Geosiphon
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''Geosiphon'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of
fungus A fungus ( : fungi or funguses) is any member of the 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, separately fr ...
in the
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
Geosiphonaceae. The genus is
monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispe ...
, containing the single species ''Geosiphon pyriformis'', first described by Kützing in 1849 as ''Botrydium pyriforme''. In 1915, Von Wettstein characterized ''Geosiphon pyriforme'' as a
multinucleate Multinucleate cells (also known as multinucleated or polynuclear cells) are eukaryotic cells that have more than one nucleus per cell, i.e., multiple nuclei share one common cytoplasm. Mitosis in multinucleate cells can occur either in a coordina ...
alga containing
endosymbiotic An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship. (The term endosymbiosis is from the Greek: ἔνδον ''endon'' "within ...
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, bl ...
, although he also noted the presence of
chitin Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...
, a component of fungal cell walls. In 1933, Knapp was the first to suggest the fungal origin of the species and described it as a
lichen A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.arbuscular mycorrhiza An arbuscular mycorrhiza (AM) (plural ''mycorrhizae'', a.k.a. ''endomycorrhiza'') is a type of mycorrhiza in which the symbiont fungus (''AM fungi'', or AMF) penetrates the cortical cells of the roots of a vascular plant forming arbuscules. ( ...
.


Life cycle

''Geosiphon pyriformis'' is known for being the
symbiont Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or parasi ...
of ''
Nostoc ''Nostoc'', also known as star jelly, troll’s butter, spit of moon, fallen star, witch's butter (not to be confused with the fungi commonly known as witches' butter), and witch’s jelly, is the most common genus of cyanobacteria found in var ...
''. The ''Geosiphon''-''
Nostoc ''Nostoc'', also known as star jelly, troll’s butter, spit of moon, fallen star, witch's butter (not to be confused with the fungi commonly known as witches' butter), and witch’s jelly, is the most common genus of cyanobacteria found in var ...
'' symbiosis, as by modern definitions, is not a
lichen A lichen ( , ) is a composite organism that arises from algae or cyanobacteria living among filaments of multiple fungi species in a mutualistic relationship.arbuscular mycorrhiza An arbuscular mycorrhiza (AM) (plural ''mycorrhizae'', a.k.a. ''endomycorrhiza'') is a type of mycorrhiza in which the symbiont fungus (''AM fungi'', or AMF) penetrates the cortical cells of the roots of a vascular plant forming arbuscules. ( ...
symbioses than to lichens. The ''Geosiphon''-''
Nostoc ''Nostoc'', also known as star jelly, troll’s butter, spit of moon, fallen star, witch's butter (not to be confused with the fungi commonly known as witches' butter), and witch’s jelly, is the most common genus of cyanobacteria found in var ...
'' symbiosis is the only known fungal endosymbiosis with cyanobacteria and is characterised by a "siphonal bladder" that is made of a swollen fungal hypha, 0.5–2 mm in size and growing on the soil surface. The upper 2/3 of the "bladder" contains the ''Nostoc'' filaments and Nostoc
heterocyst Heterocysts or heterocytes are specialized nitrogen-fixing cells formed during nitrogen starvation by some filamentous cyanobacteria, such as '' Nostoc punctiforme'', ''Cylindrospermum stagnale'', and ''Anabaena sphaerica''. They fix nitrogen fr ...
s, the lower 1/3 is filled with
lipid Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids in ...
droplets.


Reproduction

The fungal
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, ...
s are 250 
micrometre The micrometre (American and British English spelling differences#-re, -er, international spelling as used by the International Bureau of Weights and Measures; SI symbol: μm) or micrometer (American and British English spelling differences# ...
s in diameter, formed at the end of one
hypha A hypha (; ) is a long, branching, filamentous structure of a fungus, oomycete, or actinobacterium. In most fungi, hyphae are the main mode of vegetative growth, and are collectively called a mycelium. Structure A hypha consists of one or ...
, or intercalarly (within a hyphae) and resemble those of other members of the Glomeromycota, with a particular resemblance to spores of others in the
polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of conver ...
genus Glomus.


Somatic structure

''Geosiphon'' has a specialised bladder used to harbour
cyanobacteria Cyanobacteria (), also known as Cyanophyta, are a phylum of gram-negative bacteria that obtain energy via photosynthesis. The name ''cyanobacteria'' refers to their color (), which similarly forms the basis of cyanobacteria's common name, bl ...
, usually '' Nostoc punctiforme'', although other ''Nostoc'' species are known to be incorporated by the fungus. ''Geosiphon'' bladders are photosynthetically active, and are also capable of fixing nitrogen from the atmosphere. The walls of the bladder have a small pore radius (approximately 0.5  nm), which make it act as an
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 ...
barrier. This limits the free exchange of nutrients (like sugars) with the environment and increases the need to derive these carbon sources from internal sources. It has been suggested that the organization of the symbiotic interface between the fungus and ''Nostoc'' in ''G. pyriforme'' is homologous to the symbiotic interface between plant and fungus in arbuscular mycorrhiza in terms of thickness, chitin content, and ultrastructure of layers.Schuessler A, Bonfante P, Schnepf E, Mollenhauer D, Kluge, M. (1996). Characterization of the ''Geosiphon pyriforme'' symbiosome by affinity techniques: Confocal laser scanning microscopy (CLSM) and electron microscopy. ''Protoplasma'' 190(1–2): 53–67. ''G. pyriformis'' however, is not known to form arbuscular mycorrhiza. The bladder structure may be considered to be equivalent to a
symbiosome A symbiosome is a specialised compartment in a host cell that houses an endosymbiont in a symbiotic relationship. The term was first used in 1983 to describe the vacuole structure in the symbiosis between the animal host the '' Hydra'', and the en ...
, a specialized membraned structure in some plants and animals that forms a structural and functional interface between the host and its symbiont counterpart. The symbiosome may be divided into three functional areas: *the symbiosome membrane forms the border of the symbiosome. It is thought to be formed from the
plasma membrane The cell membrane (also known as the plasma membrane (PM) or cytoplasmic membrane, and historically referred to as the plasmalemma) is a biological membrane that separates and protects the interior of all cells from the outside environment (t ...
of the fungus by
invagination Invagination is the process of a surface folding in on itself to form a cavity, pouch or tube. In developmental biology, invagination is a mechanism that takes place during gastrulation. This mechanism or cell movement happens mostly in the vegeta ...
during the uptake of the symbiont.Schnepf E. (1964). Zur Feinstruktur von ''Geosiphon pyriforme''. ''Arch Micobiol''. 49: 112–31. *the symbiosome space, located between the symbiosome membrane and the endosymbiotic ''Nostoc'' cyanobacteria. *the area containing the cyanobacteria. The space between the symbiosome membrane and the ''Nostoc'' cell wall contains a 30–40 nm thick layer of carbohydrate material such as
mannose Mannose is a sugar monomer of the aldohexose series of carbohydrates. It is a C-2 epimer of glucose. Mannose is important in human metabolism, especially in the glycosylation of certain proteins. Several congenital disorders of glycosylat ...
,
fucose Fucose is a hexose deoxy sugar with the chemical formula C6H12O5. It is found on ''N''-linked glycans on the mammalian, insect and plant cell surface. Fucose is the fundamental sub-unit of the seaweed polysaccharide fucoidan. The α(1→3) li ...
, GalNAc,
sialic acid Sialic acids are a class of alpha-keto acid sugars with a nine-carbon backbone. The term "sialic acid" (from the Greek for saliva, - ''síalon'') was first introduced by Swedish biochemist Gunnar Blix in 1952. The most common member of this ...
, and
galactose Galactose (, '' galacto-'' + ''-ose'', "milk sugar"), sometimes abbreviated Gal, is a monosaccharide sugar that is about as sweet as glucose, and about 65% as sweet as sucrose. It is an aldohexose and a C-4 epimer of glucose. A galactose molecu ...
.


References


External links


The ''Geosiphon pyriformis'' symbiosis - fungus 'eats' cyanobacterium
Schuessler lab
Glomeromycota phylogeny
{{Taxonbar, from=Q144388, from2=Q18523806, from3=Q10504225 Glomeromycota Monotypic fungi genera Taxa named by Friedrich Traugott Kützing