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Raphides ( ; singular raphide or raphis) are needle-shaped crystals of monohydrate ( crystals) or calcium carbonate as aragonite ( 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; in raphide containing cells makes light microscopy difficult, though. Little is known about the mechanisms of sequestration or indeed the reason for the 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 structures. Raphides typically occur in 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 indication 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 . 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 were apparently calcium oxalate.According to this abstract in the FDA Poisonous Plant Database Agave americana contains calcium oxalate crystals, acrid oils, , and other compounds.

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 consuming animals, as they are likely to tear and harm the soft tissues of the throat or esophagus of an animal chewing on the plant's leaves. The venomous process is in two stages: mechanical pricking and injection of harmful . Typically ingestion of plants containing raphides, like those common in certain houseplants, can cause immediate numbing followed shortly by painful , vesicle formation and 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 the 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.

There is some doubt about the role of raphides in defense against herbivory. Insects are only rarely affected by them, and no more than by sand. are not harmed by them, because certain bacteria in their help to degrade .


Plants containing raphides
Common names vary. The following list is incomplete. Raphides are found in many species in the families and , but are also found in a few species in a number of other families.

:

  • Strobilanthes barbata (stem, leaf)
  • Strobilanthes callosa (stem, leaf)
  • Strobilanthes ciliata (stem, leaf)
  • Strobilanthes heyneana (leaf)
  • Strobilanthes integrifolia (stem, leaf)
  • Strobilanthes lupulina (stem, leaf)
  • Strobilanthes reticulata (stem, leaf)
  • Strobilanthes sessilis (stem, leaf)
  • Thunbergia erecta (stem, leaf)
  • Thunbergia laurifolia (stem, leaf)

:

  • (root, flower, fruit)
  • Actinidia callosa (fruit)
  • Actinidia chinensis (root, flower, fruit)
  • Actinidia chrysantha (root, fruit)
  • Actinidia deliiosa (fruit)
  • Actinidia eriantha (fruit)
  • Actinidia fulvicoma (fruit)
  • Actinidia glaucophylla (fruit)
  • Actinidia hemsleyana (root)
  • Actinidia indochinensis (fruit)
  • Actinidia lanceolata (fruit)
  • Actinidia macrosperma (fruit)
  • Actinidia polygama (fruit)
  • (root, fruit)
  • Clematoclethra scandens (flower)
  • Saurauia pittieri (flower)
  • Saurauia subspinosa (flower)

:

  • Alstroemeria angustifolia (leaf)
  • Alstroemeria aurea (leaf)
  • Alstroemeria exserens (leaf)
  • Alstroemeria hookeri (leaf)
  • Alstroemeria ligtu (leaf)
  • Alstroemeria magnifica (leaf)
  • Alstroemeria pallida (leaf)
  • Alstroemeria pelegrina (leaf)
  • Alstroemeria presliana (leaf)
  • Alstroemeria pulchra (leaf)
  • Alstroemeria versicolor (leaf)
  • Bomarea multiflora (leaf)
  • (leaf)

:

:

  • (leaf)
  • Leucojum aestivum (leaf)
  • Pancratium sickenbergeri (leaf)

:

  • Alocasia macrorrhizos (root, stem, leaf, flower)
  • Amorphophallus bulbifer (leaf)
  • Amorphophallus blumei (root, leaf)
  • Amorphophallus dunnii (leaf)
  • Amorphophallus gallaensis (leaf)
  • Amorphophallus konjac (leaf)
  • Amorphophallus krausei (leaf)
  • Amorphophallus muelleri (root, leaf)
  • Amorphophallus paeoniifolius (root, leaf)
  • Amorphophallus salmoneus (leaf)
  • Amorphophallus sutepensis (leaf)
  • Amorphophallus taurostigma (leaf)
  • Amorphophallus titanum (leaf)
  • Amorphophallus variabilis (leaf)
  • Anaphyllopsis americana (flower)
  • Anthurium andraeanum (leaf, flower)
  • (flower)
  • Anthurium crystallinum (flower)
  • Anthurium cubense (flower)
  • Anthurium halmoorei (flower)
  • Anthurium harrisii (flower)
  • Anthurium jenmannii (flower)
  • Anthurium magnificum (flower)
  • Anthurium maricense (leaf)
  • Anthurium pallidiflorum (flower)
  • Anthurium pedatoradiatum (flower)
  • Anthurium schlechtendalii (flower)
  • Anthurium signatum (flower)
  • Anthurium spectabile (flower)
  • Anubias heterophylla (flower)
  • (flower)
  • (flower)
  • (flower)
  • Cercestis dinklagei (flower)
  • Colocasia affinis (stem)
  • Colocasia gigantea (stem)
  • Colocasia esculenta (root, stem, leaf)
  • Cyrtosperma chamissonis (stem)
  • Cyrtosperma johnstonii (root, leaf, flower)
  • Cyrtosperma merkusii (stem, leaf)
  • Dieffenbachia seguine (stem, leaf, flower)
  • (leaf)
  • Monstera deliciosa (fruit)
  • Philodendron bipinnatifidum (flower)
  • Philodendron megalophyllum (flower)
  • Philodendron moonenii (flower)
  • Philodendron oblongum (flower)
  • Philodendron pedatum (flower)
  • Philodendron propinquum (flower)
  • (stem)
  • Pinellia pedatisecta (stem)
  • Pistia stratiotes (leaf)
  • Stenospermation longiopetiolatum (flower)
  • Syngonium schottianum (flower)
  • Thaumatophyllum adamantinum (flower)
  • Thaumatophyllum brasiliense (flower)
  • Thaumatophyllum corcovadense (flower)
  • Thaumatophyllum dardanianum (flower)
  • Thaumatophyllum lundii (flower)
  • Thaumatophyllum palaudicola (flower)
  • Thaumatophyllum petraeum (flower)
  • Thaumatophyllum saxicola (flower)
  • Thaumatophyllum speciosum (flower)
  • Thaumatophyllum tweedianum (flower)
  • Thaumatophyllum uliginosum (flower)
  • Thaumatophyllum undulatum (flower)
  • Thaumatophyllum williamsii (flower)
  • Typhonium flagelliforme (stem)
  • Xanthosoma sagittifolium (root, stem, leaf)
  • Zantedeschia aethiopia (flower)

:

:

  • Acrocomia aculeata (root, leaf, flower)
  • (root, leaf, flower)
  • Acrocomia emensis (root, leaf, flower)
  • Acrocomia glaucensis (root, leaf, flower)
  • Acrocomia hassleri (root, leaf, flower)
  • Acrocomia intumescens (root, leaf, flower)
  • (root, leaf, flower)
  • Allagoptera campestris (leaf)
  • Ammandra decasperma (pseudopedicel)
  • (pseudopedicel)
  • Archontophoenix alexandrae (embryo)
  • (embryo)
  • (embryo)
  • Areca novohibernica (embryo)
  • (embryo)
  • (fruit)
  • Brassiophoenix dryomphloeides (embryo)
  • Butia buenopolensis (leaf, flower)
  • (leaf)
  • (leaf)
  • Butia catarinensis (leaf)
  • (leaf)
  • Butia leptospatha (leaf)
  • (leaf)
  • Butia microspadix (leaf)
  • (leaf)
  • Calyptrocalyx albertisianus (fruit)
  • Calyptrocalyx hollrungii (embryo)
  • Calyptrocalyx lepidotus (fruit)
  • Calyptrocalyx spicatus (embryo)
  • (embryo)
  • (root, stem, leaf, flower, fruit)
  • Caryota rumphiana (stem, leaf)
  • Chamaedorea tepejilote (embryo)
  • (root, stem, leaf)
  • Copernicia tectorum (flower, fruit)
  • Cyrtostachys renda (embryo)
  • Drymophloeus pachycladus (embryo)
  • Drymophloeus subdistichus (embryo)
  • (embryo)
  • Dypsis leptocheilos (embryo)
  • (embryo)
  • (embryo)
  • (fruit)
  • Hydriastele rostrata (embryo)
  • Hyophorbe verschaffeltii
  • '' sp. (embryo)
  • Laccospadix australasicus (fruit)
  • Linospadix microcaryus (fruit)
  • Linospadix monostachyos (fruit)
  • Livistonia saribus (embryo)
  • Normanbya normanbyi (embryo)
  • Oenocarpus bacaba (embryo)
  • Oenocarpus bataua (embryo)
  • Oncosperma horridum (embryo)
  • Oncosperma tigillarium (embryo)
  • Pelagodoxa henryana (fruit)
  • (embryo)
  • Pinanga negrosensis (embryo)
  • (embryo)
  • Pinanga scortechinii (embryo)
  • Prestoea acuminata (embryo)
  • Ptychosperma caryotoides (embryo)
  • (embryo)
  • (embryo)

:

  • Asplenium cardiophyllum (leaf epidermis)

:

:

:

:

  • (stem)
  • Impatiens balfourii (stem, leaf, flower)
  • Impatiens niamniamensis (stem)
  • Impatiens parviflora (flower)
  • Impatiens thomsonii (stem, leaf, flower)

:

:

  • very widespread

:

  • widespread

:

:

  • Cochliostema odoratissimum (stem, leaf, flower)
  • Commelina coelestis (pollen)
  • Commelina tuberosa (pollen)
  • Dichorisandra thyrsiflora (anther)
  • (root, leaf, flower)
  • Tinantia pringlei (root, leaf, flower)
  • Tradescantia ohiensis (stem)
  • Tradescantia pallida (anther)
  • Tradescantia spathacea (anther)

:

:

  • Dioscorea bulbifera (root, bulbil)
  • Dioscorea esculenta (root)
  • Dioscorea hispida (root, petiole)
  • Dioscorea japonica (root)
  • Dioscorea polystachya (root, stem)

:

  • Diospyros montana (bark, fruit)

:

:

  • (stem, leaf)
  • Medicago truncatula (leaf)
  • Pachyrhizus tuberosus (root)

:

  • Anigozanthos flavidus (leaf, flower)
  • Conostylis candicans (flower)

:

:

  • Hydrangea macrophylla (leaf)

:

  • Satureja khuzestanica (leaf)

:

:

  • Orchidantha maxillarioides (stem, leaf)

:

  • very widespread

:

:

  • Bougainvillea glabra (leaf)
  • Bougainvillea spectabilis (leaf)

:

  • very widespread

:

  • Epilobium tibetanum (leaf)
  • Ludwigia peploides (leaf)

:

  • very widespread

:

'':

  • Passiflora cincinnata (fruit)

:

  • Heimholtzia glaberrima

:

:

  • Cenchrus echinatus

:

  • Rheum rhabarbarum

:

  • Monochoria africana (anther)
  • Pontederia cordata (root)
  • Pontederia crassipes (stem)

:

  • Roridula gorgonias (flower)

:

:

  • very widespread

:

  • Pelliciera rhizophorae (stem, flower)
  • Pentamerista neotropica (flower)
  • Tetramerista glabra (stem)

:

:

  • (leaf)
  • (leaf)
  • Cissus quadrangularis (leaf)
  • (leaf)
  • Cissus rotundifolia (leaf)
  • Parthenocissus quinquefolia (leaf)
  • Parthenocissus tricuspidata (leaf)
  • Parthenocissus vitacea (leaf)
  • Rhoicissus tomentosa (leaf)
  • Tetrastigma voinierianum (leaf)
  • Vitis berlandieri (root)
  • (anther)
  • (leaf)


See also


Bibliography
  • See Supporting Information.

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