Bittersweet nightshade ( Solanum dulcamara ), illustration

Systematics
 Eudicotyledons Nuclear eudicotyledons Asterids Euasterids I Order : Nightshade (Solanales) Family : Nightshade family
Scientific name
Solanaceae
Yuss.

The nightshade family (Solanaceae) are a family of flowering plants (Magnoliopsida). About 90 to 100 genera belong to it , the number of associated species is given as about 2,700. The largest genus within the family is the nightshade ( Solanum ), which usually includes around 1,000 to 2,300 species. Within the family there are important food plants as well as ornamentals ; Due to the content of alkaloids and steroids , they are also considered to be important medicinal , intoxicating and cult plants . Characteristic features are above all the five-fold flowers with overgrown sepals , partly overgrown petals , five stamens and mostly two overgrown carpels . The fruits of the nightshade family are mostly berries or capsule fruits .

## description

### Habitus

Drawing of the black nightshade ( Solanum nigrum )
Planting of Virginia tobacco ( Nicotiana tabacum )

Nightshade plants are annual , biennial , perennial or perennial plants that can grow herbaceous and, more rarely, woody. Usually they reach heights of 0.5 to 4 m, but there are also representatives that reach up to 15 m in length as lianas or as small trees 5 to 10 m, in exceptional cases up to 25 m in height. There are also Pygmean representatives (e.g. Solanum euacanthum or Petunia patagonica ), which only reach heights of 5 to 20 cm. Nightshade plants usually grow upright, sometimes climbing, epiphytic or hemiepiphytic , often also myrmecophilous . There are some low lying representatives (especially the genera Lycianthes and Exodeconus ), rarely are like the mandrake ( Mandragora ) Rosette formation can be observed. The stem axis is usually massive, but sometimes also hollow, for example in the genera Markea , Poison Berries ( Nicandra ), Deprea or Witheringia . The structure of the shoots is often difficult to see through due to adhesions and shifts in the axes and leaves.

Nightshades form of different root types thick inter alia, fleshy taproots, for example, the Alraunen ( mandrake are known), root systems with adventitious roots (at the Leptoglossis as well as various types of jujubes ( Physalis ) and nightshade ( Solanum )), with extreme swellings (in the genus Lycianthes ) or tubers and stolons (especially in the Solanum -Sektion Petota ). Rhizomes also occur rarely , among others in the Salpichroa and Nectouxia .

SEM image of a tomato leaf ( Solanum lycopersicum ). The different trichome distribution on the top and bottom is particularly noticeable .

Many nightshade plants form hairs made up of trichomes, especially on the leaves and sprouts, and occasionally also on the flowers . Since this hairiness is very different, it serves as an important morphological feature for determination and classification. A common form are simple, glandular trichomes. As in the Solanum section Rhynchantherum , the Untertribus Nierembergiinae, these can have a single-celled head or a multicellular head - as is the case with various types of tobacco ( Nicotiana ). Branched trichomes can either branch like a tree or be occupied by whorl-like branches, the former is found in the genera Sessea and Juanulloa , the latter occurs in the Anthocercidoideae; Star -shaped, sea urchin-shaped or shield-shaped trichome heads can also be found in the Solanum sub -genus Brevantherum. Spines occur only in the Solanum subgenus Leptostemonum . With younger shoots of the Saracha , brownish, tree-like branching, multicellular emergences appear. Crystal sand is found mainly in the plants of the Solanoideae subfamily, including the Atropeae, Jaboroseae, Solaneae, Datureae, Lycieae and Hyoscyameae tribes .

### leaves

The alternate leaves are usually entire, often irregularly serrated or split. They are usually simple, occasionally also composed, then pinnate unpaired or tripartite, always without stipules . Occasionally thick and leathery leaves appear. The leaves are single, sometimes in whorls of three leaves or in clusters of three to six leaves. There are both sessile leaves and those with petioles.

### Inflorescences and flowers

Potato inflorescence ( Solanum tuberosum )
Night jasmine inflorescence ( Cestrum nocturnum )

Some of the flowers are single, but mostly in differently shaped inflorescences, sometimes with up to 200 flowers. The flowers or inflorescences are in the shoot armpits (axillary), outside the armpits (extra-axillary), the leaves opposite, terminal (this often translucent axis time groups that loose panicles or tight grape shaped) or many-flowered terminal panicles, sometimes also in clumps in groups. In the subgenus Lyciosolanum of the nightshade ( Solanum ), the flower stalks are missing, stem bloom is only known from the genus Dyssochroma .

The longest inflorescences are found in the genera Cuatresia (up to 25 cm) and Merinthopodium (up to 90 cm). In most cases the flowers of the nightshade family are hermaphroditic, only in exceptional cases there are dioecious plants, including at least one species each in Dunalia and Withania , two species in Symonanthus and four species each in the genera Deprea and the buckthorn ( Lycium ). In the genre of schizanthus ( Schizanthus ) there are also andromonoezische plants, which means they have both hermaphroditic and male flowers on one plant. The flowers are usually five-fold, rarely four or six to nine.

The flower formula is bis , deviations are mentioned below. ${\ displaystyle \ star}$${\ displaystyle \ downarrow K _ {(5)} \; {[C _ {(5)} \; A_ {5}]} \; \ G _ {\ underline {(2)}}}$

#### Chalice

The calyx is mostly radial symmetry, only rarely monosymmetric ( zygomorphic ) as in the angel's trumpets ( Brugmansia ). The sepals are fused with one another, the edge of the calyx has entire margins or five to ten straight teeth. In most cases the size of the calyx remains constant after the flowering phase, but an enlargement of the calyx is very often found in the family. Sometimes this enlargement is so great that the calyx enlarges around the berry or capsule until it is almost closed and almost completely encloses the fruit. This calyx enlargement occurs, for example, in the genera of the bladder cherries ( Physalis ) or Quincula . Another type of calyx enlargement occurs in the genera Chamaesaracha and Leucophysalis , here the calyx lies close to the pericarp of the fruit, but is usually open at the top. The calyx seldom bulges outwards and thus releases the ripe fruit, this occurs mainly in the genus Jaltomata .

#### Crown

Poison berry blossom ( Nicandra physalodes )
Aubergine blossom ( Solanum melongena )
Blossom of Capsicum baccatum
Angel's Trumpet Flowers ( Brugmansia sp. )
Flowers of a Juanulloa mexicana

The petals , like the sepals, are fused together. Sometimes they are very small with 2.5 to 8 mm, as in the genus Melananthus , but can also be 100 to 370 mm long, for example in the genus Solandra . The crowns are usually radial symmetry, only rarely zygomorphic, for example in the tribe Browallieae and in the genera Rahowardiana or Schultesianthus , sometimes the crown is even two-lipped ( schizanthus ). The main flower shapes are: wheel-shaped, star-shaped, tubular, funnel-shaped and over-cup-shaped.

#### Androeceum

The male flower part, called Androeceum , usually consists of five, only very rarely four ( Nothocestrum ) or two ( Schizanthus ) stamens . They stand in only one circle, are not grown together. They are arranged between the petals and are fused with them. Many representatives have stamens of different lengths in one flower, but stamens of the same length are similarly frequent. Some of the stamens protrude beyond the rest of the flower ( Vestia , Dunalia ), but they can also lie within the flower ( Lycianthes , Juanulloa etc.)

##### Anthers

The anthers ( anthers ) usually consist of two counters. There are both small ( Deprea , hammer bushes ( Cestrum ), Tribus Schwenckieae etc.) and large anthers (6 to 13 mm in Solandra or 12 to 40 mm in angel's trumpets ( Brugmansia )) with a length of 0.2 to 2 mm . They are usually straight, exceptions are the genus Normania and the species Solanum pennellii , which have curved anthers. The counters are generally the same size, with exceptions, such as Schwenckia , Melananthus , Heteranthia or Normania curved. Due to the stunting of one of the counters, the anthers of the tribe Browallieae are clearly asymmetrical. Mostly the anthers are hairless, in the genera hammer bushes ( Cestrum ) and Hawkesiophyton they are covered with papillae , in some genera there are representatives with - compared to the anthers - relatively large, simple trichomes on the anthers (Datureae, poison berries ( Nicandra ), Streptosolen , Solanum pennellii and in tomatoes ( Solanum lycopersicum )).

##### Stamens

The stamens are usually straight and cylindrical or slightly compressed. An exception is the genus Browallia with curved and flattened stamens. The stamens are similar in length or longer than the anthers. Deviating from this are the genera Nothocestrum with greatly reduced, almost non-existent stamens, Hawkesiophyton with very short stamens, Nectouxia with laminar enlarged stamens and Vestia with very long stamens. The length of the stamens of a flower is generally the same, but stamens of different lengths also occur, for example in Lycianthes , Capsicum campylopodium , Fabiana , Vestia and others. Didynamia (the occurrence of two different types of stamens within a flower) occurs among others in Anthocercis , Crenidium , Cyphanthera and Duboisia as well as in the subfamily Salpiglossideae. Sometimes the stamens or their upper part are bent towards the inside of the flower (among others in the Atropeae and Mandragoreae tribes and in various species of the Jaborosa genus ), are inclined at an angle ( Schultesianthus , Solandra ) or widened at the upper end (in the Jaboroseae tribe). Furthermore, stamens bent over to the knee appear both at the upper end (including in the poisonous berries ( Nicandra ), hammer bushes ( Cestrum ), petunias ( Petunia ), Fabiana , Sessea and Trianaea ) and at the lower end (Petunias ( Petunia ), Fabiana and Streptosolen ), and hook-shaped stamens (Jaboroseae).

##### Pollen

The pollen grains of the nightshade family come in very many different shapes, so that they can also be used as an important morphological feature for determination. The first point of differentiation is the size of the pollen grains - they can be small ( Latua , Hawkesiophyton , Fabiana , Tribus Lycieae and others), medium-sized ( Sessea , hammer bushes ( Cestrum ), Juanulloa , Rahowardiana and others) and also large ( Metternichia , Vestia , Merinthopodium , Weißbecher ( Nierembergia )) be. The absolute sizes range from approx. 20 µm for the small to approx. 70 µm for the large pollen grains. There are also major differences in the appearance of the pollen grain surface. The outer pollen wall (exine) can have Ubisch bodies (a layer of platelets on the pollen grain surface) ( Markea sessiliflora , Markea venosa , as well as various types of Schultesianthus ) or not ( Trianaea , Juanulloa , Dyssochroma , Solandra , Rahowardiana ), can be prickly ( Metternichia , Alraunen ( Mandragora )), wrinkled ( Sessea ), reticulated-wrinkled ( Merinthopodium ), grooved or smooth (hammer bushes ( Cestrum )), smooth ( Rahowardiana ), smooth or slightly grained ( Nothocestrum ), scaly ( Hawkesiophyton , Juanulloa ), doubled or grooved ( trumpet tongues ( Salpiglossis )), finely prickly ( Lycianthes ), papillary or warty ( Normania ) as well as reticulated ( Dyssochroma ).

#### Gynoeceum

In the flowers of most nightshade plants, the female flower part, the gynoeceum , consists of two fused carpels , which are mostly at an angle to the median plane of the flower. However, with the genus of poison berries ( Nicandra ) and two species of the genus Jaborosa and Trianaea there are also Gymnoeceen with three to five carpels, the species Iochroma umbellatum has four, the genus Nolana five carpels and the genus Melananthus probably only has one carpel. In some cultivated forms, for example the tomato ( Solanum lycopersicum ), larger numbers of carpels are also found.

##### Carpels

The overgrown carpels form an upper ovary , only the genera thorn apples ( Datura ), Solandra and Nothocestrum have ovarian nodes below. Except for small glandular ( Athenaea ) or rigid ( Browallia ) trichomes , and small fleshy thorns (thorn apples ( Datura )), the ovaries are bare. Normally the ovary has as many ovary compartments as carpels, with the exception of the Trianaea with eight to ten and the Solandra with four ovarian compartments. There are also ovaries in two genera, sometimes with four ovary compartments: in the Grabowskia in the upper half, in the Vassobia in parts of the lower half.

##### Nectaries

Normally, at the bottom of the ovary there are nectar glands , which are hidden , as in the Benthamiella and some goat's thorns ( Lycium ), or as in the white cups ( Nierembergia ) and the Untertribus Solaninae. If nectar glands are present, they are generally ring-shaped and slightly protruding, only in the poison berries ( Nicandra ) they are enclosed. However, there are also two variants of inversed nectar glands: pelvic-inversed, without lobes or incisions in the genus Schwenckia or two-lobed-everted with two lobes and two incisions, as can be found in the Untertribus Nicotianinae and the genera Bouchetia , Phrodus and some others . Another variant of the nectar glands is the thick, pillow-shaped Protoschwenckia .

##### Ovules

The ovules are on a rather fleshy placenta and can be turned upside down (e.g. in Metternichia and the white cups ( Nierembergia )), turned upside down to crooked ( Phrodus , Grabowskia and Vassobia ), half turned (hammer bushes ( Cestrum ) and others) or half crooked ( peppers ( Capsicum ), split flowers ( Schizanthus ), box thorns ( Lycium ) and others). Normally there are many ovules in each ovary compartment, but in some cases there are significantly fewer, for example in Grabowskia with two pairs of ovules in each ovary, or in the goat's thorns ( Lycium ) with only one ovule pair per ovary. The exception is the single ovule as it occurs in the genus Melananthus .

##### Stylus

The shape of the stylus is usually cylindrical, sometimes compressed ( Bouchetia ) or with two lateral outgrowths at the upper end (subfamily Salpiglossoideae and Subertribus Leptoglossinae), occasionally terminal ( Salpichroa , Saracha , Eriolarynx and others) or in an intermediate form between terminal and gymnobasic ( Vassobia , Jaborosa , Paprika ( Capsicum ), Dunalia and others). The stylus is usually massive, only occasionally hollow with a stylus channel (Trumpet tongues ( Salpiglossis ), Bouchetia and others), usually smooth or very wrinkled ( Browallia ). It is rarely connected to the ovary ( Withania , Triguera , Tubocapsicum ). Sometimes two different stylus lengths appear on the same plant ( Aureliana , Athenaea , Capsicum baccatum var. Umbilicatum and others). Exceptions are also Discopodium and Jaborosa ameghinoi , on whose stylus anthroric trichomes can be found.

##### scar

The stigma is usually disc-shaped and slightly head-shaped or head-shaped-lobed (poisonous berries ( Nicandra ), mandrake ( Mandragora )), rarely almost spherical or hemispherical as in the genus Paprika ( Capsicum ), sometimes saddle-shaped (Datureae, Juanulloeae, Nothocestrum and others). Exceptions are five-part scars such as Jaborosa odonelliana or bilobed scars ( Normania ). Within the family, there are very small and inconspicuous scars (split flowers ( schizanthus )), but also relatively large scars ( ectozoma ). They are usually glandular and moist, with the exception of the glandless stigmas of the schizanthus and Nierembergia linariaefolia and the dry unicellular glands of Solandra .

### fruit

Capsule fruits of the common thorn apple ( Datura stramonium )

The fruits are mostly multi-seeded berries or differently shaped split capsules (in the genus Markea , the subfamily Cestroideae and others), sometimes there are also intermediate forms between the two types of fruit. Exceptions are the lid capsules in the tribe Hyoscyameae, the stone fruits in some representatives of the tribe Lycieae, and the collective fruits of the genus Nolana .

Although the capsule is the more original form of fruit in evolutionary terms, today they only occur in a few basal clades and in the genus of thorn apples ( datura ). According to phylogenetic studies, the predominant berries in the family have three different sources, which indicates either a monophyletic clade in which the property of berry formation has been lost three times, or a three-fold development of the same property.

Often the fruits are not very large, sometimes less than 1 cm in diameter, but cultivated varieties of tomatoes and aubergines , for example, can have fruits that weigh several kilograms. But even with wild-growing representatives very large fruits can occur, for example the fruit of Solanum lycocarpon has a diameter of up to 15 cm.

In berry-like fruits, the pericarp and placenta can enlarge to such an extent that they merge completely, for example in the genera Tubocapsicum , Acnistus or Iochroma , as well as in tomatoes ( Solanum lycopersicum ) and other species of nightshade ( Solanum ). However, it also often happens that the two structures do not merge, so that a cavity is created inside the fruit, for example in peppers ( Capsicum ), Schultesieanthus , Lycianthes rantonnei and others.

The pericarp may be thick and juicy and contain stone cells ( witheringia , acnistus , some bladder cherries ( physalis )) or not (most bladder cherries ( physalis ), jaltomata , tomatoes ( Solanum lycopersicum )). However, it can also be made thin and without stone cells, as in the genera Chamaesaracha and Quincula, or only have small stone cells, as in the genus Darcyanthus . In exceptional cases, the pericarp is fragile and breaks easily into irregular pieces, for example in the case of quincula and chamaesaracha .

The capsule fruits open either by breaking the septum with two or four flaps ( thorn apples ( Datura )), cracks in the septum ( petunias ( Petunia ), Fabiana ) or septum-like gaps in the septum (in the subfamily Salpiglossoideae and the genera Metternichia , tobacco ( Nicotiana )) and the subfamily Anthocercidoideae).

The number of seeds per fruit fluctuates greatly: while in the tobacco genus ( Nicotiana ) up to 5000 seeds can be found, in the petunias ( Petunia ) it is up to 1200, in the Fabiana around 30 to 50 and in the Metternichia four to five. In the Melananthus only a seed can be found per fruit.

### Seeds

Schematic representation of a nightshade seed
Black nightshade seeds ( Solanum nigrum )
Night Jasmine seeds ( Cestrum nocturnum )

The size of the seeds is between 0.75 mm ( Darcyanthus ) or 0.6 to 1 mm ( Schwenckia , in Schwenckia micrantha only 0.3 to 0.4 mm) and 7 to 8 mm ( Jasminosolanum ) large seeds. The shape is more or less compressed, disc or kidney-shaped (in the subfamily Solanoideae and in the genera Combera , split flowers ( Schizanthus ), trumpet tongues ( Salpiglossis )), lenticular to kidney-shaped (in the nightshade - ( Solanum ) subgenus Leptostemonum ), boomerang-shaped or bacilliform (many members of the subfamily Juanulloideae), elongated and thin ( Trianaea ) or relatively thick and not compressed in various shapes (within the subfamily Cestroideae).

#### Seed coat

The seed coats appear in the most varied of variants: In the genera Sessea and Oryctes , a thin, peripheral, woody wing is a unique feature; in some species of the Leptostemonum , a subgenus of the nightshade ( Solanum ), a broad wing is formed; sometimes the seed coat is remarkably thick, as in the angel's trumpet ( Brugmansia ). The surface can be smooth ( Melananthus ), reticulated ( Juanulloa ), bumpy ( Solanum chamaesarachidium , Capsicophysalis ) or honeycomb ( Acnistus , Witheringia ).

The cells of the seed coats can be thick-walled ( Schizanthus ), with a wavy or wavy edge ( Ectozoma , Schizanthus ), deep ( Triguera , Witheringia , Jaborosa ) or flat ( Brachistus ), egg-shaped or reticulated-egg-shaped ( Hyoscyamus) ) be. A special case are the seeds of tomatoes ( Solanum lycopersicum ), the outermost layer of the seed coat is slimy and, when this dries, makes the seeds appear covered with trichome-like hairs.

#### embryo

An important feature for the morphological determination and systematization of nightshade family is the embryo contained in the seed . It can be thick ( Schultesianthus ) or slender ( Markea ), straight, sometimes long (in Metternichia : 17 to 19 mm), sometimes short (in Sessea just under 2 to 3 mm), slightly curved (in the subfamilies Anthocercidoideae, Cestroideae and Juanulloideae ), worm-shaped ( Ectozoma , Anthocercis ), ring-shaped (Tribus Benthamielleae) or helical to almost helical (in the subfamilies Solanoideae, Salpiglossoideae and the genera of the split flowers ( Schizanthus ) and Solandra ).

There are also different ways in which the cotyledons can be shaped: three species of the tribe Cestreae and the genera Merinthopodium , Markea and Juanulloa have embryos with wide cotyledons, while the rest of the family has cotyledons that are just as wide as the rest of the embryo. Furthermore, the ratio of the size of embryo and cotyledons within the family is different: In the subfamily Anthocercidoideae, the cotyledons are only one sixth to one eighth as long as the rest of the embryo, in all other subfamilies there are cotyledons that are just as long up to 2.5 to three times shorter than the rest of the embryo. Much attention in morphological studies of the seeds is also given to the way in which the cotyledons are arranged within the seed. In the subfamilies Solanoideae, Cestroideae, Salpiglossoideae and Schizanthoideae, the cotyledons are on top or slightly crooked, whereas the Juanulloideae have adjoining, in rare cases, crooked cotyledons.

Oily endosperms are very rare within the family. This characteristic is found mainly in the subfamily Juanulloideae and in the genus Metternichia . The development of the endosperm is usually cellular, with the exception of the genus schizanthus with nuclear development.

## distribution

Sketch of the distribution area of ​​the nightshade family (green)

The species of the nightshade family are widely distributed all over the world. There are some cosmopolitan genera such as buckthorn ( Lycium ), bladder cherries ( Physalis ) and nightshade ( Solanum ), but also genera that only occur in individual flora kingdoms. There are some endemic genera, such as Nothocestrum in Hawaii , Normania in the Canary Islands and Combera and Benthamiella in Patagonia . The genera Bouchetia , Grabowskia , Leptoglossis , Leucophysalis , Weißbecher ( Nierembergia ) and the petunias ( Petunia ) have disjoint distribution areas .

The diversity of the nightshade family of South America exceeds that of all other continents and sub-continents. In addition to the cosmopolitan common genera are in the Andes only occurring there genres to find 13 more in the Andes and Southeast South America three genera. One genus ( Sessea ) occurs both in South America and on the Antilles , there are 14 endemics and the genera mentioned above with disjoint distribution areas. With a large number of wild species, South America is also the gene center of important cultivated plants such as potatoes, peppers, tobacco and tomatoes.

If one assumes the number of existing genera, the nightshade family in Africa are relatively weakly represented with only eight genera. With the nightshade ( Solanum ) and goat thorns ( Lycium ) one can find two of the three cosmopolitan genera; in addition, with Triguera and the mandrake ( Mandragora ), two genera that Africa has in common with Europe. In addition, there are two genera, henbane ( Hyoscyamus ) and Withania , which occur in Asia, Europe and Africa and a single type of tobacco ( Nicotiana ) from Namibia , as well as the endemic genus Discopodium .

In Asia , on the one hand, the three cosmopolitan genera occur, and the genus Tubocapsicum , which only occurs in Asia . In addition, the deadly nightshade ( Atropa ) and the mandrakes ( Mandragora ), which can also be found in Europe, and the genera Lycianthes , henbane ( Hyoscyamus ) and Withania, which are common to America, exist here . Thus there are a total of nine of the nightshade genera in Asia.

The subfamily Anthocercidoideae with seven genera occurs exclusively in Australia . There are also 18 endemic species of the tobacco genus ( Nicotiana ) and a large number of species from other genera.

## Chromosome number

More than 50% of the examined nightshade species have a base chromosome number of , which also and often occur. The greatest variance can be found within the subfamily Cestroideae, in which all chromosome numbers from to appear. Large differences also occur in the subfamily Solanoideae, here in addition to the frequent chromosome numbers of (in some species of paprika ( Capsicum ) and a cultivar of tomato ( Solanum lycopersicum )), ( Solanum bullatum ) and (nightshade ( Solanum ), subgenus Archaesolanum ). The other subfamilies have chromosome numbers of (Juanulloideae), (Salpiglossoideae), (Schizanthoideae, Anthocercidoideae) or (Anthocercidoideae). ${\ displaystyle x = 12}$${\ displaystyle x = 7}$${\ displaystyle x = 13}$${\ displaystyle x = 7}$${\ displaystyle x = 13}$${\ displaystyle x = 10,12,14,17}$${\ displaystyle x = 13}$${\ displaystyle x = 15}$${\ displaystyle x = 23}$${\ displaystyle x = 12}$${\ displaystyle x = 11}$${\ displaystyle x = 10}$${\ displaystyle x = 9}$

It is not uncommon for polyploidy to occur within the family, this multiplication of the number of chromosomes is known from the white cups ( Nierembergia ), Withania , bladder cherries ( Physalis ), quincula , Chamaesaracha , nightshade ( Solanum ) - sections Solanum and Petota and subfamilies Leptostemonum and Archaesolanum , Alraesolanum ( Mandragora ) and goat thorns ( Lycium ). According to two reports, plants from the nightshade ( Solanum ) section of Solanum have been found to have octoploid sets of chromosomes with chromosomes. ${\ displaystyle 2n = 8x = 96}$

## Systematics

### External system

The nightshade family are classified in the order of the nightshade family, where they form a sister clade to the bindweed family (Convolvulaceae). Both families in turn form a monophyletic taxon, which is a sister clade to the families Hydroleaceae , Sphenocleaceae and Montiniaceae .

 Solanaceae

### Internal system

The number of genera within the family is between 90 and 100, the number of species stated varies between 2,300 and 9,000 to 10,000 species, depending on the source and author. An estimate from 2007 is based on 2716 recognized species.

The last taxonomic description of the family, which is complete except for the species level, was published in 1852 by Michel Félix Dunal ; later works usually only consider a small, botanically or regionally limited part of the family, or are only complete down to the genus level. More recent, phylogenetic studies are not yet fully available due to the size of the genus, so that new knowledge about the family system and thus further changes must be expected in the future.

The following system is based on the work of Richard Olmstead et al. from the year 2008, changes to this are marked with individual evidence:

## ingredients

Mainly because of the large number and the different areas of application of food and medicinal plants within the nightshade family, the family was examined relatively early in phytochemical studies. With the investigation of new species, more and more ingredients were found, so that the work on this family remained interesting and thus a very large number of phytochemical studies on the nightshade family are available.

Especially alkaloids and steroids have a significant position as a characteristic phytochemicals within the family.

### Alkaloids

A total of nine alkaloid groups were detected within the family, with the tropane alkaloids (example atropine ) being the most common group and appearing in at least 33 genera in five subfamilies (Solanoideae, Cestroideae, Salpiglossoideae, Schizanthoideae and Anthoceridoideae). Other proven alkaloid groups are steroid alkaloids , pyrrole alkaloids , Pyrazolalkaloide , Pyridinalkaloide , imidazole alkaloids , aliphatic alkaloids or alkaloids amines and amides , quinoline alkaloids and indole alkaloids .

The best-known alkaloid of the nightshade family is the pyridine alkaloid nicotine from the tobacco plant ( Nicotiana sp. ), Other known alkaloids are hyoscyamine , atropine , scopolamine and capsaicin .

Due to the special pharmacological properties of these alkaloids, the methods of extraction from the different parts of the plant and the chemical properties of the individual compounds have been described in detailed overviews.

In 19th century psychiatry , these alkaloids, in various mixtures and dosages, played an important role as therapeutic agents.

### Steroids

Most nightshade steroids are primarily classified as primary ingredients, only a few can be counted as secondary ingredients. Above all, phytosterols such as cholesterol , β-sitosterol , stigmasterol and campesterol as well as their glycosides and esters, but also steroid lactones in numerous variants are available within the whole family.

One of the phytochemically most interesting groups of steroid lactones is that of the withanolides , of which over 300 have so far been isolated from the subfamily Solanoideae, but not a single one in any of the other subfamilies. Similar to alkaloids, they serve the plant to ward off predators .

### Other ingredients

The occurrence of coumarins , which are also known from the umbelliferae , is characteristic of the nightshade family . Coumarin-free species are not known within the nightshade family. Nightshade plants rarely contain large amounts of essential oils ; iridoid compounds do not seem to run in the family. Nightshade plants form polyphenols , but not real tannins . In terms of flavonoids , kaempferol and quercetin mainly contain flavonols ; flavones are less common.

## Importance to humans

### food

French fries , one of the many
potato- based foods

Many nightshade plants are used as food by humans. Although the fruits are usually harvested, the most important food crop, the potato , uses a different part of the plant, namely the tuber that grows underground. In 2005 the world production of potatoes was 324.5 million tons (388 million tons in 2017). Other important foods among the nightshade family are tomatoes with an annual production of 124.7 million tons (2017: 182.3 million tons), eggplants with 30.8 million tons (2017: 52.3 million tons) and peppers or Chillies with 24.7 million tons (2017: 36 million tons) of fresh and 2.6 million tons (2017: 4.6 million tons) of dried fruit. All information relates to the year 2005, resp. supplemented with information on 2017.

Other members of the family used as food crops, whose production figures do not come close to those mentioned above, are some species of the nightshade genus, such as pepino , tamarillo , lulo , various types of bladder cherries , and more rarely wolfberry or Jaltomata .

It is occasionally reported that even species treated as poisonous plants are used as food. For example, according to various publications, the leaves and young shoots of the black nightshade are prepared as vegetables. The toxin content is often reduced by repeated boiling or adding ingredients that may detoxify, such as milk. The ripe fruits of the black nightshade and related species should also be eaten occasionally, sometimes after they have been previously cooked.

The three most important food plants among the nightshade family - potatoes, tomatoes and peppers - originally come from South and Central America, where some of them have been used as food for several thousand years. Remnants of potato peelings were found during excavations in Chile and date back to about 11,000 BC. Dated. The oldest known evidence of the cultivation and thus breeding forms of nightshade family is about 6,000 years old and comes from types of paprika.

The time at which the aubergine was imported into Europe via the Arab world cannot be precisely determined. It is likely that the Roman and Greek cultures did not yet know the plant, its use in the Arab world has been documented since the 11th century. The first description of the aubergine from Europe comes from the Historia stirpium  (1542) by Leonhart Fuchs , who already mentions its use as food there.

Above all, the plants imported from America were initially mostly grown as exotic ornamental plants, the culinary value was often only discovered after a long time. In Europe, however, the potato in particular, but also the tomato, achieved an important role as food by the 18th century, so that both plants were brought across the Atlantic again by European emigrants to cultivate them in North America. The increased dependency on potatoes as a food was particularly evident during the Great Famine in Ireland in the middle of the 19th century, which was triggered by several poor harvests of the previously common potato monocultures caused by disease and pests .

### Mystical plants, stimulants and intoxicants

Young man with a pipe (Michel Gobin, 17th century)
Representation of the mandrake root as a human body, 7th century

Many of the alkaloids contained in nightshade plants provide protection against predators, as they are often poisonous and have an effect on the central nervous system , especially in mammals and humans , and trigger hallucinations or drug psychoses , but can also lead to death. Evidence about the use of nightshade plants as an intoxicant is already known from the ancient cultures of the Greeks , Romans , Arabs and Hebrews , but reports on methods of use for producing intoxicating states have also come down from many other cultures.

Nightshade plants known as intoxicants include the common mandrake ( Mandragora officinarum ), the deadly nightshade ( Atropa belladonna ), the black henbane ( Hyoscyamus niger ), various thorn apples ( Datura sp. ) And the angel's trumpets ( Brugmansia ). The economically most important pleasure and intoxicating plant among the nightshade plants, however, is tobacco ( Nicotiana tabacum and others), whose world harvest in 2005 was 6.6 million tons unprocessed.

Various methods are described to achieve the intoxicating effect, including eating different parts of plants, smoking leaves and fruits, rubbing them with ointments made from plant extracts, adding fruits and seeds to drinks.

The common mandrake ( Mandragora officinarum ), whose branched taproot has often been compared with the shape of a human body, was particularly important as a mystical plant . In one of the oldest stories in the Bible, Genesis 30: 14-16  EU , a plant called dudai is mentioned, which is very likely to be identical to the mandrake . The plant is first mentioned in ancient Greece from around 400 BC. Known, Theophrastus mentioned around 230 BC. In addition to medical uses, it can also be used as an aphrodisiac . Further mentions of the plant can also be found in records from the Roman Empire, after its collapse the mandrake is initially little mentioned. Only between 1200 and 1600 did the plant regain its mystical and spiritual importance; it was valued as a talisman . However, there are also various myths surrounding the plant. It is often reported that the plant has the power to kill people trying to dig up the root. With the increasing persecution of witches , the mandrake appears again and again as an ingredient in the so-called witch's ointments , and other nightshade plants such as henbane , thorn apple or deadly nightshade can also be found in this context.

The legend of Odysseus , whose companions are turned into pigs by the sorceress Kirke , is often traced back to a gift of henbane ( Hyoscyamus ) and the hallucinations caused by it. Henbane is also listed as an addition to baths in medieval bathing rooms to encourage freedom of movement. The seeds were also used as an additive to beer. This was often forbidden, for example by a police order from 1507 from Eichstätt or the Bavarian purity law of 1516. Henbane gained further dubious fame through the murder trial of Hawley Crippen in 1910 , who killed his wife with hyoscine , the plant's poisonous alkaloid . The case received particular attention, on the one hand, because the arrest of Crippen was made possible by using telegram communication between Europe and America for the first time . On the other hand, the investigation and evidence in court is considered the first use of forensic medicine : The toxicologist Dr. William Willcox extracted the alkaloid that led to the poisoning from the stomach, the intestines, the kidneys and the liver of the corpse and was able to prove that it was hyoscine based on the boiling point.

At the end of the 1990s, studies of drug consumption behavior among adolescents revealed an increased use of plant hallucinogens. According to the survey results, herbal “ fashion drugs ” from the 1970s, such as peyote cactus or ayahuasca, were hardly used anymore . However, in addition to the increased use of various psychoactive mushrooms (e.g. Psilocybe sp. ), An increase in the consumption of nightshade plants such as angel's trumpets and thorn apple was found. These intoxicants are often wrongly classified as harmless by the users.

### Use in medicine

The development of the medicinal use of the nightshade family is closely linked to its history as an intoxicant, and its historically documented use can often be poorly classified into one of the categories. A first documented purely medical use of the nightshade family dates back to the first century AD by the Greek doctor Dioscurides , who describes how sweet wine mixed with mandrake root is used to anesthetize patients prior to surgical interventions. Many nightshade plants are known in various cultures in folk medicine, for example henbane is used for pain relief, whooping cough, ulcers or abdominal infections. Almost worldwide there is evidence of the use of the black nightshade ( Solanum nigrum ) and related species as a means of various diseases, above all against fever and inflammation of the digestive tract. In Brazil, the Dama da Noite ( Cestrum laevigatum ) is not only considered an intoxicant, but also an antiseptic , sedative , emollient ( soothing agent ) and liver stimulant . For a long time, burning and inhaling the smoke from Datura leaves was also considered a remedy for asthma.

The most well-known current area of ​​application of the nightshade family is the use of capsaicin extracts obtained from paprika ( Capsicum ) to promote blood circulation, among other things in rheumatism. Although the name of the ABC patch still refers to the ingredient belladonna (belladonna ( Atropa belladonna )), it is no longer used today. However, extracts from the are deadly nightshade in the field of ophthalmology for pupil dilation and gastrointestinal diseases. Furthermore, extracts from the seeds of the thorn apple are used against asthma, extracts from the bittersweet nightshade ( Solanum dulcamara ) against eczema and rheumatism, and the nicotine of various tobacco products to wean smokers, for example with nicotine patches or chewing gum. Potato starch is used as an additive to medicinal powders. Henbane is seldom used in medicine today, and extracts can only be found in a few asthma medicines and ointments.

### Ornamental plant

Petunia hybrids ( Petunia ) as a hanging plant

Many members of the family are valued as ornamental plants due to the mostly numerous and multi-colored, sometimes unusually shaped flowers of the nightshade family. Various hybrids of petunias such as the Surfinia petunias are among the most popular and economically most important balcony flowers. Angel's trumpets are grown in pots due to their exceptionally large flowers. Species and hybrids of tobacco known as ornamental tobacco are characterized by striking, strongly scented flowers in different colors. In recent years Lycianthes rantonnei , also known as the gentian bush or the potato tree, has become very popular as a container plant due to its many dark blue flowers. Various colorful flowering shrubs, such as from the genera Lycium or schizanthus ( Schizanthus ) can be used especially in warmer layers for green surface design. But also because of the decorative-looking fruits of some nightshade plants, these are grown in ornamental forms, for example the lantern flower ( Physalis alkegengi ), various varieties of paprika , the coral bush ( Solanum pseudocapsicum ) or the cow udder plant ( Solanum mammosum ).

## Origin of name

Both the German and the scientific name of the family are derived from the name of the genus nightshade ( Solanum ). The additions -aceae or -gewächse indicate the family status .

The name Nachtschatten is derived from the Old High German nahtscato or Middle High German seamschade . There are several theories for the interpretation of the name. On the one hand, “night shade” could mean the dark berries of the black nightshade , on the other hand the medicinal effect of the plants is also a possible derivation. Otto Brunfels wrote in 1532 in his Contrafayt Kreüterbuch : “This herb would also be used in other ways, against the damage that the witches inflict on the people, and the sometimes white, still opportunity of the damage encountered, not on special supersticion and magic. That is why it is called Nightshade in sonderheyt . ” Johann Christoph Adelung (1808) sees the origin of the name in connection with the headache (damage) caused by the flowers of the plants, which are strongly scented at night.

The name Solanum was taken over by Linnaeus from other authors, the meaning at that time included, among other things, deadly nightshade ( Atropa ), paprika ( Capsicum ), thorn apple ( Datura ), bladder cherries ( Physalis ) and nightshade ( Solanum ). But also very different groups of plants that name were subordinated to some extent, for example, magic flowers ( Mirabilis ), oneberries ( Paris ) and pokeweed ( Phytolacca ). The origin of the scientific name as well as that of the German name has not been clarified. The connection to the Latin sōl (sun), which is mentioned by some authors, is not to be assumed according to Exactly , the derivation from the Latin sōlārī (comfort, soothe) is more likely , which could indicate the medicinal effects of small doses of nightshade family.

## Sources and further information

### Main sources

• Armando T. Hunziker: The Genera of Solanaceae. ARG Gantner Verlag KG, Ruggell, Liechtenstein 2001, ISBN 3-904144-77-4 .
• JG Hawkes et al. (Editors): Solanaceae III: Taxonomy, Chemistry, Evolution . Royal Botanic Gardens, Kew, 1991, ISBN 0-947643-31-1 .
• M. Nee et al. (Editors): Solanaceae IV, Advances in Biology and Utilization . Royal Botanic Gardens, Kew, 1999, ISBN 1-900347-90-3 .

Note: In order to ensure uniformity in the designation of the sub-taxa, the systematic according to Hunziker The Genera of Solanaceae is used in the article . Later, generally recognized rearrangements, for example the classification of the genera Lycopersicon and Cyphomandra in the genus Solanum , were also taken into account.

### Individual evidence

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17. a b c The Colonial Williamsburg Foundation: Solanaceae ( Memento of the original from March 12, 2007 in the Internet Archive ) Info: The archive link was inserted automatically and has not yet been checked. Please check the original and archive link according to the instructions and then remove this notice. . Retrieved March 16, 2007.
18. a b The Colonial Williamsburg Foundation: Root Crops ( Memento of the original from August 5, 2007 in the Internet Archive ) Info: The archive link was inserted automatically and has not yet been checked. Please check the original and archive link according to the instructions and then remove this notice. . Retrieved March 16, 2007.
19. Linda Perry et al .: Starch Fossils and the Domestication and Dispersal of Chili Peppers (Capsicum spp. L.) in the Americas . In: Science , February 16, Vol. 315, No. 5814, 2007. pp. 986-988, doi : 10.1126 / science.1136914
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21. a b M. R. Lee: Solanaceae III: henbane, hags and Hawley Harvey Crippen ( Memento of the original of September 28, 2007 in the Internet Archive ) Info: The archive link was automatically inserted and not yet checked. Please check the original and archive link according to the instructions and then remove this notice. . In: Journal of the Royal College of Physician of Edinburgh , Volume 36, August 2006.
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29. Quoted from Marzell, Volume 4, page 366.
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