Passiflora

Passion flowers
Passiflora × kewensis
P. platyloba fruit , often confused with P. quadrangularis
Scientific classification
Kingdom:	Plantae
Division:	Magnoliophyta
Class:	Magnoliopsida
Subclass:	Rosidae
(unranked):	Eurosids I
Order:	Malpighiales
Family:	Passifloraceae
Genus:	Passiflora L.
Species
About 500, see list
Synonyms
Anthactinia Bory ex M.Roem. Asephananthes Bory Astrophea Lam. ex M.Roem. Baldwinia Raf. Ceratosepalum Oerst. Cieca Medik. Decaloba M.Roem. Disemma Labill. Granadilla Mill. Hollrungia K.Schum. Monactineirma Bory Murucuja Medik. Pentaria M.Roem. Poggendorffia H.Karst. Tacsonia Juss. Tetrapathaea (DC.) Rchb. Tetrastylis Barb.Rodr.

The passion flowers or passion vines (Passiflora) are a genus of about 500 species of flowering plants, the namesakes of the family Passifloraceae. They are mostly vines, with some being shrubs, and a few species being herbaceous. For information about the fruit of the passiflora plant, see passionfruit. The monotypic genus Hollrungia seems to be inseparable from Passiflora, but further study is needed.

Distribution

The family Passifloraceae is found worldwide except in Antarctica, and Passiflora is absent from Africa though many other members of the family Passifloraceae exist there (e.g. the more plesiomorphic Adenia).

Nine species of Passiflora are native to the USA, found from Ohio to the north, west to California and south to the Florida Keys. Most other species are found in South America, China, and Southern Asia, New Guinea, four or more species in Australia and a single endemic species in New Zealand. New species continue to be identified: for example, P. pardifolia and P. xishuangbannaensis have only been known to the scientific community since 2006 and 2005, respectively.

Species of Passiflora have been naturalised beyond their native ranges. For example, Blue Passion Flower (P. caerulea) now grows wild in Spain. The purple passionfruit (P. edulis) and its yellow relative flavicarpa have been introduced in many tropical regions as commercial crops.

Ecology

Stinking Passion Flower or Wild water lemon (P. foetida) bracts with the insect-catching hairs.

The decorative passion flowers have a unique flower structure, which in most cases requires a large bee to effectively pollinate. In the American tropics, wooden beams are mounted very near passionfruit plantings to encourage carpenter bees to nest. The size and structure of flowers of other Passiflora species is optimized for pollination by hummingbirds (especially hermits like Phaethornis) , bumble bees, wasps or bats, while yet others are self-pollinating. The Sword-billed Hummingbird (Ensifera ensifera) with its immensely elongated bill has co-evolved with certain passion flowers, such as P. mixta.

The Sword-billed Hummingbird (Ensifera ensifera) is adapted to feed on Passiflora mixta and similar flowers.

Yellow Passion Flower (P. lutea) pollen is apparently the only pollen eaten by the unusual bee Anthemurgus passiflorae. However, these bees simply collect the pollen, but do not pollinate the flowers.

Passiflora species are important sources of nectar for many insects. The leaves are used as food plants by the larva of the swift moth Cibyra serta and many longwing butterflies (Heliconiinae). Well-known species among the latter are the American Sara Longwing (Heliconius sara) and the Asian Leopard Lacewing (Cethosia cyane). The caterpillars of the Postman Butterfly (Heliconius melpomene) prefer P. menispermifolia and P. oerstedii when available; those of the Zebra Longwing (Heliconius charithonia) feed on Yellow Passion Flower, Two-flowered Passion Flower (P. biflora), and Corky-stemmed Passion Flower (P. suberosa). Those of the Banded Orange (Dryadula phaetusa) are found on P. tetrastylis, those of the Julia Butterfly (Dryas iulia) on Yellow Passion Flower and P. affinis, and those of the Gulf Fritillary (Agraulis vanillae) on Yellow Passion Flower, Stinking Passion Flower (P. foetida) and Maypop (P. incarnata).

To prevent the butterflies from laying too many eggs on any single plant, some passion flowers bear small colored nubs which resemble the butterflies' eggs and seem to fool them into believing that more eggs have already been deposited on a plant than actually is the case. Also, many Passiflora species produce sweet nutrient-rich liquid from glands on their leaf stems. These fluids attract ants which will kill and eat many pests that they happen to find feeding on the passion flowers.

The bracts of the Stinking Passion Flower are covered by hairs which exude a sticky fluid. Many small insects get stuck to this and get digested to nutrient-rich goo by proteases and acid phosphatases. Since the insects usually killed are rarely major pests, this passion flower seems to be a protocarnivorous plant.

Banana Passion Flower or "banana poka" (P. tarminiana), originally from Central Brazil, is an invasive weed, especially on the islands of Hawaii. It is commonly spread by feral pigs eating the fruits. It overgrows and smothers stands of endemic vegetation, mainly on roadsides. Blue Passion Flower (P. caerulea) is holding its own in Spain these days, and it probably needs to be watched so that unwanted spreading can be curtailed.

On the other hand, some species are endangered due to unsustainable logging and other forms of habitat destruction. For example, the Chilean Passion Flower (P. pinnatistipula) is a rare vine growing in the Andes from Venezuela to Chile between 2,500 and 3,800 meters altitude, and in Coastal Central Chile, where it occurs in woody Chilean Mediterranean forests. P. pinnatistipula has a round fruit, unusual in Tacsonia group species like Banana Passion Flower and P. mixta, with their elongated tubes and brightly red to rose-colored petals.

Notable and sometimes economically significant pathogens of Passiflora are several sac fungi of the genus Septoria (including S. passiflorae), the undescribed proteobacterium called "Pseudomonas tomato" (pv. passiflorae), the Potyvirus Passionfruit woodiness virus, and the Carlavirus Passiflora latent virus.

Use by humans

Hundreds of hybrids have been named and hybridizing is currently being done extensively for flowers, foliage and fruit. A number of species of Passiflora are cultivated outside their natural range because of their beautiful flowers.

During Victorian times the flower (which in all but a few species lasts only one day) was very popular and many hybrids were created using Winged-stem Passion Flower (P. alata) and Blue Passion Flower (P. caerulea) and other tropical species.

Many cool-growing Passiflora from the Andes Mountains can be grown successfully for their beautiful flowers and fruit in cooler Mediterranean climates, such as the Monterey Bay and San Francisco in California and along the Western Coast of the U.S. into Canada. One Blue Passion Flower or hybrid even grew to large size at Malmö Central Station in Sweden.

Passion flowers have been a subject of studies investigating extranuclear inheritance; paternal inheritance of chloroplast DNA has been documented in this genus. The plastome of the Two-flowered Passion Flower (P. biflora) has been sequenced.

The French name for this plant has lent itself to La Famille Passiflore, a highly successful children's book series by Geneviève Huriet, and an animated series based upon it. These have been translated into English as Beechwood Bunny Tales and The Bellflower Bunnies, respectively.

Fruit

Most species have round or elongated edible fruit from two to eight inches long and an inch to two inches across, depending upon the species or cultivar.

• The Passion fruit or maracujá (P. edulis) is cultivated extensively in the Caribbean and south Florida and South Africa for its fruit, which is used as a source of juice. A small purple fruit which wrinkles easily and a larger shiny yellow to orange fruit are traded under this name. The latter is usually considered just a variety flavicarpa, but seems to be more distinct in fact.

• Sweet Granadilla (P. ligularis) is another widely-grown species. In large parts of Africa and Australia it is the plant called "passionfruit": confusingly, in South African English the latter species is more often called "granadilla" (without an adjective). Its fruit is somewhat intermediate between the two sold as P. edulis.

• Maypop (P. incarnata), a common species in the southeastern US. This is a subtropical representative of this mostly tropical family. However, unlike the more tropical cousins, this particular species is hardy enough to withstand the cold down to -4°F (-20°C) before its roots die (it is native as far north as Pennsylvania and has been cultivated as far north as Boston and Chicago.) The fruit is sweet, yellowish, and roughly the size of a chicken's egg; it enjoys some popularity as a native plant with edible fruit and few pests.

• Giant Granadilla (Giant Tumbo or badea, P. quadrangularis), Water Lemon (P. laurifolia) and Sweet Calabash (P. maliformis) are Passiflora species locally famed for their fruit, but not widely known elsewhere yet.

• Wild Maracuja are the fruit of P. foetida, which are popular in Southeast Asia.

• Banana passionfruits are the very elongated fruits of P. tripartita var. mollissima and P. tarminiana. These are locally eaten, but its invasive properties make it hardly worthwhile to grow at least the latter species on purpose.

Medical and entheogenic uses

Chrysin, a commercially important flavone found in the blue passion flower, P. caerulea.

P. incarnata (maypop) leaves and roots have a long history of use among Native Americans in North America and were adapted by the European colonists. The fresh or dried leaves of maypop are used to make a tea that is used to treat insomnia, hysteria, and epilepsy, and is also valued for its analgesic properties. P. edulis (passion fruit) and a few other species are used in Central and South America for similar purposes. Once dried, the leaves can also be smoked.

Harman, a harmala alkaloid found in many species of Passiflora.

Many species have been found to contain beta-carboline harmala alkaloids. which are MAO inhibitors with anti-depressant properties. The flower and fruit have only traces of these chemicals, but the leaves and the roots are often more potent and have been used to enhance the effects of mind-altering drugs. The most common of these alkaloids is harman (1-methyl-9H-b-carboline), but harmaline (4,9-Dihydro-7-methoxy-1-methyl-3H-pyrido[3,4-b]indole), harmalol (1-methyl-2,3,4,9-tetrahydropyrido[3,4-b]indol-7-one), harmine (7-Methoxy-1-methyl-9H-pyrido[3,4-b]indole) and harmol were found. The species known to bear such alkaloids include: P. actinea, P. alata (winged-stem passion flower), P. alba, P. bryonioides (cupped passion flower), P. caerulea (blue passion flower), P. capsularis, P. decaisneana, P. edulis (passion fruit), P. eichleriana, P. foetida (stinking passion flower), P. incarnata (maypop), P. quadrangularis (giant granadilla), P. ruberosa, P. subpeltata and P. warmingii

Other compounds found in passion flowers are coumarins (e.g. scopoletin and umbelliferone), maltol, phytosterols (e.g. lutenin) and cyanogenic glycosides (e.g. gynocardin) which render some species, i.e. P. adenopoda, somewhat poisonous. Many flavonoids and their glycosides have been found in Passiflora, including apigenin, benzoflavone, homoorientin, 7-isoorientin, isoshaftoside, isovitexin (or saponaretin), kaempferol, lucenin, luteolin, n-orientin, passiflorine (named after the genus), quercetin, rutin, saponarin, shaftoside, vicenin and vitexin. Maypop, Blue Passion Flower (P. caerulea), and perhaps others contain chrysin, a flavone with confirmed anxiolytic and anti-inflammatory, supposed aromatase inhibitor properties. Also documented to occur at least in some Passiflora in quantity are the hydrocarbon nonacosane and the anthocyanidin pelargonidin-3-diglycoside.

As regards organic acids, the genus is rich in formic, butyric, linoleic, linolenic, malic, myristic, oleic and palmitic acids as well as phenolic compounds, and the amino acid α-alanine. Esters like ethyl butyrate, ethyl caproate, n-hexyl butyrate and n-hexyl caproate give the fruits their flavor and appetizing smell. Sugars, contained mainly in the fruit, are most significantly d-fructose, d-glucose and raffinose. Among enzymes, Passiflora was found to be rich in catalase, pectin methylesterase and phenolase.

The medical utility of very few species of Passiflora has been scientifically studied. In initial trials for treatment of generalized anxiety disorder, maypop extract performed as well as oxazepam but with fewer short-term side effects. It was recommended to follow up with long-term studies.

Side effects

One study on mice showed that Passiflora alata has a genotoxic effect on cells. P. incarnata are generally considered as safe.

Etymology and names

Popularly, passion flowers and especially passion fruit are frequently used with sexual or romantic innuendo, giving rise to such uses as a one-time soft drink named Purple Passion. The "Passion" in "passion flower" does not refer to sex and love, however, but to the passion of Jesus in Christian theology. In the 15th and 16th centuries, Spanish Christian missionaries adopted the unique physical structures of this plant, particularly the numbers of its various flower parts, as symbols of the last days of Jesus and especially his crucifixion:

• The pointed tips of the leaves were taken to represent the Holy Lance.
• The tendrils represent the whips used in the flagellation of Christ.
• The ten petals and sepals represent the ten faithful apostles (less St. Peter the denier and Judas Iscariot the betrayer).
• The flower's radial filaments, which can number more than a hundred and vary from flower to flower, represent the crown of thorns.
• The chalice-shaped ovary with its receptacle represents a hammer or the Holy Grail
• The 3 stigmas represent the 3 nails and the 5 anthers below them the 5 wounds (four by the nails and one by the lance).
• The blue and white colors of many species' flowers represent Heaven and Purity.

The flower has been given names related to this symbolism throughout Europe since that time. In Spain, it is known as espina de Cristo ("Christ's thorn"). German names include Christus-Krone ("Christ's crown"), Christus-Strauss ("Christ's bouquet"), Dorn-Krone ("crown of thorns"), Jesus-Leiden ("Jesus' passion"), Marter ("passion") or Muttergottes-Stern ("Mother of God's star").

Outside the Christian heartland, the regularly-shaped flowers have reminded people of the face of a clock; in Israel they are known as "clock-flower" (שעונית), and in Japan they are called tokeisō (時計草, "clock plant"). In Hawaiian, they are called lilikoʻi; lī is a string used for tying fabric together, such as a shoelace, and liko means "to spring forth leaves".

In India, blue passionflowers are called Krishnakamala in Karnataka and Maharashtra, while in UP and generally north it is colloquially called "Paanch Paandav". The flower's structure lends itself to the interpretation along the lines of five Pandavas, the Divine Krishna at centre, and the opposing hundred at the edges. The colour blue is moreover associated with Krishna as colour of his aura.

In northern Peru and Bolivia, the banana passionfruits are known as tumbos. This is one possible source of the name of the Tumbes region of Peru.

Taxonomy

Passiflora is the most speciose genus of both the family Passifloraceae and the tribe Passifloreae. With over 530 species an extensive hierarchy of infrageneric ranks is required to represent the relationships of the species, and the infrageneric classification of Passiflora not only uses the widely used ranks of subgenus, section and series, but also the rank of supersection.

The New World species of Passiflora were divided among 22 subgenera by Killip (1938). More recent work reduces these to 4 - Astrophea (Americas, 57 species), Deidamioides (Americas, 17 species), Passiflora (Americas, >200 species) and Decaloba (Americas, Asia and Australasia, >200 species). Other studies have shown that the segregate Old World genera Hollrungia and Tetrapathaea are nested within Passiflora, and form a fifth subgenus (Tetrapathaea).

The Old World species form two clades - supersection Disemma (part of subgenus Decaloba) and subgenus Tetrapathaea. The former is composed of 21 species divided into sections Disemma (3 Australian species), Holrungiella (1 New Guinean species) and Octandranthus (17 south and east Asian species).

The remaining (New World) species of subgenus Decaloba are divided into 7 supersections. Supersection Pterosperma includes 4 species from Central America and southern Mexico. Supersection Hahniopathanthus includes 5 species from Central America, Mexico and northernmost South America. Supersection Cicea includes 19 species, with apetalous flowers. Supersection Bryonioides includes 21 species, with a distribution centered on Mexico. Supersection Auriculata includes 8 species from South America, one of which is also found in Central America. Supersection Multiflora includes 19 species. Supersection Decaloba includes 123 species.

Gallery

• 
  Passiflora xishuangbannaensis, another recently-described species, is found in Xishuangbanna Dai Autonomous Prefecture
• 
  The caterpillars of Heliconius charithonia, like many of their relatives, are fond of Passiflora lutea leaves whose poison protects the caterpillars from predators.
• 
  Flower of Passiflora × belotii, a horticultural hybrid.
• 
  Blue Passion Flower (Passiflora caerulea)
• 
  Pink-white passion flower Passiflora caerulea 'Hartwiesiana'
• 
  Passiflora pardifolia was only described in 2006
• 
  Passiflora xishuangbannaensis, another recently-described species, is found in Xishuangbanna Dai Autonomous Prefecture
• 
  Stinking Passion Flower or Wild water lemon (P. foetida) bracts with the insect-catching hairs.
• 
  An Passiflora incarnata. One of the most common of Passion flowers.
• 
  Passiflora entwine around this 1880 Baxter process illustration by Joseph Martin Kronheim.

See also

• List of culinary fruits

References

• Akhondzadeh, Shahin; Naghavi, H.R.; Vazirian, M.; Shayeganpour, A.; Rashidi, H. & Khani, M. (2001): Passionflower in the treatment of generalized anxiety: a pilot double-blind randomized controlled trial with oxazepam. Journal of Clinical Pharmacy and Therapeutics 26(5): 363-367. doi:10.1046/j.1365-2710.2001.00367.x PDF fulltext
• Dana, E.D.; Sanz-Elorza, M. & Sobrino, E. [2001]: Plant Invaders in Spain Check-List. PDF fulltext
• Dhawan, Kamaldeep; Kumar, Suresh & Sharma, Anupam (2002): Beneficial Effects of Chrysin and Benzoflavone on Virility in 2-Year-Old Male Rats. Journal of Medicinal Food 5(1): 43-48. doi:10.1089/109662002753723214 (HTML abstract)
• Drugs.com [2008]: Passion Flower. Retrieved 2008-NOV-01.
• Duke, James A. [2008]: Dr. Duke's Phytochemical and Ethnobotanical Databases – Passiflora spp. Retrieved 2008-NOV-01.
• Hansen, A. Katie; Escobar, Linda K.; Gilbert, Lawrence E. & Jansen, Robert K. (2006): Paternal, maternal, and biparental inheritance of the chloroplast genome in Passiflora (Passifloraceae): implications for phylogenic studies. Botany 94(1): 42-46. PDF fulltext
• Marzell, Heinrich (1927): Deutsches Wörterbuch der Pflanzennamen ["German Plant Name Dictionary"]. Leipzig.
• Pukui, Mary Kawena; Elbert, Samuel Hoyt; Mookini, Esther T. & Nishizawa, Yu Mapuana (1992): New Pocket Hawaiian Dictionary with a Concise Grammars and Given Names in Hawaiian. University of Hawaii Press, Honolulu. ISBN 0-8248-1392-8
• Petersen, Elly (1966): Passionsblume ["Passion flowers"]. In: Praktisches Gartenlexikon der Büchergilde (2nd ed.): 270-271 [in German]. Büchergilde Gutenberg. Frankfurt am Main, Vienna, Zürich.
• Radhamani, T.R.; Sudarshana, L. & Krishnan, R. (1995): Defence and carnivory: dual roles of bracts in Passiflora foetida. Journal of Biosciences 20(5): 657-664. doi:10.1007/BF02703305 PDF fulltext
• University of Maryland Medical Center (UMMC) (2008): Passionflower. Retrieved 2008-NOV-01.

External links

• The Passiflora Society International
• Passiflora: passionflowers of the world
• Killip, The American Species of Passifloraceae, Fieldiana, Bot. 19 (1938)
• Passiflora online
• Passiflora edulis
• Passiflora Picture Gallery
• Chilean Passiflora pictures
• A list of Heliconius Butterflies and the Passiflora species their larvae consume
• "Passiflora" at the Encyclopedia of Life