Papaveraceae

Papaveraceae
Corn poppy (Papaver rhoeas)
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Eudicots
Order:	Ranunculales
Family:	Papaveraceae Juss.[1]
Genera
See text

The Papaveraceae

The simple leaves are alternate or sometimes whorled. They have petioles and are not enclosed by a sheath. The leaves are usually lobed or pinnatifid (i.e. consisting of several not entirely separate leaflets), or much divided. There are no stipules.

The plants are

The non-fleshy fruit is usually a capsule, breaking open at maturity to release the seeds 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 of Platystemon is a schizocarp.

The basic chromosome number, x, is 6, 7, 8, 9, 10 and 11, up to 12n = 84 (dodecaploidy) in species of Papaver, Argemone and Meconopsis.

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. Papaver somniferum is also found to have anticancer properties mainly against cervical and colon cancers as it was found to disrupt cell membrane of tumor cells. The opium poppy (Papaver somniferum) shares a common ancestor with Macleya cordata, and Eschschloizia californica. Papaver somniferum evolved from Aquilegia coerulea then Nelumbo nucifera and Macadamia intergrifolia. The furthest known ancestor to Papaver somniferum is Oryza sativa.

Taxonomy

The APG III system (2009; unchanged from the APG II system of 2003 and the APG system of 1998) places the family in the order Ranunculales, in the clade eudicots.^[1] The Papaveraceae differ from the rest of the Ranunculales in some important characteristics but they share others such as the presence of isoquinoline-derived alkaloids. Based on molecular and morphological data, the family forms a clade with the families Lardizabalaceae, Circaeasteraceae, Menispermaceae, Berberidaceae and Ranunculaceae.^[6]

Genera

The broad circumscription of Papaveraceae in the APG III system includes three taxa that have previously been separated into different families: the Papaveraceae sensu stricto, the Fumariaceae and the Pteridophyllaceae.^[1] Thus the Cronquist system of 1981 recognised the Fumariaceae as a separate family, despite their close phylogenetic relationship to the Papaveraceae sensu 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 of Pteridophyllum.^[8][6] The subtribes are given by the Germplasm 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]

Fumarioideae

• Subfamily Fumarioideae Eaton

• Tribe Hypecoeae Dumort.

• Hypecoum L. – Mediterranean region to Mongolia and Western China.
• Pteridophyllum Siebold & Zucc. – Japan

• Tribe Fumarieae Dumort.

• Subtribe Corydalinae

• Adlumia Raf. ex DC. – Eastern North America, Korea, China
• Capnoides Mill. – Northern North America
• Corydalis DC. nom. cons. – Eurasia, North America, East Africa
• Dactylicapnos Wall. – Himalayas
• Dicentra Bernh. nom. cons. – Eastern Asia, North America
• Ehrendorferia Fukuhara & Lidén – Western United States
• Ichtyoselmis Lidén & Fukuhara – China
• Lamprocapnos Endl. – China, Korea

• Subtribe Fumariinae

• Ceratocapnos Durieu – South west of Europe, north west of Africa
• Cryptocapnos Rech.f. – Central Afghanistan
• Cysticapnos Mill. – South Africa
• Discocapnos Cham. & Schltdl. – South Africa
• Fumaria L. – Mediterranean region, Himalayas, East Africa
• Fumariola Korsh. – Central Asia
• Platycapnos (DC.) Bernh. – Western Mediterranean region
• Pseudofumaria Medik. – Italy, Balkans
• Rupicapnos Pomel – North west Africa
• Sarcocapnos DC. – Spain, Morocco, Algeria
• Trigonocapnos Schltr. – South Africa

• Photos of Fumarioideae (the fumewort subfamily)
• 
  Hypecoum procumbens
• 
  Scrambled eggs (Corydalis aurea)
• 
  Fumewort (Corydalis solida)
• 
  Ferny corydalis (Corydalis cheilanthifolia)
• 
  Pale corydalis (Capnoides sempervirens)
• 
  Climbing bleeding-heart (Dactylicapnos scandens or Dicentra scandens)
• 
  Dutchman's breeches (Dicentra cucullaria)
• 
  Fringed bleeding-heart (Dicentra eximia)
• 
  Komakusa (Dicentra peregrina)
• 
  Golden eardrops (Ehrendorferia chrysantha or Dicentra chrysantha)
• 
  Asian bleeding-heart (Lamprocapnos spectabilis or Dicentra spectabilis)
• 
  Earth smoke (Fumaria officinalis)
• 
  Yellow corydalis (Pseudofumaria lutea or Corydalis lutea)
• 
  Zapatitos de la Virgen (Sarcocapnos enneaphylla)

Papaveroideae

• Subfamily Papaveroideae Eaton

• Tribe Eschscholzieae Baill.

• Dendromecon Benth. – California.
• Eschscholzia Cham. – Western North America.
• Hunnemannia Sweet – Eastern Mexico.

• Tribe Chelidonieae Dumort.

• Bocconia L. – Central and southern America, Antilles
• Chelidonium L. – Eurasia
• Coreanomecon Nakai – Korea
• Dicranostigma Hook.f. & Thomson – Central Asia
• Eomecon Hance – Eastern China
• Glaucium Mill. – Europe to Central Asia
• Hylomecon Maxim. – Eastern Asia
• Macleaya R.Br. – Eastern Asia
• Sanguinaria L. – Eastern North America
• Stylophorum Nutt. – Eastern North America, Eastern Asia

• Tribe Platystemoneae Spach

• Hesperomecon Greene – Western North America
• Meconella Nutt. – Western North America
• Platystemon Benth. – Western North America

• Tribe Papavereae Dumort.

• Arctomecon Torr. & Frém. – Western North America
• Argemone L. – North America, Antilles, central and southern America, Hawaii
• Canbya Parry – Western North America
• Cathcartia Hook.f. – China and the Himalayas, split from Meconopsis
• Meconopsis Vig. – Central southern Asia, western Europe; paraphyletic^[13]
• Papaver L. – Northern hemisphere, South Africa, Cape Verde; paraphyletic^[13]
• Roemeria Medik. – Mediterranean region, south west Asia
• Romneya Harv. – California
• Stylomecon G. Taylor
  – California

• Photos of Papaveroideae (the poppy subfamily)
• 
  Channel Island tree poppy (Dendromecon harfordii)
• 
  Tulip poppy (Hunnemannia fumariifolia)
• 
  Plume poppy (Bocconia frutescens)
• 
  Greater celandine (Chelidonium majus)
• 
  Eastern horned poppy (Dicranostigma erectum)
• 
  Red horned poppy (Glaucium corniculatum)
• 
  Snow-poppy (Eomecon chionantha)
• 
  Forest poppy (Hylomecon vernalis)
• 
  Bloodroot (Sanguinaria canadensis)
• 
  Wood poppy (Stylophorum diphyllum)
• 
  Creamcups (Platystemon californicus)
• 
  Bearpaw poppy (Arctomecon californica)
• 
  Flat-bud prickly poppy (Arctomecon munita)
• 
  Pygmy poppy (Canbya candida)
• 
  Arctic poppy (Papaver nudicaule)
• 
  Persian poppy (Papaver bracteatum)
• 
  Opium poppy (Papaver somniferum)
• 
  Matilija poppy (Romneya coulteri)

Ecology

Pollination is

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 are ruderal 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.

Phytochemistry

Alkaloids: The isoquinolinic alkaloids present in the family are well known. They are derived from berberine, 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 of aporphine, morphinan, pavine, isopavine, narceine and rhoeadine.

Others: Other characteristic substances contained within these species include: meconic acid and chelidonic acid, as well as cyanogenic glycoside compounds derived from tyrosine: dhurrin and triglochinin; in the Fumarioideae while the Chelidonieae contain the free amino acid δ-acetylornithine.

Papaverites, a fossil fruit from the Eocene of Germany, may be associated with Papaveraceae.^[15] Chesters et al. (1967) mentions Papaver pictum from the Oligocene of England.^[16]

Cultivation

California poppy (Eschscholzia californica)

Himalayan blue poppy (Meconopsis betonicifolia

)

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 of World War I.

References

Bibliography

• Angiosperm Phylogeny Group (2009). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III". Botanical Journal of the Linnean Society. 161 (2): 105–121.
  hdl:10654/18083
  .
• 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.
  S2CID 28548382
  .
• Carolan, James C.; Hook, Ingrid L.I.; Chase, Mark W.; Kadereit, Joachim W. & Hodkinson, Trevor R. (2006). "Phylogenetics of Papaver and Related Genera Based on DNA Sequences from ITS Nuclear Ribosomal DNA and Plastid trnL Intron and trnL–F Intergenic Spacers". Annals of Botany. 98 (1): 141–155.
  PMID 16675606
  .
• doi:10.11646/phytotaxa.261.3.1
  .
• Friedel, Jean (1927). "Filiation des Papavéracées". Bulletin de la Société Botanique de France (in French). 74 (4): 673–688.
  doi:10.1080/00378941.1927.10833678
  .
• 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 and rbcL sequences, trnK restriction sites, and morphological characters". Systematic Botany. 22 (3): 575–590.
  JSTOR 2419829
  .
• Kadereit, J.W. (1993). "Papaveraceae". In Kubitzki, Rohwer & Bittrich (1993)
• Kadereit, J.W.; Schwarzbach, A.E.; Jork, K.B. (1997). "The phylogeny of Papaver s.l. (Papaveraceae): polyphyly or monophyly?". Plant Systematics and Evolution. 204 (1–2): 75–98.
  S2CID 20019969
  .
• Kubitzki, K.; Rohwer, J.G.; Bittrich, V., eds. (1993). The Families and Genera of Vascular Plants. II. Flowering Plants - Dicotyledons. Berlin: Springer-Verlag.
  ISBN 978-3-540-55509-4
  .
• Lidén, M. (1993a). "Fumariaceae". In Kubitzki, Rohwer & Bittrich (1993)
• Lidén, M. (1993b). "Pteridophyllaceae". In Kubitzki, Rohwer & Bittrich (1993)
• Lidén, M.; Fukuhara, T.; Rylander, J. & Oxelman, B. (1997). "Phylogeny and classification of Fumariaceae, with emphasis on Dicentra s.l., based on the plastid gene rps16 intron". Plant Systematics and Evolution. 206 (1–4): 411–420.
  S2CID 24285424
  .
• Smith, Una R. (2001). "Revision of the Cretaceous Fossil Genus Palaeoaster (Papaveraceae) and Clarification of Pertinent Species of Eriocaulon, Palaeoaster and Sterculiocarpus". Novon. 11 (2): 258–260.
  JSTOR 3393064
  .
• ISBN 9780521707725
  .
• 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.
  doi:10.1016/j.ppees.2009.01.001
  .
• Watson, L. & Dallwitz, M.J. (1992–2006). "The families of flowering plants: descriptions, illustrations, identification, and information retrieval. Version: 29th July 2006". Archived from the original on 3 January 2007. Retrieved 27 December 2006.
• Willemstein, S. C. (1987). An Evolutionary Basis for Pollination Ecology. E. J. Brill / Leiden University Press Leiden.
  ISBN 90-04-08457-6
  . (pbk.)

Colombo, M. L., & Bosisio, E. (1996). PHARMACOLOGICAL ACTIVITIES OFCHELIDONIUM MAJUSL. (PAPAVERACEAE). Pharmacological Research, 33(2), 127–134. https://doi.org/10.1006/phrs.1996.0019

Erdtman (1952, 1969), Faegri and Iversen (1975), Henderson (1965), Kuprianova and Alyoshina (1972, 1978), PAPAVERACEAE https://www.sciencedirect.com/sdfe/pdf/download/eid/1-s2.0-0034666779900344/first-page-pdf

Güler, D. A., Aydın, A., Koyuncu, M., Parmaksız, İ., & Tekin, Ş. (2016). Anticancer Activity of Papaver Somniferum. Journal of the Turkish Chemical Society, Section A: Chemistry, 3(3). https://doi.org/10.18596/jotcsa.43273

Labanca, F., Ovesnà, J., & Milella, L. (2018). Papaver somniferum L. taxonomy, uses and new insight in poppy alkaloid pathways. Phytochemistry Reviews, 17(4), 853–871. https://doi.org/10.1007/s11101-018-9563-3

Li, Y., Winzer, T., He, Z., & Graham, I. A. (2020). Over 100 Million Years of Enzyme Evolution Underpinning the Production of Morphine in the Papaveraceae Family of Flowering Plants. Plant Communications, 1(2), 100029. https://doi.org/10.1016/j.xplc.2020.100029

Native Plants (PAPAVERACEAE, THE POPPY FAMILY, n.d.,) https://nativeplants.org/wp-content/uploads/papaveraceae.pdf

Nemat, O. (2023). MAKING THE POPPY BAN STICK. World Today, 79(4), 19–22.

Labanca, F., Ovesnà, J., & Milella, L. (2018). Papaver somniferum L. taxonomy, uses and new insight in poppy alkaloid pathways. Phytochemistry Reviews, 17(4), 853–871. https://doi.org/10.1007/s11101-018-9563-3

External links

Wikimedia Commons has media related to Papaveraceae.

• Papaveraceae in Topwalks
• Papaveraceae Archived 20 March 2006 at the Wayback Machine [sensu stricto] in L. Watson and M.J. Dallwitz (1992 onwards). The families of flowering plants Archived 3 January 2007 at the Wayback Machine.
• Papaveraceae [sensu lato] in the Flora of North America
• links at CSDL
• Family Papaveraceae Archived 11 October 2012 at the Wayback Machine Flowers in Israel
• Mapa de Pteridophylloideae
• Mapa de Papaveroideae
• Mapa de Fumarioideae