Papaver somniferum L. taxonomy, uses and new insight in poppy alkaloid pathways


Since ancient times, opium poppy (Papaver somniferum L.) is known for its medicinal properties, related to its secondary metabolite content. Its most important secondary metabolites, called benzylisoquinoline alkaloids (BIAs), are still essential in pharmaceutical field. Few of them, like morphine, have specific clinical application but also effects on CNS. Not all poppy cultivars are able to biosynthesize morphine in high amount, making this plant useful for other purposes like food uses. For this reason it is crucial to deeply understand the origin of poppy, its possible use and have a deep knowledge of the BIA biosynthesis. These aspects are crucial for the final use of P. somniferum. This review aims to summarize the state-of-the-art on its taxonomy and origin beside its uses and BIA biosynthetic pathways, its most important metabolites. The review focuses on conflicting or unsolved questions about enzymatic localization, role of different plant organs in the biosynthesis, and storage and external conditions that influence the alkaloid production, highlighting the significant involvement of transcription factors. Behind this review, there is the firm belief that only a deep knowledge of alkaloid biosynthetic processes could lead to the characterization of undefined step and to the development of engineering cultivars optimizing the potential uses of P. somniferum. The goal is answer in more sustainable way to ever-increasing worldwide request of such products, in particular morphine and derivates, obtaining high morphine content cultivars useful for pharmaceutical market or no morphine producing cultivars appreciated as food. Devising cultivars with different BIA content could lead to decrease, or even avoid, illicit use and illegal extraction, confining only low alkaloid content cultivars to consumers market.

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Tyrosine/tyramine 3-hydroxylase


3′-hydroxyl-N-methylcoclaurine 4′-O-methyltransferase


4 Hydroxyphenylacetic acid


4-Hydroxyphenylpuruvate decarboxylase


Norcoclaurine 6-O-methyltransferase


Reticuline 7-O-methyltransferase


Adenylyl cyclase


Amplified fragment length polymorphism


Adenosine monophosphate


Adenosine monophosphate cyclic


Berberine bridge enzyme


Benzylisoquinoline alkaloids


Canadine synthase


Cheilanthifoline synthase


Coclaurine N-methyltransferase


Central nervous system


Codeine O-demethylase


Columbamine O-methyltransferase


Codeinone reductase


Community Plant Variety Office


Cytochrome P450 enzyme family


N-methylcanadine 1-hydroxylase


(S)-N-methylcoclaurine 3′-hydroxylase isozyme






Dihydrosanguinarine oxidase


European Food Safety Authority


Endoplasmic reticulum


Expressed sequence tag


Fourier-transform–ion-cyclotron resonance–mass spectrometry


Gas chromatography


G protein–coupled receptors


High performance liquid chromatography


Methyl jasmonate


Monoterpenoid indole alkaloid


Major latex proteins


N-methylstylopine 14-hydroxylase


Norreticuline 7-O-methyltransferase


Nucleus accumbens


Norcoclaurine synthase


N-methylcoclaurine 3′-hydroxylase


Noscapine synthase

Nuclear factor-κB:

Nuclear factor kappa-light-chain-enhancer of activated B cells

OR receptors:

Receptors for endogenous opiates


Opioid receptor-like


Protopine 6-hydroxylase


Protein kinase A


Protein kinase C




Random amplified polymorphic DNA


Restriction fragment length polymorphism


Reactive oxygen species


Restriction site polymorphism


Rostral ventromedial medulla


Salutaridinol 7-O-acetyltransferase


Salutaridine reductase


Sanguinarine reductase


(7S)-salutaridinol 7-O-acetyltransferase


Single nucleotide polymorphism


Scoulerine 9-O-methyltransferase




Stylopine synthase




Short tandem repeat


Salutaridine synthase


Sstylopine synthase


Thebaine 6-O-demethylase


Transcription factor binding bites


Transcription factors


Tetrahydroprotoberberine N-methyltransferase


Tyrosine/DOPA decarboxylase


Ultra high performance liquid chromatography–electrospray ionization–quadrupole time-of-flight–mass spectrometry


Variable number of tandem repeat or minisatellites


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Authors would like to thank the Czech Minstry of Agriculture, Projects NAZV QK1720263 and RO0417 for the support.

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Correspondence to Luigi Milella.

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Labanca, F., Ovesnà, J. & Milella, L. Papaver somniferum L. taxonomy, uses and new insight in poppy alkaloid pathways. Phytochem Rev 17, 853–871 (2018).

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  • Benzylisoquinoline alkaloid
  • BIAs
  • Biosynthesis
  • Poppy
  • Phylogenetic
  • Transcription factors