Lichen-Forming Fungi

Division: Ascomycota

Division: Basidiomycota

Lichens are not a natural taxonomic group. Lichens represent a fungal “life style.”

What are lichens?
• Ascomycete and basidiomycete fungi with an obligate symbiotic mode of nutrition with a photosynthetic partner • Unlike other symbiotic relationships, the lichen thallus (composite thallus) has a unique morphology unlike either partner growing alone • About 20,000 species known worldwide

Composite Thallus . Mycobiont.Photobiont.

The Mycobiont • Most lichen species are ascomycetes from the Series Discomycetes (apothecial forms) and Pyrenomycetes (perithecial forms) • A few are basidiomycetes related to mushrooms and bracket fungi .

Isolated Mycobiont Composite thallus .

or erythritol depending on the algal genus involved . sorbitol. the mycobiont receives glucose – in those with green algal partners. the mycobiont receives a sugar alcohol such as ribitol.Fungal Nutrition • Photosynthetic partner is the source of nutrients (carbon) for fungal metabolism – in those with cyanobacterial partners.

Hyalococcus and Trentepohlia • Some partners are cyanobacteria – Common genera include Nostoc and Stigonema • About 40 genera occur as symbionts .The Photobiont (phycobiont) • Most lichen partners are green algae (Division Chlorophyta) – Common genera include Trebouxia.

Lichen Algae .

Trebouxia (Chlorophyta) Trebuoxia is one of the most common photobionts in lichens .

crustose – 2 cm/yr.000 years old! . more for fruticose • Some lichens are estimated to be over 3. foliose.Lichen Growth and Longevity • Lichens grow VERY SLOWLY with some species only growing mm in hundreds of years (extreme cold environments) – 1 mm/yr.

The Nature of the Symbiosis • The mycobiont is an obligate symbiont and usually can only associate with one algal genus (but more than one species) • The algal genera are often found freeliving and may occur in hundreds of different lichens • The name of a lichen is the name of the fungal partner .

) • Most lichens are probably an example of controlled parasitism • Other species may represent a balance closer to mutualism • “dual nature” of the lichens was not proposed until 1867 by S. Schwendener (and at first rejected by many biologists) .The Nature of the Symbiosis (cont.

Mycobiont-Photobiont Interface .

Foliose. Squamulose. Umbilicate.Growth Forms: Crustose. and Fruticose .

but poorly understood taxonomically Lecanora .Crustose Lichens • Thought to be the most primitive (ancient) type • Only an upper cortex and medulla penetrating the substrate • Many taxa.

) Acarospora .Crustose Lichen (cont.

Squamulose Lichens • Aggregation of small scale-like pieces called squamules • Often with more tissue differentiation than crustose forms Psora .

Foliose Lichens • Lobed thalli with upper and lower cortex present • Usually with attachment rhizines • Common and often conspicuous Parmotrema .

Foliose Lichen Section rhizine .

Foliose Thallus Section Upper Cortex Medulla Lower Cortex .

) Melanelia Peltigera .Foliose Lichens (cont.

Umbilicate Lichens • Thallus attached by a single strand called the umbilicus • This growth form has evolved in many different groups Umbilicaria .

Fruticose Lichens • Outer cortex surrounds algal layer and inner medulla • Pendulous or erect “shrubby” forms Letharia .

) Usnea Teloschistes .Fruticose Lichens (cont.

Fruticose Lichens (cont.) Cladina Cetraria .

) Calicium polyporeum Cladonia pleurota .Fruticose Lichens (cont.

squarrose.Lichen Morphology • Cilia – various hair-like extensions from the margins of lobes • Rhizines – root-like attachment organs on the underside (simple. dichotomous) .

Marginal Cilia Parmotrema reticulatum Cilia on apothecia .

Bulbate Cilia Bulbothrix .

Rhizines (dichotomous) Parmotrema sp. with yellow medulla .

Section of Rhizine .

Lichen Morphology (cont.) • Tomentum – cottony. felt-like mats of fungal material on the upper or lower surface .

) • Cyphellae and Pseudocyphellae – “air pores” on upper or lower surface probably for gas exchange Cyphella on lower surface of Sticta .Lichen Morphology (cont.

Pseudocyphellae Punctelia .

Lichen Morphology (cont.) • Soredia – asexual. granular propagules composed of fungal and algal partners – produced in soralia in various shapes and in various positions on lichen thalli .

Yellow Soredia Pseudocyphellaria .Marginal.

Soredia and Marginal Cilia Parmotrema .

Lichen Morphology (cont.) • Isidia – finger-like outgrowth of the upper cortex or outer cortex (fruticose lichens) – can function (like soredia) as asexual reproductive fragments Bulbothrix .

) Isidia on Xanthoparmelia Isidia and cilia on Parmotrema .Isidia (cont.

Ascocarp Structure • Standard structures found in most apothecia (in a typical foliose lichen) .

Lichen Thallus Section (with apothecium) Physcia .

Sexual Reproductive Structures • LECANORINE APOTHECIA – apothecium has a thalloid exciple with algal cells – exciple is same color as the thallus and different from the disc Lecanora .

Lecanorine Apothecia Haematomma Hypogymnia .

Sexual Reproductive Structures (cont.) • LECIDEINE APOTHECIA – margin of the apothecium is a proper exciple with no algal cells – disc and exciple are same color Porpidia .

Sexual Reproductive Structures


– an elongated or branched apothecium – members commonly known as “graphids”


Sexual Reproductive Structures


– bottle-like ascocarps of members of the Series Pyrenomycetes

Reproductive Structures (cont.)
– Some lichen-forming taxa also come from the Series Loculoascomycetes (ascolocular taxa)


freeliving ascomycetes • The major difference is the necessity for encountering. and forming a symbiotic composite thallus with the proper photosynthetic partner .Life Cycle • Life cycle of most lichen-forming fungi is the same as that of the saprophytic. recognizing.

isidia) rather than by production of huge numbers of conidia • Some taxa produce pycnidia with microconidia which (theoretically) could also reproduce a lichen • These microconidia could also function as a male gamete in a sexual process .Life Cycle (cont.) • Asexual reproduction is frequently by fragmentation (soredia.

they are deposited externally on the hyphae of the cortex and/or medulla .Lichen Chemistry • Lichens produce a large number (over 500) of unique secondary metabolites often called lichen acids • Produced by the mycobiont.

and other fabrics (crimson-purple Roman togas of the ancient world) • Also used to prepare the original “litmus paper” and still today in the perfume industry (“moss and “leather” fragrances) .) • Historically these substances have been used to manufacture dyes for wool (“Harris Tweed” of Scottish fame).Lichen Chemistry (cont.

Dyes from Arctic Lichens .

Wool Dyeing in Scotland .

Litmus Production in Europe .

Perfume Manufacture Collection of “oak moss” and “tree moss” (species of Pseudevernia and Evernia) .

especially against gram+ bacteria • Usnic acid used in some topical ointments in Europe and Asia .Lichen Chemistry (cont.) • Some substances have antimicrobial effects.

depsidones. aliphatic acids (majority of lichen compounds) .Lichen Chemistry (cont.) • Lichen acids are derived from three chemical pathways: – shikimic acid pathway • pulvinic acid derivatives (yellow pigments) – mevalonic acid pathway • terpenes – acetyl-polymalonyl pathway • depsides. anthraquinones. usnic acid. xanthones.

Lichen Acids a lichen depside a lichen depsidone .

) A yellow-green pigment common in the lichen cortex .Lichen Acids (cont.

Identification of Lichen Acids • • • • Classic Spot Tests Microcrystal Tests Paper and Thin Layer Chromatography High Performance Liquid Chromatography (HPLC) .

CLASSIC SPOT TESTS – Initial discovery and use was by William Nylander in Europe in the 1860s – devised the thallus “spot tests” still used today: K (KOH). Asahina in the 1930s – techniques still useful and commonly used today . C (“Clorox”). KC (combination) – Pd (para-phenylenediamine) test was added by A.

Spot Test “K+ red” test on lichen medulla Xanthoparmelia .

MICROCRYSTAL TESTS – Introduced in the 1930s and 1940s by Y. Asahina of Japan – More accurate than spot tests .

CHROMATOGRAPHY – The 1950s and 1960s saw the introduction of paper and thin layer chromatography for a new level of accuracy in identifying lichen products – Chicita Culberson (Duke University) introduced standardized techniques which are still used routinely today .

Thin Layer Chromatography (cont.) .

HPLC – For serious lichen chemists. HPLC (high performance liquid chromatography) has proven to be the best way to find small traces of chemicals and to quantify results of chemical studies – New substances are still being discovered every year and their structures determined .

Chemical Systematics .

Biochemical Systematics • Do differences in chemical composition mean two individuals belong to different species? • Do other observations (ecological. geographical) correlate with the observed chemical differences? . morphological.

) Distribution of chemical strains of Pseudevernia furfuracea Two species or one? Varieties? Subspecies? Pseudevernia .Biochemical Systematics (cont.

Biochemical Systematics (cont.)

T. subuliformis vs. T. vermicularis

Biochemical Systematics

Adaptive Role of Lichen Acids • Resistance to decay • Release of carbohydrates from the photobiont • Defense against predation • UV protection

tardigrades. both physical and chemical weathering • Soil binding • Nitrogen fixation (cyanobacterial photobionts) . caribou and reindeer) • Soil formation (pedogenesis).Ecosystem Roles • Primary producers and food source (arthropods.

Ecosystem Roles (cont.) .

Distribution and Ecology • Lichen occur on all continents in terrestrial systems from the Arctic to the tropics • Only a very few species are known to be “aquatic” .

Distribution and Ecology (cont.) • Lichens occur on many substrates – rocks (saxicolous) of both acidic and basic chemistry (distinct floras on each) – bark and wood (corticolous and lignicolous) with often distinctions between hardwoods and conifers – soil (terricolous) and plant debris .

roof shingles etc……. stained glass.Distribution and Ecology (cont.) – on other lichens (lichenicolous) – on leaves of many tropical plants (foliicolous) – even inside rocks in Antarctica (cryptoendolithic lichens) – also found on man-made substrates: concrete. ancient canon balls. ..

Distribution and Ecology (cont.) Lichen damage to stained glass in France .

Foliicolous Lichens .

Distribution and Ecology (cont.) • Even on insects! .

Cu. Hg. and Co . nitrous oxides. ozone.Response to Pollution • Lichens are extremely sensitive to air pollutants including SO2 and acid rain. Ni. Zn. Cr. and PAN • Lichen growth can also be negatively effected by high levels of “heavy metal” pollution from elements such as Cd.

Pollution (cont.) • Because of their sensitivity pollutants. lichens serve as “early warning” organisms • Now lichen populations are frequently surveyed as part of pollution monitoring projects around the world .

Air Pollution and Lichens .

000 school children in England and Scotland .“Our Mucky Air” Prepared one summer by 15.

because of their poikilohydric nature (unable to regulate uptake of minerals or water) • Lichens act as a depositional reservoir especially for elements from nuclear testing such as 90Sr and 137Cs . in part.Radionuclide Accumulation • Lichens also are known accumulators of various elements.

) • These carcinogens/mutagens have been traced to humans populations through the Arctic lichens…. Fallout Lichens Caribou Eskimos .Radionuclide Accumulation (cont.

Lichenometry • Use of lichen growth rates to “date” rock surfaces • Used in geology and anthropology On Easter Island .

Lichenometry (cont.) • Dating of rock surfaces by geomorphologists is often done using the lichen genus Rhizocarpon .

helsinki.html • Lichenolgical Oragnizations – .uk/bto/microbes/ – http://www.html http://teaching.abls.WWW Links • Lichens in General – – – – http://www.htm#M .html • Lichen Illustrations (Line Drawings) – Links ( • Lichens and Air Pollution – – – – http://www.htm

htm • Internet Directory (Lower Plants) – Links (cont.unibayreuth.html .) • Foliicolous Lichens – http://www.

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