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Raphides (pronounced /ˈræfɪˌdiz/, singular raphide /ˈreɪfʌɪd/ or raphis) are needle-shaped crystals of calcium oxalate monohydrate (
prismatic An optical prism is a transparent optical element with flat, polished surfaces that are designed to refract light. At least one surface must be angled — elements with two parallel surfaces are ''not'' prisms. The most familiar type of optical ...
monoclinic In crystallography, the monoclinic crystal system is one of the seven crystal systems. A crystal system is described by three vectors. In the monoclinic system, the crystal is described by vectors of unequal lengths, as in the orthorhombic s ...
crystals) or calcium carbonate as aragonite ( dipyramidal orthorhombic crystals), found in more than 200 families of plants. Both ends are needle-like, but raphides tend to be blunt at one end and sharp at the other.


Calcium oxalate in plants

Many plants accumulate calcium oxalate crystals in response to surplus calcium, which is found throughout the natural environment. The crystals are produced in a variety of shapes. The crystal morphology depends on the taxonomic group of the plant. In one study of over 100 species, it was found that calcium oxalate accounted for 6.3% of plant dry weight. Crystal morphology and the distribution of raphides (in roots or leaves or tubers etc.) is similar in some taxa but different in others leaving possible opportunities for plant key characteristics and systematic identification; mucilage in raphide containing cells makes light microscopy difficult, though. Little is known about the mechanisms of sequestration or indeed the reason for accumulation of raphides but it is most likely as a defense mechanism against herbivory. It has also been suggested that in some cases raphides may help form plant skeletal structure. Raphides typically occur in parenchyma cells in aerial organs especially the leaves, and are generally confined to the mesophyll. As the leaf area increases, so does the number of raphides, the process starting in even young leaves. The first indications that the cell will contain crystals is shown when the cells enlarge with a larger nucleus. Raphides are found in specialized plant cells or crystal chambers called
idioblast An idioblast is an isolated plant cell that differs from neighboring tissues. They have various functions such as storage of reserves, excretory materials, pigments, and minerals. They could contain oil, latex, gum, resin, tannin or pigments etc. ...
s. Also, Electron micrographs have shown that raphide needle crystals are normally four sided or H-shaped (with a groove down both sides) or with a hexagonal cross section and some are barbed. Wattendorf (1976) suggested that all circular sectioned raphides, as visible in a light microscope, are probably hexagonal in cross section The hexagonal crystals reported by Wattendorf in ''
Agave americana ''Agave americana'', common names century plant, maguey, or American aloe, is a species of flowering plant in the family Asparagaceae, native to Mexico and the United States in Texas. It is cultivated worldwide as an ornamental plant, and has b ...
'' were apparently calcium oxalate. Microscopy using polarized light shows bright opalescence with raphides. Plants like '' Tradescantia pallida'' also accumulate calcium oxalate crystals in response to heavy metals stress.


Harmful effects

Raphides can produce severe toxic reactions by facilitating the passage of toxin through the herbivore's skin when the tissue containing the raphides also contains toxins. The lethal dose to mice is around 15 mg/kg. Raphides seem to be a defense mechanism against plant predators, as they are likely to tear and harm the soft tissues of the throat or esophagus of a plant predator chewing on the plant's leaves. The venomous process is in two stages: mechanical pricking and injection of harmful protease. Typically ingestion of plants containing raphides, like those common in certain houseplants, can cause immediate numbing followed shortly by painful edema, vesicle formation and
dysphagia Dysphagia is difficulty in swallowing. Although classified under "symptoms and signs" in ICD-10, in some contexts it is classified as a condition in its own right. It may be a sensation that suggests difficulty in the passage of solids or liq ...
accompanied by painful stinging and burning to the mouth and throat with symptoms occurring for up to two weeks. Airway assessment and management are of the highest priority, as are extensive irrigation and analgesics in eye exposure. Raphides cannot normally be destroyed by boiling; that requires an acidic environment or chemical solvents like ether. Plants containing large amounts of raphides are generally acrid and unpalatable. However, it is not always possible to detect the presence of raphides through taste alone. In some tubers such as Indian turnip which contain large quantities of raphides, the roots are not unpalatable when cooked because the raphides are bound within a matrix of starch which prevents the tongue from sensing their presence. Some other plants store raphides in mucilaginous environments and also do not taste acrid.


Plants containing raphides

Common names vary. The following list is incomplete. Raphides are found in many species in the families
Araceae The Araceae are a family of monocotyledonous flowering plants in which flowers are borne on a type of inflorescence called a spadix. The spadix is usually accompanied by, and sometimes partially enclosed in, a spathe (or leaf-like bract). A ...
and
Commelinaceae Commelinaceae is a family of flowering plants. In less formal contexts, the group is referred to as the dayflower family or spiderwort family. It is one of five families in the order Commelinales and by far the largest of these with about 731 kno ...
, but are also found in a few species in a number of other families.
Actinidiaceae The Actinidiaceae are a small family of flowering plants. The family has three genera and about 360 species and is a member of the order Ericales. Distribution They are temperate and subtropical woody vines, shrubs, and trees, native to As ...
: * '' Actinidia'' (Kiwifruit)
Amaranthaceae Amaranthaceae is a family of flowering plants commonly known as the amaranth family, in reference to its type genus ''Amaranthus''. It includes the former goosefoot family Chenopodiaceae and contains about 165 genera and 2,040 species, making it ...
: * '' Beta'' (Beets, Swiss Chard) * ''
Spinacia ''Spinacia'' is a flowering plant genus in the subfamily Chenopodioideae of the family Amaranthaceae. The most common member is spinach. Description The species in genus ''Spinacia'' are annual or biennial herbs. Plants are always glabrous. ...
'' (Spinach)
Araceae The Araceae are a family of monocotyledonous flowering plants in which flowers are borne on a type of inflorescence called a spadix. The spadix is usually accompanied by, and sometimes partially enclosed in, a spathe (or leaf-like bract). A ...
: * ''
Alocasia ''Alocasia'' is a genus of rhizomatous or tuberous, broad-leaved, perennial, flowering plants from the family Araceae. There are about 90 accepted species native to tropical and subtropical Asia and eastern Australia. Around the world, many gro ...
'' (Baibing in NE India) * ''
Arisaema ''Arisaema'' is a large and diverse genus of the flowering plant family Araceae. The largest concentration of species is in China and Japan, with other species native to other parts of southern Asia as well as eastern and central Africa, Mexico ...
'' (Cobra Lily) * ''
Arum ''Arum'' is a genus of flowering plants in the family Araceae, native to Europe, northern Africa, and western and central Asia, with the highest species diversity in the Mediterranean region. Frequently called arum lilies, they are not closely ...
'' (Cuckoopint) * ''
Caladium ''Caladium'' is a genus of flowering plants in the family Araceae. They are often known by the common name elephant ear (which they share with the closely related genera '' Alocasia'', ''Colocasia'', and ''Xanthosoma''), heart of Jesus, and ange ...
'' (Angel wings) * ''
Colocasia ''Colocasia'' is a genus of flowering plants in the family Araceae, native to southeastern Asia and the Indian subcontinent. Some species are widely cultivated and naturalized in other tropical and subtropical regions. The names elephant-ear ...
'' (Taro, Cocoyam, Eddoe) * ''
Dieffenbachia ''Dieffenbachia'' , commonly known as dumb cane or leopard lily, is a genus of tropical flowering plants in the family Araceae. It is native to the New World Tropics from Mexico and the West Indies south to Argentina. Some species are widely cul ...
'' (Dumb Cane) * ''
Epipremnum ''Epipremnum'' is a genus of flowering plants in the family (biology), family Araceae, found in tropics, tropical forests from China, the Himalayas, and Southeast Asia to Australia the western Pacific. They are evergreen perennial plant, perennia ...
'' (Devil's Ivy, Pothos) * ''
Monstera ''Monstera'' is a genus of 59 species of flowering plants in the arum family, Araceae, native to tropical regions of the Americas. Etymology The genus is named from the Latin word for "monstrous" or "abnormal", and refers to the unusual leaves ...
'' (Swiss Cheese Plant) * ''
Philodendron ''Philodendron'' is a large genus of flowering plants in the family Araceae. , the World Checklist of Selected Plant Families accepted 489 species; other sources accept different numbers. Regardless of number of species, the genus is the second- ...
'' (Elephant Ears) * ''
Spathiphyllum ''Spathiphyllum'' is a genus of about 47 species of monocotyledonous flowering plants in the family Araceae, native to tropical regions of the Americas and southeastern Asia. Certain species of ''Spathiphyllum'' are commonly known as spath or pea ...
'' (Peace Lily) * ''
Zantedeschia ''Zantedeschia'' is a genus of eight species of herbaceous, perennial, flowering plants in the family Araceae, native to southern Africa from South Africa north to Malawi. The genus has been introduced on all continents except Antarctica. Com ...
'' (Calla Lily)
Araliaceae The Araliaceae are a family of flowering plants composed of about 43 genera and around 1500 species consisting of primarily woody plants and some herbaceous plants. The morphology of Araliaceae varies widely, but it is predominantly distinguish ...
: * '' Brassaia'' (Umbrella Tree) Arecaceae: * ''
Caryota ''Caryota'' is a genus of palm trees. They are often known as fishtail palms because of the shape of their leaves. There are about 13 species native to Asia ( China, India, Indonesia, etc.), northern Australia, and the South Pacific. One of ...
'' (Fishtail Palm) * ''
Ptychosperma ''Ptychosperma'' is a genus of flowering plant in the family Arecaceae. Most are native to Australia and/or New Guinea, with a few in the Solomon Islands and in Maluku Province of eastern Indonesia. Some have been cultivated abroad as house or ...
'' * '' Arenga'' * '' Hyophorbe verschaffeltii'' (Spindle palm) Asparagaceae: * ''
Agave ''Agave'' (; ; ) is a genus of monocots native to the hot and arid regions of the Americas and the Caribbean, although some ''Agave'' species are also native to tropical areas of North America, such as Mexico. The genus is primarily known for ...
'' (Agave) * '' Chlorophytum orchidastrum'' (Green Orange) Bromeliaceae: *
Ananas comosus ''Ananas'' is a plant genus in the family Bromeliaceae. It is native to South America. The genus contains ''Ananas comosus'', the pineapple. Species The genus ''Ananas'' includes only two species: Gallery File:Pineapple.plantation.jpg, Pinea ...
(Pineapple) Commelinaceae: * ''
Commelina ''Commelina'' is a genus of approximately 170 species commonly called dayflowers due to the short lives of their flowers. They are less often known as widow's tears. It is by far the largest genus of its family, Commelinaceae. The Swedish taxonom ...
'' (Dayflower) * ''
Tradescantia ''Tradescantia'' () is a genus of 85 species of herbaceous perennial wildflowers in the family Commelinaceae, native to the Americas from southern Canada to northern Argentina, including the West Indies. Members of the genus are known by many com ...
'' (Wandering Jew) * ''
Triceratella ''Triceratella'' is a genus of annual monocotyledonous flowering plants in the dayflower family. The genus consists of a single species, ''Triceratella drummondii''. It is known to occur in Zimbabwe and Mozambique, but has only been collected tw ...
''
Onagraceae The Onagraceae are a family of flowering plants known as the willowherb family or evening primrose family. They include about 650 species of herbs, shrubs, and treesFuchsia ''Fuchsia'' () is a genus of flowering plants that consists mostly of shrubs or small trees. The first to be scientifically described, ''Fuchsia triphylla'', was discovered on the Caribbean island of Hispaniola (Haiti and the Dominican Republic) ...
'' (Fuchsia)
Polygonaceae The Polygonaceae are a family of flowering plants known informally as the knotweed family or smartweed—buckwheat family in the United States. The name is based on the genus '' Polygonum'', and was first used by Antoine Laurent de Jussieu in 1 ...
: * ''
Rheum rhabarbarum ''Rheum rhabarbarum'' is a species of flowering plant in the family Polygonaceae, native to a region stretching from southern Siberia to north and central China. It has been harvested from the wild for centuries for its root, which was harvested ...
'' (Rhubarb)
Rubiaceae The Rubiaceae are a family of flowering plants, commonly known as the coffee, madder, or bedstraw family. It consists of terrestrial trees, shrubs, lianas, or herbs that are recognizable by simple, opposite leaves with interpetiolar stipules a ...
: * '' Hillia''
Vitaceae The Vitaceae are a family of flowering plants, with 14 genera and around 910 known species, including common plants such as grapevines (''Vitis'' spp.) and Virginia creeper (''Parthenocissus quinquefolia''). The family name is derived from the ge ...
: * ''
Parthenocissus quinquefolia ''Parthenocissus quinquefolia'', known as Virginia creeper, Victoria creeper, five-leaved ivy, or five-finger, is a species of flowering vine in the grape family, Vitaceae. It is native to eastern and central North America, from southeastern C ...
'' (Virginia creeper)


See also

* Calcium oxalate * Druse *
Plant defense against herbivory Plant defense against herbivory or host-plant resistance (HPR) describes a range of adaptations evolved by plants which improve their survival and reproduction by reducing the impact of herbivores. Plants can sense being touched, and they can ...


References

{{reflist Plant physiology Oxalate minerals Organic minerals Plant toxins