View source for Piper (plant)

{{Short description|Genus of plants}}
{{DISPLAYTITLE:''Piper'' (plant)}}
{{Automatic taxobox
| fossil_range = {{fossil range|Maastrichtian|present|earliest=Late Cretaceous|[[Maastrichtian]] - present|ref=<ref name=Martinez2015>{{Cite journal |last1=Martínez |first1=Camila |last2=Carvalho |first2=Mónica R. |last3=Madriñán |first3=Santiago |last4=Jaramillo |first4=Carlos A. |date=2015 |title=A Late Cretaceous''Piper''(Piperaceae) from Colombia and diversification patterns for the genus |journal=American Journal of Botany |volume=102 |issue=2 |pages=273–289 |doi=10.3732/ajb.1400427 |pmid=25667080 |issn=0002-9122|doi-access=free }}</ref>}}
| image = Piper magnificum0.jpg
| image_caption = Lacquered pepper (''[[Piper magnificum|P. magnificum]]'') [[inflorescence]]
| taxon = Piper
| authority = [[Carl Linnaeus|L.]]
| subdivision_ranks = Species
| subdivision = 1000–2000; see [[List of Piper species|list]]
| synonyms = *''Anderssoniopiper'' <small>Trel.</small>
*''Arctottonia'' <small>Trel.</small>
*''Artanthe'' <small>Miq.</small>
*''Chavica'' <small>Miq.</small>
*''Discipiper'' <small>Trel. & Stehlé</small>
*''Lepianthes'' <small>Raf.</small>
*''Lindeniopiper'' <small>Trel.</small>
*''Macropiper'' <small>Miq.</small>
*''Ottonia'' <small>Spreng.</small>
*''Pleiostachyopiper'' <small>Trel.</small>
*''Pleistachyopiper'' <small>Trel.</small>
*''Pothomorphe'' <small>Miq.</small>
*''Trianaeopiper'' <small>Trel.</small>
}}

'''''Piper''''', the '''pepper plants''' or '''pepper vines''', is an economically and ecologically important [[genus]] in the family [[Piperaceae]]. The genus contains about 1,000–2,000 species of shrubs, herbs, and [[liana]]s, many of which are dominant species in their native habitat. The diversification of this taxon is of interest to understanding the evolution of plants.

The [[scientific name]] ''Piper'' and the common name "pepper" are derived from the [[Sanskrit]] term ''pippali'', denoting the [[long pepper]] (''P.&nbsp;longum'').

== Description ==
Most ''Piper'' species are either [[herbaceous]] or [[vine]]s; some grow as [[shrub]]s or almost as small [[tree]]s.

== Taxonomy ==
Pepper plants belong to the [[magnoliids]], which are [[angiosperm]]s but neither [[monocots]] nor [[eudicots]]. Their family, Piperaceae, is most closely related to the lizardtail family ([[Saururaceae]]), which in fact generally look like smaller, more delicate and [[wikt:Special:Search/amphibious|amphibious]] pepper plants. Both families have characteristic tail-shaped [[inflorescence]]s covered in tiny flowers. A somewhat less close relative is the pipevine family ([[Aristolochiaceae]]). A well-known and very close relative – also part of the Piperaceae – are the radiator plants of the genus ''[[Peperomia]]''.

=== Evolution ===
The earliest fossil of ''Piper'' is of †''Piper margaritae'' from the [[Late Cretaceous]] ([[Maastrichtian]]) of [[Colombia]]. ''P.&nbsp;margaritae'' appears to be nested within the clade ''Schilleria'', indicating extensive Cretaceous diversification of ''Piper'' into the multiple extant clades, coinciding with the final breakup of [[Gondwana]]. This contrasts with previous theories assuming a younger radiation of the genus. An earlier potential record is of †''Piper arcuatile'' from the [[Cenomanian]] to [[Santonian]] [[Kaltag Formation]] of [[Yukon]], although this affinity to ''Piper'' is not entirely reliable.<ref name="Martinez2015" /><ref>{{Cite journal |last1=Hollick |first1=Arthur |last2=Martin |first2=G.C. |date=1930 |title=The Upper Cretaceous floras of Alaska, with a description of the plant-bearing beds |url=http://dx.doi.org/10.3133/pp159 |journal=Professional Paper |doi=10.3133/pp159 |issn=2330-7102 |url-access=subscription}}</ref><ref>{{Cite web |title=Yukok_Koyukuk_2_Geology |url=http://clamp.ibcas.ac.cn/arcticfossils/Yukon_Koyukuk2_Geology.html |access-date=2023-06-19 |website=clamp.ibcas.ac.cn}}</ref>

=== Species ===
{{See also|List of Piper species}}
The largest number of ''Piper'' species are found in the Americas (about 700 species), with about 300 species from Southern Asia. There are smaller groups of species from the South Pacific (about 40 species) and Africa (about 15 species). The American, Asian, and South Pacific groups each appear to be [[monophyletic]]; the affinity of the African species is unclear.<ref name="jaramillo">{{cite journal |author=M. Alejandra Jaramillo |author2=Paul S. Manos |year=2001 |title=Phylogeny and patterns of floral diversity in the genus ''Piper'' (Piperaceae) |journal=[[American Journal of Botany]] |volume=88 |issue=4 |pages=706–716 |doi=10.2307/2657072 |jstor=2657072 |pmid=11302858}}</ref>

Some species are sometimes [[segregate (taxonomy)|segregated]] into the genera ''Pothomorphe'', ''Macropiper'', ''Ottonia'', ''Arctottonia'', ''Sarcorhachis'', ''Trianaeopiper'', and ''Zippelia'', but other sources keep them in ''Piper''.<ref name="jaramillo" />

The species called "''Piper aggregatum''" and "''P.&nbsp;fasciculatum''" are actually ''[[Lacistema aggregatum]]'', a plant from the family [[Lacistemataceae]].

==Distribution and ecology==
''Piper'' species have a [[tropical|pantropical]] distribution, and are most commonly found in the understory of lowland [[tropical forest]]s, but can also occur in clearings and in higher elevation life zones such as [[cloud forest]]s; one species &ndash; the Japanese Pepper (''[[Piper kadsura|P.&nbsp;kadsura]]'') from southern [[Japan]] and southernmost [[Korea]] &ndash; is subtropical and can tolerate light winter [[frost]]. Peppers are often dominant species where they are found.

A few species, commonly called "ant pipers" (e.g. ''[[Piper cenocladum]]''), live in a [[Mutualism (biology)|mutualism]] with [[ant]]s. The fruit of the ''Piper'' plant, called a peppercorn when it is round and pea-sized, as is usual, is distributed in the wild mainly by [[bird]]s, but small fruit-eating [[mammal]]s – e.g. [[bat]]s of the genus ''[[Carollia]]'' – are also important. Despite the high content of chemicals that are noxious to [[herbivore]]s, some have [[evolved]] the ability to withstand the chemical defences of pepper plants, for example the [[Sematurinae|sematurine]] [[moth]] ''[[Homidiana subpicta]]'' or some [[flea beetle]]s of the genus ''[[Lanka (beetle)|Lanka]]''. The latter can be significant [[pest (organism)|pests]] to pepper growers.

==Uses==
Many pepper plants make good [[ornamental plant|ornamentals]] for gardens in [[subtropical]] or warmer regions. Pepper vines can be used much as [[ivy]] in [[temperate climate]]s, while other species, like lacquered pepper (''[[Piper magnificum|P.&nbsp;magnificum]]'') grow as sizeable, compact and attractive shrubs with tough and shiny leaves. Smaller species, like Celebes pepper (''[[Piper ornatum|P.&nbsp;ornatum]]'') with its finely patterned leaves, are also suitable as indoor [[pot plant]]s.

Unsustainable [[logging]] of tropical [[primary forest]]s is threatening a number of peppers. The extent of the effect of such wholesale [[habitat destruction]] on the genus is unknown, but in the forests of [[Ecuador]] – the only larger region for which comprehensive data exists<ref>[[International Union for the Conservation of Nature and Natural Resources]] (IUCN) (2007): ''[http://www.iucnredlist.org 2007 IUCN Red List of Threatened Species] {{Webarchive|url=https://web.archive.org/web/20180508171911/http://www.iucnredlist.org/ |date=2018-05-08 }}''. IUCN, Gland.</ref> – more than a dozen species are known to be on the brink of [[extinction]]. On the other hand, other ''Piper'' species (e.g. spiked pepper, ''[[Piper aduncum|P.&nbsp;aduncum]]'') have been widely distributed as a result of human activity and are a major [[invasive species]] in certain areas.

The most significant human use of ''Piper'' is not for its looks however, but ultimately for the wide range of powerful [[secondary compounds]] found particularly in the fruits.

===Culinary===
Culinary use of pepper plants is attested perhaps as early as 9,000 years ago. Peppercorn remains were found among the food refuse left by [[Hoabinhian]] artisans at [[Spirit Cave (Thailand)|Spirit Cave]], [[Thailand]]. It is likely that these plants were collected from the wild rather than deliberately grown.<ref>{{cite journal |author=Chester F. Gorman |year=1969 |title=Hoabinhian: a pebble tool complex with early plant associations in Southeast Asia |journal=[[Science (journal)|Science]] |volume=163 |issue=3868 |pages=671–673 |doi=10.1126/science.163.3868.671 |pmid=17742735|bibcode=1969Sci...163..671G |s2cid=34052655 }}</ref><ref>{{cite journal |author=Chester F. Gorman |year=1971 |title=The Hoabinhian and after: subsistence patterns in Southeast Asia during the Late Pleistocene and Early Recent periods |journal=[[World Archaeology]] |volume=2 |issue=3 |pages=300–320 |jstor=124120 |doi=10.1080/00438243.1971.9979482}}</ref>

[[File:Pfeffer-Gewürz.jpg|thumb|[[Black pepper]] (''Piper nigrum'') corns, from left to right:<br />Green (pickled unripe fruits)<br />White (dried ripe seeds)<br />Black (dried unripe fruits)]]
Use of peppercorns as pungent spice is significant on an international scale. By [[classical antiquity]], there was a vigorous [[spice trade|trade of spices]] including black pepper (''[[Piper nigrum|P.&nbsp;nigrum]]'') from [[South Asia]] to [[Europe]]. The ''[[Apicius]]'', a recipe collection complied about 400 AD, mentions "pepper" as a spice for most main dishes.<ref>{{cite book |title=Apicius: Cookery and Dining in Imperial Rome |url=http://www.gutenberg.org/files/29728/29728-h/29728-h.htm |website=Project Gutenburg |access-date=December 9, 2018|date=2009-08-19 }}</ref> In the late [[Roman Empire]], black pepper was expensive, but was available readily enough to be used more frequently than [[salt]]{{efn|''Apicius'' generally uses ''[[garum]]'' [[fish sauce]] instead; raw [[brine]] and large quantities of [[herb]]s were also employed by many.}} or [[sugar]].

As Europe moved into the [[Early Middle Ages]], trade routes deteriorated and the use of pepper declined somewhat, but peppercorns, storing easily and having a high mass per volume, never ceased to be a profitable trade item. In the [[Middle Ages]], international traders were nicknamed ''Pfeffersäcke'' ("pepper-sacks") in [[Germany|German]] towns of the [[Hanseatic League]] and elsewhere. Later, wars were fought by European powers, between themselves and in complex alliances and enmities with [[Indian Ocean]] states, in part about control of the supply of spices, perhaps the most [[archetypal]] being black pepper fruit. Today, peppercorns of the three preparations (green, white and black) are one of the most widely used spices of plant origin worldwide.

Due to the wide distribution of ''Piper'', the fruit of other species are also important spices, many of them internationally. [[Long pepper]] (''P.&nbsp;longum''), is possibly the second-most popular ''Piper'' spice internationally; it has a rather [[Chili pepper|chili]]-like "heat" and the whole [[inflorescence]] is used as the fruits are tiny. [[Cubeb]] (''P.&nbsp;cubeba''), also known as tailed pepper, played a major role in the spice trade. Reputedly [[Philip IV of Spain]] suppressed trade in cubeb peppercorns at the end of the 1630s to capitalize on his share of the black pepper trade.<ref>{{cite book |author=John Parkinson |year=1640 |title=Theatrum Botanicum: the Theater of Plants |url=https://archive.org/details/gri_33125008297760 |publisher=T. Cotes |location=London|author-link=John Parkinson (botanist) }}</ref> It remains a significant spice around the [[Indian Ocean]] region today, however. West African pepper (''[[Piper guineense|P.&nbsp;guineense]]''), is commonly used in [[West African cuisine]], and is sometimes used in the [[East Africa]]n ''[[berbere]]'' spice mix. This species, despite being traded more extensively in earlier times, is less common outside Africa today.

Not only the seeds of ''Piper'' are used in cooking. West African Pepper leaves, known locally as ''uziza'', are used as a flavoring vegetable in [[Nigerian cuisine|Nigerian]] stews. In [[Mexico|Mexican]]-influenced cooking, ''hoja santa'' or Mexican pepperleaf (''[[Piper auritum|P.&nbsp;auritum]]'') has a variety of uses. In [[Southeast Asia]], leaves of two species of ''Piper'' have major importance in cooking: [[lolot]] (''P.&nbsp;lolot'') is used to wrap meat for grilling in the [[Indochina]] region, while wild betel (''[[Piper sarmentosum|P.&nbsp;sarmentosum]]'') is used raw or cooked as a vegetable in [[Malay cuisine|Malay]] and [[Thai cuisine]];<ref>{{cite book |author=Charmaine Solomon |year=1998 |chapter=Cha plu |title=Encyclopedia of Asian Food |publisher=Periplus Editions |chapter-url=http://www.asiafood.org/glossary_1.cfm?alpha=C&wordid=3252&startno=1&endno=25}}</ref><ref>{{cite web |year=2008 |url=http://www.earthcare.com.au/other.htm |title=Tropical root crops |publisher=Earthcare |access-date=15 July 2008 |url-status=dead |archive-url=https://web.archive.org/web/20080719012840/http://www.earthcare.com.au/other.htm |archive-date=19 July 2008 }}</ref> The stems and roots of ''[[Piper chaba]]'' are used as a spice in [[Bangladeshi cuisine]].

===Medicine===
[[File:Faikava.jpg|thumb|At a [[kava culture#Tonga|Kava club in Tonga]] ({{circa|2005}})]]

[[Cubeb]] (''P. cubeba'') has been used in [[folk medicine]] and [[herbalism]] as well as, particularly in the early 20th century, as a [[cigarette]] flavoring. ''[[Piper darienense|P.&nbsp;darienense]]'' is used medically by the [[Guna people]] of the [[Panama]]-[[Colombia]] border region, and elsewhere it is used to intoxicate fish which then can be easily caught. [[Spiked pepper]], often called ''matico'' appears to have strong [[disinfectant]] and [[antibiotic]] properties. Black pepper (''P.&nbsp;nigrum'') [[essential oil]] is sometimes used in herbalism, and long pepper (''P.&nbsp;longum'') is similarly employed in [[Ayurveda]], where it was an ingredient of ''Triphala Guggulu'' and (together with black pepper) of ''Trikatu'' [[Pill (pharmacy)|pills]], used for ''[[rasayana]]'' (rejuvenating and [[Body cleansing|detoxifying]]) purposes.

One ''Piper'' species has gained large-scale use as a [[stimulant]]. [[Betel]] (''P.&nbsp;betle'') leaves are used to wrap [[betel palm]] nut slices; its sap helps release the stimulating effect of these "cookies" which are widely known as ''pan'' in [[India]].

Conversely, another ''Piper'' species, [[kava]] (''P.&nbsp;methysticum''), is used for its [[depressant]] and [[euphoriant]] effects. In the [[Pacific]] region, where it has been widely spread as a [[Canoe plants|canoe plant]], kava is used to produce a calming and socializing drink somewhat similar to alcohol and [[benzodiazepines]] but without many of the negative side effects and less of an [[Substance use disorder|addiction]] risk. It has also become popular elsewhere in recent decades, and is used as a medical plant. However, pills that contain parts of the whole plant have occasionally shown a strong [[hepatotoxic]] effect, which has led to the banning of kava in many countries. On the other hand, the traditional preparation of the root as a calming drink appears to pose little, if any, such hazard.<ref>{{cite journal |author=L. Sorrentino |author2=A. Capasso |author3=M. Schmidt |year=2006 |title=Safety of ethanolic kava extract: results of a study of chronic toxicity in rats |journal=[[Phytomedicine (journal)|Phytomedicine]] |volume=13 |issue=8 |pages=542–549 |doi=10.1016/j.phymed.2006.01.006 |pmid=16904878}}</ref><ref>{{cite journal |author=Steven T. S. Lim |author2=Klaus Dragull |author3=Chung-Shih Tang |author4=Harry C. Bittenbender |author5=Jimmy T. Efird |author6=Pratibha V. Nerurkar |year=2007 |title=Effects of kava alkaloid, pipermethystine, and kavalactones on oxidative stress and cytochrome P450 in F-344 rats |journal=[[Toxicological Sciences]] |volume=97 |issue=1 |pages=214–221 |doi=10.1093/toxsci/kfm035 |pmid=17329236  |doi-access=free }}</ref>{{update after|2017|02|28}}

===In science===
The genus contains species suitable for studying [[natural history]], [[molecular biology]], [[natural product]]s [[chemistry]], [[community ecology]], and [[evolutionary biology]].<ref>{{cite book |author=L. A. Dyer |author2=A. N. Palmer |year=2004 |title=''Piper'': a Model Genus for Studies of Evolution, Chemical Ecology, and Trophic Interactions |publisher=Kluwer Academic Publishers |location=Boston, MA |isbn=978-0-306-48498-8 |url=http://wolfweb.unr.edu/~ldyer/KI059-Dyer-all.pdf}}</ref>

[[File:Piperine crystals.jpg|thumb|[[Crystal]]lized [[piperine]], extracted from black pepper (''P.&nbsp;nigrum'')]]

''Piper'' is a [[model organism|model genus]] for research in [[ecology and evolutionary biology]]. The diversity and ecological importance of the genus makes it a strong candidate for ecological and evolutionary studies. Most research has focused on the economically important species ''P.&nbsp;nigrum'' (black pepper), ''P.&nbsp;methysticum'' (kava), and ''P.&nbsp;betle'' (betel). A recent study based on DNA sequence analysis suggest that ''P.&nbsp;nigrum'' originated in the [[Western Ghats]] hot spot in India.<ref>{{Cite journal|last1=Sen|first1=Sandeep|last2=Dayanandan|first2=Selvadurai|last3=Davis|first3=Thomson|last4=Ganesan|first4=Rengaian|last5=Jagadish|first5=M.R.|last6=Mathew|first6=P.J.|last7=Ravikanth|first7=Gudasalamani|date=September 2019|title=Origin and evolution of the genus Piper in Peninsular India|journal=Molecular Phylogenetics and Evolution|volume=138|pages=102–113|doi=10.1016/j.ympev.2019.05.033|pmid=31132521|s2cid=167218493 |issn=1055-7903|url=https://spectrum.library.concordia.ca/985469/1/Origin-and-evolution-of-the-genus-Piper-in-P_2019_Molecular-Phylogenetics-an.pdf }}</ref>

The obligate and facultative{{Explain|date=February 2023}} [[ant]] [[mutualists]] found in some ''Piper'' species have a strong influence on their biology, making them ideal systems for research on the evolution of [[symbioses]] and the effect of mutualisms on biotic communities.{{citation needed|date=February 2023}}

Important [[secondary metabolite]]s found in pepper plants are [[piperine]] and [[chavicine]], which were first isolated from Black Pepper, and reported to have antibiotic activities. Preliminary research reports has shown that piperine has an antibacterial activity against various bacteria such as ''S.&nbsp;aureus'',<ref>{{cite journal |last1=Khan |first1=IA. |display-authors=etal | title = Piperine, a Phytochemical Potentiator of Ciprofloxacin against Staphylococcus aureus| journal = Antimicrobial Agents and Chemotherapy | volume = 50 | issue = 2 | pages = 810–812 | pmid = 16436753 | doi=10.1128/aac.50.2.810-812.2006 | pmc=1366922 | year=2006}}</ref><ref>{{cite journal |last1=Kumar |first1=A. |display-authors=etal | title = Novel structural analogues of piperine as inhibitors of the NorA efflux pump of Staphylococcus aureus| journal = Journal of Antimicrobial Chemotherapy | volume = 61 |issue= 6 | pages = 1270–1276 | pmid = 18334493 | doi=10.1093/jac/dkn088 | year=2008| doi-access= free }}</ref> ''Streptococcus mutans'',<ref>{{cite journal |last1=Dwivedi |first1=D. |display-authors=etal | title = Effects of the natural compounds embelin and piperine on the biofilm-producing property of Streptococcus mutans | journal = Journal of Traditional and Complementary Medicine | volume = 6 | issue = 1 | pages = 57–61 | pmid = 26870681 | doi=10.1016/j.jtcme.2014.11.025 | pmc=4738039 | year=2016}}</ref> and gastric cancer pathogen ''Helicobacter pylori'' <ref>{{cite journal | last1= Tharmalingam |first1= N. |last2=Kim |first2=SH. |last3=Park |first3=M. | title = Inhibitory effect of piperine on Helicobacter pylori growth and adhesion to gastric adenocarcinoma cells | journal = Infectious Agents and Cancer | volume = 16 | issue = 9 | pages = 43 | doi = 10.1186/1750-9378-9-43 | pmid = 25584066 | pmc=4290101 | year=2014 |doi-access= free }}</ref> and decreased ''H. pylori'' toxin entry to gastric epithelial cells.<ref>{{cite journal | last1= Tharmalingam |first1= N. |last2=Park |first2=M. |last3=LEE |first3=M.H | title = Piperine treatment suppresses Helicobacter pylori toxin entry in to gastric epithelium and minimizes β-catenin mediated oncogenesis and IL-8 secretion in vitro | journal = American Journal of Translational Research | volume = 15 | issue = 8 | pages = 885–98 | pmid = 27158376 | pmc=4846933 | year=2016}}</ref> The [[piperidine]] [[functional group]] is named after the former, and [[piperazine]] (which is not found in ''P.&nbsp;nigrum'' in noticeable quantities) was in turn named after piperidine.

The significant secondary metabolites of kava are [[kavalactone]]s and [[flavokawain]]s. [[Pipermethystine]] is suspected to be the main [[hepatotoxic]] compound in this plant's stems and leaves.

===Repelling insects===
Studies have been done to determine the effectiveness of piper leaves to repel different types of insects.<ref>{{Cite journal |last1=Scott |first1=I. M. |last2=Jensen |first2=H. |last3=Nicol |first3=R. |last4=Lesage |first4=L. |last5=Bradbury |first5=R. |last6=Sánchez-Vindas |first6=P. |last7=Poveda |first7=L. |last8=Arnason |first8=J. T. |last9=Philogène |first9=B. J. R. |date=August 2004 |title=Efficacy of Piper (Piperaceae) extracts for control of common home and garden insect pests |journal=Journal of Economic Entomology |volume=97 |issue=4 |pages=1390–1403 |doi=10.1093/jee/97.4.1390 |issn=0022-0493 |pmid=15384353|hdl=11056/23254 |hdl-access=free }}</ref>

[[Capuchin monkey]]s have been recorded by [[BBC Earth]] rubbing the piper leaves on them to repel insects.<ref>{{Citation |title=Monkey Insect Repellent {{!}} The Life of Mammals {{!}} BBC Earth |url=https://www.youtube.com/watch?v=9rhgafQCCIg |access-date=2023-09-25 |language=en}}</ref>

==Notes==
{{Notelist}}

==References==
{{Reflist|32em}}
* {{Cite journal|first1=Sandeep|last1=Sen|last2=Dayanandan|first2=Selvadurai|last3=Davis|first3=Thomson|last4=Ganesan|first4=Rengaian|last5=Jagadish|first5=M.R.|last6=Mathew|first6=P.J.|last7=Ravikanth|first7=Gudasalamani|date=September 2019|title=Origin and evolution of the genus Piper in Peninsular India|journal=Molecular Phylogenetics and Evolution|volume=138|pages=102–113|doi=10.1016/j.ympev.2019.05.033|pmid=31132521|s2cid=167218493 |issn=1055-7903|url=https://spectrum.library.concordia.ca/985469/1/Origin-and-evolution-of-the-genus-Piper-in-P_2019_Molecular-Phylogenetics-an.pdf }}

==External links==
{{Commons category|Piper}}
{{Wikispecies|Piper}}
*[http://delta-intkey.com/angio/ The families of flowering plants]: descriptions, illustrations, identification, information retrieval.
* [http://home.kku.ac.th/raccha/Piper.html Piper species in Thailand]

{{Taxonbar|from=Q159536}}
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[[Category:Piper (plant)| ]]
[[Category:Pantropical flora]]
[[Category:Piperales genera]]