The inhibitory effects of plant additives on biogenic amine formation in fermented foods – a review

Figures & data

Figure 1. Main microbial BA formation pathways in fermented food and fate of ingested BAs in human gastrointestinal tract. Abbreviations: TDC – tyrosine decarboxylase, LDC – lysine decarboxylase, HDC – histidine decarboxylase, ODC – ornithine decarboxylase, ADC – arginine decarboxylase, ADI – arginine deiminase, ARG – arginase, AGM – agmatinase, AgDI – agmatine deiminase. Prepared based on Benkerroum 2016; Erdag, Merhan, and Yildiz 2018; Ladero et al. 2010. Created with BioRender.com.

Figure 2. Methods to reduce BAs in fermented products. Created with canva.com.

Table 1. Antimicrobial activity of plant compounds and their effects on BA accumulation - in vitro studies.

Plant material		Extraction solvent	Culture medium for BA accumulation testing	Method for the determination of antimicrobial activity	Target microorganisms	Main effects on BA content in the culture	Reference
Common name	Scientific name
Clove	Syzygium aromaticum L.	Direct addition of sterile clove powder	Mackerel (Scomber japonicus) extract supplemented with NaCl (1 or 2%)	Total plate count method	Enterobacter aerogenes ATCC 43175 (histamine-producing)	Tested concentration of clove amounted to 0.5%.After 24 h of culturing, in medium supplemented with 1% NaCl histamine concentration was almost two-fold lower because of clove addition, while in the medium containing 2% of NaCl, the production of this amine was entirely inhibited.Likewise, cadaverine production was totally suppressed in the bacteria cultured in 2% NaCl medium supplemented with clove. In the culture containing 1% NaCl, cadaverine was at the similar level regardless of clove addition.Culturing of E. aerogenes ATCC 43175 in the medium containing 2% NaCl and clove resulted in bacteria growth reduced by 8 or 7 log orders in comparison to culturing in the medium containing 2% of NaCl or 0.5% of clove alone, respectively.	Wendakoon and Sakaguchi 1993
GingerGarlicGreen onionRed pepperClovesCinnamon	Zingiber officinale Roscoe Allium sativum L. Allium fistulosum L. Capsicum annuum L. Syzygium aromaticum L. Cinnamomum cassia L.	Ethanol/water (95/5 v/v)	TSB (tryptic soy broth) supplemented with 0.5% L-histidine hydrochloride monohydrate, L-tyrosine, L-ornithine hydrochloride, L-lysine hydrochloride, and 0.0005% pyridoxal-HCl	Agar diffusion test	Four isolates of the Bacillus licheniformis species (i.e., No. 1043, 1056, 1542 – strong producers of histamine, putrescine and cadaverine, and No. 1553 – strong producer of histamine, tyramine, and putrescine) originated from Myeolchi-jeot (fermented anchovy)	Application of 1 mL of garlic extract (ca. 16%) resulted in the reduction of putrescine, cadaverine, histamine, and tyramine concentrations by 11.2%, 18.4%, 11.7%, and 30.9%, respectively, compared to control culture. Application of 0.5 mL of this extract (ca. 8%) inhibited spermidine formation the most efficiently − 17.4% reduction. For remained tested extracts obtained reduction of BAs in experimental culture medium was as follows:- green onion: 33.28% for tyramine (0.5 mL addition) and 10.87% for spermidine (1.0 mL)- red pepper: 10.73% for cadaverine, 32.49% for tyramine, and 27.34% for spermidine (0.5 mL)- ginger: 12.77% for putrescine (1.0 mL), 26.27% for tyramine (0.5 mL), and 27.17% for spermidine (0.1 mL)- cinnamon: 20.57% for putrescine (0.1 mL) and 23.66% for tyramine (0.5 mL)- cloves: 28.30% for tyramine (0.1 mL) and 19.82% for spermidine (1.0 mL). Moreover, the extract of garlic showed the highest antimicrobial activity against the amine producers tested.	Mah, Kim, and Hwang 2009
BlackberryJuniper	Rubus fruticosus L. Juniperus oxycedrus L.	Extraction of phenolics (PEs) and essential oils (EOs)PEs: 50% ethanol using the MAE (advanced microwave extraction system)EOs: hydrodistillation; pentane and diethyl ether (1:1 v/v) were used for trapping the volatile compounds	BHI (brain heart infusion broth)	MIC by broth microdilution method	Enterococcus faecium strain FC12 (tyramine-producing) originated from cheese	Plant additives were applied at sub-lethal concentrations (50% of MIC), namely 1 mg/mL except for R. fruticosus L. EO, that was added at concentration 0.75 mg/mL. Accumulation of tyramine followed the growth kinetics of tested bacteria. After 96 h of the culturing, tyramine content was in the range from ca. 120 mg/L (cultures with the EOs) to ca. 160 mg/L (control culture), so its maximum reduction reached approx. 25%. 2-phenylethylamine concentration ranged from ca. 15 mg/L (culture with R. fruticosus L. EO) to 85.3 mg/L (control), which represents approx. 82% reduction.The EOs showed a higher antimicrobial effect than PEs (especially R. fruticosus L. added at 0.75 mg/mL), what was confirmed in flow cytometry.	Montanari et al. 2022

Table 2. Antimicrobial activity of plant compounds against BA-producing bacteria - in vitro studies.

Plant material		Extraction solvent	Method for the determination of antimicrobial activity	Target microorganisms	Main antimicrobial effects in the culture	Reference
Common name	Scientific name
GarlicGingerCinnamon Star anise	Allium sativum L. Zingiber officinale Roscoe Cinnamomum cassia L. Illicium verum L.	Ethanol/water (95/5 v/v)	Agar diffusion test	Enterobacter cloacae isolate 17, Citrobacter freundii isolate 27, and Klebsiella pneumoniae isolate 54 – BA producers isolated from fish sauce	No tested additive exhibited antibacterial activity against E. cloacae.Garlic and star anise extracts significantly inhibited K. pneumoniae and C. freundii, respectively.	Zhou et al. 2016
Six species of brown algae	Ecklonia cava Eisenia bicyclis Sargassum sagamianumSargassum thunbergiiMyagropsis myagroidesSargassum horneri	Dried and milled algae were extracted with 94% ethanol or water at a ratio of 1:10 (w/v)	Disk diffusion method and minimum inhibitory concentration (MIC) test	Morganella morganii IFO 3848, ATCC 25830 – histamine producer	An Ecklonia cava ethanol extract (ECEE) exhibited the strongest antimicrobial activity (MIC = 2 mg/mL), as well as inhibitory effect against the activity of crude histidine decarboxylase of M. morganii (reduction by 33.79%; ECEE concentration: 1 mg/mL).The ethanolic extract of Eisenia bicyclis and Sargassum thunbergii also significantly reduced the activity of the tested enzyme (19.82 and 12.91%, respectively).	Kim et al. 2014
Schizandra chinensis Baillon (SCB) Prunus mume Siebold et Zuccarini (ZUCCA) Rosmarinus officinalis L. (RO) Thuja orientalis Rosa Polyphenol of green tea (PGT)Grapefruit seed extract (GSE) Glycyrrhiza uralensis (GU) Prunus mume Siebold & Zucc. (ZUCC)		Ethanol/water (70/30 v/v) extraction, vacuum-evaporation and freeze drying.PGT and GSE were commercial samples	Agar diffusion test (ADT) and minimum inhibitory concentration (MIC) tests	Enterococcus faecium – tyramine producer isolated from Cheonggukjang – a traditional Korean fermented soy food	SCB, ZUCCA, GU, ZUCC, GSE, and PGT exhibited strong antimicrobial activity against E. faecium, based on the ADT. According to the MIC test, the extracts of SCB, ZUCC, RO, PGT, and GSE were more potent antimicrobial substances (≤ 5 mg/mL) than ZUCCA, GU, Thuja orientalis, and Rosa(≤ 10 mg/mL).	Oh et al. 2012

Table 3. Effect of plant-origin additives on BAs in fermented meat products.

Fermented product	Applied plant additive and its dose	Form of additive	Main effects on BA content	Reference
Fermented mutton sausage	Spices extracts:star anise amomum tsao-ko clovecassiafennelbay leaf nutmeg	ethanolic extracts (95% ethanol) evaporated and dissolved in water	The greatest reduction of tyramine and histamine was obtained in the samples with clove extract addition − 18.7 and 9.6% in comparison to control, respectively. Clove extract was the most efficient also regarding 2-phenylethylamine and tryptamine – reduction by 21.8 and 24.6%, respectively. Spermidine and putrescine were reduced by 27.5 and 19.3%, respectively, because of cassia addition.	Jia et al. 2020
Dry fermented pork sausages	Salvia hispanica (Chia) seed (1 or 2%) Nigella sativa (Black Cumin) seed (1 or 2%)	grounded seeds, particle size <0.3 mm	Significantly increased tyramine and putrescine concentrations in comparison to control, as well as reduced spermine concentration.Application of 2% of chia seed significantly increased spermidine content.Agmatine content was reduced to values below limit of detection.	Borrajo et al. 2021
Smoked horse meat sausage	Thyme essential oil	purchased	Application of thyme essential oil significantly reduced putrescine (by 13% bigger diameter and by 22% smaller diameter), cadaverine (23% and 24%, respectively), histamine (24% and 33%, respectively) and tyramine content (17% and 18%, respectively), as well as total BAs (16% and 18%, respectively).Spermidine content increased by 16% (bigger diam) and 10% (smaller diam), while spermine increased by 3% either in the sausages of bigger or smaller diameter.	Huang et al. 2021
Smoked horsemeat sausage	Thyme essential oil (at concentration 0.156%) + Proteus bacillus starter cultureThyme microcapsules (at concentration 0.156%) + Proteus bacillus starter culture	purchased	Application of thyme microcapsules or thyme essential oil resulted in the significant reduction of histamine content by 66.15 or 49.22%, respectively, in comparison to control.Use of thyme microcapsules was more efficient and reduced histamine concentration by 33.35% compared to the samples with thyme essential oil.	Yu et al. 2021
Fermented pork sausages	Rose polyphenols at concentrations:1 mg/g meat2 mg/g meat3 mg/g meat	vacuum dried water extracts	Application of 3 mg/kg of rose polyphenols (RPs) resulted in a significant reduction of phenylethylamine, putrescine, and histamine in comparison to control (without rose polyphenols) and other variants after 24 d of fermentation.Addition of RPs at the levels of 2 mg/g or 3 mg/g significantly decreased tyramine concentration.Addition of RPs at the levels of 1, 2 or 3 mg/g significantly reduced cadaverine and spermidine contents.	Zhang et al. 2017
Sucuk – Turkish dry-fermented sausage	Urtica dioica extract:300 mg/kg600 mg/kg Hibiscus sabdariffa extract:300 mg/kg600 mg/kg	aqueous extracts	Application of 300 mg/kg of Urtica dioica extract reduced histamine, putrescine, and tyramine concentrations by 62.86, 91.12, and 14.42%, respectively, in comparison to control, after 15 days of fermentation.Application of 600 mg/kg of Urtica dioica extract reduced histamine, putrescine, and tyramine contents by 70.19, 90.41, and 19.49%, respectively.Addition of 300 mg/kg of Hibiscus sabdariffa extract reduced histamine, putrescine, and tyramine concentrations by 46.43, 82.59, and 27.35%, respectively.Addition of 600 mg/kg of Hibiscus sabdariffa extract reduced histamine, putrescine, and tyramine by 40.85, 76.91, and 21.50%, respectively.	Karabacak and Bozkurt 2008
Turkish dry-fermented sausage	Thymbra spicata oil 300 ppmSesame (Sesamum indicum L.) oil 300 ppm	essential oil obtained by steam distillation of air dried Thymbra spicata, dried over anhydrous sodium sulfate purchased roasted sesame oil	Application of Thymbra spicata oil and sesame oil decreased putrescine content by average 34.89 and 27.7%, respectively, in comparison to control, after 16 days of fermentation. Histamine concentration was reduced by 12.60 and 8.65%, respectively, while tyramine by 12.87 and 13.50%, respectively. Applied additives reduced analyzed BAs more efficiently than BHT – an antioxidant commonly used in the food industry.	Bozkurt 2007
Sucuk – Turkish dry-fermented sausage	Green tea extract 300 ppm Thymbra spicata oil 300 ppmGreen tea extract 150 ppm + Thymbra spicata oil 150 ppm	water green tea extract essential oil obtained by steam distillation of air dried Thymbra spicata, dried over anhydrous sodium sulfate	Application of green tea extract, Thymbra spicata oil or mix of them reduced average putrescine content during 15 d of fermentation by 34.42, 22.70, and 29.21%, respectively, in comparison to control.After 15 d of fermentation histamine concentration was the lowest in the samples with green tea extract. Application of Thymbra spicata oil or mixed T. spicata and GTE also resulted in the decrease of histamine content to ca. 200 ppm, while the concentration of this amine in the control sample was 275.3 mg/kg.Tyramine content was reduced by 24.25% because of the application of green tea extract.Examined additives reduced analyzed BAs contents more efficiently than BHT.	Bozkurt 2006
Harbin dry sausage	Seasonings extracts at concentration 0.3 g/kg:cinnamoncloveanise	ethanolic extracts (95% ethanol) concentrated with removal of ethanol, then freeze-dried and ground into a powder	Application of the extracts significantly reduced the content of six analyzed BAs in comparison to control, whereby the biggest difference compared to the other variants was obtained thanks to cinnamon extract use.The efficiency in BAs reduction of tested extracts was as follows: cinnamon > anise > clove. In comparison to control, after 9 d of fermentation, addition of cinnamon extract resulted in the reduction of putrescine by 18.93%, while cinnamon, anise, and clove extracts reduced histamine content by 21.89, 17.42, and 13.44%, respectively.	Sun et al. 2018
Fermented pork sausage	Tea polyphenol 0.3%Ginger essential oil 0.3%Anise essential oil 0.3%Cinnamon essential oil 0.3%	purchased	Anise EO was the most effective in the limitation of histamine, putrescine, tyramine and cadaverine formation.Reduction of histamine content by 31.9, 34.74, 28.22, and 19.02% because of application of ginger, anise, cinnamon EOs, and tea polyphenols, respectively.Reduction of tyramine content by 16.64, 23.76, 15.55, and 10.12% because of application of ginger, anise, cinnamon EOs, and tea polyphenols, respectively.Reduction of putrescine content by 39.31% due to addition of anise EO.Reduction of cadaverine content by 40.53, 42.68, 20.68, and 29.83% because of application of ginger, anise, cinnamon EOs, and tea polyphenols, respectively.Significant reduction of total content of 4 analyzed BAs by 28.58, 34.87, 18.58, and 21.31% in comparison to control because of application of ginger, anise, cinnamon EOs, and tea polyphenols, respectively.	Wang et al. 2021
Traditional Chinese smoked horsemeat sausage	plant extracts: tea polyphenols 0.19% and essential oils of cinnamon 0.025%, cloves 0.04%, ginger 0.024%, and anise 0.025%starter cultures (Lactobacillus sakei and Staphylococcus xylosus) + plant extracts (tea polyphenols 0.19% and essential oils of cinnamon 0.025%, cloves 0.04%, ginger 0.024%, and anise 0.025%)	EOs produced by supercritical CO2 extraction tea polyphenols – purchased	BA levels were reduced stronger by plant extracts than starter cultures applied alone. Synergistic effects were noticed when plant extracts and starter cultures were applied together.After 28 d of ripening:Tryptamine concentration was reduced by 23.76 and 74.92%; 2-phenylethylamine by 30.15 and 33.70%; putrescine by 71.97 and 84.97%; cadaverine by 75.37 and 81,00%; histamine by 36.48 and 100%; tyramine by 84.84 and 90.70%, when plant extracts and combination of plant extracts + starter cultures were applied, respectively. No significant differences occurred concerning spermine content. In comparison to control, spermidine concentration was higher by 10.31% in the samples with plant extracts addition and higher by 9.16% when plant extracts + starter cultures were applied.After 4 months of storage vacuum or non-vacuum packaged:Higher reduction of the content of tryptamine, 2-phenylethylamine, putrescine, and cadaverine after storage in vacuum package. Percentage reduction of aforementioned BAs was as follows: 56.96, 76.03, 86.04, and 70.51% in the variant with plant extracts alone, and 71.51, 77.07, 91.27, and 70.47% when both plant extracts and starter cultures were applied.Histamine reduction was the highest (93.52% in comparison to control) when both plant extracts and starter cultures were added, and the product was stored in vacuum package. In the variant where plant extracts were applied alone the reduction of histamine was higher in non-vacuum-packed product (50.99% compared to control).Tyramine reduction was the highest (90.28% in comparison to control) when both plant extracts and starter cultures were added, and the product was stored in non-vacuum package. In the variant where plant extracts were applied alone the reduction of tyramine was higher in vacuum-packed product (77.50% compared to control).Spermine contents were higher than in control, when plant extracts alone were applied both in vacuum and non-vacuum packaged sausages, 10.37 and 12.98% respectively. The highest reduction of spermine resulted from application of plant extracts in combination with starter cultures and vacuum-packaging (55.54% in comparison to control).Spermidine content was reduced the most (9.85% in comparison to control) when plant extracts were applied alone, and the product was stored in non-vacuum package.	Lu et al. 2015
Fermented sausage	Cava lees 5%Cava lees 5% + L. sakei CTC494Cava lees phenolic extract 0.3%Cava lees phenolic extract 0.3% + L. sakei CTC494Cava lees phenolic extract 0.3% + L. sakei BAP110	liquid DMSO phenolic extract prepared through water: ethanol (50:50) extraction of lyophilized, ground cava lees	After 21 days of ripening:Due to the addition of 5% Cava lees to spontaneously fermented sausages tyramine and cadaverine contents were reduced by 100% and 76.50%, respectively. Putrescine, spermidine and spermine contents were not altered significantly, while histamine was not detected.Samples contaminated with aminogenic Salmonella strains producing cadaverine, putrescine and histamine; after 21 days of ripening:In spontaneous fermentation, 5% addition of Cava lees resulted in the reduction of cadaverine and putrescine by 62 and 78%, respectively. Tyramine content was not altered significantly.Inoculation with non-aminogenic L. sakei CTC494 strain resulted in absence, or low BAs content, regardless of the addition of Cava lees.In spontaneous fermentation, the addition of 0.3% cava lees phenolic extract resulted in the reduction of cadaverine and putrescine concentrations by 21 and 40%, respectively. Tyramine content was not altered significantly.Inoculation with non-aminogenic L. sakei CTC494 strain inhibited the production of putrescine, cadaverine, and tyramine regardless of cava lees extract addition.Inoculation with non-aminogenic L. sakei BAP110 inhibited the formation of cadaverine and putrescine regardless of cava lees extract addition. Tyramine was not reduced completely either in the control (inoculated with L. sakei BAP110) or enriched with cava lees extract sample (L. sakei BAP110 + 0.3% cava lees extract). Its content was at similar level in both variants.	Hernández-Macias et al. 2021
Nham: fermented pork product	Ginger extract (20%) 34.4 U/g amine oxidase activity	minced ginger	Total biogenic amines content was reduced by 85.90% compared to control, after 7 d of fermentation. Putrescine, cadaverine, histamine, and tyramine were reduced by 95.31, 92.62, 79.32, and 79.76%, respectively.	Kongkiattikajorn 2015
Smoke – cured bacon	Grape seed extract 1 g/kg	purchased	In comparison to control, addition of grape seed extract resulted in the reduction of tyramine, cadaverine, putrescine, 2-phenylethylamine, and histamine concentration by 30.7, 37.1, 29.4, 34.5, and 73.6%, respectively.	Wang, Zhang, and Ren 2018
Aeros thasou – traditional fermented pork sausage (Greece)	Satureja thymbra EO (5 g/kg) + L. sakei	no data	In comparison to control, concentrations of tyramine, putrescine, and histamine were reduced by 62, 100, and 71%, respectively.	Latorre-Moratalla et al. 2010
Beef sausages	Punica granatum L. 1% Salvia eremophila 1%	Extraction of the ground peel of P. granatum or aerial parts of S. eremophila with aqueous methanol (80% v/v) in the absence and presence of acetic acid (HOAc, 5% v/v), filtration and vacuum-concentration	After 14 days of ripening samples supplemented with P. granatum extract contained the highest concentration of histamine, cadaverine, and putrescine among the tested variants. The levels of those BAs were ca. 2.6 times higher than in the samples without extracts, supplemented with 120 ppm of nitrite, that contained the lowest level of examined BAs.Samples fortified with S. eremophila extract contained ca. 1.5 times higher concentrations of tested BAs.Application of each extract individually in combination with 60 ppm of nitrite was more effective regarding inhibition of BAs accumulation compared to the application of plant extracts alone.	Abedi et al. 2023
Chinese sausages	Tea polyphenol (TP) 0.05% (g/g)Epigallocatechin gallate (EGCG) 0.05% (g/g)palmitoyl-TP (pTP) 0.05% (g/g) palmitoyl-EGCG (pEGCG) 0.05% (g/g)	purchased, powder	In comparison to control, after 2 days of fermentation and 4 wk of ripening:The total concentration of six analyzed BAs was reduced by 22.64, 36.89, 44.15, and 55.08% in the samples supplemented with TP, EGCG, pTP, and pEGCG, respectively.Application of pEGCG was the most effective regarding putrescine, tyramine, spermine, and spermidine. Their contents were reduced by 40.52, 61.25, 44.17, and 40.02%, respectively. Additionally, the concentrations of cadaverine and histamine in the samples with pEGCG decreased by 74.02 and 3.89%, respectively.Histamine content was reduced the most efficiently in the samples fortified with TP (9.22%), while cadaverine in pTP group (ca. 10 times lower than in control).	Zhou, Mo, Tang, et al. 2023
Chinese sausages	Tea polyphenol (TP) 0.05% (g/g) Epigallocatechin gallate (EGCG) 0.05% (g/g) palmitoyl-TP (pTP) 0.05% (g/g) palmitoyl-EGCG (pEGCG) 0.05% (g/g)	purchased	In comparison to control, after 23 days of fermentation:Total BAs content was reduced by 47.41, 37.17, 37.19, and 26.24% in the samples fortified with EGCG, pEGCG, pTP, and TP, respectively.EGCG and pEGCG were the most efficient regarding tyramine (reduced by approx. 41%), as well as histamine and putrescine contents reduction, while pTP addition resulted in the lowest content of spermidine.Cadaverine concentration was reduced by 76.72, 80.23, 88.53, and 63.51% following fermentation with TP, EGCG, pTP, and pEGCG, respectively.Spermine content was reduced by 20.66, 46.64, 28.51, and 27.13% in TP, EGCG, pTP, and pEGCG groups, respectively.2-phenylethylamine content decreased to not detected values in each treatment group.	Zhou, Mo, Wang, et al. 2023
Smoked horsemeat sausage	Coreopsis tinctoria Nutt. essential oil (CEO): 0.312% Coreopsis tinctoria Nutt. essential oil microcapsules (CEOM): 0.312%	purchased	In comparison to control, after 28 days of fermentation:Histamine was reduced by 63.37% and 80.42% in the samples supplemented with CEO and CEOM, respectively.Tyramine was reduced by 58.22% and 65.25% in the samples supplemented with CEO and CEOM, respectively.Putrescine was reduced by ca. 35.00% and 62.04% in the samples supplemented with CEO and CEOM, respectively.Cadaverine content was ca. 3 and 6 times lower in CEO and CEOM groups, respectively.Tryptamine content was ca. 1.5 and 2 times lower in CEO and CEOM groups, respectively.There was no significant difference regarding concentrations of spermine, spermidine and 2-phenylethylamine.	Li et al. 2023

Table 4. Effect of plant-origin additives on BAs in fermented fish and seafood.

Fermented product	Applied plant additive and its dose	Form of additive	Main effects on BA content	Reference
Myeolchi-jeot fermented anchovy	Garlic extract	ethanolic extract (95% ethanol)	In a model system the total content of analyzed BAs in the samples with garlic extract was reduced by 8.7% compared to control.	Mah, Kim, and Hwang 2009
Fermented fish sauce	Ethanolic extracts of:GingerGarlicCinnamonStar anise	ethanolic extracts (95% ethanol)	In comparison to control, garlic and star anise extracts significantly reduced histamine content by 30.49 and 8.09%, respectively, while application of ginger or cinnamon extracts increased histamine concentration by 4.82 and 13.36%, respectively.Ginger, garlic, and star anise extracts significantly reduced putrescine concentration by 11.79, 17.65, and 32.24%, respectively. Application of cinnamon extract resulted in the decrease of putrescine content by 3.22%.Tyramine concentration was reduced significantly by 26.03 and 22.79%, because of star anise and garlic extracts application, respectively. The use of cinnamon or ginger extracts reduced tyramine concentration by 2.92 or 13.12%, respectively.Addition of garlic, ginger, and star anise extracts reduced spermidine content by 37.20, 33.45, and 27.25%, respectively. Application of cinnamon extract reduced spermidine content by 5.65%.Total amount of four BAs was significantly reduced by 27.17, 23.25, and 13.49% thanks to application of garlic, star anise, and ginger ethanolic extracts, respectively.	Zhou et al. 2016
Fermented sardine	Plant + cell-free extracts from LAB:thyme extract + cell-free extract from Lactobacillus plantarum FI8595 (TH-LP)thyme extract + cell-free extract from Pediococcus acidolactici ATCC 25741 (TH-PA) laurel extract + cell-free extract from Lactobacillus plantarum FI8595 (LA-LP)laurel extract + cell-free extract from Pediococcus acidolactici ATCC 25741 (LA-PA)	plant extracts obtained through carbon dioxide supercritical fluid extraction water cell-free extracts	The use of thyme extract with cell-free extract from Pediococcus acidolactici ATCC 25741 resulted in lower cadaverine concentration compared to the control and other variants after 8 wk of storage.After 8 wk, putrescine reached the highest value in TH-LP combination, while the lowest content of this BA was obtained in LA-PA variant.After 8 wk, spermine concentration was the lowest in LA-LP and LA-PA and the highest in TH-LP. Spermidine was present in smaller amounts, the highest value after 8 wk of storage was measured in TH-LP variant.Histamine content was insignificantly higher in PA after 8 wk of storage.Similarly, tyramine concentration was the highest in PA after 8 wk, while significantly lower in LP.Plant extracts induced lower concentrations of dopamine and serotonin after 8 wk of storage.	Kuley et al. 2018
Shrimp paste	tea polyphenols 0.3% 6-gingerol 0.3% tea polyphenols 0.15% + 6-gingerol 0.15%	purchased	The combination of tea polyphenols (0.15%) and 6-gingerol (0.15%) was the most effective in inhibiting the formation of BAs. After 160 d of storage the contents of putrescine, cadaverine, histamine, phenylethylamine, tryptamine, and tyramine were reduced by 75.05, 62.82, 72.89, 68.17, 61.00, and 65.85%, respectively, compared to control.Application of tea polyphenols or 6-gingerol alone also resulted in a significant reduction of BAs in comparison to control. 6-gingerol was even more effective than tea polyphenols regarding the concentrations of cadaverine, histamine, and tyramine. It reduced the content of aforementioned BAs by 59.15, 68.17, and 62.97%, respectively. The concentrations of putrescine, phenylethylamine, and tryptamine were reduced by the addition of 6-gingerol by 67.70, 59.03, and 54.01%, and did not differ significantly from the values obtained when tea polyphenols alone were applied.The application of tea polyphenols alone resulted in the reduction of putrescine, cadaverine, histamine, phenylethylamine, tryptamine, and tyramine by 63.64, 55.48, 63.06, 58.75, 56.06, and 59.79%, respectively.Spermine and spermidine were not detected in shrimp paste samples during the experiment.	Cai, Liu, et al. 2015
Chouguiyu – fermented fish product	Papain:10 U/g50 U/g200 U/g400 U/g	purchased	Addition of papain significantly reduced BA concentration in comparison to control (no papain addition) regardless of applied dose. Application of 200 U of papain per 1 g of fish reduced total BA content by 89.47% and was the most efficient amongst tested variants.	Xu et al. 2022

Table 5. Effect of plant-origin additives on BAs in fermented dairy products.

Fermented product	Applied plant additive and its dose	Form of additive	Main effects on BA content	Reference
Gouda cheese	Zataria multiflora Boiss. essential oil:0.05% v/v0.1% v/v0.2% v/v0.4% v/v	EO of air-dried aerial parts of Z. multiflora	After 90 days of ripening significant reduction of tyramine and histamine in comparison to control occurred in the samples where 0.1, 0.2 or 0.4% of EO were applied. The higher the concentration of EO added, the greatest the reduction of analyzed BAs. Tyramine content was reduced by 5.05, 21.94, and 34.40%, while histamine content was reduced by 14.00, 29.61, and 46.60% in the samples with 0.1, 0.2, and 0.4% of EO addition, respectively.	Es’haghi Gorji et al. 2014
Unripened curd cheese	Fermented plants at the concentration of 3% w/v: Satureja montana L. sakei:Traditional submerged fermentation (SMF)Solid state fermentation (SSF) P. acidilactici SMFSSF P. pentosaceus SMFSSF Rhaponticum carthamoides L. sakei SMFSSF P. acidilactici SMFSSF P. pentosaceus SMFSSFNon-fermented plants: Satureja montana Rhaponticum carthamoides	fermented plants: liquid fermented LAB bioproducts prepared with the use of dried, ground plants Non-fermented plants: dried and ground to particle size of 0.5 − 2.0 mm	Either type of plant, bacteria strain or type of fermentation affected the content of BAs in the final product.Generally, both fermented and non-fermented plants addition resulted in a higher total BA concentration in cheese than in control samples – without plant additives.Cadaverine contributed most to the total BAs content in all samples with fermented plants addition, as well as in the sample with non-fermented Rhaponticum carthamoides.The highest total BA content was analyzed in cheese with the addition of Rhaponticum carthamoides fermented with L. sakei under traditional submerged fermentation conditions and was above 500 times higher than in control. Cadaverine represented above 80% of the sum of 8 analyzed BAs in the aforementioned sample.The lowest concentration of total BAs amongst the samples with plant additives was obtained in cheese with Rhaponticum carthamoides fermented with L. sakei under solid state fermentation and was almost 11 times higher than in control.Tyramine was absent in almost all samples with plant additives.The highest histamine concentration was analyzed in the sample with non-fermented Satureja montana addition and reached 25.17 mg/kg.	Mozuriene et al. 2016
Cheddar cheese	Bunium persicum Boiss. essential oil (BPEO) [% (v/v)]:0.050.10.20.4	hydrodistillation of EO	The higher the BPEO concentration, the more effectively BA accumulation was inhibited.In comparison to control, after 90 days of ripening histamine and tyramine contents were reduced by 48.59% and 35.94%, respectively, in the samples containing 0.4% BPEO.	Raoofi, Noori, and Khanjari 2023

Table 6. Effect of plant-origin additives on BAs in plant-based fermented food.

Fermented product	Applied plant additive and its dose	Form of additive	Main effects on BA content	Reference
Sauerkraut	onion (350 g/kg)caraway (10 g/kg)	fresh, shredded onion caraway seeds	After 14 days of fermentation: Fermentation at 18 °C resulted in the reduction of cadaverine (by 21.77% in the samples with caraway and 54.43% in the samples with onion) and tyramine (by 45.42 and 38.79%, respectively) in comparison to control, as well as reduction of phenylethylamine (by 22.70%) in the samples with onion.Fermentation at 31 °C resulted in the reduction of tryptamine (by 59.61 and 52.51% in the samples with caraway and onion, respectively).After 12 wk of storage: significantly reduced content of cadaverine and phenylethylamine in the samples fermented at 18 °C with onion and significantly higher concentration of putrescine, cadaverine and spermidine in the samples fermented at 18 °C with caraway. Storage of samples fermented at 31 °C resulted in a significant reduction of putrescine (when caraway was added) and spermine (in the samples with onion).	Majcherczyk and Surówka 2019
Tarhana – fermented cereal product	Grapeseed extract [g/kg]:4816	powder	Application of 8 or 16 g/kg of the extract resulted in a significant reduction of putrescine (by 84% and 90%, respectively), cadaverine (to values below limit of detection), as well as total amount of analyzed BAs (by 77% and 83%, respectively) in comparison to control (without grape seed extract) after 6 months.	Akan and Ocak 2019
Fermented soybean paste	Grapefruit seed extract (300 mg/kg)Catechins (3 g/kg) –epicatechinepicatechin gallateepigallocatechinepigallocatechin gallate	purchased grapefruit seed extract catechins – powder	Addition of grapefruit seed extract resulted in a 5% reduction of total BAs and 16% reduction of putrescine content after 40 days of storage.Application of epicatechin, epicatechin gallate, epigallocatechin, and epigallocatechin gallate resulted in a reduction of total BA content by 29, 35, 44, and 37%, respectively. Putrescine concentration was reduced by 64, 60, 76, and 73%, respectively, and cadaverine by 24, 34, 42, and 37%, respectively.	Lee et al. 2018
Gangjang – Korean soy sauce	200 g of coriander per 2 kg Meju blocks	fresh coriander	Significant reduction of histamine, putrescine, and tyramine by 48.03, 43.96, and 57.74%, respectively.	Mannaa, Seo, and Park 2020
Meju – fermented soybean-based product	Allium sativum (Garlic clove):1% (GAM1)10% (GAM10) Ginkgo biloba (leaves):1% (GIM1)10% (GIM10) Nelumbo nucifera (Lotus leaves):1% (LOM1)10% (LOM10)Mix of A. sativum, G. biloba and N. nucifera (1:1:1)1% (MIM1)10% (MIM10)	ethanolic extracts	Mostly, BA concentration was higher in the samples with plant extracts addition than in control. An exception was putrescine concentration in GAM1, GAM10, and LOM1, histamine in GAM1, and GAM10, as well as spermine in GIM1, GIM10, MIM1, and MIM10. Sum of nine analyzed BAs was the highest in the LOM10 sample and was almost 6 times higher than total BAs in control. The lowest difference compared to control regarding total BAs represented GAM1 sample (0.5% higher sum of BAs).	Shukla et al. 2018
Doenjang – traditional Korean fermented soybean paste	Ground garlic:2%6%10%Heat-dried garlic:1%2%3%Freeze-dried garlic:1%2%3%	paste of fresh garlic dried garlic	After 2 months of fermentation:Reduction of the sum of 9 analyzed BAs in almost all tested variants, except 3% heat-dried garlic. Despite the highest cadaverine concentration, above 5 times higher than in control, application of 1% heat-dried garlic was the most effective in the reduction of the total BA content (by 51.34% in comparison to control). Concentration of spermidine was at similar level in all tested variants and remained in the range 1.87–1.92 mg/100g.Spermine content was reduced to not detectable value after application of 1 or 2% heat-dried garlic and 1% freeze-dried garlic.Application of 1% heat-dried garlic resulted in the greatest reduction of histamine and tyramine, by 66.24 and 86.55%, respectively, compared to control.The greatest reduction of tryptamine and putrescine, by 51.53 and 55.56%, respectively, was obtained because of an application of 3% freeze-dried garlic.Samples with 1 or 3% freeze-dried garlic were characterized by maximum reduction of cadaverine 13.19%.Application of 6% ground garlic resulted in the greatest reduction of 2-phenylethylamine content, by 37.13%.Agmatine remained below limit of detection in all tested variants.	Bahuguna et al. 2019
Kimchi	White radish ground:0 g4.5 g18 gRed pepper powder:0 g2 g8 gGarlic ground:0 g1.5 g6 gGinger ground:0 g0.25 g1 gWelsh onion ground:0 g1.5 g6 gSticky rice porridge:0 g1.75 g7 g	ground fresh white radish, garlic, ginger, and Welsh onion red pepper – powder all additives purchased from kimchi manufacturer	After 7 d of fermentation:Amongst plant additives the only one that reduced total BA content (by 4.46% in comparison to control) was red pepper powder applied at the highest level (8 g).Histamine was present in kimchi mainly due to the addition of fish sauce. None of the plant additives reduced the concentration of this BA.Tyramine concentration was reduced in the samples with zero or low white radish, zero or high garlic, low or high red pepper powder, low Welsh onion, and high sticky rice porridge addition. Amongst them the greatest reduction resulted from lack of white radish addition – decrease by 43.03% in comparison to control.Cadaverine was reduced in the samples without white radish or with high level of garlic, whereby high garlic addition was more effective. Concentration of cadaverine decreased by 28.16% in comparison to control.Content of putrescine was reduced along with zero ginger, low sticky rice porridge or high red pepper powder addition. The biggest reduction (by 42.38%) resulted from lack of ginger.Low white radish addition caused the reduction of phenylethylamine and spermidine contents by 15.38 and 9.89%, respectively, and was the most effective amongst tested combinations regarding these amines.High white radish addition reduced the content of agmatine and spermine by 30.43 and 22.71%, respectively.Tryptamine content was reduced the most when high level of sticky rice porridge was added − 34.98% less of this amine than in control.	Kim, Dang, and Ha 2022
Doenjang – fermented Korean soybean paste	Extracts-based Meju (EMD) / traditional Meju with plant extracts added later (TMD): Allium sativum (Garlic clove):1%10% Ginkgo biloba (leaves):1%10% Nelumbo nucifera (Lotus leaves):1%10%Mix of A. sativum, G. biloba and N. nucifera (1:1:1)1%10%	dried extracts obtained by freeze-dried ethanolic extraction	After 12 months of fermentation:In TMD, each plant extract caused the reduction of total BA content compared to control – without extracts.In EMD only Doenjang with 1% of garlic or lotus extracts showed higher sum of nine analyzed BAs than in control. Application of 10% garlic extract resulted in the lowest content of total BAs.	Shukla et al. 2019
Bread	Cava lees:1%2%5%	lyophilized, ground	Addition of cava lees did not alter the contents of putrescine and spermidine, regardless of applied level. Tyramine, histamine, and spermine were absent in all samples. Cadaverine was absent in control bread sample while its content reached almost 3 mg/kg in all tested variants containing cava lees.	Hernández-Macias et al. 2021
Tarhana: a cereal based fermented food	Pomegranate seed extract (PSE):0.5%1.0%2.0%	powder, purchased	Addition of 0.5, 1.0, and 2.0 % of PSE resulted in the following percentage reduction of the total BA content: 36.33, 42.49, and 52.54%, respectively.After 6 months of storage putrescine concentration was reduced significantly only when 2.0% of PSE was applied and the reduction reached 30.92% in comparison to control sample – without PSE. Cadaverine content was reduced by 13.04, 23.33, and 33.69%, while tyramine by 21.08, 24.71, and 17.85%, respectively, when 0.5, 1.0, and 2.0% of PSE was added. Significant reduction of histamine was obtained only in the samples with 0.5 or 1.0% of PSE − 19.86, and 23.0% less than in the control, respectively. Spermidine content was reduced to not detectable value in the samples supplemented with 0.5 or 2.0% of PSE. 66.68% reduction of this amine was calculated for the sample with 1.0% of PSE. There was no significant difference between samples regarding spermine concentration.	Erol and Özdestan Ocak 2020
Cheonggukjang – a traditional Korean fermented soy food	Schizandra chinensis Baillon extract (5 mg/kg) Rosmarinus officinalis L. extract (5 mg/kg) Prunus mume Siebold & Zucc (5 mg/kg) Prunus mume Siebold et Zuccarini (5 mg/kg)	extraction with 70% ethanol, vacuum-evaporation and freeze drying	In comparison to control, tyramine content was reduced by 94.74, 90.85, 86.33, and 81.93% in the samples supplemented with extracts of Rosmarinus officinalis L., Schizandra chinensis Baillon, Prunus mume Siebold et Zuccarini, and Prunus mume Siebold & Zucc, respectively.	Oh et al. 2012
Soy sauce	Phytochemicals at concentration 200 mg/kg:QuercetinRutinGallic acidFerulic acidTea polyphenol Camellia oleifera Bamboo leaf extract	Except for Camellia oleifera extract all the phytochemicals were purchased. Camellia oleifera extract was prepared through extraction with 70% (v/v) aqueous ethanol and vacuum drying via N2	After 60 days of fermentation total BA content was reduced by 24.46, 29.35, and 23.95% in the samples supplemented with quercetin, tea polyphenols, and Camellia oleifera extract, respectively. Application of other phytochemicals resulted in the lower than 20% reduction of total BAs, except for samples with rutin, in which total BA level increased in comparison to control.After 120 days of the experiment total BA content was similar in the control and quercetin-supplemented samples.	Zhou, Zhang, et al. 2023

Figure 3. Main mechanisms of antimicrobial effects of plant additives on BA production. Created with BioRender.com.

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