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''Mandragora'' is a
plant Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they are predominantly Photosynthesis, photosynthetic. This means that they obtain their energy from sunlight, using chloroplasts derived from endosymbiosis with c ...
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
belonging to the nightshade family (
Solanaceae Solanaceae (), commonly known as the nightshades, is a family of flowering plants in the order Solanales. It contains approximately 2,700 species, several of which are used as agricultural crops, medicinal plants, and ornamental plants. Many me ...
). Members of the genus are known as
mandrake A mandrake is the root of a plant, historically derived either from plants of the genus '' Mandragora'' (in the family Solanaceae) found in the Mediterranean region, or from other species, such as '' Bryonia alba'' (the English mandrake, in the ...
s. Between three and five
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
are placed in the genus. The one or two species found around the
Mediterranean The Mediterranean Sea ( ) is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean basin and almost completely enclosed by land: on the east by the Levant in West Asia, on the north by Anatolia in West Asia and Southern ...
constitute the mandrake of ancient writers such as
Dioscorides Pedanius Dioscorides (, ; 40–90 AD), "the father of pharmacognosy", was a Greek physician, pharmacologist, botanist, and author of (in the original , , both meaning "On Materia medica, Medical Material") , a 5-volume Greek encyclopedic phar ...
. Two or three further species are found eastwards into China. All are
perennial In horticulture, the term perennial ('' per-'' + '' -ennial'', "through the year") is used to differentiate a plant from shorter-lived annuals and biennials. It has thus been defined as a plant that lives more than 2 years. The term is also ...
herbaceous plant Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials. Definitions of "herb" and "herbaceous" The fourth edition o ...
s, with large tap roots and leaves in the form of a rosette. Individual flowers are bell-shaped, whitish through to violet, and followed by yellow or orange
berries A berry is a small, pulpy, and often edible fruit. Typically, berries are juicy, rounded, brightly colored, sweet, sour or tart, and do not have a stone fruit, stone or pit (fruit), pit although many wikt:pip#Etymology 2, pips or seeds may be p ...
. Like many members of the Solanaceae, species of ''Mandragora'' contain highly biologically active
alkaloid Alkaloids are a broad class of natural product, naturally occurring organic compounds that contain at least one nitrogen atom. Some synthetic compounds of similar structure may also be termed alkaloids. Alkaloids are produced by a large varie ...
s that make the plants poisonous. Their roots in particular have a long use in
traditional medicine Traditional medicine (also known as indigenous medicine or folk medicine) refers to the knowledge, skills, and practices rooted in the cultural beliefs of various societies, especially Indigenous groups, used for maintaining health and treatin ...
. Mandrakes are involved in many myths and superstitions.


Description

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
sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 Etymology The term ''sepalum'' ...
s are joined at the base, as are the
petal Petals are modified leaves that form an inner whorl surrounding the reproductive parts of flowers. They are often brightly coloured or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''corol ...
s, both in the shape of a lobed bell. The
stamen The stamen (: stamina or stamens) is a part consisting of the male reproductive organs of a flower. Collectively, the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filament ...
s are shorter than the petals, joined to the floral tube towards the base. The
ovary The ovary () is a gonad in the female reproductive system that produces ova; when released, an ovum travels through the fallopian tube/ oviduct into the uterus. There is an ovary on the left and the right side of the body. The ovaries are end ...
has two chambers (
locule A locule (: locules) or loculus (; : loculi) is a small cavity or compartment within an organ or part of an organism (animal, plant, or fungus). In angiosperms (flowering plants), the term ''locule'' usually refers to a chamber within an ovary ...
s). After fertilization, a yellow or orange fruit forms ( botanically a berry).


Taxonomy

The genus ''Mandragora'' was first used in 1753 by
Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
in the first edition of ''
Species Plantarum ' (Latin for "The Species of Plants") is a book by Carl Linnaeus, originally published in 1753, which lists every species of plant known at the time, classified into genus, genera. It is the first work to consistently apply binomial nomenclature ...
'' where the Mediterranean species '' Mandragora officinarum'' was described, which is thus the
type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...
of the genus. (Linnaeus later changed his mind and in 1759 placed ''M. officinarum'' in the genus '' Atropa'' as ''A. mandragora''.) Jackson and Berry (1979) and Ungricht et al. (1998) 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, a distinction used in 1764 when Garsault published the names ''Mandragora mas'' and ''Mandragora foemina''. 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 Antonio Bertoloni (February 8, 1775 in Sarzana – April 17, 1869 in Bologna) was an Italian physician and botanist who made extensive studies of Italian plants. He also collected notable samples of Central American flora. Biography He studied m ...
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 The ''Flora Europaea'' is a 5-volume encyclopedia of plants, published between 1964 and 1993 by Cambridge University Press. The aim was to describe all the national Floras of Europe in a single, authoritative publication to help readers identify ...
'' listed two Mediterranean species of ''Mandragora'': ''M. officinarum'' and ''M. autumnalis''. 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. Other sources divide ''M. officinarum'' ''
sensu lato ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...
'' 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''.


Classification and phylogeny

Traditionally, ''Mandragora'' has been considered to be closely related to '' Atropa'' and '' Lycium'', being grouped together in the same tribe or subtribe as at least the first of these genera. Molecular phylogenetic studies suggest that the genus belongs in the large subfamily Solanoideae, but that within this subfamily, it is one of a number of isolated genera with no immediate relatives. It has thus been placed in its own tribe, Mandragoreae. Within the genus, studies have used different circumscriptions of the Mediterranean mandrakes. Two studies that separate plants found in the
Levant The Levant ( ) is the subregion that borders the Eastern Mediterranean, Eastern Mediterranean sea to the west, and forms the core of West Asia and the political term, Middle East, ''Middle East''. In its narrowest sense, which is in use toda ...
('' Mandragora autumnalis'') from those found in the rest of the Mediterranean area ('' Mandragora officinarum'') suggest that there are two clades in the genus - one based in the Mediterranean and beyond to Turkmenistan and Iran, and one in the Sino-Himalayan region. A simplified cladogram based on these studies is shown below. In one of the studies, ''M. chinghaiensis'' was embedded within ''M. caulescens''. The Solanaceae are primarily a
New World The term "New World" is used to describe the majority of lands of Earth's Western Hemisphere, particularly the Americas, and sometimes Oceania."America." ''The Oxford Companion to the English Language'' (). McArthur, Tom, ed., 1992. New York: ...
family. ''Mandragora'' is suggested to have originated around 20 million years ago, arriving in
Eurasia Eurasia ( , ) is a continental area on Earth, comprising all of Europe and Asia. According to some geographers, Physical geography, physiographically, Eurasia is a single supercontinent. The concept of Europe and Asia as distinct continents d ...
through the agency of birds, with the main split between the species occurring around 10 million years ago.


Species

, major online plant databases (such as
Tropicos Tropicos is an online botanical database containing taxonomic information on plants, mainly from the Neotropical realm ( Central, and South America South America is a continent entirely in the Western Hemisphere and mostly in the Sou ...
,
The Plant List The Plant List was a list of botanical names of species of plants created by the Royal Botanic Gardens, Kew and the Missouri Botanical Garden and launched in 2010. It was intended to be a comprehensive record of all known names of plant specie ...
, and GRIN Taxonomy for Plants) accept different numbers of species in the genus ''Mandragora''. Three species are accepted in a 1998 review of the genus and by GRIN. Other sources keep ''M. autumnalis'' and ''M. chinghaiensis'' as separate species. *''Mandragora officinarum'' L. and '' Mandragora autumnalis'' Bertol.
Central and southern
Portugal Portugal, officially the Portuguese Republic, is a country on the Iberian Peninsula in Southwestern Europe. Featuring Cabo da Roca, the westernmost point in continental Europe, Portugal borders Spain to its north and east, with which it share ...
and throughout the
Mediterranean The Mediterranean Sea ( ) is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean basin and almost completely enclosed by land: on the east by the Levant in West Asia, on the north by Anatolia in West Asia and Southern ...
area, eastwards to
Syria Syria, officially the Syrian Arab Republic, is a country in West Asia located in the Eastern Mediterranean and the Levant. It borders the Mediterranean Sea to the west, Turkey to Syria–Turkey border, the north, Iraq to Iraq–Syria border, t ...
and
Jordan Jordan, officially the Hashemite Kingdom of Jordan, is a country in the Southern Levant region of West Asia. Jordan is bordered by Syria to the north, Iraq to the east, Saudi Arabia to the south, and Israel and the occupied Palestinian ter ...
. Virtually stemless; petals long, greenish white through blue to violet; berry globose to ovoid, yellow to orange when ripe.
''M. autumnalis'' may be included within ''M. officinarum'' or considered a separate species. Older sources consider ''M. autumnalis'' to the main species found in the Mediterranean with ''M. officinarum'' confined to northern Italy and parts of the coast of former Yugoslavia. Some more recent sources distinguish plants found in the
Levant The Levant ( ) is the subregion that borders the Eastern Mediterranean, Eastern Mediterranean sea to the west, and forms the core of West Asia and the political term, Middle East, ''Middle East''. In its narrowest sense, which is in use toda ...
as ''Mandragora autumnalis'', one difference being that the seeds are more than twice as large as those of ''M. officinarum''. *'' Mandragora turcomanica'' Mizg.
Turkmenistan Turkmenistan is a landlocked country in Central Asia bordered by Kazakhstan to the northwest, Uzbekistan to the north, east and northeast, Afghanistan to the southeast, Iran to the south and southwest and the Caspian Sea to the west. Ash ...
,
Iran Iran, officially the Islamic Republic of Iran (IRI) and also known as Persia, is a country in West Asia. It borders Iraq to the west, Turkey, Azerbaijan, and Armenia to the northwest, the Caspian Sea to the north, Turkmenistan to the nort ...
( Golestan Province). Stemless; petals long, violet; berry yellow, strongly aromatic. *'' Mandragora caulescens'' C.B.Clarke (including ''Mandragora chinghaiensis'' Kuang & A.M.Lu, ''Mandragora tibetica'' Grubov)
India India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...
,
Nepal Nepal, officially the Federal Democratic Republic of Nepal, is a landlocked country in South Asia. It is mainly situated in the Himalayas, but also includes parts of the Indo-Gangetic Plain. It borders the Tibet Autonomous Region of China Ch ...
,
Bhutan Bhutan, officially the Kingdom of Bhutan, is a landlocked country in South Asia, in the Eastern Himalayas between China to the north and northwest and India to the south and southeast. With a population of over 727,145 and a territory of , ...
and parts of China (south-east
Qinghai Qinghai is an inland Provinces of China, province in Northwestern China. It is the largest provinces of China, province of China (excluding autonomous regions) by area and has the third smallest population. Its capital and largest city is Xin ...
, west
Sichuan Sichuan is a province in Southwestern China, occupying the Sichuan Basin and Tibetan Plateau—between the Jinsha River to the west, the Daba Mountains to the north, and the Yunnan–Guizhou Plateau to the south. Its capital city is Cheng ...
, east Xizang (Tibet), north-west
Yunnan Yunnan; is an inland Provinces of China, province in Southwestern China. The province spans approximately and has a population of 47.2 million (as of 2020). The capital of the province is Kunming. The province borders the Chinese provinces ...
). Stems sometimes present; petals dark purple or yellow; berry globose. Considerably variable in size and appearance, possibly justifying dividing the taxon into subspecies or even species.


Toxicity

Like many members of the Solanaceae, species of ''Mandragora'' contain highly biologically active alkaloids,
tropane alkaloid Tropane alkaloids are a class of bicyclic .2.1alkaloids and secondary metabolites that contain a tropane ring in their chemical structure. Tropane alkaloids occur naturally in many members of the plant family Solanaceae. Certain tropane alkaloi ...
s in particular. Hanuš et al. reviewed the
phytochemistry Phytochemistry is the study of phytochemicals, which are chemicals derived from plants. Phytochemists strive to describe the structures of the large number of secondary metabolites found in plants, the functions of these compounds in human and ...
of ''Mandragora'' species. More than 80 substances have been identified; their paper gives the detailed chemical structure of 37 of them. The different parts of the plant contain different proportions and concentrations of alkaloids, with the roots having the highest concentrations. Alkaloids present include
atropine Atropine is a tropane alkaloid and anticholinergic medication used to treat certain types of nerve agent and pesticide poisonings as well as some types of slow heart rate, and to decrease saliva production during surgery. It is typically give ...
, apoatropine, belladonnine, cuscohygrine,
hyoscyamine Hyoscyamine (also known as daturine or duboisine) is a naturally occurring tropane alkaloid and plant toxin. It is a secondary metabolite found in certain plants of the family Solanaceae, including Hyoscyamus niger, henbane, Mandragora officina ...
, scopolamine (hyoscine), 3α-tigloyloxytropane, and 3α,6β-ditigloyloxytropane. ''M. caulescens'' and ''M. turcomanica'' are also reported to contain anisodamine. Clinical reports of the effects of consumption of plants described as ''M. autumnalis'' (''M. offinarum'' ''
sensu lato ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...
'') include severe symptoms similar to those of atropine poisoning, including blurred vision, dilation of the pupils (
mydriasis Mydriasis is the Pupillary dilation, dilation of the pupil, usually having a non-physiological cause, or sometimes a physiological pupillary response. Non-physiological causes of mydriasis include disease, Physical trauma, trauma, or the use of c ...
), dryness of the mouth, difficulty in urinating, dizziness, headache, vomiting, blushing, and a rapid heart rate (
tachycardia Tachycardia, also called tachyarrhythmia, is a heart rate that exceeds the normal resting rate. In general, a resting heart rate over 100 beats per minute is accepted as tachycardia in adults. Heart rates above the resting rate may be normal ...
). Hyperactivity and hallucinations also occurred in the majority of patients.


Uses

''Mandragora'' species have a long use in traditional medicine, extracts being used for their real or supposed
aphrodisiac An aphrodisiac is a substance that increases libido, sexual desire, sexual attraction, sexual pleasure, or sexual behavior. These substances range from a variety of plants, spices, and foods to synthetic chemicals. Natural aphrodisiacs, such as ...
,
hypnotic A hypnotic (from Ancient Greek, Greek ''Hypnos'', sleep), also known as a somnifacient or soporific, and commonly known as sleeping pills, are a class of psychoactive drugs whose primary function is to sleep induction, induce sleep and to trea ...
,
emetic Vomiting (also known as emesis, puking and throwing up) is the forceful expulsion of the contents of one's stomach through the mouth and sometimes the nose. Vomiting can be the result of ailments like food poisoning, gastroenteritis, preg ...
, purgative,
sedative A sedative or tranquilliser is a substance that induces sedation by reducing irritability or Psychomotor agitation, excitement. They are central nervous system (CNS) Depressant, depressants and interact with brain activity, causing its decelera ...
, and pain-killing effects. Tropane alkaloids are known to be effective as
analgesic An analgesic drug, also called simply an analgesic, antalgic, pain reliever, or painkiller, is any member of the group of drugs used for pain management. Analgesics are conceptually distinct from anesthetics, which temporarily reduce, and in s ...
s and
anaesthetic An anesthetic (American English) or anaesthetic (British English; see spelling differences) is a drug used to induce anesthesia ⁠— ⁠in other words, to result in a temporary loss of sensation or awareness. They may be divided into t ...
s, and can be used to increase circulation and dilate pupils, among other effects. Hyoscine and anisodamine are used medicinally in China. Continued use of ''M. autumnalis'' in
folk medicine Traditional medicine (also known as indigenous medicine or folk medicine) refers to the knowledge, skills, and practices rooted in the cultural beliefs of various societies, especially Indigenous groups, used for maintaining health and treatin ...
was reported in
Sicily Sicily (Italian language, Italian and ), officially the Sicilian Region (), is an island in the central Mediterranean Sea, south of the Italian Peninsula in continental Europe and is one of the 20 regions of Italy, regions of Italy. With 4. ...
in 2014. ''M. caulescens'' (as ''M. chinghaiensis'') has been listed as a Chinese medicinal plant needing conservation. The presence of
deliriant Deliriants are a subclass of hallucinogen. The term was coined in the early 1980s to distinguish these drugs from psychedelics such as LSD and dissociatives such as ketamine, due to their primary effect of causing delirium, as opposed to th ...
and hallucinogenic alkaloids and the sometimes vaguely
humanoid A humanoid (; from English ''human'' and '' -oid'' "resembling") is a non-human entity with human form or characteristics. By the 20th century, the term came to describe fossils which were morphologically similar, but not identical, to those of ...
shape of their roots have led to mandrakes being associated with a variety of myths and superstitious practices throughout history. 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.


References

{{Authority control Solanoideae Solanaceae genera Oneirogens