Fermentation and Germination Improve Nutritional Value of Cereals and Legumes through Activation of Endogenous Enzymes
Open accessAbstract
Cereals and legumes are outstanding sources of macronutrients, micronutrients, phytochemicals, as well as antinutritional factors. These components present a complex system enabling interactions with different components within food matrices. The interactions result in insoluble complexes with reduced bioaccessibility of nutrients through binding and entrapment thereby limiting their release from food matrices. The interactions of nutrients with antinutritional factors are the main factor hindering nutrients release. Trypsin inhibitors and phytates inherent in cereals and legumes reduce protein digestibility and mineral release, respectively. Interaction of phytates and phenolic compounds with minerals is significant in cereals and legumes. Fermentation and germination are commonly used to disrupt these interactions and make nutrients and phytochemicals free and accessible to digestive enzymes. This paper presents a review on traditional fermentation and germination processes as a means to address myriad interactions through activation of endogenous enzymes such as α-amylase, pullulanase, phytase, and other glucosidases. These enzymes degrade antinutritional factors and break down complex macronutrients to their simple and more digestible forms.
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Fermentation promotes the microbial degradation of antinutritional compounds by producing phytases and proteolytic enzymes, while also increasing the release of bioactive compounds and antioxidant activity.
“Additionally, fermentation further promotes the microbial degradation of antinutritional compounds by producing phytases and proteolytic enzymes, while also increasing the release of bioactive compounds and antioxidant activity [34].”
From Valorization of Microalgae as Multifunctional Protein Sources in Plant-Based Systems: Structural, Functional, and Processing Challenges (2026) · cites this paper as [34]
The Protein section (page unavailable) describes proteolytic enzymes in fermenting L. plantarum and reductions in antinutritional compounds. The Minerals section says phytase degrades phytate, but the paper emphasizes activation of phytase already present in grains rather than microbial production of phytase. The Phytochemicals section reports release of bound compounds and potential increases in antioxidant properties, while noting that effects vary. (LLM, conf=0.85)
Experimental — accuracy not yet measuredGermination activates endogenous enzymes including phytases, proteases, and α-galactosidases, which degrade phytic acid, protease inhibitors, and raffinose-family oligosaccharides, thereby enhancing mineral bioavailability and protein digestibility.
“Germination activates endogenous enzymes, including phytases, proteases, and α-galactosidases, which degrade phytic acid, protease inhibitors, and raffinose-family oligosaccharides, thereby enhancing mineral bioavailability and protein digestibility [34,35].”
From Valorization of Microalgae as Multifunctional Protein Sources in Plant-Based Systems: Structural, Functional, and Processing Challenges (2026) · cites this paper as [34]
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“The increase could be due to leaching of the antinutritional factors that bind the minerals (Idris et al., 2007).”
BackgroundEffect of germination and fermentation on nutrients in maize flour
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BackgroundResultThe hepatic glycogenoreticular system
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Effect of fermentation and processing on in vitro mineral estimation of selected fermented foods.
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“It is therefore difficult to make conclusion based on these findings because they used different models and sources of β- carotene. β- Carotene bioavailability is dependent on genotype and processing method used to develop test food (Ortiz et al., 2017; Phorbee et al., 2013).”
EIF5A1 Isoform A Can Modulate the Metabolism in HeLa Cells
Pereira, K. D., Tamborlin, L., Meneguello, L., Bomfim, L. H. M., de Proença, A. R. G., Melo, C. F. O. R., Catharino, R. R., Silveira, L. R., & Luchessi, A. D. (2017) · The FASEB Journal · doi:10.1096/fasebj.31.1_supplement.lb208 · Cited by 13 · see citing works
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Plant food anti-nutritional factors and their reduction strategies: an overview
Samtiya, M., Aluko, R. E., & Dhewa, T. (2020) · Food Production Processing and Nutrition · doi:10.1186/s43014-020-0020-5 · Cited by 1,228 · see citing works
Microbial Fermentation and Its Role in Quality Improvement of Fermented Foods
Sharma, R., Garg, P., Kumar, P., Bhatia, S. K., & Kulshrestha, S. (2020) · Fermentation · doi:10.3390/fermentation6040106 · Cited by 824 · see citing works
“Natural fermentation promotes enzymatic degradation of phytates by providing optimum pH conditions present in a complex form with polyvalent cations such as zinc, iron, magnesium, and calcium [1].”
BackgroundInfluentialProtein Digestibility of Cereal Products
Joye, I. (2019) · Foods · doi:10.3390/foods8060199 · Cited by 343 · see citing works
Health-Promoting Components in Fermented Foods: An Up-to-Date Systematic Review
Melini, F., Melini, V., Luziatelli, F., Ficca, A. G., & Ruzzi, M. (2019) · Nutrients · doi:10.3390/nu11051189 · Cited by 395 · see citing works
Grains – a major source of sustainable protein for health
Poutanen, K. S., Kårlund, A., Gómez-Gallego, C., Johansson, D. P., Scheers, N., Marklinder, I., Eriksen, A. K., Silventoinen, P., Nordlund, E., Sözer, N., et al. (2021) · Nutrition Reviews · doi:10.1093/nutrit/nuab084 · Cited by 318 · see citing works
“Germination of oats and sorghum has also been shown to improve protein solubility,(87,88) and to improve protein digestibility, either by increasing protein solubility or by removal of antinutrients.(89) Exogenous enzymes are widely applied in food processing, eg, enzymatic crosslinking of oat proteins is used as a to…”
Fermentation of Cereals and Legumes: Impact on Nutritional Constituents and Nutrient Bioavailability
Adebo, J. A., Njobeh, P. B., Gbashi, S., Oyedeji, A. B., Ogundele, O. M., Oyeyinka, S. A., & Adebo, O. A. (2022) · Fermentation · doi:10.3390/fermentation8020063 · Cited by 283 · see citing works
“Because of these limitations, starter cultures such as LAB, yeasts, fungi, Bacillus species and other microorganisms have been isolated from fermented products and adopted to make the fermentation process more reliable, controlled and reproducible [1,9,10].”
BackgroundGraham, A. E., & Ledesma‐Amaro, R. (2023) · Nature Communications · doi:10.1038/s41467-023-37891-1 · Cited by 271 · see citing works
Fermented foods and gastrointestinal health: underlying mechanisms
Mukherjee, A., Breselge, S., Dimidi, E., Marco, M. L., & Cotter, P. D. (2023) · Nature Reviews Gastroenterology & Hepatology · doi:10.1038/s41575-023-00869-x · Cited by 264 · see citing works
“This effect is somewhat counteracted by fermentation-associated degradation of other mineral chelators such as oxalates and phytates 136 .”
Vetriventhan, M., Azevedo, V., Upadhyaya, H. D., Nirmalakumari, A., Kane‐Potaka, J., Anitha, S., Ceasar, S. A., Muthamilarasan, M., Bhat, B. V., Hariprasanna, K., et al. (2020) · The Nucleus · doi:10.1007/s13237-020-00322-3 · Cited by 222 · see citing works
Enhancing Micronutrients Bioavailability through Fermentation of Plant-Based Foods: A Concise Review
Samtiya, M., Aluko, R. E., Puniya, A. K., & Dhewa, T. (2021) · Fermentation · doi:10.3390/fermentation7020063 · Cited by 215 · see citing works
“However, plant protein digestibility increases during the fermentation process [40].”
BackgroundInfluentialGaland, S. G., Asensio‐Grau, A., Calvo‐Lerma, J., Heredia, A. B., & Andrés, A. (2021) · Food Research International · doi:10.1016/j.foodres.2021.110398 · Cited by 216 · see citing works
Sawant, S. S., Park, H.-Y., Sim, E.-Y., Kim, H., & Choi, H.-S. (2025) · Fermentation · doi:10.3390/fermentation11010015 · Cited by 238 · see citing works
“Fermentation offers a solution by degrading these complexes, thereby freeing minerals and making them more accessible [44].”
Anti‐nutritional compounds in pulses: Implications and alleviation methods
Kumar, Y., Basu, S., Goswami, D., Devi, M., Shivhare, U. S., & Vishwakarma, R. K. (2021) · Legume Science · doi:10.1002/leg3.111 · Cited by 195 · see citing works
“It has also been reported to be an alternative method for reduction of ANCs in pulses (Maleki & Razavi, 2020; Nkhata et al., 2018).”
MethodThe Application of Fermentation Technology in Traditional Chinese Medicine: A Review
Li, L., Wang, L., Fan, W., Jiang, Y., Zhang, C., Li, J., Peng, W., & Wu, C. (2020) · The American Journal of Chinese Medicine · doi:10.1142/s0192415x20500433 · Cited by 185 · see citing works
Cichońska, P., & Ziarno, M. (2021) · Microorganisms · doi:10.3390/microorganisms10010091 · Cited by 177 · see citing works
“Fermentation reduces the content of thermostable ANFs in legumes, which leads directly to an increase in digestibility of proteins and bioavailability of certain nutrients [77,78].”
BackgroundThe enormity of the zinc deficiency problem and available solutions; an overview
Khan, S. T., Malik, A., Al‐Warthan, A., & Shaik, M. R. (2021) · Arabian Journal of Chemistry · doi:10.1016/j.arabjc.2021.103668 · Cited by 169 · see citing works
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Fermented foods: An update on evidence-based health benefits and future perspectives
Diez‐Ozaeta, I., & Astiazaran, O. J. (2022) · Food Research International · doi:10.1016/j.foodres.2022.111133 · Cited by 168 · see citing works
Abbaspour, N. (2024) · Applied Food Research · doi:10.1016/j.afres.2024.100468 · Cited by 171 · see citing works
Two Faces of Fermented Foods—The Benefits and Threats of Its Consumption
Skowron, K., Budzyńska, A., Grudlewska‐Buda, K., Wiktorczyk-Kapischke, N., Andrzejewska, M., Wałecka-Zacharska, E., & Gospodarek-Komkowska, E. (2022) · Frontiers in Microbiology · doi:10.3389/fmicb.2022.845166 · Cited by 173 · see citing works
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Thakur, P., Kumar, K., Ahmed, N., Chauhan, D., Rizvi, Q. U. E. H., Jan, S., Singh, T. P., & Dhaliwal, H. S. (2021) · Current Research in Food Science · doi:10.1016/j.crfs.2021.11.019 · Cited by 142 · see citing works
Lisciani, S., Marconi, S., Donne, C. L., Camilli, E., Aguzzi, A., Gabrielli, P., Gambelli, L., Kunert, K. J., Marais, D., Vorster, J., et al. (2024) · Frontiers in Nutrition · doi:10.3389/fnut.2024.1385232 · Cited by 156 · see citing works
Fabbri, L. P., Cavallero, A., Vidotto, F., & Gabriele, M. (2024) · Foods · doi:10.3390/foods13213369 · Cited by 129 · see citing works
Microbial Fermentation for Improving the Sensory, Nutritional and Functional Attributes of Legumes
Senanayake, D., Torley, P. J., Chandrapala, J., & Terefe, N. S. (2023) · Fermentation · doi:10.3390/fermentation9070635 · Cited by 124 · see citing works
Rudzki, L., Stone, T. W., Maes, M. H. J., Misiak, B., Samochowiec, J., & Szulc, A. (2021) · Progress in Neuro-Psychopharmacology and Biological Psychiatry · doi:10.1016/j.pnpbp.2020.110240 · Cited by 123 · see citing works
Chinma, C. E., Abu, J. O., Asikwe, B. N., Sunday, T., & Adebo, O. A. (2020) · LWT · doi:10.1016/j.lwt.2020.110749 · Cited by 121 · see citing works
Current Functionality and Potential Improvements of Non-Alcoholic Fermented Cereal Beverages
Ignat, M. V., Salanță, L. C., Pop, O. L., Pop, C. R., Tofană, M., Mudura, E., Coldea, T. E., Borşa, A., & Pasqualone, A. (2020) · Foods · doi:10.3390/foods9081031 · Cited by 119 · see citing works
“Certain amino acids may be synthesized, and the availability of B group vitamins may be improved [99].”
BackgroundSustainable plant-based ingredients as wheat flour substitutes in bread making
Wang, Y., & Jian, C. (2022) · npj Science of Food · doi:10.1038/s41538-022-00163-1 · Cited by 131 · see citing works
Azeez, S., Chinma, C. E., Bassey, S. O., Eze, U. R., Makinde, A. F., Sakariyah, A. A., Okubanjo, S. S., Danbaba, N., & Adebo, O. A. (2021) · LWT · doi:10.1016/j.lwt.2021.112734 · Cited by 115 · see citing works
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Atudorei, D., Stroe, S.-G., & Codină, G. G. (2021) · Plants · doi:10.3390/plants10030592 · Cited by 116 · see citing works
“Each increase is attributed to the three processes—namely, the decrease in the amount of phytic acid, the activation of phytase and the release of minerals found in legumes in bound form (in the constitution of different complex substances) [65,68,70,73,74].”
BackgroundSiddik, M. A. B., Julien, B. B., Islam, S. M., & Francis, D. S. (2024) · Reviews in Aquaculture · doi:10.1111/raq.12894 · Cited by 125 · see citing works
Overview on pulse proteins for future foods: ingredient development and novel applications
Rajpurohit, B., & Li, Y. (2023) · Journal of Future Foods · doi:10.1016/j.jfutfo.2023.03.005 · Cited by 102 · see citing works
Using Pulses in Baked Products: Lights, Shadows, and Potential Solutions
Bresciani, A., & Marti, A. (2019) · Foods · doi:10.3390/foods8100451 · Cited by 101 · see citing works
The Use of Sprouts to Improve the Nutritional Value of Food Products: A Brief Review
Miyahira, R. F., Lopes, J. D. O., & Antunes, A. E. C. (2021) · Plant Foods for Human Nutrition · doi:10.1007/s11130-021-00888-6 · Cited by 104 · see citing works
Impact of processing and storage on protein digestibility and bioavailability of legumes
Gu, J., Amrit, B. K., Wu, H., Lu, P., Nawaz, M. A., Barrow, C. J., Dunshea, F. R., & Suleria, H. A. R. (2022) · Food Reviews International · doi:10.1080/87559129.2022.2039690 · Cited by 111 · see citing works
“The fermentation effects on proteins differ considerably may be attributed to different experimental aims, treatment times and differences in the original protein or amino acid profiles of the studies.([86]) Some studies found improvements in protein and amino acid contents after the fermentation, while some others re…”
BackgroundInfluentialEfficacy of germination and probiotic fermentation on underutilized cereal and millet grains
Budhwar, S., Sethi, K., & Chakraborty, M. (2020) · Food Production Processing and Nutrition · doi:10.1186/s43014-020-00026-w · Cited by 99 · see citing works
Kamau, E. H., Nkhata, S. G., & Ayua, E. (2020) · Food Science & Nutrition · doi:10.1002/fsn3.1473 · Cited by 98 · see citing works
“They are found embedded in food matrices thereby reducing their bioavailability (Nkhata et al., 2018).”
BackgroundInfluentialSelf-citationDhiman, S., Kaur, S., Thakur, B., Singh, P., & Tripathi, M. (2025) · Fermentation · doi:10.3390/fermentation11060346 · Cited by 108 · see citing works
Mishra, B., Mishra, A. K., Mohanta, Y. K., Yadavalli, R., Agrawal, D. C., Reddy, H. P., Gorrepati, R., Reddy, C. N., Mandal, S. K., Shamim, M. Z., et al. (2024) · Food Production Processing and Nutrition · doi:10.1186/s43014-023-00200-w · Cited by 96 · see citing works
“(Nkhata et al. 2018).”
Multiple Antioxidative and Bioactive Molecules of Oats (Avena sativa L.) in Human Health
Kim, I.-S., Hwang, C., Yang, W., & Kim, C. (2021) · Antioxidants · doi:10.3390/antiox10091454 · Cited by 95 · see citing works
Emkani, M., Oliete, B., & Saurel, R. (2021) · Foods · doi:10.3390/foods10030549 · Cited by 93 · see citing works
“However, selection of the right microorganism is necessary, since some microorganisms including yeasts and molds might concern food safety [30].”
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Duraiswamy, A., Nancy Mano Sneha, A., Sherina Jebakani, K., Selvaraj, S., Pramitha, L., Ramchander, S., Petchiammal, K. I., Sharmili, K. S., Thinakaran, J., Samundeswari, R., et al. (2023) · Frontiers in Plant Science · doi:10.3389/fpls.2022.1070398 · Cited by 94 · see citing works
“This is an effective process for lowering the anti-nutrient content in plant-based foods (Nkhata et al., 2018).”
BackgroundSoybean fermentation: Microbial ecology and starter culture technology
Elhalis, H., Chin, X. H., & Chow, Y. Y. S. (2023) · Critical Reviews in Food Science and Nutrition · doi:10.1080/10408398.2023.2188951 · Cited by 99 · see citing works
“During seed germination, the endogenous proteases, which were shown to be more active at 35 °C, were primarily responsible for the decline in trypsin inhibitor levels (Avilés-Gaxiola, Chuck-Hernández, and Serna Saldívar 2018; V. Kumar et al. 2006; Nkhata et al. 2018).”
Al-Qabba, M. M., El-Mowafy, M. A., Althwab, S. A., Alfheeaid, H. A., Aljutaily, T., & Barakat, H. (2020) · Nutrients · doi:10.3390/nu12102904 · Cited by 91 · see citing works
“The decomposition of phytochemicals, such as tannins, ascorbic acid, phytic acid, and antioxidant peptides might be the main reason behind the reduction in TPC and AOA after 6 days of germination [34].”
BackgroundMaldonado‐Alvarado, P., Pavón-Vargas, D. J., Abarca-Robles, J., Valencia‐Chamorro, S., & Haros, C. M. (2023) · Foods · doi:10.3390/foods12020389 · Cited by 87 · see citing works
“The effect of germination has been studied in cereals and pseudocereals [13,14].”
BackgroundYan, J., Xue, Q., Chen, W., Wang, K., Peng, D., Jiang, J., Li, P., & Du, B. (2022) · Food Research International · doi:10.1016/j.foodres.2022.111125 · Cited by 84 · see citing works
Reduction of FODMAP content by bioprocessing
Nyyssölä, A., Ellilä, S., Nordlund, E., & Poutanen, K. S. (2020) · Trends in Food Science & Technology · doi:10.1016/j.tifs.2020.03.004 · Cited by 80 · see citing works
The effects of ultrasound on the growth, nutritional quality and microbiological quality of sprouts
Liu, H., Li, Z., Zhang, X., Liu, Y., Hu, J., Yang, C., & Zhao, X. (2021) · Trends in Food Science & Technology · doi:10.1016/j.tifs.2021.02.065 · Cited by 80 · see citing works
Bera, I., O’Sullivan, M., Flynn, D., & Shields, D. C. (2023) · Molecules · doi:10.3390/molecules28073204 · Cited by 86 · see citing works
Food processing to reduce antinutrients in plant-based foods
Faizal, F. A., Ahmad, N. H., Yaacob, J. S., Lim, S. A. H., & Rahim, M. H. A. (2023) · International Food Research Journal · doi:10.47836/ifrj.30.1.02 · Cited by 80 · see citing works
“It is a common practice in countries such as South America and Africa, due to their dependency on agricultural products such as cereals and legumes as their staple foods (Nkhata et al., 2018).”
BackgroundSingh, P., Pandey, V. K., Sultan, Z., Singh, R., & Dar, A. H. (2023) · Journal of Agriculture and Food Research · doi:10.1016/j.jafr.2023.100902 · Cited by 79 · see citing works
Jikah, A. N., & Edo, G. I. (2023) · Journal of the Science of Food and Agriculture · doi:10.1002/jsfa.12892 · Cited by 81 · see citing works
Sustainable Strategies for Increasing Legume Consumption: Culinary and Educational Approaches
Amoah, I., Ascione, A., Muthanna, F. M. S., Feraco, A., Camajani, E., Gorini, S., Armani, A., Caprio, M., & Lombardo, M. (2023) · Foods · doi:10.3390/foods12112265 · Cited by 76 · see citing works
Abioye, V. F., Babarinde, G. O., Ogunlakin, G., Adejuyitan, J. A., Olatunde, S. J., & Abioye, A. O. (2022) · Heliyon · doi:10.1016/j.heliyon.2022.e12310 · Cited by 75 · see citing works
Craddock, J. C., Genoni, A., Strutt, E. F., & Goldman, D. (2021) · Current Nutrition Reports · doi:10.1007/s13668-020-00348-8 · Cited by 77 · see citing works
Nie, J., Fu, X., Wang, L., Xu, J., & Gao, X. (2022) · Food Chemistry · doi:10.1016/j.foodchem.2022.133990 · Cited by 74 · see citing works
Navyashree, N., Sengar, A. S., Sunil, C. K., & Natarajan, V. (2021) · Food Chemistry · doi:10.1016/j.foodchem.2021.131665 · Cited by 72 · see citing works
Lactic Acid Bacteria in Dairy Foods: Prime Sources of Antimicrobial Compounds
Ahansaz, N., Tarrah, A., Pakroo, S., Corich, V., & Giacomini, A. (2023) · Fermentation · doi:10.3390/fermentation9110964 · Cited by 71 · see citing works
The prospects of African yam bean: past and future importance
George, T. T., Obilana, A. O., & Oyeyinka, S. A. (2020) · Heliyon · doi:10.1016/j.heliyon.2020.e05458 · Cited by 72 · see citing works
Application of Probiotic Yeasts on Candida Species Associated Infection
Kunyeit, L., Anu‐Appaiah, K. A., & Rao, R. P. (2020) · Journal of Fungi · doi:10.3390/jof6040189 · Cited by 82 · see citing works
“For example, functional activity of microorganisms in food helps enhance the bio-availability of micronutrients, improving the sensory quality and shelf life of the food, degrading anti-nutritive factors (such as trypsin inhibitors and phytate degradation), enriching antioxidant and antimicrobial compounds, and fortif…”
BackgroundMesfin, N., Belay, A., & Amare, E. (2021) · Heliyon · doi:10.1016/j.heliyon.2021.e08081 · Cited by 68 · see citing works
Alrosan, M., Tan, T., Easa, A. M., Gammoh, S., & Alu'datt, M. H. (2022) · Food Chemistry · doi:10.1016/j.foodchem.2022.132386 · Cited by 68 · see citing works
Kohli, V., & Singha, S. (2024) · Discover Food · doi:10.1007/s44187-024-00076-w · Cited by 68 · see citing works
“The processing methods such as germination and fermentation, can also disrupt cellular integrity by reducing the compactness of the intracellular environment which may act as an extra obstruction for the passage of digestive enzymes [17, 22].”
Thompson, H. O., Önning, G., Holmgren, K., Strandler, H. S., & Hultberg, M. (2020) · Plant Foods for Human Nutrition · doi:10.1007/s11130-020-00806-2 · Cited by 65 · see citing works
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Plant-Based Meat Proteins: Processing, Nutrition Composition, and Future Prospects
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“…polyphenols, and minerals, reducing the content of antinutrients through activation of endogenous enzymes (α-amylase, pullulanase, and phytase), improving the digestibility of macronutrients, and increasing bioavailability of micronutrients and bioactives (Nkhata et al. 2018; Zhang et al. 2015).”
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- Authors
- Nkhata, S. G., Ayua, E., Kamau, E. H., & Shingiro, J.
- Affiliations
- Department of Agriculture Extension Services Lilongwe Malawi
- University of Eldoret (KE)
- Rwanda Agriculture Board (RW)
- Corresponding author
- Nkhata, Smith G.
- Venue
- Food Science & Nutrition
- Publisher
- Wiley
- Published
- 2018-10-16
- Volume
- 6
- Issue
- 8
- Pages
- 2446-2458
- ISSN
- 2048-7177 (ISSN-L)
- PMC
- PMC6261201
- OpenAlex
- W2896888197
- Cited by
- 876
- References
- 100
- Citations by year
- 2026: 124 · 2025: 143 · 2024: 159 · 2023: 154 · 2022: 129 · 2021: 103 · 2020: 57 · 2019: 7
- FWCI
- 4.25
- Type
- journal article
- Language
- en
- Open access
- hybrid
- doi.org
- Licence
- Locations
- Food Science & Nutrition (publishedVersion, cc-by)
- Europe PMC (PubMed Central) (publishedVersion, other-oa)
- PubMed Central (publishedVersion, cc-by)
- PubMed (publishedVersion)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion)
- Informationszentrum Lebenswissenschaften (ZB MED) (submittedVersion, cc-by)
- Topics
- Life Sciences › Agricultural and Biological Sciences › Plant Science › Phytase and its Applications
- Health Sciences › Nursing › Nutrition and Dietetics › Food composition and properties
- Life Sciences › Agricultural and Biological Sciences › Food Science › Food and Agricultural Sciences
- Keywords
- cereal fermentation, Fabaceae, antinutritional factors, phytic acid, trypsin inhibitor, phenolic compounds, mineral bioaccessibility, protein digestibility, endogenous enzymes, phytase, alpha-amylase, pullulanase, glucosidase, nutrient bioavailability, phytochemicals
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