Black Bread Mold
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''Rhizopus stolonifer'' is commonly known as white bread mold. It is a member of ''
Zygomycota Zygomycota, or zygote fungi, is a former division or phylum of the kingdom Fungi. The members are now part of two phyla: the Mucoromycota and Zoopagomycota. Approximately 1060 species are known. They are mostly terrestrial in habitat, living ...
'' and considered the most important species in the genus ''
Rhizopus ''Rhizopus'' is a genus of common saprophytic fungi on plants and specialized parasites on animals. They are found in a wide variety of organic substances, including "mature fruits and vegetables", jellies, syrups, leather, bread, peanuts, and ...
''. It is one of the most common fungi in the world and has a global distribution although it is most commonly found in tropical and subtropical regions. It is a common agent of decomposition of stored foods. Like other members of the genus ''Rhizopus'', ''R. stolonifer'' grows rapidly, mostly in indoor environments.


History

This fungus was first discovered by the German scientist
Christian Gottfried Ehrenberg Christian Gottfried Ehrenberg (19 April 1795 – 27 June 1876) was a German Natural history, naturalist, zoologist, comparative anatomist, geologist, and microscopy, microscopist. Ehrenberg was an Evangelicalism, evangelist and was considered to ...
in 1818 as ''Rhizopus nigricans''. The name was changed in 1902 to ''Rhizopus stolonifer'' by the French mycologist J. P. Vuillemin.


Habitat and ecology

''Rhizopus stolonifer'' is a worldwide distributed species. It is found on all types of mouldy materials. It is often one of the first
mold A mold () or mould () is one of the structures certain fungi can form. The dust-like, colored appearance of molds is due to the formation of spores containing fungal secondary metabolites. The spores are the dispersal units of the fungi. Not ...
s to appear on stale bread. It can exist in the soil as well as in the air. A variety of natural substrata are colonized by this species because ''R. stolonifer'' can tolerate broad variations in the concentration of essential nutrients and can use carbon and nitrogen combined in diverse forms. In the laboratory, this fungus grows well on different media, including those that contain
ammonium salt The ammonium cation is a positively-charged polyatomic ion with the chemical formula or . It is formed by the protonation of ammonia (). Ammonium is also a general name for positively charged or protonated substituted amines and quaternary ...
s or amino compounds. However, ''R. stolonifer'' will not grow on Czapek's agar because it cannot utilize nitrogen in the form of nitrate. Rhizopus lives in hyphae and matured spores.


Growth and physiology

This species is known as a
saprotroph Saprotrophic nutrition or lysotrophic nutrition is a process of chemoheterotrophic extracellular digestion involved in the processing of decayed (dead or waste) organic matter. It occurs in saprotrophs, and is most often associated with fungi ( ...
and plays an important role in the early colonization of substrata in soil. Nonetheless, it can also behave as a
parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...
of plant tissues causing a rot of vegetables and fruits. Like other species of ''Rhizopus'', ''R. stolonifer'' grows rapidly and spreads by means of the stolons. The stolons provide an aerial structure for the growth of the
mycelium Mycelium (plural mycelia) is a root-like structure of a fungus consisting of a mass of branching, thread-like hyphae. Fungal colonies composed of mycelium are found in and on soil and many other substrates. A typical single spore germinates ...
and the occupation of large areas. They can climb vertically as well as horizontally. Sporangiophores of ''R. stolonifer'' can be up to 2.5 mm long and about 20 μm in diameter. The spores are shaped differently depending on the available nutrients. They can be ovate, polygonal or angular. The optimal temperature for growth varies between 25 and 30 °C. The thermal death point, which is defined as the lowest temperature that can kill all cells in ten minutes, is 60 °C. ''Rhizopus stolonifer'' can grow in acidic environments with a pH of as low as 2.2. The pH range can vary from 2.2 to 9.6.
Ultraviolet irradiation Ultraviolet (UV) is a form of electromagnetic radiation with wavelength from 10 nm (with a corresponding frequency around 30  PHz) to 400 nm (750  THz), shorter than that of visible light, but longer than X-rays. UV radiation i ...
can delay spore germination.


Reproduction

''Rhizopus stolonifer'' can reproduce asexually and sexually. It is a
heterothallic Heterothallic species have sexes that reside in different individuals. The term is applied particularly to distinguish heterothallic fungi, which require two compatible partners to produce sexual spores, from homothallic ones, which are capable ...
species. Sexual reproduction occurs when compatible mating strains are paired, ultimately giving rise to
zygospore A zygospore is a diploid reproductive stage in the life cycle of many fungi and protists. Zygospores are created by the nuclear fusion of haploid cells. In fungi, zygospores are formed in zygosporangia after the fusion of specialized budding struc ...
s. The sporangiophore contains both '+' and '−' mating type strains.
Meiosis Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately r ...
is delayed until the germination of the zygospores. The gametogenia often differ in size, regardless of mating type. This difference in size is not due to sex but presumably due to nutrition.


Disease and prevention

The disease caused by this fungus occurs mainly on ripe fruits, such as strawberries, melon and peach, which are more susceptible to wounds and have a higher sugar content. After a couple of days, the infected fruits become soft and release juices with an acidic odour. When the humidity and temperature are favourable, the mycelial growth occurs rapidly at the surface of the infected fruit and the disease causes the development of long mycelial stolons with black
sporangia A sporangium (; from Late Latin, ) is an enclosure in which spores are formed. It can be composed of a single cell or can be multicellular. Virtually all plants, fungi, and many other lineages form sporangia at some point in their life cyc ...
and spores. When the fungus germinates, it produces different kinds of
esterase An esterase is a hydrolase enzyme that splits esters into an acid and an alcohol in a chemical reaction with water called hydrolysis. A wide range of different esterases exist that differ in their substrate specificity, their protein structure ...
s, including
cutinase The enzyme cutinase (systematic name: cutin hydrolase,''EC 3.1.1.74 is a member of the hydrolase family. It catalyzes the following reaction: R1COOR2 + H2O -> R1COOH + R2OH In biological systems, the reactant carboxylic ester is a constituent of ...
, which help the fungus to penetrate the plant cell wall. The disease can also affect other adjacent healthy fruits when distributed by wind or insect activity. Some species of ''Syncephalis'' can reduce the asexual reproduction of ''R. stolonifer'' and therefore may delay or even prevent the post-harvest disease caused by this fungus. Fengycin, which is an anti-fungal complex, also induces the fungal cell death via necrosis and apoptosis. The treatment of sweet potatoes with sodium orthophenyl phenol (Stopmold B) and dichloran (Botran W) have effectively reduced storage rot. ''Rhizopus stolonifer'' is an
opportunistic Opportunism is the practice of taking advantage of circumstances – with little regard for principles or with what the consequences are for others. Opportunist actions are expedient actions guided primarily by self-interested motives. The term ...
agent of disease and hence will only cause infection in people with a weakened immunity.
Zygomycosis Zygomycosis is the broadest term to refer to infections caused by ''bread mold fungi'' of the zygomycota phylum. However, because zygomycota has been identified as polyphyletic, and is not included in modern fungal classification systems, the dise ...
is the main disease that might be caused by this fungus in humans and while it is not entirely understood yet, this disease is very dangerous and can be fatal. The action of smelling spoiled food may be a source of inhalation exposure to the mold.


Importance

''Rhizopus stolonifer'' is important economically as an agent of post-harvest storage decay. While ''
Saccharomyces cerevisiae ''Saccharomyces cerevisiae'' () (brewer's yeast or baker's yeast) is a species of yeast (single-celled fungus microorganisms). The species has been instrumental in winemaking, baking, and brewing since ancient times. It is believed to have b ...
'' is the most important source of industrial alcohol, ''R. stolonifer'' and other species of ''Rhizopus'' also produce
ethyl alcohol Ethanol (abbr. EtOH; also called ethyl alcohol, grain alcohol, drinking alcohol, or simply alcohol) is an organic compound. It is an alcohol with the chemical formula . Its formula can be also written as or (an ethyl group linked to a hyd ...
which is the most important fermentation product. ''Rhizopus stolonifer'' is also used commercially to manufacture
fumaric acid Fumaric acid is an organic compound with the formula HO2CCH=CHCO2H. A white solid, fumaric acid occurs widely in nature. It has a fruit-like taste and has been used as a food additive. Its E number is E297. The salts and esters are known as fu ...
and
lactic acid Lactic acid is an organic acid. It has a molecular formula . It is white in the solid state and it is miscible with water. When in the dissolved state, it forms a colorless solution. Production includes both artificial synthesis as well as nat ...
of high purity. The presence of
zinc Zinc is a chemical element with the symbol Zn and atomic number 30. Zinc is a slightly brittle metal at room temperature and has a shiny-greyish appearance when oxidation is removed. It is the first element in group 12 (IIB) of the periodi ...
reduces the amount of fumaric acid produced by the fungus.


References

{{Taxonbar, from=Q1500736 Mucoraceae Fungi described in 1818