Digitalis lanata
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''Digitalis lanata'', vernacularly often called woolly foxglove or Grecian foxglove, is a species of
foxglove ''Digitalis'' ( or ) is a genus of about 20 species of herbaceous perennial plants, shrubs, and biennials, commonly called foxgloves. ''Digitalis'' is native to Europe, western Asia, and northwestern Africa. The flowers are tubular in shap ...
, a flowering plant in the plantain family
Plantaginaceae Plantaginaceae, the plantain family, is a large, diverse family of flowering plants in the order Lamiales that includes common flowers such as snapdragon and foxglove. It is unrelated to the banana-like fruit also called "plantain." In older cl ...
. It gets its name due to the woolly
indumentum In biology, an indumentum (Latin, literally: "garment") is a covering of trichomes (fine "hairs") on a plant Davis, Peter Hadland and Heywood, Vernon Hilton (1963) ''Principles of angiosperm taxonomy'' Van Nostrandpage, Princeton, New Jersey, pa ...
of the leaves. ''D. lanata'', like other foxglove species, is toxic in all parts of the plant. Symptoms of digitalis poisoning include nausea, vomiting, severe headache, dilated pupils, problems with eyesight, and convulsions at the worst level of toxicity. The plant is also harmful to other animals.


Description

''Digitalis lanata'' is a
biennial Biennial means (an event) lasting for two years or occurring every two years. The related term biennium is used in reference to a period of two years. In particular, it can refer to: * Biennial plant, a plant which blooms in its second year and th ...
or
perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wid ...
growing from a woody, horizontal rootstock. There is a tidy rosette before the spike goes up, and it is neatly arranged around the purple-tinged stems. The plant commonly forms a single, upright, more or less uniformly leafy stem that is partly ascending at the base. The plant send up these stems and flowers in usually in its second year. The stems are 0.3 to 0.6 meters in height, or about 13 to 26 inches. The leaves are moderately green in colour, woolly, veined, and covered with white hairs on the underside. They have a very bitter taste. The lower cauline leaves are 6 to 12 cm (sometimes to 20 cm) long and 1.5 to 3.5 cm wide, the upper cauline leaves are 4 to 10 cm long and
lanceolate The following is a list of terms which are used to describe leaf morphology in the description and taxonomy of plants. Leaves may be simple (a single leaf blade or lamina) or compound (with several leaflets). The edge of the leaf may be regular ...
shaped, usually with entire margins, and with a distinctive mid-rib. The leaves along the stalks are alternate. The lower stem leaves wither by early flowering. Both flowers and stems are also woolly or hairy. The
inflorescence An inflorescence is a group or cluster of flowers arranged on a Plant stem, stem that is composed of a main branch or a complicated arrangement of branches. Morphology (biology), Morphologically, it is the modified part of the shoot of sperma ...
axis An axis (plural ''axes'') is an imaginary line around which an object rotates or is symmetrical. Axis may also refer to: Mathematics * Axis of rotation: see rotation around a fixed axis * Axis (mathematics), a designator for a Cartesian-coordinat ...
is densely covered with densely matted woolly hairs (
tomentose Trichomes (); ) are fine outgrowths or appendages on plants, algae, lichens, and certain protists. They are of diverse structure and function. Examples are hairs, glandular hairs, scales, and papillae. A covering of any kind of hair on a pl ...
), and the flowers are densely arranged into a raceme that is pyramidal in shape. The flowers are tubular and bell shaped, pale yellow to whitish with brown or violet lines (
ferruginous The adjective ferruginous may mean: * Containing iron, applied to water, oil, and other non-metals * Having rust on the surface * With the rust (color) See also * Ferrous, containing iron (for metals and alloys) or iron(II) cations * Ferric, cont ...
reticulated markings), and the centre lobe of the lower lip is 8 to 13 mm long. The fruit is a conical capsule with a blunt end topped by a short beak. The seeds which develop within are quadrangular or prismatic in shape, and are about 0.6mm broad and 1.1 to 1.3mm long.


Distribution

It is native to
Anatolia Anatolia, tr, Anadolu Yarımadası), and the Anatolian plateau, also known as Asia Minor, is a large peninsula in Western Asia and the westernmost protrusion of the Asian continent. It constitutes the major part of modern-day Turkey. The ...
and the
Balkans The Balkans ( ), also known as the Balkan Peninsula, is a geographical area in southeastern Europe with various geographical and historical definitions. The region takes its name from the Balkan Mountains that stretch throughout the who ...
, where it is found in
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,
Bulgaria Bulgaria (; bg, България, Bǎlgariya), officially the Republic of Bulgaria,, ) is a country in Southeast Europe. It is situated on the eastern flank of the Balkans, and is bordered by Romania to the north, Serbia and North Macedo ...
,
Croatia , image_flag = Flag of Croatia.svg , image_coat = Coat of arms of Croatia.svg , anthem = "Lijepa naša domovino"("Our Beautiful Homeland") , image_map = , map_caption = , capit ...
,
Greece Greece,, or , romanized: ', officially the Hellenic Republic, is a country in Southeast Europe. It is situated on the southern tip of the Balkans, and is located at the crossroads of Europe, Asia, and Africa. Greece shares land borders ...
,
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(European Turkey),
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,
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,
Kosovo Kosovo ( sq, Kosova or ; sr-Cyrl, Косово ), officially the Republic of Kosovo ( sq, Republika e Kosovës, links=no; sr, Република Косово, Republika Kosovo, links=no), is a partially recognised state in Southeast Euro ...
and
Romania Romania ( ; ro, România ) is a country located at the crossroads of Central, Eastern, and Southeastern Europe. It borders Bulgaria to the south, Ukraine to the north, Hungary to the west, Serbia to the southwest, Moldova to the east, and ...
, eastwards to northern
Moldova Moldova ( , ; ), officially the Republic of Moldova ( ro, Republica Moldova), is a landlocked country in Eastern Europe. It is bordered by Romania to the west and Ukraine to the north, east, and south. The unrecognised state of Transnistr ...
. It has been introduced to Ukrainian Galicia,
Austria Austria, , bar, Östareich officially the Republic of Austria, is a country in the southern part of Central Europe, lying in the Eastern Alps. It is a federation of nine states, one of which is the capital, Vienna, the most populous ...
, the
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,
Estonia Estonia, formally the Republic of Estonia, is a country by the Baltic Sea in Northern Europe. It is bordered to the north by the Gulf of Finland across from Finland, to the west by the sea across from Sweden, to the south by Latvia, a ...
and
Germany Germany,, officially the Federal Republic of Germany, is a country in Central Europe. It is the second most populous country in Europe after Russia, and the most populous member state of the European Union. Germany is situated betwe ...
. Its presence in
Slovakia Slovakia (; sk, Slovensko ), officially the Slovak Republic ( sk, Slovenská republika, links=no ), is a landlocked country in Central Europe. It is bordered by Poland to the north, Ukraine to the east, Hungary to the south, Austria to the s ...
is questionable. In the US it has
naturalised Naturalization (or naturalisation) is the legal act or process by which a non-citizen of a country may acquire citizenship or nationality of that country. It may be done automatically by a statute, i.e., without any effort on the part of the i ...
in a number of northeastern states. Although it has yet only spread to a few counties per state, it is still considered potentially invasive or a noxious weed. The state of Minnesota considers it a noxious weed, and it is an invasive species of grasslands and woodlands in
Wisconsin Wisconsin () is a state in the upper Midwestern United States. Wisconsin is the 25th-largest state by total area and the 20th-most populous. It is bordered by Minnesota to the west, Iowa to the southwest, Illinois to the south, Lake M ...
, and
Kansas Kansas () is a state in the Midwestern United States. Its capital is Topeka, and its largest city is Wichita. Kansas is a landlocked state bordered by Nebraska to the north; Missouri to the east; Oklahoma to the south; and Colorado to th ...
. As of 2001, it may not be imported into Kansas. It has also naturalised in the US states of
Connecticut Connecticut () is the southernmost state in the New England region of the Northeastern United States. It is bordered by Rhode Island to the east, Massachusetts to the north, New York to the west, and Long Island Sound to the south. Its capita ...
,
Indiana Indiana () is a U.S. state in the Midwestern United States. It is the 38th-largest by area and the 17th-most populous of the 50 States. Its capital and largest city is Indianapolis. Indiana was admitted to the United States as the 19th s ...
,
Maine Maine () is a state in the New England and Northeastern regions of the United States. It borders New Hampshire to the west, the Gulf of Maine to the southeast, and the Canadian provinces of New Brunswick and Quebec to the northeast and ...
,
Maryland Maryland ( ) is a state in the Mid-Atlantic region of the United States. It shares borders with Virginia, West Virginia, and the District of Columbia to its south and west; Pennsylvania to its north; and Delaware and the Atlantic Ocean to ...
,
Massachusetts Massachusetts (Massachusett: ''Muhsachuweesut Massachusett_writing_systems.html" ;"title="nowiki/> məhswatʃəwiːsət.html" ;"title="Massachusett writing systems">məhswatʃəwiːsət">Massachusett writing systems">məhswatʃəwiːsət'' En ...
,
Michigan Michigan () is a U.S. state, state in the Great Lakes region, Great Lakes region of the Upper Midwest, upper Midwestern United States. With a population of nearly 10.12 million and an area of nearly , Michigan is the List of U.S. states and ...
,
Minnesota Minnesota () is a state in the upper midwestern region of the United States. It is the 12th largest U.S. state in area and the 22nd most populous, with over 5.75 million residents. Minnesota is home to western prairies, now given over to ...
,
Nebraska Nebraska () is a state in the Midwestern region of the United States. It is bordered by South Dakota to the north; Iowa to the east and Missouri to the southeast, both across the Missouri River; Kansas to the south; Colorado to the sout ...
,
New Hampshire New Hampshire is a state in the New England region of the northeastern United States. It is bordered by Massachusetts to the south, Vermont to the west, Maine and the Gulf of Maine to the east, and the Canadian province of Quebec to the nor ...
,
New Jersey New Jersey is a state in the Mid-Atlantic and Northeastern regions of the United States. It is bordered on the north and east by the state of New York; on the east, southeast, and south by the Atlantic Ocean; on the west by the Delaware ...
, New York,
Ohio Ohio () is a state in the Midwestern region of the United States. Of the fifty U.S. states, it is the 34th-largest by area, and with a population of nearly 11.8 million, is the seventh-most populous and tenth-most densely populated. The sta ...
,
Pennsylvania Pennsylvania (; ( Pennsylvania Dutch: )), officially the Commonwealth of Pennsylvania, is a state spanning the Mid-Atlantic, Northeastern, Appalachian, and Great Lakes regions of the United States. It borders Delaware to its southeast, ...
,
Vermont Vermont () is a state in the northeast New England region of the United States. Vermont is bordered by the states of Massachusetts to the south, New Hampshire to the east, and New York to the west, and the Canadian province of Quebec to ...
and
West Virginia West Virginia is a state in the Appalachian, Mid-Atlantic and Southeastern regions of the United States.The Census Bureau and the Association of American Geographers classify West Virginia as part of the Southern United States while the B ...
. In Canada it has become naturalised in southern
Ontario Ontario ( ; ) is one of the thirteen provinces and territories of Canada.Ontario is located in the geographic eastern half of Canada, but it has historically and politically been considered to be part of Central Canada. Located in Central C ...
and
Quebec Quebec ( ; )According to the Canadian government, ''Québec'' (with the acute accent) is the official name in Canadian French and ''Quebec'' (without the accent) is the province's official name in Canadian English is one of the thirtee ...
. Other areas where it might be naturalised are
Vietnam Vietnam or Viet Nam ( vi, Việt Nam, ), officially the Socialist Republic of Vietnam,., group="n" is a country in Southeast Asia, at the eastern edge of mainland Southeast Asia, with an area of and population of 96 million, making i ...
and the western
Himalayas The Himalayas, or Himalaya (; ; ), is a mountain range in Asia, separating the plains of the Indian subcontinent from the Tibetan Plateau. The range has some of the planet's highest peaks, including the very highest, Mount Everest. Over 10 ...
.


Taxonomy

The word ''lanatus'' means 'woolly'.


Ecology

In
Minnesota Minnesota () is a state in the upper midwestern region of the United States. It is the 12th largest U.S. state in area and the 22nd most populous, with over 5.75 million residents. Minnesota is home to western prairies, now given over to ...
it is in leaf all year, in flower in June and July, and the seeds ripen in early-mid September. The flowers are perfect (having both male and female organs). Bees
pollinate Pollination is the transfer of pollen from an anther of a plant to the stigma of a plant, later enabling fertilisation and the production of seeds, most often by an animal or by wind. Pollinating agents can be animals such as insects, birds, a ...
the flowers. In the Ukraine and Moldova it flowers in July and August. The species is adapted to sunny and warm sites. It prefers part shade and humus-rich soil, but also grows in sandy, stony loamy, and clay soils under dry or moist conditions. During the first vegetation period, only the leaf rosette is developed, flowering follows during the second vegetation period. Therefore, especially during the first year, there is a high resistance against drought stress. The leaves remain somewhat
turgid Turgor pressure is the force within the cell that pushes the plasma membrane against the cell wall. It is also called ''hydrostatic pressure'', and is defined as the pressure in a fluid measured at a certain point within itself when at equilibriu ...
even under very low leaf
water potential Water potential is the potential energy of water per unit volume relative to pure water in reference conditions. Water potential quantifies the tendency of water to move from one area to another due to osmosis, gravity, mechanical pressure and mat ...
, due to
osmotic Osmosis (, ) is the spontaneous net movement or diffusion of solvent molecules through a selectively-permeable membrane from a region of high water potential (region of lower solute concentration) to a region of low water potential (region o ...
adaptation by synthesis of non-ionic substances in the leaves. The drought induced diminishing of
photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored i ...
is reversible after a few hours following watering of the plants. Drought stress also reduces the
quantum yield The quantum yield (Φ) of a radiation-induced process is the number of times a specific event occurs per photon absorbed by the system. Applications Fluorescence spectroscopy The fluorescence quantum yield is defined as the ratio of the numb ...
of
photosystem Photosystems are functional and structural units of protein complexes involved in photosynthesis. Together they carry out the primary photochemistry of photosynthesis: the absorption of light and the transfer of energy and electrons. Photosystems ...
2. More than 70 bitter glycosides with cardiac activity, with five different
aglycone An aglycone (aglycon or genin) is the compound remaining after the glycosyl group on a glycoside is replaced by a hydrogen atom. For example, the aglycone of a cardiac glycoside would be a steroid molecule. Detection A way to identify aglycone ...
es
digitoxigenin Digitoxigenin, a cardenolide, is the aglycone of digitoxin. Digitoxigenin can be used to prepare actodigin. In Lednicer's book on steroids, it is made from deoxycholic acid Deoxycholic acid is a bile acid. Deoxycholic acid is one of the se ...
, gitoxigenin,
digoxigenin Digoxigenin (DIG) is a steroid found exclusively in the flowers and leaves of the plants '' Digitalis purpurea'', '' Digitalis orientalis'' and '' Digitalis lanata'' (foxgloves), where it is attached to sugars, to form the glycosides (e.g. Lana ...
, diginatigenin and gitaloxigenin in the leaves act as a protection against herbivores. Yield and concentration of these cardiac glycosides are enhanced by greenhouse cultivation through enhanced temperature and enhanced carbon dioxide concentration.


Uses


Medical

''Digitalis lanata'' contains a powerful cardiac glycoside that may be used by patients with heart conditions. Digoxin (digitalin) is a drug that is extracted from ''D. lanata''. It is used to treat some heart conditions such as atrial fibrillation and cardiac arrhythmias. Its cardiotonic effects slows atrioventricular conduction so that the heartbeat slows down and slightly increases contraction power (positive
inotropic An inotrope is an agent that alters the force or energy of muscular contractions. Negatively inotropic agents weaken the force of muscular contractions. Positively inotropic agents increase the strength of muscular contraction. The term ''inotro ...
effect). Because of the improved circulation in congestive heart failure caused by fast atrial fibrillation, the kidneys can function better, which stimulates the flow of urine, which lowers the volume of the blood and lessens the load on the heart. The medical uses of ''Digitalis'' glycosides in
veterinary Veterinary medicine is the branch of medicine that deals with the prevention, management, diagnosis, and treatment of disease, disorder, and injury in animals. Along with this, it deals with animal rearing, husbandry, breeding, research on nutri ...
care has been controversial because of its relatively low therapeutic to toxicity profile, divergent physiological responses, variation in side effects, and the difficulty in demonstrating positive therapeutic responses. Cardiac glycosides from ''D. lanata'' and other natural sources have toxic effects, producing ventricular automaticity and vagal effects, resulting in AV nodal blocking. Acute toxicity produces nausea, vomiting, lethargy, confusion, and weakness.
William Withering William Withering FRS (17 March 1741 – 6 October 1799) was an English botanist, geologist, chemist, physician and first systematic investigator of the bioactivity of digitalis. Withering was born in Wellington, Shropshire, the son of a surg ...
is credited as the first to clinically investigate the plant as a therapy for
dropsy Edema, also spelled oedema, and also known as fluid retention, dropsy, hydropsy and swelling, is the build-up of fluid in the body's tissue. Most commonly, the legs or arms are affected. Symptoms may include skin which feels tight, the area ma ...
, writing a book in 1785 about the potential medical uses of ''D. purpurea'' extracts after his human trials. Digitalin was not discovered until the mid-19th century by two French scientists Homolle Ouevenne and Theodore Ouevenne. It was not until 1875 that Oscar Schmiedberg identified digoxin in the plant. It was first isolated in the 1930s in Britain by Sydney Smith. Today it is still extracted from the plant because industrial synthesis is too expensive and difficult. However, it is becoming less frequently used due to the narrow therapeutic margin and high potential for severe side effects. Digoxin is being replaced by newer drugs including beta blockers,
angiotensin Angiotensin is a peptide hormone that causes vasoconstriction and an increase in blood pressure. It is part of the renin–angiotensin system, which regulates blood pressure. Angiotensin also stimulates the release of aldosterone from the adren ...
-converting enzyme inhibitors, and the calcium channel blocking agents. As new pharmacotherapeutic agents arise, the use of digitalis preparations is expected to continue to decline. Asylums used digitalis extracts throughout the 19th century as sedatives along with hyoscine and opiates. The commercial production of digoxin from ''D. lanata'' involves growing the plant from seed for two years, harvesting and drying it in a silo, then the leaves are crushed into a powder, and the compound is extracted and purified using chemical processes.


Other

Digoxigenin Digoxigenin (DIG) is a steroid found exclusively in the flowers and leaves of the plants '' Digitalis purpurea'', '' Digitalis orientalis'' and '' Digitalis lanata'' (foxgloves), where it is attached to sugars, to form the glycosides (e.g. Lana ...
(DIG) is a steroid found in the flowers and leaves of ''
Digitalis ''Digitalis'' ( or ) is a genus of about 20 species of herbaceous perennial plants, shrubs, and biennials, commonly called foxgloves. ''Digitalis'' is native to Europe, western Asia, and northwestern Africa. The flowers are tubular in shap ...
'' species, and is extracted from ''D. lanata''. Digoxigenin can be used as a
molecular probe A molecular probe is a group of atoms or molecules used in molecular biology or chemistry to study the properties of other molecules or structures. If some measurable property of the molecular probe used changes when it interacts with the analyte ...
to detect
mRNA In molecular biology, messenger ribonucleic acid (mRNA) is a single-stranded molecule of RNA that corresponds to the genetic sequence of a gene, and is read by a ribosome in the process of synthesizing a protein. mRNA is created during the ...
in situ and label DNA, RNA, and
oligonucleotides Oligonucleotides are short DNA or RNA molecules, oligomers, that have a wide range of applications in genetic testing, research, and forensics. Commonly made in the laboratory by solid-phase chemical synthesis, these small bits of nucleic acids ...
. It can easily be attached to nucleotides such as
uridine Uridine (symbol U or Urd) is a glycosylated pyrimidine analog containing uracil attached to a ribose ring (or more specifically, a ribofuranose) via a β-N1-glycosidic bond. The analog is one of the five standard nucleosides which make up nucle ...
by chemical modifications. DIG molecules are often linked to nucleotides; DIG-labelled uridine can then be incorporated into RNA via ''
in vitro ''In vitro'' (meaning in glass, or ''in the glass'') studies are performed with microorganisms, cells, or biological molecules outside their normal biological context. Colloquially called " test-tube experiments", these studies in biology ...
''
transcription Transcription refers to the process of converting sounds (voice, music etc.) into letters or musical notes, or producing a copy of something in another medium, including: Genetics * Transcription (biology), the copying of DNA into RNA, the fir ...
. Once hybridisation occurs, RNA with the incorporated DIG-U can be detected with anti-DIG antibodies conjugated to alkaline phosphatase. To reveal the hybridised transcripts, a
chromogen In chemistry, the term chromogen refers to a colourless (or faintly coloured) chemical compound that can be converted by chemical reaction into a compound which can be described as "coloured". There is no universally agreed definition of the term. ...
can be used which reacts with the alkaline phosphatase to produce a coloured precipitate.


Pictures

Image:Digitalis lanata (digitale laineuse).JPG, Image:Digitalis lanata 001.JPG, Image:Digitalis lanata. Wooly Foxglove (30755195067).jpg, Image:Digitalis lanata LC0053.jpg, Image:Digitalis lanata 002.JPG, Image:Digitalis lanata 003.JPG, Image:Digitalis lanata 04.jpg, Image:Digitalis lanata 004.JPG,


References


Further reading

*https://www.drugs.com/npp/digitalis.html *http://www.discoveriesinmedicine.com/Com-En/Digitalis.html


External links


Plants for a Future database
{{Taxonbar, from=Q157838 lanata Taxa named by Jakob Friedrich Ehrhart