ThePapaveraceae,/pəˌpævəˈreɪsiˌiː/[2] informally known as thepoppy family, are an economically importantfamily of about 42genera and approximately 775 knownspecies[3] offlowering plants in the orderRanunculales. The family iscosmopolitan, occurring intemperate andsubtropical climates (mostly in the northern hemisphere) like Eastern Asia as well as California in North America. It is almost unknown in thetropics. Most areherbaceous plants, but a few areshrubs and smalltrees. The family currently includes two groups that have been considered to be separate families: Fumariaceae and Pteridophyllaceae.Papaver is the classical name for poppy in Latin.[4]
Papaveraceae are known for diverse and colorful flowers with distinctive sepals. The plants may be annual, biennial, orperennial. Usuallyherbaceous, a few species form shrubs or evergreen trees. All parts contain a well-developed system of latex ducts called "laticifers", that produce milkylatex, a watery white, yellow or red juice.
Theleaves are alternate or sometimes whorled and havepetioles and nostipules. They are usually lobed or pinnatifid (i.e. consisting of several not entirely separate leaflets), or much divided.
The plants arehermaphroditic andpollinated mostly byinsects (entomophilous), but nectaries are lacking. A few are wind pollinated (anemophilous). There is a distinctcalyx andcorolla, except inMacleaya where the corolla is lacking. The flowers are medium-sized or large. The terminal flowers are solitary in many species. In others the terminalinflorescence is cymose orracemose. The flowers are odourless and regular.
There are manystamens, mostly 16 to 60, arranged in two separate whorls, the outer one with stamens alternating with petals, the inner one opposite, or numerous in the subfamilyPapaveroideae.[5]: 86 The gynoecium consists of a compoundpistil with 2 to 100 carpels. Theovary is superior andunilocular. The ovary is either stemless (sessile) or on a short stem (stipitate). The sepals of the plant typically number half of the petals for example two sepals accompany 4 petals or 3 sepals accompany 6 petals. The pistils and stamens are hidden inside the petals.
The non-fleshyfruit is usually acapsule, breaking open at maturity to release theseeds through pores (poricidal), through the partitions between the cells (septicidal), or by means of valves (valvular). The numerous seeds are small. Their nutritive tissue (endosperm) is oily and farinose. The fruit ofPlatystemon is aschizocarp.
The Papaveraceae family includes many plants that produce alkaloids, including opium poppy (Papaver somniferum). Opium is derived from the latex of the opium poppy seed pods and has been used for centuries due to its psychoactive properties. The main alkaloids found in opium, such as morphine, and codeine have huge impact on pharmaceuticals and a big importance on many countries that produce it on a large-scale including Afghanistan. Afghanistan has an economic dependence on opium cultivation making it hard to stop large scale production of these flowers ultimately increasing illegal production. Researchers are understanding how alkaloids are made in poppy plants to develop poppy plants with specific alkaloid levels. There is high genetic variability among poppy cultivars and environmental factors like wounding and methyl jasmonate treatment induce higher alkaloid production. Challenges still remain in understanding what effects alkaloid production for pharmaceutical purposes, highlighting importance of research in this field.[citation needed]
The broadcircumscription of Papaveraceae in the APG III system includes three taxa that have previously been separated into different families: the Papaveraceaesensu stricto, the Fumariaceae and the Pteridophyllaceae.[1] Thus theCronquist system of 1981 recognised the Fumariaceae as a separate family, despite their close phylogenetic relationship to the Papaveraceaesensu stricto. The three former families may be treated as subfamilies. One morphological and molecular study concluded that the former family Pteridophyllaceae has a basal position with a subsequent division into two terminal clades each containing one of the subfamilies Fumarioideae and Papaveroideae, which are clearly monophyletic.[7] A more recent study includes the former Pteridophyllaceae in the Fumarioideae, dividing the Papaveraceae into only two subfamilies.[8]
The internal division of the Fumarioideae shown below follows Lidén (1993),[9] with the exception of the placement ofPteridophyllum.[8][6] The subtribes are given by theGermplasm Resources Information Network.[10] The division of the Papaveroideae follows Hoot et al. (1997).[7] In the latter study, the tribe Eschscholzieae would be the basal clade and sister group to the rest of the subfamily, which is divided into a different terminal clade (Chelidonieae) and into its sister group, formed by the Papavereae and Platystemoneae, whose separation is not based on the data presented by these authors. For discussions of subfamilies, see Carolan et al. (2006)[11] and Blattner & Kadereit (1999).[12]
Pollination isentomophile (basically byflies andwasps and bees, less often bybeetles), except inBocconia andMacleaya. In Papaveroideae, the reward is pollen as there is no nectar. The visual attractant is the petals that are usually brightly coloured and often have basalguides, sometimes the attractant can also be theandroecium as the petals do not last long. Some species, mostly those from the arctic and alpine regions, reinforce their attraction with floral fragrance (for example,Papaver alpinum smells of cloves), which in the case ofRomneya drugs the insects. Theanthers andstigmas mature at the same time, butBocconia is clearlyprotogynous, the stigmas emerge from thecalyx that encloses them.Autopollination is common and in some cases (for example,Roemeria hybrida) it occurs before the bud opens (cleistogamy). The presence of anaril suggests dispersion of seeds by ants (myrmecochory), once they have been expelled by the fruit. In the case ofBocconia the seeds remain attached to thereplums after the capsule'svalves have fallen leaving their brilliant red or orange arils exposed, which attract birds to feed on them, facilitating their dispersal (ornithochory). Seeds that lack an aril appear to be dispersed by the wind (anemochory) for capsules that open, in the other cases they are freed when the fruit decomposes. ManyFumarioideae species have explosive fruits (ballistic), whileRupicapnos andSarcocapnos species arechasmophytes, growing on rocks, and their fruit'speduncles andpedicels aregeotropic and they lengthen so that the seeds bury into the base of the plant.
The Papaveroideae typically grow in cooler and wooded areas, forming part of the undergrowth. They have adapted to arctic and alpine habitats and to arid, Mediterranean areas, many species areruderal and segetal (growing in cornfields).Pteridophyllum grows in the undergrowth of woods of needle-leaved trees between 1,000 and 2,000 metres (3,300 and 6,600 ft). The Fumarioideae are basically found in open, rocky, alpine landscapes or vertical or overhanging cracks, while some species are ruderal or segetal.
Alkaloids: Theisoquinolinic alkaloids present in the family are well known. They are derived fromberberine, tetrahydroberberine,protopine and benzophenanthridine in Papaveroideae, and from spirobenzylisoquinoline and cularine in Fumarioideae, as well as from other groups that give them pharmacological properties: derivatives ofaporphine,morphinan,pavine,isopavine,narceine andrhoeadine.
Others: Other characteristic substances contained within these species include:meconic acid andchelidonic acid, as well ascyanogenic glycoside compounds derived fromtyrosine:dhurrin and triglochinin; in the Fumarioideae while the Chelidonieae contain the free amino acid δ-acetylornithine.
Many of these plants arepoisonous. TheMexican prickly poppy is poisonous if taken internally and may causeoedema andglaucoma. Even if an animal, such as agoat, should persist in grazing on this plant, not only will the animal suffer but so will those who drink itsmilk, because the poisons are passed along in the milk.
The fossils of the lateCretaceous poppyPalaeoaster inquirenda from the Western Interior of North America occurs from 74.5 million year old deposits in theFruitland Formation inNew Mexico to 64.5 million year old deposits in theHell Creek Formation inNorth Dakota. Dehiscent fruit fossils ofPalaeoaster have been found at the excavation site for the well knownTyrannosaurus rex specimen BHI 3033. The seed capsule ofPalaeoaster has some similarities to that of the extant poppy genusRomneya.[14]
Papaverites, a fossil fruit from theEocene ofGermany, may be associated with Papaveraceae.[15] Chesters et al. (1967) mentionsPapaver pictum from theOligocene ofEngland.[16]
The family is well known for its striking flowers, with many species grown asornamental plants, includingCalifornia poppy (Eschscholtzia californica, the Californiastate flower), the stunning blueHimalayan poppies (Meconopsis), several species ofPapaver, and the wildflowerbloodroot. Only two species are of economic importance for the production ofopium and its derivatives for pharmaceutical use:Papaver somniferum is cultivated legally in order to obtainmorphine and otheropiates, andPapaver bracteatum, forthebaine.Papaver somniferum is also the source of thepoppy seeds used incooking andbaking, andpoppy seed oil. The illegal cultivation of poppies in Asia for the production of opium andheroin is virtually equal to the legal production in the rest of the world. Some Funarioideae have a limited use in gardening, withLamprocapnos spectabilis ("bleeding heart"), andPseudofumaria lutea ("yellow corydalis") commonly used. Chinese traditional medicine used the boiled and dried tubers ofCorydalis yanhusuo ("yanhusuo").
The opium poppy and corn poppy are symbols, respectively, of sleep and death. In Great Britain, Canada, the United States, and Australia the corn poppy is worn in remembrance ofWorld War I.
Blattner, Frank R. & Kadereit, Joachim W. (1999). "Morphological evolution and ecological diversification of the forest-dwelling poppies (Papaveraceae: Chelidonioideae) as deduced from a molecular phylogeny of the ITS region".Plant Systematics and Evolution.219 (3–4):181–197.Bibcode:1999PSyEv.219..181B.doi:10.1007/bf00985578.S2CID28548382.
Hoot, S.B.; Kadereit, J.W.; Blattner, F.R.; Jork, K.B.; Schwarzbach, A.E. & Crane, P.R. (1997). "Data congruence and phylogeny of the Papaveraceae s.l. based on four data sets:atpB andrbcL sequences,trnK restriction sites, and morphological characters".Systematic Botany.22 (3):575–590.doi:10.2307/2419829.JSTOR2419829.
Lidén, M.; Fukuhara, T.; Rylander, J. & Oxelman, B. (1997). "Phylogeny and classification of Fumariaceae, with emphasis onDicentra s.l., based on the plastid generps16 intron".Plant Systematics and Evolution.206 (1–4):411–420.Bibcode:1997PSyEv.206..411L.doi:10.1007/bf00987960.S2CID24285424.
Wang, W.; Lu, A.-M.; Ren, Y.; Endress, M.E. & Chen, Z.-D. (2009). "Phylogeny and classification of Ranunculales: Evidence from four molecular loci and morphological data".Perspectives in Plant Ecology, Evolution and Systematics.11 (2):81–110.Bibcode:2009PPEES..11...81W.doi:10.1016/j.ppees.2009.01.001.
Labanca, Fabiana; Ovesnà, Jaroslava; Milella, Luigi (August 2018). "Papaver somniferum L. taxonomy, uses and new insight in poppy alkaloid pathways".Phytochemistry Reviews.17 (4):853–871.Bibcode:2018PChRv..17..853L.doi:10.1007/s11101-018-9563-3.
Nemat, O (2023). "MAKING THE POPPY BAN STICK".World Today.79 (4):19–22.
Labanca, Fabiana; Ovesnà, Jaroslava; Milella, Luigi (August 2018). "Papaver somniferum L. taxonomy, uses and new insight in poppy alkaloid pathways".Phytochemistry Reviews.17 (4):853–871.Bibcode:2018PChRv..17..853L.doi:10.1007/s11101-018-9563-3.
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