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Lu, Y.;  Xing, S.;  He, L.;  Li, C.;  Wang, X.;  Zeng, X.;  Dai, Y. Characterization, High-Density Fermentation, and  Production of  Lactobacilli. Encyclopedia. Available online: https://encyclopedia.pub/entry/29960 (accessed on 12 July 2025).
Lu Y,  Xing S,  He L,  Li C,  Wang X,  Zeng X, et al. Characterization, High-Density Fermentation, and  Production of  Lactobacilli. Encyclopedia. Available at: https://encyclopedia.pub/entry/29960. Accessed July 12, 2025.
Lu, Yun, Shuqi Xing, Laping He, Cuiqin Li, Xiao Wang, Xuefeng Zeng, Yifeng Dai. "Characterization, High-Density Fermentation, and  Production of  Lactobacilli" Encyclopedia, https://encyclopedia.pub/entry/29960 (accessed July 12, 2025).
Lu, Y.,  Xing, S.,  He, L.,  Li, C.,  Wang, X.,  Zeng, X., & Dai, Y. (2022, October 18). Characterization, High-Density Fermentation, and  Production of  Lactobacilli. In Encyclopedia. https://encyclopedia.pub/entry/29960
Lu, Yun, et al. "Characterization, High-Density Fermentation, and  Production of  Lactobacilli." Encyclopedia. Web. 18 October, 2022.
Characterization, High-Density Fermentation, and  Production of  Lactobacilli
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This entry is adapted from the peer-reviewed paper 10.3390/foods11193063

Lactobacilli have been widely concerned for decades. Bacteria of the genus Lactobacillus have been commonly employed in fermented food to improve the appearance, smell, and taste of food or prolong its shelf-life. They comprise 261 species (by March 2020) that are highly diverse at the phenotypic, ecological, and genotypic levels. Some Lactobacilli strains have been documented to be essential probiotics, which are defined as a group of living microorganisms that are beneficial to the health of the host when ingested in sufficiency. The viability and stability of Lactobacilli in the food industry and gastrointestinal environment are critical challenges at the industrial scale. The new production equipment and technology of DVS starter of Lactobacilli strains will have the potential for large-scale application, for example, developing low-temperature spray drying, freezing granulation drying, and spray freeze-drying.

Lactobacilli strains probiotics characterization performance improvement production of DVS starter

1. Introduction

Lactobacilli are Gram-positive rod, non-spore-forming, catalase-negative bacteria that commonly colonize the human intestine and have essential physiological functions in the human body [1][2]. Microscopically, these bacteria represent non-motile, thin rods that differ from long to short. Sometimes, they are present in coryneform, bent morphology, or chains. Lactobacilli comprise 261 species (by March 2020) that are highly diverse at the phenotypic, ecological, and genotypic levels [3]. Moreover, Lactobacilli have been documented to be important probiotics, defined as a group of living microorganisms that are beneficial to the host’s health when ingested in sufficiency [4].
Lactobacilli may be added as starters, which are used to ferment and produce specific changes in the chemical composition and sensory properties of foods [5]. It can produce amylase, protease, dehydrogenase, decarboxylase, β-glucosidase, and peptidase during the fermentation, thereby can be widely used in the food industry for the production of yogurt [6], cheese [7], sourdough [8], sausages [9], cucumber pickles [10], olives [11], sauerkraut [12], and so on. However, there is some diversity in the Lactobacilli used in fermented food, depending on the food matrix. One example is Lactobacillus plantarum, which is used as a starter culture in meat and wine (malolactic) fermentation [5], whereas L. bulgaricus can be found as the primary starter culture in yogurt fermentation [13].

2. Screening out Lactobacilli Strains

Lactobacilli used in the fermented food industry are diverse and many. Due to their excellent fermentation performance, they are extensively used to ferment food based on various raw materials, including milk, meat, cereals, fruits, vegetables, and seafood. Their commercial products, including probiotics, have ample market space [14]. So, screening out one or several new strains with excellent fermentation performance and potential probiotic properties is very meaningful work. High-throughput screening technology is a method for the quick selection of certain strains of Lactobacilli species with outstanding performance (such as extracellular polysaccharides, bacteriocin, gamma amino acid, butyric acid, short-chain fatty acid, etc.) [15]. These Lactobacilli strains can be traditionally isolated from a wide range of sources, such as human and animal mucosal membranes, plants or material of plant origin, and fermented food.
Researchers constructed a phylogenetic tree containing 64 Lactobacilli species based on the related Lactobacilli 16S rRNA gene sequence derived from National Center for Biotechnology Information (https://www.ncbi.nlm.nih.gov/). It shows that the Lactobacilli species used in the food industry are not clustered in a particular branch but are almost evenly distributed in all branches of the Lactobacilli phylogenetic tree. Based on this, the other species in this genus may also have potential use value in the field of food, livestock breeding, and medicine. Given that Lactobacilli have strain specificity, the traditionally safe Lactobacilli strains cannot ensure the safety of all strains in the species. Thus, the identification and detection of Lactobacilli are of significance.
Generally, microorganisms always contain desirable and undesirable characteristics. A qualified candidate for use in the food industry initially requires a desirable characterization, especially safety. For Lactobacilli strains, researchers expect to select strains with probiotic properties. These characteristics of new Lactobacilli strains need to be evaluated by in vitro and in vivo experiments to understand their potential probiotic properties and industrial uses.

3. Identification and Safety Assessment of Lactobacilli Strains

The safety status of Lactobacilli used in food production has become the focus of the attention of manufacturers and government departments. Although European Qualified Presumption of Safety regulations have identified microorganisms that can be safely used in food [16], some safety aspects must be evaluated before the newly screened cultures are applied to fermented food. For a newly screened Lactobacilli strain, its inherent characteristics need to be tested in vitro. First, researchers need to identify the isolate to make sure it is what researchers want and safe, and it can preliminarily predict whether the strains of this genus can be fully put into production [17]. 
The identification methods of Lactobacilli use phenotypic methods and molecular identification methods. In contrast to phenotypic approaches, molecular identification and characterization tools can distinguish even between closely related groups of species, which are indistinguishable based on phenotype, which is far more consistent, quick, trustworthy, and reproducible [18][19]. The most commonly employed molecular techniques for the identification of Lactobacilli can be divided into two groups: species-specific identification techniques (including amplified ribosomal DNA restriction analysis (ARDRA) and 16S and 23S rRNA sequencing) and strain-specific identification techniques (including ribotyping, restriction enzyme analysis (REA) with pulsed-field gel electrophoresis (PFGE), genetic probes/DNA dot blot, multiplex PCR using specific primers, randomly amplified polymorphic DNA (RAPD), amplified fragment length polymorphism (AFLP), PCR-denaturing gradient gel electrophoresis (DGGE), and fluorescent in situ hybridization (FISH)) [20][21][22][23][24][25][26][27][28][29][30][31][32][33][34]. Taxonomy and phylogeny of the genus Lactobacillus have been recognized as rather complicated, because of a great number of species with a diverse group of species [3]. It is clear that for reliable species determination within this genus, a polyphasic approach based primarily on one or more molecular methods is required. Additionally, the International Committee on Systematic Bacteriology has acknowledged polyphasic taxonomy as a trustworthy method for describing species and revising the current nomenclature of specific bacterial groupings. In a recent study, Zheng et al. [3] proposed reclassification of the genus Lactobacillus into 25 genera including the emended genus Lactobacillus, which includes host-adapted organisms that have been referred to as the Lactobacillus delbrueckii group, Paralactobacillus and 23 novel genera including Holzapfelia, Amylolactobacillus, Bombilactobacillus, Companilactobacillus, Lapidilactobacillus, Agrilactobacillus, Schleiferilactobacillus, Loigolactobacilus, Lacticaseibacillus, Latilactobacillus, Dellaglioa, Liquorilactobacillus, Ligilactobacillus, Lactiplantibacillus, Furfurilactobacillus, Paucilactobacillus, Limosilactobacillus, Fructilactobacillus, Acetilactobacillus, Apilactobacillus, Levilactobacillus, Secundilactobacillus, and Lentilactobacillus. This reclassification reflects the phylogenetic position of the micro-organisms and groups Lactobacilli into robust clades with shared ecological and metabolic properties that can anticipate the addition of new species shortly. Then, the relevant antibiotic susceptibility is usually determined and evaluated according to the protocol provided by the European Food Safety Agency (EFSA) [35]. Microdilution broth tests on test tubes, disk diffusion [36], and commercial ready-to-use kits [37] have been used to determine the physical sensitivity of known antibiotics to newly screened strains. Hemolytic activity was also investigated [38]. The production of various enzymes should also be evaluated. Maybe they are the cause of pathogenicity. Strains should be tested for known human toxins (e.g., cytolysin) by appropriate in vitro analysis. Detecting the toxicity of pathogenic genes and metabolites is also conducted; several Lactobacilli can decarboxylate and reduce amino acids in food to produce biogenic amines, which can cause poisoning symptoms if they accumulate in excess amounts in the body [39]. These in vitro experimental analyses are simple and rapid in determining the safety of a newly screened strain and avoid the use of harmful strains. For example, a hemolytic and toxin-producing strain can easily be excluded from further analysis [38]. The false negative strains created by in vitro experimental research are concerning. Therefore, further in vivo experiments are needed, including animal models and clinical applications [11][40].

4. Potential Probiotic Functionalities of Lactobacilli Strains

Some Lactobacilli have been reported as strains with high probiotic potential and support efforts to improve probiotic quality, such as L. salivarius strains BCRC14759 and BCRC 12574, with the highest exopolysaccharide production [41], L. johnsonii ZLJ010, with better adaptation to the gut environment and its probiotic functionalities [42], and L. helveticus D75 and D76 that can inhibit the growth of pathogens and pathobionts [43]. However, Lactobacilli strains in the probiotic market are still limited, and Lactobacilli strains with potential probiotic properties should be explored. An important aspect is to evaluate the selected Lactobacilli in vitro and find their probiotic potential. Some in vitro probiotic performance evaluations of the strains include survival under stress (low pH, high bile salt, high osmotic pressure, high oxygen, oxidation, starvation, etc.), adhesion ability, and antibacterial, antioxidation, cholesterol-lowering, and anticancer activities.
As probiotics, Lactobacilli colonizing the intestine to reach 1 × 106 CFU is necessary for its probiotic effect [44]. Lactobacilli can survive in the robust acid environment in the gastric juice and high bile salt concentration in the small intestine, which are two criteria for screening good probiotic Lactobacilli strains. The acid and bile salt tolerance of Lactobacilli strains use the rate of viable bacteria incubated in various acid pH environments as an indicator in in vitro assays. Additionally, many studies conducted artificially simulated gastric juice tolerance and animal model tests of probiotic Lactobacilli [14][45][46]. The survival rate was used as an index to evaluate probiotic Lactobacilli’s acid and bile salt tolerance.
Adhesion is another of the essential characteristics of probiotic bacteria that contributes to the colonization of probiotics in the gastrointestinal tract [47]. The ability of the bacteria to stick with hydrocarbons determines the extent of adhesion to the epithelial cells in the gastrointestinal tract, known as cell surface hydrophobicity [48]. The direct method of cell surface hydrophobicity of bacteria is to determine the change of absorbance of the supernatant of bacterial cell solution at 600 nm after treatment with hydrocarbons such as n-hexadecane and toluene. More precisely, the adhesion of Lactobacilli strains to mucin has also been determined [49]. Moreover, commercial kits for determining these mucins have been reported and can be used for high-throughput screening [50].
Intestinal epithelial cell (IEC) lines are often presumed to better represent conditions in the tissues of the GIT. Several studies have been conducted using human epithelial cell lines (such as HT-29, HT-29MTX, and Caco-2) to screen the adhesion of probiotic strains [47]. Additionally, other studies have focused on the self-aggregation of probiotics [51], which is also related to adhesion.
Lactobacilli strains can secrete lactic acid and other organic acids, lowering the environment’s pH and thereby inhibiting other microorganisms’ growth [52]. Additionally, Lactobacilli strains produce medicinal probiotic metabolites and bacteriocin BACs, often used as biological preservatives in the food industry, arousing people’s attention [53]. These metabolites have antagonistic activity against bacteria genetically similar to producing bacteria, which are immune to their own BACs. BACs have also been considered biologically active molecules with potential activities for human health, such as use as antiviral and anticancer drugs. BACs are extracellular antimicrobial peptides synthesized by ribosomes. They have extensive antibacterial activity and are a safe alternative to antibiotics. As a result, the shelf life of naturally fermented foods has increased. Therefore, screening high-yield BAC probiotic Lactobacilli strains from naturally fermented food should be an option. Researchers have also studied the production and characterization of BACs by different probiotics [12]. Additionally, the combined culture of different probiotics may produce new antibacterial products [54][55].
Some reports show that Lactobacilli strains have antioxidant activity and can be used as antioxidants in food, stabilizing food’s color, flavor, and taste [56]. Additionally, Lactobacilli strains can reduce the oxidative stress injury of Caco-2 cells and improve the antioxidant capacity under oxidative stress. Firstly, the tolerance of Lactobacilli strains to hydrogen peroxide was studied [57]. The antioxidant capacity of Lactobacilli strains was evaluated by measuring the hydroxyl radical scavenging capacity of cell-free extracts of these strains. These strains can produce metabolites such as superoxide dismutase, glutathione, and extracellular polysaccharide to inhibit oxidation.

5. Fermentation Performance of Lactobacilli Strains

As a lactic acid starter, Lactobacilli strains should be tolerant to harsh conditions, such as temperature changes, osmotic pressure (high fat and protein concentration in milk and meat and high salt in kimchi), and lactic acid accumulation. These characteristics can ensure the rapid adaptation and growth of microorganisms to bring good physical properties and taste to the products. Due to different food components, some of them are used for specific food manufacturing, such as yogurt and cheese (L. delbrueckii), fermented vegetables (L. plantarum and L. pentosus), and fermented meat (L. pentosus).
The diversity of lactic acid food produced by Lactobacilli strains requires that the fermentation characteristics of these strains are different [58]. For example, Lactobacilli strains used in meat processing should be able to improve the flavor of end products without producing biogenic amines, because these compounds are produced by the deacidification of free amino acids and have toxic effects on human intestines. Studies have revealed that Lactobacilli strains with protein hydrolytic activity [59], which belong to homogeneous fermentation, can significantly reduce the biogenic amines of fermented sausage. The production of bacteriocin by Lactobacilli strains, for example, is another feature of evaluating the development of meat products by Lactobacilli strains. It can inhibit the growth of pathogenic bacteria and increase the shelf life of products. As mentioned above, the antibacterial activity of Lactobacilli strains was screened to resist various pathogens evaluating the production of nisin against Listeria monocytogenes, Clostridium perfringens, Bacillus cereus, and Staphylococcus aureus [52][60][61][62].
Lactobacilli strains produce large amounts of lactic acid, a non-volatile, odorless compound that contributes to the aroma of the product [63]. Therefore, the production of another fermentation performance flavor molecule was evaluated by gas chromatography–mass spectrometry. The main aroma components include aldehydes, organic acids, higher alcohols, esters, carboxylic acids, and ketones [64][65]. Lactobacilli strains convert precursor molecules into aromatic compounds by secreting various extracellular enzymes [59][66][67]. In a protein-rich environment, proteolytic enzymes play a major role in forming aromatic molecules from the amino acids released by complex proteins. For example, milk is rich in casein, and Lactobacilli strains used in yogurt and cheese convert these precursor molecules into flavor substances. Lipid degradation also plays a vital role in the aroma formation of fermented meat and dairy products [68].

6. Health Functions of Lactobacilli Strains

From the functional characteristics, the existence of Lactobacilli strains in the intestinal microbiota is related to the host’s health status [69][70]. Clinical research showed that the reduction or increase in the proportion of this genus in the human body would produce health problems, such as irritable bowel syndrome [71], human immunodeficiency virus [72], obesity [73], type 2 diabetes [74], etc. Several strains with probiotic properties of specific metabolites have been successfully included in various functional foods. Although the mechanism of their role is uncertain, the health-promoting effect of these strains is related to the strains themselves and their active metabolic substances (such as extracellular polysaccharide, bacteriocin, polypeptide, short-chain fatty acid, etc.). Several meta-analyses and systematic research [75][76][77][78][79][80] support the health effects of probiotic Lactobacilli strains and specific metabolites produced by Lactobacilli strains in treatment cases, including acute rotavirus diarrhea in children, antibiotic-related diarrhea in children, Helicobacter pylori infection, allergic rhinitis, high blood pressure, hyperlipidemia, and other diseases. Additionally, based on its excellent physiological characteristics and probiotic function, Lactobacillus has become an important direction in the field of probiotic function. Given the specificity of the Lactobacilli strains, the new strains may have potential unique functional characteristics. 

7. Performance Development and Improvement of Lactobacilli Strains

As mentioned above, numerous species of Lactobacilli are used in food production, including improving traditional food and developing new products. On the one hand, Lactobacilli strains can enhance the quality of fermented food and, on the other hand, prolong the storage period of food as a preservative. Therefore, the excellent characteristics of Lactobacilli strains are the key to their application in the food industry. However, Lactobacilli strains have specificity themselves. Different strains of the same species show significant differences; therefore, new characteristics can be found [81]. Thus, scholars are committed to screening new Lactobacilli strains.
On the one hand, the growth of naturally screened Lactobacilli strains is limited by physical and chemical factors, such as pH [82][83], oxygen [84][85], osmotic stress [86], temperature [83], carbohydrate substrates[87], and other factors [88][89]. On the other hand, the yields of beneficial metabolites of naturally screened Lactobacilli strains, such as lactic acid [90], γ-aminobutyric acid [91][92], extracellular polysaccharide [92], and bacteriocin [93] are relatively low and cannot meet the requirements of industrial production. Therefore, reasonable breeding strategies are used to improve the performance of Lactobacilli strains with potential application in the food industry.
One method is mutagenesis breeding. Mutation breeding of Lactobacilli strains can change the genetic structure and function of Lactobacilli strains, and then screen mutants to obtain the required high-yield and high-quality strains [94]. It is the most basic modern breeding method. The breeding speed is fast, the cost is low, the time is short, and the method is simple, mainly including physical, chemical, and biological mutagens. Chemical mutagenesis primarily uses nitrosoguanidine, diethyl sulfate, and other chemicals. These chemicals are harmful to the human body. Thus, they are not widely used in the food industry. A limited number of studies focused on the biological mutagenesis of Lactobacilli strains, mainly involving transposon mutations [95][96]. Physical mutagenesis of Lactobacilli strains commonly uses ultraviolet [97] or microwave radiation [98]. Given the possible tolerance of traditional radiation technology of Lactobacilli strains, new mutation technologies, such as heavy ion beam irradiation and plasma mutation breeding, have recently appeared [99][100][101]. The operation of traditional mutation breeding is simple, and the experimental conditions are not high; the mutation is random, and the workload is enormous despite the introduction of high-throughput screening technology in the mutation process [99][102][103].
Another method is metabolic engineering, a continuation, and upgrade of gene engineering technology. This method can directionally change the functional characteristics of Lactobacilli strains and compensate for the shortcomings of classical mutagenesis screening [104][105][106][107][108]. The metabolic strategies of Lactobacilli mainly focus on the changes in pyruvate metabolism to produce essential fermentation end products, such as sweeteners, spices, aromatic compounds, and complex biosynthetic pathways, leading to the production of extracellular polysaccharides and vitamins [109]. Currently, the most commonly used methods for metabolic engineering of Lactobacilli include whole-genome amplification [110], genome shuffling [111][112], and genome editing (plasmid-based homologous recombination, Red/RecET-mediated double-stranded DNA recombination, and single-stranded DNA recombination) [113][114]. However, the safety of these methods for metabolic engineering to change the metabolic characteristics of Lactobacilli is worth considering and is not accepted by the European Union [115].

8. Role of Lactobacilli Strains in Food Production

The primary role of Lactobacilli strains in dairy processing (such as yogurt, dahi, kefir, koumiss, and cheese) is not only to improve the nutritional value but also to produce lactic acid, butyric acid, a variety of amino acids, and vitamins and other metabolites, resulting in a unique food flavor. Additionally, these strains use dairy products as a carrier to promote human health due to their probiotic effect [116]. The application of Lactobacilli strains to meat products can improve the appearance of meat products, promote the improvement of taste, inhibit the growth of spoilage bacteria, reduce the generation of nitrite and greatly improve the overall quality of meat products [117].
In turn, fermented foods as a carrier play a role in transporting and storing these excellent strains. On the one hand, these strains were found in traditional fermented foods, which characterized their excellent properties. On the other hand, these strains were intensively inoculated into conventional fermented food to improve product control. Fermented fruits can be produced by natural fermentation of the surface flora spontaneously formed (such as Lactobacilli and Pediococcus spp.) or inoculated with fermentation starter (such as L. plantarum, L. rhamnosus, and L. acidophilus). Food nutritionists are developing a new generation of fermented fruit products with special biological and unique sensory characteristics [5][118][119]. Fermented vegetable products can positively impact human health because they are rich in substances beneficial to human beings (such as dietary fiber, minerals, antioxidants, and vitamins). The principle is to use Lactobacilli strains attached to vegetables and several artificially selected excellent strains to carry out a series of microbial fermentations and finally obtain the finished pickle. The Lactobacilli contained in kimchi can promote human gastrointestinal peristalsis, reduce fat, and enhance immunity [10][65].
The fermentation of probiotic strains with excellent performance has attracted people’s attention. The screened new strains are often used in the development of new products. In recent years, several Lactobacilli strains have been widely used in various functional foods due to their unique physiological efficacy and flavor, such as active Lactobacilli drinks and solid drinks [64][120]. With the deepening of relevant research, Lactobacilli will be used in human health conditioning treatments as a probiotic functional food to a greater extent, and the application direction will be more extensive.
Lactobacilli can also be applied to preserving food, such as meat, fruit, vegetables, seafood, etc. These Lactobacilli strains are used as biological preservatives due to the following manifestations: (1) produce organic acids, such as lactic acid and acetic acid, to inhibit the growth and reproduction of most spoilage bacteria; (2) H2O2 production activates the catalase thiocyanate system in milk; (3) produce small proteins or peptides similar to bacteriocin, etc., [121][122].

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Yun Lu
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Shuqi Xing
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Laping He
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Cuiqin Li
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Xiao Wang
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Xuefeng Zeng
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Yifeng Dai
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