Mandragora (genus)

Species of Mandragora are perennial herbaceous plants. They have large vertical tap-roots, sometimes forked. Their stems are short or virtually absent. The leaves form a rosette at the base of the plant. The flowers are sometimes borne on a short stalk (scape), and are solitary, with whorls of five parts. The sepals are joined at the base, as are the petals, both in the shape of a lobed bell. The stamens are shorter than the petals, joined to the floral tube towards the base. The ovary has two chambers (locules). After fertilization, a yellow or orange fruit forms (botanically a berry).[1][2]

Mandragora officinarum with violet flowers

The genus Mandragora was first used in 1753 by Carl Linnaeus in the first edition of Species Plantarum where the Mediterranean species Mandragora officinarum was described,[3][4] which is thus the type species of the genus.[5] (Linnaeus later changed his mind and in 1759 placed M. officinarum in the genus Atropa as A. mandragora.[6])

Jackson and Berry (1979)[5] and Ungricht et al. (1998)[7] have documented the subsequent confusion over the number of Mediterranean species of Mandragora and their scientific names. Dioscorides was among those who distinguished between "male" and "female" mandrakes,[5] a distinction used in 1764 when Garsault published the names Mandragora mas and Mandragora foemina.[7] The size and shape of the fruit and the colour and time of appearance of the flowers have been used to distinguish possible species. In the 1820s, Antonio Bertoloni used flowering time to name two species as Mandragora vernalis, the spring-flowering mandrake, and Mandragora autumnalis, the autumn-flowering mandrake. Identifying the former as Linnaeus's M. officinarum, works such as Flora Europaea listed two Mediterranean species of Mandragora: M. officinarum and M. autumnalis.[2][5] Using statistical analysis of morphological characters, Ungricht et al. found no distinct clusters among the specimens they examined and concluded that Linnaeus's M. officinarum is a single, variable species.[7] Other sources divide M. officinarum sensu lato differently. Plants from the western Mediterranean, from Turkey westwards to the Iberian peninsula and Morocco, are placed in M. officinarum; plants from the eastern Mediterranean, from Syria to Israel, are placed in M. autumnalis.[8]

Atropine is a mixture of D and L isomers, the L isomer being hyoscyamine

Like many members of the Solanaceae, species of Mandragora contain highly biologically active alkaloids, tropane alkaloids in particular. Hanuš et al. reviewed the phytochemistry of Mandragora species. More than 80 substances have been identified; their paper gives the detailed chemical structure of 37 of them.[18] The different parts of the plant contain different proportions and concentrations of alkaloids, with the roots having the highest concentration. Alkaloids present include atropine, apoatropine, belladonnine, cuscohygrine, hyoscyamine, scopolamine (hyoscine), 3α-tigloyloxytropane and 3α,6β-ditigloyloxytropane.[5][7] M. caulescens and M. turcomanica are also reported to contain anisodamine.[17][19]

Clinical reports of the effects of consumption of plants described as Mandragora autumnalis (Mandragora offinarum s.l.) include severe symptoms similar to those of atropine poisoning, including blurred vision, dilation of the pupils (mydriasis), dryness of the mouth, difficulty in urinating, dizziness, headache, vomiting, blushing and a rapid heart rate (tachycardia). Hyperactivity and hallucinations also occurred in the majority of patients.[20][21]

Mandragora species have a long use in traditional medicine, extracts being used for their real or supposed aphrodisiac, hypnotic, emetic, purgative, sedative and pain-killing effects. Tropane alkaloids are known to be effective as analgesics and anaesthetics, and can be used to increase circulation and dilate pupils, among other effects.[7] Hyoscine and anisodamine are used medicinally in China.[17] Continued use of M. autumnalis in folk medicine was reported in Sicily in 2014.[22] M. caulescens (as M. chinghaiensis) has been listed as a Chinese medicinal plant needing conservation.[23]

The presence of deliriant and hallucinogenic alkaloids and the sometimes vaguely humanoid shape of their roots have led to mandrakes being associated with a variety of myths and superstitious practices throughout history.[7] However, the plants used in this way are not always species of Mandragora; for example, bryony (Bryonia) is explicitly mentioned as the source of a "mandrake" or "mandragora" in some sources.[24]