REVIEW ARTICLE

Lactic acid bacteria: A bio-green preservative against mycotoxins for food safety and shelf-life extension

Sneh Punia Bangar¹^*, Nitya Sharma², Aastha Bhardwaj³, Yuthana Phimolsiripol⁴

¹Department of Food, Nutrition and Packaging Sciences, Clemson University, Clemson, SC, USA;

²Food Customization Research Lab, Centre for Rural Development and Technology, Indian Institute of Technology Delhi, New Delhi, India;

³Department of Food Technology, Jamia Hamdard, New Delhi, India;

⁴Faculty of Agro-Industry, Chiang Mai University, Chiang Mai, Thailand

Abstract

Mycotoxins produced from Aspergillus, Penicillium, and Fusarium cause food spoilages during handling and storage, owing to immense economic losses and serious human health concerns including immunosuppression and carcinogenic effects. Furthermore, these species are also known to produce mycotoxins. Aflatoxin B1 (AFB₁), zearalenone (ZEA), ochratoxin A (OTA), and deoxynivalenol (DON) are the most commonly occurring mycotoxins. The removal of mycotoxins from the contaminated food using lactic acid bacterias (LABs) has been proposed as a green, inexpensive, safe, and promising mycotoxin decontamination strategy. LABs can control the mycotoxin production following a series of steps, including, adsorption, metabolite interaction, and biodegradation. This article provides systematic review of LABs as bio-green preservative with anti-mycotoxin potential for sustainable food safety. This consolidated review may be of technical importance to understand detoxification mechanisms and potential interaction of compounds originated with mycotoxin degradation for target food before incorporation by the food industry.

Key words: anti-mycotoxin, food safety, lactic acid bacteria, metabolite interaction, shelf-life

^*Corresponding Author: Sneh Punia Bangar, Department of Food, Nutrition and Packaging Sciences, Clemson University, Clemson, NC, USA. Email: [email protected]

Received: 23 November 2021; Accepted: 14 March 2022; Published: 13 April 2022

DOI: 10.15586/qas.v14i2.1014

© 2022 Codon Publications

This is an Open Access article distributed under the terms of the Creative Commons Attribution-NonCommercial-ShareAlike 4.0 International (CC BY-NC-SA 4.0). License (http://creativecommons.org/licenses/by-nc-sa/4.0/)

Introduction

Fungal spoilage is a major challenge for food industries, leading to food sensory defects, food waste, economic losses, and public health concerns due to their toxins. Plant-pathogenic fungi are responsible for up to 20% loss of the global harvest yield, which is sufficient to take care of about 600 million individuals every year. In addition, fungal diseases of the five most cultivated food crops worldwide were assessed to annihilate about 125 million tons of produce annually (Almeida et al., 2019). Certain fungal species secrete toxic secondary metabolites (mycotoxins), such as aflatoxins, ochratoxins, fumonisins, etc., which cause a major food safety issue for humans and livestock. According to the Food and Agriculture Organization report, 25% of the world’s food crops are badly affected by mycotoxins during cultivation or storage (USDA, 2016). Thus, fungal mycotoxins in food are major concerns for producers, purchasers, researchers, and regulatory agencies. Various food products such as fruits, cereals, nuts, pulses, etc., have demonstrated the prevalence of mycotoxins as shown in Table 1.

Table 1. Mycotoxins found in foods.

Fungus	Positive samples (%)	Mycotoxins	Detected foods	References
Aspergillus flavus	14.06	Aflatoxins B1	Dry nuts	Macri et al. (2021)
Aspergillus parasiticus	67.14	Aflatoxins B1	Vegetable oil	Poormohammadi et al. (2021)
	41.1	Aflatoxins M1	Bovine milk	Pandey et al. (2021)
Aspergillus niger, Aspergillus flavus, and Fusarium sp.	70	Aflatoxins B1	Dry fruits	Awan et al. (2021)
Aspergillus niger, Aspergillus tubingensis, and Aspergillus flavus	5.26	Ochratoxin A	Palm dates	Nikolchina and Rodrigues (2021)
	0-100	Ochratoxin A	Salami	Tolosa et al. (2020)
	7.61	Ochratoxin A	Milk	Turkoglu and Keyvan (2019)
Monascus spp.	Raw = 69.0 Dietary supplements = 35.1 Processed products = 5.7	Citrinin	Fermented red rice	TwaruZek et al. (2021)
Aspergillus spp. Penicillium spp.	0-62 0-69	Citrinin Ochratoxin A	Supermarket food samples	Meerpoel et al. (2021)
	30.8 17.5 33.3 Not detected	Aflatoxin B₁,T-2 toxin, Ochratoxin A, Deoxynivalenol	Dried shrimp, dried fish, and dried mussel products	Deng et al. (2020)
Penicillum expansum Penicillum cyclopium	9.0	Patulin	Dried fruits	Przybylska et al. (2021)
Penicillum expansum Penicillum cyclopium	40	Patulin	Strawberry	A-Reda and Sahib (2021)
	21.8	Patulin	Mango	Hussain et al. (2020)
Fusarium poae, Fusarium equiseti,	8.2-12.3	Fumonisin B1	Cornmeal	Massarolo et al. (2021)
Fusarium acuminatum , Fusarium sporotrichioides , Fusarium graminearum , Fusarium cerealis, Fusarium culmorum	63.0	Fumonisin B1	Whole wheat	Iqbal et al. (2020)
	Wheat/wheat flour = 4 Maize = 20 Paddy rice = 55	Deoxynivalenol	Wheat, maize, paddy rice, wheat flour	Golge and Kabak (2020)

Owing to the grave concerns over mycotoxins, there is an urgent need to establish alternative, eco-safe, and cost- effective approaches to overcome food mycotoxin contam-ination. As of late, the utilization of bio-preservatives, microorganisms, or their antimicrobial components for food preservation, has received a flood of interest due to increasing demands from consumers to embrace more natural food preservation approaches instead of depending on manufactured synthetic compounds. Lactic acid bacteria (LABs) are ideal probiotic candidates for food as fungal antagonists (Nielsen et al., 2021). LABs are utilized in traditional food fermentations and are considered as Generally Regarded as Safe (GRAS) and Qualified Presumption of Safety (QPS) by the American Food and Drug Agency (FDA) and the European Food Safety Authority (EFSA), respectively (Mora-Villalobos et al., 2020). LABs are considered as “green preservatives” due to their potential to inhibit fungal growth in foods. Organic acids, diacetyl, bioactive anti-mycotic peptides, fatty acids, carboxylic acids, bacteriocins, hydrogen peroxide (H₂O₂), lactones, alcohols, and reuterin are the reported antifungal compounds produced by LABs (Sadiq et al., 2019). This review provides a concise overview of the anti-mycotoxin potential of LABs as green biopreservative, along with its application in various food products.

LABs

In order to address the two consumer health concerns caused by (1) fungal growth and mycotoxin release in foods and (2) use of chemical preservative in foods, there is a great demand to develop safe and effective antifungal methods to improve or replace the current chemical and physical treatments. Biological control is a strategy that uses microorganisms or their metabolites to inhibit the growth and proliferation of pathogens. Using LAB as a green preservative is one of the most effective alternative owing to their potential to release antifungal metabolites against various fungal species.

LABs is a term given to a group of gram-positive bacteria that are characterized as catalase-negative, non-motile, and non-spore forming. The main fermentation products obtained from species LAB homofermentatives is lactic acid, while LAB heterofermentatives produce lactic acid along with carbon dioxide and ethanol/acetate. Various studies have promoted the use of LAB as a natural preservative that can effectively replace chemical preservatives in foods, and can also provide health-promoting and probiotic properties (Nasrollahzadeh et al., 2022a). Owing to the GRAS and QPS status of LAB, further exploring their potential as a biopreservative is now greatly appealing researchers over any other microorganism (Nasrollahzadeh et al., 2022b). Also, the LABs are easy to culture and maintain, and since they are naturally present in the gut they are more effective against mycotoxins (Muhialdin et al., 2020).

The LABs comprise genera, for example, Lactobacillus, Lactococcus, Streptococcus, Leuconostoc, Pediococcus, Enterococcus, Oenococcus, and Weissella. Different LAB strains with antifungal activity can be obtained from multiple sources, including Lactobacillus kefiri M4 and Pediococcus acidilactici MRS-7 from kefir; Pediococcus acidilactici, Limosilactobacillus fermentum, and Lactiplantibacillus plantarum from traditional fermented milk; Lactobacillus sucicola, Weissella paramesenteroides, Pediococcus acidilactici from citrus; and L. plantarum, Lacticaseibacillus paracasei, and Lactiplantibacillus pentosus from fermented beverages. These strains have been proved to be a promising tool to enhance the shelf-life of cereals, fruits and vegetables, nuts and seeds, bakery products, etc. An elaborate and latest prior-art of several LABs well acknowledged for their antifungal potential is given in Table 2.

Table 2. Anti-mycotoxin potential of LABs.

Fungal species	LAB species for myco- toxin removal	Substrate	Culture conditions	Detection and quantification technique	Percent mycotoxin reduction	Proposed mechanism	References
Ochratoxin A
Aspergiiius niger, Aspergiiius carbonarius	Pediococcus pentosaceus	Grapes	Grape samples were diluted in sterile saline solution and then plated on MRS agar at 37°C for 48 h; ochratoxin removal potential assessed in both MRS and PBS	HPLC-FLD	84% in MRS and 25% in PBS	Biodegradation; Various metabolic compounds and metabolites of LAB contributed to the antifungal activity.	Taroub etal. (2019)
Aspergiiius fiavus, Aspergiiius parasiticus, Aspergiiius niduians Aspergiiius ochraceus	L. piantarum	Table cream	Inoculation at a 5.2 log CFU/mL in MRS broth and incubation at 37°C for 4 days anaerobically	HPLC-FLD	58%	Adsorption to bacterial cell wall components, especially to polysaccharides and peptidoglycans as well as teichoic and lipoteichoic acids	Hashemi and Gholamhosseinpour (2019)
Aspergiiius parasiticus	L. piantarum, L. brevis, Levilactobacillus spp.	Brazilian artisanal cheese	LAB strains inoculated in MRS broth washed, resuspended culture was adjusted to 0.5 MCFarland (-108 CFU/mL) and diluted in 0.9% NaCI to obtain a final cell density of 5.0 log₁₀ CFUxg^-1; mycotoxin removal potential assessed in PPB	Liquid chromatography with fluorescence detector	≈ 50-90%	Adsorption to bacterial cell walls through ion exchange, complexation, and hydrophobic iterations	Moller et al. (2021)
Aflatoxin
Aspergiiius fiavus, Aspergiiius parasiticus, and Aspergiiius nomius	L. iactis ssp. cremoris, L. rhamnosus, L. ssp. iactis	Skimmed milk	Bacterial cells re-activated in MRS broth and isolated in MRS agar; bacterial cell wall isolates	Ultraperformance liquid chromatography	81.4,56.8, and 50.8%, respectively.	Adsorption; Sequestering property of clays	Muaz et al. (2021)
Aspergiiius fiavus and Aspergiiius carbonarius	L. kefiri	Kombucha beverage	LAB was isolated on MRS agar and incubated at 30°C under anaerobic conditions for 5 days.	HPLC-FLD	97.22 and 95.27% of AFB, and AFB₂, respectively	Biodegradation and adsorption (mechanisms involving hydroxylation, epoxidation, reduction, and dehydrogenation)	Taheur et al. (2019)
Not known	L. rhamnosus, L. iactis	Frescal cheese	1.0 x 10¹⁰ cells/g of the starter culture as procured by the manufacturer	HPLC-FLD	≈ 100%	Adsorption: Physical binding of the toxin to bacterial cell wall components, mainly peptidoglycans and polysaccharides	Gongalves et at. (2020)
Aspergillus flavus,	L. fermentum,	Fruit processing	Cultures were maintained	HPLC-FLD	≈80%	Not known	Cruz ef al. (2021)
Aspergillus parasiticus, and	L. paracasei,	by-products	aerobically on MRS agar at 4°C
Aspergillus nomius	L. plantarum		and transferred to a new media monthly. Before use in assays, each isolate was cultivated anaerobically in MRS broth at 37°C for 20-24 h, harvested by centrifugation, washed twice, and resuspended in PBS.
Aspergillus, Penicillium, and	L.s rhamnosus,	Milk	Cryopreserved LABs were	HPLC-FLD	≈ 33-100%	Adsorption, degradation: All	Martinez ef al. (2019)
Fusarium spp.	Pediococcus acidilactici,		reactivated in MRS broth and			tested strains adsorbed 19 to
	Pediococcus pentosaceus		incubated at 37°C and 5% C0₂ for 24 h.			61% AFM1 in milk
Aspergillus and Fusarium	L. plantarum spp	Corn kernels and	Incubated on MRS Broth for 12	LC-MS/MS	39-63.1%	Antifungal activity of LAB	Nazareth ef at. (2021)
species		corn ears	h at 37 °C; anaerobic conditions for 72 h at 37 °C to allow for MRS fermentation.			metabolites
Not known	L. acidophilus, bulgaricus,	Salt fermented	Individually inoculated in 100 mL	LC-MS/MS	≈ 8-35%	Physical adsorption; high-salt	Ye et al. (2020)
	and L. casei	fish product	of MRS broth, cultured to the log			stress promoted synthesis of
			phase at 37°C			detoxification factor in LAB.
Aspergillus flavus and	Pediococcus pentosaceus,	Brewer's grains	Grown in MRS broth at 37°C in	HPLC-FLD	37.6-70.7%	Adsorption: bacterial	Asurmendi ef al. (2020)
Aspergillus parasiticus	L. plantarum,		microaerobiosis for 24 h MRS			peptidoglycans and
	L. mesenteroides,					polysaccharides act as
	L. mesenteroides, L. coryniformis ssp. coryniformis					mycotoxin binders
Aspergillus flavus	L. rhamnosus,		Incubated at 37°C for 24 hr in		27.1-56.8%	Adsorption: mycotoxin binding	Danial ef al. (2020)
	L. mesenteroides, L. lactis ssp., L. casei, Streptococcus thermophilus, Bifidobacterium longum, Bifidobacterium animalis ssp.	-	MRS broth	TLC		to bacterial cell walls
Patulin
Aspergillus, Penicillium, and	L. casei,		LAB strains were activated by		95% by	Absorption and irreversible	Zheng ef al. (2020)
Byssochlamys species	L. plantarum,		growing on MRS liquid media		Lactobacillus	biotransformation
	L. fermentum, L. paracasei and L. rhamnosus	Fruit juice	at 37°C for 24 h	HPLC	casei
Fungal species	LAB species for myco- toxin removal	Substrate	Culture conditions	Detection and quantification technique	Percent mycotoxin reduction	Proposed mechanism	References
Not known	L. rhamnosus	Apple juice	PBS	^—	72.73 and 70.51% for HCL-treated and heat- treated LAB	Adsorption: binding rates increased in the presence of acid (21.37%) and heat treatments (19.15%)	Li et al. (2020)
Penicillium expansum	L. plantarum, L. fermentum	Kefir grains	Water kefir grains (10% w/v) inoculated and kept at room temperature for 2 days; ground, suspended in sterile saline; inoculated on MRS agar for 3 days at 37°C	HPLC	≈ 93%	Adsorption: smoother the bacterial cell wall exterior, and the bigger the cell wall volume and the surface area is, the higher the adsorption ability.	Bahati et al. (2021)
Penicillium expansum	L. plantarum and L. acidophilus	Syn biotic apple juice	Strain was inoculated into 10 mL MRS broth (pH 6.2) and incubated at 37°C for 48 h.	HPLC-UV	52.36 and 59.7%, respectively	Adsorption: noncovalent interaction between patulin and carbohydrates and surface layer proteins of the bacterial cell walls	Zoghi et al. (2021)
ZEA
Aspergillus parasiticus	L. plantarum, L. brevis, and Levilactobacillus spp.	Brazilian artisanal cheeses	Inoculation into MRS broth and incubation at 30°C for 24 h. Grown culture was centrifuged, and the cell pellet was resuspended in 10 mL 0.9% NaCI and centrifuged. The washed, resuspended culture was adjusted to 0.5 MCFarland (~ 10⁸ CFU/mL) and diluted in 0.9% NaCI in order to obtain a final cell density of 5.0 log₁₀ CFU g^-1.	HPLC-FLD	≈ 50-90%	Adsorption: exposure of binding sites and production of exopolysachharides by LAB.	Moller et al. (2021)
Fusarium spp.	L. acidophilusand L. delbmeckii subsp. bulgaricus	Animal liquid feed	Strains were grown overnight on MRS at 37°C under microaerophilic conditions and shaken with 125 strokes/min	UHPLC-FLD/DAD	57%	Adsorption and biodegradation	Ragoubi etal. (2021)
Fusarium spp.	L. buchneri, L. lactis, L. plantarum, L. lactis	Corn silage	LAB strains were grown at 37°C in MRS broth; cultures were then centrifuged for 5 min at 4600 x g at room temperature; supernatant was discarded. The pellet was washed three times with a saline solution (0.89 g/100 mL NaCI), diluted in the same solution.	HPLC-MS/MS	40-60%	Adsorption: LAB inoculation interacted with the fungal population to change the mycotoxin profile relative to untreated silage.	Gallo ef at. (2021)
Fusarium genus such as F. cerealis, F. graminearum	L. paracasei	Dairy products	Bacteria were cultured in MRS broth sterile medium 24 h at 37°C; culture at 3.83 McFarland (1.149 ’10⁹CFU/mL) was transferred to a sterile conical flask with ZEA in DMSO was added; Incubation at 37°C.	HPLC-ESI-MS/ MS	84.93%	Biotransformation and biosorption: physical binding of ZEA to surface components of the cell, transport of ZEA inside the cell and its accumulation, metabolization of ZEA to a less toxic form	Zloch etal. (2020)
Fumonisins
F. verticillioides	Enterococcus casseiiflavus Enterococcus faecium	Maize grains	For LAB isolation, 25 g of maize grain were placed in MRS broth and incubated at 37°C for 24 h.	HPLC	88.75%		Diaz et al. (2021)
Fusarium species	Not known	Ogi (fermented maize-based food)	Maize grains were soaked in water and allowed to ferment (steeping) for 2-4 days (48-96 h). The softened grains were then washed, wet-milled, and sieved using a muslin cloth. The sieved paste was diluted with water in a container and left to ferment (souring) for 1-2 days (24-48 h).	LC-MS/MS	47.45- 84.88%	Lactic acid fermentation reduces the levels of toxins.	Ademolaetal. (2021)
F. verticillioides and F. proliferatum	L. paracasei	Artisanal butter of Tunisia	The inoculated cultures were incubated for 3 h at 37°C in MRS in a chamber with 95% air: 5% co₂	-	Not known	Degradation; mitigation of FB, toxicities by reduction of its bioavailability in the gastrointestinal tract	Ezdini et al. (2020)
Not known	L. plantarum, L. deibrueckii subsp. Delbrueckii, Pediococcus pentosaceus	Maize based fermented cereals	LAB strains were cultivated and stored on MRS agar slants at 4°C for 3 months and for long-term conservation, cryopreserved at -80°C in 12.5% glycerol	HPLC-FLD	≈80%	Adsorption; deactivation of the mycotoxins by LAB was due to binding rather than metabolism	Dawlaletal. (2019)

ESI, Electrospray ionization; FB1, fumonisin B1; FLD, Fluorescence detector; HPLC, FHigh performance liquid chromatography; LC, Liquid chromatography; MS, Mass spectroscopy; MRS, de Man, Rogosa, and Sharpe agar; PBS, Phosphate-buffered saline buffer solution; TLC, Thin layer chromatography; UV, Ultraviolet detection; ZEA, Zearalenone.

Mycotoxin Detoxification Using LAB

Mycotoxin detoxification in foods by LABs can be achieved either through viable cells and their metabolites, or by particular enzymes obtained by certain LAB strains. It has been hypothesized that the development of fungal mycotoxins is encouraged under unfavorable environmental conditions and can be arrested by observing competition for available space and nutrients by the viable cells of LABs (Sadiq et al., 2019). These viable cells of LABs are capable of releasing acids and antifungal bioactive metabolites, such as lactic acid, benzoic and propionic acid, formic acid, butyric acid, hexanoic and caproic acid, phenyllactic acid, hydrogen peroxide, monohydroxy octadecenoic acid, carbon dioxide, cyclic dipeptides, phenolic compounds, bacteriocins, fungicins, reuterine, ethanol, diacetyl, and hydroxyl fatty acids (Ruggirello et al., 2019), all of which are associated to arresting the fungal activity. On the other hand, certain strains of LABs produce proteolytic enzymes that hydrolyze cell wall proteinases into polypeptides, peptide transporters (that carry peptides in the cell), and intracellular peptidases (that degrade peptides into amino acids) (Muhialdin et al., 2020). However, the mechanisms of reduction in mycotoxins have certain uncertainties; for example, the same phenomena of decrease in toxin concentration is ambivalent, as the conventional analytical methods cannot determine whether the mycotoxins have been adrift or have been masked by being temporarily bound to other elements in food (du Plessis et al., 2020). Therefore, in order to understand the reduction in fungal growth and mycotoxin levels, this section reviews the possible mechanisms reported for various mycotoxins. As the literature suggests, the possible reduction in mycotoxins is mainly due to a series of steps, including, adsorption, metabolite interaction, and biodegradation (Figure 1).

Figure 1. Possible binding mechanisms of mycotoxins to LAB cell wall components.

Binding/adsorption, interaction, and degradation of

(1) Ochratoxin A (OTA): Some authors have reported the adsorption of mycotoxins onto LAB cell walls as a probable mechanism for their anti-mycotoxin potential. OTA degradation was observed by binding OTA to LAB strains cell wall components, owing to the surface hydrophobicity, electron donor–acceptor association, and Lewis acid–base interaction. This binding capacity can be further increased through mutagenesis/genetic manipulation or supplementation with binding promoting compounds (Sadiq et al., 2019). The most common LAB species known to adsorb OTA are L. plantarum (Hashemi and Gholamhosseinpour, 2019) and Lactobacillus brevis. However, the effect of L. plantarum against OTA produced from Aspergillus parasiticus depended on the medium pH, as the maximum OTA reduction was observed at pH 3.0 compared to pH 6.5 (Møller et al., 2021). Similar results were observed by Taheur et al. (2021), where LAB species Lactobacillus kefiri diminished the OTA content produced from Aspergillus flavus and Aspergillus carbonarius in agar medium by 75% in bacteria supernatant (CFS), which was significantly affected and reduced to 17% when the pH was neutralized to 7. The authors inferred that the residual OTA amount in culture media was directly influenced by the pH, fungal strain, and bacterial species. Taheur et al. (2021) also examined the in vitro OTA degradation and absorption by LAB. They demonstrated that the decrease in the mycotoxins was mainly due to the inhibition of fungal growth, followed by adsorption. Du et al. (2021) also suggested the involvement of microbial catabolism and adsorption as potential mechanisms for the anti-mycotoxigenic activity of LABs, in Tibetian kefir grains, with the dominant LAB species of Lactobacillus kefiranofaciens.

(2) Aflatoxins: Likewise, aflatoxins have also been observed to have binding potential to LAB cell walls through a reversible noncovalent interaction, independent of the cell activity (Liu et al., 2020). Peptidoglycans, carbohydrates, teichoic acids or proteins, form an inherent part of the LAB cell wall that interact with functional groups and bind to the toxin through physical adsorption, ion exchange, and complexation (Asurmendi et al., 2020; Chlebicz and Śliżewska, 2020). Many authors have studied and validated the involvement of adsorption and degradation of aflatoxins under the influence of LAB strains as decontaminating agents (Kademi et al., 2019). Martínez et al. (2019) reported adsorption of AFM1 by Lacticaseibacillus rhamnosus in Artemia salina.

Similar adsorption and degradation phenomena to reduce aflatoxin AFB₁ were observed by Taheur et al. (2020) in black tea. The authors revealed that the binding was attributed to the adsorption of the toxin. At the same time, the degradation was carried out by processes like hydroxylation, epoxidation, reduction, and dehydrogenation based on the degrading agent. Asurmendi et al. (2020) stated that aflatoxin detoxification is more of a bonding process and less of a metabolic degradation process (Asurmendi et al., 2020). The binding, however, is dependent on existing environmental conditions (Mosallaie et al., 2019). As evidence to this statement, a recent study found that the production ability of aflatoxins (AFB₁, AFB₂, AFG₁, and AFG₂) by A. parasiticus was significantly affected by the applied conditions like pH (YES agar, YES broth, and MRS agar), time of treatment, heat-killed LAB strains (four strains of L. plantarum, two strains of L. brevis, four strains of Lactobacillus spp.), and the matrix used (milk or buffer) (Møller et al., 2021). Danial et al. (2020) also stated that AFG₁ adsorption distinctions by different LAB strains are mainly due to varying cell divider components (comprising a peptidoglycan framework and a proteinaceous layer; polysaccharides) and cell envelope structures.

Another effect of environmental conditions was depicted by Ye et al. (2020), where they emphasized the presence of high salt stress in the environment that promoted the synthesis of detoxification factors in LAB strains Lactobacillus acidophilus, Lactobacillus bulgaricus, and Lacticaseibacillus casei by increasing their metabolism against aflatoxin B₁ and causing physical absorption. Interestingly, heat-killed and acid-killed cells have been shown to have the highest binding capacity with aflatoxins. For example, heat-killed and acid-killed LAB cells from L. rhamnosus, Lactococcus lactis ssp. lactis, and L. lactis ssp. cremoris in contaminated skim milk have shown exceptionally high binding ability with AFM₁ (Muaz et al., 2021). This was attributed to the denaturation of membrane proteins, peptidoglycans, and degradation of polysaccharides components of the cell wall, thereby changing their hydrophobicity and respective binding capacities. Muaz et al. (2021) also found that the addition of an additive, sorbitan monostearate (SM), further increased the binding capacity of heat-killed LAB strains as the hydrophobic end of SM gets attached to the hydrophobic sites of LAB cells, such as peptidoglycan and teichoic acids, thus leaving the other hydrophilic end of SM to bind to hydroxy groups of AFM₁. Similar trends have been observed for detoxifying yogurt that evidenced enhanced binding of AFM1 to viable LAB strains by adding inulin as an additive (Sevim et al., 2019). Inulin supplementation promoted the growth and viability of mixed LAB inoculations (B. bifidum-Bifidobacterium animalis, L. plantarum-B. bifidum, L. plantarum-Bifidobacterium animalis) at extended storage periods.

(3) Fuminosins: Among various fumonisins, fumonisin B₁ (FB₁), and fumonisin B₂ (FB₂) are the major feed contaminants that adversely affect livestock and human health. The interaction and adsorption of FB₁ and FB₂ depend on the cell wall components and functional groups, specifically peptidoglycan and similar compounds (Sadiq et al., 2019). The decrease in fumonisin is mainly due to its quick binding ability to the peptidoglycan layer, which is considered to be the most credible binding site (Chlebicz and Śliżewska, 2020). Apart from this, reduction in pH with lactic acid production also leads to the transformation of fumonisin, leading to less toxicity (Ademola et al., 2021). Diaz et al. (2021) characteristically found that FB₁ produced from a phytopathogenic fungus responsible for maize gain contamination in the silo storage structure, Fusarium verticillioides, could be inhibited with a heterogeneous mixture of volatile organic compounds (diacetyl, acetoin, acetic acid, etc.) produced from LAB strain E. casseliflavus as a result of its metabolic activity. The authors also revealed that acetoin has potential in mycotoxin biosynthesis.

Dawlal et al. (2019) visualized and quantified the interaction between fumonisins and LAB strains and found that LAB metabolism was not required for interaction and binding with fumonisins without biodegradation, as both viable and nonviable LAB cells showed binding capacity, nonviable cells having the higher binding ratio. This was reasoned as the heat treatment promoted denaturation or disintegration of LABs, which opened up the available sites for higher fumonisin binding. Similarly, in viable cells, electrostatic potential favored the binding interaction between fumonisins and LABs. Apart from varying cell structure and components of LAB strains, the variance in fumonisin molecules’ structural conformation and charge also contributed to the binding and interaction. FB₁ and FB₂ carry different surface electrostatic potentials, chemical structure (FB₁ has an additional hydroxyl group in C10), and physical structure, making them preferential binding. The authors reviewed that LAB cells and fumonisin binding interaction were mainly mediated by long-range (steric and electrostatic interactions) and short-range (Van der Waals, Lewis acid–base, hydrogen bonding, and biospecific interactions) forces. However, the study failed to visualize this discrepancy as both fumonisins had the same fluorescing.

(4) Patulin: Like other mycotoxins, patulin reduction using LABs is also based on its adsorption in the cell wall and degradation by intracellular or extracellular enzymes (Zheng et al., 2020). Patulin adsorption is mainly observed as binding with the LAB cell wall protein, including thiol, esters, and alkaline amino acids. The main functional groups involved are C–O, OH, C– – O, COO–, C–N, and/or N–H (Wei et al., 2020). Ngea et al. (2021) critiqued the ability of LAB cells to reduce patulin in apple juice to be affected by critical environmental factors, including cell density, cell viability, patulin initial concentration, pH, and incubation time. The extent of patulin reduction is closely related to LAB cell surface area’s physical and chemical properties, cell wall volume, nitrogen–carbon (N/C) ratio, hydrophobicity, and functional groups. Large surface area, adsorptive selectivity, and large functional groups make nonviable cells more efficient in patulin reduction than viable cells (Bahati et al., 2021; Sajid et al., 2019). Exposure of LAB cells to conditions such as high temperature, acidic environments, etc., brings about structural changes to the cell walls that reduce the glycan layer crosslinking and increase cell wall permeability. Interestingly, Li et al. (2020) revealed that hydrochloric acid-treated LAB strains had significantly higher patulin detoxification ability and stability than heat-treated (121°C) LAB cells, owing to the disruption of hydrophobic interaction. This was speculated due to changes in cell wall structures that attain different degrees of cross-linking, thereby obstructing the toxin release. Additionally, additives like fructooligosaccharides, ascorbic acid, and citric acid to the apple juice demonstrated enhanced patulin binding by L. plantarum by reducing pH that stimulated S-layer proteins synthesis in the LAB cell wall (Zoghi et al., 2019).

(5) Zearalenone (ZEA): Last but not least, LAB-assisted ZEA removal involves either interaction with LAB cell wall components like peptidoglycans and surface proteins or interaction with intracellular proteins followed by absorption into the LAB cell wall (Sadiq et al., 2019). According to Złoch et al. (2020), ZEA neutralization by Lactobacillus paracasei cells is a nonlinear two-step process involving biosorption/binding techniques of ZEA by L. paracasei cells, as well as metabolization and biotransformation of ZEA to less toxic α-ZOL, β-ZOL forms. ZEA removal depends on cell wall protein type and structure, thus making it a strain-specific process. Out of the 17 strains of plant-derived LAB, L. plantarum isolated from wild spider flower pickle possessed the highest ZEA removal capability (Adunphatcharaphon et al., 2021). As per the results obtained by the authors, LAB cell wall polysaccharides did not affect ZEA removal stating non-involvement of hydrogen bonds in the interaction between LAB strain and ZEA. As far as lipids were concerned, lipase-treated LAB cells showed a significant reduction in ZEA as lipase hydrolyzed the ester bond lipid, causing a change in lipid structure. In addition, the presence of hydrophobic interactions was confirmed with a dominance of C–OH, C–C, and C–O–C functional groups of polysaccharides and single form bending functional groups of bonds in CH₂ and CH₃ present in teichoic acids, peptidoglycan, lipopolysaccharides, and phospholipids (Adunphatcharaphon et al., 2021). An adsorption–desorption study by Ragoubi et al. (2021) exposed that the viable cells of LAB caused ZEA biodegradation in PBS medium, with ZEA being the only carbon source. However, no related metabolites like, α and β-zearalenol, zearalenone, and its reduced metabolites were detected in inoculated PBS at the end of the incubation period, stating the absence of biodegradation. On the contrary, Adunphatcharaphon et al. (2021) showed that heat-treated nonviable Lactobacillus plantarum cells had a higher capacity to reduce ZEA. Still, since no ZEA degradation products were detected, the authors suggested the properties of heat-inactivated LAB cells for ZEA reduction and not biotransformation.

Mycotoxin Reduction in Foods Using LABs

LABs have been used for bio-preservation as an innovative approach to foods, including dairy products, bakery products, juices, meat, fruits and vegetables, and feeds (Table 3), for thousands of years due to their inhibitory properties. The contamination can occur at various stages during the manufacturing process. In cereal grains, mycotoxins can be formed by several spoilage-indicating molds, including, above all, Aspergillus spp. and Penicillium spp. Even in dry grains, mycotoxins can also be formed if either moisture migration due to temperature changes creates condensation points with a higher water content (hot spot theory) or moisture is formed through the respiratory activity of grains weevils or other grain pests (mites, larvae of flour moths) and then secondary mold growth occurs, and often not noticed. The stability of the silage (animal feed) cannot be estimated in advance, and therefore can be contaminated in similar proportions as grains (Liu et al., 2018).

Table 3. Practical food applications of LABs to the extended shelf life of food products.

Lactic acid bacteria	Strain	Possible compounds	Food/Feed product	Targeted fungi/mycotoxins	Extended shelf-life/lnhibition	References
		derived from metabolic activity			percentage
Limosilactobacillus reuteri		p-Coumaric acid	Bread	Fusarium culmorum	Compared to control < 4 days	Schmidt et al. (2018)
		Azelaic acid Reuterin/Acrolein	Whole wheat	Aspergillus niger	extra <7 days	Sadeghi et al. (2019)
		Reutericyclin Hydrogen peroxide	sourdough	Aspergillus niger	extra <7 days	Sadeghi et al. (2019)
		Reutericyclin Hydrogen peroxide	sourdough
Limosilactobacillus reuteri	R29R2Δ	(mentioned above)	Quinoa and rice bread	Mold growth	2-4 days	Axel et al. (2016)
andLevilactobacillus brevis		4-Hyd roxyphenyllactic acid Hyd rogen peroxide
Lactobacillus hammesii	R29	-	Bread	Mold growth	2-6 days	Black et al. (2013)
	-	-	Flaxseed sourdough	Aspergillus niger and	Compared to control < 2 days	Quattrini et al. (2018)
			bread	Penicillium roqueforti	extra
Lactiplantibacillus plantamm	UFG 121	Benzoic acid	Bread	Fusarium culmorum	No growth of the targeted	Russo et al. (2017)
		p-Coumaric acid			fungus up to 7 days
	-	Cyclic dipeptides 3-Phenyllactic acid		Aspergillus parasiticus	<3-4 days extra	Saladino et al. (2016)
	CCFM259	2-Flydroxy4-methypentanoic acid	Chinese steamed bread	Penicillium roqueforti	Up to 7 days no growth	Yan et al. (2017)
	CRL 778	Methylhydantoin	Quinoa flour-based	Mold growth	Significant increased compared	Dallagnol et al. (2015)
		Mevalonolactone	bread		to control
	C10	d-Dodecalactone Hyd rogen peroxide	Muskmelon fruit	Trichothecium roseum	Inhibited pink rot	Lv et al. (2018)
	-	d-Dodecalactone Hyd rogen peroxide	Grape berries	Aspergillus carbonarius	-	Lappa et al. (2018)
	E3 E4			B. cinerea and A. ochraceus	reduced the growth	Dopazo et al. (2021)
			Cottage cheese	Penicillium commune	> 25 days	Cheong et al. (2014)
	5BG		Spanish-style table olives	Fungal growth	> 5 months	Lavermicocca et al. (2018)
	TK9		Citrus and apple juice	Old-growth	<3-4 days extra	Zhang et al. (2016)
	UFG 121		Oat-based beverage	Fusarium culmorum	> 21 days	Russo et al. (2017)
	LR/14		Grains	Mucor racemosus, Rhizopus, stolonifer, Penicillium chrysogenum, and Aspergillus niger	2.5 years	Gupta and Srivastava (2014)
	YML007		Animal feed	Mold growth	30 days	Rather et al. (2014)
Lactiplantibacillus plantarum	–	–	Fermented milk: Doogh (Iranian traditional)	AFM1	99%	Sokoutifar et al.(2018)
Lactiplantibacillus plantarum	–	–	Skim milk	AFM1		Panwar et al.(2018)
Lactiplantibacillus plantarum Lactobacillus acidophilus	–	–	Apple juice	PT	90%	Zoghi et al.(2019)
Lactiplantibacillus plantarum in combination with Lacticaseibacillus rhamnosusand/or Lactobacillus harbinensis	L244CIRM-BIA1113 L172	3-Phenyllactic acid Hydrogen peroxide	Sour cream	Rhodotorula mucilaginosa, Penicillium commune, and Mucor racemosus	Inhibition of targeted fungi from 2 to 24 h	Salas et al.(2018)
Lactiplantibacillus plantarum in combination with other lactobacilli	–	Hydrogen peroxide Vanillic acid	Caciotta cheese	Penicilliumchrysogenum ATCC 9179 and Aspergillus flavusATCC 46283	≥ 30 days	Cosentino et al.(2018)
Limosilactobacillus fermentum	C14		Bread	Mucor sp.	Up to 25 days	Barman et al.(2017)
	GA715		Banana	Fungal growth	Increased three times	Wayah and Philip (2018)
	YML014		Tomato puree	Aspergillus niger, Aspergillus flavus, and Penicillium expansum	Increased by nine days at 25°C	Adedokun et al.(2016)
Lactobacillus citreumand Levilactobacillus brevis	L123 Lu35		Pain au lait or plain cakes	Penicillium corylophilum	Significant increase	Le Lay et al.(2016)
Lactobacillus amylovorus	DSM19280	Cinnamic acid derivatives Salicylic acid	Gluten-free quinoa sourdough bread	Mold growth	compared to control < 2 days extra	Axel et al.(2016) Ryan et al.(2011)
Lacticaseibacillus rhamnosus	GG	3-Phenyllactic acid	Whole pears	Fungal growth	Up to 9 days	Zudaire et al.(2018)
Lacticaseibacillus rhamnosus	–	–	Apple juice	Mycotoxin PT	80%	Hatab et al.(2012)
Lactobacillus delbrueckii spp. Bulgaricus Lacticaseibacillus rhamnosus, and Bifidobacterium lactis – in combination			Milk	AFM1	12%	Corassin et al.(2013)
Lacticaseibacillus rhamnosus alone or in combination with Bifidobacterium animalis subsp. lactis	A238 A026		Cottage cheese	Penicillium chrysogenum	Up to 21 days	Fernandez et al.(2017)
Lacticaseibacillus casei	AST18	2-hydroxy-4-methypentanoic acid	Yogurt	Penicillium sp.	compared to control < 4 days extra	Li et al.(2013)
Lactobacillus harbinensis	K.V9.3.1Np		Yogurt	Mold growth	Up to 6 weeks	Delavenne et al.(2013)
Lactobacillus parafarraginis	–		Silage	Yeast	Stability up to 144 h	Liu et al.(2018) Broberg et al.(2007)
Lactobacillus helveticus	KLDS 1.8701		Soybean milk	Penicillium sp.	Up to 21 days	Bian et al.(2016)
Lactococcuspiscium			Food matrixes	Brochothrix thermosfacta		Fall et al.(2012); Saraoui et al.(2016); Leroi et al.(2015)
	CNCM1-4031		Shrimps	L. monocytogenes		Saraoui et al.(2016)
	All strains	–	Salmon and cod juices	Photobacterium phosphoreum, B. thermosphacta, S. baltica, and L. monocytogenes	Strongly inhibited	Wiernasz et al.(2017)
	All strains	–	Salmon and cod juices	Shewanella baltica, Serratia proteamaculans, H. Alvei, and L. sakei	Broadly inhibited
Leuconostoc gelidum	All strains		Salmon juice	Shewanella baltica	Strongly inhibited
			Fish juices	L. monocytogenes	Highly inhibited
			Fish juice	P. phosphoreum	Slightly inhibited
			Cooked peeled shrimps	P. phosphoreum, B. thermosphacta, S. proteamaculans	Slightly inhibited	Leroi et al.(2015)
genus Lactobacillus			Ready-to-eat (RTE) seafood	pathogenic bacteria commonly reported in RTE	High antimicrobial activity	Sahnouni et al.(2016)
L. sakei	cocktail (3 strains/genomic diversity)		Ground beef	E. coli/Salmonella		Chaillou et al.(2014)
Lactococcus garviae		H2O2	Milk Cheese	S. aureus	Higher inhibition	Delbes Paus et al.(2010); Delpech et al.(2015); Delpech et al.(2017)
Lacticaseibacillus paracasei subsp. paracasei	KU517839	Bacteriocin	Cheese Cheese made from pasteurized milk	S. aureus		Heredia-Castro et al.(2015);Yoon et al.(2016); Madi and Boushaba (2017)
Carnobacterium divergens	V41	Bacteriocin: divercin	Smoked salmon	L. monocytogenes		Richard et al.(2003)

Aflatoxin M1: AFM1; Aflatoxin B1: AFB1; Ochratoxin A: OTA; Patulin.Conclusion and Future Perspectives

For the food and feed industry, the production of compounds derived from LAB metabolic activity is of high importance; their antimicrobial spectrum has inhibitory potential against spoilage organisms such as fungi (especially by mycotoxigenic fungi), yeasts, Gram-negative and Gram-positive bacteria, protozoa, retarding microbial growth y extending considerably shelf-life as shown in Table 3 (Strack et al., 2020). LABs inhibit microbial decay by generating antagonist metabolic products or establishing antimicrobial compounds. Organic acids, mainly lactic acid followed by acetic acid, are the main metabolites of LAB, but depend on the LAB strains, their mechanism of action, and carbohydrates as substrate, other kinds of antimicrobial substances, namely low molecular weight metabolites (reuterin, reutericyclin, diacetyl, fatty acids), hydrogen peroxide, antifungal compounds (propionate, phenyl-lactate, hydroxyphenyl-lactate, and 3-hydroxy fatty acids) can be produced (compounds metabolized by different LAB strains listed in Table 3) (Wang et al., 2021). They have been used for food bio-preservation as an innovative approach for dairy products, bakery products, juices, meat, fruits, and vegetables. Various mechanisms that impart the anti-mycotoxin effect in foods by LAB fermentation are presented in Figure 2. Table 3 shows a consolidated overview of the selected recent applications of LABs in various food products.

Figure 2. Anti-mycotoxin effect of LAB fermentation in foods.

Conclusion and Future Perspectives

Bio-preservation, including the use of LABs and their active metabolites, is a natural tool to prevent fungal growth, prolong shelf-life, and increase the safety of foods. LABs are GRAS and possess a large potential for bio-preservation due to their production of antimicrobial compounds. LABs effectively reduce mycotoxin production by fungi via adsorption of mycotoxin with LABs cell surface components, degradation of fungal mycotoxins, and inhibition of mycotoxin production. However, the antifungal and anti-mycotoxin potential of LABs depends on pH, initial viable count, growth medium and condition, and incubation temperature and time. Due to antifungal and anti-mycotoxin agents, LABs could be an ideal bio-preservative candidate for sustainable food systems, including dairy products, fruits and vegetables, cereal grains, bakery goods, nuts and seeds, and meat and meat products.

Mycotoxin detoxification capacity of LABs through cell wall bindings is an effective way for the removal of mycotoxins from food and feed. However, possible in vivo release of bound toxins during the detoxification process can be a matter of human health concern. Thus regulating the environmental conditions that can lead to this release is an important aspect. Further studies on the effect of several factors, like pH, growth medium, initial bacterial count, incubation time and temperature, growth condition (single or mixed), bacterial state (viable or nonviable), on the detoxifcation mechanism can help better understand the anti-mycotoxin activity of LAB. Additionally, understanding the LAB’s detoxification capacity and potential interaction of compounds obtained with mycotoxin degradation for particular food products may also serve as an important source of data, that can be further industrialized for the food sector.

Conflict of Interest

There are no conflicts of interest to declare.

Funding

This research was partially supported by Chiang Mai University, Thailand, under the Cluster of Agro Bio-Circular-Green Industry (Agro-BCG).

Ethical Approval

Ethics approval was not required for this research.

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