Bacillus spp
Occurrence of Bacillus species has been reported on edible seaweeds and products, including cultivated kelp (Blikra et al. 2018; Lytou et al. 2021) and wild-harvested seaweed (Gupta et al. 2010) in Europe, and ready-to-eat (RTE) seaweed-based foods (Martelli et al. 2021). Although Bacillus levels on fresh seaweed are generally low, further growth may occur during improper handling and storage. Modelling studies of RTE kimbap (popular Korean dish made of cooked rice and various fillings in a sheet of nori) show that manual preparation and room-temperature storage strongly promote B. cereus proliferation (Park et al. 2005). Illness is linked to their toxin production, typically occurring above ~10⁵ colony forming units (CFU) per gram (Salkinoja-Salonen et al. 1999; Granum and Braid-Parker 2000). Bacillus spores are highly resistant, tolerating over 100 °C and pH below 3.0 for several minutes (Setlow 2006), though they cannot reproduce under such conditions. Once favourable conditions return, spores may germinate, grow, and produce toxins. After heat treatment, non-spore-forming bacteria are generally eliminated, and spore-forming bacteria can proliferate faster than in untreated products. For seaweed, similar precautions as for rice are recommended, i.e., cooked product should be cooled quickly to prevent outgrowth of B. cereus or Clostridium perfringens.
Vibrio spp
Pathogenic Vibrio species, i.e., V. parahaemolyticus and V. vulnificus, have been detected on both wild and cultivated seaweeds across different species and growing conditions (Mahmud et al. 2007; Mahmud et al. 2008; Musa and Wei 2008; Kudaka et al. 2008; Barberi et al. 2020). Their occurrence indicates a potential food safety risk when seaweeds are consumed raw and underscores the need for strict hygiene during post-harvest handling, particularly during summer when Vibrio levels peak. Vibrio alginolyticus, a potential but uncommon foodborne pathogen, was isolated from fresh A. esculenta cultivated in Scotland, but not from dried samples (Lytou et al. 2021), while no Vibrio spp. were found in wild P. palmata from Ireland (Moore et al. 2002). Vibrio parahaemolyticus and V. vulnificus are well-known causes of illness from other seafoods such as prawns and oysters (Honda and Iida 1993; Sumner and Ross 2002), but cases of food poisoning linked to Vibrio spp. in seaweed are extremely rare. Vibrio spp. are sensitive to thermal processing but may survive mild drying. For instance, low levels (below 10 CFU g-1) were found in low-temperature dried Ulva lactuca from Turkey, suggesting incomplete inactivation (Karacalar and Gamze 2008). Therefore, suitable handling procedures are crucial, particularly for seaweeds consumed raw or lightly processed.
Aeromonas spp
Aeromonas spp. are potential foodborne pathogens which may occur on seaweeds and survive or grow at chilled temperatures. Most studies have dealt with A. hydrofila, which have been implicated in many seafood-borne outbreaks, but evidence of seaweed-specific outbreaks is scarce. However, based on their indigenous aquatic prevalence, Aeromonas spp. could be expected to colonize seaweeds and possibly follow the raw material to processing. Occasional detections of Aeromonas spp. in e.g., air-dried Ulva reticulata from Malaysia (Vairappan and Suzuki 2000), and fresh Chondrus crispus and Chondracanthus teedii from Italy (Ziino et al. 2010) suggest the need for attention to hygiene and cold-chain handling.
Escherichia coli, Salmonella spp., Listeria monocytogenes, Staphylococcus aureus and other pathogens
Studies show variable contamination levels, with generally low counts in most regions but occasional detections near pollution sources. In Maine, USA, E. coli and Vibrio spp. were found on farmed S. latissima from multiple sites although at low levels (Barberi et al. 2020). The same study also reported the detection of Salmonella and enterohemorrhagic E. coli in several kelp samples subjected to a microbiological enrichment step (i.e., to detect microbes present in small numbers) prior to characterisation using molecular methods. Several European studies did not detect gastrointestinal pathogens on wild or farmed seaweeds (Liot et al. 1993; Moore et al. 2002; Gupta et al. 2010; Duinker et al. 2020). In Norway and Scotland, L. monocytogenes, Salmonella, E. coli, and S. aureus were largely absent (Blikra et al. 2018; Lytou et al. 2021), with one A. esculenta sample positive for L. monocytogenes likely contaminated during handling (Lytou et al. 2021). RTE products containing seaweed tend to show higher contamination rates, likely due to handling failures rather than the seaweed itself. Hygiene surveys of Korean kimbap frequently reported S. aureus, B. cereus, and occasionally Salmonella (Park et al. 2005; Cho et al. 2008; Kim et al. 2008).
Other potential hazardous microorganisms include:
Campylobacter jejuni and Yersinia enterocolitica: rarely detected but may grow under refrigeration upon contamination.
Clostridium spp.: occasionally detected in nori, but not the pathogenic species C. botulinum nor C. perfringens (Choi et al. 2014).
Shigella spp.: detected in a minority of kimbap samples using enrichment methods (Cho et al. 2008).
Yeasts and moulds: generally absent from fresh seaweed; may appear during prolonged storage of dried products (Stévant et al. 2020).
Viruses
Norovirus and Hepatitis A virus are the main foodborne viruses relevant to seaweed. Several outbreaks, in Asia (Kusumi et al. 2017; Sakon et al. 2018) linked to dried, shredded, or uncooked nori contaminated during processing or handling, and in Norway from a contaminated wakame salad (Folkehelseinstituttet 2019). Norovirus is environmentally robust and can remain infectious for months under dry conditions. Heat treatments ≥ 90 °C for longer than 90 seconds effectively inactivate enteric viruses, but post-processing contamination remains a key risk (Bosch et al. 2018). Refrigeration and freezing do not eliminate viral infectivity.
3.8. Allergens
Allergens in seaweeds
To date, seaweed protein is not among the established food allergens, and there is limited data on the allergenicity of seaweed proteins by themselves (Garciarena et al. 2022). Proteins extracted from Ulva sp. were annotated to known allergens using sequence similarity (Polikovsky et al. 2019), however, assessing the allergenic potential of novel proteins is a complex process, as complete allergens need to bind IgE‐antibodies, elicit an allergic reaction, and have de novo sensitisation capacity (Verhoeckx et al. 2019). On the other hand, seaweed farming infrastructure in the marine environment creates habitats for a variety of fish and shellfish species. The marine food allergens crustacean and mollusc tropomyosin and fish parvalbumin have previously been detected in edible seaweeds (Motoyama et al. 2007; Mildenberger et al. 2022). The European food regulation imposes the labelling of specified allergenic foods when used or added to food products and encourages the precautionary labelling of unintentionally occurring allergens (EU No 1169/2011). Therefore, to enable the broad inclusion of seaweeds in food products, the allergenic potential of seaweed-based ingredients must be better characterized in relation to different seasons and locations.
Regulation and product labelling
The European food regulation (EU No 1169/2011; article 36) also states that voluntary information should not be misleading or ambiguous, and should be based on scientific data to not unnecessarily restrict the available food products for allergenic customers. The Voluntary Incidental Trace Allergen Labelling (VITAL®) approach was developed over time to establish a program of risk-based precautionary allergen labelling (PAL) (Allergen Bureau 2024a). The key parameter in the current risk assessment framework (VITAL 4.0) is the eliciting dose (ED), specifically ED05, which is used as the reference dose. ED05 represents the amount of protein from a given allergenic food below which only 5% of the allergic population is expected to experience objective symptoms (Allergen Bureau 2024b). ED05 has been adopted in place of the more conservative ED01 used in the previous VITAL 3.0 framework with the aim to increase food choice for allergic consumers while maintaining a minimal risk to public health. With ED05, it is still extremely unlikely that life-threatening reactions occur, and allergic symptoms will be mild or moderate for most (>95%) of the 5 % of reacting allergic consumers (Allergen Bureau 2024b). VITAL 4.0 additionally includes a reference dose for molluscs, which was missing in the previous framework. Consumer exposure to allergenic proteins further depends on the quantity of the food typically consumed in a single eating occasion (referred to as the reference amount). Action levels (ALs) are defined to guide when precautionary allergen labelling (PAL) should be used. AL1 indicates that PAL is not recommended, while AL2 means that PAL is warranted. The cut-off between AL1 and AL2 corresponds to the amount of allergenic protein in a typical serving (reference amount) that would expose consumers to the reference dose (ED05), i.e., the level at which allergic reactions could occur (FAO and WHO 2023; Allergen Bureau 2024a). The recommended reference doses for seafood allergens from current and previous VITAL risk assessment frameworks, as well as AL transition points and the highest measured levels of this study are summarised in Table 6.