Role of taste receptors in salty taste perception of minerals and amino acids and developments in salt reduction strategies: A review

Figures & data

Figure 1. Depiction of tongue and taste buds in rats: (1) taste receptor cells within the papillae on the tongue and cross section of three types of papillae- circumvallate, foliate, fungiform. (2) Cross section of a taste bud with its components. (3) Cross section of ENaC channel present on the taste bud. Rats have 3 ENaC subunits, while in humans there is an additional ENaC-δ subunit present (adapted from McCaughey 2019; McCaughey and Scott 1998).

Figure 2. Transient receptor potential vanilloid-1 (TRPV1) [adapted from: Ahern et al. (2005), and Aroke et al. (2020)].

Table 1. Key receptors activated by minerals and amino acid and the taste qualities elicited by them.

Receptor family	Receptor	Stimuli	Taste qualities	References
Ion Channels	ENaC	Sodium salt	Salty taste, intensity dependent on the anion	Bigiani (2020); Mattes (1997); Behrens and Meyerhof (2015); McCaughey (2019); McCaughey and Scott (1998) Stähler et al. (2008)
Transient Receptor Potential (TRP) (Ion Channel)	TRPV-1 TRPV1 (and/or) TRPA1 (i.e., TRP ankyrin 1)	Harmful stimuli like capsaicin, protons, and noxious heat, Cations (Ca, Mg, K, Zn, Cu, Fe) Herbs and spices (TRPV1 agonists, e.g., capsaicin and resiniferatoxin, N-geranyl cyclopropyl carboxamide)	Salty, Sweet, Bitter, Umami Salty, Sweet, Bitter, Umami	Ahern et al. (2005); Lyall et al. (2004); Riera et al. (2009); Ishida et al. (2002) Rhyu, Kim, and Lyall (2021)
	TRPM5	Cations, particularly of mineral salts (extremely sensitive to calcium)	Sweet, Bitter, Umami	Talavera et al. (2008); Dutta Banik et al. (2018); Hofmann et al. (2003).
G Protein coupled receptors (GPCR)	Heterodimer T1R3/T1R2* T1R3	Sugar and sugar alcohols Amino acid (MSG) + nucleotides Calcium	Sweet Umami Salty/Sour/Chalky	Li et al. (2002); Nelson et al. (2001) Tordoff et al. (2012); Tordoff (1996)
	T1R1^+	Amino acid (MSG) + nucleotides	Umami	Nelson et al. (2002)
	TAS2R7	Divalent cations (calcium, magnesium, zinc, copper, manganese, aluminum)	Bitter	Wang et al. (2019)
	CaSR	Calcium ions Tripeptide glutathione (Glu-Cys-Gly)	Salty/Sour/Chalky Kokumi/Umami/Salty/Sweet (Majorly kokumi taste, however, might mildly elicit the other mentioned taste qualities)	Tordoff et al. (2008) Ahmad and Dalziel (2020); Ohsu et al. (2010)
	mGluR4 mGluR1	Glutamate	Umami	Chaudhari, Landin, and Roper (2000)
	Family C, subtype 6 A (hGPRC6A) (h = human)	L-amino acid	Umami^ (^known so far)	Pallante et al. (2021); Jørgensen and Bräuner-Osborne (2020)
Transmembrane channel like 4 (TMC4)	mTMC4, hTMC4 (m = mouse, h = human)	Chloride anion	Salty	Kasahara et al. (2022)

* T1R3 + T1R2 works as a heterodimer sweet receptor|⁺T1R1 + T1R3work as a heterodimer umami taste receptor.

Table 2. Available salt substitutes and their applications.

Author and year	Food matrix	Substitute	Sensory method	Results
Mineral salts, amino acid and their derivatives
Nakagawa et al. (2014)	Aqueous solutions	Mixture of NaCl (335 mM) + KCl (335 mM) + Sodium aspartate (1.7 mM)	Discrimination test (Binomial comparison method) to evaluate saltiness; untrained panel (n = 10)	Sodium aspartate enhanced the salty taste of solution containing both NaCl & KCl. Sodium aspartate reduced the bitterness of KCl. The study proposed that the ionic channels perceiving Na & K undergoes changes when both the ions are present in the same system, which is further changed in the presence of sodium aspartate, leading to an increase the perception of saltiness.
Vidal et al. (2020)	Low sodium salted meat	NaCl, KCl, (control 1: 100% NaCl and control 2: 25% NaCl + 75% KCl) & CaCl2. Seven treatments with 75% NaCl replacement by KCl + different flavor enhancers (Lysine, Taurine, Arginine, Inosinate + Guanylate, yeast extract, i.e., Bionis YE MXE NS, and Bionis SFE 201, Purac Arome NA4)	Overall acceptance and overall difference test.; untrained panel (n = 120).	Lysine and yeast extracts (Bionis YE MXE NS) led to the best results in reducing sensorial defects (predominantly bitterness) caused by KCl / CaCl2
Dos Santos Alves et al. (2017)	Low-fat Bologna-type sausages	KCl + amino acid (lysine) + flavor enhancers (50% IMP + 50% GMP), liquid smoke, other spices and herbs like black pepper coriander, garlic	Liking (9-point hedonic scale) and attribute rating (color, aroma, flavor, texture,); untrained panel (n = 100)	50% replacement of NaCl with KCl – no negative impact on the technological, physiochemical, and microbiological properties. Sensorial defects suppressed after the addition of lysine and/or liquid smoke.
Campagnol et al. (2011)	Fermented cooked sausages	KCl + amino acids (lysine, taurine), flavor enhancers 5’-ribonucleotide disodium inosinate and disodium guanylate (50% IMP + 50%GMP)	Liking (9-point hedonic scale) and attribute rating (color, aroma, taste and texture); untrained panel (n = 60).	50% replacement of NaCl with KCl to avoid manufacturing defects. Addition of amino acids and flavor enhancers compensated the sensory defects.
Mitchell, Brunton, and Wilkinson (2011)	Frozen ready meal (chilli con carne),	5 different combinations used: KCl, KCl + Lysine, KCl + autolyzed yeast, whole cell yeast, nucleotide yeast extract	Paired comparison & preference test (30 trained panelists), discrimination test (triangle test) (20 trained panelists), ranking test (15 trained panelists)	40% NaCl reduction achieved by gradual salt reduction in recipe. 60% NaCl reduction achieved by substituting NaCl with nucleotide yeast extract.
(Rocha et al. 2020)	Aqueous solutions	Flavor enhancer [disodium inosinate (IMP), monosodium glutamate (MSG), disodium guanylate (GMP), monoammonium glutamate (MAG)]	Temporal profile analysis (TI and TDS); Trained panel (n = 15)	MAG & MSG enhanced salty taste in sodium reduced solutions more effectively than IMP and GMP. These flavor enhancers elicited both salty and umami taste. Intensity of umami taste was majorly affected by the concentration of flavor enhancers.
Guo et al. (2020)	Reconstructed Ham	35.8% Na reduction + 0.2 to 0.8% Lysine	Liking (9-point hedonic scales) (mouthfeel, appearance, taste and global acceptance); trained panel (n = 13).	0.8% Lysine improved the water holding capacity, textural and sensory acceptance of low salt reconstructed ham. No significant increase in the perception of saltiness. i.e., addition of lysine increased the acceptance of the low salt foods but did not increase the saltiness perception.
da Silva et al. (2020)	Low-fat and low-sodium bologna-type sausages	KCl + Arginine & histidine (as flavor enhancers)	Liking (9-point hedonic scales) (color, aroma, flavor, texture, overall acceptance); untrained panel (n = 100)	Reduction of 60% NaCl by KCl, but KCl caused textural, sensorial defects and emulsion instability. Addition of arginine and histidine reduced these defects.
Grummer et al. (2012)	Cheddar cheese	KCl, MgCl2, or CaCl2	Descriptive Analysis; trained panel (n = 9)	CaCl2 and MgCl2 substitution resulted in detectable off notes like bitter, metallic, earthy, and soapy flavors. Cheddar cheese manufactured with KCl in combination with NaCl resulted in closest to the NaCl control cheese.
Horita et al. (2011)	Reduced-fat mortadella	CaCl2, MgCl2, KCl	Liking (9-point hedonic scale); untrained panel (n = 112)	Different combinations of these salts were used in addition to NaCl. Best sensory attributes were observed with 50% NaCl + 25% KCl + 25% CaCl2. CaCl2 had a negative impact on the emulsion stability and cooking yield. MgCl2 did not affect the emulsion stability or textural attribute, but the sensory attributes were compromised.
Herbs and Spices
Ghawi, Rowland, and Methven (2014)	Tomato soup	Basil modification- basil, black pepper, celery and garlic Cumin & coriander- cumin, coriander, ground celery seed and garlic Oregano modification-oregano, bay leaves, garlic, celery, and black pepper.	Profiling (trained panel; n = 10), Unstructured line-scale scoring of liking (overall liking, liking of appearance, flavor, texture, and aftertaste) & JAR (just-about-right flavor intensity), untrained panel (n = 148)	Direct incorporation of herbs and spices into a salt reduced tomato soups decreased consumer liking in comparison to the full salt soup. However, liking of the herb and spice reduced salt soups increased significantly over repeated tasting days (exposure) whereas liking for the reduced salt soups without herbs and spices did not. 53% reduction in salt achieved with enhanced perception of saltiness on using blend of herbs and spices (especially, the oregano blend).
Mitchell et al. (2013)	Chilled Ready meals: chicken supreme (a chicken and mushroom dish in a cream and wine sauce), a cottage pie (a savory minced beef dish topped with potato), and a chunky vegetable soup	Oregano, sage, rosemary, garlic, pimento, and clove. Lactoferrin whey proteins	Liking (9-point hedonic scale); untrained panel (n = 60)	Addition of these herbs lead to increased acceptability of salt reduced meals as they act as natural flavor enhancers. Off notes associated with whey proteins was not observed, there was increased consumer likeability. Can be used as a natural flavor enhancer.
Lopes et al. (2014)	Margarine	10 treatments with 1.5 g/100g spice mixture (A or B) and varied sodium content (0%, 25%, 50%, 75%, 100%) Mixture A: green onion, garlic, marjoram, thyme. Mixture B: lemon, oregano, basil, thyme (All spices in dehydrated form)	Liking Test (9-point hedonic scale); untrained panel (n = 62)	Addition of spice mix (A or B) increased the overall purchase intent, taste and flavor of margarine. Spice mix addition showed promising results with up to 50 (-75% sodium reduction Additionally, the spice mix (both A and B) also significantly increased the antioxidant activity of margarine
Wang, Lee, and Lee (2014)	Fresh chicken noodle soup, retorted canned chicken noodle soup, fresh tomato soup, canned tomato soup	Oregano, basil, marjoram, thyme, bay leaf, carrot, celery, onion	Overall liking (9-point hedonic scale), untrained panel (n1=95; n2=101)	Less salt was added in soups with herb blends and overall liking was increased, especially in fresh soups, the overall liking was not very significant for canned soups. But high levels of herbs reduced the overall acceptance of the soups.
Plant extracts
Lee (2011)	Plant aqueous extract	Magnolia, kukoshi, mushroom, mint, sea tangle, Salicornia (saltwort), ginger, yam, green onion, radish, onion, garlic, nappa cabbage	Intensity scale rating (0 to 15) of taste attributes; trained panel (n = 8)	Salicornia (saltwort), sea tangle, kukoshi were the three extracts scored high for umami and salty taste by the panel and was further selected for analysis. The sodium content of these plant extracts was 57% of that of NaCl (on dry weight basis).
Pires-Cabral, Pires-Cabral, and Quintas (2021)	Fermentation of white cabbage	Salicornia ramosissima; three fermentation trials; A (Control)- 80 g cabbage + 2.4 g NaCl (∼1.15% Na) B (with Salicornia).- 80 g cabbage + 24 g Salicornia (∼0.34% Na) C(salt + Salicornia)- 80 g cabbage + 0.4 g NaCl + 24 g Salicornia (∼0.49%Na)	No sensory	Salicornia substitution was effective in natural fermentation with desirable pH and acidity and no undesirable bacterial activity was observed. Conversely, it led to decrease in the population of fungi and coliforms to below detectable limits and had improved antioxidant capacity. Overall had a positive impact on the microbial and physical aspects of the fermented cabbage.
Others
Smith, Metzger, and Drake (2016)	Cream of broccoli soup	Whey permeates from cheddar, mozzarella, cottage cheese, milk and de-lactosed (DLC) permeates (with a salt and no salt control)	Descriptive Analysis; trained panel (n = 8)	Cottage cheese and de-lactosed (DLC) permeate was the saltiest. Cottage cheese has high lactic acid and lower lactose concentration. DLC had high mineral and organic acid content. However, they elicited other flavors (cardboard flavor, sour taste) in the soup, which was unacceptable by the consumers.
Zhang et al. (2014)	Aqueous extracts of seafood shells	Cuttle fish bone, clamshell, oyster shell, arabia cowry shell	Degree of difference test; semi trained panel (n = 20)	Cuttlefish bone saltiest among other shells. At same level of saltiness, its Na content was 73% of that of NaCl. Cuttlefish bone also had higher mineral (K, Mg, Ca, Zn, P) and amino acid (e.g., Lys., His., Arg.) content as compared to other shells. Note: This study also included some marine plants (Seaweed, kelp, semen cassiae, inula flower, sanna leaf.), however, they were not selected by the panelists during the screening process.
García-Lomillo et al. (2017)	Beef patties	Red wine pomace seasoning (2% w/w) added to control patties containing 2%, 1.5% and 1% (w/w salt/base-meat mass)	No sensory analysis	Red wine pomace seasoning reduced microbial growth, produced patties with higher K, Ca, fiber and phenol levels, enabled salt reduction without effecting the shelf life. Potential to be used as a flavor enhancer in salt reduced products while maintaining shelf life.
Desmond (2006)	Biscuits, snacks and savory dishes	Mycoscent (product derived from Mycoprotein)	Not available	50% sodium reduction achieved in biscuits and other snacks and 25% reduction in savory dishes. No information available on the physical or sensory properties of the products.

McCaughey, S. A. 2019. Dietary salt and flavour: Mechanisms of taste perception and physiological controls. In Reducing salt in foods. 2nd ed., ed. C. Beeren, K. Groves and P. M. Titoria, 45–70. Woodhead Publishing.

 Google Scholar

McCaughey, S. A., and T. R. Scott. 1998. The taste of sodium. Neuroscience & Biobehavioral Reviews 22 (5):663–76. doi:10.1016/S0149-7634(97)00067-5.

 PubMed Web of Science ®Google Scholar

Ahern, G. P., I. M. Brooks, R. L. Miyares, and X. b Wang. 2005. Extracellular cations sensitize and gate capsaicin receptor TRPV1 modulating pain signaling. The Journal of Neuroscience: The Official Journal of the Society for Neuroscience 25 (21):5109–16. doi:10.1523/jneurosci.0237-05.2005.

 PubMed Web of Science ®Google Scholar

Aroke, E. N., K. L. Powell-Roach, R. B. Jaime-Lara, M. Tesfaye, A. Roy, P. Jackson, and P. V. Joseph. 2020. Taste the pain: The role of TRP channels in pain and taste perception. International Journal of Molecular Sciences 21 (16):5929. doi:10.3390/ijms21165929.

 PubMed Web of Science ®Google Scholar

Bigiani, A. 2020. Does ENaC work as sodium taste receptor in humans? Nutrients 12 (4):1195. doi:10.3390/nu12041195.

 PubMed Web of Science ®Google Scholar

Mattes, R. D. 1997. The taste for salt in humans. The American Journal of Clinical Nutrition 65 (2 Suppl):692S–7S. doi:10.1093/ajcn/65.2.692s.

 PubMed Web of Science ®Google Scholar

Behrens, M., and W. Meyerhof. 2015. Taste receptors. In Flavour Development, Analysis and Perception in Food and Beverages, ed. J. K. Parker, J. S. Elmore, and L. Methven, 297–329. Woodhead Publishing.

 Google Scholar

Stähler, F., K. Riedel, S. Demgensky, K. Neumann, A. Dunkel, A. Täubert, B. Raab, M. Behrens, J.-D. Raguse, T. Hofmann, et al. 2008. A role of the epithelial sodium channel in human salt taste transduction? Chemosensory Perception 1 (1):78–90. doi:10.1007/s12078-008-9006-4.

 Web of Science ®Google Scholar

Lyall, V., G. L. Heck, A. K. Vinnikova, S. Ghosh, T. H. T. Phan, R. I. Alam, O. F. Russell, S. A. Malik, J. W. Bigbee, and J. A. DeSimone. 2004. The mammalian amiloride-insensitive non-specific salt taste receptor is a vanilloid receptor-1 variant. The Journal of Physiology 558 (Pt 1):147–59. doi:10.1113/jphysiol.2004.065656.

 PubMed Web of Science ®Google Scholar

Riera, C. E., H. Vogel, S. A. Simon, S. Damak, and J. Le Coutre. 2009. Sensory attributes of complex tasting divalent salts are mediated by TRPM5 and TRPV1 channels. The Journal of Neuroscience: The Official Journal of the Society for Neuroscience 29 (8):2654–62. doi:10.1523/jneurosci.4694-08.2009.

 PubMed Web of Science ®Google Scholar

Ishida, Y., S. Ugawa, T. Ueda, S. Murakami, and S. Shimada. 2002. Vanilloid receptor subtype-1 (VR1) is specifically localized to taste papillae. Brain Research. Molecular Brain Research 107 (1):17–22. doi:10.1016/S0169-328X(02)00441-2.

 PubMedGoogle Scholar

Rhyu, M.-R., Y. Kim, and V. Lyall. 2021. Interactions between chemesthesis and taste: Role of TRPA1 and TRPV1. International Journal of Molecular Sciences 22 (7):3360. doi:10.3390/ijms22073360.

 PubMed Web of Science ®Google Scholar

Talavera, K., K. Yasumatsu, R. Yoshida, R. F. Margolskee, T. Voets, Y. Ninomiya, and B. Nilius. 2008. "The taste transduction channel TRPM5 is a locus for bitter-sweet taste interactions. FASEB Journal: Official Publication of the Federation of American Societies for Experimental Biology 22 (5):1343–55. doi:10.1096/fj.07-9591com.

 PubMed Web of Science ®Google Scholar

Dutta Banik, D., L. E. Martin, M. Freichel, A.-M. Torregrossa, and K. F. Medler. 2018. TRPM4 and TRPM5 are both required for normal signaling in taste receptor cells. Proceedings of the National Academy of Sciences 115 (4):E772. doi:10.1073/pnas.1718802115.

 PubMed Web of Science ®Google Scholar

Hofmann, T., V. Chubanov, T. Gudermann, and C. Montell. 2003. TRPM5 is a voltage-modulated and Ca2+-activated monovalent selective cation channel. Current Biology 13 (13):1153–8. doi:10.1016/S0960-9822(03)00431-7.

 PubMed Web of Science ®Google Scholar

Li, X., L. Staszewski, H. Xu, K. Durick, M. Zoller, and E. Adler. 2002. Human receptors for sweet and umami taste. Proceedings of the National Academy of Sciences 99 (7):4692–6. doi:10.1073/pnas.072090199.

 PubMed Web of Science ®Google Scholar

Nelson, G., M. A. Hoon, J. Chandrashekar, Y. Zhang, N. J. Ryba, and C. S. Zuker. 2001. Mammalian sweet taste receptors. Cell 106 (3):381–90. doi:10.1016/s0092-8674(01)00451-2.

 PubMed Web of Science ®Google Scholar

Tordoff, M. G. L. K., Alarcón, S. Valmeki, and P. Jiang. 2012. T1R3: A human calcium taste receptor. Scientific Reports 2 (1):496. doi:10.1038/srep00496.

 PubMedGoogle Scholar

Tordoff, M. G. 1996. Some basic psychophysics of calcium salt solutions. Chemical Senses 21 (4):417–24. doi:10.1093/chemse/21.4.417.

 PubMed Web of Science ®Google Scholar

Nelson, G., J. Chandrashekar, M. A. Hoon, L. Feng, G. Zhao, N. J. P. Ryba, and C. S. Zuker. 2002. An amino-acid taste receptor. Nature 416 (6877):199–202. doi:10.1038/nature726.

 PubMed Web of Science ®Google Scholar

Wang, Y., A. L. Zajac, W. Lei, C. M. Christensen, R. F. Margolskee, C. Bouysset, J. Golebiowski, H. Zhao, S. Fiorucci, and P. Jiang. 2019. Metal ions activate the human taste receptor TAS2R7. Chemical Senses 44 (5):339–47. doi:10.1093/chemse/bjz024.

 PubMed Web of Science ®Google Scholar

Tordoff, M. G., H. Shao, L. K. Alarcón, R. F. Margolskee, B. Mosinger, A. A. Bachmanov, D. R. Reed, and S. McCaughey. 2008. Involvement of T1R3 in calcium-magnesium taste. Physiological Genomics 34 (3):338–48. doi:10.1152/physiolgenomics.90200.2008.

 PubMed Web of Science ®Google Scholar

Ahmad, R., and J. E. Dalziel. 2020. G protein-coupled receptors in taste physiology and pharmacology. Frontiers in Pharmacology 11 (1771):587664. doi:10.3389/fphar.2020.587664.

 PubMedGoogle Scholar

Ohsu, T., Y. Amino, H. Nagasaki, T. Yamanaka, S. Takeshita, T. Hatanaka, Y. Maruyama, N. Miyamura, and Y. Eto. 2010. Involvement of the calcium-sensing receptor in human taste perception. The Journal of Biological Chemistry 285 (2):1016–22. doi:10.1074/jbc.M109.029165.

 PubMed Web of Science ®Google Scholar

Chaudhari, N., A. M. Landin, and S. D. Roper. 2000. A metabotropic glutamate receptor variant functions as a taste receptor. Nature Neuroscience 3 (2):113–9. doi:10.1038/72053.

 PubMed Web of Science ®Google Scholar

Pallante, L., M. Malavolta, G. Grasso, A. Korfiati, S. Mavroudi, B. Mavkov, A. Kalogeras, C. Alexakos, V. Martos, D. Amoroso, et al. 2021. On the human taste perception: Molecular-level understanding empowered by computational methods. Trends in Food Science & Technology 116:445–59. doi:10.1016/j.tifs.2021.07.013.

 Web of Science ®Google Scholar

Jørgensen, C. V., and H. Bräuner-Osborne. 2020. Pharmacology and physiological function of the orphan GPRC6A receptor. Basic & Clinical Pharmacology & Toxicology 126 Suppl 6 (S6):77–87. doi:10.1111/bcpt.13397.

 PubMedGoogle Scholar

Kasahara, Y., M. Narukawa, A. Takeuchi, M. Tominaga, K. Abe, and T. Asakura. 2022. Molecular logic of salt taste reception in special reference to transmembrane channel-like 4 (TMC4). The Journal of Physiological Sciences: JPS 72 (1):31. doi:10.1186/s12576-022-00856-y.

 PubMed Web of Science ®Google Scholar

Nakagawa, T., J. Kohori, S. Koike, Y. Katsuragi, and T. Shoji. 2014. Sodium aspartate as a specific enhancer of salty taste perception-sodium aspartate is a possible candidate to decrease excessive intake of dietary salt. Chemical Senses 39 (9):781–6. doi:10.1093/chemse/bju051.

 PubMed Web of Science ®Google Scholar

Vidal, V. A. S., J. B. Santana, C. S. Paglarini, M. A. A. P. da Silva, M. Q. Freitas, E. A. Esmerino, A. G. Cruz, and M. A. R. Pollonio. 2020. Adding lysine and yeast extract improves sensory properties of low sodium salted meat. Meat Science 159:107911. doi:10.1016/j.meatsci.2019.107911.

 PubMed Web of Science ®Google Scholar

Dos Santos Alves, L. A. A., J. M. Lorenzo, C. A. Gonçalves, B. A. Dos Santos, R. T. Heck, A. J. Cichoski, and P. C. B. Campagnol. 2017. Impact of lysine and liquid smoke as flavor enhancers on the quality of low-fat Bologna-type sausages with 50% replacement of NaCl by KCl. Meat Science 123:50–6. doi:10.1016/j.meatsci.2016.09.001.

 PubMed Web of Science ®Google Scholar

Campagnol, P. C. B., B. A. dos Santos, M. A. Morgano, N. N. Terra, and M. A. R. Pollonio. 2011. Application of lysine, taurine, disodium inosinate and disodium guanylate in fermented cooked sausages with 50% replacement of NaCl by KCl. Meat Science 87 (3):239–43. doi:10.1016/j.meatsci.2010.10.018.

 PubMed Web of Science ®Google Scholar

Mitchell, M., N. P. Brunton, and M. G. Wilkinson. 2011. Current salt reduction strategies and their effect on sensory acceptability: A study with reduced salt ready-meals. European Food Research and Technology 232 (3):529–39. doi:10.1007/s00217-010-1420-6.

 Web of Science ®Google Scholar

Rocha, R. A. R., M. N. Ribeiro, G. A. Silva, L. C. R. Rocha, A. C. M. Pinheiro, C. A. Nunes, J. de, and D. S. Carneiro. 2020. Temporal profile of flavor enhancers MAG, MSG, GMP, and IMP, and their ability to enhance salty taste, in different reductions of sodium chloride. Journal of Food Science 85 (5):1565–75. doi:10.1111/1750-3841.15121.

 PubMed Web of Science ®Google Scholar

Guo, X., S. Tao, J. Pan, X. Lin, C. Ji, H. Liang, X. Dong, and S. Li. 2020. Effects of L-Lysine on the physiochemical properties and sensory characteristics of salt-reduced reconstructed ham. Meat Science 166:108133. doi:10.1016/j.meatsci.2020.108133.

 PubMed Web of Science ®Google Scholar

da Silva, S. L., J. M. Lorenzo, J. M. Machado, M. Manfio, A. J. Cichoski, L. L. M. Fries, M. A. Morgano, and P. C. B. Campagnol. 2020. Application of arginine and histidine to improve the technological and sensory properties of low-fat and low-sodium bologna-type sausages produced with high levels of KCl. Meat Science 159:107939. doi:10.1016/j.meatsci.2019.107939.

 PubMed Web of Science ®Google Scholar

Grummer, J., M. Karalus, K. Zhang, Z. Vickers, and T. C. Schoenfuss. 2012. Manufacture of reduced-sodium Cheddar-style cheese with mineral salt replacers. Journal of Dairy Science 95 (6):2830–9. doi:10.3168/jds.2011-4851.

 PubMed Web of Science ®Google Scholar

Horita, C. N., M. A. Morgano, R. M. S. Celeghini, and M. A. R. Pollonio. 2011. Physico-chemical and sensory properties of reduced-fat mortadella prepared with blends of calcium, magnesium and potassium chloride as partial substitutes for sodium chloride. Meat Science 89 (4):426–33. doi:10.1016/j.meatsci.2011.05.010.

 PubMed Web of Science ®Google Scholar

Ghawi, S. K., I. Rowland, and L. Methven. 2014. Enhancing consumer liking of low salt tomato soup over repeated exposure by herb and spice seasonings. Appetite 81:20–9. doi:10.1016/j.appet.2014.05.029.

 PubMed Web of Science ®Google Scholar

Mitchell, M., N. P. Brutnon, R. J. Fitzgerald, and M. G. Wilkinson. 2013. The use of herbs, spices, and whey proteins as natural flavor enhancers and their effect on the sensory acceptability of reduced-salt chilled ready-meals. Journal of Culinary Science & Technology 11 (3):222–40. doi:10.1080/15428052.2013.769869.

 Google Scholar

Lopes, C. d O., M. d F. P. Barcelos, N. A. A. Dias, J. d D. S. Carneiro, and W. C. de Abreu. 2014. Effect of the addition of spices on reducing the sodium content and increasing the antioxidant activity of margarine. LWT - Food Science and Technology 58 (1):63–70. doi:10.1016/j.lwt.2014.02.029.

 Web of Science ®Google Scholar

Wang, C., Y. Lee, and S.-Y. Lee. 2014. Consumer acceptance of model soup system with varying levels of herbs and salt. Journal of Food Science 79 (10):S2098–S2106. doi:10.1111/1750-3841.12637.

 PubMed Web of Science ®Google Scholar

Lee, G. H. 2011. A salt substitute with low sodium content from plant aqueous extracts. Food Research International 44 (2):537–43. doi:10.1016/j.foodres.2010.11.018.

 Web of Science ®Google Scholar

Pires-Cabral, P., P. Pires-Cabral, and C. Quintas. 2021. Salicornia ramosissima as a salt substitute in the fermentation of white cabbage. Journal of Food Science and Technology 59 (2):597–605. doi:10.1007/s13197-021-05047-y.

 PubMed Web of Science ®Google Scholar

Smith, S. T., L. Metzger, and M. A. Drake. 2016. Evaluation of whey, milk, and delactosed permeates as salt substitutes. Journal of Dairy Science 99 (11):8687–98. doi:10.3168/jds.2016-10904.

 PubMed Web of Science ®Google Scholar

Zhang, Y., Q. Cheng, Y. Yao, X. Guo, R. Wang, and Z. Peng. 2014. A preliminary study: Saltiness and sodium content of aqueous extracts from plants and marine animal shells. European Food Research and Technology 238 (4):565–71. doi:10.1007/s00217-013-2136-1.

 Web of Science ®Google Scholar

García-Lomillo, J., M. a Luisa González-SanJosé, R. Del Pino-García, M. a Dolores Rivero-Pérez, and P. Muñiz-Rodríguez. 2017. Alternative natural seasoning to improve the microbial stability of low-salt beef patties. Food Chemistry 227:122–8. doi:10.1016/j.foodchem.2017.01.070.

 PubMed Web of Science ®Google Scholar

Desmond, E. 2006. Reducing salt: A challenge for the meat industry. Meat Science 74 (1):188–96. doi:10.1016/j.meatsci.2006.04.014.

 PubMed Web of Science ®Google Scholar