Reframing seafood quality and safety through synergistic cold plasma strategies

2.1. Thermal preservation

Heat treatment is one of the most widely used conventional preservation strategies to ensure microbial safety and extend the shelf life of seafood products. Depending on the desired shelf life, microbial risk profile, and intended mode of consumption, different heat-based approaches, such as cook–chill processing, pasteurization, and retort sterilization, are used in seafood systems.

Cook–chill processing involves cooking seafood products, rapidly cooling them within a defined time frame, storing them under refrigerated conditions, and reheating them prior to consumption. This approach is particularly relevant for ready-to-eat seafood products, as it provides convenience while delaying microbial spoilage and extending refrigerated shelf life (Pramandari et al., 2023). Pasteurization is another important thermal process used for products such as crab meat, surimi, and caviar. It commonly involves mild heat treatments below 100°C and aims to suppress microbial activity while maintaining product acceptability. However, pasteurization may not fully inactivate endogenous enzymes, and thermal load may adversely affect colour, texture, and flavour, particularly in delicate seafood matrices (Rosnes et al., 2011; Ravi et al., 2025). More intensive treatments, such as sterilization and retort processing, are used to produce shelf-stable seafood products and to control risks associated with spore-forming pathogens, particularly Clostridium botulinum. Although these treatments provide wide microbiological safety margins, the high temperatures and prolonged processing times required may result in sensory deterioration, nutrient losses, and pronounced textural changes in canned or retorted seafood products (Jimenez et al., 2024).

The major limitation of heat-based preservation is the unavoidable trade-off between microbial safety and product quality. Thermal processing can alter texture, colour, water-holding capacity, protein structure, lipid stability, and nutritional quality in fish and other seafood products, depending on the intensity and duration of heating (Hassoun et al., 2020a). Seafood muscle is highly sensitive to thermal processing because heat-induced denaturation and aggregation of myofibrillar proteins can reduce water-holding capacity and lead to undesirable textural changes such as firmness, dryness, or structural breakdown. These effects are especially critical in delicate fish species, where even moderate thermal loads may reduce juiciness, chewiness, and consumer acceptability (Strzelczak et al., 2021; Tan et al., 2024). In addition, seafood products are rich in polyunsaturated fatty acids and are therefore susceptible to heat-induced lipid oxidation. Elevated temperatures may accelerate the formation of secondary oxidation products, including aldehydes and ketones, which contribute to rancid flavours, off-odours, and overall sensory quality loss (Aubourg, 2023).

Thermal processing may also compromise the nutritional value of seafood. Long-chain omega-3 fatty acids, particularly eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA), are sensitive to heat and oxygen, and their degradation during high-temperature or long-duration processing can reduce one of the main health-promoting attributes of seafood products (Suryati et al., 2022, 2025). Heat-sensitive vitamins, bioactive compounds, and some amino acids may also be partially degraded during intensive treatments such as sterilization, potentially reducing nutritional quality and bioavailability (Jimenez et al., 2024). In crustaceans, heat-induced denaturation of carotenoprotein complexes releases astaxanthin, producing the characteristic red colouration of cooked shrimp; however, excessive or uncontrolled heating can lead to pigment degradation, uneven colour development, aroma loss, and reduced visual appeal (Yang et al., 2015).

Overall, thermal preservation methods remain reliable, cost-effective, and widely used tools for microbial inactivation in seafood products. However, because they inevitably cook the product and may cause texture deterioration, lipid oxidation, nutrient loss, pigment degradation, and undesirable sensory changes, they are not suitable for maintaining the fresh-like characteristics of raw or minimally processed seafood. These limitations highlight the need for non-thermal or minimally intensive preservation technologies that can improve microbial safety while better preserving the sensory and nutritional quality of seafood.

2.2. Traditional preservation methods

Traditional preservation methods, including salting, drying, smoking, marinating, and fermenting, have long been used to extend the shelf life of fish and other seafood products. These approaches have been applied for centuries and continue to be used in both traditional production systems and modern processing facilities, with improved process control and standardization. Their long-standing use reflects their accessibility, relatively low cost, cultural acceptance, and effectiveness in delaying spoilage under conditions where advanced preservation infrastructure or continuous cold-chain systems may be limited.

The preservative effect of these methods is mainly due to reduced water activity, lowered pH, inhibited microbial growth, and slowed biochemical spoilage reactions. Salting and drying restrict microbial proliferation by decreasing the availability of free water in the product (Rani et al., 2022), whereas smoking contributes to preservation through dehydration and the antimicrobial and antioxidant effects of smoke-derived compounds (Messina et al., 2021). Marination inhibits microbial growth and delays spoilage by combining organic acids and salt, lowering product pH and reducing microbial activity (Aguilar & Pengson, 2024). Fermentation improves product stability through acidification, the activity of competitive or starter microflora, and the production of antimicrobial metabolites, thereby contributing to microbial control and shelf-life extension (Li et al., 2024).

Despite these advantages, traditional preservation methods have important limitations. Because they often rely on substantial changes in water activity, salt concentration, acidity, smoke exposure, or microbial fermentation, they may considerably alter the original sensory and nutritional characteristics of seafood. Salting and drying may result in excessive saltiness, texture hardening, dehydration-related quality loss, and reduced consumer acceptance among individuals seeking fresh-like products. Smoking can provide desirable flavour, colour, and antioxidant effects; however, excessive or poorly controlled smoking can lead to significant sensory changes and the accumulation of smoke-derived contaminants, such as polycyclic aromatic hydrocarbons, which require careful process control in smoked seafood products (Tongo et al., 2017). Marinated and fermented seafood products may also differ substantially from fresh seafood in terms of acidity, flavour profile, texture, and appearance. Therefore, although these approaches remain accessible, economical, and culturally important, they are generally less suitable for preserving the fresh-like attributes of raw or minimally processed seafood products.

Modern consumers increasingly demand seafood products that are safe, minimally processed, nutritious, and close to their original fresh characteristics. In this context, traditional preservation methods may not fully meet current expectations because they extend shelf life largely by transforming the product rather than maintaining its fresh state. These limitations highlight the need for alternative or complementary technologies that can improve microbial safety and shelf life while better preserving the intrinsic sensory and nutritional qualities of seafood (Abel et al., 2021).

2.3. Chilled and frozen storage

Low-temperature storage is one of the most widely used strategies for delaying seafood spoilage and maintaining product quality. Chilled and frozen storage are extensively applied in the seafood industry because they reduce microbial proliferation and slow enzymatic and oxidative reactions associated with quality deterioration (Rezende et al., 2022; Bejaoui et al., 2021; Tavares et al., 2021). However, their effectiveness depends strongly on the initial quality of the raw material, rapid postharvest cooling, hygienic handling, and strict temperature control throughout the supply chain.

Chilled storage generally maintains seafood at low temperatures, typically near 0°C, although refrigeration conditions may vary depending on the product and distribution system. At these temperatures, the activity of endogenous enzymes and spoilage microorganisms is reduced, thereby delaying the onset of quality deterioration (Sharifian et al., 2014). However, refrigeration does not completely arrest microbial growth or enzymatic activity. Psychrotrophic spoilage bacteria and some cold-tolerant pathogens may continue to grow slowly during refrigerated storage, limiting the effectiveness of chilling as a long-term preservation strategy (Sharifian et al., 2014; Mao et al., 2023; Kontominas et al., 2021). Therefore, chilled storage is mainly suitable for short-term preservation and often requires complementary hurdles to improve microbial safety and shelf-life stability.

Frozen storage, typically at temperatures at or below −18°C, is used for long-term preservation of seafood. Freezing immobilizes a large proportion of water as ice, reduces water availability, and prevents microbial growth during storage, although it does not necessarily inactivate all microorganisms. It also markedly slows enzymatic and oxidative reactions, thereby helping to preserve the quality and nutritional value of seafood over extended periods (Rezende et al., 2022; Bejaoui et al., 2021; Syanya et al., 2023). Nevertheless, frozen storage does not completely prevent quality deterioration, particularly when storage temperatures fluctuate or when products are rich in polyunsaturated fatty acids (Tavares et al., 2021; Dawson et al., 2018).

A critical factor in frozen storage is the formation and control of ice crystals. Rapid freezing produces smaller ice crystals, causing less structural damage and better preserving texture, flavour, and overall product integrity (Brown & Dave, 2021; Sommers et al., 2011). In contrast, slow freezing promotes the formation of larger ice crystals, which may rupture cell membranes, increase drip loss during thawing, and negatively affect texture and juiciness (Brown & Dave, 2021; Bejaoui et al., 2021; Hassoun et al., 2020b). In addition, temperature fluctuations during storage and distribution can promote ice recrystallization, moisture migration, protein denaturation, lipid oxidation, freezer burn, and sensory deterioration, including changes in texture, colour, odour, and flavour (Dawson et al., 2018; Hassoun et al., 2020b; Fadiji et al., 2021).

Despite their widespread use, chilled and frozen storage have several limitations. Refrigeration only delays spoilage and cannot fully prevent psychrotrophic growth, enzymatic degradation, formation of volatile spoilage compounds, or lipid oxidation during extended storage (Tavares et al., 2021; Kontominas et al., 2021). Frozen storage provides longer shelf life but may still compromise fresh-like quality through ice crystal damage, drip loss after thawing, protein denaturation, lipid oxidation, and cold-chain-related quality losses. Moreover, both methods depend on an uninterrupted cold chain, which increases energy demand, operational costs, and vulnerability to temperature abuse during transportation, storage, and retail (Fadiji et al., 2021; Heard & Miller, 2016). Therefore, although chilled and frozen storage remain essential preservation strategies, they are not sufficient as stand-alone approaches to ensure microbial safety and fresh-like quality throughout the seafood supply chain. These limitations support the use of complementary preservation technologies, including non-thermal approaches such as CP.

2.4. Chemical preservatives

Chemical preservatives are substances, either synthetic or naturally derived, that are intentionally added to foods to delay spoilage, inhibit microbial growth, prevent undesirable biochemical reactions, and extend shelf life while maintaining product safety and quality (Gökoğlu, 2019). In seafood products, their application is particularly relevant because fish and shellfish are highly susceptible to rapid quality loss caused by microbial growth, endogenous enzymatic activity, and chemical oxidation, with microbial spoilage generally representing the dominant deterioration mechanism in fresh and lightly preserved products (DeWitt & Oliveira, 2016; Tavares et al., 2021; Huang et al., 2021). Chemical preservatives have therefore been widely used in seafood systems because they are generally cost-effective, easy to incorporate into existing processing operations, and effective in controlling microbial growth, delaying oxidation, suppressing enzymatic reactions, and reducing losses during storage and distribution (Gökoğlu, 2019; Giannakourou & Tsironi, 2021; Huang et al., 2021).

The most widely used chemical preservatives in seafood products include weak organic acids and their salts, sulphiting agents, nitrites/nitrates, and antioxidants (Syamdidi, 2012). Organic acids such as sorbic, benzoic, acetic, lactic, and citric acids inhibit microorganisms mainly by entering microbial cells in their undissociated form, lowering intracellular pH, and disrupting enzyme activity and energy metabolism. Sulphites are commonly applied in crustaceans to prevent melanosis through the inhibition of polyphenol oxidase, whereas nitrites and nitrates are used in certain processed seafood products to suppress anaerobic pathogens, particularly Clostridium botulinum. Antioxidants such as ascorbic acid, citric acid, tocopherols, butylated hydroxyanisole (BHA) and butylated hydroxytoluene (BHT) are used to retard lipid oxidation and preserve sensory quality, especially in fatty fish species. Therefore, the preservative action of these compounds is based on multiple mechanisms, including antimicrobial activity, enzyme inhibition, pH reduction, and oxidative stability improvement (Kontominas et al., 2021; Smyth et al., 2018; Wińska et al., 2019).

Despite their technological advantages, chemical preservatives are increasingly limited by consumer, regulatory, and efficacy-related concerns. The perception of synthetic preservatives as “chemical” or “artificial” has reduced consumer acceptance and increased demand for minimally processed and clean-label seafood products (Gökoğlu, 2019; Mei et al., 2019; Kontominas et al., 2021). In addition, some preservatives, particularly sulphiting agents, may be associated with hypersensitivity reactions and potential toxicity, leading to strict regulatory limits on their use (D’Amore et al., 2020). The efficacy of chemical preservatives may also be limited by the complex seafood matrix, the persistence of psychrotrophic spoilage microorganisms during refrigerated storage, and the product-dependent performance of organic acids and other antimicrobial treatments in fish systems (Smyth et al., 2018; Leyva Salas et al., 2017; DeWitt & Oliveira, 2016). Therefore, chemical preservatives alone may not fully satisfy current requirements for seafood safety, quality preservation, and consumer-driven clean-label expectations.

Growing consumer concerns regarding synthetic preservatives have shifted interest towards natural antimicrobial compounds as alternative or complementary strategies for seafood preservation. Plant extracts, essential oils, bacteriocins, organic acids, chitosan, and algae- or mushroom-derived compounds have been investigated for their antimicrobial and antioxidant potential in fish and shellfish products (Gökoğlu, 2019; Mei et al., 2019; Kontominas et al., 2021). Nevertheless, their practical application remains challenging because their composition and activity may vary depending on source, extraction method, purity, and processing conditions. In addition, many natural antimicrobials, particularly essential oils and plant extracts, may cause undesirable changes in odour, flavour, colour, or texture, which is a critical limitation for seafood products with delicate sensory characteristics (Tongnuanchan & Benjakul, 2014; Nieto, 2020; Kontominas et al., 2021). Their antimicrobial performance may also be reduced in complex seafood matrices compared with in vitro systems, and meaningful pathogen reduction often requires relatively high concentrations or combination with other preservation hurdles (Mei et al., 2019; Huang et al., 2021). Therefore, although natural preservatives align well with clean-label expectations, their limitations in terms of standardization, sensory acceptability, matrix-dependent efficacy, and limited microbial inactivation have driven the development of non-thermal, residue-free technologies such as CP.

2.5. Packaging

Packaging is an integral component of seafood preservation because it modulates the storage environment and influences both product quality and safety during shelf life. Packaging systems range from simple protective materials that limit exposure to oxygen, light, moisture, odours, microorganisms, and physical damage to more advanced technologies designed to extend shelf life through product-specific atmospheric conditions. This function is particularly important for highly perishable seafood products, which are susceptible to rapid microbial spoilage, lipid oxidation, moisture loss, and sensory deterioration (Kontominas et al., 2021). Although various active, intelligent, and functional packaging concepts have been developed, this section focuses on vacuum packaging, MAP, and sous-vide processing because these approaches are widely used in seafood preservation and provide a relevant framework for discussing the limitations of packaging-based strategies.

For most fish and seafood products, packaging strategies aim to reduce oxygen availability in order to slow aerobic spoilage and lipid oxidation. In vacuum packaging, air is removed from the package, and the low-oxygen environment is maintained using packaging materials with low gas permeability. This approach can delay the growth of aerobic spoilage microorganisms and reduce oxidative reactions. However, its effectiveness depends strongly on the product’s initial microbial quality, hygienic processing, appropriate packaging materials, and uninterrupted refrigeration. Moreover, reduced-oxygen environments may support the growth of facultative or obligate anaerobic microorganisms when temperature control is inadequate or additional hurdles are absent (Abel et al., 2021).

MAP involves replacing air in the package with a product-specific gas mixture, typically containing elevated CO₂ levels, reduced O₂, and, in some cases, N₂ as a filler gas. In seafood systems, CO₂ is the primary antimicrobial component because it dissolves into the product’s aqueous and lipid phases, forming carbonic acid, lowering pH, and inhibiting microbial growth, thereby extending shelf life (Gökoğlu, 2020). Nevertheless, MAP also has limitations. Excessive CO₂ absorption into the product may cause package collapse, drip formation, or undesirable visual changes, while insufficient gas control may reduce preservative effectiveness (Silbande et al., 2016). Therefore, gas composition, gas-to-product ratio, packaging material permeability, product fat content, and storage temperature must be carefully optimized for each seafood product.

A major safety concern associated with vacuum- and MAP-packaged seafood is the potential growth and toxin production of non-proteolytic Clostridium botulinum types B, E, and F under reduced-oxygen conditions, particularly when refrigeration is inadequate (Koukou et al., 2022). This risk is critical because botulinum toxin may be produced before obvious sensory spoilage becomes detectable. Reduced-oxygen packaging has also been associated with concerns related to Yersinia enterocolitica, Listeria monocytogenes, and histamine formation in susceptible fish products (European Food Safety Authority [EFSA], 2015). These risks indicate that packaging systems should not be considered standalone lethal treatments. Instead, their safe use requires high-quality raw materials, validated processing conditions, Hazard Analysis and Critical Control Point (HACCP)-based control, strict cold-chain management, and additional hurdles where necessary (Kontominas et al., 2021). Similarly, Gökoğlu (2020) emphasized that packaging techniques are generally insufficient when applied alone and should be combined with other preservation methods.

Sous-vide is a packaging-based preservation and preparation method that combines vacuum packaging with controlled, low-temperature heat treatment. This method is suitable for ready-to-eat seafood products because it provides convenience, reduces post-processing contamination, and helps maintain sensory quality during refrigerated storage. However, sous-vide converts the product into a cooked state and is therefore not applicable to the preservation of raw or fresh seafood. Moreover, because sous-vide involves a mild heat treatment combined with anaerobic packaging, careful control of time–temperature parameters and refrigerated storage is essential to ensure safety. For this reason, sous-vide products may require additional hurdles or complementary non-thermal technologies to improve pathogen control and enhance shelf-life stability (Mol et al., 2026; Coşansu et al., 2022; Onyeaka et al., 2022).

Overall, packaging-based preservation strategies play an important role in extending the shelf life and maintaining the quality of seafood products. However, they primarily modify the product environment rather than directly eliminating microorganisms. Their effectiveness depends strongly on raw material quality, packaging design, gas composition, storage temperature, and cold-chain integrity. Therefore, although vacuum packaging, MAP, and sous-vide processing are valuable tools in seafood preservation, they should be integrated with additional preservation hurdles to achieve effective pathogen control, minimize spoilage, and maintain fresh-like quality. This need for complementary strategies supports the application of non-thermal technologies such as CP, particularly in combination with packaging systems.

2.6. Innovative preservation technologies

A variety of innovative preservation technologies have been investigated to improve microbial safety and extend the shelf life of seafood products while minimizing the quality losses associated with conventional processing. These approaches include light-based treatments, ionizing irradiation, pulsed electric field (PEF), ohmic heating, ultrasound, high-pressure processing, ozone, and other non-thermal or minimally intensive technologies (Chacha et al., 2021). Although they differ in their mechanisms of action, these technologies share a common objective: achieving microbial control while preserving the sensory and nutritional quality of seafood products.

Light-based applications, such as ultraviolet-C (UV-C) and pulsed light, are attractive because of their short treatment times and ability to achieve surface decontamination. However, their effectiveness is often limited by low penetration depth, shadowing effects, treatment non-uniformity, and reduced performance on irregular, wet, or rough seafood surfaces. Ionizing irradiation technologies, including gamma rays, electron beams, and X-rays, provide deeper penetration and have demonstrated strong antimicrobial effectiveness. Nevertheless, their application may be constrained by consumer acceptance, regulatory requirements, specialized facility infrastructure, operational complexity, and cost (Monteiro et al., 2023).

PEF technology is another promising approach capable of microbial inactivation with relatively limited thermal impact, particularly in liquid or pumpable products. However, its performance is strongly influenced by product conductivity, matrix composition, electrode configuration, treatment chamber design, and process parameters, which may limit its consistent application across diverse seafood matrices (Ahmed et al., 2025). Ohmic heating provides rapid, relatively uniform volumetric heating compared with conventional methods; however, because it remains a heat-based process, it may still induce thermal quality changes. Moreover, variations in electrical conductivity among seafood products and the complexity of equipment design can restrict its practical implementation (Abel et al., 2021).

Ultrasound has been widely investigated because of its cavitation-induced mechanical effects and potential to enhance microbial inactivation or improve mass transfer. However, excessive ultrasound treatment may negatively affect product quality, and effective pathogen reduction often requires its combination with other hurdle technologies (Rathnayake et al., 2025). Similarly, other electrical discharge systems, including arc discharge, have been explored for microbial control, but their high energy requirements, limited process control, and possible thermal effects may restrict their suitability for delicate seafood products.

Overall, innovative preservation technologies offer valuable opportunities to improve seafood safety and quality. However, their limitations, including limited penetration, matrix dependency, treatment non-uniformity, equipment complexity, regulatory barriers, cost, and potential quality effects, indicate that no single technology can fully address the preservation challenges of seafood products. Within this context, CP represents a promising non-thermal approach because it enables microbial decontamination through reactive species generated at near-ambient temperatures and can be integrated into multi-hurdle preservation concepts, particularly in combination with packaging, natural antioxidants, plasma-activated liquids, or other non-thermal treatments (Kontominas et al., 2021).

3.1. Fundamentals and key advantages of CP

The term “plasma” was first introduced by Langmuir in 1928 to describe ionized gases, which are often referred to as the fourth state of matter (Rathod et al., 2021). Plasma is generated when sufficient energy is supplied to a gas, resulting in partial ionization and the formation of a reactive mixture containing electrons, ions, radicals, excited atoms and molecules, photons, and short- and long-lived reactive species. In CP, the system operates under non-equilibrium conditions, where electrons possess high kinetic energy while the overall gas temperature remains close to ambient levels. This feature enables the generation of chemically reactive species without substantial bulk heating, making CP particularly attractive for heat-sensitive food matrices such as seafood. As illustrated in Figure 1, CP can be generated in situ from electricity and working gases, producing reactive species under non-thermal conditions without the direct addition of conventional chemical preservatives.

In food applications, CP can be generated using various electrical discharge systems, including dielectric barrier discharge (DBD), corona discharge, radiofrequency discharge, microwave discharge, gliding arc discharge, and atmospheric-pressure plasma jets. These systems differ in energy input, plasma chemistry, reactive species generation, treatment configuration, and scalability. Among them, DBD and plasma jet systems are among the most widely investigated for food applications because of their operational flexibility, relatively simple configuration, and compatibility with direct, indirect, and in-package treatments (Aaliya et al., 2021; Bigi et al., 2023; Ojha et al., 2021).

The antimicrobial activity of CP is mainly attributed to the generation of reactive oxygen species (ROS) and reactive nitrogen species (RNS), together with other plasma-associated components such as charged particles, ultraviolet photons, and ozone. These agents can act individually or synergistically to damage microbial cell envelopes, intracellular proteins, nucleic acids, and other essential cellular structures, leading to microbial inactivation (Shintani et al., 2010). The effectiveness of CP depends on multiple process and product factors, including applied voltage, frequency, treatment time, gas composition, humidity, distance from the plasma source, post-treatment exposure time, package atmosphere, sample geometry, and surface characteristics (Olatunde et al., 2021).

One of the major advantages of CP is its application versatility. It can be applied directly to food surfaces, indirectly through plasma-treated gases or liquids, or inside sealed packages as in-package plasma treatment. Direct exposure can provide strong microbial inactivation because reactive species are generated close to the target surface; however, its effectiveness may be limited by surface irregularities and the short lifetime of some reactive species. Indirect approaches, including plasma-treated gases or plasma-activated liquids, may offer more uniform exposure for complex geometries but may exhibit reduced reactivity depending on the stability and transport of active species. In-package plasma is particularly attractive for seafood because it can reduce post-treatment recontamination risk and enables microbial control within the final package, thereby supporting integration with existing cold-chain and packaging systems (Zhang et al., 2017; Olatunde et al., 2021).

For seafood preservation, CP is especially relevant because it can be applied under near-ambient temperature conditions and therefore has the potential to improve microbial safety while better preserving fresh-like characteristics than conventional heat-based treatments. Under optimized conditions, CP may avoid cooking effects and minimize undesirable changes in sensory and nutritional quality. In addition, it does not require conventional chemical preservatives and can be generated using air or other low-cost gases, which supports its compatibility with clean-label and sustainability-oriented processing concepts (Ojha et al., 2021; Aaliya et al., 2021; Sarangapani et al., 2018). However, these advantages depend strongly on appropriate process optimization, since excessive treatment intensity or unsuitable plasma chemistry may promote lipid oxidation, protein modification, or sensory changes in sensitive seafood matrices.

The adjustability of key process parameters, such as voltage, frequency, treatment duration, gas composition, humidity, source-to-sample distance, and package atmosphere, allows CP treatments to be tailored to specific seafood products and target microorganisms. This process flexibility provides a basis for combining CP with other preservation technologies to achieve synergistic effects within multi-hurdle processing strategies. Therefore, CP should not be viewed only as a standalone decontamination method, but also as a platform technology that can be integrated with packaging, natural antimicrobials, antioxidants, PAW, or other non-thermal treatments to improve seafood safety and shelf-life stability (Olatunde et al., 2021).

3.2. Microbial inactivation mechanisms of CP in seafood matrices

Microbial contamination in seafood is frequently concentrated on external and exposed surfaces, including the skin, gills, shells, and cut muscle surfaces, due to post-harvest handling, processing, and storage conditions (Sheng & Wang, 2021). This surface-dominated contamination profile is particularly relevant for CP applications because plasma-generated reactive species act primarily at or near the treated surface. Therefore, despite its limited penetration depth, CP can be well matched to seafood decontamination when microbial targets are located on accessible surfaces. However, the irregular topography, high moisture content, mucus layer, organic residues, and tissue heterogeneity of seafood matrices may influence the transport, lifetime, and reactivity of plasma-generated species, thereby affecting antimicrobial performance.

The antimicrobial activity of CP is mainly attributed to the combined action of ROS, RNS, charged particles, electric fields, ultraviolet photons, and ozone. These components may act individually or synergistically to induce oxidative and structural damage in microbial cells (Shintani et al., 2010; Rathod et al., 2021). ROS such as atomic oxygen, hydroxyl radicals, singlet oxygen, superoxide, hydrogen peroxide, and ozone can oxidize membrane lipids, proteins, enzymes, and nucleic acids, leading to membrane permeabilization, leakage of intracellular components, metabolic disruption, and loss of viability (Rathod et al., 2021; Bourke et al., 2017). RNS, including nitric oxide and nitrogen oxides, may further contribute to microbial inactivation through nitrosative stress, intracellular acidification, protein modification, and nucleic acid damage (Rathod et al., 2021).

In seafood matrices, high surface moisture and water activity play an important role in plasma–microbe interactions. Moist surfaces can promote the formation and transfer of secondary reactive species through plasma–liquid interactions, including hydrogen peroxide, nitrite, nitrate, peroxynitrite-related species, and acidic products. These species may extend antimicrobial activity beyond the immediate plasma exposure period, particularly in indirect treatments or plasma-activated liquids. At the same time, organic matter, proteins, lipids, salts, and mucus may consume or quench reactive species, reducing their availability for microbial inactivation. Therefore, the seafood matrix can both support and limit plasma efficacy depending on product composition, surface condition, and treatment configuration.

Atmospheric CP inactivates microorganisms through multiple, interconnected mechanisms involving oxidative stress, UV-induced effects, electrostatic disruption, and biofilm matrix damage. The accumulation of charged particles can weaken cell integrity through electroporation, while the synergistic interaction of reactive species enhances cell membrane erosion and lesion formation, thereby increasing overall inactivation efficiency (Bigi et al., 2023; Chen et al., 2019; Coutinho et al., 2021). CP may also contribute to the disruption of microbial biofilms, which are important in seafood processing environments and on product-contact surfaces. Biofilm-associated cells are typically more resistant to conventional decontamination strategies because extracellular polymeric substances, reduced metabolic activity, biofilm architecture, and matrix composition can limit antimicrobial penetration. Plasma-generated reactive species, UV photons, and charged particles may damage both microbial cells and the extracellular biofilm matrix, thereby reducing biofilm viability and attachment (Nwabor et al., 2022; Puligundla & Mok, 2017; Rao et al., 2020). However, biofilm architecture, thickness, species composition, and matrix composition may still limit the penetration of reactive species and reduce treatment efficacy.

The extent of microbial inactivation is strongly influenced by process-related parameters, microbial characteristics, and product factors. Process variables such as applied voltage, frequency, treatment duration, working gas composition, humidity, source-to-sample distance, exposure mode, and package atmosphere directly influence the type and concentration of reactive species generated during treatment (Olatunde et al., 2021; Bigi et al., 2023). Microbial factors, including species, strain variability, Gram type, spore formation, physiological state, and biofilm formation, also determine susceptibility to plasma exposure. Gram-positive bacteria and bacterial spores are often more resistant than Gram-negative vegetative cells because of their thicker cell wall structures and protective layers (Bigi et al., 2023).

Product-related factors are particularly important in seafood applications. Surface roughness, tissue porosity, surface moisture, fat content, protein level, salt concentration, and the presence of skin, shell, mucus, or exudates may alter plasma distribution and reactive species availability. Rough, folded, or porous surfaces can shield microorganisms from direct plasma exposure, whereas high organic load may reduce antimicrobial efficiency by reacting with plasma-generated species before they reach microbial targets (Bourke et al., 2017; Olatunde et al., 2021). Consequently, microbial inactivation in seafood matrices is highly product-specific and requires careful optimization for each species, product format, and processing objective.

Overall, CP inactivates microorganisms in seafood through multiple, interconnected mechanisms involving oxidative stress, nitrosative stress, membrane disruption, intracellular damage, UV-associated effects, and biofilm matrix disruption. Its surface-oriented action makes it especially relevant for seafood products where contamination is predominantly located on exposed surfaces (Sheng & Wang, 2021; Olatunde et al., 2021). Nevertheless, treatment efficacy is governed by the interaction between plasma parameters, microbial resistance, and seafood matrix properties. Therefore, successful application requires product-specific optimization to achieve microbial safety while minimizing undesirable quality changes.

3.3. Impact of CP on quality attributes and oxidative stability of seafood

ROS and RNS generated during CP treatment may induce oxidative modifications in fish proteins. These reactive species can oxidize the protein backbone and amino acid side chains, leading to the formation of protein–protein cross-links or protein fragmentation. Such structural alterations may affect protein solubility, water-holding capacity, gel-forming ability, and digestibility, thereby inducing changes in the technological and sensory quality of seafood products. Moderate protein modification may improve some functional properties, such as gel formation and texture stability; however, excessive oxidation can result in protein degradation and reduced nutritional value. The extent of CP-induced protein oxidation depends on processing parameters such as treatment time, gas composition, applied voltage, and plasma type (Olatunde et al., 2021). Therefore, product-specific optimization of CP parameters is essential to minimize adverse quality changes while maximizing potential functional benefits. Under optimized conditions, quality deterioration may remain limited, whereas uncontrolled treatments may lead to increased protein oxidation.

In addition to protein oxidation, plasma-generated reactive species can induce various physicochemical changes in seafood products, potentially limiting the practical applicability of this technology if not properly controlled. Ozone, one of the reactive species produced during CP treatment with oxygen-containing gas mixtures, exhibits a high oxidation potential. This reactive species can initiate reactions that negatively affect meat quality, particularly colour attributes. Treatment time has been identified as a major factor influencing colour changes in CP-treated seafood, attributed to increased ROS and RNS generation with prolonged exposure (Olatunde et al., 2021). Accordingly, increased total colour difference (ΔE) values have been reported with longer treatment durations (Choi et al., 2016). Furthermore, protein modifications induced by plasma-generated reactive species may influence the water-holding capacity of treated seafood products. However, the effects of CP on water-holding capacity in seafood remain incompletely understood, highlighting the need for further investigation (Olatunde et al., 2021). For example, low-field proton nuclear magnetic resonance analysis showed that in-package DBD plasma treatment reduced “trapped water” within the myofibrillar network, suggesting that plasma treatment may influence water distribution and texture-related quality attributes in fish muscle (Paulsen et al., 2022).

During CP treatment, ROS and RNS may also promote oxidative conversion and structural modification of unsaturated fatty acids into lipid peroxides (Olatunde & Benjakul, 2022). Fish lipids are rich in polyunsaturated fatty acids, and their oxidation can adversely affect sensory properties (Gavahian et al., 2018). Therefore, lipid oxidation should be carefully considered during prolonged CP treatments, particularly in fatty fish species (Paulsen et al., 2022). As CP applications may accelerate lipid oxidation under certain conditions, process parameters must be optimized to ensure effective microbial inactivation while preserving quality and sensory attributes. In this context, natural antimicrobial or antioxidant compounds, such as plant extracts and essential oils, have been suggested as complementary strategies to mitigate oxidative and sensory deterioration (Olatunde et al., 2021).

Various reactions, particularly lipid oxidation, may negatively affect the sensory characteristics of CP-treated seafood products. Therefore, appropriate processing parameters and application conditions that induce minimal or negligible sensory changes must be selected to ensure microbial safety and extend shelf life. CP treatment may lead to the development of undesirable odours and flavours, such as rancid, oxidized, or overly fishy, especially in high-fat fish species, which can result in consumer rejection. Fishy odour is one of the primary sensory attributes leading to consumer disapproval and may be intensified by plasma-generated reactive species. Sensory properties of CP-treated seafood may be significantly affected under harsh processing conditions or when high concentrations of active species are generated, a phenomenon largely associated with accelerated lipid oxidation in seafood matrices (Olatunde & Benjakul, 2018).

A wide variety of CP systems have been developed, and most studies evaluate custom-designed experimental setups. Significant variability in treatment distance, exposure time, discharge configuration, gas composition, and energy input may lead to different quality outcomes across seafood matrices (Hajhoseini et al., 2020). Consequently, the effects of CP on seafood quality should be interpreted in relation to both processing conditions and product characteristics.

Overall, the effects of CP on the quality attributes and oxidative stability of seafood are directly linked to the intensity of reactive species, exposure duration, and the processing conditions used for microbial inactivation. Observed changes in protein oxidation, lipid oxidation, colour attributes, water distribution, and sensory quality indicate a delicate balance between effective microbial control and quality preservation. When process parameters are appropriately optimized, microbial safety can be improved while maintaining acceptable quality; however, uncontrolled or overly intensive treatments may adversely affect product acceptance. Therefore, CP should not be evaluated solely on the basis of antimicrobial efficacy but also on the controllability of its oxidative and sensory effects. These considerations support the use of CP as part of integrated preservation and multi-hurdle strategies rather than as a standalone intervention, particularly for oxidation-sensitive seafood products.

4.1. Chemical and quality-related limitations

Although CP is generally considered a non-thermal and residue-minimizing technology, its antimicrobial action is based on reactive oxygen and nitrogen species, which may also interact with seafood components. This represents one of the major limitations of CP in seafood processing, because fish and shellfish are highly susceptible to oxidative quality deterioration due to their high contents of polyunsaturated fatty acids and oxidation-sensitive proteins (Tavares et al., 2021). In particular, plasma-generated reactive species may accelerate lipid oxidation, leading to rancidity and the development of undesirable off-flavours and off-odours, especially in fatty fish species (Coutinho et al., 2018; Liao et al., 2020).

In addition to lipid oxidation, CP treatment may induce protein oxidation, colour changes, texture modification, and alterations in water-holding capacity, depending on the treatment intensity and seafood matrix. Such effects are influenced by processing parameters, including gas composition, applied voltage, treatment time, distance from the plasma source, and package atmosphere. Therefore, microbial inactivation and quality preservation must be carefully balanced, as conditions that improve antimicrobial efficacy may simultaneously increase oxidative or sensory deterioration.

Another important limitation is the limited predictability of chemical reactions occurring in complex seafood matrices. Interactions among reactive species, lipids, proteins, pigments, salts, endogenous enzymes, and surface exudates are not yet fully understood, which may complicate the interpretation and standardization of treatment outcomes (Bigi et al., 2023). Consequently, the chemical and quality-related effects of CP are highly product- and process-dependent. These limitations highlight the need for product-specific optimization and the use of complementary strategies, such as antioxidants, protective packaging, or combined hurdle approaches, to minimize oxidative damage while maintaining microbial safety.

4.2. Physical and technological limitations

CP technology is primarily effective at or near the treated surface because of the limited penetration capacity of plasma-generated reactive species. This restricts its ability to inactivate microorganisms embedded within internal tissue structures, protected by surface irregularities, or located beneath multi-layered microbial aggregates. This limitation becomes particularly important under high microbial load conditions, where overlapping cells, organic residues, or biofilm-like structures may shield underlying microorganisms, reducing overall decontamination efficiency (Coutinho et al., 2018; Speranza et al., 2021).

The physical characteristics of seafood products further complicate treatment uniformity. Porous, irregular, folded, or complex surfaces can hinder the homogeneous distribution of reactive species and create shadowing or shielding effects. In seafood systems, structural diversity such as scales, skin, shells, exoskeletons, gills, and differences among whole, filleted, sliced, or minced products may substantially influence plasma exposure and antimicrobial efficacy. Therefore, the effectiveness of CP applications depends heavily on product geometry, surface topography, matrix composition, microbial species, and environmental conditions.

From a technological perspective, CP may be less suitable for large-volume, thick, irregularly shaped, or structurally complex products when uniform treatment of all contaminated sites is required. Substantial variability among plasma generation systems, discharge configurations, operating gases, treatment distances, exposure times, and energy inputs also makes comparison across studies difficult and complicates process standardization (Speranza et al., 2021). These issues are particularly relevant for industrial-scale seafood applications, where reproducible treatment intensity, product throughput, operator safety, equipment design, and integration with packaging or cold-chain systems must be considered (Priyadarshini et al., 2019).

Although CP has shown promising antimicrobial effects across various food systems, its application in fish and seafood is less well validated than in other food matrices. Further seafood-specific studies are therefore needed to evaluate process scalability, treatment uniformity, equipment feasibility, and product-dependent efficacy under realistic processing and storage conditions. These physical and technological limitations indicate that CP should be optimized for each seafood product and, where necessary, combined with complementary preservation hurdles to achieve consistent microbial safety.

4.3. Process and industrial limitations

One of the major barriers to the industrial implementation of CP technology is scale-up. Although CP systems have shown promising results in laboratory conditions, developing equipment suitable for continuous, large-scale, and reproducible food processing remains challenging. Scale-up requires not only increasing treatment capacity but also maintaining uniform plasma distribution, consistent reactive species generation, and comparable antimicrobial efficacy across larger product surfaces or volumes (Andoni et al., 2021). This issue is particularly relevant for large or irregular seafood products, where achieving homogeneous plasma exposure across broad, uneven, or moist surfaces can be difficult.

System stability during long-term operation is another important technological limitation. Factors such as electrode degradation, gas contamination, component wear, reactor geometry, gas-flow dynamics, and fluctuations in operating conditions may affect plasma chemistry and process performance. As a result, treatment outcomes may vary among systems or even within the same system over prolonged operation. Although various reactor designs and process configurations have been proposed to improve scalability and treatment consistency, practical implementation in industrial seafood processing remains limited. Further research is therefore needed to improve system robustness, process monitoring, and reproducibility under realistic production conditions (Sasikumar et al., 2025).

Energy use, equipment complexity, and cost also influence industrial feasibility. Plasma generation requires controlled power input, gas handling, reactor design, and monitoring systems; therefore, industrial adoption depends on achieving an acceptable balance between microbial reduction, treatment time, energy efficiency, throughput, and product quality. The need for advanced power supplies, optimized reactor configurations, and precise control of voltage, frequency, gas composition, flow rate, humidity, and treatment distance may increase installation and operational costs. These requirements may be particularly challenging for small- and medium-sized seafood processors, limiting broader adoption unless simpler, robust, and cost-effective systems are developed (Tagrida et al., 2024; Sasikumar et al., 2025).

Beyond engineering and economic constraints, microbiological limitations should also be considered. CP may cause sublethal injury in some microorganisms, and exposure to non-thermal processes may induce a viable but non-culturable (VBNC) state under certain conditions. VBNC cells remain viable and metabolically active but may not be detected by conventional culture-based methods, leading to an underestimation of microbial risk in treated foods (Schottroff et al., 2018; Chakka et al., 2021). In addition, plasma susceptibility varies among microorganisms depending on species, strain, cell structure, physiological state, and biofilm formation; therefore, some spoilage or stress-adapted organisms may exhibit greater resistance than expected (Tagrida et al., 2024).

Overall, the industrial application of CP in seafood processing is limited not only by antimicrobial efficacy but also by scalability, system stability, energy efficiency, equipment cost, process monitoring, and microbiological validation. These limitations highlight the need for standardized reporting of plasma parameters, product-specific validation studies, and pilot-scale trials under realistic seafood processing and cold-chain conditions.

10.1. Oxidation-mitigating combinations

One major limitation of CP applications in seafood is that ROS and RNS, while contributing to microbial inactivation, may also induce lipid and protein oxidation. This issue is particularly critical in fish and shellfish rich in polyunsaturated fatty acids, where oxidative reactions may lead to discoloration, texture deterioration, sensory degradation, nutritional losses, and reduced shelf life. Therefore, combining CP with natural antioxidants, plant extracts, pigments, oligosaccharides, or other bioactive compounds has emerged as a key strategy to preserve antimicrobial efficacy while minimizing oxidative quality losses (Figure 2). As summarized in Table 2, antioxidant-supported CP combinations have mainly been designed to limit lipid and protein oxidation while maintaining microbial control and sensory quality in seafood products.

Figure 2

Source: Authors’ contribution, based on the literature cited in the manuscript.

The main objective of oxidation-mitigating combinations is to counterbalance plasma-induced oxidative stress without compromising microbial control. Several studies have demonstrated that antioxidant-supported plasma treatments can simultaneously delay microbial spoilage and chemical quality deterioration. For example, Gao et al. (2024) reported that PAW combined with coconut exocarp flavonoids (CF) delayed microbial spoilage, lipid oxidation, and total volatile basic nitrogen (TVB-N) accumulation, while improving colour and texture stability. Importantly, this study also emphasized the role of treatment sequence, showing that flavonoid pre-treatment followed by PAW application was more effective. Similarly, Seyedalangi et al. (2024) showed that CP combined with phycocyanin suppressed microbial growth, delayed the formation of TVB-N and trimethylamine nitrogen (TMA-N), enhanced antioxidant activity, and helped preserve fatty acids.

Natural antioxidant extracts have also been successfully used to reduce oxidation-related quality losses in crustaceans and fish. In Pacific white shrimp, Shiekh & Benjakul (2020) reported that Chamuang leaf extract (CLE) combined with CP reduced microbial load and melanosis, delayed increases in TVB-N, pH, thiobarbituric acid reactive substances (TBARS), and peroxide value, limited the oxidation of sensitive fatty acids such as EPA and DHA, preserved sensory quality, and extended shelf life to approximately 15 days. Similarly, in Asian sea bass, Olatunde et al. (2019) showed that CP combined with ascorbic acid and ethanolic coconut husk extract (ECHE) reduced microbial load while delaying protein oxidation and degradation, thereby preserving fatty acid profiles and extending shelf life. These findings indicate that natural antioxidants do not function merely as supportive additives but as critical quality-protective hurdles that counteract oxidative stress generated during plasma treatment.

Oligosaccharide-based combinations represent another important group of strategies to mitigate oxidation. Singh & Benjakul (2020) and Singh et al. (2020) demonstrated that CP combined with chitooligosaccharides (COS) reduced microbial load while decreasing TBARS, peroxide value, carbonyl content, and polyunsaturated fatty acids (PUFA) oxidation, thereby maintaining sensory acceptability. Similarly, Yu et al. (2023) reported that CP combined with chitosan oligosaccharide delayed increases in total viable counts, TVB-N, and pH while preserving moisture, water activity, and shelf life. These results suggest that chitooligosaccharides (COS) may provide dual antimicrobial and antioxidant activity, making them particularly suitable for seafood matrices that are vulnerable to oxidative deterioration.

Encapsulation-based strategies may further enhance the performance of antioxidant-plasma combinations. Olatunde et al. (2020a, 2020b) demonstrated that pre-treatment with liposome-encapsulated coconut husk extract reduced lipid and protein oxidation compared with plasma alone, prevented discoloration, and maintained sensory quality. In this context, liposomal encapsulation may improve the stability, dispersion, and effectiveness of bioactive compounds, thereby minimizing undesirable quality changes during plasma exposure.

The mechanistic basis of oxidation mitigation is supported by findings from model lipid systems. Wang et al. (2024a), using a fish oil model, showed that CP increased peroxide value, malondialdehyde formation, singlet oxygen generation, and unsaturated fatty acid loss. However, incorporating antioxidants such as curcumin, tea polyphenols, vitamin E, and β-carotene effectively reduced these oxidative changes, with vitamin E showing the strongest protective effect. These findings confirm that antioxidant systems can directly interfere with plasma-induced lipid oxidation pathways, particularly in PUFA-rich matrices.

Overall, oxidation-mitigating combinations demonstrate that the quality limitations of CP can be managed through the strategic use of antioxidants, plant extracts, phycocyanin, COS, and encapsulated bioactives. As summarized in Figure 2, these strategies primarily rely on natural antioxidants and bioactive compounds to scavenge reactive species, delay lipid and protein oxidation, preserve sensitive fatty acids, reduce discolouration, and maintain sensory acceptability. PAW systems may also contribute to oxidative stability depending on treatment conditions and application sequence. However, the success of these combinations depends strongly on matrix composition, antioxidant type, concentration, delivery system, and treatment order. Therefore, oxidation-mitigating combinations are particularly relevant for high-fat fish species, shrimp, and other PUFA-rich seafood products, where microbial safety must be achieved without accelerating oxidative quality loss.

10.2. Microbial inactivation-enhancing combinations

One of the major advantages of CP is its strong antimicrobial activity; however, a standalone application may not always achieve the desired level of microbial inactivation or may require prolonged treatment times. This can reduce process efficiency and increase the risk of oxidative or sensory deterioration. Therefore, combining CP with pre-treatment methods, non-thermal physical technologies, photonic applications, and chemically active agents has emerged as an important strategy to enhance microbial inactivation efficiency while limiting excessive plasma exposure. As shown in Table 3, combinations involving PEF, UV-based treatments, ultrasound, PAW, peracetic acid, and photocatalytic systems have been used to improve microbial inactivation through complementary stress mechanisms.

The primary objective of studies within this group is to increase microbial susceptibility to plasma-generated reactive species by weakening cellular structures or imposing additional stress. Pre-treatment technologies such as PEF are particularly relevant in this context because they can induce membrane permeabilization and structural damage, thereby facilitating the action of subsequent plasma treatment. Shiekh et al. (2021a, b) demonstrated that PEF pre-treatment enhanced the antimicrobial efficacy of CP in seafood systems by increasing microbial cell membrane permeability. In these studies, the inclusion of CLE also contributed to oxidative stability, delayed melanosis, reduced microbial load, and extended shelf life while maintaining product quality. Similarly, Ahmad et al. (2025) showed that a multi-hurdle strategy combining PEF, vacuum impregnation (VI), plant extracts, MAP, and CP provided superior preservation performance by suppressing microbial growth and supporting product stability.

Combinations with light-based or photonic technologies represent another important group of microbial inactivation-enhancing strategies. Colejo et al. (2018) reported that combined UV-C and CP treatment resulted in greater microbial inactivation than either treatment alone, suggesting that UV-induced cellular damage and plasma-generated reactive species can act in a complementary manner. However, high-intensity applications may lead to antagonistic effects or increased lipid oxidation, indicating that treatment intensity must be carefully optimized. Similarly, Zorzi et al. (2023) showed that UV-A, combined with plasma-enhanced reactive species, improved microbial inactivation, particularly in liquid systems.

Advanced antimicrobial and chemically active systems can further enhance plasma-based microbial inactivation through multi-target mechanisms. Lee et al. (2022) demonstrated that sequential UV-activated ZnO treatment followed by in-package CP significantly improved microbial inactivation compared with either treatment alone, highlighting the synergistic potential of combining photocatalytic and plasma-based mechanisms. Similarly, Zhao et al. (2021) reported that ultrasound, PAW, and peracetic acid increased the generation of reactive oxygen and nitrogen species and improved microbial inactivation. However, the authors also emphasized that combined treatments do not always produce proportional increases in microbial reduction, confirming the need for process optimization.

Overall, microbial inactivation-enhancing combinations improve CP performance by increasing cell membrane permeability, promoting reactive species generation, and imposing multi-target cellular damage. As summarized in Figure 2, these strategies primarily rely on non-thermal physical technologies, UV- and light-assisted treatments, and advanced antimicrobial or chemical systems. Although these approaches can improve decontamination efficiency and reduce the need for prolonged plasma exposure, their effectiveness depends on treatment sequence, intensity, matrix characteristics, and microbial resistance. Therefore, microbial inactivation-enhancing combinations should be optimized not only for maximum log reduction, but also for oxidative stability, sensory acceptability, and shelf-life performance in seafood systems.

10.3. Shelf-life-oriented and stability-enhancing strategies

In CP applications, achieving microbial inactivation alone is not sufficient; maintaining product stability after treatment, preventing recontamination, and minimizing quality deterioration during storage are also critical. For this reason, the combination of CP with MAP, biodegradable packaging films, edible coatings, and PAW has emerged as a promising strategy for extending the shelf life of seafood products. Shelf-life-oriented and stability-enhancing CP-based hurdle strategies reported for seafood products are summarized in Table 4.

The main objective of studies within this group is to preserve physicochemical and sensory stability after plasma treatment, slow microbial regrowth, and maintain product quality throughout storage. In particular, combinations involving MAP have attracted considerable attention. Tagrida and Benjakul (2022) and Tagrida et al. (2021) reported that CP combined with liposome-encapsulated plant extracts under MAP effectively suppressed microbial growth, maintained sensory acceptability, and extended the shelf life of tilapia samples to approximately 12 days. These studies demonstrate that packaging systems function not only as physical barriers but also as important components of multi-hurdle preservation strategies when integrated with plasma and bioactive compounds.

Edible coating and biopolymer-based systems also play an important role in maintaining storage stability. Jeon et al. (2022) demonstrated that in-package CP combined with whey protein isolate (WPI) coatings containing malic acid enhanced the inactivation and growth inhibition of Salmonella and Listeria monocytogenes during refrigerated storage. The study further showed that CP accelerated the diffusion of malic acid from the coating matrix, thereby improving antimicrobial efficacy. Similarly, Chaijan et al. (2022) reported that WPI–ginger extract coating combined with PAW reduced discolouration, drip loss, and textural deterioration while limiting lipid, protein, and myoglobin oxidation. These findings suggest that coating systems act not merely as passive barriers but as active quality-preserving systems.

Studies focusing on the optimization of packaging and storage conditions have also been reported. Qiu et al. (2026) evaluated different packaging systems following PAW pre-treatment and showed that MAP combinations with high CO₂ concentrations effectively slowed microbial growth, reduced TVB-N accumulation, and improved protein stability. However, excessively high CO₂ levels were also associated with package deformation and potential sensory defects. These findings highlight that shelf-life-oriented combination strategies should consider not only microbial safety but also packaging stability and consumer acceptability.

PAW systems have likewise gained attention in this context. Wang et al. (2024c) reported that the combination of PAW and CP delayed microbial growth, reduced Pseudomonas levels, improved water-holding capacity, and significantly extended shelf life. In contrast, Wang et al. (2024b) showed that PAW + CP treatment could increase certain volatile aroma compounds while simultaneously promoting lipid oxidation. These findings suggest that although PAW-based systems show considerable promise for improving storage stability, treatment conditions must be carefully optimized.

Chemically assisted storage systems also significantly enhance shelf life and product stability. Zhu et al. (2023) demonstrated that plasma-activated slightly acidic electrolyzed water (SAEW) enhanced antimicrobial efficacy while preserving colour and texture and limiting lipid oxidation. Likewise, Tahmouzi et al. (2025) reported that biodegradable films containing silver nanoparticles (AgNPs) improved barrier properties, reduced microbial load, and delayed the formation of TBARS and total volatile nitrogen (TVN) when combined with CP treatment. Such systems demonstrate the considerable potential of integrating active packaging technologies with plasma applications for future industrial seafood preservation.

Collectively, shelf-life-oriented and stability-enhancing combinations demonstrate that CP can be more effective when integrated with systems that continue to protect the product after treatment. As summarized in Figure 2, these strategies are predominantly associated with packaging technologies, edible coating/biopolymer systems, and PAW applications. Packaging systems and biodegradable films contribute to post-treatment stability by reducing recontamination and slowing microbial regrowth, while edible coatings act as both physical barriers and carriers of active antimicrobial or antioxidant compounds. Plasma treatment may further enhance the diffusion and effectiveness of these active agents, whereas PAW-based systems can support storage stability by improving microbial control and preserving physicochemical quality. Therefore, shelf-life-oriented CP combinations rely on complementary preservation mechanisms to maintain product quality and extend storage stability in seafood systems.