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Passiflora

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Passiflora, known also as the passion flowers or passion vines, is a genus of about 550 species of flowering plants, the type genus of the family Passifloraceae.

Passiflora species are widely cultivated for their striking flowers, flavorful fruits, traditional medicinal uses, and roles in dietary supplements and ayahuasca analogs, with several ornamental hybrids earning Royal Horticultural Society awards.

DescriptionEdit

They are mostly tendril-bearing vines, with some being shrubs or trees. They can be woody or herbaceous.<ref name = "Ulmer & McDougal, 2004" />

Passion flowers produce regular and usually showy flowers with a distinctive corona. There can be as many as eight concentric coronal series, as in the case of P. xiikzodz.<ref name = "Ulmer & McDougal, 2004">Template:Cite book</ref> and Passiflora alata. The hallmark of the genus is the androgynophore, a central column to which the stamens and pistil are attached, which can be very long in some species such as Passiflora coactilis. The flower is pentamerous (except for a few Southeast Asian species) and ripens into an indehiscent fruit with numerous seeds.

The fruit ranges from Template:Convert long and Template:Convert across, depending upon the species or cultivar.

ChemistryEdit

Template:Multiple image Many species of Passiflora have been found to contain beta-carboline harmala alkaloids,<ref name="drugs" /><ref name="Duke">Duke (2008)</ref><ref name="Meuninck">Template:Cite book</ref> some of which are MAO inhibitors. The flower and fruit have only traces of these chemicals, but the leaves and the roots often contain more.<ref name="Meuninck" /> The most common of these alkaloids is harman, but harmaline, harmalol, harmine, and harmol are also present.<ref name="drugs" /><ref name="Duke" /> The species known to bear such alkaloids include: P. actinia, 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. suberosa, P. subpeltata and P. warmingii.<ref name="Duke" />

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, Template:Nowrap, isoshaftoside, isovitexin (or saponaretin), kaempferol, lucenin, luteolin, Template:Nowrap, passiflorine (named after the genus), quercetin, rutin, saponarin, shaftoside, vicenin and vitexin. Maypop, blue passion flower (P. caerulea), and perhaps others contain the flavone chrysin. Also documented to occur at least in some Passiflora in quantity are the hydrocarbon nonacosane and the anthocyanidin pelargonidin-3-diglycoside.<ref name="drugs" /><ref name="Duke" /><ref name="dd2008">Dhawan, et al. (2002)</ref>

The genus is rich in organic acids including formic, butyric, linoleic, linolenic, malic, myristic, oleic and palmitic acids as well as phenolic compounds, and the amino acid Template:Nowrap. 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 Template:Nowrap, Template:Nowrap and raffinose. Among enzymes, Passiflora was found to be rich in catalase, pectin methylesterase and phenolase.<ref name="drugs" /><ref name="Duke" />

TaxonomyEdit

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Passiflora is the most species rich genus of both the family Passifloraceae and the tribe Passifloreae. With over 550 species, an extensive hierarchy of infrageneric ranks is required to represent the relationships of the species. 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 first divided among 22 subgenera by Killip (1938) in the first monograph of the genus.<ref name="Killip 1938" /> More recent work has reduced these to 4, which are commonly accepted today (in order from most basally to most recently branching):<ref>Template:Cite journal</ref>

  • Astrophea (Americas, ~60 species), trees and shrubs with simple, unlobed leaves
  • Passiflora (Americas, ~250 species), woody vines with large flowers and elaborate corolla
  • Deidamioides (Americas, 13 species), woody or herbaceous vines
  • Decaloba (Americas, Asia and Australasia, ~230 species), herbaceous vines with palmately veined leaves

Some studies have shown that the segregate Old World genera Hollrungia and Tetrapathaea are nested within Passiflora, and form a fifth subgenus (Tetrapathaea).<ref>Template:Cite journal</ref> Other studies support the current four subgenus classification.<ref>Template:Cite journal</ref>

Relationships below the subgenus level are not known with certainty and are an active area of research. 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 (three Australian species), Holrungiella (one New Guinean species) and Octandranthus (seventeen south and east Asian species).<ref>Shawn Elizabeth Krosnick, PhD thesis, Phylogenetic relationships and patterns of morphological evolution in the Old Word species of Passiflora (subgenus Decaloba: supersection Disemma and subgenus Tetrapathaea) Template:Webarchive</ref>

The remaining (New World) species of subgenus Decaloba are divided into seven supersections. Supersection Pterosperma includes four species from Central America and southern Mexico. Supersection Hahniopathanthus includes five species from Central America, Mexico and northernmost South America. Supersection Cicea includes nineteen species, with apetalous flowers. Supersection Bryonioides includes twenty-one species, with a distribution centered on Mexico. Supersection Auriculata includes eight species from South America, one of which is also found in Central America. Supersection Multiflora includes nineteen species. Supersection Decaloba includes 123 species.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

DistributionEdit

Passiflora has a largely neotropic distribution, unlike other genera in the family Passifloraceae, which includes more Old World species (such as the genus Adenia). The vast majority of Passiflora are found in Mexico, Central America, the United States and South America, although there are additional representatives in Southeast Asia and Oceania.<ref>Template:Cite journal</ref> New species continue to be identified: for example, P. xishuangbannaensis and P. pardifolia have only been known to the scientific community since 2005 and 2006, respectively.

Some species of Passiflora have been naturalized beyond their native ranges. For example, the blue passion flower (P. caerulea) now grows wild in Spain.<ref name=danaetal2001>Template:Cite journal</ref> The purple passionfruit (P. edulis) and its yellow relative flavicarpa have been introduced in many tropical regions as commercial crops.

EcologyEdit

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Passion flowers have floral structures adapted for biotic pollination. Pollinators of Passiflora include bumblebees, carpenter bees (e.g., Xylocopa sonorina), wasps, bats, and hummingbirds (especially hermits such as Phaethornis); some others are additionally capable of self-pollination. Passiflora often exhibit high levels of pollinator specificity, which has led to frequent coevolution across the genus. The sword-billed hummingbird (Ensifera ensifera) is a notable example: it, with its immensely elongated bill, is the sole pollinator of 37 species of high Andean Passiflora in the supersection Tacsonia.<ref>Template:Cite journal</ref>

The leaves are used for feeding by the larvae of a number of species of Lepidoptera. Famously, they are exclusively targeted by many butterfly species of the tribe Heliconiini. The many defensive adaptations visible on Passiflora include diverse leaf shapes (which help disguise their identity), colored nubs (which mimic butterfly eggs and can deter Heliconians from ovipositing on a seemingly crowded leaf), extrafloral nectaries, trichomes, variegation, and chemical defenses.<ref>Template:Cite journal</ref> These, combined with adaptations on the part of the butterflies, were important in the foundation of coevolutionary theory.<ref>Template:Cite journal</ref><ref>Template:Cite journal</ref>

Recent studies have shown that passiflora both grow faster and protect themselves better in high-nitrogen soils. In low-nitrogen environments, passiflora focus on growth rather than defense and are more vulnerable to herbivores.<ref>Template:Cite journal</ref>

The following lepidoptera larvae are known to feed on Passiflora:

The generally high pollinator and parasite specificity in Passiflora may have led to the tremendous morphological variation in the genus. It is thought to have among the highest foliar diversity among all plant genera,<ref name="Killip 1938">Template:Cite book</ref> with leaf shapes ranging from unlobed to five-lobed frequently found on the same plant.<ref>Template:Cite journal</ref> Coevolution can be a major driver of speciation, and may be responsible for the radiation of certain clades of Passiflora such as Tacsonia.

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.<ref name="Radhamani et al 1995">Template:Cite journal</ref>

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 an invasive species in Spain and considered likely to threaten ecosystems there.<ref name = danaetal2001 />

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 Tropical Andes southwards from Venezuela between Template:Convert in altitude, and in Coastal Central Chile, where it only occurs in a few tens of square kilometres of fog forest by the sea, near Zapallar. 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.Template:Citation needed

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.

Adverse effectsEdit

Passion flower is not recommended during pregnancy because it may induce contractions.<ref name="nih" /><ref name="drugs" /> Consuming passion flower products may cause drowsiness, nausea, dizziness, abnormal heart rhythms, asthma, or rhinitis.<ref name="nih" /><ref name="drugs" />

UsesEdit

OrnamentalEdit

File:Flickr - ggallice - Passiflora, Gainesville.jpg
Passiflora incarnata, one of the most common passion flowers

A number of species of Passiflora are cultivated outside their natural range for both their flowers and fruit. Hundreds of hybrids have been named; hybridizing is currently being done extensively for flowers, foliage and fruit. The following hybrids and cultivars have gained the Royal Horticultural Society's Award of Garden Merit:

  • 'Amethyst'<ref>{{#invoke:citation/CS1|citation

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During the Victorian era the flower (which in all but a few species lasts only one day) was very popular, and many hybrids were created using the winged-stem passion flower (P. alata), the 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.<ref>Petersen (1966)</ref>

Passion flowers have been a subject of studies investigating extranuclear inheritance; paternal inheritance of chloroplast DNA has been documented in this genus.<ref>E.g. Hansen et al. (2006)</ref> The plastome of the two-flowered passion flower (P. biflora) has been sequenced.

FruitEdit

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File:Passion fruits - whole and halved.jpg
Ripe fruits of Passiflora edulis

Most species have round or elongated edible fruit.

  • The passion fruit or {{#invoke:Lang|lang}} (P. edulis) is cultivated extensively in the Caribbean, South America, south Florida and South Africa for its fruit, which is used as a source of juice. A small pink fruit that wrinkles easily and a larger shiny yellow to orange fruit are traded under this name. The latter is usually considered just a variety of flavicarpa, but seems to be more distinct.Template:Citation needed
  • Sweet granadilla (P. ligularis) is another widely grown species. In large parts of Africa 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 Template:Convert 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,<ref>Template:Cite EB1911</ref> but not widely known elsewhere Template:As of.Template:Cn
  • The blue passionflower (Passiflora caerulea) produces bright orange fruit with numerous seeds. While the fruit is edible, it is often described as being bland in comparison to other edible passionfruit, or with a flavour vaguely similar to blackberries.<ref>{{#invoke:citation/CS1|citation

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Ayahuasca analogEdit

A native source of beta-carbolines (e.g., passion flower in North America) is mixed with Desmanthus illinoensis (Illinois bundleflower) root bark to produce a hallucinogenic drink called prairiehuasca, which is an analog of the shamanic brew ayahuasca.<ref>Template:Cite book</ref>

Traditional medicine and dietary supplementEdit

Passiflora incarnata (maypop) leaves and roots have a long history of use as a traditional medicine by Native Americans in North America and were adapted by European colonists.<ref name="nih">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref name="drugs">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> The fresh or dried leaves of maypop are used to make a tea that is used as a sedative.<ref name=nih/> Passionflower as dried powder or an extract is used as a dietary supplement.<ref name=nih/> There is insufficient clinical evidence for using passionflower to treat any medical condition.<ref name=nih/><ref name=drugs/>

Passionflower is classified as generally recognized as safe for use as a food ingredient in the U.S.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref>

In cultureEdit

File:Passiflora caerulea (makro close-up).jpg
Blue passion flower (P. caerulea) showing most elements of the Christian symbolism
File:Passiflora Krishnakamal Karnataka India.jpg
Passiflora 'Soi Fah', Template:Aka Template:Transliteration in India

The passion in passion flower purportedly refers to the passion of Jesus in Christian theology;<ref name="Kostka">{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> the word passion comes from the Latin {{#invoke:Lang|lang}}, meaning 'suffering'. 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:<ref name="Hammer2015">Template:Cite book</ref>

The flower has been given names related to this symbolism throughout Europe since the 15th century. In Spain, it is known as {{#invoke:Lang|lang}} ('thorn of Christ'). Older Germanic names<ref>Marzell (1927)</ref> include {{#invoke:Lang|lang}} ('Christ's crown'), {{#invoke:Lang|lang}} ('Christ's bouquet'),<ref>"Christ's flower" is a mistranslation of Marzell (1927)</ref> {{#invoke:Lang|lang}} ('crown of thorns'), {{#invoke:Lang|lang}} ('Jesus' passion'), {{#invoke:Lang|lang}} ('passion')<ref>"Martyr" is a mistranslation of Marzell (1927)</ref> or {{#invoke:Lang|lang}} ('Mother of God's star').<ref>Muttergottes-Schuzchen (or -Schurzchen) is a nonsensical misreading of Marzell (1927)</ref>

Outside the Roman Catholic heartland, the regularly shaped flowers have reminded people of the face of a clock. In Israel they are known as "clock-flower" ({{#invoke:Lang|lang}}) and in Greece as "clock plant" ({{#invoke:Lang|lang}}); in Japan too, they are known as Template:Nihongo3. In Hawaiian, they are called {{#invoke:Lang|lang}};<ref>Template:Hawaiian Dictionaries</ref> {{#invoke:Lang|lang}} is a string used for tying fabric together, such as a shoelace, and {{#invoke:Lang|lang}} means 'to spring forth leave'.<ref>Pukui et al. (1992)</ref>

In India, it is known as Krishnakamala because of its relation to the Epic Mahabharata. The 100 peripheral petals represent the 100 Kauravas, while the 5 yellow petals in between stand for the Pandavas. The sensitive, but firm green bulb on top is Draupadi, or Krishna.

GalleryEdit

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See alsoEdit

ReferencesEdit

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External linksEdit

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Template:Carnivorous plants Template:Passiflora Template:Taxonbar

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