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{{Short description|Species of fungus}}
{{Speciesbox
{{Speciesbox
| image = Rhizopus_stolonifer4.JPG
| image = Rhizopus_stolonifer4.JPG
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| species = stolonifer
| species = stolonifer
| authority = Vuillemin (1902)
| authority = Vuillemin (1902)
| synonyms = *''Rhizopus nigricans'' <small> Ehrenberg (1820)</small>
| synonyms = *''Rhizopus nigricans'' <small>Ehrenberg (1820)</small>
*''Mucor stolonifer'' <small> Ehrenberg (1818)</small>
*''Mucor stolonifer'' <small>Ehrenberg (1818)</small>
*Rhizopus artocarpi var. luxurians <small>Schroet (1886)</small>
*Rhizopus artocarpi var. luxurians <small>Schroet (1886)</small>
*Mucor niger <small> Geodoelst (1902) </small>
*Mucor niger <small>Geodoelst (1902) </small>
*Rhizopus niger <small> Ciaglinski & Hewelke (1893)</small>
*Rhizopus niger <small>Ciaglinski & Hewelke (1893)</small>
}}
}}


'''''Rhizopus stolonifer''''' is commonly known as '''bear bread mold'''.<ref name = Ellis/> It is a member of ''[[Zygomycota]]'' and considered the most important species in the genus ''[[Rhizopus]]''.<ref name = Webster2007/> 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.<ref name = Moore-Landecker1972/> It is a common agent of decomposition of stored foods.<ref name = Alexopoulos1952/> Like other members of the genus ''Rhizopus'', ''R. stolonifer'' grows rapidly, mostly in indoor environments.<ref name = Benny2016/>
'''''Rhizopus stolonifer''''' is commonly known as '''black bread mold'''.<ref name = Ellis/> It is a member of ''[[Zygomycota]]'' and considered the most important species in the genus ''[[Rhizopus]]''.<ref name = Webster2007/> 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.<ref name = Moore-Landecker1972/> It is a common agent of decomposition of stored foods.<ref name = Alexopoulos1952/> Like other members of the genus ''Rhizopus'', ''R. stolonifer'' grows rapidly, mostly in indoor environments.<ref name = Benny2016/>


==History==
==History==
Line 18: Line 19:


==Habitat and ecology==
==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]]s to appear on stale bread.<ref name = onions1981/> 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.<ref name = onions1981/>
''Rhizopus stolonifer'' has a [[cosmopolitan distribution]] and is found on all types of mouldy materials. It is often one of the first [[Mold (fungus)|mold]]s to appear on stale bread.<ref name = onions1981/> 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.<ref name = onions1981/>


In the laboratory, this fungus grows well on different media, including those that contain [[ammonium salt]]s or amino compounds.<ref name = onions1981/> However, ''R. stolonifer'' will not grow on [[Czapek medium|Czapek’s agar]] (CZA) because it cannot utilize nitrogen in the form of nitrate.<ref name = onions1981/>
In the laboratory, this fungus grows well on different media, including those that contain [[ammonium salt]]s or amino compounds.<ref name = onions1981/> However, ''R. stolonifer'' will not grow on [[Czapek medium|Czapek's agar]] because it cannot utilize nitrogen in the form of nitrate.<ref name = onions1981/> Rhizopus lives in [[Hypha|hyphae]] and matured spores.


== Growth and physiology ==
== Growth and physiology ==
[[File:Rhizopus_stolonifer2.jpg|thumb|left|200px|Gem-shaped sporangiospores of ''R. stolonifer'' showing characteristic striations.]]This species is known as a [[saprotroph]] and plays an important role in the early colonization of substrata in soil. Nonetheless, it can also behave as a [[Parasitism|parasite]] of plant tissues causing a rot of vegetables and fruits.<ref name = Webster2007/> Like other species of ''Rhizopus'', ''R. stolonifer'' grows rapidly and spreads by means of the [[stolon]]s.<ref name = Webster2007/> The stolons provide an aerial structure for the growth of the [[mycelium]] and the occupation of large areas. They can climb vertically as well as horizontally.<ref name = Vanbreuseghem1965/> Sporangiophores of ''R. stolonifer'' can be up to 2.5&nbsp;mm long and about 20&nbsp;μm in diameter.<ref name = onions1981/> The [[spore]]s are shaped differently depending on the available nutrients. They can be ovate, polygonal or angular.<ref name = onions1981/> The optimal temperature for growth varies between 25-30&nbsp;°C.<ref name = Vanbreuseghem1965/> The thermal death point, which is defined as the lowest temperature that can kill all cells in ten minutes, is 60&nbsp;°C.<ref name = Vanbreuseghem1965/> ''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.<ref name = Skinner1930/> [[Ultraviolet irradiation]] can delay spore [[germination]].<ref name = Ainsworth1965/>
[[File:Rhizopus_stolonifer2.jpg|thumb|left|200px|Gem-shaped sporangiospores of ''R. stolonifer'' showing characteristic striations.]]This species is known as a [[saprotroph]] and plays an important role in the early colonization of substrata in soil. Nonetheless, it can also behave as a [[Parasitism|parasite]] of plant tissues causing a rot of vegetables and fruits.<ref name = Webster2007/> Like other species of ''Rhizopus'', ''R. stolonifer'' grows rapidly and spreads by means of the [[stolon]]s.<ref name = Webster2007/> The stolons provide an aerial structure for the growth of the [[mycelium]] and the occupation of large areas. They can climb vertically as well as horizontally.<ref name = Vanbreuseghem1965/> ''Rhizopus'' species periodically produce rhizoids, which anchor it to the substrate and unbranched aerial sporangiophores.<ref>{{Cite web |title=Home - Rhizopus stolonifer NRRL 66455 v1.0 |url=https://mycocosm.jgi.doe.gov/Rhisto1/Rhisto1.home.html |access-date=2023-05-03 |website=mycocosm.jgi.doe.gov}}</ref> Sporangiophores of ''R. stolonifer'' can be up to 2.5&nbsp;mm long and about 20&nbsp;μm in diameter.<ref name = onions1981/> The [[spore]]s are shaped differently depending on the available nutrients. They can be ovate, polygonal or angular.<ref name = onions1981/> The optimal temperature for growth varies between 25 and 30&nbsp;°C.<ref name = Vanbreuseghem1965/> The thermal death point, which is defined as the lowest temperature that can kill all cells in ten minutes, is 60&nbsp;°C.<ref name = Vanbreuseghem1965/> ''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.<ref name = Skinner1930/> [[Ultraviolet irradiation]] can delay spore [[germination]].<ref name = Ainsworth1965/>


==Reproduction==
==Reproduction==
''Rhizopus stolonifer'' can reproduce asexually and sexually. It is a [[Heterothallism|heterothallic]] species.<ref name = Moore-Landecker1972/> Sexual reproduction occurs when compatible mating strains are paired, ultimately giving rise to [[zygospore]]s. The sporangiophore contains both '+' and '−' mating type strains.<ref name = Brown/> [[Meiosis]] is delayed until the germination of the zygospores. The [[Gametogonium|gametogenia]] often differ in size, regardless of mating type. This difference in size is not due to sex but presumably due to nutrition.<ref name = Gwynne-Vaughan1927/>
''Rhizopus stolonifer'' can reproduce asexually and sexually. It is a [[Heterothallism|heterothallic]] species.<ref name = Moore-Landecker1972/> Sexual reproduction occurs when compatible haploid mating types in the form of hyphae make contact with each other, producing diploid [[zygospore]]s. The sporangiophore contains both '+' and '−' mating type strains in the form of sporangiospores.<ref name = Brown/> [[Meiosis]] is delayed until the germination of the zygospores. The [[Gametogonium|gametogenia]] often differ in size, regardless of mating type. This difference in size is not due to sex but presumably due to nutrition.<ref name = Gwynne-Vaughan1927/>


==Disease and prevention==
==Disease and prevention==
[[File:Rhizopus_stolonifer_on_a_strawberry_showing_black_sporangia.jpg|thumb|right|250px|''R. stolonifer'' colonizing a strawberry.]]
[[File:Rhizopus_stolonifer_on_a_strawberry_showing_black_sporangia.jpg|thumb|right|250px|''R. stolonifer'' colonizing a strawberry.]]
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.<ref name = Baggio2016/> After a couple of days, the infected fruits become soft and release juices with an acidic odour.<ref name = Baggio2016/> 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 [[Sporangium|sporangia]] and spores.<ref name = Baggio2016/> When the fungus germinates, it produces different kinds of [[esterase]]s, including [[cutinase]], which help the fungus to penetrate the plant cell wall.<ref name = Baggio2016/> The disease can also affect other adjacent healthy fruits when distributed by wind or insect activity.<ref name = Alexopoulos1952/>
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.<ref name = Baggio2016/> ''R. stolonifer'' can also cause soft rot of many vegetables, flowers, bulbs, corms, and seeds.<ref>{{Cite web |last=Scheck |first=Heather |date=2020-09-29 |title=Rhizopus stolonifer (Ehrenb.: Fr.) Vuill. 1902 Bulb rot |url=https://blogs.cdfa.ca.gov/Section3162/?p=7605 |access-date=2023-05-03 |website=Pest Rating Proposals and Final Ratings |language=en-US}}</ref> After a couple of days, the infected fruits become soft and release juices with an acidic odour.<ref name = Baggio2016/> 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 [[Sporangium|sporangia]] and spores.<ref name = Baggio2016/> When the fungus germinates, it produces different kinds of [[esterase]]s, including [[cutinase]], which help the fungus to penetrate the plant cell wall.<ref name = Baggio2016/> Once spores germinate and puncture the cell wall, various enzymes are released to break down glucose, fructose, sucrose, and other sugars, producing fumaric acid, lactic acid, and ethanol.<ref>{{Cite journal |last1=Zaveri |first1=Anjali |last2=Edwards |first2=Jacqueline |last3=Rochfort |first3=Simone |date=January 2022 |title=Production of Primary Metabolites by Rhizopus stolonifer, Causal Agent of Almond Hull Rot Disease |journal=Molecules |language=en |volume=27 |issue=21 |pages=7199 |doi=10.3390/molecules27217199 |doi-access=free |issn=1420-3049 |pmc=9657676 |pmid=36364023}}</ref> The disease can also affect other adjacent healthy fruits when distributed [[Anemochory|by wind]] or insect activity.<ref name = Alexopoulos1952/>


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.<ref name = baker1977/> Fengycin, which is an anti-fungal complex, also induces the fungal cell death via [[necrosis]] and [[apoptosis]].<ref name = Tang2014/> The treatment of sweet potatoes with [[sodium orthophenyl phenol]] (Stopmold B) and dichloran (Botran W) have effectively reduced storage rot.<ref name = Sarbhoy1966/>
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.<ref name = baker1977/> Fengycin, which is an anti-fungal complex, also induces the fungal cell death via [[necrosis]] and [[apoptosis]].<ref name = Tang2014/> The treatment of sweet potatoes with [[sodium orthophenyl phenol]] (Stopmold B) and dichloran (Botran W) have effectively reduced storage rot.<ref name = Sarbhoy1966/>


''Rhizopus stolonifer'' is an [[Opportunism|opportunistic]] agent of disease and hence will only cause infection in people with a weakened immunity. [[Zygomycosis]] 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.<ref name = Brown/> The action of smelling spoiled food may be a source of inhalation exposure to the mold.<ref name = Brown/>
''Rhizopus stolonifer'' is an [[Opportunism|opportunistic]] agent of disease and hence will only cause infection in people with a weakened immunity. [[Zygomycosis]] 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.<ref name = Brown/> More commonly, Rhizopus stolonifer may cause respiratory infections, sinusitis, and otomycosis.<ref>{{Cite journal |last1=Maness |first1=Lisa R |last2=Zubov |first2=Tanya |date=2019-01-28 |title=The Inhibitory Effect of Essential Oils on''Rhizopus stolonifer, Trichophyton mentagrophytes'', and''Microsporum gypseum'' |url=https://doi.org/10.1093/labmed/lmy080 |journal=Laboratory Medicine |volume=50 |issue=2 |pages=e18–e22 |doi=10.1093/labmed/lmy080 |issn=0007-5027 |pmc=6452308 |pmid=30690523}}</ref> The action of smelling spoiled food may be a source of inhalation exposure to the mold.<ref name = Brown/>


==Importance==
==Importance==
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{{Reflist|30em|refs=<ref name = Webster2007>{{cite book | last1 = Webster | first1 = John | last2 = Weber | first2 = Roland | title = Introduction to fungi | date = 2007 | publisher = Cambridge university press | location = Cambridge, UK | isbn = 978-0-511-27783-2 | edition = 3rd}}</ref>
{{Reflist|30em|refs=<ref name = Webster2007>{{cite book | last1 = Webster | first1 = John | last2 = Weber | first2 = Roland | title = Introduction to fungi | date = 2007 | publisher = Cambridge university press | location = Cambridge, UK | isbn = 978-0-511-27783-2 | edition = 3rd}}</ref>


<ref name = Alexopoulos1952>{{cite book | last1 = Alexopoulos | first1 = C.J. | title = Introductory mycology | year = 1952 | publisher = Academic Press | location = London, UK | isbn = 9780122204029 | edition = 2nd}}</ref>
<ref name = Alexopoulos1952>{{cite book | last1 = Alexopoulos | first1 = C.J. | title = Introductory mycology | year = 1952 | publisher = Academic Press | location = London, UK | isbn = 9780122204029 | edition = 2nd | url-access = registration | url = https://archive.org/details/tudesdesciencer00putsgoog }}</ref>


<ref name = Moore-Landecker1972>{{cite book | last1 = Moore-Landecker | first1 = E. | title = Fundamentals of the Fungi | year = 1972 | publisher = Prentice Hall | location = New Jersey| isbn = 0133392678 }}</ref>
<ref name = Moore-Landecker1972>{{cite book | last1 = Moore-Landecker | first1 = E. | title = Fundamentals of the Fungi | year = 1972 | publisher = Prentice Hall | location = New Jersey | isbn = 0133392678 | url-access = registration | url = https://archive.org/details/fundamentalsoffu0000moor_m8o7 }}</ref>


<ref name = Vanbreuseghem1965>{{cite book | last1 = Vanbreuseghem | first1 = R. | last2 = Langeron | first2 = M. | title = Outline of mycology | year = 1965 | publisher = Pitman publishing | isbn = 0273417207 | edition= 2nd}}</ref>
<ref name = Vanbreuseghem1965>{{cite book | last1 = Vanbreuseghem | first1 = R. | last2 = Langeron | first2 = M. | title = Outline of mycology | year = 1965 | publisher = Pitman publishing | isbn = 0273417207 | edition= 2nd}}</ref>


<ref name = Baggio2016>{{cite journal | last1 = Baggio | first1 = J.S. | last2 = Goncalves | first2 = F.P. | last3 = Lourenco| first3 = S.A.| last4 = Tanaka | first4= A.O. |last5=Pascholati | first5 = S.F.| last6= Amorim | first6= L. |title = Direct penetration of Rhizopus stolonifer into stone fruits causing rhizopus rot | year = 2016 | journal= Plant Pathology}}</ref>
<ref name = Baggio2016>{{cite journal | last1 = Baggio | first1 = J.S. | last2 = Goncalves | first2 = F.P. | last3 = Lourenco| first3 = S.A.| last4 = Tanaka | first4= A.O. |last5=Pascholati | first5 = S.F.| last6= Amorim | first6= L. |title = Direct penetration of Rhizopus stolonifer into stone fruits causing rhizopus rot | year = 2016 | journal= Plant Pathology| volume = 65 | issue = 4 | pages = 633–642 | doi = 10.1111/ppa.12434 }}</ref>


<ref name="baker1977">{{cite journal|last1=Baker|first1=KL|last2=Beneke|first2=ES|last3=Hooper|first3=GR|last4=Fields|first4=WG|title=Host Range and Axenic Culture of the Mycoparasite Syncephalis sphaerica (Mucorales)|journal=Mycologia|date=1977|volume=69|issue=5|pages=1008–1015}}</ref>
<ref name="baker1977">{{cite journal|last1=Baker|first1=KL|last2=Beneke|first2=ES|last3=Hooper|first3=GR|last4=Fields|first4=WG|title=Host Range and Axenic Culture of the Mycoparasite Syncephalis sphaerica (Mucorales)|journal=Mycologia|date=1977|volume=69|issue=5|pages=1008–1015|doi=10.1080/00275514.1977.12020152}}</ref>


<ref name = onions1981>{{cite book | last1 = Onions | first1 = A.H.S. | last2 = Allsopp | first2 = D. | last3 = Eggins | first3 = H.O.W. | title = Smith's introduction to industrial mycology | date = 1981 | publisher = Arnold | location = London, UK | isbn = 0-7131-2811-9 | edition = 7th}}</ref>
<ref name = onions1981>{{cite book | last1 = Onions | first1 = A.H.S. | last2 = Allsopp | first2 = D. | last3 = Eggins | first3 = H.O.W. | title = Smith's introduction to industrial mycology | date = 1981 | publisher = Arnold | location = London, UK | isbn = 0-7131-2811-9 | edition = 7th}}</ref>


<ref name = Ainsworth1965>{{cite book | last1 = Ainsworth | first1 = G.C. | last2= Sussman | first2= A.S. |title = The Fungi An Advanced Treatise | year = 1965 | publisher = Academic Press | ASIN = B002M3UZVM | volume = 1}}</ref>
<ref name = Ainsworth1965>{{cite book | last1 = Ainsworth | first1 = G.C. | last2= Sussman | first2= A.S. |title = The Fungi An Advanced Treatise | year = 1965 | publisher = Academic Press | asin = B002M3UZVM | volume = 1}}</ref>


<ref name = Brown>{{cite web | last = Brown | first = Ryan | title = Mold: a study of common fungi | url= https://www.microscopy-uk.org.uk/mag/artnov14macro/Brown_Ryan_Mold.pdf}}</ref>
<ref name = Brown>{{cite web | last = Brown | first = Ryan | title = Mold: a study of common fungi | url= https://www.microscopy-uk.org.uk/mag/artnov14macro/Brown_Ryan_Mold.pdf}}</ref>


<ref name = Campbell2013>{{cite book | last1 =Campbell | first1 = C.K. | last2 = Johnson | first2 = E.M. | last3 = Warnock| first3 = D.W. |title = Identification of pathogenic fungi | year = 2013 | publisher = Wiley-Blackwell| location = New Jersey| ASIN= B0000CIFGO}}</ref>
<ref name = Campbell2013>{{cite book | last1 =Campbell | first1 = C.K. | last2 = Johnson | first2 = E.M. | last3 = Warnock| first3 = D.W. |title = Identification of pathogenic fungi | year = 2013 | publisher = Wiley-Blackwell| location = New Jersey| asin= B0000CIFGO}}</ref>


<ref name="Sarbhoy1966">{{cite journal|last1=Sarbhoy|first1=AK|title=Rhizopus stolonifer|journal=CMI Descriptions of Pathogenic Fungi and Bacteria|date=1966|volume=110|pages=1–2}}</ref>
<ref name="Sarbhoy1966">{{cite journal|last1=Sarbhoy|first1=AK|title=Rhizopus stolonifer|journal=CMI Descriptions of Pathogenic Fungi and Bacteria|date=1966|volume=110|pages=1–2}}</ref>
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<ref name = Tang2014>{{cite journal | last1 = Tang | first1 = Q. | last2 = Bie | first2 = X. | last3 = Lu| first3 = Z. | last4 = Lv | first4= F. |last5=Tao | first5 = Y.| last6= Qu | first6= X. |title = Effects of Fengycin from Bacillus subtilis fmbJ on Apoptosis and Necrosis in Rhizopus stolonifer | year = 2014| journal= Journal of Microbiology | volume=2}}</ref>
<ref name = Tang2014>{{cite journal | last1 = Tang | first1 = Q. | last2 = Bie | first2 = X. | last3 = Lu| first3 = Z. | last4 = Lv | first4= F. |last5=Tao | first5 = Y.| last6= Qu | first6= X. |title = Effects of Fengycin from Bacillus subtilis fmbJ on Apoptosis and Necrosis in Rhizopus stolonifer | year = 2014| journal= Journal of Microbiology | volume=2}}</ref>


<ref name = Gwynne-Vaughan1927>{{cite book | last1 =Gwynne-Vaughan | first1 = H.C.I. | last2 = Barnes | first2 = B. |title = structure and development of the fungi | year = 1927 | publisher = The university press | location = Cambridge}}</ref>
<ref name = Gwynne-Vaughan1927>{{cite book | last1 =Gwynne-Vaughan | first1 = H.C.I. | last2 = Barnes | first2 = B. |title = structure and development of the fungi | url =https://archive.org/details/structuredevelop0000gwyn_b0y0 | url-access =registration | year = 1927 | publisher = The university press | location = Cambridge}}</ref>


<ref name = Ellis>{{cite web | last = Ellis | first = D. | title = Fungal Descriptions and Antifungal Susceptibility | url= https://mycology.adelaide.edu.au/descriptions/zygomycetes/rhizopus/}}</ref>
<ref name = Ellis>{{cite web | last = Ellis | first = D. | title = Fungal Descriptions and Antifungal Susceptibility | url= https://mycology.adelaide.edu.au/descriptions/zygomycetes/rhizopus/}}</ref>


<ref name = Benny2016>{{cite journal | last1 = Benny | first1 = G.L. | last2 = Smith | first2 = M.E. | last3 = Kirk| first3 = p.m.| last4 = Tretter | first4= E.D|last5=White| first5 = M.M.| title = Challenges and Future Perspectives in the Systematics of Kickxellomycotina, Mortierellomycotina, Mucoromycotina, and Zoopagomycotina | year = 2016 | journal= Biology of Microfungi}}</ref>
<ref name = Benny2016>{{cite journal | last1 = Benny | first1 = G.L. | last2 = Smith | first2 = M.E. | last3 = Kirk| first3 = p.m.| last4 = Tretter | first4= E.D|last5=White| first5 = M.M.| title = Challenges and Future Perspectives in the Systematics of Kickxellomycotina, Mortierellomycotina, Mucoromycotina, and Zoopagomycotina | year = 2016 | journal= Biology of Microfungi| series = Fungal Biology | pages = 65–126 | doi = 10.1007/978-3-319-29137-6_5 | isbn = 978-3-319-29135-2 }}</ref>


<ref name = Skinner1930>{{cite book | last1 =Skinner | first1 = C. |title = Molds, Yeasts and Actinomycetes| year = 1930 | publisher = John Wiley and Sons | ASIN= B002A1J300}}</ref>
<ref name = Skinner1930>{{cite book | last1 =Skinner | first1 = C. |title = Molds, Yeasts and Actinomycetes| url =https://archive.org/details/in.ernet.dli.2015.271321 | year = 1930 | publisher = John Wiley and Sons | asin= B002A1J300}}</ref>


}}
}}
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{{Taxonbar|from=Q1500736}}
{{Taxonbar|from=Q1500736}}


[[Category:Food microbiology]]
[[Category:Mucoraceae]]
[[Category:Mucoraceae]]
[[Category:Fungi described in 1818]]
[[Category:Fungus species]]

Latest revision as of 15:17, 22 July 2024

Rhizopus stolonifer
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Fungi
Division: Mucoromycota
Class: Mucoromycetes
Order: Mucorales
Family: Mucoraceae
Genus: Rhizopus
Species:
R. stolonifer
Binomial name
Rhizopus stolonifer
Vuillemin (1902)
Synonyms
  • Rhizopus nigricans Ehrenberg (1820)
  • Mucor stolonifer Ehrenberg (1818)
  • Rhizopus artocarpi var. luxurians Schroet (1886)
  • Mucor niger Geodoelst (1902)
  • Rhizopus niger Ciaglinski & Hewelke (1893)

Rhizopus stolonifer is commonly known as black bread mold.[1] It is a member of Zygomycota and considered the most important species in the genus Rhizopus.[2] 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.[3] It is a common agent of decomposition of stored foods.[4] Like other members of the genus Rhizopus, R. stolonifer grows rapidly, mostly in indoor environments.[5]

History

[edit]

This fungus was first discovered by the German scientist Christian Gottfried Ehrenberg in 1818 as Rhizopus nigricans. The name was changed in 1902 to Rhizopus stolonifer by the French mycologist J. P. Vuillemin.[6]

Habitat and ecology

[edit]

Rhizopus stolonifer has a cosmopolitan distribution and is found on all types of mouldy materials. It is often one of the first molds to appear on stale bread.[6] 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.[6]

In the laboratory, this fungus grows well on different media, including those that contain ammonium salts or amino compounds.[6] However, R. stolonifer will not grow on Czapek's agar because it cannot utilize nitrogen in the form of nitrate.[6] Rhizopus lives in hyphae and matured spores.

Growth and physiology

[edit]
Gem-shaped sporangiospores of R. stolonifer showing characteristic striations.

This species is known as a saprotroph and plays an important role in the early colonization of substrata in soil. Nonetheless, it can also behave as a parasite of plant tissues causing a rot of vegetables and fruits.[2] Like other species of Rhizopus, R. stolonifer grows rapidly and spreads by means of the stolons.[2] The stolons provide an aerial structure for the growth of the mycelium and the occupation of large areas. They can climb vertically as well as horizontally.[7] Rhizopus species periodically produce rhizoids, which anchor it to the substrate and unbranched aerial sporangiophores.[8] Sporangiophores of R. stolonifer can be up to 2.5 mm long and about 20 μm in diameter.[6] The spores are shaped differently depending on the available nutrients. They can be ovate, polygonal or angular.[6] The optimal temperature for growth varies between 25 and 30 °C.[7] The thermal death point, which is defined as the lowest temperature that can kill all cells in ten minutes, is 60 °C.[7] 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.[9] Ultraviolet irradiation can delay spore germination.[10]

Reproduction

[edit]

Rhizopus stolonifer can reproduce asexually and sexually. It is a heterothallic species.[3] Sexual reproduction occurs when compatible haploid mating types in the form of hyphae make contact with each other, producing diploid zygospores. The sporangiophore contains both '+' and '−' mating type strains in the form of sporangiospores.[11] Meiosis 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.[12]

Disease and prevention

[edit]
R. stolonifer colonizing a strawberry.

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.[13] R. stolonifer can also cause soft rot of many vegetables, flowers, bulbs, corms, and seeds.[14] After a couple of days, the infected fruits become soft and release juices with an acidic odour.[13] 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 and spores.[13] When the fungus germinates, it produces different kinds of esterases, including cutinase, which help the fungus to penetrate the plant cell wall.[13] Once spores germinate and puncture the cell wall, various enzymes are released to break down glucose, fructose, sucrose, and other sugars, producing fumaric acid, lactic acid, and ethanol.[15] The disease can also affect other adjacent healthy fruits when distributed by wind or insect activity.[4]

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.[16] Fengycin, which is an anti-fungal complex, also induces the fungal cell death via necrosis and apoptosis.[17] The treatment of sweet potatoes with sodium orthophenyl phenol (Stopmold B) and dichloran (Botran W) have effectively reduced storage rot.[18]

Rhizopus stolonifer is an opportunistic agent of disease and hence will only cause infection in people with a weakened immunity. Zygomycosis 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.[11] More commonly, Rhizopus stolonifer may cause respiratory infections, sinusitis, and otomycosis.[19] The action of smelling spoiled food may be a source of inhalation exposure to the mold.[11]

Importance

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Rhizopus stolonifer is important economically as an agent of post-harvest storage decay.

While Saccharomyces cerevisiae is the most important source of industrial alcohol, R. stolonifer and other species of Rhizopus also produce ethyl alcohol which is the most important fermentation product.[4] Rhizopus stolonifer is also used commercially to manufacture fumaric acid and lactic acid of high purity.[20] The presence of zinc reduces the amount of fumaric acid produced by the fungus.[10]

References

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  1. ^ Ellis, D. "Fungal Descriptions and Antifungal Susceptibility".
  2. ^ a b c Webster, John; Weber, Roland (2007). Introduction to fungi (3rd ed.). Cambridge, UK: Cambridge university press. ISBN 978-0-511-27783-2.
  3. ^ a b Moore-Landecker, E. (1972). Fundamentals of the Fungi. New Jersey: Prentice Hall. ISBN 0133392678.
  4. ^ a b c Alexopoulos, C.J. (1952). Introductory mycology (2nd ed.). London, UK: Academic Press. ISBN 9780122204029.
  5. ^ Benny, G.L.; Smith, M.E.; Kirk, p.m.; Tretter, E.D; White, M.M. (2016). "Challenges and Future Perspectives in the Systematics of Kickxellomycotina, Mortierellomycotina, Mucoromycotina, and Zoopagomycotina". Biology of Microfungi. Fungal Biology: 65–126. doi:10.1007/978-3-319-29137-6_5. ISBN 978-3-319-29135-2.
  6. ^ a b c d e f g Onions, A.H.S.; Allsopp, D.; Eggins, H.O.W. (1981). Smith's introduction to industrial mycology (7th ed.). London, UK: Arnold. ISBN 0-7131-2811-9.
  7. ^ a b c Vanbreuseghem, R.; Langeron, M. (1965). Outline of mycology (2nd ed.). Pitman publishing. ISBN 0273417207.
  8. ^ "Home - Rhizopus stolonifer NRRL 66455 v1.0". mycocosm.jgi.doe.gov. Retrieved 2023-05-03.
  9. ^ Skinner, C. (1930). Molds, Yeasts and Actinomycetes. John Wiley and Sons. ASIN B002A1J300.
  10. ^ a b Ainsworth, G.C.; Sussman, A.S. (1965). The Fungi An Advanced Treatise. Vol. 1. Academic Press. ASIN B002M3UZVM.
  11. ^ a b c Brown, Ryan. "Mold: a study of common fungi" (PDF).
  12. ^ Gwynne-Vaughan, H.C.I.; Barnes, B. (1927). structure and development of the fungi. Cambridge: The university press.
  13. ^ a b c d Baggio, J.S.; Goncalves, F.P.; Lourenco, S.A.; Tanaka, A.O.; Pascholati, S.F.; Amorim, L. (2016). "Direct penetration of Rhizopus stolonifer into stone fruits causing rhizopus rot". Plant Pathology. 65 (4): 633–642. doi:10.1111/ppa.12434.
  14. ^ Scheck, Heather (2020-09-29). "Rhizopus stolonifer (Ehrenb.: Fr.) Vuill. 1902 Bulb rot". Pest Rating Proposals and Final Ratings. Retrieved 2023-05-03.
  15. ^ Zaveri, Anjali; Edwards, Jacqueline; Rochfort, Simone (January 2022). "Production of Primary Metabolites by Rhizopus stolonifer, Causal Agent of Almond Hull Rot Disease". Molecules. 27 (21): 7199. doi:10.3390/molecules27217199. ISSN 1420-3049. PMC 9657676. PMID 36364023.
  16. ^ Baker, KL; Beneke, ES; Hooper, GR; Fields, WG (1977). "Host Range and Axenic Culture of the Mycoparasite Syncephalis sphaerica (Mucorales)". Mycologia. 69 (5): 1008–1015. doi:10.1080/00275514.1977.12020152.
  17. ^ Tang, Q.; Bie, X.; Lu, Z.; Lv, F.; Tao, Y.; Qu, X. (2014). "Effects of Fengycin from Bacillus subtilis fmbJ on Apoptosis and Necrosis in Rhizopus stolonifer". Journal of Microbiology. 2.
  18. ^ Sarbhoy, AK (1966). "Rhizopus stolonifer". CMI Descriptions of Pathogenic Fungi and Bacteria. 110: 1–2.
  19. ^ Maness, Lisa R; Zubov, Tanya (2019-01-28). "The Inhibitory Effect of Essential Oils onRhizopus stolonifer, Trichophyton mentagrophytes, andMicrosporum gypseum". Laboratory Medicine. 50 (2): e18–e22. doi:10.1093/labmed/lmy080. ISSN 0007-5027. PMC 6452308. PMID 30690523.
  20. ^ Campbell, C.K.; Johnson, E.M.; Warnock, D.W. (2013). Identification of pathogenic fungi. New Jersey: Wiley-Blackwell. ASIN B0000CIFGO.