f u n g a l b i o l o g y r e v i e w s 3 7 ( 2 0 2 1 ) 4 1 e5 8 journal homepage: www.elsevier.com/locate/fbr Review Sporobolomyces and Sporidiobolus e nonconventional yeasts for use in industries Anna M. KOTa,*, Marek KIELISZEKa, Kamil PIWOWAREKa, a _ Stanis1aw B1AZEJAK , Cassamo Ussemane MUSSAGYb a Department of Food Biotechnology and Microbiology, Institute of Food Sciences, Warsaw University of Life Sciences, Nowoursynowska 159C, 02-776, Warsaw, Poland b Department of Bioprocesses and Biotechnology, School of Pharmaceutical Sciences, Sao Paulo State University (UNESP), Araraquara, Brazil article info abstract Article history: The search for new, biotechnologically useful yeast strains has been carried out in many Received 8 April 2021 research centers in the world. Sporobolomyces and Sporidiobolus are examples of such useful Received in revised form yeasts, that can be used as a source of many valuable metabolites in industries. This article 5 June 2021 describes the modern taxonomy of these yeasts, which resulted from many years of Accepted 20 June 2021 research, including both classical microbiology and genetic analyses. Subsequently, the paper presents a review of scientific studies on the biosynthesis of various extracellular and Keywords: intracellular metabolites produced by Sporobolomyces and Sporidiobolus yeasts, which are of Biocontrol great importance in the contemporary food, feed, and pharmaceutical industries. Such me- Carotenoids tabolites include exopolysaccharides, lipids, carotenoids, enzymes, and g-decalactone. g-decalactone Aiming at developing a sustainable circular bioeconomy, this study considers two direc- Exopolysaccharides tions of use of these yeasts, i.e., as a feed additive and as an antagonist in the biocontrol Lipids of plant materials. This article is one of the first to organize the knowledge collected Microbial enzymes from published studies and present the contemporary scientific achievements and pros- Red yeast pects for the biotechnological use of Sporobolomyces and Sporidiobolus yeasts. ª 2021 The Author(s). Published by Elsevier Ltd on behalf of British Mycological Society. This is an open access article under the CC BY license (http://creativecommons.org/ licenses/by/4.0/). 1. Introduction Yeast has been used by humans for centuries to produce food products such as beer, wine, bread, and kefir, and as a source of B vitamins and proteins. Traditionally, yeasts from the genus Saccharomyces have been primarily used in industries, but in recent decades the valuable properties of other types of yeasts, referred to as ‘non-conventional’ yeasts, have also been documented. These microorganisms include Yarrowia lipolytica, Trichosporon spp., Kluyveromyces spp., Pichia spp., Candida spp., Debaryomyces spp., and Brettanomyces spp. (Navarrete and Martınez, 2020). Non-conventional yeasts also include organisms from the Sporidiobolaceae family, such as Rhodotorula spp. (anamorph) and Rhodosporidium (teleomorph), which have been described in detail in many * Corresponding author. E-mail addresses: anna_kot@sggw.edu.pl (A. M. Kot), marek_kieliszek@sggw.edu.pl (M. Kieliszek), kamil_piwowarek@sggw.edu.pl _ (K. Piwowarek), stanislaw_blazejak@sggw.edu.pl (S. B1azejak), cassamo.mussagy@unesp.br (C. U. Mussagy). https://doi.org/10.1016/j.fbr.2021.06.001 1749-4613/ª 2021 The Author(s). Published by Elsevier Ltd on behalf of British Mycological Society. This is an open access article under the CC BY license (http://creativecommons.org/licenses/by/4.0/). 42 A. M. Kot et al.  ndez-Almanza et al., 2014; Kot et al., scientific papers (Herna 2016; Wen et al., 2020). The Sporidiobolaceae family also comprises Sporobolomyces spp. (anamorph) and Sporidiobolus spp. (teleomorph), which have the ability to biosynthesize various metabolites (Fig. 1). These yeasts can find different applications in many industries, especially in the food and cosmetic industries. To the best of our knowledge, no study has described the properties of these yeasts so far. This paper presents an overview of the most important studies that have discussed the characteristics and possibilities of the biotechnological use of yeasts of the genera Sporobolomyces and Sporidiobolus. 2. History and taxonomy Yeasts belonging to the genus Sporobolomyces were first described by von Wettstein in 1885, reporting the yeast Rhodomyces kochii, cf., yeast able to produce interminal teliospores. However, the original strains did not survive, and due to that, the description was considered ambiguous (Sampaio et al., 2011). In 1924, Kluyver and van Neil studied a genus of yeasts and observed the mirror-image formation of cultures in inverted Petri dishes. The cause of this phenomenon was found to be the release of ballistospores via the droplet route, due to the spontaneous separation of a drop of fluid from the tip of the conidial shaft. This phenomenon resulted in the release of energy, causing the spore to be lifted in the air. Kluyver and van Neil called these yeasts Sporobolomyces, emphasizing the diagnostic importance of the droplet mechanism. The first species assigned to this genus was Sporobolo€ ller, 1954). In the first edition of ‘The myces roseus (see Mu Yeasts: A Taxonomic Study’, Lodder and Kreger-van Rij (1952) classified seven species belonging to the genus Sporobolomyces, on the basis of the colony color, ability to form mycelium, Fig. 1 e Biomolecules produced by Sporobolomyces and Sporidiobolus yeasts and potential application in biotechnology. and asexual reproduction by budding. In the second edition of ‘The Yeasts: A Taxonomic Study’ (Lodder, 1970), the genus was expanded to include the species that lack the ability to ferment sugars or synthesize starch-like compounds, as well as those that did not form pigmented colonies on malt agar, which resulted in the recognition of nine species. This issue was also noted with the genus Sporidiobolus, in which two species were assigned (Barnett, 2004, Sampaio et al., 2011). This genus was created by Nyland in 1949 for yeast strains isolated from red raspberry leaves in Washington state. These yeasts exhibited atypical features such as the simultaneous presence of budding cells and ballistospores, production of mycelium from cells or blastospores, terminal or intercalar production of teliospores in the mycelium, and germination of teliospores through a short steroid-type ri et tube. Nyland named them as Sporidiobolus johnsonii (Vale al., 2008). The life cycle of this yeast was described by Laffin and Cutter in 1959. The first paper reported the sigmoidal growth of S. johnsonii during exposure to UV radiation. The course of the growth curve indicated that the yeast cells were diploid. Cytological tests showed the presence of mitotic cells as well as the meiotic-type cells (Laffina and Cuttera, 1959a). Later, Laffina and Cuttera (1959b) confirmed that mitotic divisions took place during the germination of teliospores. In 1963, Ruinen reported six yeast strains isolated from tropical plants in Indonesia and assigned them to Sporobolomyces salmonicolor. The author indicated that these strains were able to form mycelium with chlamydospores, and their inclusion in the genus Sporidiobolus could be correct. Seven years later, Phaff showed that the Ruinen strains differed from the Nyland strains, including the ability to assimilate raffinose and galactitol and the inability to assimilate melezitose and a-methyl-D-glucoside. He classified these strains as Sporidiobolus ruinenii (Holzschu et al., 1981; Nakase, 2000). In 1981, Fell and Tallman described the germination of the Sporidiobolus teliospores which gives rise to a single-celled metabasidium having one or two basidiospores. Based on their studies, the authors decided to reclassify Sporidiobolus johnsonii to Sporobolomyces salmonicolor. They also showed that the strains that were described as Sporidiobolus johnsonii assimilate very similar carbon and nitrogen compounds as Sporobolomyces salmonicolor (Fell and Tallman, 1981). However, in the same year, Holzschu and colleagues showed that the two species differ significantly in their guanine and cytosine (G þ C) content and have only about 8.5 % complementary nuclear DNA base sequence (Holzschu et al., 1981). In 2000, Hamamoto and Nakase conducted a phylogenetic analysis on yeasts capable of producing ballistoconids. They determined the complete 18S rDNA sequences (1749e1807 nucleotides) for 25 Sporobolomyces species and five Sporidiobolus species. Furthermore, the authors revealed that a close taxonomic relationship existed between the genera Sporobolomyces and Rhodotorula. They showed that it is impossible to distinguish the two genera from each other based on morphology alone, as some Sporobolomyces strains may lose ballistospore-forming ability. In the fifth edition of ‘The Yeasts: A Taxonomic Study’, seven yeast species from the genus Sporidiobolus and 53 from the genus Sporobolomyces were described (Hamamoto et al., 2011, Sampaio et al., 2011). According to the taxonomic Non-conventional red yeasts for use in industries classification provided by the National Center for Biotechnology Information (as of March 2021), the two genera belong to the family Sporidiobolaceae, the order Sporidiales, the class Microbotryomycetes, the division Basidiomycota, and the kingdom Fungi. The genus Sporobolomyces includes the following 16 species: S. bannaensis, S. beijingensis, S. blumeae, S. carnicolor, S. inositophilus, S. japonicus, S. jilinensis, S. johnsonii, S. koalae, S. lactosus, S. longiusculus, S. patagonicus, S. phaffii, S. roseus, S. ruberrimus, and S. salmoneus. The genus Sporidiobolus includes the following three species: S. metaroseus, S. pararoseus, and S. salmonicolor. The differences in classification resulted from a taxonomic review by Wang et al. (2015a, b). Based on multigene sequence analysis, the researchers proposed the reorganization of the Pucciniomycotina subtype of yeasts. In line with the “One Fungus ¼ One Name” principle, new types of yeasts were separated, including those belonging to the genus Rhodosporidiobolus. In accordance with the phylogenetic analysis of seven genes and the analysis of the LSU rRNA domain, some species that were previously classified as Rhodotorula, Rhodosporidium, Sporobolomyces, and Sporidiobolus were included in the genus. Those species belonging to the new genus Rhodosporidiobolus reproduce asexually by budding, and in some cases by sexual reproduction. They can produce mycelium, pseudo-mycelium, and teliospores, which form septated basidia upon sprouting. The colonies of Rhodosporidiobolus yeasts are cream, pink, or red in color. Wang et al. (2015b) included the following nine species in this genus: R. azoricus, R. colostri, R. lusitaniae, R. fluvialis, R. microsporus, R. nylandia, R. odoratus, R. poonsookiae, and R. ruineniae. In 2020, three new species were included in the genus Rhodosporidiobolus: R. platycladi, R. jianfalingensis and R. fuzhouensis (Li et al., 2020a, b). 3. Morphology, physiology, and genetics Yeasts belonging to the genera Sporobolomyces and Sporidiobolus are commonly found in the environment. These microorganisms have low nutritional requirements and are widely classified as prototrophs. So far, they have been isolated from smoked dried sausages (Asefa et al., 2009), sedimentary rocks of glaciers in the Antarctica (Barahona et al., 2016), tree leaves (Wang and Bai, 2004; Cobban et al., 2016), nectarine fruits (Janisiewicz et al., 2010), fermented tea (Kim, 2009), Chinese miscanthus (Nakase et al., 1987), grapefruit (Sun et al.,  jcik et al., 2013), citrus fruits (Furuya et al., 2009), soils (Wo 2012), and apple must (Lorenzini et al., 2019). Colonies of Sporobolomyces appear cream, yellow-brown, salmon, orange, pink, or red in color on a solid medium (Fig. 2). The cells of this yeast are spherical, ellipsoidal, or cylindrical in shape. They reproduce asexually by polar budding, or less often by lateral or polygonal budding, but do not reproduce sexually, whereas species with this ability are classified in the genus Sporidiobolus (teleomorphic form). All species belonging to the genus Sporobolomyces produce symmetrically bilateral ballistoconidia (Hamamoto et al., 2011). The cells of Sporidiobolus yeast have an ovoid, elongated, or ellipsoidal shape. Their colonies appear orange, pink, or red on solid medium. They reproduce asexually by budding and forming ballistoconidia, or sexually by producing spherical 43 Fig. 2 e Characteristic growth of pigmented Sporidiobolus yeast strains on maltose agar (A e S. salmonicolor LOCK 275, B e S. salmonicolor CCY 19-6-4, C e S. pararoseus CCY 19-9-6, D e S. pararoseus ATCC 11386; LOCK e Culture Collection of Technical University of Lodz, Poland; CCY e Culture Collection of Yeasts, Slovakia; ATTC e American Type Culture Collection). teliospores, from which septated ovoid- or rod-shaped basidia form after germination. Terminal or intercalar teliospores can be formed on the hyphae. Some species require a partner for sexual reproduction and are referred to as heterothallic (selfsterile). For example, the strain Sporidiobolus pararoseus CBS 491 is sexually compatible with S. pararoseus CBS 484. On the other hand, strains that do not require a partner for sexual reproduction are termed homothallic (self-fertile) (Sampaio et al., 2011). Sporidiobolus and Sporobolomyces cannot ferment sugars. However, they have the ability to assimilate many compounds as a carbon source, including glucose, galactose, sucrose, maltose, trehalose, ethanol, glycerol, or raffinose. The type of compounds assimilated depends on the strain of the yeast species, which is considered useful in classical identification methods. The optimal temperature for the growth of most of their strains is 20e25  C. A characteristic feature of both Sporidiobolus and Sporobolomyces is the absence of xylose in cell wall hydrolyzates, and sugars namely mannose, glucose, galactose, and fucose are found dominant. Another characteristic feature is their positive reaction with diazole blue B in the DBB test and inability to synthesize starch-like compounds. In addition, the cells of both Sporidiobolus and Sporobolomyces produce coenzyme Q-10 (Hamamoto et al., 2011, Sampaio et al., 2011). The genomes of Sporidiobolus salmonicolor CBS 6832 (Coelho et al., 2015) and Sporobolomyces pararoseus NGR (Li et al., 2020a, b) have been described in recent studies. The total size of the genome of the former was determined as 20.52 Mb, and the G 44 A. M. Kot et al. þ C content as 61.3 %. The predicted number of genes was estimated at 5147. In the case of Sporobolomyces pararoseus NGR, the total genome size determined from the sequencing analysis was 20.9 Mb. The G þ C content of the NGR strain was lower compared to the CBS 6832 strain and amounted to 47.59 %. The predicted number of genes of the NGR strain was estimated at 5963. 4. Biomolecules produced by Sporidiobolus and Sporobolomyces yeast Exopolysaccharides Some yeast strains from the genus Sporobolomyces are capable of producing exopolysaccharides (EPS) (Table 1), cf., biopolymers produced extracellularly. These compounds accumulate on the cell surface in the form of mucus or are secreted into the culture medium. Extracellular polysaccharides protect cells against adverse conditions such as rapid temperature changes, drying, and phagocytosis, and also have cryoprotec ska-Jaroszuk et al., 2015). Aqueous tive properties (Osin solutions of EPS produced by yeast have been characterized by high viscosity and pseudoplasticity, which indicates that these compounds can be of use in the food, pharmaceutical, and cosmetic industries (Gientka et al., 2015). The synthesis of EPS is associated with the secondary metabolism of yeasts. Structure and physical properties of these compounds depend primarily on the yeast strain and the composition of the culture medium (Rusinova-Videva et al., 2010). Pavlova et al. (2004) identified that Sporobolomyces salmonicolor AL1 was the best producer of EPS out of the 38 tested strains of yeast isolated from lichen, moss, and soil collected from the region of the Bulgarian base on the Livingston Island (Antarctica). Yeast screening for the biosynthesis of polymers was carried out in media containing different sources of carbon and nitrogen. It was found that 5 % sucrose and 0.25 % ammonium sulfate were optimal for the synthesis of extracellular polysaccharides. The production of polymers was accompanied by a decrease in the pH of the culture medium from the initial value of 5.3 to 1.7e2.0 after 24 h of cultivation, which was maintained until the end of cultivation. During EPS biosynthesis, the viscosity of the culture medium increased to Table 1 e Efficiency of exopolysaccharide (EPS) and lipid biosynthesis by different strains of Sporidiobolus and Sporobolomyces. Strain Carbon and nitrogen source Exopolysaccharides Saccharose Sporobolomyces Ammonium sulfate salmonicolor AL1 Cultivation method Biosynthesis efficiency (g/L) Composition Pavlova et al. (2004) Batch 5.63 Fed-batch 10.23 Fed-batch 13.10 Fed-batch 14.30 Galactose:glucose:mannose:fucose Wang et al. (2020) 45:37:2:1 Crude glycerol Ammonium sulfate Olive oil Rice residue from canteen waste Ammonium sulfate Batch 6.4 Batch 8.35 Sporidiobolus pararoseus KM281507 Crude glycerol Ammonium sulfate Batch (Light 10000 Lux plus pure oxygen) 6.61 Sporidiobolus pararoseus CCTCC M 2010326 Glucose Corn steep liquor Fed-batch 51.0 Sporidiobolus pararoseus JD-2 Glucose Ammonium sulfate Fed-batch 47.1 Sporidiobolus metaroseus CCY 19-6-20 Glucose Ammonium chloride Oleic acid: 75.08 % Linolenic acid: 1.33 % Palmitoleic acid: 1.79 % Oleic acid: 62.13 % Palmitic acid: 20.55 % Stearic acid: 5.58 % Linolenic acid: 0.98 % Oleic acid: 81.2 % Palmitic acid: 10,3 % Stearic acid: 3.6 % Linolenic acid: 1.7 % Oleic acid: 72.5 % Palmitic acid: 15.7 % Stearic acid: 3.26 % Linolenic acid: 4.25 % Oleic acid: 69.1 % Palmitic acid: 16.3 % Stearic acid: 2.9 % Linolenic acid: 5.7 % Oleic acid: approx. 50 % Linolenic acid: approx. 20 % Palmitic acid: approx. 12 % Sporidiobolus pararoseus JD-2 Lipids Sporidiobolus pararoseus KX709872 Glucose Corn steep liquor Glucose Corn steep liquor Yeast extract Glucose Corn steep liquor 4.9 Glucose: 54.1 % Mannose: 42.6 % Fucose: 3.3 % Carbon: 35.46 % Oxygen: 49.29 % Galactose:glucose:mannose 16:8:1 References Han et al., (2016a) Han et al. (2018) Chaiyaso and Manowattana 2016 Chaiyaso et al. (2018) Manowattana et al. (2018) Han et al., (2016b) Wang et al. (2020)  et al. (2021) Kostovova Non-conventional red yeasts for use in industries a maximum of 15.37 mPa/s, and the highest amount of polysaccharides was produced (5.63 g/L) after 120 h. Addition of carbon sources at low concentrations to the medium was not suitable for EPS synthesis by Sporobolomyces salmonicolor AL1, since almost all the carbon was utilized for the production of cell biomass. Chromatographic analysis showed that the crude polysaccharide was of high purity (over 90 % carbon content) and composed of glucose (54.1 %), mannose (42.6 %), and fucose (3.3 %). Proteins and minerals were present in small amounts e 5.30 and 4.54 %, respectively. From the polysaccharide produced, pure mannan containing 98.6 % mannose was isolated. The rheological properties of this EPS were characterized using different arrays. Based on the dynamic viscosity of the medium and the flow index, it was found that this EPS can be used in the food industry as a thickening agent (Pavlova et al., 2004). In a study conducted by Han et al. (2018) investigated the rheological properties of EPS separated from the cultures of Sporidiobolus pararoseus JD-2. The maximum amount of EPS (10.23 g/L) was obtained after 72 h. Its molecular weight was determined by gel permeation chromatography to be 7.4  105 Da. Composition analysis showed that the main elements present in the separated polymer were carbon and oxygen at 35.46 and 49.29 % of dry substance, respectively. The electrokinetic potential of the polymer was 30 mV at pH 6.0, which proves that it is an anionic polymer. Selected rheological properties determined in the aqueous solutions of this polymer indicated that it exhibits similar characteristics to carboxymethyl cellulose and guar gum and can hence be used as a thickening agent in the food industry. In another work (Han et al., 2018), the authors optimized the composition of the medium and the conditions for polymer biosynthesis. The highest biosynthesis efficiency was achieved in the medium containing glucose (120 g/L), maize hydrolyzate (20 g/L), and yeast extract (10 g/L). After 72 h of batch-fed culture, an EPS yield of 13.1 g/L was obtained. Composition analysis showed that the polymer was composed of galactose, glucose, and mannose in a ratio of 16:8:1. Lipids Some yeast strains of the genera Sporobolomyces and Sporidiobolus are classified as oleaginous (Table 1) e microrganisms capable of synthesizing and accumulating lipids at above 20 % of dry cell substance (Patel et al., 2020). These microorganisms can be found in the natural environment. Ciu et al. (2012) isolated a psychrophilic yeast strain identified as Sporobolomyces roseus from the salt marshes in Nova Scotia (Canada). The highest content of lipids in dry cell substance (39.2 %) was obtained after cultivation in a medium containing 2 % glucose at 14  C. The extracted lipids were characterized by a favorable profile of fatty acids, with oleic acid (49.4 %), linoleic acid (14.3 %), and linolenic acid (6.7 %) found to be dominant. In another work (Matsui et al., 2012), Sporobolomyces carnicolor O33 (NBRC 107648) was isolated from soil in the Ibaraki Prefecture (Japan). This strain synthesized lipids at 26 % of dry cell substance in the xylose medium. Moreover, it was found that this carbon source had a positive effect on the biosynthesis of linoleic acid (21.4 %). Oleic acid (36.2 %) and palmitic acid (28.9 %) formed the highest shares in the pool 45 of fatty acids. In a study conducted by Han (2018), Sporidiobolus pararoseus JD-2 (isolate from chili sauce) was grown in a bioreactor in a medium containing glucose as a carbon source and corn steep liquor as a nitrogen source. Glucose was dosed into the medium to maintain its concentration at a level of 20e20 g/L. After 72 h of cultivation, 55 % of lipids were found in the yeast cell biomass. Following extraction, the main components of the microbial oil were identified as oleic (73.2 %), palmitic (17.3 %), and linoleic (4.24 %) acids. Additionally, 1 kg of the oil contained 1.32 g of squalene, 59 mg of b-carotene, 74 mg of g-carotene, 294 mg of torulene, 78 mg of torularhodin, and 4.07 g of ergosterol. Carotenoids In recent years, there has been growing interest in the natural methods of obtaining carotenoid pigments due to their health-promoting properties and potential applications (Mussagy et al., 2021a). Currently, chemical synthesis represent almost 80e90 % of the industry’s total demand for carotenoids, while a much lesser amount is obtained from the natural sources. Dissemination of healthy nutrition trends has increased the interest in natural carotenoids among both consumers and producers. Extraction of carotenoids from plants is expensive, and the quality of the dyes obtained is determined by the batch of raw material. Hence, microbial carotenoids are considered a valuable alternative. Several groups of microorganisms have the ability to produce carotenoids, imcluding filamentous fungi (Blakeslea trispora), algae (Dunaliella spp., Haematococcus spp.), yeasts (Phaffia spp., Rhodotorula spp., Sporobolomyces spp., Sporidiobolus spp., Rhodosporidium spp.), and bacteria (Flavobacterium spp., Micrococcus spp.). Microbial carotenoids extraction can be industrially useful when microorganisms that can synthesize a significant amount of dyes are used and the production cost can be minimized by using low-cost raw materials (e.g. industrial waste) as carbon and nitrogen sources and sustainable, biocompatible solvent for extraction/purification/polishing (Kirti et al., 2014; Mannazzu et al., 2015; Mussagy et al., 2020; Ram et al., 2020). The main carotenoids produced by the Sporobolomyces and Sporidiobolus yeasts are b-carotene, torulene, and torularhodin (Shi et al., 2013). 2-Hydroxytorularhodin and g-carotene were also identified in carotenoid extracts (Fig. 3) (Weber et al., 2005). In yeast cells, the major precursor of carotenoid biosynthesis is geranylegeranyl pyrophosphate (GGPP) containing 20 carbon atoms. Condensation of two GGPP particles, catalyzed by phytoene synthase (EC 2.5.1.32), results in phytoene, the first 40-carbon product of the pathway. After several steps, this compound is converted to lycopene, a precursor for the biosynthesis of g-carotene, b-carotene, torulene, and torularhodin (Fig. 4) (Li et al., 2017a, b; Kot et al., 2018). b-Carotene belongs to the group of carotenes, and its molecule contains two b-ionon rings. It is an antioxidant soluble in nonpolar solvents and exhibits provitamin A-like properties. These characteristics allow the use of b-carotene as an ingredient in cosmetics, vitamin preparations, and additives for animal feed (BogaczRadomska and Harasym, 2018). Torulene and torularhodin are precursors of vitamin A and display antiaging and antioxidant as well as immune-strengthening properties (Breierova 46 Fig. 3 e Structural formulas of carotenoids synthesized by Sporobolomyces and Sporidiobolus yeasts (A e b-carotene, B e torulene, C e torularhodin, D e 2-hydroxytorularhodin, E e g-carotene). et al., 2008). Torulene is a typical carotenoid of yeasts, has 13 double bonds, and is orange in color (Kot et al., 2018). It shows stronger antioxidant properties than b-carotene due to the greater number of conjugated double bonds in its structure (Mata-Gomez et al., 2014). Torularhodin is one of the few carotenoids with the properties of a carboxylic acid. It shows about 75 % of the activity of provitamin A in comparison to b-carotene. It is also characterized by high antioxidant activity due to the presence of 13 double bonds and polar nature (Kot et al., 2018). Over the last 15 y, many research teams have studied the biosynthesis of carotenoids by Sporobolomyces and Sporidiobolus and the properties of these compounds (Table 2). It was found that these yeasts can synthesize carotenoids in media containing wastes originating from various industries, which are partially utilized and thus minimize production costs. It Fig. 4 e Carotenoid biosynthesis pathway in yeast Sporidiobolus pararoseus, proposed by Li et al. (2017a, b). A. M. Kot et al. has been shown that waste animal fat can be used as a carbon source (Marova et al., 2017; Szotkowski et al., 2019). In a medium with hydrolyzed animal fat, Sporidiobolus pararoseus CCY 19-9-6 synthesized 24.8 mg/L of carotenoids. In addition, the obtained yeast biomass was rich in lipids (54.5 %) (Szotkowski et al., 2019). A significant amount of carotenoids was also synthesized by yeasts in media containing waste glycerol from biodiesel production as a carbon source. Cardoso et al. (2016) showed that the yeast strain Sporobolomyces ruberrimus H110 synthesized carotenoids at 0.51 g/L in a medium containing 3.35 % waste glycerol. The analysis of the carotenoid fraction revealed the presence of four compounds: b-carotene, g-carotene, torulene, and torularhodin e the dominant compound (69 %). Chaiyaso and Manowattana (2018) also used waste glycerol from biodiesel production as a carbon source for Sporidiobolus pararoseus KM281507. After 120 h of cultivation in a medium with 5.5 % glycerol, the yield of b-carotene biosynthesis was 15.76 mg/L, and total amount of carotenoids was 33.67 mg/L. After optimizing the temperature (24  C) and pH of the medium (5.6), the volumetric efficiency of biosynthesis increased to 27.41 and 53.70 mg/L, respectively. In the next stage, the authors determined the influence of the addition of Tween 20, Tween 40, Tween 60, Tween 80, and oleic acid on the biosynthesis of carotenoids. They found that the biosynthesis process was most effectively intensified when olive oil was added at a dose of 2 %. Under these conditions, the volumetric efficiency of the biosynthesis of b-carotene and total carotenoids was estimated at 54.43 and 70.93 mg/L, respectively. Probably, the KM281507 strain produced extracellular lipase and this enzyme hydrolyzed olive oil to free fatty acids and glycerol which were additional carbon sources. In another work (Manowattana et al., 2018), this yeast strain was cultivated in two types of bioreactors (stirred-tank and airlift). A production medium containing 5.5 % glycerol as the carbon source and ammonium sulfate as the nitrogen source was used for cultivation. Under optimized conditions (aeration rate: 6.0 vvm, dissolved oxygen: 60  5 %, light irradiation: 1000 lx), the volumetric efficiency of the biosynthesis of b-carotene and total carotenoids increased to 109.75 and 151.00 mg/L, respectively. In the next work (Manowattana et al., 2020), the parent strain KM281507 was successfully modified with ethyl methanesulfonate in combination with irradiation. The S. pararoseus E47 mutant was characterized by a biosynthetic efficiency of 128.97 mg/L (b-carotene) and 179.72 mg/L (total carotenoids) after cultivation in a medium containing 5.5 % glycerol in an airlift bioreactor. The carotenoids extracted from the biomass were microencapsulated in an alginate solution (0.5 %). Examination of the microcapsule morphology under a bright-field microscope showed that the carotenoid extract was completely covered with an alginate layer, as was confirmed by Nile red staining and fluorescence microscopy. Nonmicroencapsulated b-carotene degraded very quickly with a half-life of 13 d. The stability of b-carotene was extended by adding 1.0 % vitamin C to alginate. At 4  C, the half-life of bcarotene microencapsulated with vitamin C was found to be 63 d. This may be due to the fact that vitamin C efficiently converts the carotenoid radical cation (CARþ) into parent carotenoids (CARþ þ vitamin C / carotenoids þ vitamin Cþ). The conducted experiments thus revealed that after the Non-conventional red yeasts for use in industries 47 Table 2 e Carotenoid biosynthesis efficiency by various yeast strains of the genera Sporobolomyces and Sporidiobolus depending on the type of carbon and nitrogen source and the method of cultivation. Strain Sporidiobolus pararoseus TISTR 5213 Cultivation method Batch Carbon source Nitrogen source Glucose, malt extract Crude glycerol Yeast extract Yeast extract, ammonium sulfate Waste glycerol Yeast extract, ammonium sulfate Optimalized medium with waste glycerol Sporidiobolus pararoseus TISTR 5213 Sporidiobolus pararoseus CCT 7689 Batch Crude glycerol Yeast extract, ammonium sulfate Batch Raw glycerol Cane molasses Corn steep liquor Corn steep liquor Sporidiobolus pararoseus W8 Batch Glucose Peptone Sporidiobolus salmonicolor CBS 2636 Semicontinuous Glucose Crude glycerol Sporobolomyces ruberrimus H110 Batch Sporobolomyces ruberrimus ATCC 66500 Sporidiobolus pararoseus DAGIII Batch Glucose Pure glycerol Technical glycerol Glucose Peptone Corn maceration water, rice parboiling water Peptone, yeast extract Batch Sporidiobolus pararoseus JD-02 Cultivation conditions 25  C, 200 rpm, 120 h References 4.87 1.58 Manowattana et al. (2012) 1.77 23.9  C, 200 rpm, 5 d 24  C, pH 5.63, 120 h 16.55 27.41 Manowattana et al. (2015) 25  C, 180 rpm, 168 h pH 4.0, 180 rpm. 120 h 25  C, pH 4.0, 180 rpm, 96 h 0.78 0.52 Machado and Burkert (2015) 0.85 Cabral et al. (2011) 4.38 7.39 Colet et al. (2017) 1.77 1.56 4.81 Razavi and Marc (2006) 15.26 Saha et al. (2015) 45.08 Han et al. (2012) 15.0  Kostovova et al. (2021) 4.75 Colet et al. (2019) 3.13 Valduga et al. (2011) 27  C, pH 6.0, 210 rpm, 50 h Peptone, yeast extract, malt extract 27  C, pH 6.0, 210 rpm, Glucose Yeast extract, peptone Fed-batch Glucose Corn steep liquor Sporidiobolus pararoseus CCY 1909-06 Sporidiobolus salmonicolor CBS 2636 Batch Glucose Ammonium chloride 26  C, pH 5.3, 120 rpm, 120 h Dissolved oxygen: 20e30 %, 28  C, pH 6 23  C, pH 6.8, 96 h Fed-batch Crude glycerol Corn steep liquor, rice parboiling water Sporidiobolus salmonicolor CBS 2636 Batch Glucose, malt extract Peptone extraction process, yeast carotenoids can be stabilized by microencapsulation. After stabilization with vitamin C, yeast b-carotene microcapsules can be successfully used in the production of food, feed, and cosmetics (Manowattana et al., 2020). Ananda and Vadlani (2010) attempted to obtain a carotenoid-enriched feed product using a mixed culture of Phaffia rhodozyma ATCC 24202 and Sporobolomyces roseus ATCC 28988. They used a mixture of stillage obtained from the production of ethanol, glycerol, and corn steep as substrate. Under optimal conditions, P. rhodozyma produced 278.97 mg of astaxanthin/gd.m., while S. roseus produced 278 mg of b-carotene/gd.m.. The dried stillage is sold as fodder for cattle and poultry; therefore, enriching it with carotenoids Carotenoid biosynthesis efficiency (mg/L) 25  C, pH 4.0, 180 rpm, 1.5 vvm, 96 h 25  C, pH 4.0, 180 rpm, 1.5 vvm, 96 h will increase its nutritional value. In the next work (Ananda and Vadlani, 2011), the authors investigated the possibility of obtaining yeast feed additives using rice bran, wheat, and soybean waste products as substrates, after supplementation with 5 % glycerol. The highest concentration of astaxanthin (about 80 mg/gd.m.) was produced by P. rhodozyma and the highest concentration of b-carotene (about 836 mg/gd.m.) by S. roseus after cultivation with rice bran, as well as in a medium containing flour soybean. Assessment of the nutritional value of these bioproducts (Ananda and Vadlani, 2015) showed a reduction in crude fiber, crude protein, and amino acid content, while crude fat content was found to be increased in the medium with rice bran and decreased in the medium 48 with soybean flour. Increased levels of certain amino acids, such as hydroxyproline, hydroxylysine, and ornithine, were also reported by the authors. It should be emphasized that after the cultivation of red yeast, the trypsin inhibitor was reduced to an undetectable level in the soy flour bioproduct. Analysis of the polysaccharide content showed that the soybean flour product contained 1.01e3.23 % mannan and 2.43e3.52 % glucans, while 1.1e2.75 % mannan and 4.77e5.65 % glucans were present in the rice bran bioproduct. Results of the analysis of carotenoids and yeast cell wall polysaccharides in both bioproducts showed that they can be used in the production of compound feed with adequate nutrients based on the animal’s nutritional requirements. Li et al. (2016, 2017a, b, 2019) conducted advanced research on the process of torulene biosynthesis by the yeast Sporidiobolus pararoseus NGR (CGMCC 2.5280). This strain also synthesizes b-carotene and torularhodin at a lower yield. At the beginning (Li et al., 2016), the authors characterized phytoene desaturase (EC 1.3.5.5), one of the key enzymes involved in the pathway of carotenoid biosynthesis by yeast. This enzyme was found to be encoded by gene crtI, the genomic DNA of which is 2330-bp long and the cDNA is 1683 bp. The product of the crtI gene has a molecular weight of 62.28 kDa and consists of 560 amino acids. Phytoene desaturase has also been found to catalyze a series of enzymatic reactions involved in the conversion of phytoene to lycopene and 3,4didehydrolycopene. The conversion of intermediate products to torulene was presented by Li et al. (2017a, b). The authors attempted to investigate the effect of osmotic stress caused by the addition of sodium chloride on carotenoid biosynthesis by S. pararoseus NGR. After 5 d of incubation, the volumetric efficiency of carotenoid biosynthesis was significantly higher after cultivation in media supplemented with sodium chloride at 0.75 M (3.95 mg/L), 1 M (2.89 mg/L), and 1.5 M (1.99 mg/L) compared to a control sample without NaCl supplementation (1.63 mg/L). The osmotic stress conditions caused by sodium chloride supplementation significantly increased the efficiency of torulene biosynthesis. Lycopene and 3,4didehydrolycopene obtained as intermediates in the carotenoid biosynthesis pathway were transformed into g-carotene and torulene, respectively, by the enzymes encoded by gene crtI. The product of the crtI and crtYB genes in the medium with 0.75 M NaCl were determined to be 5.2 and 2.5 times higher, respectively, than in the control medium. The authors (Li et al., 2019) also sequenced the entire RNA transcriptome. Out of 3,849 differential expression genes (DEGs) in response to NaCl-induced osmotic stress, 2019 were found to be upregulated and 1,830 downregulated. Three carotenogenic genes e crtE, crtYB, and crtI e were identified among the DEGs. In addition, 14 genes that code for enzymes involved in the conversion of torulene to torularhodin were described: four hydroxylases, seven mono-oxygenases, and three oxidases. Carotenoids synthesized by yeast cells are intracellularly, that remaining inside of the yeast cells to guarantee their structural and functional properties (Mussagy et al., 2021a), and for efficient extraction, pre-treatments and cell wall disruption procedures are necessary. There is no exclusive method to obtain the best result for the carotenoids extraction (Mussagy et al., 2021b), the choice of the great method should guarantee not only the sustainability of the bioprocess but A. M. Kot et al. also the biological activity of the dyes recovered from natural sources. Lopez et al. (2017) tested the effect of mechanical and chemical disruption methods on the efficiency of carotenoid extraction from S. pararoseus CCT 7689 biomass. Chemical disruption was carried out using the following methods: dimethyl sulfoxide (DMSO), hydrochloric acid (4 M), acetic acid (4 M), and lactic acid (4 M). The mechanical techniques tested included the combination of ultrasound assisted method (UAE) at a frequency of 20 and 40 kHz, shaking with glass beads, grinding with diatomaceous earth, and maceration with liquid nitrogen. Among the investigated chemical techniques, the most effective extraction of carotenoids from yeast biomass was achieved using DMSO. The extraction efficiency was presented as the carotenoid content per gram of yeast dry matter, and the efficiency obtained with DMSO was 87.3 mg/gd.m.. After the use of hydrochloric, lactic, and acetic acids, the amount of carotenoids extracted from the biomass ranged only from 11.0 to 18.8 mg/gd.m.. Among the mechanical techniques tested, the most promising results were noted with the use of UAE at a frequency of 40 kHz, as well as with biomass grinding with glass beads (84.8 and 76.9 mg/ gd.m., respectively). Valduga et al. (2009) tested the effect of compressed gases, DMSO, and diatomaceous earth on the extractivity of carotenoids from S. salmonicolor CBS 2636 biomass. They also investigated the effect of the volatile organic solvents (VOCs) (DMSO, petroleum ether, hexane, ethyl acetate, chloroform, and acetone) on the efficiency of the extraction procedures. The highest volumetric yield of carotenoids (913 mg/L) was achieved after combined disintegration with liquid N2 and DMSO, followed by extraction with a mixture of acetone and methanol (7: 3, v/v). Monks et al. (2011) determined the effect of supercritical carbon dioxide and propane, as well as a combination of these methods and DMSO on the extraction of carotenoids. The highest efficiency (2.87 mg/L) was obtained when the yeast biomass was treated with supercritical CO2 (300 bar/120 min). An increase in the extractability of carotenoids from yeast biomass was also achieved by pretreatment of the biomass with lytic enzymes (Monks et al., 2013). The results of the previous studies showed that initial cell wall disruption and solvent used for extraction determines the efficiency of carotenoid recovery, and its parameters should be selected individually for each strain due to differences in the structure of the cell wall. Enzymes In 2019, the size of the global enzyme market was valued at $9.9 billion. About 80 % of the enzymes on the market were produced using microorganisms. The greatest use of enzymes has been recorded in the food, pharmaceutical, and feed industries (Grand View Research, 2020). Due to the increasing demand, manufacturers are looking for new enzymes of microbial origin. In the future, yeasts of the genera Sporobolomyces or Sporidiobolus can be used for enzyme production, as they can biosynthesize b-glucosidase (EC 3.2.1.21) (Ishikawa et al., 2005; Baffi et al., 2011, 2013), lipases (Mase et al., 2011; Ferraz et al., 2012; Smaniotto et al., 2012, 2014; Thabet et al., 2012), proteases (Kim, 2009, Bia1kowska et al., 2018), and amylases (Chaiyaso et al., 2018; Kwon et al., 2020). It was found that these Non-conventional red yeasts for use in industries yeasts are also capable of producing urease (EC 3.5.1.5) (Jahns, 1995), aldehyde reductase (EC 1.1.1.2) (Kataoka et al., 1992), carbonyl reductase (EC 1.1.1.184) (Li et al., 2009), and phenylalanine ammonia-lyase (EC 4.3.1.5) (Monge et al., 1995). b-Glucosidase b-Glucosidase (1,4-b-D-glucoside glucohydrolase, EC 3.2.1.21) catalyzes the breakdown of monoterpenes into monoterpenyl b-D-glucosides, which together with other compounds are responsible, for example, for the aroma of wine. Baffi et al. (2011) isolated Sporidiobolus pararoseus SP8A from grape skins. The yeast was cultivated in YP medium (10 g/L yeast extract, 20 g/L peptone) supplemented with various carbon compounds (glucose, sucrose, cellobiose) at doses of 5, 10, or 20 g/L. The highest b-glucosidase activity was noted after 72 h of cultivation in a medium containing 20 % of cellobiose. After isolation and purification, the enzyme was found to be the most active at 50  C and pH 5.5. Further experiments were performed using white wine and red wine which were added with 0.5 U of crude b-glucosidase and stored at 18  C (white wine) and 25  C (red wine) for 12 d. After storage, a significant increase in the concentration of geraniol, linalool, a-terpineol, and nerol was found in wines, which proves that yeast bglucosidase can be used in the production of wines to increase their aroma. The research was further continued by the authors (Baffi et al., 2013), and the yeast b-glucosidase was purified and characterized. The crude enzyme extract was purified in a four-step process involving concentration, dialysis, ultrafiltration, and ion-exchange chromatography. Electrophoretic studies showed that the purified enzyme had a molecular weight of approximately 63 kDa and an isoelectric point of 5.0. The specific activity of yeast b-glucosidase was determined as 20 U/mg. Purified b-glucosidase efficiently hydrolyzed terpene compounds, which confirmed their potential application as an additive to wines. Some strains of the species Sporobolomyces singularis can synthesize b-galactosidase-like enzyme. Ishikawa et al. (2005) obtained a yeast mutant from the parent strain S. singularis ATCC 24193 which showed a 10-fold higher b-galactosidase-like enzyme activity. After isolation and purification, the enzyme was found to belong to glycoproteins and exhibit b-glucosidase- and b-galactosidase-like activity. The molecular weight of the enzyme was 73.9 kDa, and the MichaeliseMenten constant values for o-nitrophenyl-D-galactopyranoside and p-nitrophenyl-D-glucopyranoside were determined as 5.40 and 1.96 mM, respectively. It was also found that the obtained enzyme was 35 % identical to the plant glucosidases belonging to the family of glycosyl hydrolases and can be used for the production of galactooligosaccharides. This was also confirmed in the study by Sakai et al. (2008). To bioconvert lactose to galactooligosaccharides, S. singularis cells immobilized in alginate were used, and the bioconversion process was carried out for 22 h in lactose solution (600 g/L) at various temperatures (50, 55, and 60  C) and pHs (4.0, 5.0, and 6.0) under constant stirring (80e100 rpm). The authors noted that the enzyme maintained its catalytic activity for 22 cycles (22 h/cycle, 440 h in total), and the highest yield of galacto-oligosaccharides (242 g/L) was obtained at 55  C and pH 5.5 or 6.0. 49 Lipases Lipases and esterases belong to the group of hydrolases and are used in various industries. For the industrial production of lipolytic enzymes, yeasts of the genus Candida, molds of the genus Rhizopus, and Pseudomonas bacteria are commonly used (Thabet et al., 2012). Some strains of Sporidiobolus salmonicolor (Thabet et al., 2012), Sporobolomyces ruberrimus (Ferraz et al., 2012), and Sporidiobolus pararoseus (Mase et al., 2011; Smaniotto et al., 2012, 2014) have been found to have the ability to synthesize these enzymes. Thabet et al. (2012) evaluated the lipolytic activity of 26 strains isolated in India in a factory processing sunflower oil. The highest lipolytic activity (8.2 U/ mL) against tributyrin was found after cultivation of the yeast OMV-15, which was identified as Sporidiobolus salmonicolor. After mutagenization with UV radiation, the mutants UV40 and UV70 exhibited a lipolytic activity of 18.2 and 18.9 U/mL. Chemical mutagenization with ethidium bromide allowed obtaining the OVS8 strain showing a lipase activity of 38.5 U/ mL after 96 h of cultivation in a medium at pH 6.0 and a temperature of 28  C. Ferraz et al. (2012) showed that Sporobolomyces ruberrimus isolated from soybean meal was able to efficiently synthesize lipolytic enzymes during solid-state fermentation using soybean flour, sugarcane pomace, and rice flour as substrates. Following the optimization of time and temperature of fermentation, the activity of lipolytic enzymes was determined as 130.1, 164.2, and 189.5 U/g for soybean flour, sugarcane pomace, and rice meal, respectively. Characterization of the enzyme confirmed its specificity for short-chain alcohols and fatty acids, as well as its stability and activity at 35 and 60  C. The highest activity of these lipolytic enzymes was noted at pH 6.5 and a temperature of 40  C. Smaniotto et al. (2012) isolated a strain of S. pararoseus that can biosynthesize lipolytic enzymes from soybean flour. The authors found that the most efficient biosynthesis of lipase occurred in a medium containing 5.0 % peptone, 6.8 % yeast extract, 7.0 % sodium chloride, and 1 % olive oil during cultivation at 30  C and 150 rpm. Under these conditions, the lipolytic activity was estimated at 26.9 U/mL after 72 h. In another work (Smaniott et al., 2014), lipase biosynthesis was carried out in a medium based on industrial waste and containing corn steep liquor, hydrolyzed yeast extract, waste frying oil, and sodium chloride. After 72 h of cultivation of the yeast S. pararoseus, the lipolytic enzyme extract showed an activity of 12.3 U/mL, which was significantly lower than that obtained in the conventional medium (26.9 U/mL). This was probably due to the fact that the substrate prepared from the waste contained compounds that can inhibit the lipase production process by yeast. The extracts of lipolytic enzymes were purified by fractionated precipitation with ammonium sulfate, and the enzymatic activity was determined as 154.6 and 120.9 U/g for the purified extracts from conventional and waste media, respectively. Both extracts were characterized by a high specificity for short-chain alcohols and fatty acids as well as thermal stability. Based on their results, the authors indicated the potential application of lipolytic preparations in the production of aromatic compounds. Similar observations were made by Mase et al. (2010) when they investigated the lipolytic properties of the Sporidiobolus pararoseus 25-A strain isolated from the digestive juice of Nepenthes truncata in Japan. The authors 50 A. M. Kot et al. found that this yeast synthesized lipase with a molecular weight of 37 kDa, and the enzyme exhibited optimal activity at pH 6.0 and a temperature of 60  C. They also analyzed the possibility of using this lipase in the dairy industry and found that its addition during the production of mozzarella improved the aroma and taste of the finished product. Based on these results, the authors indicated that this enzyme may be used as a substitute for pregastric esterase in the dairy industry in future. was also isolated from the sea algae Grateloupia sp. (Kwon et al., 2020). The PHGra1 enzyme produced by this strain showed the highest amylolytic activity at 55  C and pH 6.5 with the addition of 0e3.0 % NaCl, and was stable during storage at a temperature of 15e45  C. It showed more specificity for potato starch compared to corn and wheat starch. Proteases In recent years, consumers prefer foods that contain only raw materials of natural origin. This has encouraged the biotechnologists to investigate the possibilities of using yeast in the production of flavoring substances such as g-decalactone. In 1930, Derx isolated a yeast strain similar to Sporobolomyces roseus from orange leaves and described it as Sporobolomyces odorus because its culture was characterized by an aromatic smell (Tahara et al., 1972). The characteristic fruity smell was later attributed to the presence of lactones (Tahara et al., 1972, 1973). g-Decalactone (C10H18O2) was found to be the main compound responsible for the fruity odor of the S. odorus yeast culture (now Sporidiobolus salmonicolor). It is a cyclic ester containing a closed ring composed of four carbon atoms and one endocyclic oxygen atom, coupled to an adjacent ketone group. In addition to Sporidiobolus and Sporobolomyces, yeasts of the genera Yarrowia, Pichia, Candida, and Rhodotorula, among others, have also been found to biosynthesize g-decalactone. The compound is characterized by a peach aroma and taste at concentrations above 5 mg/L. It also occurs in fruits including peach, apricot, and strawberry (Braga and Belo, 2016). g-Decalactone can be obtained by b-oxidation of ricinoleic acid, the main component (approximately 90 %) of castor oil. Feron et al. (2005) found that in some strains of Sporidiobolus g-decalactone synthesis took place in microcells, but in others in mitochondria. The precursor for the biosynthesis process was 4-hydroxydecanoic acid (Feron et al., 1996). BlinPerrin et al. (2000) examined four species of yeast (S. salmonicolor, S. ruinenii, S. johnsonii, and S. pararoseus) to determine the differences in the course of b-oxidation of ricinoleic acid. Their in vitro and in vivo studies revealed that the levels and composition of intermediates in fatty acid oxidation differed between the species. Based on the obtained results, the authors hypothesized that different pathways of ricinoleic acid oxidation are involved in the yeast strains. The efficiency of g-decalactone biosynthesis depends, for example, on the producer strain, the composition of the growing medium, and the conditions as well as method of cultivation. Castor oil or ricinoleic acid esters are most often used as a substrate for the biosynthesis of g-decalactone. Lin et al. (1996) examined how replacement of this substrate with palmitic, stearic, oleic, or linoleic acid would affect the growth of S. odorus AHU 3246 and the subsequent production of lactone. Their results showed that the growth of yeast in media containing palmitic, stearic, or oleic acid was similar to that determined under the control conditions, but the presence of linoleic acid caused a significant reduction in growth. In the media containing the tested fatty acids, a decrease in the productivity of g-decalactone (about 3.5 mg/L) was noted in comparison to the medium containing ricinoleic acid (71.8 mg/L). Proteolytic enzymes belong to the class of hydrolases and catalyze the hydrolysis of peptide bonds. In the food industry, these enzymes are used mainly in the production of cheese and soy sauces, for softening meat, and as an additive to bread. In the brewing industry, they are used for the extraction of proteins from barley and malt. Proteases are also used in the production of juices to remove proteins that cause clouding (Bond, 2019). Bia1kowska et al. (2018) found that the psychrotrophic yeast strain Sporobolomyces roseus LOCK 1119 isolated from groundwater from the Luiza silver and lead mine in Zabrze (Poland) synthesized an extracellular protease which exhibited an activity of about 560 U/L in YPD medium. After optimizing the composition of the medium (glucose 60 g/L, beef extract 40 g/L, yeast extract 20 g/L, bovine serum albumin 10 g/L), the yeast strain synthesized a protease with an activity of 2059.6 U/L, while a temperature decrease from 20 to 10  C caused an increase in the enzyme activity to 2555.3 U/L. It has been shown that the enzyme belongs to the group of aspartyl proteases. The G8 protease showed the highest activity at 50  C. By using the techniques of ionexchange chromatography and gel filtration, a highly purified enzyme was obtained. The enzyme showed substrate specificity for aromatic and hydrophobic amino acids. Furthermore, the ability of the enzyme to produce peptides with antioxidant properties was tested and the highest antioxidant activity (69 %) was recorded for bovine casein hydrolyzates. The Sporidiobolus ruineniae CO-3 strain (currently Rhodosporidiobolus ruineniae) isolated from fermented tea was also found with the ability to synthesize proteolytic enzymes during cultivation (Kim, 2009). The protease activity was highest after the cultivation of this yeast in a medium containing 1 % xylose, 1 % yeast extract, and 0.3 % K2HPO4. The highest activity of the protease was found at pH 7.0 and a temperature of 50  C. Amylases Recent research (Chaiyaso et al., 2018; Kwon et al., 2020) showed that some strains of the species Sporidiobolus pararoseus have the ability to produce enzymes that can break down starch (i.e. amylases). These are a group of enzymes that hydrolyze a-glucosidic bonds in the starch molecule. The products of these reactions include, among others, dextrins, maltose, and glucose. Amylolytic enzymes are commonly used in the food, alcohol, and chemical industries. Chaiyaso et al. (2018) isolated the yeast strain S. pararoseus KX709872 from flowers and simultaneously showed that the strain synthesized a-amylase (540 mU/mL) and amyloglucosidase (23 mU/mL) in media containing waste resulting from rice production. The amylolytic yeast strain S. pararoseus g-Decalactone Non-conventional red yeasts for use in industries Fed-batch and repeated-batch cultivation methods are one of the ways to significantly increase the productivity of g-decalactone by yeasts. Fed-batch cultivation involves periodic or continuous dosing of sterile medium (or specific nutrients) into the bioreactor. The products of microbial metabolism remain inside the bioreactor until the end of the culture. On the other hand, during the repeated-batch process, a part of the medium with biomass and metabolites is removed and replaced with a fresh portion of the medium. Feed-in batch processes are used to prevent the substrate or metabolic products from inhibiting the biocatalyst activity. Such type of process was used by Lee et al. (1995) for the cultivation of S. odorus yeast (now S. salmonicolor) in a 5-L bioreactor. In the batch culture conditions, the authors obtained the highest amount of gdecalactone (54.6 mg/L) after 120 h. A significant increase in productivity was observed in the fed-batch cultures, in which the g-decalactone yield increased with the number of dosages of castor oil hydrolyzate on the 3rd, 4th, and 5th day of cultivation and after 7 d (208 mg/L). In another study, Dufosse et al. (1998) cultivated the yeast Sporidiobolus ruinenii in a 7-L bioreactor and collected a part of the medium every 10 d while feeding the same volume of fresh medium. They achieved the highest amount of g-decalactone when a third batch of fresh medium was added. The lactone content estimated at the end of the process was 5.5 g/L, and the productivity was 0.39 mg/L/day. It is known that g-decalactone is highly toxic to microorganisms, including the producer strains. The toxicity of this compound is probably related to the change in the structure and permeability of cell membranes (Wache et al., 2003). However, this phenomenon can be reduced by following yeast immobilization. For instance, Lee et al. (1999) used calcium alginate to immobilize the yeast Sporidiobolus salmonicolor CCRC 21975. The authors observed that the immobilized yeast cells were able to produce this compound, and its concentration increased during cultivation. Furthermore, they found that the initial pH of the medium significantly influenced the production of g-decalactone, which ranged from 75.0 to 114.7 mg/L and was the highest in the medium with an initial pH of 4.0. Immobilized yeast cells more efficiently synthesized lactone than free cells. One of the most important advantages of cell immobilization is that the immobilized material can be reused. The authors used the immobilized S. salmonicolor yeast cells in 13 4-day production cycles. After the first cycle, the efficiency of g-decalactone biosynthesis was 114.7 mg/L, while after the next cycle it was in the range of 67.0e104.0 mg/L. At the end of the last cycle, the production of lactone was determined as 58.4 % which was higher compared to the value obtained after the first cycle. Dufosse et al. (1999) tested three techniques to reduce the toxicity of g-decalactone toward yeast cells in a medium with castor oil. They carried out in situ trapping using various oily phases, porous hydrophobic sorbents, and b-cyclodextrins. It was found that trapping of Sporidiobolus salmonicolor cells in olive oil or Miglyol resulted in an increase in the productivity of g-decalactone as well as yeast viability. On the other hand, with the use of hydrophobic b-cyclodextrins, satisfactory cell viability was not achieved, while trapping in porous hydrophobic sorbents caused a 30 % reduction in the amount of g-decalactone, although the yeast cell viability 51 was not affected. The conducted experiments thus showed that the negative effect of g-decalactone can be reduced by using cheap oils as trapping agents. 5. Potential applications of biomolecules produced by Sporidiobolus and Sporobolomyces yeast Feed additive The use of Sporidiobolus yeast as a potential animal feed additive is a new direction of research. In 2017, Tapingkae et al., 2017 analyzed how feed supplementation with dried S. pararoseus CMU-THA52 yeast will affect the production performance and egg quality of laying hens. The feed supplemented with Saccharomyces cerevisiae at a dose of 2 g/kg was used as a reference sample in the experiments. After 12 weeks of use of the supplemented feed, the authors found no significant differences in the productivity of laying hens and the weight of eggs. Supplementation of feed with the biomass of S. pararoseus yeast increased the intensity of the yolk color. The highest intensity of yolk color was noted in the case of hens fed with fodder supplemented with 2 g/kg of Sporidiobolus yeast. Moreover, in the same group, lower levels of cholesterol and triglycerides were found in serum and egg yolk. In this work, the authors used dried yeast biomass, which brings many benefits. Compared to wet yeast biomass, the use of dried yeast reduces the cost of storage, transportation and provided consistency of quality for consumers (Rapoport et al., 2016). Yeast can also be used as an additive to fish feed. Strains that produce enzymes capable of converting tannins, which are classified as antinutritional compounds, may be of particular importance. In the feed ingredients used in aquaculture, the antinutritional compounds come from plants used as a protein source (soybean meal, rapeseed meal, pea meal, and mustard cake). To reduce the content of tannins, addition of tannase (EC 3.1.1.20) is required. Kanpiengjai et al. (2016, 2020) proposed adding live cultures of yeast Sporidiobolus ruineniae A45.2 to the feed instead of enzymes. The strain used in their study was isolated from fermented tea leaves and was found to have a unique cell wall structure that is associated with resistance to a high concentration of tannins. The strain produced thermostable cell-associated tannase that can degrade tannic acid into gallic acid (Kanpiengjai et al., 2016). In another work (Kanpiengjai et al., 2020), these yeasts were analyzed for their potential probiotic properties. In the model conditions of the gastrointestinal tract (pH 2.0, pepsin, bile salts, and pancreatin), 90 % survival of yeast cells and maintenance of tannase activity were noticed. The supernatant of S. ruineniae A45.2 culture showed antimicrobial activity against Gram-positive bacteria: Bacillus cereus, Staphylococcus aureus, and Streptococcus agalactiae. It was also characterized by high antimicrobial activity, which in the FRAP, DPPH, and ABTS assays was estimated at, respectively, 9.2  1.8, 9.0  0.9, and 9.8  0.7 mg gallic acid equivalent/mL supernatant. The obtained results showed that the S. ruineniae A45.2 strain may be a potential multifunctional feed additive. However, further animal studies should be performed to confirm the in vivo properties of this yeast. 52 A. M. Kot et al. Table 3 e Patents for the biotechnological use of the Sporidiobolus and Sporobolomyces yeast (source: World Intellectual Property Organization). Number Year Title Additional information 111793569 2020 111117903 2020 20190338327 2019 High-yield fermentation method of carotenoid high-yield Sporobolomyces strain and application of method Marine-derived Sporidiobolus pararoseus lyophilized preparation, as well as preparation method and application thereof A process for production of galacto-oligosaccharides 110396478 2019 109837317 2019 Sporobolomyces reseus and method for optimized extraction of carotenoid by using response surface method Method for synthesizing chiral bisaryl alcohol compound 109337831 2019 Heavy metal resistant microorganism and application thereof 110257266 2019 109554410 2019 109055245 2018 PCT/EP2016/068009 2017 106256910 2016 105994942 2016 Sporidiobolus pararoseus strain and application thereof in production of lactase Fed-batch feeding method for high-density fermentation of Sporidiobolus pararoseus Ocean source Sporidiobolus pararoseus and application thereof in preventing and controlling strawberry diseases A Sporobolomyces roseus strain for the production of compositions with colorant and antioxidant properties Method for removing arabinose in wheat bran oligosaccharide through Sporobolomyces singularis fermentation Preparation method of feed additive for laying ducks 2018038345 2016 PI9811518 2015 102766580 2012 102719367 2012 PCT/US2011/032275 2011 101037657 PCT/EP2006/005671 2007 2006 20050260293 2005 1584010 2005 2004313187 2004 New strain of Sporidiobolus pararoseus, food and drink, method for producing food and drink, skin cosmetics, and method for producing skin cosmetics ~ o para biocontrole das doenças de planta e me todo Composiça  s plantas proteça ~ o contra pato  genos para fornecer a Yeast strain for producing biosurfactant and application thereof Catalytic synthesis method of L-theanine by using microorganism-produced gamma-glutamyl amino carboxamide synthase Coenzyme Q10 production using Sporidiobolus johnsonii Country - China - China Galacto-oligosaccharides from Sporobolomyces singularis - United States of America China Carbonyl reductase SSCR derived from Sporobolomyces salmonicolor Microorganism: Sporobolomyces carnicolor MWT-01 - China - China - China - Spain - China Microorganism: Sporobolomyces sp. CICC33080 - China Microorganism: Sporobolomyces roseus Microorganism: Sporidiobolus salmonicolor AH3 Microorganism: Sporidiobolus pararoseus CCTCC M2012232 - Brasil China China Japan China China United States of America China Germany Huaxi Sporobolomyces and its application Metabolic engineering of Q10 production in yeasts of the genus Sporidiobolus Biocontrol for plants with Bacillus subtilis, Pseudomonas putida, and Sporobolomyces roseus - Process for producing Sporidiobolus ruineniae strains with improved coenzyme Q10 production Mutagenized Sporidiobolus ruineniae strain, process for producing the same, and method for producing improved coenzyme Q10 - United States of America China - Japan Biocontrol of plant materials Spoilage of fruits and vegetables at various stages of food distribution and sale accounts for significant financial losses. Fruit and vegetable raw materials are unstable and favor the development of various microorganisms, including pathogenic ones. Various techniques can be used to inhibit the - growth of microorganisms and extend the shelf-life of raw material. The use of biological control agents (BCAs) is an example of such techniques (Quaglia et al., 2011). Biocontrol refers to the use of one population of microorganisms to inhibit the growth of another. Selected molds, yeasts, and bacteria that are safe for humans can be used as BCAs. Of these, yeasts are particularly important because they can use Non-conventional red yeasts for use in industries numerous nutrients, propagate quickly, have a high ability to colonize dry surfaces for a long time, and produce exopolysaccharides capable of protecting their cells against adverse environmental factors (Abdelhai et al., 2019b). Strong antagonistic properties are shown by the following yeast species: Hanseniaspora uvarum, Y. lipolytica, Cryptococcus laurentii, and Metsch€ rkel nikowia pulcherrima (Qin and Tian, 2005; Liu et al., 2010; Tu et al., 2014; Zhu et al., 2019). In recent years, the possibility of using certain yeast strains of the genera Sporobolomyces and Sporidiobolus in fruit biocontrol has also been demonstrated. One example is the research conducted on the properties of the strain Sporidiobolus pararoseus Y16 by Li et al. (2017a, b) and Abdelhai et al. (2019a, b). Li et al. (2017a, b) found that S. pararoseus Y16 significantly inhibited the growth of Aspergillus niger on grapes. After the storage period, the fruit treated with yeast cells showed higher activity of polyphenol oxidase (PPO, EC 1.14.18.1), catalase (CAT, EC 1.11.1.6), phenylalanine ammonia-lyase (PAL, EC 4.3.1.5), and ascorbate peroxidase (APX, EC 1.11.1.11), which resulted from the greater expression of PPO, CAT, PAL, and APX genes. Based on this, it was concluded that increased gene expression and enzyme activity must have contributed to biocontrol. Abdelhai et al. (2019a) investigated the possibility of inhibiting the growth of Penicillium expansum using the same strain of S. pararoseus Y16 yeast and the extract of Adansonia digitata L. (100 mg/ mL). The applied combination of natural antagonistic factors significantly slowed down the development of P. expansum at 4  C for 30 d and at 20  C for 15 d. Moreover, these factors showed no negative influence on the organoleptic and quality characteristics of apples. Abdelhai et al. (2019b) showed that the antagonistic activity of S. pararoseus Y16 strain can be increased by additionally using a solution of N,N,Ntrimethylglycine at a concentration of 1 mM. With the use of these two factors, a significant inhibition was found in the spore germination of P. expansum, as well as a reduction in the size of the mycelium and the length of the hyphae. These results indicate the possibility of using the Sporidiobolus pararoseus Y16 strain as a natural additive to extend the shelf-life of grapes and apples after harvesting. The yeast Sporidiobolus pararoseus can also be used to protect strawberries against the development of gray mold. Botrytis cinerea infects leaves, flowers, and stems, as well as healthy fruits due to the production of large amounts of spores. Thus, this species causes damage both before and after harvest. Shen et al. (2019) isolated the strain S. pararoseus ZMY-1 from the mangrove swamp. This strain significantly inhibited the growth of B. cinerea on strawberries at a dose of 108  108 CFU/mL, while not affecting the fruit quality. The antagonistic effect was observed at both 20 and 4  C. Similar observations were made by Huang et al. (2012) when they investigated the antagonistic properties of Sporidiobolus pararoseus YCXT3. They found that inoculation of strawberries with a yeast suspension (105e106 CFU/mL) reduced the incidence of gray mold in the fruit from 96 to 100 % (control group) to 39e50 %. In addition, 39 volatile organic compounds were synthesized by S. pararoseus YCXT3, including 2-ethyl-1hexanol which has strong antifungal properties and was responsible for the biocontrol mechanism of the tested yeast strain. 53 Carvalho et al. (2020) found that the live yeast Sporidiobolus johnsonii AH 16-1 (strain isolated from Impatiens parviflora leaves) can be an effective BCA to protect beans (Phaseolus vulgaris L.) against bacterial blight caused by the bacterium Xanthomonas axonopodis pv. phaseoli, which can significantly reduce the crop yield. The authors showed that triple application of S. johnsonii AH 16-1 yeast reduced the disease development by 58.42 %. Some strains of Sporobolomyces yeasts have the ability to degrade the patulin e a mycotoxin with mutagenic, genotoxic, immunotoxic, teratogenic, and cytotoxic properties. It was found (Ianiri et al., 2013) that the strain Sporobolomyces sp. IAM 13481 can degrade patulin and convert it into deoxypatulinic acid and ascladiol, which are less toxic than patulin. 6. Conclusions and future perspectives This literature review of the characteristics and biotechnological use of the Sporobolomyces and Sporidiobolus yeasts allowed organizing the knowledge on modern taxonomy, which, due to the development of genetic engineering, has been constantly changing in recent years. One example is the reclassification of some previously classified Sporobolomyces and Sporidiobolus species to the new genus Rhodosporidiobolus. It should be assumed that in the coming years, further taxonomic revisions are possible, and unknown yeast species can be isolated, for example, from extreme environments. The lipids produced by the Sporobolomyces and Sporidiobolus yeasts were found to have a favorable fatty acid composition and could be used as a substitute for traditional vegetable oils in the production of biodiesel. These yeasts also synthesize carotenoids (mainly b-carotene, torulene, torularhodin), which have provitamin A, antioxidant, and antimicrobial properties. Both lipids and carotenoids can be synthesized by yeasts in the media containing agro-food waste as a source of nutrients. This leads to a reduction in the costs of media, while allowing partially management of the waste. Furthermore, Sporobolomyces and Sporidiobolus yeasts can also utilize by-products from various industries due to their ability to synthesize a wide range of enzymes such as lipases, proteases, and amylases. The valuable properties of these types of yeast include production of g-decalactone by b-oxidation of fatty acids. This compound is characterized by a peach aroma and has a great application potential in the cosmetic and food industries. Some strains of the genera Sporobolomyces and Sporidiobolus can be used in the production of preparations that can be used as an alternative to chemicals for the protection of plant materials after harvest, as they show antagonistic properties against plant pathogens such as P. expansum and B. cinerea. Some strains have also been found to break down patulin. Therefore, it is expected that research on the decomposition of mycotoxins will be continued. Many research works on the biotechnological use of Sporidiobolus and Sporobolomyces yeasts have been patented (Table 3). 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