Umbelliferae

Umbelliferae
Wild carrot ( Daucus carota subsp. Carota ), illustration
Systematics
Eudicotyledons Nuclear eudicotyledons Asterids Euasterids II Order : Umbelliferae (Apiales) Family : Umbelliferae
Scientific name
Apiaceae
Lindl.

The umbelliferae or umbelliferae (Apiaceae or Umbelliferae) are a family of plants in the order of the umbelliferae (Apiales). Most of the species are herbaceous plants with leaves that are divided several times and double umbels as an inflorescence , making them easy to assign to the family.

The family contains about 434 genera with about 3780 species , and is represented worldwide in the temperate zones . The umbelliferae include many aromatic plants and food plants , but also some very poisonous plant species, for example the water hemlock and the spotted hemlock .

Description and ecology

Vegetative characteristics

The representatives of the umbelliferae are almost exclusively perennial herbaceous plants. A few taxa, such as in the subfamily Mackinlayoideae, are lignified . The stem axis is usually hollow and nodular . The stature heights range from several meter high plants in the steppes of Central Asia ( Ferula ) to a few centimeter high cushion plants in the Antarctic ( Azorella ).

Many species form a taproot . The lateral roots arise on both sides of the xylempole, as a resin duct runs at the tip of the xylempole.

The alternate leaves are single or multiple pinnate . Only in exceptional cases do they have simple leaves ( bupleurum ). The leaves have a leaf sheath .

Inflorescences

Inflorescence (double umbel) in the forest angelica - the individual umbel can be clearly seen

The inflorescence is usually a multi-rayed double umbel , an umbel made of usually many umbel . This structure of the inflorescences is very characteristic of the umbelliferae and has also given them their old scientific name Umbelliferae (umbrella carrier).

The bracts of the umbels are tightly packed together and form the envelope (involucre), they are often only weakly developed or absent. This is where the dome stalks originate = umbel rays. The umbellules are in turn surrounded by a (often missing) envelope (involucellum). The flower stalks are not called "dome rays". Often the inflorescence forms a dome or even an area on which insects can often be found.

Simple umbels are less common. There are also species with single flowers ( Azorella ). In the case of very large species, several double umbels can be combined to form an even larger inflorescence ( giant hogweed , Heracleum mantegazzianum ).

blossoms

Flower diagram of Laserpitium

With the exception of the gynoeceum, the mostly inconspicuous flowers are five-fold and usually radially symmetrical . In some species, however, the marginal florets in particular are asymmetrical and therefore zygomorphic . There are originally five sepals , but they are often stunted or absent. The five petals are free and are mostly white, more rarely yellow, pink to purple. The petals often have a knocked-in lobule (lobulum inflexum) at the tip. Its shape as well as the shape of the front edge of the petal (flexure edge) are important determinants.

There is only one circle with five free, fertile stamens that are curved in the bud. Two carpels have become an under constant ovary grown. The two styluses (also known as beak) sit on a disc-shaped to conical, shiny, stylus cushion (Stylopodium). This serves as a nectarium , i.e. i.e., it excretes nectar . In each of the two ovary compartments there is a hanging anatropic ovule . A second withers away very early.

The structure of the flower can be summarized in the following flower formula : ${\displaystyle \star K_{5-0}\;C_{5}\;A_{5}\;G_{\overline {(2)}}}$

The flowers are mostly protandric . The pollination is usually about flies , beetles and other insects kurzrüsselige ( Entomophilie ).

Section of an infructescence with young fruits of the Alpine eyelid ( Athamanta cretensis )

Fruits and seeds

The fruit is a dry, two-part split fruit , also known as a double achene. The shape is often cylindrical with a round to elliptical cross-section. Spherical ( Coriandrum ) and double- spherical shapes ( Bifora ) are rarer . The two partial fruits (mericarpies or carpids) initially remain with the upper side on a fruit holder ( carpophor ), which is located in the middle.

Each partial fruit has five longitudinal ribs or main ribs (juga primaria) each with a vascular bundle on its free side. In between there are valleys (valleculae), in each of the walls of which there is a mostly dark schizogenic oil duct (here called oil streak) (caused by the divergence of cells). In some species, each valley has a secondary rib (jugum secundarium, for example the carrot with prickly secondary ribs). The oil ducts can also be increased ( Pimpinella ) or reduced ( Coriandrum ) or completely absent ( Conium ).

The seed consists of a very small embryo in a large, fat and protein-rich endosperm . The embryo lies at the top of the seed with the hypocotyl pointing upwards . The seed coat is glued to the pericarp.

The spread occurs through animals (epi zoochory ), the wind ( anemochory ), water ( hydrochory ), through self- propagation ( autochory ) and partly through humans ( hemerochory ).

Chemical characteristics

The main components of the essential oils can be mainly formed from terpenes or from phenylpropanoids , depending on the type . With coriander it is predominantly (+) - linalool (terpene), with caraway (+) - carvone (terpene), with fennel and aniseed anethole (phenylpropanoid).

Ingredients (selection)
( S ) - (+) - carvone
Psoralen , a linear furanocoumarin
cis -Anethol (top) and trans -Anethol (bottom)
Falcarinol , a polyacetylene

The umbelliferae are the family with the largest range of coumarin compounds. In addition to simple coumarins and hydroxycoumarins (e.g. umbelliferone ), there are also a large number of prenylated, geranylated and farnesylated coumarin derivatives. This also includes the furano and pyranocoumarins. The former can be linear or angular. Hydroxy and furanocoumarins have a deterrent effect on herbivores (deterrent), as phytoalexins and as germination inhibitors. The toxicity increases from hydroxy to linear to angular furanocoumarins. The furanocoumarins are phototoxic : when exposed to UV light, the DNA is inactivated (photosensitization). Angular furanocoumarins are more toxic than linear ones, although their phototoxicity is less. Most of the species-rich genera of the family, which are widespread in the Holarctic, contain furanocoumarins ( e.g. Bupleurum and Pimpinella with 150 species each), while many monotypical genera with limited geographical distribution do not contain furanocoumarins.

Sesquiterpene lactones are found in the family with over 100 compounds. The same basic structures (e.g. germacranolide, eudesmanolide, eremophilanolide and elemanolide) occur as in the sunflower plants, but they are stereochemically different. They are also more frequently hydroxylated and esterified, especially at C11.

Over 150 polyacetylene compounds have been detected in the umbelliferous plants . The most common are the C ₁₇ diyne dienes of the falcarinol group. The toxicity of the water hemlock ( Cicuta virosa ) and the saffron vine umbels ( Oenanthe crocata ) is based on polyacetylenes.

Alkaloids are rare. Coniin and similar piperidine derivatives occur in the spotted hemlock ( Conium maculatum ). Triterpene saponins often occur in the subfamily Saniculoideae . Typical carbohydrates are the trisaccharide umbelliferose and the sugar alcohol mannitol .

The presence of petroselinic acid as the main fatty acid testifies to the close connection between the Apiaceae and the Araliaceae .

use

Because of the essential oils, many species are used as aromatic, vegetable and medicinal plants. The fruits, leaves and roots are used. Examples are caraway ( Carum carvi ), anise ( Pimpinella anisum ), coriander ( Coriandrum sativum ), dill ( Anethum graveolens ), lovage ( Levisticum officinale ), fennel ( Foeniculum vulgare ), parsley ( Petroselinum crispum ), and celery ( Apium graveolens ) .

Carrot ( Daucus carota ) and parsnip ( Pastinaca sativa ), which are mainly grown for their carbohydrate content, are a certain exception .

Some species are very poisonous. The spotted hemlock ( Conium maculatum ) provided the poison for Socrates, who was sentenced to death . The water hemlock ( Cicuta virosa ) is also very poisonous . Dog parsley ( Aethusa cynapium ) is less poisonous , but it is often confused with parsley , which often leads to poisoning.

Many species are photosensitizing and phototoxic due to their furanocoumarins . The giant hogweed ( Heracleum mantegazzianum ) is particularly worth mentioning here . However, the psoralen contained in the phototoxic species is used in medicine as part of PUVA therapy for the treatment of skin diseases.

Occurrence

The family is distributed worldwide, but the main focus is in the northern temperate zones . In the tropics , umbelliferae are particularly widespread in the montane altitudes . The umbelliferae grow mainly in steppes, swamps, meadows and forests.

Systematics

Subfamily Mackinlayoideae: Actinotus helianthi

Subfamily Mackinlayoideae: Indian pennywort ( Centella asiatica )

Synonyms for Apiaceae Lindl. nom. cons. are: Umbelliferae Juss. nom. cons., Actinotaceae AIKonstant. & Melikyan , Ammiaceae Bercht. & J.Presl , Angelicaceae Martinov , Daucaceae Martinov , Ferulaceae Sacc. , Saniculaceae Bercht. & J. Presl .

The sister taxon of the umbelliferae within the order Apiales is the group of Pittosporaceae , Araliaceae and Myodocarpaceae . The family contains about 434 genera with about 3780 species . Since 2010 it has only been divided into three subfamilies.

• The Mackinlayoideae Plunkett & Lowry are mostly woody plants. Most species grow in the areas around the South Pacific, the genus Centella mainly in South Africa . There are about six genera with about 67 species:
  • Actinotus Labill. : All 15 species are common in Australia and one of them is also found in New Zealand .
  • Apiopetalum Baill. : The four or so species occur in New Caledonia .
  • Centella L. (Syn .: Trisanthus Lour. ): The approximately 40 species are mainly distributed in the southern hemisphere , for example:
    • Indian pennywort ( Centella asiatica (L.) Urb. )
  • Mackinlaya F. Muell. (Syn .: Anomopanax Harms ): The ten or so species occur in Sulawesi , the Philippines , New Guinea , the Solomon Islands and northeastern Australia.
  • Micropleura Lag .: There are about two species in Colombia and Chile .
  • Platysace Bunge : The approximately 26 species are widespread in Australia, but none are found in Tasmania .
  • Xanthosia Rudge : The 20 or so species are widespread in temperate Australia.

Subfamily Azorelloideae: Yareta ( Azorella yareta )

Subfamily Azorelloideae: Bolax gummifera

Subfamily Azorelloideae: Mulinum spinosum

• The Azorelloideae Plunkett & Lowry occur in South America, Australia and the Antarctic. They have a large nucellus, a tetrasporous embryo sac. The fruits have a lignified perisperm. It contains about 21 to 23 genera with about 155 species:
  • Asteriscium Cham. & Schltdl. : It contains only one type:
    • Asteriscium novarae Constance & Charpin : It occurs in Chile and Argentina .
  • Andean cushion ( Azorella Lam. , Syn .: Fragosa Ruiz & Pav. , Laretia Gillies & Hook. , Mulinum Pers. ): It contains about 70 species, including:
    • Yareta ( Azorella compacta Phil. )
  • Bolax Comm. ex Juss. : The four to five species occur in temperate southern South America in Argentina and Chile.
  • Bowlesia Ruiz & Pav .: It is mainly found in South America.
  • Dichosciadium Domin : It contains only one species:
    • Dichosciadium ranunculaceum (F.Muell.) Domin (Syn .: Azorella ranunculacea (F.Muell.) Druce , Dichopetalum ranunculaceum F.Muell. , Azorella dichopetala Benth. Nom. Illeg., Pozoa ranunculacea (F.Muell.) Drude ): It occurs with two varieties only in the Australian states of New South Wales , Victoria and Tasmania .
  • Dickinsia Franch. (Syn .: Cotylonia C. Norman ): It contains only one species:
    • Dickinsia hydrocotyloides Franch. (Syn .: Cotylonia bracteata C. Norman ): It thrives in shady, damp forests and the banks of rivers at altitudes between 1500 and 3200 meters in the Chinese provinces of Guizhou , Hubei , Hunan , Sichuan and Yunnan .
  • Diplaspis Hook. f. : The only two species occur only in the Australian states of New South Wales, Victoria and Tasmania.
  • Diposis DC. : The two or three types occur in Uruguay and Chile.
  • Domeykoa Phil .: The four or so species occur in Peru and Chile.
  • Drusa DC. : It contains only one type:
    • Drusa glandulosa (Poir.) H.Wolff ex Engl .: It occurs in the Canary Islands , Madeira , Morocco and also in Somalia .
  • Eremocharis Phil .: The nine or so species occur in Peru and Chile.
  • Gymnophyton Clos : The six or so species thrive in the Andes of Chile and Argentina.
  • Hermas L .: There are about seven species common in southern Africa.
  • Homalocarpus Hook. & Arn. : The four to six species occur in Chile.
  • Huanaca Cav. : The four or so species occur in Peru and Chile.
  • Oschatzia Walp. : The only two species occur only in the Australian states of New South Wales, Victoria and Tasmania.
  • Pozoa Lag .: The roughly two species thrive in the Andes of Chile and Argentina.
  • Schizeilema (Hook. F.) Domin : It contains only one species:
    • Schizeilema fragoseum (F.Muell.) Domin : It occurs only in the Australian states of New South Wales and Victoria.
  • Spananthe Jacq. : It contains only one type:
    • Spananthe paniculata Jacq. : It occurs in the Andes.

Subfamily Apioideae: Opopanax chironium

• The Apioideae Drude , including the taxa of the former Saniculoideae Burnett . This subfamily has no stipules. The ovules are tenuinucellate, the endocarp consists of a layer of cells and is not lignified. The leaves are usually divided several times or undivided. The inflorescences are usually double umbels. The partial fruits can have a fruit holder (carpophor) and have more or less pronounced oil ducts. They have terpenoids of the Kauren-type. With the taxa of the former Saniculoideae there are now over 400 genera with around 3500 species. They are distributed worldwide, but mainly in the northern temperate zones. Lignify a few taxa, for example some Bupleurum and Myrrhidendron species. The basic chromosome number is x = 11 (8, 9, 12). For the internal systematics of this subfamily see main article: Apioideae .

swell

• The Apiaceae family on the AP website (sections description and systematics)
• The Apiaceae family at DELTA. (Section description)
• Menglan She, Fading Pu, Zehui Pan, Mark Watson, John FM Cannon, Ingrid Holmes-Smith, Eugene V. Kljuykov, Loy R. Phillippe & Michael G. Pimenov: Apiaceae. , P. 1 - online with the same text as the printed work , In: Wu Zheng-yi & Peter H. Raven (Eds.): Flora of China , Volume 14 - Apiaceae through Ericaceae , Science Press and Missouri Botanical Garden Press, Beijing and St. Louis, 2005. ISBN 1-930723-41-5 (Sections Description and Systematics)
• JM Powell: Apiaceae at the New South Wales Flora Online . (Sections Description and Distribution)
• Anthony R. Magee, Carolina I. Calviño, Mei (Rebecca) Liu, Stephen R. Downie, Patricia M. Tilney & Ben-Erik van Wyk: New tribal delimitations for the early diverging lineages of Apiaceae subfamily Apioideae , In: Taxon , Volume 59, Issue 2, 2010, pp. 567-580: PDF-Online. (Section systematics)
• Andreas Bresinsky , Christian Körner , Joachim W. Kadereit , Gunther Neuhaus , Uwe Sonnewald : Strasburger - textbook of botany. Founded by E. Strasburger. Spektrum Akademischer Verlag, Heidelberg 2008 (36th edition) ISBN 978-3827414557
• Andreas Herde: Investigation of the coumarin pattern in fruits of selected Apiaceae . Dissertation Universität Hamburg, 2005 ( full text pdf 2.3 MB ) (sections chemical characteristics and use)

Individual evidence

1. ↑ T. Stuhlfauth, H. Fock, H. Huber, K. Klug: The distribution of fatty acids including petroselinic and tariric acids in the fruit and seed oils of the Pittosporaceae, Araliaceae, Umbelliferae, Simarubaceae and Rutaceae. In: Biochemical Systematics and Ecology. 13, 1985, pp. 447-453, doi : 10.1016 / 0305-1978 (85) 90091-2 .
2. ^ ^{A b} Apiaceae in the Germplasm Resources Information Network (GRIN), USDA , ARS , National Genetic Resources Program. National Germplasm Resources Laboratory, Beltsville, Maryland.
3. ↑ APG page on Apiales
4. ^ ^{A b} Anthony R. Magee, Carolina I. Calviño, Mei (Rebecca) Liu, Stephen R. Downie, Patricia M. Tilney & Ben-Erik van Wyk: New tribal delimitations for the early diverging lineages of Apiaceae subfamily Apioideae , In: Taxon , Volume 59, Issue 2, 2010, pp. 567-580: PDF-Online.
5. ↑ Gregory M. Plunkett, GT Chandler, PP Lowry II, SM Pinney & TS Sprenkle: Recent advances in understanding Apiales and a revised classification , In: South African Journal of Botany , Volume 70, No. 3, 2004, p. 371– 381.
6. ↑ M. Liu, B.-E. Van Wyk, PM Tilney, Gregory M. Plunkett & PP Lowry II: Evidence from fruit structure supports in general the circumscription of Apiaceae subfamily Azorelloideae , In: Plant Systematics and Evolution , Volume 280, 2009, pp. 1-13.

Supplementary literature

• Dietrich Frohne, Uwe Jensen: Systematics of the plant kingdom with special consideration of chemical characteristics and plant drugs . 4th revised edition. Gustav Fischer, Stuttgart / Jena / New York 1992, ISBN 3-437-20486-6 .
• Siegmund Seybold (Ed.): Schmeil-Fitschen interactive . CD-ROM, version 1.1. Quelle & Meyer, Wiebelsheim 2002, ISBN 3-494-01327-6 .

Web links

Commons : Umbelliferae (Apiaceae)  - Collection of images, videos and audio files

• Versatile information from the Umbellifer Resource Center. (engl.)