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. 2024 Apr 9;17(4):e14449. doi: 10.1111/1751-7915.14449

Harnessing the dual nature of Bacillus (Weizmannia) coagulans for sustainable production of biomaterials and development of functional food

Emanuela Maresca 1, Martina Aulitto 1,2,✉, Patrizia Contursi 1,3,4,5
PMCID: PMC11003712  PMID: 38593329

Abstract

Bacillus coagulans, recently renamed Weizmannia coagulans, is a spore‐forming bacterium that has garnered significant interest across various research fields, ranging from health to industrial applications. The probiotic properties of W. coagulans enhance intestinal digestion, by releasing prebiotic molecules including enzymes that facilitate the breakdown of not‐digestible carbohydrates. Notably, some enzymes from W. coagulans extend beyond digestive functions, serving as valuable biotechnological tools and contributing to more sustainable and efficient manufacturing processes. Furthermore, the homofermentative thermophilic nature of W. coagulans renders it an exceptional candidate for fermenting foods and lignocellulosic residues into L‐(+)‐lactic acid. In this review, we provide an overview of the dual nature of W. coagulans, in functional foods and for the development of bio‐based materials.

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INTRODUCTION

Bacillus coagulans was initially described in 1915 by Hammer, who isolated strain ATCC 7050 (also referred to as DSM1) from spoiled canned milk (Su et al., 2012). Recently, it has been phylogenetically reclassified as a distinct monophyletic clade called Weizmannia (Gupta et al., 2020). W. coagulans is a Gram‐positive, lactic acid‐producer, non‐pathogenic and spore‐forming bacterium (Konuray & Erginkaya, 2018) which is naturally distributed in a variety of niches as well as in food fermented from soybean, locust bean, maize, rice and others (Cho et al., 2023; Shudong et al., 2022). Since it has several health benefits, such as the production of prebiotic molecules, prevention of muscle damage during exercise, improvement of gastrointestinal disorders, ease of diarrhoea and prevention of bacterial vaginosis, it is widely used as a commercial probiotic, often in combination with other strains from Bacillaceae and Lactobacillaceae families (Bang et al., 2021; Kim et al., 2021).

Unlike other well‐known probiotics, W. coagulans thrives optimally at temperatures between 35 and 50°C and a pH close to 6 (Rhee et al., 2011). It is regarded as “the king of probiotics” because of its ability to form spores, exhibiting strong resistance and survival rate under harsh conditions, i.e., low‐oxygen environments of the gastrointestinal tract, stomach acids and bile salts as well as food processing (Zhou et al., 2020). For these reasons, W. coagulans is widely used in commercial foods such as dairy products, fermented meats and cereals (Konuray & Erginkaya, 2018). Worth‐mentioning is the secretion of extracellular carbohydrate‐ and protein‐degrading enzymes that enhance the quality and sensory properties of fermented foods (Konuray & Erginkaya, 2018; Olajuyigbe & Ehiosun, 2013). W. coagulans also produces compounds with antimicrobial activity against Gram‐positive and Gram‐negative food pathogens (Kim et al., 2021). The biotechnological potential of W. coagulans does not reside only in the food industry. Indeed, it is widely used also as a microbial cell factory for the eco‐sustainable production of lactic acid (LA), providing an alternative to chemical synthesis (Huang, Tian, et al., 2023; Huang, Wang, et al., 2023; Pamueangmun et al., 2023). The moderately thermophilic nature of W. coagulans strains reduces the risk of microbial contamination, making them naturally suited for lactate production even in open fermentations (Aulitto et al., 2019). The ability to operate at high temperature also reduces the energy consumption needed for sterilization and cooling of the fermentation system (Su & Xu, 2014). To date, several W. coagulans strains have been utilised for LA production from lignocellulosic materials due to their temperature compatibility (50–55°C) with commercial enzymes currently used for the biomass saccharification (Li et al., 2021; Pol et al., 2016; Rhee et al., 2011; Walton et al., 2010).

Some molecular biology studies have been addressed to analyse the physiological response of W. coagulans during fermentation. For instance, the steam‐explosion pre‐treatment of lignocellulosic biomasses releases toxic compounds that hinder LA production as demonstrated for the MA‐13 strain (Aulitto et al., 2017, 2019). Moreover, a comparative transcriptomic analysis revealed that sodium lactate induced downregulation and upregulation of glycolysis/gluconeogenesis and TCA cycle genes, respectively, thus affecting LA fermentation (Qin et al., 2015).

Moreover, a time‐resolved transcriptomic and proteomic study shed light on the response mechanisms of W. coagulans during NaOH‐buffered LA fermentation, highlighting the expression levels of the glycolytic and TCA cycle genes throughout the fermentation process (Huang, Tian, et al., 2023; Huang, Wang, et al., 2023).

Upon the identification of multiple strains of W. coagulans (65 strains reported on NCBI on 21 December 2023), comparative genomic studies have revealed that some strain‐specific genetic traits can be exploited for diverse biotechnological fields. For instance, pangenomic scans offer insights into the comprehensive landscape of central carbon metabolism, as well as into the absence/presence of active virulence factor genes and/or prophage sequences highlighting that not all the available strains are suitable for targeted applications, especially in the field of the food industry (Aulitto et al., 2022; Su & Xu, 2014; Sun et al., 2023). Currently, only a limited number of reports are available on the potential role of W. coagulans as a whole‐cell biofactory for the effective production of bioactive compounds (biomolecules and fermentation bioproducts). This mini‐review aims to highlight the most recent advancements in W. coagulans as a probiotic as well as a producer of prebiotics and biotechnological relevant biomolecules in the food and feed sectors. Moreover, its emergence as a model system for the eco‐sustainable synthesis of LA‐based biomaterial is discussed with relevant examples from agri‐food waste conversion.

FUNCTIONAL FOODS

Functional foods (often indicated also as ‘natural health products’ or ‘healthy foods’) exhibit beneficial effects on specific bodily functions beyond their nutritional value, such as improving general healthy conditions and/or decreasing the risk of chronic disease appearance (Henry, 2010; Kaur & Das, 2011). The global market for functional foods and beverages reached a value of $192 billion in 2020, driven by their increasing integration into consumers' daily diets worldwide (Konstantinidi & Koutelidakis, 2019). A well‐known example of functional food is yogurt enriched with fruits, vegetables, cereals and other active compounds (such as honey, moringa, aloe vera extracts and essential oils etc) (Ahmad et al., 2022). Functional foods often contain bioactive compounds that support the activity of probiotics, by preventing the growth of pathogens and/or inhibiting the production of virulence factors (Damián et al., 2022). Overall, the combination of functional foods and probiotics promotes a synergistic effect on nutritional health (Ballini et al., 2023) and offers a reliable approach to enhance the ability of the probiotics to compete with harmful bacteria, supporting immune function and improving overall well‐being (Silva et al., 2020).

Specifically, it is well known that W. coagulans contributes significantly to digestive health, nutrient absorption and human health through various mechanisms, including enzyme production, metabolite generation and modulation of the gut environment, as summarised in the Table 1 (Cao et al., 2020; Palomino et al., 2023).

TABLE 1.

Beneficial health effects of W. coagulans consumption.

Beneficial effects on the health of the host References
Interaction with gut microbiota The consumption of W. Coagulans has been associated with an increase of lactobacillus and Bifidobacteria, while also competitively outcompeting opportunistic pathogens such as Clostridium difficile Reduction of toxic metabolites Haldar and Gandhi (2016), Ianiro et al. (2016), Gibson et al. (2014), Kodali and Sen (2008)
Immunity response

  • W. coagulans PTA‐6086 enhances the activity of various human immune cells (increased phagocytic activity, maturation of mononuclear phagocytes).

  • W. coagulans reinforces the immune system and alleviates inflammation through cell‐derived mediators, modifying the production of immune‐activating, anti‐inflammatory cytokines, chemokines and growth factors.

 Benson et al. (2012), Jensen et al. (2010), Jensen et al. (2010), Sudha et al. (2015)
Amino acid absorption

  • Absorption of specific amino acids, including those involved in blood flow regulation (citrulline) and recovery (glutamine).

 Jäger et al. (2018)
Metabolite production

  • W. coagulans produces metabolites like diacetyl, short‐chain fatty acids (SCFAs) and vitamins.

  • Stimulates intestinal peristalsis, reducing harmful substances such as amines and promoting healthy bowel movements.

 Nyangale et al. (2014)
Other health benefits associated with the intake of W. coagulans
  • Constipation Relief:
Studies show that W. coagulans administration effectively relieves constipation.
  • Cholesterol Reduction:
Animal studies demonstrate that W. coagulans can reduce cholesterol levels by degrading cholesterol in blood serum.
  • Bile Acid Metabolism Improvement:

Combination of soy pulp and B. coagulans lilac‐01 improves intestinal bile acid metabolism

 Aminlari et al. (2019), Lee et al. (2016b), Minamida et al. (2015)
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W. coagulans in food industries

Lactic acid bacteria (LAB), bifidobacteria and yeasts are well‐researched and commonly used as probiotics in a wide range of food products (Palanivelu et al., 2022). However, the harsh environmental conditions encountered during food processing and storage often compromise the viability and activity of probiotics, thus affecting their effectiveness (Terpou et al., 2019). For instance, several Lactobacillus or Bifidobacterium spp. employed in functional foods formulations for the management of different intestinal disorders (Palanivelu et al., 2022), require freeze‐drying processes or powdered formulations, which have a detrimental effect on their viability. It is worth noting that most of the W. coagulans strains isolated so far exhibit thermal resistance and high survival rates in gastrointestinal traits (Zhang et al., 2024), thus enhancing its potential as a probiotic.

Interestingly, after conducting in vivo and in vitro toxicity studies, a spore‐forming strain of W. coagulans known as GBI‐30, 6086 was recognised as Generally Regarded As Safe (GRAS) by the US Food and Drug Administration (FDA) and the European Union Food Safety Authority (EFSA) in 2013 (Konuray & Erginkaya, 2018). This relevant outcome has sparked increased interest in spore‐forming probiotics because of their ability to withstand adverse environmental conditions (Ahmad et al., 2022). Moreover, reducing manufacturing costs and eliminating the need for freeze‐drying processes are interesting features in food industries. Most importantly, the addition of W. coagulans spores did not compromise the sensory or nutritional characteristics of food. Over time, W. coagulans spores have been integrated into a wide range of functional foods, from pasta, chocolate and ice cream to heat‐resistant cereal‐based products, thanks to their ability to withstand high temperatures during food processing (Figure 1; Table 2) (Amini et al., 2022; Cao et al., 2020; Fares et al., 2015; Janipour et al., 2023; Kobus‐Cisowska et al., 2019; Konuray & Erginkaya, 2018).

FIGURE 1.

FIGURE 1

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Benefits of incorporating W. coagulans in functional foods.

TABLE 2.

W. coagulans – based functional foods.

W. Coagulans strain Functional foods References
GBI‐30, 6086 Pasta Fares et al. (2015)
GBI‐30, 6086 Dark chocolate Kobus‐Cisowska et al. (2019)
n.r. Ice cream Janipour et al. (2023)
n.r. Gluten‐Free Cake Mix Amini et al. (2022)
MTCC 5856 Fermented tea Kombucha Yang et al. (2022)
GBI‐30, 6086 Orange juice Almada‐Érix et al. (2021)
GBI‐30, 6086 Yogurt Almada‐Érix et al. (2021)
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Furthermore, a recent study revealed the effectiveness of a novel organic functional beverage containing apple cider vinegar. This beverage with the supplementation of W. coagulans spores not only improved the serum lipid profile but also implemented the protection against HFD‐induced hepatic steatosis in mice (Urtasun et al., 2020).

Interestingly, W. coagulans has been found to co‐exist in complex hand‐made ecological communities, such as Kombucha. Indeed, a study by Yang et al. (2022) analysed the microbial composition of nine different kombucha beverages that were prepared using various tea varieties and different probiotic additions. The results of the study showed that W. coagulans was the most dominant strain under all the tested conditions (Yang et al., 2022). Furthermore, in 2021, a dynamic system designed to replicate the conditions of the gastrointestinal tract provided evidence that spore of W. coagulans GBI‐30, 6086 strain exhibited remarkable survival when incorporated into orange juice and yogurt (Almada‐Érix et al., 2021). Additionally, this commercially available strain has served as a reference control in studying the probiotic properties of newly isolated W. coagulans strains from corn, pickled cucumber, potato and tomato (Konuray Altun & Erginkaya, 2021).

Bioactive probiotic molecules

Multiple studies have provided evidence that W. coagulans possesses the capability to inhibit bacterial growth by balancing the microbiota through mechanisms including the production of bioactive effector molecules, such as LA and acetic acid, which serve the indirect role of antimicrobial agents by reducing the pH of the surrounding environment (Ibrahim et al., 2021). In this context, bacteriocins are small antimicrobial peptides or proteins that bacteria produce to inhibit the growth of closely related strains or other competing microorganisms. Bacteriocins are categorised into two main classes: (1) class I, undergoing post‐translational modifications, and (2) class II, existing in a cyclic not modified form (Ríos Colombo et al., 2018). Moreover, several strains of W. coagulans (ATCC 7050, BDU3, MTCC 5856) produce lactosporins, a unique class of bacteriocins that does not belong to the conventional classification and exhibiting broad antimicrobial activity as well as shield ability on skin cells against UV‐induced apoptosis (Abdhul et al., 2015; Riazi et al., 2009; Majeed et al., 2020).

Coagulins belong to class II of bacteriocins, and their mechanism of action involves the integrity disruption of the target bacterial cell membrane (Soltani et al., 2021). W. coagulans strain I4 and L1208 were found to produce a plasmid‐linked antimicrobial peptide called coagulin active against foodborne bacteria (Cao et al., 2020; Contursi et al., 2014; Notomista et al., 2015; Zhang et al., 2024) (Figure 2).

FIGURE 2.

FIGURE 2

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W. coagulans probiotic bioactive and effector molecules promoting human/animal health.

A genetic arrangement that encodes for a non‐lanthionine‐antimicrobial peptide called circularin A was found in MA‐13 and XZL9 strains by comparative genomic studies (Aulitto et al., 2019, 2022; Gabrielsen et al., 2014).

Some other strains may have lost certain genetic determinants during evolution, as observed in XZL4 where the circC gene involved in circularin A maturation was absent (Aulitto et al., 2022). Unlike circularin A and coagulin, which naturally occur as circular peptides, the presence of a gene encoding for a polypeptide antibiotic called bacitracin has been reported for MA‐13 (Aulitto et al., 2022). Interestingly, among the nine strains examined, only MA‐13 bears a complete pathway for bacitracin production including a signal transduction system (bceRS), along with adjacent efflux transporters (bceA/B) and an additional bcrC gene (Aulitto et al., 2022). Bacitracin acts by inhibiting bacterial cell wall synthesis and it is a widely used broad‐spectrum veterinary antibiotic, commonly employed as a feed additive and as a preservative in meat, aquatic, vegetable and dairy industries (Cai et al., 2020; Choyam et al., 2021).

MICROBIAL CELL FACTORY FOR ENZYME PRODUCTION

Numerous studies have addressed the utilization of W. coagulans for the cost‐effective production of valuable products suitable for diverse biotechnological applications such as hydrolytic enzymes that have been successfully applied in numerous food processing industries, including brewing, baking and dairy food (Aulitto et al., 2018; Konuray & Erginkaya, 2018; Pamueangmun et al., 2023).

Although many of the hydrolytic enzymes herein described are also produced by other microorganisms, the main advantage of exploiting those derived by a probiotic (such as W. coagulans), resides in their safe use in food manufacturing. Additionally, their potential extends to the pharmaceutical and biotechnological sectors, where they can be employed to enhance the production of desired compounds and facilitate biochemical transformations (Pamueangmun et al., 2023). A full description of W. coagulans enzymes is supplied to deepen its potential as a microbial cell factory for industrially relevant enzymes (Aulitto et al., 2021). Moreover, the robustness of the enzymes deriving from the thermophilic nature of W. coagulans as well as their capability to work over wide temperature and pH ranges, represent an added value in several industrial applications (Ou et al., 2009).

Glycosyl hydrolases involved in food processing

The main enzymes involved in starch metabolism are amylases catalysing the hydrolysis of starch and related carbohydrates such as cyclodextrin glucanotransferase (CGTase) (Lim et al., 2021). These extracellular enzymes convert starch into cyclodextrins (CDs), i.e., 6–12 glucose units joined by α‐1,4‐glycosidic bonds, through a glycosyl transferase mechanism (Lim et al., 2021). The amylase capacity to break down starch into simpler sugars makes it valuable for processes like fermentation, dough rising, starch liquefaction and saccharification (Raveendran et al., 2018). The applications of W. coagulans‐produced amylases span various industries, including food and beverage, starch processing, animal feed and more (Marzo‐Gago et al., 2024). W. coagulans B49, releases a thermostable α‐amylase when wheat bran is added to the culture medium, while W. coagulans CUMC 512, isolated from soil, produces an alkaline thermostable α‐amylase and an extracellular CGTase, the latter one exhibiting a significant ability to convert about 60% of soluble starch into CDs in 20 h at 50°C, widely utilised in food, pharmaceutical and chemical industries (Lim et al., 2021).

It is known that probiotic LAB and Bifidobacteria, typically present in the small intestine, could serve as natural suppliers of digestive enzymes, including α‐ and β‐galactosidases, the former catalysing the hydrolysis of α‐1,6‐linked‐galactose residues from galactose containing oligosaccharides, whilst the latter hydrolyzes the lactose into glucose and galactose (Kailasapathy & Chin, 2000). Several W. coagulans strains have been proven to encode for α‐galactosidases and β‐galactosidases and some of them have been biochemically characterised (Batra et al., 2002; Liu et al., 2019; Zhao et al., 2018). For instance, NRR1207 produces an α‐galactosidase that enhances the nutritional value of soybean meal by breaking down α‐galactoside linkages and making essential nutrients more accessible for absorption (Ra et al., 2018). Furthermore, among β‐galactosidases, the BcGalB enzyme from the MA‐13 strain, besides lactose hydrolysis suitable for lactose‐free dairy products, performs also a transglycosylation reaction, generating galactooligosaccharides (Aulitto et al., 2021). These latter can be incorporated into functional foods to promote gut health, allowing individuals with lactose intolerance to enjoy dairy‐based functional foods without adverse effects (Park & Oh, 2010).

Industrial applications of W. coagulans enzymes

W. coagulans exhibits a multifaceted hydrolytic enzymatic repertoire, including phytases, lipases, xylanase and proteases. Phytases, such as those from W. coagulans IDCC 1201, enhance mineral bioavailability by breaking down phytic acid in plant‐based feed ingredients (Lee et al., 2016a). Biofertiliser formulations combining W. coagulans and Pseudomonas fluorescens have been demonstrated to supply environmental benefits by reducing reliance on synthetic fertilisers and mobilizing native phosphorus for plant uptake (Yaashikaa et al., 2020).

Various W. coagulans strains produce lipases with alkaline properties, suitable for broad applicability in detergent and pharmaceutical industries. For instance, W. coagulans BTS‐3 and ZJU318 produce lipases with optimal activity in a wide range of temperatures and pH values (Al‐Ghanayem, 2021; Kumar et al., 2005). Interestingly, W. coagulans MTCC‐6375 features a unique lipase with versatile activity, demonstrating changes in optimal pH under specific incubation conditions (Kanwar et al., 2006).

Furthermore, proteases derived from W. coagulans are utilized in various industries. For instance, an extracellular protease from a psychrotrophic strain is optimized to function at 37°C and a pH of 8 (Talebi et al., 2013). A thermostable protease from W. coagulans PSB‐07 exhibits remarkable stability in organic solvents (Olajuyigbe & Ehiosun, 2013). Finally, xylanases from W. coagulans BL69 and B30 are efficient in xylan degradation, making them valuable in textile, pulp and paper industries (Heck et al., 2005; Sharma et al., 2013). A full list of the enzymes isolated and characterised is provided in Table 3 and Figure 3 and their working operational parameters are reported.

TABLE 3.

List of the biochemically characterised enzymes from W. coagulans strains, operating between 30° and 70°C and pH 4–9. Thermostability values are reported.

W. coagulans strains Enzyme T opt (°C) pHopt Thermostability (half‐life) References
ATCC 7050 α‐Galactosidase 55 6 60°C for 30 min Zhao et al. (2018)
T242 β‐Galactosidase 50 6.8 n.r. Xu et al. (2021)
RCS3 β‐Galactosidase 65 6.8 65°C for 2 h Batra et al. (2002)
NL01 β‐Galactosidase 55–60 6 60°C for 3.5 h Liu et al. (2019)
MA – 13 β‐Galactosidase 60 5 60°C for 4 h Aulitto et al. (2021)
IDCC 1201 Phytase 50 4 60°C for 3 h Lee et al. (2016a)
n.r. Cyclodextrinase 70°C 6 n.r. Lim et al. (2021)
BTS – 3 Lipase 55°C 8.5 55°C for 2 h Kumar et al. (2005)
n.r. Lipase 30°C 8 n.r. Al‐Ghanayem (2021)
n.r. Lipase 37°C 7 n.r. Alkan et al. (2007)
MTCC‐6375 Lipase 45°C 8.5 55°C for 20 min Kanwar et al. (2006)
PSB – 07 Protease 60°C 8 50/60°C for 60 min Olajuyigbe and Ehiosun (2013)
B30 Xylanase 50°C 9 n.r. Sharma et al. (2013)
BL69 Xylanase 60°C 7 n.r. Heck et al. (2005)
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FIGURE 3.

FIGURE 3

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Production of enzymes from W. coagulans and their diverse applications in different fields.

LA PRODUCTION FROM AGRI‐FOODS

W. coagulans is recognised for its proficiency in LA production, a versatile compound with widespread applications in the food industry, pharmaceuticals, medical fields and personal care products (Abdel‐Rahman et al., 2013; Huang, Tian, et al., 2023; Huang, Wang, et al., 2023). Moreover, LA serves as a building‐block for the PLA (poly lactic acid), an eco‐friendly alternative to traditional plastics made up of L‐and D‐isomers (de França et al., 2022). W. coagulans stands out for its robust metabolic pathways, ensuring efficient conversion of 5‐or 6C sugars to L‐ and/or D‐LA, and contributing to improved cost‐effectiveness (Huang, Tian, et al., 2023; Huang, Wang, et al., 2023). Some studies have documented the effective utilization of W. coagulans for converting agri‐food wastes into LA, exploring the potential of various agricultural residues, including wheat, rice and rye straw, along with sugar cane bagasse and corn stover (Hu et al., 2015; Jiang et al., 2016; Ouyang et al., 2020). Herein, we report the successful production of LA from food residues and/or leftovers by employing diverse strains of W. coagulans in different fermentation configuration (Figure 4, Table 4).

FIGURE 4.

FIGURE 4

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The process of LA production from agri‐food residues through W. coagulans and its potential advantages.

TABLE 4.

Valorization of diverse agri‐food residues through W.coagulans fermentation.

W. coagulans strains Agri‐Food residues Pre‐treatment of raw materials Process organization Lactic acid (g/L) Yield Productivity (g/L·h) References
n.r. Coffee mucilage – Batch 43.3 0.7 gLA/gsugars 1.5 Neu et al. (2016)
A559 Pasta waste Enzymatic hydrolysis SSF 47.65 0.81 gLA/gsugars n.r. López‐gómez et al. (2022)
A534 Sugar bread Enzymatic hydrolysis SHF 80 0.85 g/g 2.67 Olszewska‐Widdrat et al. (2020)
n.r. Crust bread waste Enzymatic hydrolysis SHF 62.2 0.57 gLA/gcrust bread waste 2.59 Alexandri et al. (2020)
DSM1 Bread waste Enzymatic hydrolysis Fed‐batch 155.4 0.85 gLA/gglucose 1.3 Cox et al. (2022)
JI12 Empty fruit bunch Acid‐catalysed and enzymatic hydrolysis SSF 80.6 n.r 3.4 Ye et al. (2014)
LA1507 Sweet sorghum juice Acid‐catalysed hydrolysis Batch 102 0.943 gLA/gsugars 2.9 Olszewska‐Widdrat et al. (2019)
MA‐13 Citrus waste – SSF Batch and Fed‐batch 44.8 0.96 gLA/gglucose 1.9 Aulitto et al. (2024)
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Coffee waste

Coffee residues, such as coffee pulp or spent coffee grounds, have high organic content, including carbohydrates that can serve as a substrate for LA production. Generally, during coffee production, a step of spontaneous fermentation is usually included by indigenous microbiota (e.g., yeasts and LA bacteria). For instance, Pereira et al. showed that L. plantarum LPBR01 was able to produce LA from the coffee pulp with a productivity of 1.33 g/L of LA after 24 h (Pereira et al., 2016). In this context, W. coagulans isolated from rape seed meal was employed in fermentation experiments, using coffee mucilage as a renewable nitrogen source in LA fermentation (Neu et al., 2016). Interestingly, after a relatively short fermentation time (about 30 h), the resulting LA yield was 0.7 g/g of free sugars, using initial concentrations of 10.8 g/L sucrose, 21.5 g/L glucose and 27.8 g/L galactose/fructose/xylose, with LA production of 43.3 g/L (Table 2). When the fermentation was supplemented with 5 g/L of yeast extract, only the productivity was increased, but the LA yield was not affected (Neu et al., 2016). Considering that coffee waste amounts to 10 million tons per year, its valorization as a renewable alternative to commercial nitrogen sources, not only contributes to waste reduction but also enhances the economic feasibility and sustainability of the LA production process.

Pasta and bread waste

Pasta and bakery leftovers/by‐products are significant contributors to food waste in many countries. Indeed, they typically contain 50%–70% starch, available for LA through fermentation processes. Several fermentation studies are reported and summarised in Table 4. López‐gómez et al. (2022) have reported the ability of W. coagulans to produce LA starting from enzymatically treated pasta waste. Firstly, the enzymatic saccharification was successfully used to release 0.81 g sugars per gram of dry pasta waste. After preliminary comparative screenings of various W. coagulans strains, A559 was selected as the best performer in both laboratory and pilot scales. Indeed, during the lab‐scale experiments, two fermentation modes were tested simultaneous saccharification and fermentation (SSF), and sequential hydrolysis and fermentation (SHF). This latter was proven to be more efficient compared to SSF, with a yield of 0.81 g of LA per gram of sugars (Table 2), highlighting the effectiveness of SHF for efficient LA production from pasta waste and demonstrating its potential as a sustainable and productive approach for valorizing this waste stream (López‐gómez et al., 2022). Through batch experiments, it was observed that the strain W. coagulans A534 effectively metabolises different substrates, including acid whey, sugar beet molasses, sugar bread and tapioca starch (Olszewska‐Widdrat et al., 2020). In continuous fermentations, the highest productivity was achieved using sugar beet molasses, with a remarkable value of 10.34 g/L·h. Moreover, the LA to sugar conversion yield reached 0.86 g/g demonstrating that LA can be efficiently produced in a continuous mode, regardless of the substrate employed (Olszewska‐Widdrat et al., 2020; Table 2). Bread waste, which is rich in starch, has been employed for the production of LA through SSF using different strains of Lactobacillus paracasei. From this process, a very low yield (0.054 g/g) of LA from bread waste was achieved (Sadaf et al., 2021), compared to W. coagulans. Indeed, Alexandri et al. utilised crust bread waste, an alternative bakery residue, to produce LA using different strains of W. coagulans (Alexandri et al., 2020). Through enzymatic hydrolysis, they successfully achieved a production of 62.2 g/L of LA, with a conversion yield of 0.57 g/g of crust bread waste and a productivity rate of 2.59 g/L·h. Most recently, Cox et al. demonstrated an eco‐friendly solution to valorise bread waste into LA through acidic and enzymatic hydrolysis using W. coagulans DSM1 under non‐sterile conditions (Cox et al., 2022). The concentration of LA using waste bread and enzymatic hydrolysate reached remarkable levels (102.4 g/L and 129.4 g/L of LA), with yields of 0.75 and 0.83 g LA/g glucose (0.42 g/g feedstock), respectively. Employing enzymatic hydrolysate in a fed‐batch process further enhanced the LA titre, reaching an impressive yield of 155.4 g/L, with a conversion yield of 0.85 g/g and a productivity rate of 1.30 g/L·h (Table 2). Additionally, the solid residues generated during the hydrolysis and fermentation stages were effectively utilised for the generation of biogas, with methane being the primary product. This integrated approach not only achieved an outstanding LA titre under non‐sterile conditions but also represents a proof of concept of a waste bread‐based integrated biorefinery, with the simultaneous production of valuable biogas from fermented residues.

Empty fruit bunch

Southeast Asia is a significant resource hub for palm oil production, accounting for over 80% of the global output. An interesting and valuable approach for the valorization of agricultural waste is the use of empty fruit bunch (EFB). Recently, Triwahyuni et al. focused on exploring LA production from EFB treated with alkali explosion and using cellulolytic enzymes and Lactobacillus delbrueckii through the SHF approach at 55° and 60°C. However, W. coagulans JI12 was previously demonstrated to have a remarkable ability to produce L‐LA with exceptional optical purity of up to 99.5% due to a frameshift mutation observed in the D‐LA dehydrogenase gene conferring regioselectivity (and enantiospecificify) to the encoded enzyme (Ye et al., 2014). By comparing the processes of simulated sugar mixtures and EFB hydrolysate with similar sugar compositions, it was found that the productivity of EFB hydrolysate (6.2 g/L·h) was lower than the simulated mixture (7.1 g/L·h) (Table 2). This slight decrease in productivity has to be traced back to the presence of inhibitors, such as furfural, hydroxymethylfurfural, acetic acid and other compounds that are generated during the pre‐treatment process of the EFB (Ye et al., 2014). Therefore, the exploitation of W. coagulans strains able to withstand such toxic compounds represents a solution to exploit the full potential of EFB wastes. In this context, JI12 has shown remarkable capabilities in converting both glucose and xylose into LA because of its remarkable tolerance to high concentrations of acetate (20 g/L) and furfural (4 g/L). Notably, the strain can metabolise all furfural in acid‐pre‐treated EFB whole slurry, while simultaneously producing over 80 g/L of LA through detoxification, saccharification and fermentation.

Citrus waste

Citrus waste, such as peels, pulp and other residues, offers significant potential as a renewable feedstock for LA production. Orange peel waste has been employed as a raw material for LA production utilizing Lactobacilli sp., resulting in conversion yields that ranged from 80% to 90% (Bustamante et al., 2020). Among the microbial strains, W. coagulans MA‐13 is a resilient biocatalyst capable of efficiently fermenting lignocellulosic biomass into L‐LA, even in the presence of inhibitory compounds derived from biomass. Recently, it was set up a one‐step production of L‐LA using W. coagulans MA‐13 with untreated citrus waste as a sustainable feedstock (Aulitto et al., 2024). To enhance biomass degradation, a thermophilic enzymatic cocktail was employed in conjunction with the hydrolytic capabilities of MA‐13, resulting in an impressive (up to 62%) increase in biomass degradation. Furthermore, batch and fed‐batch fermentation experiments demonstrated the complete conversion of glucose into L‐LA, achieving concentrations of up to 44.8 g/L, with a productivity of 1.9 g/L·h and yield of 0.96 g/g (Table 2).

Sorghum waste juice

Sweet sorghum juice (SSJ) is extracted by crushing the harvested stalks of Sorghum bicolour, a highly versatile grain with various culinary possibilities. Compared to cellulosic hydrolysates, the utilization of SSJ was proven to be more convenient and cost‐effective for LA production, because of its high concentration of soluble sugars, including sucrose, glucose and fructose. In the study conducted by Wang et al., they investigated the use of acid hydrolysate from SSJ for fermentative production of L‐LA (Wang et al., 2017) by employing strain LA1507. Results showed a maximum productivity of 2.90 g/L·h and an impressive yield of 0.943 g/g (Table 2) (Olszewska‐Widdrat et al., 2019).

CONCLUSIONS AND PERSPECTIVES

Relevant features that make W. coagulans a valuable microorganism for a wide array of sustainable and eco‐friendly technologies related to food manufacturing and biomaterial production are: (i) resilience towards diverse physiochemical conditions also due to the spore‐forming capability, (ii) enzymatic versatility, (iii) thermotolerance, (iv) robustness towards inhibitory compounds, (v) probiotic/prebiotic properties and (vi) LA production. Indeed, thanks to its wide spectrum of enzymatic activities and production of prebiotics/antibacterials molecules, W. coagulans as a probiotic enhances the absorption of essential nutrients in the digestive tract and extends shelf life by inhibiting the growth of harmful bacteria. Interestingly, some studies and clinical trials have explored the use of W. coagulans as part of bacteriotherapy interventions for various gastrointestinal disorders, including irritable bowel syndrome.

Bacteriotherapy, utilizing W. coagulans, is a promising approach for preventing and treating dysbiosis in different body areas like the respiratory tract, skin and ears. Advances in additive manufacturing technologies enable the creation of bacterial cell‐loaded devices for localised probiotic deployment. Techniques like electrospinning, electrospray and extrusion‐based printing facilitate the production of structures that encapsulate diverse bacteria types, including E. coli and Lactobacilli. W. coagulans can be encapsulated within these structures and deployed using advanced manufacturing methods. Leveraging 4D bioprinting, researchers can optimise W. coagulans delivery, offering potential solutions for dysbiosis treatment in various body regions.

Furthermore, its metabolic versatility and excellent substrate utilization property during fermentation, enable the efficient conversion of diverse agri‐food and lignocellulose residues into LA. New perspectives could be achieved by engineering W. coagulans to implement the production of LA and the expression of functional enzymes.

Overall, these features point to W. coagulans as an essential player in the usage of plant‐based building materials towards the development of an eco‐friendly, sustainable and greener future by reducing wastage, landfills and toxic emissions.

AUTHOR CONTRIBUTIONS

Emanuela Maresca: Conceptualization; writing – original draft; writing – review and editing. Martina Aulitto: Conceptualization; writing – original draft; writing – review and editing. Patrizia Contursi: Funding acquisition; supervision; writing – original draft; writing – review and editing.

CONFLICT OF INTEREST STATEMENT

All the authors declare no competing interests.

ETHICAL APPROVAL

This article does not contain any studies with human participants or animals performed by any of the authors.

ACKNOWLEDGEMENTS

This work was supported by the Ministry of University and Research CUP PRIN 2022 CONTURSI E53D23003010006 and to CN_00000033 MUR decree n. 1034 of 17 June 2022 CUP F13C22000720007, project title “National Biodiversity Future Center‐NBFC”.

Maresca, E. , Aulitto, M. & Contursi, P. (2024) Harnessing the dual nature of Bacillus (Weizmannia) coagulans for sustainable production of biomaterials and development of functional food. Microbial Biotechnology, 17, e14449. Available from: 10.1111/1751-7915.14449

[Correction added on 30 May 2024, after first online publication: the article title has been updated in this version.]

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