scapose, the single stem or scape bearing either a solitary flower or forming an umbel with up to 20 blooms. Species bearing a solitary flower include section Bulbocodium and most of section Pseudonarcissus. Umbellate species have a fleshy racemose inflorescence (unbranched, with short floral stalks) with 2 to 15 or 20 flowers, such as N. papyraceus (see illustration, left) and N. tazetta (see Table I). The flower arrangement on the inflorescence may be either with (pedicellate) or without (sessile) floral stalks. Prior to opening, the flower buds are enveloped and protected in a thin, dry, papery or membranous (scarious) spathe. The spathe consists of a singular bract that is ribbed, and which remains wrapped around the base of the open flower. As the bud grows, the spathe splits longitudinally. Bracteoles are small or absent. ; Flowers : The flowers of Narcissus are hermaphroditic (bisexual), have three parts (tripartite), and are sometimes fragrant (see Fragrances). The flower symmetry is actinomorphic (radial) to slightly zygomorphic (bilateral) due to declinate-ascending stamens (curving downwards, then bent up at the tip). Narcissus flowers are characterised by their, usually conspicuous, corona (trumpet). The three major floral parts (in all species except N. cavanillesii in which the corona is virtually absent - Table I: Section Tapeinanthus) are: • (i) the proximal floral tube (hypanthium), • (ii) the surrounding free tepals, and • (iii) the more distal corona (paracorolla, paraperigon, paraperigonium). All three parts may be considered to be components of the perianth (perigon, perigonium). The perianth arises above the apex of the inferior ovary, its base forming the hypanthial floral tube. The floral tube is formed by fusion of the basal segments of the tepals (proximally connate). Its shape is from an inverted cone (obconic) to funnel-shaped (funneliform) or cylindrical, and is surmounted by the more distal corona. Floral tubes can range from long and narrow (in sections Apodanthi and Jonquilla) to rudimentary (N. cavanillesii). Surrounding the floral tube and corona and reflexed (bent back) from the rest of the perianth are the six spreading tepals or floral leaves, in two whorls which may be distally ascending, reflexed (folded back), or lanceolate. Like many monocotyledons, the perianth is homochlamydeous, which is undifferentiated into separate calyx (sepals) and corolla (petals), but rather has six tepals. The three outer tepal segments may be considered sepals, and the three inner segments petals. The transition point between the floral tube and the corona is marked by the insertion of the free tepals on the fused perianth. The corona, or paracorolla, is variously described as bell-shaped (funneliform, trumpet), bowl-shaped (cupular, crateriform, cup-shaped) or disc-shaped with margins that are often frilled, and is free from the stamens. Rarely is the corona a simple callose (hardened, thickened) ring. The corona is formed during floral development as a tubular outgrowth from stamens which fuse into a tubular structure, the anthers becoming reduced. At its base, the fragrances which attract pollinators are formed. All species produce nectar at the top of the ovary. Coronal morphology varies from the tiny pigmented disk of N. serotinus (see Table I) or the rudimentary structure in N. cavanillesii to the elongated trumpets of section Pseudonarcissus (trumpet daffodils, Table I). While the perianth may point forwards, in some species such as N. cyclamineus it is folded back (reflexed, see illustration, left), while in some other species such as N. bulbocodium (Table I), it is reduced to a few barely visible pointed segments with a prominent corona. The colour of the perianth is white, yellow or bicoloured, with the exception of the night flowering N. viridiflorus, which is green. In addition, the corona of N. poeticus has a red crenulate margin (see Table I). Flower diameter varies from 12 mm (N. bulbocodium) to over 125 mm (N. nobilis=N. pseudonarcissus subsp. nobilis). Flower orientation varies from pendent or deflexed (hanging down) as in N. triandrus (see illustration, left), through declinate-ascendant as in N. alpestris = N. pseudonarcissus subsp. moschatus, horizontal (patent, spreading) such as N. gaditanus or N. poeticus, erect as in N. cavanillesii, N. serotinus and N. rupicola (Table I), or intermediate between these positions (erecto-patent). The flowers of Narcissus demonstrate exceptional floral diversity and sexual polymorphism, primarily by corona size and floral tube length, associated with pollinator groups (see for instance Figs. 1 and 2 in Graham and Barrett). Barrett and Harder (2005) describe three separate floral patterns: • "Daffodil" form • "Paperwhite" form • "Triandrus" form. The predominant patterns are the 'daffodil' and 'paperwhite' forms, while the "triandrus" form is less common. Each corresponds to a different group of pollinators (See Pollination). The "daffodil" form, which includes sections Pseudonarcissus and Bulbocodium, has a relatively short, broad or highly funnelform tube (funnel-like), which grades into an elongated corona, which is large and funnelform, forming a broad, cylindrical or trumpet-shaped perianth. Section Pseudonarcissus consists of relatively large flowers with a corolla length of around , generally solitary but rarely in inflorescences of 2–4 flowers. They have wide greenish floral tubes with funnel-shaped bright yellow coronas. The six tepals sometimes differ in colour from the corona and may be cream coloured to pale yellow. The "paperwhite" form, including sections Jonquilla, Apodanthi and Narcissus, has a relatively long, narrow tube and a short, shallow, flaring corona. The flower is horizontal and fragrant. The "triandrus" form is seen in only two species, N. albimarginatus (a Moroccan endemic) and N. triandrus. It combines features of both the "daffodil" and "paperwhite" forms, with a well-developed, long, narrow tube and an extended bell-shaped corona of almost equal length. The flowers are pendent. ; Androecium : There are six stamens in one to two rows (whorls), with the filaments separate from the corona, attached at the throat or base of the tube (epipetalous), often of two separate lengths, straight or declinate-ascending (curving downwards, then upwards). The anthers are basifixed (attached at their base). ; Gynoecium : The ovary is inferior (below the floral parts) and trilocular (three chambered) and there is a pistil with a minutely three lobed stigma and filiform (thread like) style, which is often exserted (extending beyond the tube). ; Fruit : The fruit consists of dehiscent loculicidal capsules (splitting between the locules) that are ellipsoid to subglobose (almost spherical) in shape and are papery to leathery in texture. ; Seeds : The fruit contains numerous subglobose seeds which are round and swollen with a hard coat, sometimes with an attached elaiosome. The testa is black and the pericarp dry. Most species have 12 ovules and 36 seeds, although some species such as N. bulbocodium have more, up to a maximum of 60. Seeds take five to six weeks to mature. The seeds of sections Jonquilla and Bulbocodium are wedge-shaped and matt black, while those of other sections are ovate and glossy black. A gust of wind or contact with a passing animal is sufficient to disperse the mature seeds. Chromosomes Chromosome numbers include 2n=14, 22, 26, with numerous aneuploid and polyploid derivatives. The basic chromosome number is 7, with the exception of N. tazetta, N. elegans and N. broussonetii in which it is 10 or 11; this subgenus (Hermione) was in fact characterised by this characteristic. Polyploid species include N. papyraceus (4x=22) and N. dubius (6x=50). Phytochemistry Alkaloids As with all Amaryllidaceae genera, Narcissus contains unique isoquinoline alkaloids. The first alkaloid to be identified was lycorine, from N. pseudonarcissus in 1877. These are considered a protective adaptation and are utilised in the classification of species. Nearly 100 alkaloids have been identified in the genus, about a third of all known Amaryllidaceae alkaloids, although not all species have been tested. Of the nine alkaloid ring types identified in the family, Narcissus species most commonly demonstrate the presence of alkaloids from within the Lycorine (lycorine, galanthine, pluviine) and Homolycorine (homolycorine, lycorenine) groups. Hemanthamine, tazettine, narciclasine, montanine and galantamine alkaloids are also represented. The alkaloid profile of any plant varies with time, location, and developmental stage. Narcissus also contain fructans and low molecular weight glucomannan in the leaves and plant stems. Fragrances Fragrances are predominantly monoterpene isoprenoids, with a small amount of benzenoids, although N. jonquilla has both equally represented. Another exception is N. cuatrecasasii which produces mainly fatty acid derivatives. The basic monoterpene precursor is geranyl pyrophosphate, and the commonest monoterpenes are limonene, myrcene, and trans-β-ocimene. Most benzenoids are non-methoxylated, while a few species contain methoxylated forms (ethers), e.g. N. bujei. Other ingredients include indole, isopentenoids and very small amounts of sesquiterpenes. Fragrance patterns can be correlated with pollinators, and fall into three main groups (see Pollination). ==Taxonomy==

History Early The genus Narcissus was well known to the ancient Greeks and Romans. In Greek literature Theophrastus and Dioscorides is from Greek narkissos. According to Plutarch narkissos has been connected because of the plant's narcotic properties, with narkē "numbness"; it may also be connected with hell. It is frequently linked to the myth of Narcissus, who became so obsessed with his own reflection in water that he drowned and the narcissus plant sprang from where he died. There is no evidence for the flower being named after Narcissus. Narcissus poeticus, which grows in Greece, has a fragrance that has been described as intoxicating. Furthermore, there were accounts of narcissi growing long before the story of Narcissus appeared (see Greek culture). It has also been suggested that narcissi bending over streams represent the youth admiring his reflection. The narcissus was frequently referred to as the asphodel The reason for the introduction of the initial "d" is not known. == Distribution and habitat ==

Distribution '' in Israel Although the family Amaryllidaceae are predominantly tropical or subtropical as a whole, Narcissus occurs primarily in the Mediterranean region, with a centre of diversity in the Iberian Peninsula (Spain and Portugal). N. elegans occurs on the Northwest African Coast (Morocco and Libya), as well as the coastline of Corsica, Sardinia and Italy, and N. bulbocodium between Tangier and Algiers and Tangier to Marrakesh, but also on the Iberian Peninsula. N. serotinus is found along the entire Mediterranean coast. N. tazetta occurs as far east as Iran and Kashmir. Since this is one of the oldest species found in cultivation, it is likely to have been introduced into Kashmir. N. poeticus and N. pseudonarcissus have the largest distribution ranges. N. poeticus ranges from the Pyrenees along the Romanian Carpathians to the Black Sea and along the Dalmatian coast to Greece. N. pseudonarcissus ranges from the Iberian Peninsula, via the Vosges Mountains to northern France and Belgium, and the United Kingdom where there are still wild stocks in Southern Scotland. The only occurrence in Luxembourg is located near Lellingen, in the municipality of Kiischpelt. In Germany it is found mainly in the nature reserve at Perlenbach-Fuhrtsbachtal and the Eifel National Park, where in the spring at Monschau the meadows are teeming with yellow blooms. One of the most easterly occurrences can be found at Misselberg near Nassau on the Lahn. '' growing in Međulići, near Gacko, Bosnia and Herzegovina However, unlike the above examples, most species have very restricted endemic ranges which may overlap resulting in natural hybrids. For instance in the vicinity of the Portuguese city of Porto where both N. pseudonarcissus and N. triandrus occur there are found various intersections of the two species while in a small area along part of the Portuguese Mondego river are found intersectional hybrids between N. scaberulus and N. triandrus. The biogeography demonstrates a phylogenetic association, for instance subgenus Hermione having a lowland distribution, but subgenus Narcissus section Apodanthi being montane and restricted to Morocco, Spain and Portugal. The remaining sections within subgenus Narcissus include both lowland and mountain habitats. Section Pseudonarcissus, although widely naturalised, is endemic to the Baetic Ranges of the southeastern Iberian Peninsula. Habitats Their native habitats are very varied, with different elevations, bioclimatic areas and substrates, being found predominantly in open spaces ranging from low marshes to rocky hillsides and montane pastures, and including grassland, scrub, woods, river banks and rocky crevices. Although requirements vary, overall there is a preference for acidic soils, although some species will grow on limestone. Narcissus scaberulus will grow on granite soils where it is moist in the growing season but dry in the summer, while Narcissus dubius thrives best in regions with hot and dry summers. The Pseudonarcissus group in their natural habitat prefers humid situations such as stream margins, springs, wet pastures, clearings of forests or shrublands with humid soils, and moist hillsides. These habitats tend to be discontinuous in the Mediterranean mountains, producing discrete isolated populations. In Germany, which has relatively little limestone, Narcissus pseudonarcissus grows in small groups on open mountain meadows or in mixed forests of fir, beech, oak, alder, ash and birch trees with well-drained soil. == Ecology ==

Life cycle Narcissus are long-lived perennial geophytes with winter-growing and summer-dormant bulbs Narcissus latent virus (NLV, Narcissus mild mottle virus) which causes green mottling near leaf tips, Narcissus degeneration virus (NDV), Narcissus late season yellows virus (NLSYV) which occurs after flowering, streaking the leaves and stems, Narcissus mosaic virus, Narcissus yellow stripe virus (NYSV, Narcissus yellow streak virus), Narcissus tip necrosis virus (NTNV) which produces necrosis of leaf tips after flowering and Narcissus white streak virus (NWSV). Less host specific viruses include Raspberry ringspot virus, Nerine latent virus (NeLV) =Narcissus symptomless virus, Arabis mosaic virus (ArMV), Broad Bean Wilt Viruses (BBWV) Cucumber mosaic virus (CMV), Tomato black ring virus (TBRV), Tomato ringspot virus (TomRSV) and Tobacco rattle virus (TRV). Of these viruses the most serious and prevalent are NDV, NYSV and NWSV. NDV is associated with chlorotic leaf striping in N. tazetta. Infection with NYSV produces light or grayish-green, or yellow stripes or mottles on the upper two-thirds of the leaf, which may be roughened or twisted. The flowers which may be smaller than usual may also be streaked or blotched. NWSV produces greenish-purple streaking on the leaves and stem turning white to yellow, and premature senescence reducing bulb size and yield. These viruses are primarily diseases of commercial nurseries. The growth inhibition caused by viral infection can cause substantial economic damage. ; Bacteria : Bacterial disease is uncommon in Narcissus but includes Pseudomonas (bacterial streak) and Pectobacterium carotovorum sp. carotovorum (bacterial soft rot). ; Fungi : More problematic for non-commercial plants is the fungus, Fusarium oxysporum f. sp. narcissi, which causes basal rot (rotting of the bulbs and yellowing of the leaves). This is the most serious disease of Narcissus. Since the fungus can remain in the soil for many years it is necessary to remove infected plants immediately, and to avoid planting further narcissi at that spot for a further five years. Not all species and cultivars are equally susceptible. Relatively resistant forms include N. triandrus, N. tazetta and N. jonquilla. Another fungus which attacks the bulbs, causing narcissus smoulder, is Botrytis narcissicola (Sclerotinia narcissicola) and other species of Botrytis, including Botrytis cinerea, particularly if improperly stored. Copper sulfate is used to combat the disease, and infected bulbs are burned. Blue mould rot of bulbs may be caused by infection with species of Penicillium, if they have become damaged either through mechanical injury or infestation by mites (see below). Species of Rhizopus (e.g. Rhizopus stolonifer, Rhizopus nigricans) cause bulb soft rot and Sclerotinia bulborum, black slime disease. A combination of both Peyronellaea curtisii (Stagonosporopsis curtisii) and Botrytis narcissicola causes neck rot in the bulbs. Fungi affecting the roots include Nectria radicicola (Cylindrocarpon destructans), a cause of root rot and Rosellinia necatrix causing white root rot, while others affect root and bulb, such as Aspergillus niger (black mold), and species of Trichoderma, including T. viride and T. harzianum (=T. narcissi) responsible for green mold. Other fungi affect the remainder of the plant. Another Botrytis fungus, Botrytis polyblastis (Sclerotinia polyblastis) causes brown spots on the flower buds and stems (narcissus fire), especially in damp weather and is a threat to the cut flower industry. Ramularia vallisumbrosae is a leaf spot fungus found in warmer climates, causing narcissus white mould disease. Peyronellaea curtisii, the Narcissus leaf scorch, also affects the leaves as does its synanamorph, Phoma narcissi (leaf tip blight). Aecidium narcissi causes rust lesions on leaves and stems. ; Animals : Arthropods that are Narcissus pests include insects such as three species of fly that have larvae that attack the plants, the narcissus bulb fly Merodon equestris, and two species of hoverflies, the lesser bulb flies Eumerus tuberculatus and Eumerus strigatus. The flies lay their eggs at the end of June in the ground around the narcissi, a single female fly being able to lay up to fifty eggs. The hatching larvae then burrow through the soil towards the bulbs and consume their interiors. They then overwinter in the empty bulb shell, emerging in April to pupate in the soil, from which the adult fly emerges in May. The larvae of some moths such as Korscheltellus lupulina (the common swift moth) attack Narcissus bulbs. Other arthropods include Mites such as Steneotarsonemus laticeps (Bulb scale mite), Rhizoglyphus and Histiostoma infest mainly stored bulbs and multiply particularly at high ambient temperature, but do not attack planted bulbs. Planted bulbs are susceptible to nematodes, the most serious of which is Ditylenchus dipsaci (Narcissus eelworm), the main cause of basal plate disease in which the leaves turn yellow and become misshapen. Infested bulbs have to be destroyed; where infestation is heavy avoiding planting further narcissi for another five years. Other nematodes include Aphelenchoides subtenuis, which penetrates the roots causing basal plate disease and Pratylenchus penetrans (lesion nematode) the main cause of root rot in narcissi. Other nematodes such as the longodorids (Longidorus spp. or needle nematodes and Xiphinema spp. or dagger nematodes) and the stubby-root nematodes or trichodorids (Paratrichodorus spp. and Trichodorus spp.) can also act as vectors of virus diseases, such as TBRV and TomRSV, in addition to causing stunting of the roots. Gastropods such as snails and slugs also cause damage to growth. == Conservation ==

Many of the smallest species have become extinct, requiring vigilance in the conservation of the wild species. The IUCN Red List describes five species as 'Endangered' (Narcissus alcaracensis, Narcissus bujei, Narcissus longispathus, Narcissus nevadensis, Narcissus radinganorum). In 1999 three species were considered endangered, five as vulnerable and six as rare. In response, a number of species have been granted protected species status and protected areas (meadows) have been established such as the Negraşi Daffodil Meadow in Romania, or Kempley Daffodil Meadow in the UK. These areas often host daffodil festivals in the spring. ==Cultivation==

History File:Gerard N minor serotinus.jpg|N. serotinus, John Gerard, The Herball 1597 File:Hortus Eystettensis, 1640 (BHL 45339 063) - Classis Verna 52.jpg|Narcissi, Hortus Eystettensis 1613 File:Hale Plate 42 Poetick Daffodil.jpg|N. poeticus, Thomas Hale, Eden: Or, a Compleat Body of Gardening 1757 File:Lauremberg.png|Narcissus, Peter Lauremberg 1632 File:ParkinsonNarcissus101.jpg|Narcissi, John Parkinson, Paradisus Terrestris 1629. (8. Great Double Yellow Spanish Daffodil) Of all the flowering plants, the bulbous have been the most popular for cultivation. Of these, narcissi are one of the most important spring flowering bulb plants in the world. Indigenous in Europe, the wild populations of the parent species had been known since antiquity. Narcissi have been cultivated from at least as early as the sixteenth century in the Netherlands, when large numbers of bulbs where imported from the field, particularly Narcissus hispanicus, which soon became nearly extinct in its native habitat of France and Spain, though still found in the southern part of that country. Narcissi are well suited for planting under small thickets of trees, where they can be grouped as 6–12 bulbs. They also grow well in perennial borders, especially in association with day lilies which begin to form their leaves as the narcissi flowers are fading. A number of wild species and hybrids such as "Dutch Master", "Golden Harvest", "Carlton", "Kings Court" and "Yellow Sun" naturalise well in lawns, but it is important not to mow the lawn till the leaves start to fade, since they are essential for nourishing the bulb for the next flowering season. Blue Scilla and Muscari which also naturalise well in lawns and flower at the same time as narcissus, make an attractive contrast to the yellow flowers of the latter. Unlike tulips, narcissi bulbs are not attractive to rodents and are sometimes planted near tree roots in orchards to protect them. Propagation The commonest form of commercial propagation is by twin-scaling, in which the bulbs are cut into many small pieces but with the two scales still connected by a small fragment of the basal plate. The fragments are disinfected and placed in nutrient media. Some 25–35 new plants can be produced from a single bulb after four years. Micropropagation methods are not used for commercial production but are used for establishing commercial stock. Breeding For commercial use, varieties with a minimum stem length of are sought, making them ideal for cut flowers. Florists require blooms that only open when they reach the retail outlet. For garden plants the objectives are to continually expand the colour palette and to produce hardy forms, and there is a particular demand for miniature varieties. The cultivars so produced tend to be larger and more robust than the wild types. The main species used in breeding are N. bulbocodium, N. cyclamineus, N. jonquilla, N. poeticus, N. pseudonarcissus, N. serotinus and N. tazetta. Narcissus pseudonarcissus gave rise to trumpet cultivars with coloured tepals and corona, while its subspecies N. pseudonarcissus subsp. bicolor was used for white tepaled varieties. To produce large cupped varieties, N. pseudonarcissus was crossed with N. poeticus, and to produce small cupped varieties back crossed with N. poeticus. Multiheaded varieties, often called "Poetaz" are mainly hybrids of N. poeticus and N. tazetta. Classification For horticultural purposes, all Narcissus cultivars are split into 13 divisions as first described by Kington (1998), for the Royal Horticultural Society (RHS), based partly upon flower form (shape and length of corona), number of flowers per stem, flowering period and partly upon genetic background. Division 13, which includes wild daffodils, is the exception to this scheme. Growers register new daffodil cultivars by name and colour with the Royal Horticultural Society, which is the international registration authority for the genus. Their International Daffodil Register is regularly updated with supplements available online and is searchable. The most recent supplement (2014) is the sixth (the fifth was published in 2012). More than 27,000 names were registered as of 2008, and the number has continued to grow. Registered daffodils are given a division number and colour code such as 5 W-W ("Thalia"). In horticultural usage it is common to also find an unofficial Division 14: Miniatures, which although drawn from the other 13 divisions, have their miniature size in common. Over 140 varieties have gained the Royal Horticultural Society's Award of Garden Merit (See List of Award of Garden Merit narcissus). Colour code Daffodil breeding has introduced a wide range of colours, in both the outer perianth tepal segment and the inner corona. In the registry, daffodils are coded by the colours of each of these two parts. Thus "Geranium", Tazetta (Division 8) as illustrated here with a white outer perianth and orange corona is classified as 8 W-O. == Toxicity ==

Pharmacology All Narcissus species contain the alkaloid poison lycorine, mostly in the bulb but also in the leaves. Members of the monocot subfamily Amaryllidoideae present a unique type of alkaloids, the norbelladine alkaloids, which are 4-methylcatechol derivatives combined with tyrosine. They are responsible for the poisonous properties of a number of the species. Over 200 different chemical structures of these compounds are known, of which 79 or more are known from Narcissus alone. The toxic effects of ingesting Narcissus products for both humans and animals (such as cattle, goats, pigs, and cats) have long been recognised and they have been used in suicide attempts. Ingestion of N. pseudonarcissus or N. jonquilla is followed by salivation, acute abdominal pains, nausea, vomiting, and diarrhea, then neurological and cardiac events, including trembling, convulsions, and paralysis. Death may result if large quantities are consumed. The toxicity of Narcissus varies with species, N. poeticus being more toxic than N. pseudonarcissus, for instance. The distribution of toxins within the plant also varies, for instance, there is a five times higher concentration of alkaloid in the stem of N. papyraceus than in the bulb, making it dangerous to herbivores more likely to consume the stem than the bulb, and is part of the plant's defence mechanisms. The distribution of alkaloids within tissues may also reflect defence against parasites. The bulbs can also be toxic to other nearby plants, including roses, rice, and cabbages, inhibiting growth. For instance placing cut flowers in a vase alongside other flowers shortens the life of the latter. Poisoning Many cases of poisoning or death have occurred when narcissi bulbs have been mistaken for leeks or onions and cooked and eaten. Recovery is usually complete in a few hours without any specific intervention. In more severe cases involving ingestion of large quantities of bulbs, activated carbon, salts and laxatives may be required, and for severe symptoms intravenous atropine and emetics or stomach pumping may be indicated. However, ingestion of large quantities accidentally is unusual because of a strong unpleasant taste. When narcissi were compared with a number of other plants not normally consumed by animals, narcissi were the most repellent, specifically N. pseudonarcissus. Consequently, narcissus alkaloids have been used as repellents and may also discourage fungi, molds, and bacteria. Topical effects One of the most common dermatitis problems for flower pickers, packers, florists, and gardeners, "daffodil itch", involves dryness, fissures, scaling, and erythema in the hands, often accompanied by subungual hyperkeratosis (thickening of the skin beneath the nails). It is blamed on exposure to calcium oxalate, chelidonic acid or alkaloids such as lycorine in the sap, either due to a direct irritant effect or an allergic reaction. It has long been recognised that some cultivars provoke dermatitis more readily than others. N. pseudonarcissus and the cultivars "Actaea", "Camparelle", "Gloriosa", "Grande Monarque", "Ornatus", "Princeps" and "Scilly White" are known to do so. If bulb extracts come into contact with wounds, both central nervous system and cardiac symptoms may result. The scent can also cause toxic reactions such as headaches and vomiting from N. bulbocodium. ==Uses==

Traditional medicine Despite the lethal potential of Narcissus alkaloids, they have been used for centuries as traditional medicines for a variety of complaints, including cancer. Plants thought to be N. poeticus and N. tazetta are described in the Bible in the treatment for what is thought to be cancer. In the Classical Greek world Hippocrates (c. BC 460–370) recommended a pessary prepared from narcissus oil for uterine tumors, a practice continued by Pedanius Dioscorides (c. AD 40–90) and Soranus of Ephesus (AD 98–138) in the first and second centuries AD, while the Roman Pliny the Elder (AD 23–79), advocated topical use. The bulbs of N. poeticus contain the antineoplastic agent narciclasine. This usage is also found in later Arabian, North African, Central American and Chinese medicine during the Middle Ages. In China N. tazetta var. chinensis was grown as an ornamental plant but the bulbs were applied topically to tumors in traditional folk medicine. These bulbs contain pretazettine, an active antitumor compound. Narcissus products have received a variety of other uses. The Roman physician Aulus Cornelius Celsus listed narcissus root in De Medicina among medical herbs, described as emollient, erodent, and "powerful to disperse whatever has collected in any part of the body". N. tazetta bulbs were used in Turkey as a remedy for abscesses in the belief they were antiphlogistic and analgesic. Other uses include the application to wounds, strains, painful joints, and various local ailments as an ointment called 'Narcissimum'. Powdered flowers have also been used medically, as an emetic, a decongestant and for the relief of dysentery, in the form of a syrup or infusion. The French used the flowers as an antispasmodic, the Arabs the oil for baldness and also an aphrodisiac. In the eighteenth century the Irish herbal of John K'Eogh recommended pounding the roots in honey for use on burns, bruises, dislocations and freckles, and for drawing out thorns and splinters. N. tazetta bulbs have also been used for contraception, while the flowers have been recommended for hysteria and epilepsy. In the traditional Japanese medicine of kampo, wounds were treated with narcissus root and wheat flour paste; the plant, however, does not appear in the modern kampo herb list. There is also a long history of the use of Narcissus as a stimulant and to induce trance like states and hallucinations. Sophocles referred to the narcissus as the "Chaplet of the infernal Gods", a statement frequently wrongly attributed to Socrates (see Antiquity). Biological properties Extracts of Narcissus have demonstrated a number of potentially useful biological properties including antiviral, prophage induction, antibacterial, antifungal, antimalarial, insecticidal, cytotoxic, antitumor, antimitotic, antiplatelet, hypotensive, emetic, acetylcholine esterase inhibitory, antifertility, antinociceptive, chronotropic, pheromone, plant growth inhibitor, and allelopathic. An ethanol extract of Narcissus bulbs was found effective in one mouse model of nociception, para-benzoquinone induced abdominal constriction, but not in another, the hot plate test. Most of these properties are due to alkaloids, but some are also due to mannose-binding lectins. The most-studied alkaloids in this group are galantamine (galanthamine), lycorine, narciclasine, and pretazettine. It is likely that the traditional use of narcissi for the treatment of cancer was due to the presence of isocarbostyril constituents such as narciclasine, pancratistatin and their congeners. N. poeticus contains about 0.12 g of narciclasine per kg of fresh bulbs. Acetylcholine esterase inhibition has attracted the most interest as a possible therapeutic intervention, with activity varying by a thousandfold between species, and the greatest activity seen in those that contain galantamine or epinorgalanthamine. The rodent repellant properties of Narcissus alkaloids have been utilised in horticulture to protect more vulnerable bulbs. Therapeutics Of all the alkaloids, only galantamine has made it to therapeutic use in humans, as the drug galantamine for Alzheimer's disease. Galantamine is an acetylcholine esterase inhibitor which crosses the blood brain barrier and is active within the central nervous system. Daffodils are grown commercially near Brecon in Powys, Wales, to produce galantamine. Commercial uses Throughout history the scent of narcissi has been an important ingredient of perfumes, a quality that comes from essential oils rather than alkaloids. Narcissi are also an important horticultural crop, and source of cut flowers (floriculture). The Netherlands, which is the most important source of flower bulbs worldwide is also a major centre of narcissus production. Of 16,700 hectares (ha) under cultivation for flower bulbs, narcissi account for about 1,800 hectares. In the 1990s narcissus bulb production was at 260 million, sixth in size after tulips, gladioli, irises, crocuses and lilies and in 2012 it was ranked third. About two-thirds of the area under cultivation is dedicated to about 20 of the most popular varieties. In the 2009/2010 season, 470 cultivars were produced on 1578 ha. By far the largest area cultivated is for the miniature 'Tête-à-Tête', followed at some distance by 'Carlton'. The largest production cultivars are shown in Table II. "Carlton" and "Ice Follies" (Division 2: Large cup) have a long history of cultivation, together with "Dutch Master" and "Golden Harvest" (1: yellow). "Carlton" and "Golden Harvest" were introduced in 1927, and "Ice Follies" in 1953. "Carlton", with over 9 billion bulbs (350 000 tons), is among the more numerous individual plants produced in the world. The other major areas of production are the United States, Israel which exported 25 million N. tazetta cultivar bulbs in 2003, and the United Kingdom. In the United Kingdom a total of 4100 ha were planted with bulbs, of which 3800 ha were Narcissi, the UK's most important bulb crop, much of which is for export, making this the largest global production centre, about half of the total production area. While some of the production is for forcing, most is for dry bulb production. Bulb production and forcing occurs in the East, while production in the south west is mainly for outdoor flower production. The farm gate value was estimated at £10m in 2007. Production of both bulbs and cut flowers takes place in open fields in beds or ridges, often in the same field, allowing adaptation to changing market conditions. Narcissi grow best in mild maritime climates. Compared to the United Kingdom, the harsher winters in the Netherlands require covering the fields with straw for protection. Areas with higher rainfall and temperatures are more susceptible to diseases that attack crops. Production is based on a 1 (UK) or 2 (Netherlands) year cycle. Optimal soil pH is 6.0–7.5. Prior to planting disinfection by hot water takes place, such as immersion at 44.4 °C for three hours. Bulbs are harvested for market in the summer, sorted, stored for 2–3 weeks, and then further disinfected by a hot (43.5 °C) bath. This eliminates infestations by narcissus fly and nematodes. The bulbs are then dried at a high temperature, and then stored at 15.5 °C. The initiation of new flower development in the bulb takes place in late spring before the bulbs are lifted, and is completed by mid summer while the bulbs are in storage. The optimal temperature for initiation is 20 °C followed by cooling to 13 °C. Traditionally, sales took place in the daffodil fields prior to harvesting the bulbs, but today sales are handled by Marketing Boards although still before harvesting. In the Netherlands there are special exhibition gardens for major buyers to view flowers and order bulbs, some larger ones may have more than a thousand narcissus varieties on display. While individuals can visit these gardens they cannot buy bulbs at retail, which are only available at wholesale, usually at a minimum of several hundredweight. The most famous display is at Keukenhof, although only about 100 narcissus varieties are on display there. Forcing There is also a market for forced blooms, both as cut flowers and potted flowers through the winter from Christmas to Easter, the long season requiring special preparation by growers. Cut flowers For cut flowers, bulbs larger than 12 cm in size are preferred. To bloom in December, bulbs are harvested in June to July, dried, stored for four days at 34 °C, two weeks at 30 and two weeks at 17–20 °C and then placed in cold storage for precooling at 9 degrees for about 15–16 weeks. The bulbs are then planted in light compost in crates in a greenhouse for forcing at 13–15 °C and the blooms appear in 19–30 days. Potted flowers For potted flowers a lower temperature is used for precooling (5 °C for 15 weeks), followed by 16–18 °C in a greenhouse. For later blooming (mid- and late-forcing), bulbs are harvested in July to August and the higher temperatures are omitted, being stored a 17–20 °C after harvesting and placed in cold storage at 9 °C in September for 17–18 (cut flowers) or 14–16 (potted flowers) weeks. The bulbs can then be planted in cold frames, and then forced in a greenhouse according to requirements. N. tazetta and its cultivars are an exception to this rule, requiring no cold period. Often harvested in October, bulbs are lifted in May and dried and heated to 30 °C for three weeks, then stored at 25 °C for 12 weeks and planted. Flowering can be delayed by storing at 5–10 °C. == Culture ==

Symbols '' 'Thalia', considered a grave flower The daffodil is the national flower of Wales, associated with Saint David's Day (1 March). The narcissus is also a national flower symbolising the new year or Nowruz in the Kurdish culture. In the West the narcissus is perceived as a symbol of vanity, in the East as a symbol of wealth and good fortune , while in Persian literature, the narcissus is a symbol of beautiful eyes. In western countries the daffodil is also associated with spring festivals such as Lent and its successor Easter. In Germany the wild narcissus, N. pseudonarcissus, is known as the Osterglocke or "Easter bell". In the United Kingdom the daffodil is sometimes referred to as the Lenten lily. Although prized as an ornamental flower, some people consider narcissi unlucky, because they hang their heads implying misfortune. A similar account is provided by Lucian describing the flowers in the underworld. The Persian phrase (, literally "a reddish-blue narcissus") in the classical poetries of the Persian, Urdu, Ottoman Turkish, Azerbaijani and Chagatai languages; to this day also the vernacular names of some narcissus cultivars (for example, Shahla-ye Shiraz and Shahla-ye Kazerun). As described by the poet Ghalib (1797–1869), "God has given the eye of the narcissus the power of seeing". Narcissus is also considered a symbol in Shia Islam and likewise in Iran for Mahdi. Narcissus tazetta var chinensis1.jpg|Chinese Sacred Lily Shuixiandiaoke.JPG|Chinese decorative carved Narcissus bulb Narcissus poeticus.jpg|N. poeticus symbolising the eye in Islamic culture Popular culture The word "daffodil" has been used widely in popular culture, from Dutch cars (DAF Daffodil) to films (Daffodils) to slurs against homosexuals and cross-dressers (as in the film J. Edgar, when Hoover's mother explains why real-life cross-dresser Barton Pinkus was called "Daffy" (short for "Daffodil" and the equivalent of a pansy), and admonishes, "I'd rather have a dead son than a daffodil for a son"). Festivals In some areas where narcissi are prevalent, their blooming in spring is celebrated in festivals. For instance, the slopes around Montreux, Switzerland and its associated riviera come alive with blooms each May (May Snow) at the annual Narcissi Festival. Festivals are also held in many other countries. Cancer Various cancer charities around the world, such as the American Cancer Society, Cancer Society, Cancer Council, Irish Cancer Society, and Marie Curie in the UK use the daffodil as a fundraising symbol on "Daffodil Days". == See also ==