Rhizopus oryzae

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Rhizopus oryzae
Rhizopus Oryzae.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Fungi
Division: Mucoromycota
Order: Mucorales
Family: Mucoraceae
Genus: Rhizopus
Species:
R. oryzae
Binomial name
Rhizopus oryzae
Synonyms

Rhizopus arrhizusA. Fisch., (1892)
Rhizopus stolonifer Vuillemin, (1902)
Rhizopus japonicusVuill. (1902)
Rhizopus nodosusNamsyl. (1906)
Rhizopus nodosusHanzawa (1912)

Contents

Rhizopus oryzae is a filamentous heterothallic microfungus that occurs as a saprotroph in soil, dung, and rotting vegetation. This species is very similar to Rhizopus stolonifer, but it can be distinguished by its smaller sporangia and air-dispersed sporangiospores. It differs from R. oligosporus and R. microsporus by its larger columellae and sporangiospores. [1] The many strains of R. oryzae produce a wide range of enzymes such as carbohydrate digesting enzymes and polymers along with a number of organic acids, ethanol and esters giving it useful properties within the food industries, bio-diesel production, and pharmaceutical industries. It is also an opportunistic pathogen of humans causing mucormycosis.

History and taxonomy

Rhizopus oryzae was discovered by Frits Went and Hendrik Coenraad Prinsen Geerligs in 1895. [1] The genus Rhizopus (family Mucoraceae ) was erected in 1821 by the German mycologist, Christian Gottfried Ehrenberg [2] to accommodate Mucor stolonifer and Rhizopus nigricans as distinct from the genus Mucor. [3] The genus Rhizopus is characterized by having stolons, rhizoids, sporangiophores sprouting from the points of which rhizoids were attached, globose sporangia with columellae, striated sporangiospores. [3] In the mid 1960s, researchers divided the genus based on temperature tolerance. Numerical methods were later used in the early 1970s where researchers arrived at similar conclusions. R. oryzae was relegated to a distinct section because it grew well at 37 °C but failed to grow at 45 °C. [4] In the past, strains were identified through isolating active components of the species that were commonly found in food and alcoholic drinks in Indonesia, China, and Japan. [4] There are approximately 30 synonyms, the most common being R. arrhizus. [5] Scholer popularized R. oryzae because he thought R. arrhizus represented an extreme form of R. oryzae. [4]

Growth and morphology

Rhizopus oryzae grows quickly in optimal temperatures, at 1.6 mm per hour (nearly 0.5 μm per second - enough to be able to directly visualize hyphal elongation in real-time under the microscope). [1] R. oryzae can grow in temperature of 7 °C to 44 °C and the optimum growth temperature is 37 °C. [1] [6] There is very poor growth from 10 °C to 15 °C [3] and negligible growth at 45 °C. [2] [4] There is substantial growth in media containing 1% NaCl, very poor growth at 3% NaCl, and none at 5% NaCl. R. oryzae favors slightly acidic media. Good growth is observed at a pH of 6.8; in the range of 7.7-8.1, there is very poor growth. [3] Most amino acids—with the exception of L-valine—promote R. oryzae growth, with L-tryptophan and L-tyrosine being the most effective. It also grows well on mineral nitrogen sources, except nitrate, and can utilize urea. [7]

Rhizopus oryzae has variable sporangiosphores. They can be straight or curved, swollen or branched, and the walls can be smooth or slightly rough. The colour of sporangiosphores range from pale brown to brown. Sporangiosphores grow between 210-2500 μm in length and 5-18 μm in diameter. The sporangia in R. oryzae are globose or subglobose, wall spinous and black when mature, 60-180 μm in diameter. They can be distinguishable from Rhizopus stolonifer as they have smaller sporangia and spores. [1] The optimal conditions for sporangium production are temperatures between 30 °C to 35 °C and low water levels. [7] Sporulation is stimulated by amino acids (except L-valine) when grown in light, while in darkness only L-tryptophan and L-methionine effect stimulation of growth. The columellae are globose, subglobose, or oval in shape. The wall is usually smooth and the colour is pale brown. The average diameter growth ranges from 30-110 μm. Sporangiospores are elliptical, globose, or polygonal, they are striated and grow 5-8 μm in length. Dormant and germinated sporangiospores show deep furrows and prominent ridges with a pattern that makes it distinguishable from that of R. stolonifer. The germination of sporangiospores can be induced by the combined action of L-proline and phosphate ions. L-ornithine, L-arginine, D-glucose and D-mannose are also effective. Optimal germination occurs on media containing D-glucose and mineral salts. [7] R. oryzae has abundant, root-shaped rhizoids. [3] Zygospores are produced by diploid cells when sexual reproduction occurs under nutrient poor conditions. They have colors that range from red to brown, they are spherical or laterally flattened, and ranges from 60-140μm in size. [2] In high nutrient levels, R. oryzae reproduces asexually, producing azygospores. [8] The stolons found in R. oryzae are smooth or slightly rough, almost colorless or pale brown, 5-18 μm in diameter. The chlamydospores are abundant, globose ranging in 10-24 μm in diameter, elliptical, and cylindrical. Colonies of R. oryzae are white initially, becoming brownish with age [6] and can grow to about 1 cm thick. [2]

Habitat and ecology

Rhizopus oryzae can be found in various soils across the world. For example, it has been found in India, Pakistan, New Guinea, Taiwan, Central America, Peru, Argentina, Namibia, South Africa, Iraq, Somalia, Egypt, Libya, Tunisia, Israel, Turkey, Spain, Italy, Hungary, Czech Republic, Slovakia, Germany, Ukraine, British Isles, and the USA. The soils where R. oryzae has been isolated are varied ranging from grassland, cultivated soils under lupin, corn, wheat, groundnuts, other legumes, sugar canes, rice, citrus plantations, steppe type vegetation, alkaline soils, salt-marshes, farm manure soils, to sewage filled soils. The pH of the soils where the species has been isolated typically range from 6.3 to 7.2. [7]

Rhizopus oryzae is often identified as R. arrhizus when isolated from foods. It is found in rotting fruits and vegetables where it is often called R. stolonifer. Unlike the other species such as R. stolonifer, R. oryzae is common in tropical conditions. In East Asia, it is common in peanuts. For instance, there was 21% isolation from peanut kernels from Indonesia. [1] It is present in maize, beans, sorghum, and cowpeas, pecans, hazelnuts, pistachios, wheat, barley, potatoes, sapodillas, and various other tropical foods. [1] Maize meal on which isolates of R. oryzae had been grown was found to be toxic to ducklings and rats, causing growth depression. [4]

Pathogenicity

Rhizopus oryzae is one of the most common causes of a disease known as mucormycosis, characterized by growing hyphae within and surrounding blood vessels. The causal agents of mucormycosis may also produce toxins like agroclavine which is toxic to humans, sheep and cattle. [7] This infection usually occurs in immunocompromised individuals but is rare. [9] [10] Common risk factors associated with primary cutaneous mucormycosis is ketoacidosis, neutropenia, acute lymphobloastic leukemia, lymphomas, systemic steroids, chemotherapy, and dialysis. Treatment includes amphotericin B, posaconazole, itraconazole, and fluconazole. [11] The majority of the cases of infection are rhinocerebral infections. At the same time, it has been found in literature that R. oryzae can produce antibiotic activity on some bacteria. [7]

The pathogenicity towards plants is attributed to the presence of large number of carbohydrate digesting enzymes.[ citation needed ]

Physiology and industrial uses

Rhizopus oryzae is involved in steroid transformations and it produces 4-desmethyl steroids which has been useful in the fermentation industry. The carbon sources does influence the ratio of polar and neutral lipids. The mycelium found in R. oryzae contains lipids and the highest lipid content occurs when grown on fructose. The highest unsaturated fatty acid content is observed at 30 °C and lowest at 15 °C. Proteolytic properties have been observed well under the conditions of pH 7 at 35 °C. Pyridozine and thiamine prefer proteinase production. R. oryzae can degrade aflatoxin A1 to isomeric hydroxy compounds and aflatoxin G1 to fluorescent metabolite aflatoxin A1. [7] There are various factors that influence the production of dextro-lactic acids, fumaric acid, and metabolism of R. oryzae. For examples, in 40 °C there is more favorable growth for glucose consumption, however this influenced production of d-lactic acid production negatively. Glucose concentration of 15% is needed for optimal production of d-lactic acid. Fumaric acid production was suppressed in media containing more than 6 grams of NH4NO3 per liter and is favorable to d-lactic acid production. [12]

Rhizopus oryzae is considered GRAS by the FDA and thus recognized as safe to use industrially as it can consume a range of carbon sources. [13] During fermentation. R. oryzae produce amylase, lipase, and protease activity to increase nutrient's ability to use many compounds as an energy and carbon source. [14] Historically, it has been used in fermentation, specifically to ferment soybean and create tempeh in Malaysia and Indonesia. [15] Using the same methods to create traditional tempeh, R. oryzae can be inoculated in other cooked legumes such as peas, beans, and fava beans. Similarly in tempeh making, there is an initial bacterial fermentation in legumes when they are soaked for a while before being cooked. Fermentation incubation lasts for 48 hours at 33 °C. After incubation, mycelium can be observed between the legumes creating a larger, uniform product. Overall, fruits, grains, nuts, and legumes mold-fermentation with R. oryzae produces sensory changes in foods such as creating acidity, sweetness and bitterness. R. oryzae can produce lactate from glucose at high levels, which is used as a food additive and can also degrade plastics. [16] In enzyme-modified cheese products, R. oryzae provides microbial enzymes where milk fat and proteins are broken down to create powder and paste forms of cheese. Specifically, it breaks down cheese curds and acid casein. [17]

Among finding cellulases and hemicellulases, other enzymes such as protease, urease, ribonuclease, pectate lyase, and polygalacturonase are found in cultural media of R. oryzae. Besides producing a number of enzymes, it can also produce a number of organic acids, alcohol, and esters. Cellulases in R. oryzae can be applied to biotechnology, in food, brewery and wine, animal feed, textiles and laundry, pulp and paper industries, and agriculture. R. oryzae can convert both glucose and xylose under aerobic conditions into pure L (+)-lactic acids with by-products such as xylitol, glycerol, ethanol, carbon dioxide and fungal biomass. Endo-xylanase is a key enzyme for xylan depolymerization and was produced by R. oryzae fermentation from different xylan-containing agricultural by-products such as wheat straw, wheat stems, cottons bagasse, hazelnut shells, corn cobs, and oat sawdust. Pectinases are required for extraction and clarification of fruit juices and wines, extraction of oils, flavors and pigmentation from plant material, preparation of cellulose fibers for linen, jute and hemp manufacture as well as, coffee and tea fermentations. R. oryzae can break down starch content in rice plants and therefore shows amylolytic activities. Also, it has been reported to produce extra cellular isoamylase which is used in food industries. Isoamylase was found to saccharify potato starch, arrow root, tamarind kernel, tapioca, and oat. The saccharifying ability of the enzyme is highly applicable in sugar production industries. Proteases, which can be found in R. oryzae are highly useful in commercial industries. For instance, it has increased application in food, pharmaceutical, detergent, leather, tanning industries. It is also involved in silver recovery and peptide synthesis. One strain of R. oryzae was found to secrete alkaline serine protease which shows high pH stability within 3 to 6 and poor thermos-stability. Lipase that is extracted from R. oryzae have been consumed as digestive aids without adverse reactions. Lipases hydrolyze fats and oils with subsequent release of free fatty acids such as diacylglycerols, monoacylglycerols and glycerol. Lipases have been involved in biotechnology applications because of its ability to catalyze synthetic reactions in non-aqueous solutions. One study has reported the expression of a fungal 11 alpha-steroid hydroxylase from R. oryzae which can be used to perform the 11 alpha-hydroxylation of the steroid skeleton which has simplified steroid drug production. [18] R. oryzae can produce intracellular ribonuclease in a metal ion-regulated liquid medium with the addition of calcium and molybdenum stimulating ribonuclease production. R. oryzae strain ENHE isolated from contaminated soil was found to be capable of tolerating and removing pentachlorophenol. R. oryzae is known to produce L (+)-lactic acid because the fungus cells possess better resistance to high concentration of accumulated lactic acid and lower content of nutrient requirement compared to the commonly used bacterial procedures. Thus, R. oryzae is the most efficient approached to improve lactic acid production process that facilitates multiple reuses of fungal cells for long-term lactic acid production. Ethanol is the main by-product in the fermentation process of R. oryzae during the production of L-lactic acid. R. oryzae can be used as a biocatalyst for ester production in organic solvent. Dry mycelium of four R. oryzae strains proved effective for catalysing the synthesis of different flavor esters. For example, the pineapple flavour or butyl acetate esters was produced by the esterification reactions between acetic acid and butanol by R. oryzae. This flavor compound can be used in food, cosmetic and pharmaceutical industries. Within the biodiesel industry, biodiesel fuel as fatty acid methyl ester is produced by the esterification of plant oil or animal fat with methanol. This is a renewable fuel resource compared to the traditional petroleum-based fuels. Production of biodiesel fuel from plant oils from cells of R. oryzae immobilized within biomass support particles were investigated for the methanolysis of soybean oil. Olive oil or oleic acid was found to be effective for enhancing methanolysis activity which is a promising results within the biodiesel industry.[ citation needed ]

Related Research Articles

<span class="mw-page-title-main">Lactic acid</span> Group of stereoisomers

Lactic acid is an organic acid. It has the molecular formula CH3CH(OH)COOH. 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 natural sources. Lactic acid is an alpha-hydroxy acid (AHA) due to the presence of a hydroxyl group adjacent to the carboxyl group. It is used as a synthetic intermediate in many organic synthesis industries and in various biochemical industries. The conjugate base of lactic acid is called lactate. The name of the derived acyl group is lactoyl.

<span class="mw-page-title-main">Lactic acid fermentation</span> Series of interconnected biochemical reactions

Lactic acid fermentation is a metabolic process by which glucose or other six-carbon sugars are converted into cellular energy and the metabolite lactate, which is lactic acid in solution. It is an anaerobic fermentation reaction that occurs in some bacteria and animal cells, such as muscle cells.

<span class="mw-page-title-main">Malolactic fermentation</span> Process in winemaking

Malolactic conversion is a process in winemaking in which tart-tasting malic acid, naturally present in grape must, is converted to softer-tasting lactic acid. Malolactic fermentation is most often performed as a secondary fermentation shortly after the end of the primary fermentation, but can sometimes run concurrently with it. The process is standard for most red wine production and common for some white grape varieties such as Chardonnay, where it can impart a "buttery" flavor from diacetyl, a byproduct of the reaction.

<i>Rhizopus</i> Genus of fungi

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 tobacco. They are multicellular. Some Rhizopus species are opportunistic human pathogens that often cause fatal disease called mucormycosis. This widespread genus includes at least eight species.

Industrial fermentation is the intentional use of fermentation in manufacturing processes. In addition to the mass production of fermented foods and drinks, industrial fermentation has widespread applications in chemical industry. Commodity chemicals, such as acetic acid, citric acid, and ethanol are made by fermentation. Moreover, nearly all commercially produced industrial enzymes, such as lipase, invertase and rennet, are made by fermentation with genetically modified microbes. In some cases, production of biomass itself is the objective, as is the case for single-cell proteins, baker's yeast, and starter cultures for lactic acid bacteria used in cheesemaking.

<span class="mw-page-title-main">Lactic acid bacteria</span> Order of bacteria

Lactobacillales are an order of gram-positive, low-GC, acid-tolerant, generally nonsporulating, nonrespiring, either rod-shaped (bacilli) or spherical (cocci) bacteria that share common metabolic and physiological characteristics. These bacteria, usually found in decomposing plants and milk products, produce lactic acid as the major metabolic end product of carbohydrate fermentation, giving them the common name lactic acid bacteria (LAB).

<i>Rhizopus oligosporus</i> Species of fungus

Rhizopus oligosporus is a fungus of the family Mucoraceae and is a widely used starter culture for the production of tempeh at home and industrially. As the mold grows it produces fluffy, white mycelia, binding the beans together to create an edible "cake" of partly catabolized soybeans. The domestication of the microbe is thought to have occurred in Indonesia several centuries ago.

<span class="mw-page-title-main">Fermentation</span> Metabolic process producing energy in the absence of oxygen

Fermentation is a metabolic process that produces chemical changes in organic substances through the action of enzymes. In biochemistry, it is broadly defined as the extraction of energy from carbohydrates in the absence of oxygen. In food production, it may more broadly refer to any process in which the activity of microorganisms brings about a desirable change to a foodstuff or beverage. The science of fermentation is known as zymology.

Rhizopus arrhizus is a fungus of the family Mucoraceae, characterized by sporangiophores that arise from nodes at the point where the rhizoids are formed and by a hemispherical columella. It is the most common cause of mucormycosis in humans and occasionally infects other animals.

Leuconostoc mesenteroides is a species of lactic acid bacteria associated with fermentation, under conditions of salinity and low temperatures. In some cases of vegetable and food storage, it was associated with pathogenicity. L. mesenteroides is approximately 0.5-0.7 µm in diameter and has a length of 0.7-1.2 µm, producing small grayish colonies that are typically less than 1.0 mm in diameter. It is facultatively anaerobic, Gram-positive, non-motile, non-sporogenous, and spherical. It often forms lenticular coccoid cells in pairs and chains, however, it can occasionally form short rods with rounded ends in long chains, as its shape can differ depending on what media the species is grown on. L. mesenteroides grows best at 30 °C, but can survive in temperatures ranging from 10 °C to 30 °C. Its optimum pH is 5.5, but can still show growth in pH of 4.5-7.0.

<i>Mucor mucedo</i> Species of fungus

Mucor mucedo, commonly known as the common pinmould, is a fungal plant pathogen and member of the phylum Mucoromycota and the genus Mucor. Commonly found on soil, dung, water, plants and moist foods, Mucor mucedo is a saprotrophic fungus found world-wide with 85 known strains. It is often mistaken for Rhizopus rots on fruits due to similar mould growth shape and colour. Contrastingly, however, Mucor mucedo is found to grow on a wide range of stored grains and plants, including cucumber and tomato. Discovered in Italy in 1729 by P.A. Micheli and later noted by Carl Linnaeus in 1753 in the Species Plantarum, Mucor mucedo was originally classified as Mucor vulgaris by Micheli but later classified synonymous under name Mucor mucedo. The species was redescribed as Ascophora mucedo by H.J. Tode in 1790 but this type resided in a stoloniferous habitat and was later made the type of new genus Rhizopus.

Rhizopus soft rot is a disease of the sweet potato. It is one of the most common to affect the sweet potato, happening during packing and shipping. The disease causes a watery soft rot of the internal portion of the storage root. Strategies to manage the disease include the development of resistant varieties, curing through the use of heat and humidity, and application of decay control products.

<i>Jiuqu</i> East Asian fermentation starter

Jiuqu, also simply known as qu is a type of dried fermentation starter used in the production of traditional Chinese alcoholic beverages. The word jiuqu specifically refers to a type of yeast used to make alcohol such as huangjiu, baijiu and jiuniang.

<i>Mucor plumbeus</i> Species of fungus

Mucor plumbeus is a fungus in the family Mucoraceae that is very common, abundant and distributed worldwide. Mucor plumbeus is not known to be a plant or animal pathogen; however it is able to elicit an immune response in humans by activating the complement system. This species is commonly found in various types of soils over a range of pH, although alkaline soils seem more conducive to its growth. It is also known from the roots of wheat, oat and barley. In addition, M. plumbeus is a common fungal contaminant of indoor built environments. This species shares many similarities with M. racemosus, another fungus that belongs to the family Mucoraceae which is known to cause mucormycosis. Mucor plumbeus is a common spoilage agent of cheese, apples, apple cider and yogurt.

<i>Nuruk</i> Korean fermentation starter

Nuruk (Korean: 누룩) is a traditional Korean fermentation starter. It is used to make various types of Korean alcoholic beverages including takju, cheongju, and soju. It is an essential ingredient in Shindari and is mixed with rice. Historically, it was used in a variety of provinces of Korea, including Jeju Island.

<i>Cunninghamella echinulata</i> Species of fungus

Cunninghamella echinulata is a fungal species in the genus Cunninghamella. It is an asexually reproducing fungus and a mesophile, preferring intermediate temperature ranges. C. echinulata is a common air contaminant, and is currently of interest to the biotechnology industry due to its ability to synthesize γ-linolenic acid as well as its capacity to bioconcentrate metals. This species is a soil saprotroph that forms rhizoids, preferring soils enriched in nitrogen, phosphorus and potassium. It has been reported occasionally an agent of mucormycosis following the inhalation of fungal spores. Czapek's agar is a suitable growth medium for the propagation of C. echinulata.

<span class="mw-page-title-main">Industrial microbiology</span>

Industrial microbiology is a branch of biotechnology that applies microbial sciences to create industrial products in mass quantities, often using microbial cell factories. There are multiple ways to manipulate a microorganism in order to increase maximum product yields. Introduction of mutations into an organism may be accomplished by introducing them to mutagens. Another way to increase production is by gene amplification, this is done by the use of plasmids, and vectors. The plasmids and/ or vectors are used to incorporate multiple copies of a specific gene that would allow more enzymes to be produced that eventually cause more product yield. The manipulation of organisms in order to yield a specific product has many applications to the real world like the production of some antibiotics, vitamins, enzymes, amino acids, solvents, alcohol and daily products. Microorganisms play a big role in the industry, with multiple ways to be used. Medicinally, microbes can be used for creating antibiotics in order to treat infection. Microbes can also be used for the food industry as well. Microbes are very useful in creating some of the mass produced products that are consumed by people. The chemical industry also uses microorganisms in order to synthesize amino acids and organic solvents. Microbes can also be used in an agricultural application for use as a biopesticide instead of using dangerous chemicals and or inoculants to help plant proliferation.

<i>Rhizopus stolonifer</i> Species of fungus

Rhizopus stolonifer is commonly known as black bread mold. It is a member of Zygomycota and considered the most important species in the genus Rhizopus. 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.

Aspergillus wentii is an asexual, filamentous, endosymbiotic fungus belonging to the mold genus, Aspergillus. It is a common soil fungus with a cosmopolitan distribution, although it is primarily found in subtropical regions. Found on a variety of organic materials, A. wentii is known to colonize corn, cereals, moist grains, peanuts and other ground nut crops. It is also used in the manufacture of biodiesel from lipids and is known for its ability to produce enzymes used in the food industry.

<i>Actinomucor elegans</i> Species of fungus

Actinomucor elegans was originally described by Schostakowitsch in Siberia in 1898 and reevaluated by Benjamin and Hesseltine in 1957. Commonly found in soil and used for the commercial production of tofu and other products made by soy fermentation. Its major identifying features are its spine-like projections on the sporangiophore and its ribbon-like hyphal structure when found in the tissue of a host.

References

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  9. ↑ Shirazi, Fazal; Kontoyiannis, Dimitrios P; Ibrahim, Ashraf S (April 2015). "Iron starvation induces apoptosis in rhizopus oryzae in vitro". Virulence. 6 (2): 121–126. doi:10.1080/21505594.2015.1009732. PMC   4601319 . PMID   25830548.
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  11. ↑ Rodríguez-Lobato, Erika; Ramírez-Hobak, Lourdes; Aquino-Matus, Jorge E.; Ramírez-Hinojosa, Juan P.; Lozano-Fernández, Víctor H.; Xicohtencatl-Cortes, Juan; Hernández-Castro, Rigoberto; Arenas, Roberto (3 November 2016). "Primary Cutaneous Mucormycosis Caused by Rhizopus oryzae: A Case Report and Review of Literature". Mycopathologia. 182 (3–4): 387–392. doi:10.1007/s11046-016-0084-6. PMID   27807669. S2CID   33088979.
  12. ↑ Lockwood, L.B. (1963). "The physiology of rhizopus oryzae".{{cite journal}}: Cite journal requires |journal= (help)
  13. ↑ Rani, Richa; Ghosh, Sanjoy (November 2011). "Production of phytase under solid-state fermentation using Rhizopus oryzae: Novel strain improvement approach and studies on purification and characterization". Bioresource Technology. 102 (22): 10641–10649. doi:10.1016/j.biortech.2011.08.075. PMID   21945206.
  14. ↑ Cantabrana, Igor; Perise, Ramón; Hernández, Igor (January 2015). "Uses of Rhizopus oryzae in the kitchen". International Journal of Gastronomy and Food Science. 2 (2): 103–111. doi: 10.1016/j.ijgfs.2015.01.001 .
  15. ↑ Londoño-Hernández, Liliana (18 September 2017). "Rhizopus oryzae – Ancient microbial resource with importance in modern food industry". International Journal of Food Microbiology. 257: 110–127. doi:10.1016/j.ijfoodmicro.2017.06.012. ISSN   0168-1605. PMID   28651077.
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  17. ↑ Early, R. (2012). "Dairy products and milk-based food ingredients". Natural Food Additives, Ingredients and Flavourings. Elsevier. pp. 417–445. doi:10.1533/9780857095725.2.417. ISBN   9781845698119.
  18. ↑ Petrič, Š.; Hakki, T.; Bernhardt, R.; Žigon, D.; Črešnar, B. (2010-11-01). "Discovery of a steroid 11α-hydroxylase from Rhizopus oryzae and its biotechnological application". Journal of Biotechnology. 150 (3): 428–437. doi:10.1016/j.jbiotec.2010.09.928. ISSN   0168-1656. PMID   20850485.
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