Biopreservation of ready-to-eat Atlantic salmon (Salmo salar) by lactic acid bacteria: Effect on safety and quality parameters

Biopreservation of ready-to-eat Atlantic salmon (Salmo salar) by lactic acid bacteria: Effect on safety and quality parameters | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Biopreservation of ready-to-eat Atlantic salmon (Salmo salar) by lactic acid bacteria: Effect on safety and quality parameters Jelena Stupar, Sunniva Hoel, Jørgen Lerfall, Turid Rustad, Anita Nordeng Jakobsen This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7968166/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Lactic acid bacteria (LAB) are recognised as candidates for biopreservation because of their antagonistic activities. However, the LAB strain's efficacy in and compatibility with a specific food matrix must be emphasised to develop biopreservation as an efficient hurdle for industrial applications. The aim of the present study was to investigate the biopreservative potential of four lactic acid bacteria (LAB; Carnobacterium maltaromaticum 35, C. maltaromaticum 55, C. divergens 468, and Leuconostoc gelidum 406) in vacuum-packed (VP) ready-to-eat (RTE) salmon portions stored at 4℃ for 22 days. The evaluation was based on the strain's growth properties, their effect on the microbial community structure, their ability to inhibit an artificially inoculated strain of Listeria innocua (CCUG 15531) and their effect on chemical (ATP-degradation products and biogenic amines (BA)) and physicochemical properties (pH, colour, and water-holding properties (WHP)) of the salmon portions. All LAB strains grew well, increasing their concentration from 4.2–4.6 log CFU/g to 7.2–8.6 log CFU/g. The cultures outcompeted the population of the spoilage organism Photobacterium , which predominated the microbial community of the control samples. LAB strains did not negatively affect chemical and physicochemical properties. In the absence of inoculated LAB, L. innocua increased significantly from 3.4 ± 0.1 to 5.0 ± 0.3 log CFU/g, while in the presence of C. maltaromaticum 35, no significant proliferation was observed (p = 0.19). Thus, C. maltaromaticum 35 could be an efficient hurdle to ensure safe and microbiologically stable RTE salmon products. Lactic acid bacteria Biopreservation Listeria ready-to-eat salmon Physicochemical properties Figures Figure 1 Figure 2 Figure 2 Figure 3 Figure 3 1. Introduction Pre-rigor filleted vacuum-packed (VP) fresh salmon loins are nutritious and appealing to consumers as ready-to-eat (RTE) products. However, the production lacks efficient barriers to control the growth of the pathogenic bacteria Listeria monocytogenes , recognised as the causative agent of listeriosis. Although the general contamination level of Listeria spp. in fresh salmon is low (Noseda et al., 2014 ; Svanevik et al., 2021), their ability to persist in biofilms in food production environments (Fagerlund et al., 2021 ) and their ability to tolerate stressful conditions applied in food processing and packaging (Chan & Wiedmann, 2008 ; Noseda et al., 2014 ; Tuytschaever et al., 2023 ) challenge the safety of RTE salmon products. Furthermore, fresh seafood is highly perishable (Leroi, 2010 ), and the growth of spoilage bacteria belonging to genera of e.g. Photobacterium., Brochothrix and Pseudomonas can result in the production of total volatile basic nitrogen (TVB-N), sulphur compounds, biogenic amines (BA) and other spoilage products (Rathod et al., 2021 ) lowering the product quality. Biopreservation using lactic acid bacteria (LAB) represents a natural preservation strategy to ensure product safety and microbial stability with minimal effect on physicochemical and nutritional properties (Barcenilla et al., 2022 ; Strack et al., 2020 ). Food-fermenting LAB, including members of Carnobacterium and Leuconostoc genera, are ‘Generally Recognised as Safe’ (GRAS) and are given a Qualified Presumption of Safety (QPS)’ status by the Food and Drug Administration (FDA) and European Food Safety Agency (EFSA) (EFSA BIOHAZ Panel, 2021). This has made them interesting candidates for various applications. However, studies also link LAB to the production of BA and spoilage metabolites (Emborg et al., 2002 ; Wiernasz et al., 2017 ; Yazgan et al., 2021 ), which calls for diligent strain selection. LAB, including the genera Carnobacterium and Leuconostoc , have been widely investigated for biopreservative purposes of various foods (Angiolillo et al., 2018 ; Jang et al., 2015 ; Mei et al., 2020 ; Ramos et al., 2020 ; Saraoui et al., 2017 ; Shi & Maktabdar, 2022 ; Stupar et al., 2023 ; Tahiri et al., 2009 ; Wiernasz et al., 2020 ; Wong & Li, 2023 ; Woraprayote et al., 2018 ). However, their implementation in fresh fish remains relatively unexplored (Evangelista et al., 2023 ; Gómez-Sala et al., 2019 ). Although underexplored, species from the Carnobacterium and Leuconostoc genus have demonstrated the ability to inhibit the growth of several spoilage and pathogenic bacteria by the production of antimicrobial compounds such as organic acids, hydrogen-peroxide, bacteriocins (Begrem et al., 2020 ; Rao et al., 2023 ; Stupar et al., 2023 ), and competition for nutrients and attachment sites (Nilsson et al., 2005 ; Stupar et al., 2023 ; Wiernasz et al., 2020 ). Finding suitable LAB for the biopreservation of fresh salmon is challenging due to the clean sensory profile, short shelf life and low storage temperature of the product. LAB strains selected in the present study have shown inhibitory properties against strains of L. monocytogenes and L. innocua in vitro (Stupar et al., 2021 ) and have demonstrated suitable growth properties in fresh VP salmon at 4℃ (Stupar et al., 2023 ). The aim of the present study was to apply a polyphasic approach combining microbiological, chemical, and physicochemical factors to investigate the effect of selected Carnobacterium and Leuconostoc strains on safety and quality parameters in VP salmon portions stored at 4℃ for 22 days. Inoculation trials with non-pathogenic Listeria innocua (CCUG 15531) were applied, supported by previous data confirming the applicability of this strain as a substitute for L. monocytogenes (Stupar et al., 2021 ). This is, to the best of our knowledge, the first study exploring the effect of LAB on the safety, chemical and physiochemical properties of RTE VP salmon portions. 2. Materials and methods 2.1. Experimental design The experiment aimed to evaluate the impact of LAB inoculation on VP salmon portions with respect to microbial, chemical and physicochemical quality, and safety parameters. Experimental groups (Table 1 ) included: i) control - non-inoculated salmon portions; ii) portions inoculated with a single LAB strain ( C. maltaromaticum 35 (C.m.35), C. maltaromaticum 55 (C.m.55), L. gelidum 406 (Le.g.406) or C. divergens 468 (C.d.468)); iii) portions inoculated with the target organisms ( L. innocua ); and iv) portions inoculated with both LAB and the target. Two independent batches of pre-rigor filleted salmon (Batch I and Batch II, slaughtered on different dates, Section 2.2.3 ) were used: Batch I for experiments with C.m.55, and Batch II for C.m.35, C.d.468, and Le.g.406. All samples were vacuum-packed and stored at 4°C for 22 days. Total aerobic plate counts (APC), H 2 S-producing bacteria, LAB, and L. innocua were quantified on days 0, 7, 14 and 21 of storage (Section 2.2.4 ). Microbial community composition was assessed at day 14 of storage (Section 2.3 ). Chemical and physicochemical analyses were performed on day 0 (pre-inoculation), day 8 and day 22 post-packaging. Samples for chemical analysis were stored at -80℃ until further processing. Salmon portions inoculated with Listeria were not subjected to physicochemical, chemical or microbial community analysis to prevent cross-contamination in the food processing laboratory. Table 1 Experimental design showing inoculation and analysis of vacuum-packed salmon portions investigated. All samples were stored at 4°C for 22 days. Inoculation Vacuum packed salmon Sampling days for microbiological analyses Sampling days for chemical analyses Sampling days for physicochemical analyses Groups LAB (10 4 CFU/g) Target (10 3 CFU/g) Sample size (g) Culture-dependent Culture-independent ATP-degradation products, biogenic amines Water content, pH, colour, texture Control - - 100 ± 1, 10 ± 1 0,7, 14,21 14 0, 8, 22 0, 8, 22 LAB* C.m.35 C.m.55 Le.g.406 C.d.468 - 100 ± 1, 10 ± 1 0,7, 14,21 14 0, 8, 22 0, 8, 22 LAB*+T C.m.35 C.m.55 Le.g.406 C.d.468 L.innocua 10 ± 1 0,7, 14,21 - - - T - L.innocua 10 ± 1 0,7, 14,21 - - - LAB = Lactic acid bacteria; T = Target ( L. innocua ); *LAB inoculation with a single culture of Carnobactrium maltaromaticum 35 , Carnobacterium maltaromaticum 55 , Leuconostoc gelidum 406 or Carnobacterium divergens 468. Culture-dependent analysis included analysis of aerobic plate count, H 2 S-producing bacteria, lactic acid bacteria and Listeria spp. Culture- independent analysis included analysis of the microbial community composition. 2.2. Biopreservation of fresh salmon vacuum-packed fillets 2.2.1. Selection and preparation of LAB strains LAB strains were pre-cultured in De Man, Rogosa and Sharpe (MRS) (Oxoid, Norway) for 2.5 days at 25℃ under anaerobic conditions using GasPak EZ Anaerobe container system sachets with indicator (BD, Norway). A single colony was transferred to MRS broth (Oxoid) and incubated for 2.5 days at 8℃ to promote cold adaptation. Following incubation, cultures were standardised to an optical density (OD) of 0.22 at 600 nm (Shimadzu UV 1800, Germany), corresponding to approximately 10 8 CFU/mL (Stupar et al., 2021 ). For inoculation, 0.1 or 1 mL of the diluted cold-adapted cultures was applied to the surface of each salmon portion to achieve a concentration of approximately 10 4 CFU/g. 2.2.2. Preparation of target strain L. innocua was cultivated Brain Heart Infusion (BHI) agar (Oxoid) for 24h at 37℃. A single colony was picked and incubated in BHI broth (Oxoid) for 24h at 15℃ to promote adaptation to low temperature (Stupar et al., 2021 ). The resulting culture was adjusted to an OD₆₀₀ of 0.1 (Shimadzu UV 1800; Shimadzu, Germany). An aliquot of 0.1 mL was applied to the surface of each salmon portion (Section 2.2.4 ), corresponding to an inoculation concentration of approximately 10 3 CFU/g. 2.2.3. Sample preparation and packaging Atlantic salmon ( Salmo salar ) was obtained from a local slaughterhouse and transported on ice to NTNU, where it was hand-filleted within 5 h post-mortem. Back loin portions were prepared by cutting into 10 ± 1 g pieces for microbiological and chemical analysis, and into 100 ± 1 g pieces for physicochemical analysis. All samples were stored on ice until processing the following day. Randomised pieces were placed on an absorbent pad (Tommen Gram, Norway) and inoculated with 1% (v/w) LAB and/or target on the salmon surface as described by Stupar et al. ( 2023 ). All samples were packed in vacuum (20 µm polyamide (PA)/70-µm polyethylene (PE) bag (120 9 80 mm, Star-Pack Productive, Boissy-'Aillerie, France) with a Webomatic Supermax- C vacuum machine (Webomatic, Germany) and stored at 4℃ for 21/22 days (21 days for microbiological and 22 days for chemical and physicochemical analyses). The 21/22 storage period was chosen to follow the microbial growth and potential re-growth of the target strain beyond the shelf life of commercial products (approximately 10 days). 2.2.4. Microbiological analysis The salmon sample (10 g) was homogenised with 90 g of peptone water (1 g/L peptone (Oxoid) and 8.5 g/L NaCl (VWR, Belgium)), using a stomacher (IUL Masticator, Spain). Homogenates were serially diluted and plated on media. Lyngby's Iron agar (IA; Oxoid) supplemented with 0.04% L-cysteine (Sigma-Aldrich, Norway) was used to quantify APC and H 2 S-producing bacteria by incubation for 72 ± 6h at 22℃. LABs were enumerated on MRS agar using anaerobic incubation for 2–5 days at 25℃. L. innocua was quantified on Brilliance Listeria agar (BLA; Oxoid), supplemented with Brilliance Listeria supplement (Oxoid), incubated at 37°C for 24 ± 1 h. 2.3. Culture-independent microbial community analysis Total genomic DNA was extracted from homogenates (1:10 in sterile peptone water; n = 3) collected on day 14. A sample of 5 ml was centrifuged (83 x g, 5 min, ∼ 20℃) to remove most of the fish particles. The supernatant was again centrifuged (2061 x g, 15 min, ∼20℃), and the pellet was resuspended in 1 ml of peptone water before a final centrifugation(6160 x g, 5 min, ∼20℃). The pellets were used for DNA extraction using the PowerFood® DNA isolation kit (MoBio Laboratories Inc., Carlsbad, CA, USA). Quantitative and qualitative analysis of the extracted DNA was assessed by spectrophotometry (PowerWave XS, BioTek® ) and 1% gel electrophoresis. The V1-V3 hypervariable region of the 16S rRNA gene (~ 490 bp) was amplified using primers 16S_f (5′ AGAGTTTGATCATGGCTCAG 3′) and 16S_r (5′ GTATTACCGCGGCTGCTG 3′) (Leser et al., 2002 ; Weisburg et al., 1991 ). Amplicon sequencing and microbial community analysis were performed using the INVIEW Microbiome Profiling 3.0 package (Eurofins Genomics, Germany) on an Illumina MiSeq, including amplicon library generation and standardised bioinformatic, as described by Tsoukalas et al. ( 2023 ). 2.4. Chemical analysis 2.4.1. Extraction procedure Approximately 2 g of frozen sample was grated using a stainless-steel grater and homogenised for 2 min with an Ultra Turrax T25 Basic (Janke & Kunkel IKA®-Labortechnik, Staufen, Germany) in 10 mL 7% trichloroacetic acid (TCA). Subsequently, 1.5 ml of 1 mol/l potassium hydroxide was added, vortexed, and centrifuged (16,000×g, 10 min, 4°C). The resulting supernatants were filtered through a 0.45 µm nylon filter; one portion was used for BA analysis, and the remaining portion was used for analysis of ATP-degradation products (Section 2.4.2 ). 2.4.2. Analysis of ATP-degradation products ATP-degradation products were analysed in samples collected on day 0 (prior to inoculation) and on days 8 and 22 post-packaging. Quantification was performed according to the method described by Lerfall et al. ( 2018 ), with a detection limit of 5 µmol/L. Ki-value (Karube et al., 1984 )d value (Luong et al., 1992 ) were calculated by using the following equations: H-value (%) = [Hx / (IMP + HxR + Hx)] × 100, (1) Ki-value (%) = [(HxR + Hx) / (IMP + HxR + Hx)] × 100 (2) where HxR = Inosine, Hx = Hypoxanthine, and IMP = Inosine monophosphate. 2.4.3. Biogenic amines Fish extract for BA determination was prepared as described in Section 2.4.1 . The extract was neutralised by potassium hydroxide (1M KOH), derivatised with benzyl chloride (99%, Sigma-Aldrich, CAS:98-88-4) for 20 min at room temperature. Benzylated amines were extracted twice with diethyl ether, and the combined organic phase was evaporated to dryness under nitrogen at 30°C. The residue was reconstituted in acetonitrile–water (90:10). BA quantification was performed by UHPLC as described by Lerfall et al. ( 2018 ), with a detection level of 0.05 µmol/L, using a YMC-Triart PFP (100 x 2 mm, 1.9 µm) column connected to an Agilent 1290 chromatography (Agilent Technologies, Paolo Alto, CA, USA) and an Agilent 1260 diode array UV-Vis detector. The loaded samples were detected at 254 nm using H 2 O/acetonitrile-based elution. 2.5. Physicochemical parameters All analyses (water content, drip loss, muscle pH, texture and colour analysis) were performed in triplicate on salmon pieces (100 ± 1g) from both LAB-inoculated and non-inoculated controls. Selected analyses were performed on days 0 (prior to inoculation), day 8 and day 22. 2.5.1. Water content and muscle pH Water content (WC) was determined according to Skipnes et al. ( 2007 ). Standardised muscle pieces were prepared by pressing the fish muscle with a metal cylinder (diameter 31 mm) to obtaine pieces 6 mm in height. Each piece was transversally divided into two equal parts, and the bottom piece was used for WC analysis. Samples were weighed and dried for 16h at 105℃. Drip loss (DL) was calculated as the percentage difference (%) of the weighed sample (g) to its initial weight (g). Muscle pH was measured by a portable pH meter (Hach HQ40d multi-Portable Meter, Hach, USA) with a puncture pH electrode (Hach Intellical™ PHC108, Hach, CO, USA). 2.5.2. Texture analysis Breaking force was measured using a Texture Analyser TA-XT plus (Stable Micro Systems Ltd, England) equipped with a flat-ended cylindrical probe (12.7 mm P/0.5). Breaking force was defined as the force (N) required to rupture the fillet surface, while firmness was recorded as the force at 60% compression. Data acquisition and analysis were performed using the Texture Exponent light software for Windows (version 4.12, SMS). 2.5.3. Colour analysis Color measurements of selected samples were performed using a DigiEye full system (VeriVide Ltd., Leicester, UK) equipped with a digital camera (Nikon D80, 35 mm lens, Nikon Corp., Japan). Colour parameters (L* = lightness, a*= redness, b*= yellowness, C*= Chroma and h* = hue angle) were obtained using software DigiPix (version 2.8). Total colour difference (ΔE) was calculated based on CIE (1994). Chroma (C*) and hue angle (h*) were calculated by the following Equations: C* = (a* 2 + b* 2 ) 1/2 (6) h* = arctan (b*/a*) (7) Colour changes (Δ-values) were calculated by the following Equation: Δ colour parameter at day x = Colour average measurement of LAB-inoculated sample - Colour average measurement of control sample (8) where Δcolour parameter (L*, a*, b*, C*, h*, ΔE) (n = 3) at the same storage day. 2.6. Statistical analysis Statistical analyses were conducted using IBM SPSS Statistics software (version 28, IBM Corporation, USA). Microbial counts were log-transformed and expressed as mean ± standard deviation (SD). Log-transformed bacterial counts were fitted to the Baranyi and Roberts ( 1994 ) primary growth model using the DMFit program ( www.combase.cc ). Differences between experimental groups were assessed by using one-way ANOVA with Tukey's pairwise comparisons and Independent Samples Test at a 5% significance level (p < 0.05). A general linear model (GLM) was applied to evaluate the antilisterial effect of LAB during storage and the influence of batch variations on target strain growth (p < 0.05). Pearson’s correlation was used to examine the effect of storage time on Ki-values. All experimental groups were analysed in triplicate, and results are presented as mean values ± standard deviation (SD). As no significant differences (p < 0.05) in bacterial counts were observed between Batch I and Batch II for both the Control group (non-inoculated samples) and the Target group (samples inoculated with L. innocua , positive control), data from both batches were combined and reported as mean values (n = 6) ± SD. The only exception is for the culture-independent analysis, where the controls from different batches were presented separately (n = 3 ± SD). 3. Results and Discussion 3.1. Growth characteristics of LAB in RTE VP salmon portions at 4℃ In the present study, four LAB strains, C. maltaromaticum (C.m.35), C. maltaromaticum (C.m.55), C. divergens (C.d.468) and L. gelidum (Le.g.406), were selected for biopreservation of RTE VP salmon portions for 21 days at 4℃. The strains were selected according to their origin in RTE seafood products, antimicrobial activity against Listeria spp., growth properties in salmon juice, and ability to grow in VP salmon portions at refrigerated temperature (Stupar et al., 2023 ; Stupar et al., 2021 ). All strains displayed growth in the VP salmon at 4℃ without a lag phase, increasing from the initial concentration of 4.2–4.6 log CFU/g to a final concentration of 7.2–8.6 log CFU/g during the storage period (Table 2 ). In comparison, the concentration of LAB in the control samples increased from 1.9 ± 0.2 to 6.6 ± 0.2 log CFU/g (Table 2 ). Compared to non-inoculated samples, LAB counts were significantly higher in inoculated samples, indicating their dominance during the storage period (one-way ANOVA, p < 0.05), except for C.d.468 on days 14 and 21 (Fig. 1 A). Among the applied LAB strains, Le.g.406 reached the highest maximum growth rate (µ max ) and maximum population density (Y max ), while C.d.468 had the lowest values (Table 2 ). The results align with growth kinetic parameters obtained previously for the same strains in a salmon juice model system at 4℃ (Stupar et al., 2023 ), demonstrating that the salmon juice model system is reliable for assessing growth kinetic parameters in relevant conditions. No significant differences between MRS and IA plate counts were observed for any of the inoculated or non-inoculated samples (Fig. 1 A and Fig. 1 B), except for Le.g.406 (Independent Samples Test, day 0 (p < 0.001), day 7 (p = 0.035), day 14 (p < 0.001) and day 21 (p = 0.007)), implying that bacteria quantified on IA corresponds to the inoculated LAB. Table 2 Growth kinetic parameters for selected LAB quantified on MRS agar. The growth kinetic parameters (initial count (log CFU/g), maximum growth rate, µ max (logCFU/g/day), maximum population density, Y max (log CFU/g), and lag phase (day) are estimated by the primary model of Baranyi and Roberts ( 1994 ). R 2 -coefficient of determination; SE of Fit-standard error of fit; NL- no lag phase. Group Initial count µ max Y max R 2 SE of Fit Lag phase (log CFU/g) (log CFU/g/d) (log CFU/g) (days) Control 1.9 ± 0.2 a 0.32 ± 0.03 b 6.6 ± 0.2 a 0.99 0.23 NL C.m.35 4.2 ± 0.2 b 0.42 ± 0.04 bc 8.6 ± 0.1 b 0.99 0.21 NL C.m.55 4.3 ± 0.1 b 0.45 ± 0.01 c 8.6 ± 0.1 b 0.99 0.07 NL Le.g.406 4.6 ± 0.1 b 0.48 ± 0.02 c 8.6 ± 0.1 b 0.99 0.08 NL C.d.468 4.5 ± 0.2 b 0.20 ± 0.04 a 7.2 ± 0.2 ab 0.96 0.26 NL p-value p < 0.05 p < 0.05 p = 0.05 Small letters (abc) indicate significant differences between the groups (LAB-inoculated samples and controls), with the corresponding p-value at the bottom of each column, calculated by one-way ANOVA (p < 0.05). C.m.35: C. maltaromaticum 35; C.m.55: C. maltaromaticum 55; Le.g.406: L. gelidum 406; C.d.468: C. divergens 468. Although an accurate lag-phase determination requires more sampling points, no detectable lag phase suggests that the LAB strains adapted well to the salmon matrix and the applied storage conditions (VP and 4℃), probably because they originate from refrigerated seafood products (cold-smoked salmon (CSS), sushi and gravlax) (Stupar et al., 2021 ). This is also supported by previous findings observed in salmon juice and pre-rigor filleted salmon, where more frequent measurements were conducted (Stupar et al., 2023 ). The same adaptation and dominance of artificially added C. maltaromaticum, Lactococcus piscium and L. gelidum isolated from CSS and fresh salmon was observed in inoculated gravlax (Wiernasz et al., 2020 ). However, the study of Aymerich et al. ( 2019 ) showed that LAB ( Lactobacillus sakei ), isolated from other food matrices (meat), can also be applied for the biopreservation of CSS. For biopreservative purposes, the absence of lag phase followed by a high growth rate for the inoculated LAB strain can potentially prevent indigenous spoilage microbiota and limit the production of undesirable metabolic products (Huang et al., 2021 ; Zhou et al., 2021 ). In addition, some LAB might produce several metabolic products with antagonistic effects on undesirable microbiota, where even beneficial microbiota such as endogenous LAB can be affected. Thus, the ability of the LAB strain to prevent growth of undesirable microorganisms (both spoilage microorganisms and pathogens) and, at the same time, co-exist with beneficial microbiota is crucial for its implementation in food products (Woo et al., 2021 ). No H 2 S-producing bacteria, representing fish spoilage indicator bacteria, were observed during storage, except for control samples at the end of storage (0.58 log CFU/g). Some LAB strains are associated with food spoilage due to H 2 S-production when metabolising cysteine (Leroi, 2010 ). Moreover, the low level of H 2 S-producing bacteria in the non-inoculated samples at the end of storage confirmed that the VP salmon was not spoiled by H 2 S-producing bacteria, also observed by Rode and Hovda ( 2016 ) during 26 days of storage of VP salmon. In general, different levels of this bacterial group are reported for VP salmon. The study of Jakobsen et al. ( 2022 ) detected low counts of H 2 S-producing bacteria (< 3.8 log CFU/g) in VP salmon fillets stored at 4℃. On the contrary, the study of Hansen et al. ( 2009 ) reported the levels of H 2 S-producing bacteria to be above 6 log CFU/g after 15 days in VP salmon fillets at 1.2℃. In addition, microbial contamination is highly dependent on conditions during harvesting, degree of cross-contamination and type of processing and packaging (Chan et al., 2021 ; Rollini et al., 2016 ). By applying LAB biopreservation, microbial contaminants on salmon fillets can probably be inhibited by non-specific competition between populations (Jameson-effect interaction), thereby prolonging salmon shelf life and quality (Bolívar et al., 2021 ). 3.2. Effect of LAB on microbial community composition in VP salmon The effect of the four LAB strains (C.m.35, C.m.55, Le.g.406, and C.d.468) on the microbial community composition of RTE VP salmon was evaluated in samples collected on day 14. Illumina MiSeq sequencing generated 1 244 429 raw sequence reads, of which 1 244 101 sequences were obtained after quality processing, chimaera detection and filtering. A total of 909 382 (73.1%) sequences were assigned to operational taxonomic units (OTUs), ranging from 5 283 to 24 321 reads per sample. Endogenous LAB was present in control samples of both Batch I and II, but the relative abundance of LAB was significantly higher in Control 1 (Independent Samples Test, p = 0.003). The same significant batch-to-batch variation in the abundance of endogenous LAB was observed using culture-dependent analysis (Independent Sample Test, p < 0.001) (data not shown). The microbial community composition of Control 1 was dominated by LAB represented by unclassified Lactobacillales (relative abundance of 48.7 ± 12.3%) and Leuconostoc (5 ± 0.8%), and in addition, Photobacterium (47 ± 12.9%), while Control 2 was dominated by Photobacterium with a relative abundance of 98.1 ± 2.4% (Fig. 2 ). In LAB-inoculated samples, the community composition on day 14 corresponded to the inoculated LAB. The relative abundance of Leuconostoc in the samples inoculated with strain Le.g.406 was 96.5 ± 1.74% and thus significantly different from Control 2 where 1.76 ± 2.1 of unclassified Lactobacillales was detected (Independent Sample Test, p < 0.001). The relative abundance of Carnobacterium in the samples inoculated with strain C.d.468 was 64.7 ± 17.5% and thus significantly different from Control 2 (Independent Sample Test, p = 0.003). In samples inoculated with C.m.35 and C.m.55, 81 ± 3.8 and 76.8 ± 12.5% of the microbial population were classified at the order level ( Lactobacillales ), respectively (Fig. 2 ). Although the LAB population could not be assigned to the genus level, it can be assumed that the dominating OTU, assigned as Lactobacillales , results from the inoculated strains. Furthermore, Leuconostoc constituted 10.3 ± 2.1 and 7.9 ± 0.7%, while the relative abundance of Photobacterium was 8.4 ± 6.2 and 15.5 ± 12.9% in samples inoculated with C.m.35 and C.m.55, respectively (Fig. 2 ). It is evident the chosen sequencing approach holds certain limitations regarding the taxonomic resolution of bacteria belonging to the order Lactobacillales (O’Callaghan et al., 2021 ; Rintala et al., 2017 ), and a combined approach with full-length sequencing of the 16S rRNA gene could enable more accurate identification of the dominating LAB genera. The results demonstrated that inoculated LAB affected the salmon’s natural microbial community composition by outcompeting Photobacterium. Species belonging to Photobacterium , e.g. Photobacterium phosphoreum , are commonly recognised as fish spoilers (Antunes-Rohling et al., 2019 ; Dalgaard et al., 1998 ; Tsoukalas et al., 2024 ) due to a strong correlation with the production of BA (Bjornsdottir-Butler et al., 2018 ; Li et al., 2023 ), and ability to reduce trimethylamine oxide (TMAO) to the undesirable odourant trimethylamine (TMA) (Jääskeläinen et al., 2019 ; López-Caballero et al., 2002 ; Sørensen et al., 2020 ). Moreover, Photobacterium is shown to be an H 2 S-producer under certain conditions (Tsoukalas et al., 2023 ), but this was not confirmed in the present study as only low counts (< 1 log CFU/g) of H 2 S-producing bacteria were detected in control samples at the end of storage. Accordingly, the absence of H 2 S-production by Photobacterium has also been observed by Kuuliala et al. ( 2018 ). Photobacterium is one of the most abundant genera constituting the microbial community of chilled, packaged fresh salmon (Hansen et al., 2009 ; Jääskeläinen et al., 2019 ; Li et al., 2023 ; Macé et al., 2012 ). Thus, the observed ability of the applied LAB strains to outcompete Photobacterium is of significant importance as inhibition of this spoilage bacterium is likely to extend the shelf-life of chilled MAP and VP fish (Dalgaard et al., 1998 ; Tsoukalas et al., 2023 ). 3.3. ATP-degradation products The ATP-degradation products were analysed on day 0 (prior to inoculation) and days 8 and 22. The ATP-degradation products AMP and IMP are associated with the umami taste (Johnson et al., 2013 ; Mouritsen, 2023 ), while loss of freshness and unpleasant flavours of the fish are related to IMP degradation to Ino and Hx (Yu et al., 2021 ). The Ki-value (Karube et al., 1984 ), highly correlated to the K-value, is a freshness indicator mainly recommended for the assessment of early changes during fish storage (Hong et al., 2017 ), while H-values are more effective in expressing the influence of bacterial enzymes on the freshness and spoilage of salmon. No significant effect on the Ki-value and H-value (Table 3 ) was observed when applying biopreservation. The initial Ki-value in control samples was 34.5 ± 2.8, with a significant increase at the end of storage, reaching 99.9 ± 0.2 (r = 0.919, p < 0.001). No adverse effect of inoculated LAB on the accumulation of off-flavours and thus on Ki- and H-value suggests the high potential of these strains to be used in RTE salmon products. Table 3 ATP-degradation products. Concentration of inosine monophosphate (IMP (µmol/g)), inosine (Ino (µmol/g)), hypoxanthine (Hx (µmol/g)), Ki-value, and H-value (%) in lactic acid bacteria (LAB)-inoculated (n = 3) and non-inoculated (control) samples (n = 6) in vacuum-packed salmon portions on days 8 and 22. Group Day after packaging IMP Ino Hx H-value Ki-value (µmol/g) (µmol/g) (µmol/g) (%) (%) Control 8 0.8 ± 0.6 3.1 ± 0.6 3.5 ± 0.6 47 ± 12 90 ± 7 C.m.35 8 0.9 ± 0.9 4.1 ± 0.6 3.1 ± 0.2 38 ± 6 89 ± 9 C.m.55 8 1.7 ± 1.3 3.3 ± 0.6 2.7 ± 0.7 37 ± 16 80 ± 15 Le.g.406 8 1.5 ± 0.3 4.2 ± 0.1 2.8 ± 0.3 33 ± 3 82 ± 3 C.d.468 8 0.6 ± 0.2 3.1 ± 0.4 3.8 ± 0.8 51 ± 9 93 ± 2 p-value p = 0.28 p = 0.05 p = 0.17 p = 0.20 p = 0.28 Control 22 ND 1.4 ± 1.1 5.2 ± 1.2 78 ± 14 99.9 ± 0.2 C.m.35 22 ND 0.7 ± 0.4 6.7 ± 0.7 90 ± 6 99.8 ± 0.2 C.m.55 22 ND 0.7 ± 0.3 6.7 ± 0.3 90 ± 4 100 ± 0.0 Le.g.406 22 ND 0.3 ± 0.2 6.4 ± 0.3 95 ± 3 100 ± 0.0 C.d.468 22 ND 1.7 ± 0.2 5.4 ± 0.5 75 ± 4 100 ± 0.0 p-value - p = 0.10 p = 0.07 p = 0.07 p = 0.75 C.m.35: C. maltaromaticum 35; C.m.55: C. maltaromaticum 55; Le.g.406: L. gelidum 406; C.d.468: C. divergens 468. ND: Not detected 3.4. Biogenic amines The content of five BAs (cadaverine, spermidine, tryptamine, spermine, and tyramine) was quantified in VP fresh salmon portions on day 0 (prior to inoculation), 8 and 22. Low cadaverine levels were detected on day 8 (< 1 µg/g), and day 22 (0.2 ± 0.3–1.2 ± 0.1 µg/g) in all samples (Additional file 1). Spermidine (0.2 ± 0.3 µg/g) and tryptamine (0.4 ± 0.4 µg/g) were only detected in the samples inoculated with C.m.55 on day 8, while these BAs were not detected on day 22. The low quantities of BAs in these samples indicate that they maintained acceptable quality even after 22 days of storage. BAs are of concern either due to their toxicological effect on humans (Wójcik et al., 2021 ) and/or their correlation with spoilage by producing off-odours (Katikou et al., 2006 ). The present results showed no effect of the inoculated LAB on BA formation, which is essential for their potential industrial implementation. Although low quantities of spermidine and tryptamine were observed in samples inoculated with C.m.55, this should be taken with caution when considering this strain for biopreservation of processed salmon products or products within favourable conditions for decarboxylation activity (Biji et al., 2016 ). Moreover, the study of Özogul and Hamed ( 2018 ) emphasised that LAB can inhibit the growth of foodborne bacteria and thereby prevent their BA formation, however, other LAB can stimulate the BA production of foodborne pathogens. Thus, as the effect is strain and food matrix-dependent (Özogul & Hamed, 2018 ), it must be investigated as a criterion for strain selection for biopreservative purposes. 3.5. LAB effect on physiochemical properties of VP fresh salmon portions Changes in WC, DL and pH during the storage period at 4℃ are presented in Fig. 3 A, 3 B, and 3 C. No significant effect of inoculated LAB on WC, DL and pH was observed during the storage period in fresh VP salmon portions (one-way ANOVA, p < 0.05). Colour and textural changes in VP fresh salmon portions were analysed on days 8 and 22. No significant differences were found between LAB-inoculated and corresponding control samples in colour parameters (lightness (L*), redness (a*), yellowness (b*), chroma (C*), and total colour difference (ΔE)) (Table 4 ), breaking force and firmness (F60%) during the storage period (Additional file 1). The only difference was observed for the hue angle (h*) on day 22 (Table 4 ). Table 4 Differences in colour parameters (lightness (L*), redness (a*), yellowness (b*), chroma (C*), hue angle (h*) and total colour difference (ΔE)) in LAB-inoculated samples compared to the control at the same day, expressed as Δ-values (n = 3) ± SD. Data with * represents negative Δ-values. Group Day after packaging ΔL* Δa* Δb* Δc* Δh* Δ(ΔE) C.m.35 8 *3.6 ± 1.6 3.3 ± 1.0 3.2 ± 0.7 4.7 ± 1.2 *0.0 ± 0.3 1.4 ± 0.9 C.m.55 8 *2.2 ± 3.0 2.3 ± 3.3 2.4 ± 4.4 3.3 ± 5.4 0.1 ± 1.2 2.1 ± 1.9 Le.g.406 8 *3.6 ± 3.0 2.6 ± 3.6 2.7 ± 5.1 3.7 ± 6.1 *0.0 ± 1.3 0.8 ± 2.0 C.d.468 8 1.3 ± 3.8 *1.6 ± 4.7 *0.9 ± 5.0 *1.8 ± 6.9 0.6 ± 0.2 0.3 ± 2.3 p-value p = 0.22 p = 0.36 p = 0.63 p = 0.50 p = 0.78 p = 0.69 C.m.35 22 1.6 ± 0.8 *1.7 ± 0.6 2.6 ± 1.6 0.8 ± 1.3 8.2 ± 2.6 b *8.9 ± 6.2 C.m.55 22 1.3 ± 0.5 *1.7 ± 1.0 1.9 ± 1.1 0.2 ± 1.4 7.3 ± 0.9 b *9.5 ± 7.6 Le.g.406 22 1.2 ± 0.4 1.0 ± 3.6 0.9 ± 4.7 1.3 ± 5.8 *0.1 ± 1.5 a *0.1 ± 1.4 C.d.468 22 2.9 ± 5.1 *1.8 ± 1.9 *0.9 ± 1.6 *1.9 ± 2.4 0.7 ± 0.7 a *0.2 ± 1.8 p-value p = 0.86 p = 0.36 p = 0.46 p = 0.66 p < 0.05 p = 0.08 Small letters (ab) indicate significant differences between groups on the same day with the corresponding p-value at the bottom of each column, calculated by one-way ANOVA (p < 0.05). C.m.35: C. maltaromaticum 35; C.m.55: C. maltaromaticum 55; Le.g.406: L. gelidum 406; C.d.468: C. divergens 468. Overall, the selected LAB strains are considered suitable candidates for use in fresh VP salmon portions due to their demonstrated antagonistic activity and ability to maintain product quality. Similar results are also confirmed in other food products. In the study of Danielski et al. ( 2020 ), C. maltaromaticum did not affect the physicochemical properties of cooked ham, while in the study of Boulares et al. ( 2017 ), reduced water- and lipid loss were achieved for sea bass inoculated with combinations of Lactococcus lactic, Lactobacillus plantarum ( Lactiplantibacillus plantarum ), and C. piscicola ( C. maltaromaticum ). In the study of Mozuriene et al. ( 2016 ), an improved effect on the colour and texture of pork meat was observed by lacto-fermented potato tuber juice treatment ( Pediococcus acidilactici, P. pentosaceus and Latilactobacillus sakei ). However, higher drip loss and lower water holding capacity were observed for LAB-treated samples compared to the control in the same study (Mozuriene et al., 2016 ). Overall, adding LAB to the VP fresh salmon portions did not affect any of the measured physiochemical parameters. As these parameters are tightly connected (e.g. LAB could provide a proteolytic effect by degrading muscle proteins as well as connective tissue, which in turn impacts the water-holding properties), preserving the physicochemical properties without significant changes in the pH of the fresh salmon is crucial for consumer acceptance. 3.6. Antilisterial effect of inoculated LAB strains The selected LAB strain's antibacterial activity in VP fresh salmon portions was examined. In the absence of inoculated LAB, L. innocua proliferated from an initial concentration of 3.4 ± 0.1 to 5.0 ± 0.3 log CFU/g during 21 days of storage at 4℃ (Table 5 ). However, in the presence of C.m.35, L. innocua was not able to proliferate in the product. On the last day of storage, there was no significant difference in the concentration of Listeria compared to the initial concentration (p > 0.05) (Table 5 ). In fact, significantly lower counts of L. innocua were found on day 14 (p = 0.001) and day 21 (p = 0.004) compared to samples inoculated with only L. innocua (Table 5 ). Although both C.m.35 and C.d.468 are recognised as producers of bacteriocin-like substances (BLS) (Stupar et al., 2023 ), the same inhibition of L. innocua was not observed for the C.d.468 or the two non-bacteriocin-producing strains (Table 5 ). Even though all LAB strains performed total inhibition of L. innocua in vitro (Stupar et al., 2023 ; Stupar et al., 2021 ), a lower effect was observed in salmon portions, indicating that food matrix, initial concentrations, and pH can govern inter-species competition (Bolívar et al., 2021 ; Mellefont et al., 2008 ). Noteworthy lower initial concentrations of LAB (~ 10 4 CFU/g) and higher target concentration (~ 10 3 CFU/g) were used in the present study compared to the in vitro studies in pH-adjusted salmon juice (Stupar et al., 2023 ; Stupar et al., 2021 ). Previously, it was shown that most strains use nutrient competition as the main mechanism against the target organism (Stupar et al., 2023 ), assuming the Jameson effect, where higher LAB concentrations play an essential role (Mellefont et al., 2008 ). Thus, a higher initial LAB concentration and/or a combination of several LAB strains might express more efficacy against undesirable microbiota. For example, in the study of Boulares et al. ( 2017 ), the highest reduction of L. monocytogenes was > 3 log CFU/g in fresh VP sea bass stored for 21 days at 4℃, achieved by combining two or three LAB strains at high initial concentrations (8 log CFU/g of each). Moreover, Lactobacillus casei demonstrated bacteriostatic and bactericidal effects when inoculated at concentrations of 6 and 8 log CFU/g, respectively, against L. innocua in VP CSS (Vescovo et al., 2006 ). In the present study, a lower initial LAB concentration was applied to reduce the risk of negative effects on physicochemical properties of fresh salmon, but at the same time aiming to achieve sufficient antimicrobial activity. Combined with other hurdle technologies, the applied approach might be even more powerful. Furthermore, the lack of antilisterial effect of the BLS-producing strain C.d.468 during storage (GLM, p = 0.599) might also be connected to non-optimal conditions for the growth and bacteriocin activity of the applied strain (Fig. 1 A). The observed bacteriostatic effect of strain C.m.35 is an effective hurdle for the multiplication of L. innocua . The present study simulated a worst-case scenario, given the unrealistically high initial target concentrations (Svanevik et al., 2021) and the application of a target strain demonstrably more resistant to biopreservation than L. monocytogenes (Stupar et al., 2021 ). Thus, given the expected low contamination level of L. monocytogenes in fresh salmon (Svanevik et al., 2021), inhibition of 1–2 log CFU/g would be sufficient to keep the level of this pathogen within the safe zone during refrigerated storage. On the other hand, more studies, including a range of L. monocytogenes strains from various serotypes regularly detected in the salmon product and /or the salmon processing environment, should be conducted to give a broader insight into the LAB strain's antilisterial effect and their ability to act as a protective hurdle in RTE salmon products. Despite the QPS status of C. maltaromaticum , C. divergens and Leuconostoc spp. (EFSA BIOHAZ Panel, 2021) the strains should be further safety tested before any industrial application. In particular, safety documentation for food applications of L. gelidum strains is deficient. The application of protective cultures in raw salmon products will probably be defined as “novel food” in the EU and must be approved by food safety authorities before industrial applications. Although no negative effect of LAB inoculation was observed on quality parameters in the current study, the sensory aspects must be further elucidated using a sensory panel, and a consumer acceptance analysis must be conducted before industrial applications. Table 5 Proliferation of Listeria innocua (Target) in vacuum-packed salmon portions in the presence of selected LAB for 21 days at 4℃. The results are presented as mean log CFU/g ± SD (n = 3 for target in coculture with LAB and n = 6 for target-monoculture). Group Day after packaging p-value 0 7 14 21 C.m.35 + T 3.3 ± 0.1 Ib 3.4 ± 0.3 Ia 3.4 ± 0.4 Ia 4.0 ± 0.5 Ia = 0.19 C.m.55 + T 3.0 ± 0.0 Ia 3.7 ± 0.3 IIab 4.1 ± 0.3 II,IIIab 4.4 ± 0.2 IIIab < 0.05 Le.g.406 + T 3.4 ± 0.0 Ib 4.0 ± 0.3 I,IIab 4.1 ± 0.4 II, IIIab 4.7 ± 0.1 IIIab < 0.05 C.d.468 + T 3.3 ± 0.2 Ib 4.3 ± 0.4 IIb 4.6 ± 0.0 II,IIIb 5.0 ± 0.1 IIIb < 0.05 Target 3.4 ± 0.1 Ib 4.0 ± 0.2 IIab 4.7 ± 0.3 IIIb 5.0 ± 0.3 IIIb < 0.05 p-value < 0.05 < 0.05 < 0.05 < 0.05 Small letters ( ab ) indicate significant differences between the groups on the same day, with the corresponding p-value at the bottom of each column. Roman numbers ( I, II, III ) indicate significant differences in a group at all tested days, with the corresponding p-value in the same row. Significant differences are calculated by one-way ANOVA, Tukey HSD, p < 0.05. C.m.35: C. maltaromaticum 35; C.m.55: C. maltaromaticum 55; Le.g.406: L. gelidum 406; C.d.468: C. divergens 468. T = Target; Target = L. innocua Conclusion In the present study, four LAB strains were selected for biopreservation of VP fresh salmon portions. The LAB strains demonstrated good adaptation to the salmon matrix and did not negatively affect the physicochemical and chemical properties of VP salmon portions at 4℃ for 22 days of storage. Moreover, the selected LAB strains are considered safe in terms of no significant production of biogenic amines. The strain C. maltaromaticum 35 (C.m.35) demonstrated a bacteriostatic effect on L. innocua , which makes it an interesting candidate for controlling L. monocytogenes in ready-to-eat salmon. In a future perspective, the effect of this strain could be optimised by combining it with other LAB strains or applying additional shelf-life-improving strategies. Declarations Authors’ contributions Jelena Stupar: Formal analysis, Investigation, Data curation, Methodology, Visualization, Writing-Original Draft, Writing-review and editing; Sunniva Hoel: Supervision, Conceptualization, Methodology, Visualization, Writing-Review & Editing; Jørgen Lerfall: Supervision, Conceptualization, Methodology, Visualization, Writing-Review & Editing; Turid Rustad: Supervision, Conceptualization, Methodology, Visualization, Writing-Review & Editing; Anita Nordeng Jakobsen: Supervision, Conceptualization, Methodology, Visualization, Writing-Review & Editing, Project Administration. Acknowledgements The authors would like to thank the technical staff at the Department of Biotechnology and Food Science at NTNU for practical help in the microbiological, analytical and food processing laboratory. Ethics declarations Not applicable. Ethics approval and consent to participate Not applicable. 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Supplementary Files Additionalfile1.docx Additional files File name: Additional file 1.doc Title of data: Content of biogenic amines in vacuum-packed salmon portions stored at 4℃ Description: Content of biogenic amines in experimental groups evaluated on day 8 nd 22 of cold storage. Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. 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07:07:11","extension":"html","order_by":43,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":259952,"visible":true,"origin":"","legend":"","description":"","filename":"earlyproof.html","url":"https://assets-eu.researchsquare.com/files/rs-7968166/v1/630ba1c67334d7c1b5c2108d.html"},{"id":97765402,"identity":"a44b1482-6c47-460c-9c77-3f45cdbd6e1d","added_by":"auto","created_at":"2025-12-09 07:07:10","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":151081,"visible":true,"origin":"","legend":"\u003cp\u003eSee image above for figure legend\u003c/p\u003e","description":"","filename":"Figure1A.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7968166/v1/be3cd4accd80f3d4be868ab3.jpg"},{"id":97897498,"identity":"dbed3c7d-116e-4c4d-80aa-b1c4bcaab72f","added_by":"auto","created_at":"2025-12-10 15:37:53","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":35968,"visible":true,"origin":"","legend":"\u003cp\u003eGenus relative abundance in LAB-inoculated and non-inoculated control samples on day 14. \u003cem\u003eC. maltaromaticum \u003c/em\u003e35 (C.m.35), \u003cem\u003eC. maltaromaticum \u003c/em\u003e55 (C.m.55), \u003cem\u003eL. gelidum \u003c/em\u003e406 (Le.g.406), \u003cem\u003eC. divergens \u003c/em\u003e468 (C.d.468), Control 1 corresponds to experiments with C.m.55 and Control 2 to experiments with C.m.35, Le.g.406, and C.d.468 (n=3). \u0026nbsp;All genera with \u0026gt;1% relative abundance are shown, while less than 1% are assembled in the category ‘‘other’’.\u003c/p\u003e","description":"","filename":"Figure2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7968166/v1/0924e423b97c6498cf5e5962.jpg"},{"id":97810211,"identity":"b7f0ff76-482c-4e9e-b0f3-cf87db85073c","added_by":"auto","created_at":"2025-12-09 15:41:24","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":35968,"visible":true,"origin":"","legend":"\u003cp\u003eGenus relative abundance in LAB-inoculated and non-inoculated control samples on day 14. \u003cem\u003eC. maltaromaticum \u003c/em\u003e35 (C.m.35), \u003cem\u003eC. maltaromaticum \u003c/em\u003e55 (C.m.55), \u003cem\u003eL. gelidum \u003c/em\u003e406 (Le.g.406), \u003cem\u003eC. divergens \u003c/em\u003e468 (C.d.468), Control 1 corresponds to experiments with C.m.55 and Control 2 to experiments with C.m.35, Le.g.406, and C.d.468 (n=3). \u0026nbsp;All genera with \u0026gt;1% relative abundance are shown, while less than 1% are assembled in the category ‘‘other’’.\u003c/p\u003e","description":"","filename":"Figure2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7968166/v1/36e133fbf328ddada846f0c6.jpg"},{"id":97897191,"identity":"5d7c11ad-b1ac-47c8-8df5-69863b4d0a53","added_by":"auto","created_at":"2025-12-10 15:37:33","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":109166,"visible":true,"origin":"","legend":"\u003cp\u003eSee image above for figure legend\u003c/p\u003e","description":"","filename":"Figure3A.jpg","url":"https://assets-eu.researchsquare.com/files/rs-7968166/v1/b94a95e24a8d3510f9e852c8.jpg"},{"id":97810302,"identity":"afd10cce-bc34-42be-9ff4-f215ff8dc588","added_by":"auto","created_at":"2025-12-09 15:44:18","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":370079,"visible":true,"origin":"","legend":"\u003cp\u003eSee image above for figure legend\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-7968166/v1/26faf9f29b1d71f30625e723.png"},{"id":103129541,"identity":"212d5599-71f0-4872-8cbb-c89b649335cc","added_by":"auto","created_at":"2026-02-21 12:40:45","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2179798,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7968166/v1/79f894d9-290f-4663-af0d-9ce5aa1f6173.pdf"},{"id":97896405,"identity":"b552db02-4dca-4209-aa3e-78f3196ad7cd","added_by":"auto","created_at":"2025-12-10 15:36:29","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":29146,"visible":true,"origin":"","legend":"\u003cp\u003eAdditional files\u003c/p\u003e\n\u003cp\u003eFile name: Additional file 1.doc\u003c/p\u003e\n\u003cp\u003eTitle of data: Content of biogenic amines in vacuum-packed salmon portions stored at 4℃\u003c/p\u003e\n\u003cp\u003eDescription: Content of biogenic amines in experimental groups evaluated on day 8 nd 22 of cold storage.\u003c/p\u003e","description":"","filename":"Additionalfile1.docx","url":"https://assets-eu.researchsquare.com/files/rs-7968166/v1/a26b9eb5b96e1e289c316304.docx"}],"financialInterests":"\u003cp\u003eNo competing interests reported.\u003c/p\u003e\n\u003cp\u003eAdditional file 1 not available with this version.\u003c/p\u003e","formattedTitle":"Biopreservation of ready-to-eat Atlantic salmon (Salmo salar) by lactic acid bacteria: Effect on safety and quality parameters","fulltext":[{"header":"1. Introduction","content":"\u003cp\u003e\u003cem\u003ePre-rigor\u003c/em\u003e filleted vacuum-packed (VP) fresh salmon loins are nutritious and appealing to consumers as ready-to-eat (RTE) products. However, the production lacks efficient barriers to control the growth of the pathogenic bacteria \u003cem\u003eListeria monocytogenes\u003c/em\u003e, recognised as the causative agent of listeriosis. Although the general contamination level of \u003cem\u003eListeria\u003c/em\u003e spp. in fresh salmon is low (Noseda et al., \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Svanevik et al., 2021), their ability to persist in biofilms in food production environments (Fagerlund et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and their ability to tolerate stressful conditions applied in food processing and packaging (Chan \u0026amp; Wiedmann, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Noseda et al., \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Tuytschaever et al., \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) challenge the safety of RTE salmon products. Furthermore, fresh seafood is highly perishable (Leroi, \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2010\u003c/span\u003e), and the growth of spoilage bacteria belonging to genera of e.g. \u003cem\u003ePhotobacterium., Brochothrix\u003c/em\u003e and \u003cem\u003ePseudomonas\u003c/em\u003e can result in the production of total volatile basic nitrogen (TVB-N), sulphur compounds, biogenic amines (BA) and other spoilage products (Rathod et al., \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) lowering the product quality.\u003c/p\u003e\u003cp\u003eBiopreservation using lactic acid bacteria (LAB) represents a natural preservation strategy to ensure product safety and microbial stability with minimal effect on physicochemical and nutritional properties (Barcenilla et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Strack et al., \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Food-fermenting LAB, including members of \u003cem\u003eCarnobacterium\u003c/em\u003e and \u003cem\u003eLeuconostoc\u003c/em\u003e genera, are \u0026lsquo;Generally Recognised as Safe\u0026rsquo; (GRAS) and are given a Qualified Presumption of Safety (QPS)\u0026rsquo; status by the Food and Drug Administration (FDA) and European Food Safety Agency (EFSA) (EFSA BIOHAZ Panel, 2021). This has made them interesting candidates for various applications. However, studies also link LAB to the production of BA and spoilage metabolites (Emborg et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Wiernasz et al., \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Yazgan et al., \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), which calls for diligent strain selection.\u003c/p\u003e\u003cp\u003eLAB, including the genera \u003cem\u003eCarnobacterium\u003c/em\u003e and \u003cem\u003eLeuconostoc\u003c/em\u003e, have been widely investigated for biopreservative purposes of various foods (Angiolillo et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Jang et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Mei et al., \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Ramos et al., \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Saraoui et al., \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Shi \u0026amp; Maktabdar, \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Stupar et al., \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Tahiri et al., \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Wiernasz et al., \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Wong \u0026amp; Li, \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Woraprayote et al., \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). However, their implementation in fresh fish remains relatively unexplored (Evangelista et al., \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; G\u0026oacute;mez-Sala et al., \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Although underexplored, species from the \u003cem\u003eCarnobacterium\u003c/em\u003e and \u003cem\u003eLeuconostoc\u003c/em\u003e genus have demonstrated the ability to inhibit the growth of several spoilage and pathogenic bacteria by the production of antimicrobial compounds such as organic acids, hydrogen-peroxide, bacteriocins (Begrem et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Rao et al., \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Stupar et al., \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), and competition for nutrients and attachment sites (Nilsson et al., \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Stupar et al., \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Wiernasz et al., \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eFinding suitable LAB for the biopreservation of fresh salmon is challenging due to the clean sensory profile, short shelf life and low storage temperature of the product. LAB strains selected in the present study have shown inhibitory properties against strains of \u003cem\u003eL. monocytogenes\u003c/em\u003e and \u003cem\u003eL. innocua in vitro\u003c/em\u003e (Stupar et al., \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and have demonstrated suitable growth properties in fresh VP salmon at 4℃ (Stupar et al., \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The aim of the present study was to apply a polyphasic approach combining microbiological, chemical, and physicochemical factors to investigate the effect of selected \u003cem\u003eCarnobacterium\u003c/em\u003e and \u003cem\u003eLeuconostoc\u003c/em\u003e strains on safety and quality parameters in VP salmon portions stored at 4℃ for 22 days. Inoculation trials with non-pathogenic \u003cem\u003eListeria innocua\u003c/em\u003e (CCUG 15531) were applied, supported by previous data confirming the applicability of this strain as a substitute for \u003cem\u003eL. monocytogenes\u003c/em\u003e (Stupar et al., \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). This is, to the best of our knowledge, the first study exploring the effect of LAB on the safety, chemical and physiochemical properties of RTE VP salmon portions.\u003c/p\u003e"},{"header":"2. Materials and methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e\u003ch2\u003e2.1. Experimental design\u003c/h2\u003e\u003cp\u003eThe experiment aimed to evaluate the impact of LAB inoculation on VP salmon portions with respect to microbial, chemical and physicochemical quality, and safety parameters. Experimental groups (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e) included: i) control - non-inoculated salmon portions; ii) portions inoculated with a single LAB strain (\u003cem\u003eC. maltaromaticum\u003c/em\u003e 35 (C.m.35), \u003cem\u003eC. maltaromaticum\u003c/em\u003e 55 (C.m.55), \u003cem\u003eL. gelidum\u003c/em\u003e 406 (Le.g.406) or \u003cem\u003eC. divergens\u003c/em\u003e 468 (C.d.468)); iii) portions inoculated with the target organisms (\u003cem\u003eL. innocua\u003c/em\u003e); and iv) portions inoculated with both LAB and the target. Two independent batches of pre-rigor filleted salmon (Batch I and Batch II, slaughtered on different dates, Section \u003cspan refid=\"Sec7\" class=\"InternalRef\"\u003e2.2.3\u003c/span\u003e) were used: Batch I for experiments with C.m.55, and Batch II for C.m.35, C.d.468, and Le.g.406. All samples were vacuum-packed and stored at 4\u0026deg;C for 22 days. Total aerobic plate counts (APC), H\u003csub\u003e2\u003c/sub\u003eS-producing bacteria, LAB, and \u003cem\u003eL. innocua\u003c/em\u003e were quantified on days 0, 7, 14 and 21 of storage (Section \u003cspan refid=\"Sec8\" class=\"InternalRef\"\u003e2.2.4\u003c/span\u003e). Microbial community composition was assessed at day 14 of storage (Section \u003cspan refid=\"Sec9\" class=\"InternalRef\"\u003e2.3\u003c/span\u003e). Chemical and physicochemical analyses were performed on day 0 (pre-inoculation), day 8 and day 22 post-packaging. Samples for chemical analysis were stored at -80℃ until further processing. Salmon portions inoculated with \u003cem\u003eListeria\u003c/em\u003e were not subjected to physicochemical, chemical or microbial community analysis to prevent cross-contamination in the food processing laboratory.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eExperimental design showing inoculation and analysis of vacuum-packed salmon portions investigated. All samples were stored at 4\u0026deg;C for 22 days.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"8\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u003cp\u003eInoculation\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eVacuum packed salmon\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"2\" nameend=\"c6\" namest=\"c5\"\u003e\u003cp\u003eSampling days for microbiological analyses\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003eSampling days for chemical analyses\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c8\"\u003e\u003cp\u003eSampling days for physicochemical analyses\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eGroups\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eLAB\u003c/p\u003e\u003cp\u003e(10\u003csup\u003e4\u003c/sup\u003e CFU/g)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eTarget\u003c/p\u003e\u003cp\u003e(10\u003csup\u003e3\u003c/sup\u003e CFU/g)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eSample size\u003c/p\u003e\u003cp\u003e(g)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eCulture-dependent\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eCulture-independent\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003eATP-degradation products, biogenic amines\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c8\"\u003e\u003cp\u003eWater content,\u003c/p\u003e\u003cp\u003epH, colour,\u003c/p\u003e\u003cp\u003etexture\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eControl\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e\u003cp\u003e100\u0026thinsp;\u0026plusmn;\u0026thinsp;1, 10\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0,7, 14,21\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e14\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0, 8, 22\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0, 8, 22\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eLAB*\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eC.m.35 C.m.55\u003c/p\u003e\u003cp\u003eLe.g.406 C.d.468\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e\u003cp\u003e100\u0026thinsp;\u0026plusmn;\u0026thinsp;1, 10\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0,7, 14,21\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e14\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0, 8, 22\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0, 8, 22\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eLAB*+T\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003eC.m.35 C.m.55\u003c/p\u003e\u003cp\u003eLe.g.406 C.d.468\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cem\u003eL.innocua\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e\u003cp\u003e10\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0,7, 14,21\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eT\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cem\u003eL.innocua\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e\u003cp\u003e10\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0,7, 14,21\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eLAB\u0026thinsp;=\u0026thinsp;Lactic acid bacteria; T\u0026thinsp;=\u0026thinsp;Target (\u003cem\u003eL. innocua\u003c/em\u003e); *LAB inoculation with a single culture of \u003cem\u003eCarnobactrium maltaromaticum 35\u003c/em\u003e, \u003cem\u003eCarnobacterium maltaromaticum 55\u003c/em\u003e, \u003cem\u003eLeuconostoc gelidum\u003c/em\u003e 406 or \u003cem\u003eCarnobacterium divergens\u003c/em\u003e 468. Culture-dependent analysis included analysis of aerobic plate count, H\u003csub\u003e2\u003c/sub\u003eS-producing bacteria, lactic acid bacteria and \u003cem\u003eListeria\u003c/em\u003e spp. Culture- independent analysis included analysis of the microbial community composition.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec4\" class=\"Section2\"\u003e\u003ch2\u003e2.2. Biopreservation of fresh salmon vacuum-packed fillets\u003c/h2\u003e\u003cdiv id=\"Sec5\" class=\"Section3\"\u003e\u003ch2\u003e2.2.1. Selection and preparation of LAB strains\u003c/h2\u003e\u003cp\u003eLAB strains were pre-cultured in De Man, Rogosa and Sharpe (MRS) (Oxoid, Norway) for 2.5 days at 25℃ under anaerobic conditions using GasPak EZ Anaerobe container system sachets with indicator (BD, Norway). A single colony was transferred to MRS broth (Oxoid) and incubated for 2.5 days at 8℃ to promote cold adaptation. Following incubation, cultures were standardised to an optical density (OD) of 0.22 at 600 nm (Shimadzu UV 1800, Germany), corresponding to approximately 10\u003csup\u003e8\u003c/sup\u003e CFU/mL (Stupar et al., \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). For inoculation, 0.1 or 1 mL of the diluted cold-adapted cultures was applied to the surface of each salmon portion to achieve a concentration of approximately 10\u003csup\u003e4\u003c/sup\u003e CFU/g.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec6\" class=\"Section3\"\u003e\u003ch2\u003e2.2.2. Preparation of target strain\u003c/h2\u003e\u003cp\u003e\u003cem\u003eL. innocua\u003c/em\u003e was cultivated Brain Heart Infusion (BHI) agar (Oxoid) for 24h at 37℃. A single colony was picked and incubated in BHI broth (Oxoid) for 24h at 15℃ to promote adaptation to low temperature (Stupar et al., \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The resulting culture was adjusted to an OD₆₀₀ of 0.1 (Shimadzu UV 1800; Shimadzu, Germany). An aliquot of 0.1 mL was applied to the surface of each salmon portion (Section \u003cspan refid=\"Sec8\" class=\"InternalRef\"\u003e2.2.4\u003c/span\u003e), corresponding to an inoculation concentration of approximately 10\u003csup\u003e3\u003c/sup\u003e CFU/g.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec7\" class=\"Section3\"\u003e\u003ch2\u003e2.2.3. Sample preparation and packaging\u003c/h2\u003e\u003cp\u003eAtlantic salmon (\u003cem\u003eSalmo salar\u003c/em\u003e) was obtained from a local slaughterhouse and transported on ice to NTNU, where it was hand-filleted within 5 h post-mortem. Back loin portions were prepared by cutting into 10\u0026thinsp;\u0026plusmn;\u0026thinsp;1 g pieces for microbiological and chemical analysis, and into 100\u0026thinsp;\u0026plusmn;\u0026thinsp;1 g pieces for physicochemical analysis. All samples were stored on ice until processing the following day. Randomised pieces were placed on an absorbent pad (Tommen Gram, Norway) and inoculated with 1% (v/w) LAB and/or target on the salmon surface as described by Stupar et al. (\u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). All samples were packed in vacuum (20 \u0026micro;m polyamide (PA)/70-\u0026micro;m polyethylene (PE) bag (120 9 80 mm, Star-Pack Productive, Boissy-'Aillerie, France) with a Webomatic Supermax- C vacuum machine (Webomatic, Germany) and stored at 4℃ for 21/22 days (21 days for microbiological and 22 days for chemical and physicochemical analyses). The 21/22 storage period was chosen to follow the microbial growth and potential re-growth of the target strain beyond the shelf life of commercial products (approximately 10 days).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec8\" class=\"Section3\"\u003e\u003ch2\u003e2.2.4. Microbiological analysis\u003c/h2\u003e\u003cp\u003eThe salmon sample (10 g) was homogenised with 90 g of peptone water (1 g/L peptone (Oxoid) and 8.5 g/L NaCl (VWR, Belgium)), using a stomacher (IUL Masticator, Spain). Homogenates were serially diluted and plated on media. Lyngby's Iron agar (IA; Oxoid) supplemented with 0.04% L-cysteine (Sigma-Aldrich, Norway) was used to quantify APC and H\u003csub\u003e2\u003c/sub\u003eS-producing bacteria by incubation for 72\u0026thinsp;\u0026plusmn;\u0026thinsp;6h at 22℃. LABs were enumerated on MRS agar using anaerobic incubation for 2\u0026ndash;5 days at 25℃. \u003cem\u003eL. innocua\u003c/em\u003e was quantified on Brilliance Listeria agar (BLA; Oxoid), supplemented with Brilliance Listeria supplement (Oxoid), incubated at 37\u0026deg;C for 24\u0026thinsp;\u0026plusmn;\u0026thinsp;1 h.\u003c/p\u003e\u003c/div\u003e\u003c/div\u003e\u003cdiv id=\"Sec9\" class=\"Section2\"\u003e\u003ch2\u003e2.3. Culture-independent microbial community analysis\u003c/h2\u003e\u003cp\u003eTotal genomic DNA was extracted from homogenates (1:10 in sterile peptone water; n\u0026thinsp;=\u0026thinsp;3) collected on day 14. A sample of 5 ml was centrifuged (83 x g, 5 min, \u0026sim; 20℃) to remove most of the fish particles. The supernatant was again centrifuged (2061 x g, 15 min, \u0026sim;20℃), and the pellet was resuspended in 1 ml of peptone water before a final centrifugation(6160 x g, 5 min, \u0026sim;20℃). The pellets were used for DNA extraction using the PowerFood\u0026reg; DNA isolation kit (MoBio Laboratories Inc., Carlsbad, CA, USA). Quantitative and qualitative analysis of the extracted DNA was assessed by spectrophotometry (PowerWave XS, BioTek\u0026reg;\u003csup\u003e)\u003c/sup\u003e and 1% gel electrophoresis. The V1-V3 hypervariable region of the 16S rRNA gene (~\u0026thinsp;490 bp) was amplified using primers 16S_f (5\u0026prime; AGAGTTTGATCATGGCTCAG 3\u0026prime;) and 16S_r (5\u0026prime; GTATTACCGCGGCTGCTG 3\u0026prime;) (Leser et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Weisburg et al., \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e1991\u003c/span\u003e). Amplicon sequencing and microbial community analysis were performed using the INVIEW Microbiome Profiling 3.0 package (Eurofins Genomics, Germany) on an Illumina MiSeq, including amplicon library generation and standardised bioinformatic, as described by Tsoukalas et al. (\u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec10\" class=\"Section2\"\u003e\u003ch2\u003e2.4. Chemical analysis\u003c/h2\u003e\u003cdiv id=\"Sec11\" class=\"Section3\"\u003e\u003ch2\u003e2.4.1. Extraction procedure\u003c/h2\u003e\u003cp\u003eApproximately 2 g of frozen sample was grated using a stainless-steel grater and homogenised for 2 min with an Ultra Turrax T25 Basic (Janke \u0026amp; Kunkel IKA\u0026reg;-Labortechnik, Staufen, Germany) in 10 mL 7% trichloroacetic acid (TCA). Subsequently, 1.5 ml of 1 mol/l potassium hydroxide was added, vortexed, and centrifuged (16,000\u0026times;g, 10 min, 4\u0026deg;C). The resulting supernatants were filtered through a 0.45 \u0026micro;m nylon filter; one portion was used for BA analysis, and the remaining portion was used for analysis of ATP-degradation products (Section \u003cspan refid=\"Sec12\" class=\"InternalRef\"\u003e2.4.2\u003c/span\u003e).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec12\" class=\"Section3\"\u003e\u003ch2\u003e2.4.2. Analysis of ATP-degradation products\u003c/h2\u003e\u003cp\u003eATP-degradation products were analysed in samples collected on day 0 (prior to inoculation) and on days 8 and 22 post-packaging. Quantification was performed according to the method described by Lerfall et al. (\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), with a detection limit of 5 \u0026micro;mol/L. Ki-value (Karube et al., \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e1984\u003c/span\u003e)d value (Luong et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e1992\u003c/span\u003e) were calculated by using the following equations:\u003c/p\u003e\u003cp\u003eH-value (%) = [Hx / (IMP\u0026thinsp;+\u0026thinsp;HxR\u0026thinsp;+\u0026thinsp;Hx)] \u0026times; 100, (1)\u003c/p\u003e\u003cp\u003eKi-value (%) = [(HxR\u0026thinsp;+\u0026thinsp;Hx) / (IMP\u0026thinsp;+\u0026thinsp;HxR\u0026thinsp;+\u0026thinsp;Hx)] \u0026times; 100 (2)\u003c/p\u003e\u003cp\u003ewhere HxR\u0026thinsp;=\u0026thinsp;Inosine, Hx\u0026thinsp;=\u0026thinsp;Hypoxanthine, and IMP\u0026thinsp;=\u0026thinsp;Inosine monophosphate.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec13\" class=\"Section3\"\u003e\u003ch2\u003e2.4.3. Biogenic amines\u003c/h2\u003e\u003cp\u003eFish extract for BA determination was prepared as described in Section \u003cspan refid=\"Sec11\" class=\"InternalRef\"\u003e2.4.1\u003c/span\u003e. The extract was neutralised by potassium hydroxide (1M KOH), derivatised with benzyl chloride (99%, Sigma-Aldrich, CAS:98-88-4) for 20 min at room temperature. Benzylated amines were extracted twice with diethyl ether, and the combined organic phase was evaporated to dryness under nitrogen at 30\u0026deg;C. The residue was reconstituted in acetonitrile\u0026ndash;water (90:10). BA quantification was performed by UHPLC as described by Lerfall et al. (\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), with a detection level of 0.05 \u0026micro;mol/L, using a YMC-Triart PFP (100 x 2 mm, 1.9 \u0026micro;m) column connected to an Agilent 1290 chromatography (Agilent Technologies, Paolo Alto, CA, USA) and an Agilent 1260 diode array UV-Vis detector. The loaded samples were detected at 254 nm using H\u003csub\u003e2\u003c/sub\u003eO/acetonitrile-based elution.\u003c/p\u003e\u003c/div\u003e\u003c/div\u003e\u003cdiv id=\"Sec14\" class=\"Section2\"\u003e\u003ch2\u003e2.5. Physicochemical parameters\u003c/h2\u003e\u003cp\u003eAll analyses (water content, drip loss, muscle pH, texture and colour analysis) were performed in triplicate on salmon pieces (100\u0026thinsp;\u0026plusmn;\u0026thinsp;1g) from both LAB-inoculated and non-inoculated controls. Selected analyses were performed on days 0 (prior to inoculation), day 8 and day 22.\u003c/p\u003e\u003cdiv id=\"Sec15\" class=\"Section3\"\u003e\u003ch2\u003e2.5.1. Water content and muscle pH\u003c/h2\u003e\u003cp\u003eWater content (WC) was determined according to Skipnes et al. (\u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). Standardised muscle pieces were prepared by pressing the fish muscle with a metal cylinder (diameter 31 mm) to obtaine pieces 6 mm in height. Each piece was transversally divided into two equal parts, and the bottom piece was used for WC analysis. Samples were weighed and dried for 16h at 105℃. Drip loss (DL) was calculated as the percentage difference (%) of the weighed sample (g) to its initial weight (g). Muscle pH was measured by a portable pH meter (Hach HQ40d multi-Portable Meter, Hach, USA) with a puncture pH electrode (Hach Intellical\u0026trade; PHC108, Hach, CO, USA).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec16\" class=\"Section3\"\u003e\u003ch2\u003e2.5.2. Texture analysis\u003c/h2\u003e\u003cp\u003eBreaking force was measured using a Texture Analyser TA-XT plus (Stable Micro Systems Ltd, England) equipped with a flat-ended cylindrical probe (12.7 mm P/0.5). Breaking force was defined as the force (N) required to rupture the fillet surface, while firmness was recorded as the force at 60% compression. Data acquisition and analysis were performed using the Texture Exponent light software for Windows (version 4.12, SMS).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec17\" class=\"Section3\"\u003e\u003ch2\u003e2.5.3. Colour analysis\u003c/h2\u003e\u003cp\u003eColor measurements of selected samples were performed using a DigiEye full system (VeriVide Ltd., Leicester, UK) equipped with a digital camera (Nikon D80, 35 mm lens, Nikon Corp., Japan). Colour parameters (L* = lightness, a*= redness, b*= yellowness, C*= Chroma and h* = hue angle) were obtained using software DigiPix (version 2.8). Total colour difference (ΔE) was calculated based on CIE (1994). Chroma (C*) and hue angle (h*) were calculated by the following Equations:\u003c/p\u003e\u003cp\u003eC* = (a*\u003csup\u003e2\u003c/sup\u003e + b*\u003csup\u003e2\u003c/sup\u003e)\u003csup\u003e1/2\u003c/sup\u003e (6)\u003c/p\u003e\u003cp\u003eh* = arctan (b*/a*) (7)\u003c/p\u003e\u003cp\u003eColour changes (Δ-values) were calculated by the following Equation:\u003cdiv class=\"BlockQuote\"\u003e\u003cp\u003eΔ colour parameter at day x\u0026thinsp;=\u0026thinsp;Colour average measurement of LAB-inoculated sample - Colour average measurement of control sample (8)\u003c/p\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003ewhere Δcolour parameter (L*, a*, b*, C*, h*, ΔE) (n\u0026thinsp;=\u0026thinsp;3) at the same storage day.\u003c/p\u003e\u003c/div\u003e\u003c/div\u003e\u003cdiv id=\"Sec18\" class=\"Section2\"\u003e\u003ch2\u003e2.6. Statistical analysis\u003c/h2\u003e\u003cp\u003eStatistical analyses were conducted using IBM SPSS Statistics software (version 28, IBM Corporation, USA). Microbial counts were log-transformed and expressed as mean\u0026thinsp;\u0026plusmn;\u0026thinsp;standard deviation (SD). Log-transformed bacterial counts were fitted to the Baranyi and Roberts (\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1994\u003c/span\u003e) primary growth model using the DMFit program (\u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003e\u003ca href=\"http://www.combase.cc\" target=\"_blank\"\u003ewww.combase.cc\u003c/a\u003e\u003c/span\u003e\u003cspan address=\"http://www.combase.cc\" targettype=\"URL\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e). Differences between experimental groups were assessed by using one-way ANOVA with Tukey's pairwise comparisons and Independent Samples Test at a 5% significance level (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05). A general linear model (GLM) was applied to evaluate the antilisterial effect of LAB during storage and the influence of batch variations on target strain growth (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05). Pearson\u0026rsquo;s correlation was used to examine the effect of storage time on Ki-values. All experimental groups were analysed in triplicate, and results are presented as mean values\u0026thinsp;\u0026plusmn;\u0026thinsp;standard deviation (SD). As no significant differences (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05) in bacterial counts were observed between Batch I and Batch II for both the Control group (non-inoculated samples) and the Target group (samples inoculated with \u003cem\u003eL. innocua\u003c/em\u003e, positive control), data from both batches were combined and reported as mean values (n\u0026thinsp;=\u0026thinsp;6)\u0026thinsp;\u0026plusmn;\u0026thinsp;SD. The only exception is for the culture-independent analysis, where the controls from different batches were presented separately (n\u0026thinsp;=\u0026thinsp;3\u0026thinsp;\u0026plusmn;\u0026thinsp;SD).\u003c/p\u003e\u003c/div\u003e"},{"header":"3. Results and Discussion","content":"\u003cdiv id=\"Sec20\" class=\"Section2\"\u003e\u003ch2\u003e3.1. Growth characteristics of LAB in RTE VP salmon portions at 4℃\u003c/h2\u003e\u003cp\u003eIn the present study, four LAB strains, \u003cem\u003eC. maltaromaticum\u003c/em\u003e (C.m.35), \u003cem\u003eC. maltaromaticum\u003c/em\u003e (C.m.55), \u003cem\u003eC. divergens\u003c/em\u003e (C.d.468) and \u003cem\u003eL. gelidum\u003c/em\u003e (Le.g.406), were selected for biopreservation of RTE VP salmon portions for 21 days at 4℃. The strains were selected according to their origin in RTE seafood products, antimicrobial activity against \u003cem\u003eListeria\u003c/em\u003e spp., growth properties in salmon juice, and ability to grow in VP salmon portions at refrigerated temperature (Stupar et al., \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Stupar et al., \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). All strains displayed growth in the VP salmon at 4℃ without a lag phase, increasing from the initial concentration of 4.2\u0026ndash;4.6 log CFU/g to a final concentration of 7.2\u0026ndash;8.6 log CFU/g during the storage period (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). In comparison, the concentration of LAB in the control samples increased from 1.9\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2 to 6.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2 log CFU/g (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Compared to non-inoculated samples, LAB counts were significantly higher in inoculated samples, indicating their dominance during the storage period (one-way ANOVA, p\u0026thinsp;\u0026lt;\u0026thinsp;0.05), except for C.d.468 on days 14 and 21 (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eA).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eAmong the applied LAB strains, Le.g.406 reached the highest maximum growth rate (\u0026micro;\u003csub\u003emax\u003c/sub\u003e) and maximum population density (Y\u003csub\u003emax\u003c/sub\u003e), while C.d.468 had the lowest values (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The results align with growth kinetic parameters obtained previously for the same strains in a salmon juice model system at 4℃ (Stupar et al., \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), demonstrating that the salmon juice model system is reliable for assessing growth kinetic parameters in relevant conditions. No significant differences between MRS and IA plate counts were observed for any of the inoculated or non-inoculated samples (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eA and Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eB), except for Le.g.406 (Independent Samples Test, day 0 (p\u0026thinsp;\u0026lt;\u0026thinsp;0.001), day 7 (p\u0026thinsp;=\u0026thinsp;0.035), day 14 (p\u0026thinsp;\u0026lt;\u0026thinsp;0.001) and day 21 (p\u0026thinsp;=\u0026thinsp;0.007)), implying that bacteria quantified on IA corresponds to the inoculated LAB.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eGrowth kinetic parameters for selected LAB quantified on MRS agar. The growth kinetic parameters (initial count (log CFU/g), maximum growth rate, \u0026micro;\u003csub\u003emax\u003c/sub\u003e (logCFU/g/day), maximum population density, Y\u003csub\u003emax\u003c/sub\u003e (log CFU/g), and lag phase (day) are estimated by the primary model of Baranyi and Roberts (\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1994\u003c/span\u003e). R\u003csup\u003e2\u003c/sup\u003e-coefficient of determination; SE of Fit-standard error of fit; NL- no lag phase.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"7\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eGroup\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eInitial count\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u0026micro;\u003csub\u003emax\u003c/sub\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eY\u003csub\u003emax\u003c/sub\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eR\u003csup\u003e2\u003c/sup\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eSE of Fit\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003eLag phase\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003e(log CFU/g)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003e(log CFU/g/d)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003e(log CFU/g)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003e(days)\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eControl\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e1.9\u0026nbsp;\u0026plusmn;\u0026nbsp;0.2\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.32\u0026nbsp;\u0026plusmn;\u0026nbsp;0.03\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e6.6\u0026nbsp;\u0026plusmn;\u0026nbsp;0.2\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0.99\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0.23\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eNL\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC.m.35\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e4.2\u0026nbsp;\u0026plusmn;\u0026nbsp;0.2\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.42\u0026nbsp;\u0026plusmn;\u0026nbsp;0.04\u003csup\u003ebc\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e8.6\u0026nbsp;\u0026plusmn;\u0026nbsp;0.1\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0.99\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0.21\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eNL\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC.m.55\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e4.3\u0026nbsp;\u0026plusmn;\u0026nbsp;0.1\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.45\u0026nbsp;\u0026plusmn;\u0026nbsp;0.01\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e8.6\u0026nbsp;\u0026plusmn;\u0026nbsp;0.1\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0.99\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0.07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eNL\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eLe.g.406\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e4.6\u0026nbsp;\u0026plusmn;\u0026nbsp;0.1\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.48\u0026nbsp;\u0026plusmn;\u0026nbsp;0.02\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e8.6\u0026nbsp;\u0026plusmn;\u0026nbsp;0.1\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0.99\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0.08\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eNL\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC.d.468\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e4.5\u0026nbsp;\u0026plusmn;\u0026nbsp;0.2\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.20\u0026nbsp;\u0026plusmn;\u0026nbsp;0.04\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e7.2\u0026nbsp;\u0026plusmn;\u0026nbsp;0.2\u003csup\u003eab\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e\u003cp\u003e0.96\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e0.26\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003eNL\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ep-value\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003ep\u0026thinsp;\u0026lt;\u0026thinsp;0.05\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ep\u0026thinsp;\u0026lt;\u0026thinsp;0.05\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ep\u0026thinsp;=\u0026thinsp;0.05\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eSmall letters (abc) indicate significant differences between the groups (LAB-inoculated samples and controls), with the corresponding p-value at the bottom of each column, calculated by one-way ANOVA (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05).\u003c/p\u003e\u003cp\u003eC.m.35: \u003cem\u003eC. maltaromaticum\u003c/em\u003e 35; C.m.55: \u003cem\u003eC. maltaromaticum\u003c/em\u003e 55; Le.g.406: \u003cem\u003eL. gelidum\u003c/em\u003e 406; C.d.468: \u003cem\u003eC. divergens\u003c/em\u003e 468.\u003c/p\u003e\u003cp\u003eAlthough an accurate lag-phase determination requires more sampling points, no detectable lag phase suggests that the LAB strains adapted well to the salmon matrix and the applied storage conditions (VP and 4℃), probably because they originate from refrigerated seafood products (cold-smoked salmon (CSS), sushi and gravlax) (Stupar et al., \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). This is also supported by previous findings observed in salmon juice and pre-rigor filleted salmon, where more frequent measurements were conducted (Stupar et al., \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). The same adaptation and dominance of artificially added \u003cem\u003eC. maltaromaticum, Lactococcus piscium\u003c/em\u003e and \u003cem\u003eL. gelidum\u003c/em\u003e isolated from CSS and fresh salmon was observed in inoculated gravlax (Wiernasz et al., \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). However, the study of Aymerich et al. (\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) showed that LAB (\u003cem\u003eLactobacillus sakei\u003c/em\u003e), isolated from other food matrices (meat), can also be applied for the biopreservation of CSS.\u003c/p\u003e\u003cp\u003eFor biopreservative purposes, the absence of lag phase followed by a high growth rate for the inoculated LAB strain can potentially prevent indigenous spoilage microbiota and limit the production of undesirable metabolic products (Huang et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Zhou et al., \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). In addition, some LAB might produce several metabolic products with antagonistic effects on undesirable microbiota, where even beneficial microbiota such as endogenous LAB can be affected. Thus, the ability of the LAB strain to prevent growth of undesirable microorganisms (both spoilage microorganisms and pathogens) and, at the same time, co-exist with beneficial microbiota is crucial for its implementation in food products (Woo et al., \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eNo H\u003csub\u003e2\u003c/sub\u003eS-producing bacteria, representing fish spoilage indicator bacteria, were observed during storage, except for control samples at the end of storage (0.58 log CFU/g). Some LAB strains are associated with food spoilage due to H\u003csub\u003e2\u003c/sub\u003eS-production when metabolising cysteine (Leroi, \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Moreover, the low level of H\u003csub\u003e2\u003c/sub\u003eS-producing bacteria in the non-inoculated samples at the end of storage confirmed that the VP salmon was not spoiled by H\u003csub\u003e2\u003c/sub\u003eS-producing bacteria, also observed by Rode and Hovda (\u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) during 26 days of storage of VP salmon. In general, different levels of this bacterial group are reported for VP salmon. The study of Jakobsen et al. (\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) detected low counts of H\u003csub\u003e2\u003c/sub\u003eS-producing bacteria (\u0026lt;\u0026thinsp;3.8 log CFU/g) in VP salmon fillets stored at 4℃. On the contrary, the study of Hansen et al. (\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2009\u003c/span\u003e) reported the levels of H\u003csub\u003e2\u003c/sub\u003eS-producing bacteria to be above 6 log CFU/g after 15 days in VP salmon fillets at 1.2℃. In addition, microbial contamination is highly dependent on conditions during harvesting, degree of cross-contamination and type of processing and packaging (Chan et al., \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Rollini et al., \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). By applying LAB biopreservation, microbial contaminants on salmon fillets can probably be inhibited by non-specific competition between populations (Jameson-effect interaction), thereby prolonging salmon shelf life and quality (Bol\u0026iacute;var et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec21\" class=\"Section2\"\u003e\u003ch2\u003e3.2. Effect of LAB on microbial community composition in VP salmon\u003c/h2\u003e\u003cp\u003eThe effect of the four LAB strains (C.m.35, C.m.55, Le.g.406, and C.d.468) on the microbial community composition of RTE VP salmon was evaluated in samples collected on day 14. Illumina MiSeq sequencing generated 1 244 429 raw sequence reads, of which 1 244 101 sequences were obtained after quality processing, chimaera detection and filtering. A total of 909 382 (73.1%) sequences were assigned to operational taxonomic units (OTUs), ranging from 5 283 to 24 321 reads per sample. Endogenous LAB was present in control samples of both Batch I and II, but the relative abundance of LAB was significantly higher in Control 1 (Independent Samples Test, p\u0026thinsp;=\u0026thinsp;0.003). The same significant batch-to-batch variation in the abundance of endogenous LAB was observed using culture-dependent analysis (Independent Sample Test, p\u0026thinsp;\u0026lt;\u0026thinsp;0.001) (data not shown).\u003c/p\u003e\u003cp\u003eThe microbial community composition of Control 1 was dominated by LAB represented by unclassified \u003cem\u003eLactobacillales\u003c/em\u003e (relative abundance of 48.7\u0026thinsp;\u0026plusmn;\u0026thinsp;12.3%) and \u003cem\u003eLeuconostoc\u003c/em\u003e (5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.8%), and in addition, \u003cem\u003ePhotobacterium\u003c/em\u003e (47\u0026thinsp;\u0026plusmn;\u0026thinsp;12.9%), while Control 2 was dominated by \u003cem\u003ePhotobacterium\u003c/em\u003e with a relative abundance of 98.1\u0026thinsp;\u0026plusmn;\u0026thinsp;2.4% (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eIn LAB-inoculated samples, the community composition on day 14 corresponded to the inoculated LAB. The relative abundance of \u003cem\u003eLeuconostoc\u003c/em\u003e in the samples inoculated with strain Le.g.406 was 96.5\u0026thinsp;\u0026plusmn;\u0026thinsp;1.74% and thus significantly different from Control 2 where 1.76\u0026thinsp;\u0026plusmn;\u0026thinsp;2.1 of unclassified \u003cem\u003eLactobacillales\u003c/em\u003e was detected (Independent Sample Test, p\u0026thinsp;\u0026lt;\u0026thinsp;0.001). The relative abundance of \u003cem\u003eCarnobacterium\u003c/em\u003e in the samples inoculated with strain C.d.468 was 64.7\u0026thinsp;\u0026plusmn;\u0026thinsp;17.5% and thus significantly different from Control 2 (Independent Sample Test, p\u0026thinsp;=\u0026thinsp;0.003). In samples inoculated with C.m.35 and C.m.55, 81\u0026thinsp;\u0026plusmn;\u0026thinsp;3.8 and 76.8\u0026thinsp;\u0026plusmn;\u0026thinsp;12.5% of the microbial population were classified at the order level (\u003cem\u003eLactobacillales\u003c/em\u003e), respectively (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Although the LAB population could not be assigned to the genus level, it can be assumed that the dominating OTU, assigned as \u003cem\u003eLactobacillales\u003c/em\u003e, results from the inoculated strains. Furthermore, \u003cem\u003eLeuconostoc\u003c/em\u003e constituted 10.3\u0026thinsp;\u0026plusmn;\u0026thinsp;2.1 and 7.9\u0026thinsp;\u0026plusmn;\u0026thinsp;0.7%, while the relative abundance of \u003cem\u003ePhotobacterium\u003c/em\u003e was 8.4\u0026thinsp;\u0026plusmn;\u0026thinsp;6.2 and 15.5\u0026thinsp;\u0026plusmn;\u0026thinsp;12.9% in samples inoculated with C.m.35 and C.m.55, respectively (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). It is evident the chosen sequencing approach holds certain limitations regarding the taxonomic resolution of bacteria belonging to the order \u003cem\u003eLactobacillales\u003c/em\u003e (O\u0026rsquo;Callaghan et al., \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Rintala et al., \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), and a combined approach with full-length sequencing of the 16S rRNA gene could enable more accurate identification of the dominating LAB genera.\u003c/p\u003e\u003cp\u003eThe results demonstrated that inoculated LAB affected the salmon\u0026rsquo;s natural microbial community composition by outcompeting \u003cem\u003ePhotobacterium.\u003c/em\u003e Species belonging to \u003cem\u003ePhotobacterium\u003c/em\u003e, e.g. \u003cem\u003ePhotobacterium phosphoreum\u003c/em\u003e, are commonly recognised as fish spoilers (Antunes-Rohling et al., \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Dalgaard et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Tsoukalas et al., \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) due to a strong correlation with the production of BA (Bjornsdottir-Butler et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Li et al., \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), and ability to reduce trimethylamine oxide (TMAO) to the undesirable odourant trimethylamine (TMA) (J\u0026auml;\u0026auml;skel\u0026auml;inen et al., \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; L\u0026oacute;pez-Caballero et al., \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; S\u0026oslash;rensen et al., \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Moreover, \u003cem\u003ePhotobacterium\u003c/em\u003e is shown to be an H\u003csub\u003e2\u003c/sub\u003eS-producer under certain conditions (Tsoukalas et al., \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), but this was not confirmed in the present study as only low counts (\u0026lt;\u0026thinsp;1 log CFU/g) of H\u003csub\u003e2\u003c/sub\u003eS-producing bacteria were detected in control samples at the end of storage. Accordingly, the absence of H\u003csub\u003e2\u003c/sub\u003eS-production by \u003cem\u003ePhotobacterium\u003c/em\u003e has also been observed by Kuuliala et al. (\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). \u003cem\u003ePhotobacterium\u003c/em\u003e is one of the most abundant genera constituting the microbial community of chilled, packaged fresh salmon (Hansen et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; J\u0026auml;\u0026auml;skel\u0026auml;inen et al., \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Li et al., \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Mac\u0026eacute; et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Thus, the observed ability of the applied LAB strains to outcompete \u003cem\u003ePhotobacterium\u003c/em\u003e is of significant importance as inhibition of this spoilage bacterium is likely to extend the shelf-life of chilled MAP and VP fish (Dalgaard et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Tsoukalas et al., \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec22\" class=\"Section2\"\u003e\u003ch2\u003e3.3. ATP-degradation products\u003c/h2\u003e\u003cp\u003eThe ATP-degradation products were analysed on day 0 (prior to inoculation) and days 8 and 22. The ATP-degradation products AMP and IMP are associated with the umami taste (Johnson et al., \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Mouritsen, \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), while loss of freshness and unpleasant flavours of the fish are related to IMP degradation to Ino and Hx (Yu et al., \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The Ki-value (Karube et al., \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e1984\u003c/span\u003e), highly correlated to the K-value, is a freshness indicator mainly recommended for the assessment of early changes during fish storage (Hong et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), while H-values are more effective in expressing the influence of bacterial enzymes on the freshness and spoilage of salmon. No significant effect on the Ki-value and H-value (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e) was observed when applying biopreservation. The initial Ki-value in control samples was 34.5\u0026thinsp;\u0026plusmn;\u0026thinsp;2.8, with a significant increase at the end of storage, reaching 99.9\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2 (r\u0026thinsp;=\u0026thinsp;0.919, p\u0026thinsp;\u0026lt;\u0026thinsp;0.001). No adverse effect of inoculated LAB on the accumulation of off-flavours and thus on Ki- and H-value suggests the high potential of these strains to be used in RTE salmon products.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eATP-degradation products. Concentration of inosine monophosphate (IMP (\u0026micro;mol/g)), inosine (Ino (\u0026micro;mol/g)), hypoxanthine (Hx (\u0026micro;mol/g)), Ki-value, and H-value (%) in lactic acid bacteria (LAB)-inoculated (n\u0026thinsp;=\u0026thinsp;3) and non-inoculated (control) samples (n\u0026thinsp;=\u0026thinsp;6) in vacuum-packed salmon portions on days 8 and 22.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"7\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eGroup\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eDay after packaging\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eIMP\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eIno\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eHx\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eH-value\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003eKi-value\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003e(\u0026micro;mol/g)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003e(\u0026micro;mol/g)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003e(\u0026micro;mol/g)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003e(%)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003e(%)\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eControl\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e3.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e3.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e47\u0026thinsp;\u0026plusmn;\u0026thinsp;12\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e90\u0026thinsp;\u0026plusmn;\u0026thinsp;7\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC.m.35\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.9\u0026thinsp;\u0026plusmn;\u0026thinsp;0.9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e3.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e38\u0026thinsp;\u0026plusmn;\u0026thinsp;6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e89\u0026thinsp;\u0026plusmn;\u0026thinsp;9\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC.m.55\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.7\u0026thinsp;\u0026plusmn;\u0026thinsp;1.3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e3.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e37\u0026thinsp;\u0026plusmn;\u0026thinsp;16\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e80\u0026thinsp;\u0026plusmn;\u0026thinsp;15\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eLe.g.406\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.5\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e33\u0026thinsp;\u0026plusmn;\u0026thinsp;3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e82\u0026thinsp;\u0026plusmn;\u0026thinsp;3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC.d.468\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e3.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e3.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e51\u0026thinsp;\u0026plusmn;\u0026thinsp;9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e93\u0026thinsp;\u0026plusmn;\u0026thinsp;2\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ep-value\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ep\u0026thinsp;=\u0026thinsp;0.28\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ep\u0026thinsp;=\u0026thinsp;0.05\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ep\u0026thinsp;=\u0026thinsp;0.17\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003ep\u0026thinsp;=\u0026thinsp;0.20\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003ep\u0026thinsp;=\u0026thinsp;0.28\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eControl\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e22\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1.4\u0026thinsp;\u0026plusmn;\u0026thinsp;1.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e5.2\u0026thinsp;\u0026plusmn;\u0026thinsp;1.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e78\u0026thinsp;\u0026plusmn;\u0026thinsp;14\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e99.9\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC.m.35\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e22\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e6.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e90\u0026thinsp;\u0026plusmn;\u0026thinsp;6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e99.8\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC.m.55\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e22\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e6.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e90\u0026thinsp;\u0026plusmn;\u0026thinsp;4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e100\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eLe.g.406\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e22\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e0.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e6.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e95\u0026thinsp;\u0026plusmn;\u0026thinsp;3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e100\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC.d.468\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e22\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003eND\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e5.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e75\u0026thinsp;\u0026plusmn;\u0026thinsp;4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e100\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ep-value\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ep\u0026thinsp;=\u0026thinsp;0.10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ep\u0026thinsp;=\u0026thinsp;0.07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003ep\u0026thinsp;=\u0026thinsp;0.07\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003ep\u0026thinsp;=\u0026thinsp;0.75\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eC.m.35: \u003cem\u003eC. maltaromaticum\u003c/em\u003e 35; C.m.55: \u003cem\u003eC. maltaromaticum\u003c/em\u003e 55; Le.g.406: \u003cem\u003eL. gelidum\u003c/em\u003e 406; C.d.468: \u003cem\u003eC. divergens\u003c/em\u003e 468.\u003c/p\u003e\u003cp\u003eND: Not detected\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec23\" class=\"Section2\"\u003e\u003ch2\u003e3.4. Biogenic amines\u003c/h2\u003e\u003cp\u003eThe content of five BAs (cadaverine, spermidine, tryptamine, spermine, and tyramine) was quantified in VP fresh salmon portions on day 0 (prior to inoculation), 8 and 22. Low cadaverine levels were detected on day 8 (\u0026lt;\u0026thinsp;1 \u0026micro;g/g), and day 22 (0.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3\u0026ndash;1.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1 \u0026micro;g/g) in all samples (Additional file 1). Spermidine (0.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3 \u0026micro;g/g) and tryptamine (0.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4 \u0026micro;g/g) were only detected in the samples inoculated with C.m.55 on day 8, while these BAs were not detected on day 22. The low quantities of BAs in these samples indicate that they maintained acceptable quality even after 22 days of storage.\u003c/p\u003e\u003cp\u003eBAs are of concern either due to their toxicological effect on humans (W\u0026oacute;jcik et al., \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and/or their correlation with spoilage by producing off-odours (Katikou et al., \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). The present results showed no effect of the inoculated LAB on BA formation, which is essential for their potential industrial implementation. Although low quantities of spermidine and tryptamine were observed in samples inoculated with C.m.55, this should be taken with caution when considering this strain for biopreservation of processed salmon products or products within favourable conditions for decarboxylation activity (Biji et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Moreover, the study of \u0026Ouml;zogul and Hamed (\u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2018\u003c/span\u003e) emphasised that LAB can inhibit the growth of foodborne bacteria and thereby prevent their BA formation, however, other LAB can stimulate the BA production of foodborne pathogens. Thus, as the effect is strain and food matrix-dependent (\u0026Ouml;zogul \u0026amp; Hamed, \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2018\u003c/span\u003e), it must be investigated as a criterion for strain selection for biopreservative purposes.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec24\" class=\"Section2\"\u003e\u003ch2\u003e3.5. LAB effect on physiochemical properties of VP fresh salmon portions\u003c/h2\u003e\u003cp\u003eChanges in WC, DL and pH during the storage period at 4℃ are presented in Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eA, \u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eB, and \u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eC. No significant effect of inoculated LAB on WC, DL and pH was observed during the storage period in fresh VP salmon portions (one-way ANOVA, p\u0026thinsp;\u0026lt;\u0026thinsp;0.05). Colour and textural changes in VP fresh salmon portions were analysed on days 8 and 22. No significant differences were found between LAB-inoculated and corresponding control samples in colour parameters (lightness (L*), redness (a*), yellowness (b*), chroma (C*), and total colour difference (ΔE)) (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e), breaking force and firmness (F60%) during the storage period (Additional file 1). The only difference was observed for the hue angle (h*) on day 22 (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eDifferences in colour parameters (lightness (L*), redness (a*), yellowness (b*), chroma (C*), hue angle (h*) and total colour difference (ΔE)) in LAB-inoculated samples compared to the control at the same day, expressed as Δ-values (n\u0026thinsp;=\u0026thinsp;3)\u0026thinsp;\u0026plusmn;\u0026thinsp;SD. Data with * represents negative Δ-values.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"8\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eGroup\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eDay after packaging\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eΔL*\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eΔa*\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003eΔb*\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\"\u003e\u003cp\u003eΔc*\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c7\"\u003e\u003cp\u003eΔh*\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c8\"\u003e\u003cp\u003eΔ(ΔE)\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC.m.35\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e*3.6\u0026thinsp;\u0026plusmn;\u0026thinsp;1.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e3.3\u0026thinsp;\u0026plusmn;\u0026thinsp;1.0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e3.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e4.7\u0026thinsp;\u0026plusmn;\u0026thinsp;1.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e*0.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e1.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.9\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC.m.55\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e*2.2\u0026thinsp;\u0026plusmn;\u0026thinsp;3.0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e2.3\u0026thinsp;\u0026plusmn;\u0026thinsp;3.3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2.4\u0026thinsp;\u0026plusmn;\u0026thinsp;4.4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e3.3\u0026thinsp;\u0026plusmn;\u0026thinsp;5.4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.1\u0026thinsp;\u0026plusmn;\u0026thinsp;1.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e2.1\u0026thinsp;\u0026plusmn;\u0026thinsp;1.9\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eLe.g.406\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e*3.6\u0026thinsp;\u0026plusmn;\u0026thinsp;3.0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e2.6\u0026thinsp;\u0026plusmn;\u0026thinsp;3.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2.7\u0026thinsp;\u0026plusmn;\u0026thinsp;5.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e3.7\u0026thinsp;\u0026plusmn;\u0026thinsp;6.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e*0.0\u0026thinsp;\u0026plusmn;\u0026thinsp;1.3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.8\u0026thinsp;\u0026plusmn;\u0026thinsp;2.0\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC.d.468\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.3\u0026thinsp;\u0026plusmn;\u0026thinsp;3.8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e*1.6\u0026thinsp;\u0026plusmn;\u0026thinsp;4.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e*0.9\u0026thinsp;\u0026plusmn;\u0026thinsp;5.0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e*1.8\u0026thinsp;\u0026plusmn;\u0026thinsp;6.9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e0.3\u0026thinsp;\u0026plusmn;\u0026thinsp;2.3\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ep-value\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ep\u0026thinsp;=\u0026thinsp;0.22\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ep\u0026thinsp;=\u0026thinsp;0.36\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ep\u0026thinsp;=\u0026thinsp;0.63\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003ep\u0026thinsp;=\u0026thinsp;0.50\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003ep\u0026thinsp;=\u0026thinsp;0.78\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ep\u0026thinsp;=\u0026thinsp;0.69\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC.m.35\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e22\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e*1.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e2.6\u0026thinsp;\u0026plusmn;\u0026thinsp;1.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.8\u0026thinsp;\u0026plusmn;\u0026thinsp;1.3\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e8.2\u0026thinsp;\u0026plusmn;\u0026thinsp;2.6\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e*8.9\u0026thinsp;\u0026plusmn;\u0026thinsp;6.2\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC.m.55\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e22\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.5\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e*1.7\u0026thinsp;\u0026plusmn;\u0026thinsp;1.0\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e1.9\u0026thinsp;\u0026plusmn;\u0026thinsp;1.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e0.2\u0026thinsp;\u0026plusmn;\u0026thinsp;1.4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e7.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.9\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e*9.5\u0026thinsp;\u0026plusmn;\u0026thinsp;7.6\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eLe.g.406\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e22\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.2\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e1.0\u0026thinsp;\u0026plusmn;\u0026thinsp;3.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e0.9\u0026thinsp;\u0026plusmn;\u0026thinsp;4.7\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e1.3\u0026thinsp;\u0026plusmn;\u0026thinsp;5.8\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e*0.1\u0026thinsp;\u0026plusmn;\u0026thinsp;1.5\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e*0.1\u0026thinsp;\u0026plusmn;\u0026thinsp;1.4\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC.d.468\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e\u003cp\u003e22\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e2.9\u0026thinsp;\u0026plusmn;\u0026thinsp;5.1\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e*1.8\u0026thinsp;\u0026plusmn;\u0026thinsp;1.9\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e*0.9\u0026thinsp;\u0026plusmn;\u0026thinsp;1.6\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003e*1.9\u0026thinsp;\u0026plusmn;\u0026thinsp;2.4\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003e0.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.7\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003e*0.2\u0026thinsp;\u0026plusmn;\u0026thinsp;1.8\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ep-value\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003ep\u0026thinsp;=\u0026thinsp;0.86\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003ep\u0026thinsp;=\u0026thinsp;0.36\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003ep\u0026thinsp;=\u0026thinsp;0.46\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u003cp\u003ep\u0026thinsp;=\u0026thinsp;0.66\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c7\"\u003e\u003cp\u003ep\u0026thinsp;\u0026lt;\u0026thinsp;0.05\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c8\"\u003e\u003cp\u003ep\u0026thinsp;=\u0026thinsp;0.08\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eSmall letters (ab) indicate significant differences between groups on the same day with the corresponding p-value at the bottom of each column, calculated by one-way ANOVA (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05).\u003c/p\u003e\u003cp\u003eC.m.35: \u003cem\u003eC. maltaromaticum\u003c/em\u003e 35; C.m.55: \u003cem\u003eC. maltaromaticum\u003c/em\u003e 55; Le.g.406: \u003cem\u003eL. gelidum\u003c/em\u003e 406; C.d.468: \u003cem\u003eC. divergens\u003c/em\u003e 468.\u003c/p\u003e\u003cp\u003eOverall, the selected LAB strains are considered suitable candidates for use in fresh VP salmon portions due to their demonstrated antagonistic activity and ability to maintain product quality. Similar results are also confirmed in other food products. In the study of Danielski et al. (\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), C. \u003cem\u003emaltaromaticum\u003c/em\u003e did not affect the physicochemical properties of cooked ham, while in the study of Boulares et al. (\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), reduced water- and lipid loss were achieved for sea bass inoculated with combinations of \u003cem\u003eLactococcus lactic, Lactobacillus plantarum\u003c/em\u003e (\u003cem\u003eLactiplantibacillus plantarum\u003c/em\u003e), and \u003cem\u003eC. piscicola\u003c/em\u003e (\u003cem\u003eC. maltaromaticum\u003c/em\u003e). In the study of Mozuriene et al. (\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), an improved effect on the colour and texture of pork meat was observed by lacto-fermented potato tuber juice treatment (\u003cem\u003ePediococcus acidilactici, P. pentosaceus\u003c/em\u003e and \u003cem\u003eLatilactobacillus sakei\u003c/em\u003e). However, higher drip loss and lower water holding capacity were observed for LAB-treated samples compared to the control in the same study (Mozuriene et al., \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2016\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eOverall, adding LAB to the VP fresh salmon portions did not affect any of the measured physiochemical parameters. As these parameters are tightly connected (e.g. LAB could provide a proteolytic effect by degrading muscle proteins as well as connective tissue, which in turn impacts the water-holding properties), preserving the physicochemical properties without significant changes in the pH of the fresh salmon is crucial for consumer acceptance.\u003c/p\u003e\u003c/div\u003e\u003cdiv id=\"Sec25\" class=\"Section2\"\u003e\u003ch2\u003e3.6. Antilisterial effect of inoculated LAB strains\u003c/h2\u003e\u003cp\u003eThe selected LAB strain's antibacterial activity in VP fresh salmon portions was examined. In the absence of inoculated LAB, \u003cem\u003eL. innocua\u003c/em\u003e proliferated from an initial concentration of 3.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1 to 5.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3 log CFU/g during 21 days of storage at 4℃ (Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). However, in the presence of C.m.35, \u003cem\u003eL. innocua\u003c/em\u003e was not able to proliferate in the product. On the last day of storage, there was no significant difference in the concentration of \u003cem\u003eListeria\u003c/em\u003e compared to the initial concentration (p\u0026thinsp;\u0026gt;\u0026thinsp;0.05) (Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). In fact, significantly lower counts of \u003cem\u003eL. innocua\u003c/em\u003e were found on day 14 (p\u0026thinsp;=\u0026thinsp;0.001) and day 21 (p\u0026thinsp;=\u0026thinsp;0.004) compared to samples inoculated with only \u003cem\u003eL. innocua\u003c/em\u003e (Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). Although both C.m.35 and C.d.468 are recognised as producers of bacteriocin-like substances (BLS) (Stupar et al., \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), the same inhibition of \u003cem\u003eL. innocua\u003c/em\u003e was not observed for the C.d.468 or the two non-bacteriocin-producing strains (Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e). Even though all LAB strains performed total inhibition of \u003cem\u003eL. innocua in vitro\u003c/em\u003e (Stupar et al., \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Stupar et al., \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), a lower effect was observed in salmon portions, indicating that food matrix, initial concentrations, and pH can govern inter-species competition (Bol\u0026iacute;var et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Mellefont et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Noteworthy lower initial concentrations of LAB (~\u0026thinsp;10\u003csup\u003e4\u003c/sup\u003e CFU/g) and higher target concentration (~\u0026thinsp;10\u003csup\u003e3\u003c/sup\u003e CFU/g) were used in the present study compared to the \u003cem\u003ein vitro\u003c/em\u003e studies in pH-adjusted salmon juice (Stupar et al., \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Stupar et al., \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Previously, it was shown that most strains use nutrient competition as the main mechanism against the target organism (Stupar et al., \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), assuming the Jameson effect, where higher LAB concentrations play an essential role (Mellefont et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2008\u003c/span\u003e). Thus, a higher initial LAB concentration and/or a combination of several LAB strains might express more efficacy against undesirable microbiota. For example, in the study of Boulares et al. (\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), the highest reduction of \u003cem\u003eL. monocytogenes\u003c/em\u003e was \u0026gt;\u0026thinsp;3 log CFU/g in fresh VP sea bass stored for 21 days at 4℃, achieved by combining two or three LAB strains at high initial concentrations (8 log CFU/g of each). Moreover, \u003cem\u003eLactobacillus casei\u003c/em\u003e demonstrated bacteriostatic and bactericidal effects when inoculated at concentrations of 6 and 8 log CFU/g, respectively, against \u003cem\u003eL. innocua\u003c/em\u003e in VP CSS (Vescovo et al., \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). In the present study, a lower initial LAB concentration was applied to reduce the risk of negative effects on physicochemical properties of fresh salmon, but at the same time aiming to achieve sufficient antimicrobial activity. Combined with other hurdle technologies, the applied approach might be even more powerful. Furthermore, the lack of antilisterial effect of the BLS-producing strain C.d.468 during storage (GLM, p\u0026thinsp;=\u0026thinsp;0.599) might also be connected to non-optimal conditions for the growth and bacteriocin activity of the applied strain (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eA).\u003c/p\u003e\u003cp\u003eThe observed bacteriostatic effect of strain C.m.35 is an effective hurdle for the multiplication of \u003cem\u003eL. innocua\u003c/em\u003e. The present study simulated a worst-case scenario, given the unrealistically high initial target concentrations (Svanevik et al., 2021) and the application of a target strain demonstrably more resistant to biopreservation than \u003cem\u003eL. monocytogenes\u003c/em\u003e (Stupar et al., \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Thus, given the expected low contamination level of \u003cem\u003eL. monocytogenes\u003c/em\u003e in fresh salmon (Svanevik et al., 2021), inhibition of 1\u0026ndash;2 log CFU/g would be sufficient to keep the level of this pathogen within the safe zone during refrigerated storage. On the other hand, more studies, including a range of \u003cem\u003eL. monocytogenes\u003c/em\u003e strains from various serotypes regularly detected in the salmon product and /or the salmon processing environment, should be conducted to give a broader insight into the LAB strain's antilisterial effect and their ability to act as a protective hurdle in RTE salmon products. Despite the QPS status of \u003cem\u003eC. maltaromaticum\u003c/em\u003e, \u003cem\u003eC. divergens\u003c/em\u003e and \u003cem\u003eLeuconostoc\u003c/em\u003e spp. (EFSA BIOHAZ Panel, 2021) the strains should be further safety tested before any industrial application. In particular, safety documentation for food applications of \u003cem\u003eL. gelidum\u003c/em\u003e strains is deficient. The application of protective cultures in raw salmon products will probably be defined as \u0026ldquo;novel food\u0026rdquo; in the EU and must be approved by food safety authorities before industrial applications. Although no negative effect of LAB inoculation was observed on quality parameters in the current study, the sensory aspects must be further elucidated using a sensory panel, and a consumer acceptance analysis must be conducted before industrial applications.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab5\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 5\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eProliferation of \u003cem\u003eListeria innocua\u003c/em\u003e (Target) in vacuum-packed salmon portions in the presence of selected LAB for 21 days at 4℃. The results are presented as mean log CFU/g\u0026thinsp;\u0026plusmn;\u0026thinsp;SD (n\u0026thinsp;=\u0026thinsp;3 for target in coculture with LAB and n\u0026thinsp;=\u0026thinsp;6 for target-monoculture).\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"6\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003eGroup\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colspan=\"4\" nameend=\"c5\" namest=\"c2\"\u003e\u003cp\u003eDay after packaging\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c6\" morerows=\"1\" rowspan=\"2\"\u003e\u003cp\u003ep-value\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003e7\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003e14\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c5\"\u003e\u003cp\u003e21\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC.m.35\u0026thinsp;+\u0026thinsp;T\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e3.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1\u003csup\u003eIb\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e3.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3\u003csup\u003eIa\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e3.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4\u003csup\u003eIa\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e4.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.5\u003csup\u003eIa\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e=\u0026thinsp;0.19\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC.m.55\u0026thinsp;+\u0026thinsp;T\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e3.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0\u003csup\u003eIa\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e3.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3\u003csup\u003eIIab\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3\u003csup\u003eII,IIIab\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e4.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2\u003csup\u003eIIIab\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e\u0026lt;\u0026thinsp;0.05\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eLe.g.406\u0026thinsp;+\u0026thinsp;T\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e3.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0\u003csup\u003eIb\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e4.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3\u003csup\u003eI,IIab\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4.1\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4\u003csup\u003eII, IIIab\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e4.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1\u003csup\u003eIIIab\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e\u0026lt;\u0026thinsp;0.05\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eC.d.468\u0026thinsp;+\u0026thinsp;T\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e3.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2\u003csup\u003eIb\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e4.3\u0026thinsp;\u0026plusmn;\u0026thinsp;0.4\u003csup\u003eIIb\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4.6\u0026thinsp;\u0026plusmn;\u0026thinsp;0.0\u003csup\u003eII,IIIb\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e5.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1\u003csup\u003eIIIb\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e\u0026lt;\u0026thinsp;0.05\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eTarget\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e3.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1\u003csup\u003eIb\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e4.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.2\u003csup\u003eIIab\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e4.7\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3\u003csup\u003eIIIb\u003c/sup\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e5.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3\u003csup\u003eIIIb\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e\u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e\u003cp\u003e\u0026lt;\u0026thinsp;0.05\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ep-value\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u0026lt;\u0026thinsp;0.05\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u0026lt;\u0026thinsp;0.05\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u0026lt;\u0026thinsp;0.05\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c5\"\u003e\u003cp\u003e\u0026lt;\u0026thinsp;0.05\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003ctfoot\u003e\u003ctr\u003e\u003ctd colspan=\"6\"\u003eSmall letters (\u003csup\u003eab\u003c/sup\u003e) indicate significant differences between the groups on the same day, with the corresponding p-value at the bottom of each column. Roman numbers (\u003csup\u003eI, II, III\u003c/sup\u003e) indicate significant differences in a group at all tested days, with the corresponding p-value in the same row. Significant differences are calculated by one-way ANOVA, Tukey HSD, p\u0026thinsp;\u0026lt;\u0026thinsp;0.05. C.m.35: \u003cem\u003eC. maltaromaticum\u003c/em\u003e 35; C.m.55: \u003cem\u003eC. maltaromaticum\u003c/em\u003e 55; Le.g.406: \u003cem\u003eL. gelidum\u003c/em\u003e 406; C.d.468: \u003cem\u003eC. divergens\u003c/em\u003e 468. T\u0026thinsp;=\u0026thinsp;Target; Target\u0026thinsp;=\u0026thinsp;\u003cem\u003eL. innocua\u003c/em\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tfoot\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003c/div\u003e"},{"header":"Conclusion","content":"\u003cp\u003eIn the present study, four LAB strains were selected for biopreservation of VP fresh salmon portions. The LAB strains demonstrated good adaptation to the salmon matrix and did not negatively affect the physicochemical and chemical properties of VP salmon portions at 4℃ for 22 days of storage. Moreover, the selected LAB strains are considered safe in terms of no significant production of biogenic amines. The strain \u003cem\u003eC. maltaromaticum\u003c/em\u003e 35 (C.m.35) demonstrated a bacteriostatic effect on \u003cem\u003eL. innocua\u003c/em\u003e, which makes it an interesting candidate for controlling \u003cem\u003eL. monocytogenes\u003c/em\u003e in ready-to-eat salmon. In a future perspective, the effect of this strain could be optimised by combining it with other LAB strains or applying additional shelf-life-improving strategies.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003eAuthors\u0026rsquo; contributions\u003c/p\u003e\n\u003cp\u003eJelena Stupar: Formal analysis, Investigation, Data curation, Methodology, Visualization, Writing-Original Draft, Writing-review and editing; Sunniva Hoel: Supervision, Conceptualization, Methodology, Visualization, Writing-Review \u0026amp; Editing; J\u0026oslash;rgen Lerfall: Supervision, Conceptualization, Methodology, Visualization, Writing-Review \u0026amp; Editing; Turid Rustad: Supervision, Conceptualization, Methodology, Visualization, Writing-Review \u0026amp; Editing; \u0026nbsp;Anita Nordeng Jakobsen: Supervision, Conceptualization, Methodology, Visualization, Writing-Review \u0026amp; Editing, Project Administration.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eAcknowledgements\u003c/p\u003e\n\u003cp\u003eThe authors would like to thank the technical staff at the Department of Biotechnology and Food Science at NTNU for practical help in the microbiological, analytical and food processing laboratory.\u003c/p\u003e\n\u003cp\u003eEthics declarations\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003eEthics approval and consent to participate\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003eConsent for publication\u003c/p\u003e\n\u003cp\u003eAll authors have read and agreed to the published version of the manuscript.\u003c/p\u003e\n\u003cp\u003eCompeting interests\u003c/p\u003e\n\u003cp\u003eThe authors have no known conflicts of interest to declare.\u003c/p\u003e\n\u003cp\u003eFunding sources\u003c/p\u003e\n\u003cp\u003eThis research did not receive any specific grant from funding agencies in the public, commercial, or not-for-profit sectors.\u003c/p\u003e\n\u003cp\u003eData availability\u003c/p\u003e\n\u003cp\u003eData will be made available on request.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAngiolillo, L., Conte, A., \u0026amp; Del Nobile, M. 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Effect of lactic acid bacteria on mackerel (\u003cem\u003ePneumatophorus japonicus\u003c/em\u003e) seasoning quality and flavor during fermentation. \u003cem\u003eFood Bioscience\u003c/em\u003e, \u003cem\u003e41\u003c/em\u003e, 100971. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/https://doi.org/10.1016/j.fbio.2021.100971\u003c/span\u003e\u003cspan address=\"10.1016/j.fbio.2021.100971\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Lactic acid bacteria, Biopreservation, Listeria, ready-to-eat salmon, Physicochemical properties","lastPublishedDoi":"10.21203/rs.3.rs-7968166/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7968166/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eLactic acid bacteria (LAB) are recognised as candidates for biopreservation because of their antagonistic activities. However, the LAB strain's efficacy in and compatibility with a specific food matrix must be emphasised to develop biopreservation as an efficient hurdle for industrial applications. The aim of the present study was to investigate the biopreservative potential of four lactic acid bacteria (LAB; \u003cem\u003eCarnobacterium maltaromaticum\u003c/em\u003e 35, \u003cem\u003eC. maltaromaticum\u003c/em\u003e 55, \u003cem\u003eC. divergens\u003c/em\u003e 468, and \u003cem\u003eLeuconostoc gelidum\u003c/em\u003e 406) in vacuum-packed (VP) ready-to-eat (RTE) salmon portions stored at 4℃ for 22 days. The evaluation was based on the strain's growth properties, their effect on the microbial community structure, their ability to inhibit an artificially inoculated strain of \u003cem\u003eListeria innocua\u003c/em\u003e (CCUG 15531) and their effect on chemical (ATP-degradation products and biogenic amines (BA)) and physicochemical properties (pH, colour, and water-holding properties (WHP)) of the salmon portions. All LAB strains grew well, increasing their concentration from 4.2\u0026ndash;4.6 log CFU/g to 7.2\u0026ndash;8.6 log CFU/g. The cultures outcompeted the population of the spoilage organism \u003cem\u003ePhotobacterium\u003c/em\u003e, which predominated the microbial community of the control samples. LAB strains did not negatively affect chemical and physicochemical properties. In the absence of inoculated LAB, \u003cem\u003eL. innocua\u003c/em\u003e increased significantly from 3.4\u0026thinsp;\u0026plusmn;\u0026thinsp;0.1 to 5.0\u0026thinsp;\u0026plusmn;\u0026thinsp;0.3 log CFU/g, while in the presence of \u003cem\u003eC. maltaromaticum\u003c/em\u003e 35, no significant proliferation was observed (p\u0026thinsp;=\u0026thinsp;0.19). Thus, \u003cem\u003eC. maltaromaticum\u003c/em\u003e 35 could be an efficient hurdle to ensure safe and microbiologically stable RTE salmon products.\u003c/p\u003e","manuscriptTitle":"Biopreservation of ready-to-eat Atlantic salmon (Salmo salar) by lactic acid bacteria: Effect on safety and quality parameters","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-12-09 07:07:05","doi":"10.21203/rs.3.rs-7968166/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"c0219566-4045-41a8-aaa7-9e9fc29d2868","owner":[],"postedDate":"December 9th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2026-02-21T12:40:09+00:00","versionOfRecord":[],"versionCreatedAt":"2025-12-09 07:07:05","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-7968166","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7968166","identity":"rs-7968166","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}