Lobaria pulmonaria
Encyclopedia
Lobaria pulmonaria is a large epiphytic
Epiphyte
An epiphyte is a plant that grows upon another plant non-parasitically or sometimes upon some other object , derives its moisture and nutrients from the air and rain and sometimes from debris accumulating around it, and is found in the temperate zone and in the...

 lichen
Lichen
Lichens are composite organisms consisting of a symbiotic organism composed of a fungus with a photosynthetic partner , usually either a green alga or cyanobacterium...

 consisting of an ascomycete fungus
Fungus
A fungus is a member of a large group of eukaryotic organisms that includes microorganisms such as yeasts and molds , as well as the more familiar mushrooms. These organisms are classified as a kingdom, Fungi, which is separate from plants, animals, and bacteria...

 and a green algal partner living together in a symbiotic
Symbiosis
Symbiosis is close and often long-term interaction between different biological species. In 1877 Bennett used the word symbiosis to describe the mutualistic relationship in lichens...

 relationship with a cyanobacterium—a symbiosis involving members of three kingdoms
Kingdom (biology)
In biology, kingdom is a taxonomic rank, which is either the highest rank or in the more recent three-domain system, the rank below domain. Kingdoms are divided into smaller groups called phyla or divisions in botany...

 of organisms. Commonly known by various names like tree lungwort, lung lichen, lung moss, lungwort lichen, oak lungs or oak lungwort, it is sensitive to air pollution
Air pollution
Air pollution is the introduction of chemicals, particulate matter, or biological materials that cause harm or discomfort to humans or other living organisms, or cause damage to the natural environment or built environment, into the atmosphere....

 and is also negatively affected by habitat loss and changes in forestry
Forestry
Forestry is the interdisciplinary profession embracing the science, art, and craft of creating, managing, using, and conserving forests and associated resources in a sustainable manner to meet desired goals, needs, and values for human benefit. Forestry is practiced in plantations and natural stands...

 practices. Its population has declined across Europe
Europe
Europe is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally 'divided' from Asia to its east by the watershed divides of the Ural and Caucasus Mountains, the Ural River, the Caspian and Black Seas, and the waterways connecting...

 and L. pulmonaria is considered endangered
Endangered species
An endangered species is a population of organisms which is at risk of becoming extinct because it is either few in numbers, or threatened by changing environmental or predation parameters...

 in many lowland
Lowland
In physical geography, a lowland is any broad expanse of land with a general low level. The term is thus applied to the landward portion of the upward slope from oceanic depths to continental highlands, to a region of depression in the interior of a mountainous region, to a plain of denudation, or...

 areas. The species has a history of use in traditional medicine
Traditional medicine
Traditional medicine comprises unscientific knowledge systems that developed over generations within various societies before the era of modern medicine...

s, and recent research has corroborated some medicinal properties of lichen extracts.

Description

It is a foliose lichen and its leaf-like thallus is green, leathery and lobed with a pattern of ridges and depressions on the upper surface. Bright green under moist conditions, it becomes brownish and papery when dry. This species often has a fine layers of hairs, a tomentum, on its lower surface. The cortex, the outer protective layer on the thallus surface, is roughly comparable to the epidermis of a green plant. The thallus is typically 5 – in diameter, with individual lobes 1 – wide and up to 7 cm long. The asexual reproductive structures soredia and isidia are present on the thallus surface. Minute (0.5–1.5 mm in diameter) cephalodia—pockets of cyanobacteria—are often present on the lower surface of the thallus; these spots are conspicuously darker than the green surface of the thallus. Like other foliose lichens, the thallus is only loosely attached to the surface on which it grows.

Photobionts

The thallus contains internal structures known as cephalodia
Cephalodium
Cephalodia are small gall-like structures found in some species of lichens that contain cyanobacterial symbionts. Cephalodia can occur within the tissues of the lichen, or on its upper or lower surface. Lichens that have both green algal and cyanobacterial symbionts restrict the cyanobacteria to...

, characteristic of three-membered lichen symbioses involving two photobionts (the photosynthetic symbionts in the fungal-algal lichen relationship). These internal cephalodia, found between the "ribs" of the thallus surface, arise when blue-green algae (from the genus Nostoc
Nostoc
Nostoc is a genus of cyanobacteria found in a variety of environmental niches that forms colonies composed of filaments of moniliform cells in a gelatinous sheath.The name "Nostoc" was invented by Paracelsus...

) on the thallus surface are enveloped during mycobiont growth. Structurally, cephalodia consist of dense aggregates of Nostoc cells surrounded by thin-walled hypha
Hypha
A hypha is a long, branching filamentous structure of a fungus, and also of unrelated Actinobacteria. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium; yeasts are unicellular fungi that do not grow as hyphae.-Structure:A hypha consists of one or...

e—this delimits them from the rest of the thallus which contains a loose structure of thick-walled hyphae. Blue-green cyanobacteria can fix atmospheric nitrogen
Nitrogen fixation
Nitrogen fixation is the natural process, either biological or abiotic, by which nitrogen in the atmosphere is converted into ammonia . This process is essential for life because fixed nitrogen is required to biosynthesize the basic building blocks of life, e.g., nucleotides for DNA and RNA and...

, enhancing nutrient availability for the lichen. The other photobiont of L. pulmonaria is the green algae Dictyochloropsis reticulata
Dictyochloropsis reticulata
Dictyochloropsis reticulata is a species of green algae in the Microthamniales. It is a known as a photobiont with several lichen species, like Lobaria pulmonaria, but also as a free-living soil alga as well...

.

Reproduction

L. pulmonaria has the ability to form both vegetative propagation
Vegetative
Vegetative describes vegetation.Vegetative may also refer to:*Vegetative reproduction, a type of asexual reproduction for plants*Persistent vegetative state, a condition of people with severe brain damage...

 and sexual propagules at an age of about 25 years. In sexual reproduction, the species produces small reddish-brown discs known as apothecia
Ascocarp
An ascocarp, or ascoma , is the fruiting body of an ascomycete fungus. It consists of very tightly interwoven hyphae and may contain millions of asci, each of which typically contains eight ascospores...

 containing asci
Ascus
An ascus is the sexual spore-bearing cell produced in ascomycete fungi. On average, asci normally contain eight ascospores, produced by a meiotic cell division followed, in most species, by a mitotic cell division. However, asci in some genera or species can number one , two, four, or multiples...

, from which spore
Spore
In biology, a spore is a reproductive structure that is adapted for dispersal and surviving for extended periods of time in unfavorable conditions. Spores form part of the life cycles of many bacteria, plants, algae, fungi and some protozoa. According to scientist Dr...

s are forcibly released into the air (like ballistospore
Ballistospore
A spore that is shot off. In fungi most types of basidiospores, formed on basidia are discharged into the air from the tips of sterigmata. These actively discharged spores are a type of ballistospore....

s). Based on studies of ascospore germination
Germination
Germination is the process in which a plant or fungus emerges from a seed or spore, respectively, and begins growth. The most common example of germination is the sprouting of a seedling from a seed of an angiosperm or gymnosperm. However the growth of a sporeling from a spore, for example the...

, it has been suggested that L. pulmonaria spores use some mechanism to inhibit germination—the inhibition is lifted when the spores are grown in a synthetic growth medium containing an adsorbent like bovine serum albumin
Bovine serum albumin
Bovine serum albumin is a serum albumin protein derived from cows. It is often used as a protein concentration standard....

 or α-cyclodextrin
Cyclodextrin
Cyclodextrins are a family of compounds made up of sugar molecules bound together in a ring ....

.

Dispersal by vegetative propagules (via soredia or isidia) has been determined as the predominant mode of reproduction in L. pulmonaria. In this method, the protruding propagules become dry and brittle during the regular wet/dry cycles of the lichen, and can easily crumble off the thallus. These fragments may develop into new thalli, either at the same locale or at a new site after dispersal by wind or rain. A number of steps are required for the development of the vegetative propagules, including the degeneration of the thallus cortex, replication of green algal cells, and entanglement of fungal hyphae with the green algal cells. This steps lead to an increase in internal pressure which eventually breaks through the cortex. Continued growth leads to these granules being pushed upwards and out of the thallus surface.

Distribution and habitat

It has a wide distribution in Europe
Europe
Europe is, by convention, one of the world's seven continents. Comprising the westernmost peninsula of Eurasia, Europe is generally 'divided' from Asia to its east by the watershed divides of the Ural and Caucasus Mountains, the Ural River, the Caspian and Black Seas, and the waterways connecting...

, Asia
Asia
Asia is the world's largest and most populous continent, located primarily in the eastern and northern hemispheres. It covers 8.7% of the Earth's total surface area and with approximately 3.879 billion people, it hosts 60% of the world's current human population...

, North America
North America
North America is a continent wholly within the Northern Hemisphere and almost wholly within the Western Hemisphere. It is also considered a northern subcontinent of the Americas...

 and Africa
Africa
Africa is the world's second largest and second most populous continent, after Asia. At about 30.2 million km² including adjacent islands, it covers 6% of the Earth's total surface area and 20.4% of the total land area...

, preferring damp habitats with high rainfall, especially coast
Coast
A coastline or seashore is the area where land meets the sea or ocean. A precise line that can be called a coastline cannot be determined due to the dynamic nature of tides. The term "coastal zone" can be used instead, which is a spatial zone where interaction of the sea and land processes occurs...

al areas. It is the most widely distributed and most common Lobaria species in North America. Associated with old-growth forests, its presence and abundance may be used as an indicator of forest age, at least in the Interior Cedar-Hemlock biogeoclimatic zone in eastern British Columbia
British Columbia
British Columbia is the westernmost of Canada's provinces and is known for its natural beauty, as reflected in its Latin motto, Splendor sine occasu . Its name was chosen by Queen Victoria in 1858...

. It is also found in pasture-woodlands. It usually grows on the bark of broad-leaved trees such as oak
Oak
An oak is a tree or shrub in the genus Quercus , of which about 600 species exist. "Oak" may also appear in the names of species in related genera, notably Lithocarpus...

, beech
Beech
Beech is a genus of ten species of deciduous trees in the family Fagaceae, native to temperate Europe, Asia and North America.-Habit:...

 and maple
Maple
Acer is a genus of trees or shrubs commonly known as maple.Maples are variously classified in a family of their own, the Aceraceae, or together with the Hippocastanaceae included in the family Sapindaceae. Modern classifications, including the Angiosperm Phylogeny Group system, favour inclusion in...

 but will also grow on rocks. In the laboratory, L. pulmonaria has been grown on nylon
Nylon
Nylon is a generic designation for a family of synthetic polymers known generically as polyamides, first produced on February 28, 1935, by Wallace Carothers at DuPont's research facility at the DuPont Experimental Station...

 microfilaments.
Various environmental factors are thought to affect the distribution of L. pulmonaria, such as temperature, moisture (average humidity, rapidity and frequency of wet-dry cycles), sunlight exposure, and levels of air pollution. Attempts to quantitatively evaluate the contribution of these factors to lichen growth is difficult because differences in the original environment from which the lichen thalli are collected will greatly affect heat and desiccation tolerances.

Due to declining population, L. pulmonaria is considered to be rare or threatened in many parts of the world, especially in lowland areas of Europe. The decline has been attributed to industrial forestry
Forestry
Forestry is the interdisciplinary profession embracing the science, art, and craft of creating, managing, using, and conserving forests and associated resources in a sustainable manner to meet desired goals, needs, and values for human benefit. Forestry is practiced in plantations and natural stands...

 and air pollution
Air pollution
Air pollution is the introduction of chemicals, particulate matter, or biological materials that cause harm or discomfort to humans or other living organisms, or cause damage to the natural environment or built environment, into the atmosphere....

, particularly acid rain
Acid rain
Acid rain is a rain or any other form of precipitation that is unusually acidic, meaning that it possesses elevated levels of hydrogen ions . It can have harmful effects on plants, aquatic animals, and infrastructure. Acid rain is caused by emissions of carbon dioxide, sulfur dioxide and nitrogen...

. L. pulmonaria, like other lichens containing a blue-green algal component, are particularly susceptible to the effects of acid rain, because the subsequent decrease in pH
PH
In chemistry, pH is a measure of the acidity or basicity of an aqueous solution. Pure water is said to be neutral, with a pH close to 7.0 at . Solutions with a pH less than 7 are said to be acidic and solutions with a pH greater than 7 are basic or alkaline...

 reduces nitrogen fixation
Nitrogen fixation
Nitrogen fixation is the natural process, either biological or abiotic, by which nitrogen in the atmosphere is converted into ammonia . This process is essential for life because fixed nitrogen is required to biosynthesize the basic building blocks of life, e.g., nucleotides for DNA and RNA and...

 through inhibition of the algal nitrogenase
Nitrogenase
Nitrogenases are enzymes used by some organisms to fix atmospheric nitrogen gas . It is the only known family of enzymes that accomplish this process. Dinitrogen is quite inert because of the strength of its N-N triple bond...

 enzyme.

Chemical compounds

L. pulmonaria is known to contain a variety of acids common to lichens, such as stictic acid, desmethyl stictic acid, gyrophoric acid, tenuiorin, constictic acid, norstictic acid, peristictic acid, and methylnorstictic acid. These compounds, collectively known as depsidones, are known to be involved in defense against grazing
Grazing
Grazing generally describes a type of feeding, in which a herbivore feeds on plants , and also on other multicellular autotrophs...

 herbivores like lichen-feeding molluscs. It also contains the sugar alcohol
Sugar alcohol
A sugar alcohol is a hydrogenated form of carbohydrate, whose carbonyl group has been reduced to a primary or secondary hydroxyl group . Sugar alcohols have the general formula Hn+1H, whereas sugars have HnHCO...

s D-arabitol
Arabitol
Arabitol or arabinitol is a sugar alcohol. It can be formed by the reduction of either arabinose or lyxose. Some organic acid tests check for the presence of D-arabinitol, which may indicate overgrowth of intestinal parasites such as Candida albicans or other yeast/fungus species....

, volemitol
Volemitol
Volemitol is a naturally occurring seven-carbon sugar alcohol. It is a substance widely distributed in plants, red algae, fungi, mosses, and lichens. It was also found in lipopolysaccharides from E. coli. In certain higher plants, such as Primula, volemitol plays several important physiological roles...

, in addition to several carotenoid
Carotenoid
Carotenoids are tetraterpenoid organic pigments that are naturally occurring in the chloroplasts and chromoplasts of plants and some other photosynthetic organisms like algae, some bacteria, and some types of fungus. Carotenoids can be synthesized fats and other basic organic metabolic building...

s (total content > 10 mg/kg), such as alpha carotene, beta carotene, and beta cryptoxanthin
Cryptoxanthin
Cryptoxanthin is a natural carotenoid pigment. It has been isolated from a variety of sources including the petals and flowers of plants in the genus Physalis, orange rind, papaya, egg yolk, butter, apples, and bovine blood serum.-Chemistry:...

. The upper cortex of the lichen contains melanin
Melanin
Melanin is a pigment that is ubiquitous in nature, being found in most organisms . In animals melanin pigments are derivatives of the amino acid tyrosine. The most common form of biological melanin is eumelanin, a brown-black polymer of dihydroxyindole carboxylic acids, and their reduced forms...

s that screen UV
Ultraviolet
Ultraviolet light is electromagnetic radiation with a wavelength shorter than that of visible light, but longer than X-rays, in the range 10 nm to 400 nm, and energies from 3 eV to 124 eV...

 and PAR radiation
Photosynthetically active radiation
Photosynthetically active radiation, often abbreviated PAR, designates the spectral range of solar radiation from 400 to 700 nanometers that photosynthetic organisms are able to use in the process of photosynthesis. This spectral region corresponds more or less with the range of light visible to...

 from the photobiont. The synthesis of melanin pigments in the lichen increases in response to greater solar irradiation, and shade-adapted thalli are greenish-grey in the air-dry state, while sun-exposed thalli can be dark brown in color. This adaptation helps protect the photosymbiont D. reticulata, known to be relatively intolerant to high light levels.

Also known to be present are various steroid
Steroid
A steroid is a type of organic compound that contains a characteristic arrangement of four cycloalkane rings that are joined to each other. Examples of steroids include the dietary fat cholesterol, the sex hormones estradiol and testosterone, and the anti-inflammatory drug dexamethasone.The core...

s, namely ergosterol
Ergosterol
Ergosterol is a sterol found in fungi, and named for ergot, a common name for the members of the fungal genus Claviceps from which ergosterol was first isolated. Ergosterol does not occur in plant or animal cells...

, episterol
Episterol
Episterol is a sterol involved in the biosynthesis of steroids. Episterol is converted from 24-methylenelophenol. Episterol is converted to 5-dehydroepisterol by the enzyme lathosterol oxidase. Episterol is also known to be a precursor to ergosterol....

, fecosterol and lichesterol.

Medicinal

Its shape somewhat resembles the tissue inside lung
Lung
The lung is the essential respiration organ in many air-breathing animals, including most tetrapods, a few fish and a few snails. In mammals and the more complex life forms, the two lungs are located near the backbone on either side of the heart...

s and therefore it is thought to be a remedy for lung diseases based on the doctrine of signatures
Doctrine of signatures
The doctrine of signatures is a philosophy shared by herbalists from the time of Dioscurides and Galen. This doctrine states that herbs that resemble various parts of the body can be used to treat ailments of that part of the body. Examples include the plants liverwort; snakeroot, an antidote for...

. The lichen's common English names are derived from this association. Gerard's
John Gerard
John Gerard aka John Gerarde was an English herbalist notable for his herbal garden and botany writing. In 1597 he published a large and heavily illustrated "Herball, or Generall Historie of Plantes", which went on to be the most widely circulated botany book in English in the 17th century...

 book The Herball or General Historie of plants (1597) recommends L. pulmonaria as medicinally valuable. It is still used for asthma
Asthma
Asthma is the common chronic inflammatory disease of the airways characterized by variable and recurring symptoms, reversible airflow obstruction, and bronchospasm. Symptoms include wheezing, coughing, chest tightness, and shortness of breath...

, urinary incontinence
Urinary incontinence
Urinary incontinence is any involuntary leakage of urine. It is a common and distressing problem, which may have a profound impact on quality of life. Urinary incontinence almost always results from an underlying treatable medical condition but is under-reported to medical practitioners...

 and lack of appetite. In India
India
India , officially the Republic of India , is a country in South Asia. It is the seventh-largest country by geographical area, the second-most populous country with over 1.2 billion people, and the most populous democracy in the world...

 it is used as a traditional medicine to treat hemorrhages and eczema
Eczema
Eczema is a form of dermatitis, or inflammation of the epidermis . In England, an estimated 5.7 million or about one in every nine people have been diagnosed with the disease by a clinician at some point in their lives.The term eczema is broadly applied to a range of persistent skin conditions...

, and it is used as a remedy for coughing up blood by the Hesquiaht
Hesquiaht First Nation
The Hesquiaht First Nation is a Nuu-chah-nulth First Nations government based on the west coast of Vancouver Island in British Columbia, Canada. It is a member of the Nuu-chah-nulth Tribal Council.- Introduction :...

 in British Columbia
British Columbia
British Columbia is the westernmost of Canada's provinces and is known for its natural beauty, as reflected in its Latin motto, Splendor sine occasu . Its name was chosen by Queen Victoria in 1858...

, Canada
Canada
Canada is a North American country consisting of ten provinces and three territories. Located in the northern part of the continent, it extends from the Atlantic Ocean in the east to the Pacific Ocean in the west, and northward into the Arctic Ocean...

. An ethnophytotherapeutical survey of the high Molise
Molise
Molise is a region of Southern Italy, the second smallest of the regions. It was formerly part of the region of Abruzzi e Molise and now a separate entity...

 region in central-southern Italy
Italy
Italy , officially the Italian Republic languages]] under the European Charter for Regional or Minority Languages. In each of these, Italy's official name is as follows:;;;;;;;;), is a unitary parliamentary republic in South-Central Europe. To the north it borders France, Switzerland, Austria and...

 revealed that L. pulmonaria is used as an antiseptic
Antiseptic
Antiseptics are antimicrobial substances that are applied to living tissue/skin to reduce the possibility of infection, sepsis, or putrefaction...

, and is rubbed on wounds.

A hot-water extract
Extract
An extract is a substance made by extracting a part of a raw material, often by using a solvent such as ethanol or water. Extracts may be sold as tinctures or in powder form....

 prepared using this species has been shown to have anti-inflammatory
Anti-inflammatory
Anti-inflammatory refers to the property of a substance or treatment that reduces inflammation. Anti-inflammatory drugs make up about half of analgesics, remedying pain by reducing inflammation as opposed to opioids, which affect the central nervous system....

 and ulcer
Peptic ulcer
A peptic ulcer, also known as PUD or peptic ulcer disease, is the most common ulcer of an area of the gastrointestinal tract that is usually acidic and thus extremely painful. It is defined as mucosal erosions equal to or greater than 0.5 cm...

-preventing activities. Also, methanol extracts were shown to have a protective effect on the gastrointestinal system of rats, possibly by reducing oxidative stress
Oxidative stress
Oxidative stress represents an imbalance between the production and manifestation of reactive oxygen species and a biological system's ability to readily detoxify the reactive intermediates or to repair the resulting damage...

 and reducing the inflammatory
Inflammation
Inflammation is part of the complex biological response of vascular tissues to harmful stimuli, such as pathogens, damaged cells, or irritants. Inflammation is a protective attempt by the organism to remove the injurious stimuli and to initiate the healing process...

 effects of neutrophils. Furthermore, methanol extracts also have potent antioxidative activity and reducing power, probably due to the presence of phenolic compounds.

Other uses

L. pulmonaria has also been used to produce an orange dye
Dye
A dye is a colored substance that has an affinity to the substrate to which it is being applied. The dye is generally applied in an aqueous solution, and requires a mordant to improve the fastness of the dye on the fiber....

 for wool, in the tanning
Tanning
Tanning is the making of leather from the skins of animals which does not easily decompose. Traditionally, tanning used tannin, an acidic chemical compound from which the tanning process draws its name . Coloring may occur during tanning...

 of leather
Leather
Leather is a durable and flexible material created via the tanning of putrescible animal rawhide and skin, primarily cattlehide. It can be produced through different manufacturing processes, ranging from cottage industry to heavy industry.-Forms:...

, in the manufacture of perfume
Perfume
Perfume is a mixture of fragrant essential oils and/or aroma compounds, fixatives, and solvents used to give the human body, animals, objects, and living spaces "a pleasant scent"...

s and as an ingredient in brewing
Brewing
Brewing is the production of beer through steeping a starch source in water and then fermenting with yeast. Brewing has taken place since around the 6th millennium BCE, and archeological evidence suggests that this technique was used in ancient Egypt...

.

External links

The source of this article is wikipedia, the free encyclopedia.  The text of this article is licensed under the GFDL.
 
x
OK