Ancient Grains

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1		Mahsa Majzoobi	--	4140	2023-06-21 10:24:36	\|
2	layout & references	Sirius Huang	Meta information modification	4140	2023-06-21 10:48:01	\|

This entry is adapted from the peer-reviewed paper 10.3390/foods12112213

In terms of genetic background, cereals are dived into “modern” and “ancient” cereals. Ancient grains were long-forgotten due to the dominance of modern grains, but have been rediscovered as highly nutritious, healthy and resilient grains for solving the nutrition demand and food supply chain problems.

sustainable grains combating hunger malnutrition ancient cereals

1. Ancient Grains

Ancient cereals are those species of grains that have not been subjected to any selection or breeding by humans and have maintained specific genetic properties from their wild ancestors, such as ear height, low harvest index, brittle rachis and brittle individual variation ^[1]. Ancient grains include varieties of wheat (Spelt, Khorasan wheat or Kamut, Einkorn and Emmer); green wheat, barley; wild rice, oats; sorghum; millets, and pseudocereals of teff, amaranth; buckwheat and quinoa. In some references, freekeh and bulgur have been considered ancient grains even though they are made from ordinary wheat ^[2].

Many ancient grains are ancestors of modern grains. For example, the crossing between a diploid species of chamois (Aegilops tauschii Coss.) and Emmer (ancient wheat) resulted in Spelt, which was mutated over several generations to convert into common wheat ^[3].

At the dawn of civilisation, ancient cereals used to provide a vital food source in the human diet. However, over the centuries, the selection of domesticated species with higher production yields and improved techno-functional properties has led to a dramatic decline in the production of other grains and the dominance of only a few grains—known as leading cereals—including wheat, rice, corn and barley ^[4]. This has generated significant food security concerns, especially with increasing the adverse impacts of climate change and supply chain disruptions due to the global pandemic and geopolitical and socioeconomic issues ^[5]. However, in recent years, ancient grains are regaining worldwide attention for a variety of reasons. The production of ancient grains is regarded as being environmentally friendly, generating low carbon footprints as they require less irrigation, pesticides and fertilisers compared to many normal grains. Ancient grains are also suitable for climate-smart agriculture since they can tolerate harsh growing conditions ^[6]. In addition, ancient grains are recognised as rich sources of nutrients and bioactive compounds with numerous health benefits ^[1]. Therefore, they are a key player in developing sustainable food systems and are well-positioned to tackle food insecurity caused by the ongoing climate change.

2. Physicochemical, Nutritional Profile and Health Benefits of the Ancient Grains

2.1. Wheat

Archaeological evidence shows that wheat most likely appeared first in Lebanon, Syria, Turkey, Egypt and Ethiopia. The domestication of wheat is likely to have begun around 10,000 years ago in the Fertile Crescent, and since then, wheat has been regarded as the most cultivated crop in the world ^[7]. The most common ancient wheat species include Einkorn (Triticum monococcum), Emmer (Triticum dicoccum), Khorasan (Triticum turgidum ssp. turanicum) and Spelt (Triticum spelta). Wheat grains have lengths mainly between 5 and 9 mm and shapes that may vary from spherical to flattened. The 1000-kernel weight of Spelt is ~44 g, which is much higher than that of Einkorn (~28 g) ^[8]. Einkorn and Emmer wheat are typically composed of 53–72% carbohydrates (mainly starch), 12.5–12.7% protein, 10.6–12.5% dietary fibre, 2.1% lipids and 1–3% minerals. The main interest in the worldwide adaptation of ancient wheat species could be related to their high protein contents and production yield and their high tolerance to many biotic and abiotic stresses. The high-yielding modern wheat produced by breeding programs often have lower protein content than ancient grains. Higher protein contents (~18%) in Einkorn wheat than other cultivars of Emmer (~15%) and Spelt (~13%) have been reported ^[3]. Ancient wheat species contain slightly lower carbohydrate contents than modern wheat. Within the ancient wheat group, Spelt and Einkorn have the lowest carbohydrate contents (~67–69%). The starch content of ancient wheat species is often lower than modern cultivars, and its composition varies greatly from modern wheat. For instance, Einkorn has lower resistant starch content (25.6 g/kg) than modern wheat (30–88 g/kg), whereas Spelt, Emmer and Einkorn contain 30–32% rapidly digestible starch, 26–59% slowly digestible starch and 2.3–2.4% resistant starch ^[9]. Einkorn showed a higher content of lipids compared to common wheat. Modern wheat varieties may have a rich content of mineral and dietary fibre compared to Einkorn and Emmer wheat. Ancient wheat species also contain fewer anti-nutrients than common wheat. The phytic acid contents of the Einkorn and Emmer wheat were between 1594 and 1863 mg/100 g ^[10].

A comparison between old and modern wheat cultivars showed higher health-relevant benefits of old cultivars. It has been indicated that the consumption of bakery products made with Khorasan wheat can enhance the immune functions in patients with severe symptoms and sleep disorders ^[3].

2.2. Green Wheat (Freekeh)

Premature green wheat, or freekeh, is an ancient whole grain with a history spanning thousands of years. Green wheat is produced from wheat harvested early, at the end of the milky stage, when culms and spikes are green. Grain shape, plumpness and greenness determine the quality of freekeh. Green wheat has a high initial moisture content that varies from 40–45% (wet basis), but during the drying process, it loses about 40% of its weight ^[11]^[12]^[13]. Depending on moisture content, kernel length, width and thickness differ from 6.24 to 6.66 mm, 3.65 to 4.22 mm and 3.43 to 3.85 mm, respectively. In addition, the mass of 1000 seeds varies from 15 to 51 g at different maturation stages ^[14]. To produce green wheat, often, immature durum wheat (Triticum durum) and, sometimes, immature bread wheat (Triticum aeisvum) are used. The Zenit and Diyarbakır spp. durum wheat is favoured for this purpose ^[11]^[15].

Green wheat is used as a raw material in the production of many foods and healthy drinks. Roasted green wheat, which is commonly known as freekah (also known as frekeh or frikah), has been a popular staple food in Middle Eastern, North African, and Chinese cuisines for centuries. Roasting improves the flavour of the grains; however, it causes huge losses to their nutritional quality ^[11].

Green wheat contains 73–80% carbohydrates, 11–15% protein and 12–19% dietary fibre. The starch content of green wheat is 45% and 68%, and its resistant starch content is about 8.0 to 10%. Due to its higher resistant starch and dietary fibre content and lower GI (52–54) compared to wheat, green wheat is more suitable for people with diabetes and for weight control.

Green wheat has a significantly greater proportion of essential amino acids, particularly lysine, methionine and threonine, and has better protein digestibility than normal wheat. Its total fatty-acid content varies from 1.32 to 2.7%, which is higher than that of yellow wheat. Palmitic acid is the dominant saturated fatty acid, and linoleic acid is the dominant unsaturated fatty acid ^[15].

The total mineral content in green wheat is higher than that in mature yellow wheat. Green wheat is a rich source of bioactive compounds. The total phenolic content, flavonoid content and antioxidant properties of green wheat are about twice that of wheat. Nevertheless, green wheat contains antinutrient compounds, such as phytate (660–700 mg/100 g), which is formed during the maturation of the seeds ^[11]^[15]. The freekeh grains harvested at earlier stages have the lowest phytic acid and phytate contents, which are nutritionally quite desirable ^[14]. The food applications of freekeh are limited to some traditional and homemade foods; however, due to increased knowledge about the nutritional and health benefits of green wheat, an increase in the global consumption of green wheat is expected. A few studies have shown the applications of green wheat in the formulation of healthy foods. For example, it has been found that the inclusion of green-wheat flour in the preparation of noodles can enhance the quality of the noodles and reduce their predicted GI ^[11].

2.3. Barley

Barley is a highly nutritious and adaptable ancient grain crop with growing cultivation all over the world. It is globally cultivated as the fourth most popular cereal in terms of production after wheat, rice and corn. Barley may have originated in Southeast Asia, including China, Tibet and Nepal ^[16]. There is limited information on the domestication of barley grains. A study reported the genome sequences of ancient barley grains excavated at Yoram Cave in the Judean Desert in Israel ^[17]. Barley grains are generally larger than wheat, with a 1000-kernel weight of about 40–45 g, and appear with a bright, light-yellow colour. Typical barley cultivars have distinct two-layered cells with adherent hulls departed at harvest maturity. However, hull-less varieties of barley have a low prevalence but are cultivated from certain seeds ^[18]. Today, more than 70% of barley grains are used for animal feed, about 20% are used for malting and brewing industries, and only a very small fraction is directly used in the human diet ^[16].

Generally, barley contains protein (10–17%), carbohydrates (~65–68%), lipids (2–4%), dietary fibres (18–22%), β-glucan (4–9%), minerals (1.5–2.5%) and vitamins (~2%). It contains ~14–20% rapidly digestible starch, ~20–25% slowly digestible starch and about 2.2% resistant starch that would help regulate the rate of glucose release in barley-containing foods in the human body. Barley kernel contains several bioactive compounds, including β-glucans, lignans, phytosterols and polyphenols. The relatively high β-glucan content present in barley helps to lower serum cholesterol levels and control blood glucose and insulin resistance. Barley is also a good potential source of a range of vitamins, including B1 (0.35 mg/100 g), B2 (0.091 mg/100 g) and E (0.85–3.15 mg/100 g). More recently, research has focused on the nutritional profiles of germinated barley grains as a food ingredient that could be rich in antioxidant compounds useful in functional food applications ^[19].

2.4. Oats

Oat (Avena L., Poaceae family) is a valuable cereal crop in many countries with a primary usage for animal feed, but due to its health benefits, its food applications are growing rapidly. However, the world production of oat for human food is still lower than other grains due to the lower yield and high cost of production and transport (due to the low density of oat grains) ^[11]. Oats have a 1000-kernel weight of about 34–35 g and have been grown from ancient times in many parts of the world, particularly in Northern and Eastern Europe ^[20].

Oats contain carbohydrates (75–80%), protein (10–15%), lipids (3–8%) and β-glucan (4%). In contrast to other cereals, a distinguished feature of oat grains is their high protein content and distinct and balanced amino acid composition. The amino acid composition of oat grains is superior to that of other cereals because its major storage protein is globulin, with higher concentrations of essential amino acids such as lysine than other cereals ^[21]. Oats are rich in carbohydrates, including ~60% starch with about 15% rapidly digestible starch, 8–9% slowly digestible starch and 76% resistant starch ^[22].

2.5. Sorghum

Sorghum is a drought-tolerant cereal belonging to the Poaceae grass family and originating in the northeast quadrant of Africa. It is the world’s fifth most important cereal after wheat, rice, maize and barley, with over 58.7 million tons of total production in 2020. The United States is the most significant producer of this crop, followed by Nigeria, Ethiopia, India, Mexico and China ^[23]^[24]. Sorghum is a very genetically diverse crop, with over 24 diverse species identified to date. Notable among these is S. bicolor, known for its food use and considered one of the most important species in modern commercial breeding programs. S. bicolor originated from its wild progenitor Sorghum bicolor L. Moench subsp. Verticilliflorum. Sorghum bicolor (L.) Moench is categorised into five major races: bicolor (the primitive type), guinea, caudatum, kafir and durra with various physical and biochemical properties ^[25]. Sorghum varieties have been classified based on different characteristics. However, based on the end-use applications, sorghum is classified into five groups, including sweet sorghum (syrup and biofuel), grain (biofuel, human food and animal feed), fibre, forage/fodder (animal feed) and broomcorn (broom-making) ^[24].

Sorghum has small seeds with pigmented pericarp, and the most commercially available varieties are black, white and red ^[26]^[27]. White sorghum is used for food products, while red sorghum is utilised primarily in the alcohol distillation industry ^[28]. Sorghum grains are ovoid with one end more pointed; the grain diameter ranges between 4 and 8 mm, and the mean weight of 1000 grains varies from 20 to 60 g. Starch is the main component of sorghum (about 70%); however, sorghum grains show the highest content of resistant starch (4–21%) and lowest starch digestibility (~19–37% rapidly digestible starch) and glycemic index among cereal crops ^[29].

The major protein fractions in sorghum are prolamins (kafirins), followed by glutelins; however, it has a low content of essential amino acids such as lysine, methionine and isoleucine ^[30].

The lipid in sorghum grains is made up of saturated fats and a high concentration of unsaturated fatty acids. Sorghum, especially red sorghum, is a rich source of various phytochemicals, mainly phenolic acid (mostly ferulic acid), flavonoids and tannins, with substantial health-promoting effects.

Sorghum grains, especially pigmented grains, have limited applications as human foods due to the presence of condensed tannins contributing to bitter taste, phytates, cyanogenic glycosides and trypsin inhibitors, which are considered the major antinutritional factors. However, varying food-processing methods such as sprouting, cooking, fermentation, steaming and flaking can reduce the antioxidants in sorghum ^[30]. In addition, low-tannin sorghum varieties have been identified and bred that have been used as an alternative for corn to feed animals ^[31]. It is also possible to reduce the tannin content of sorghum using food-processing methods such as milling followed by soaking in 0.3% Na₂CO₃ solution for 8 h ^[32]. Novel applications of sorghum include the production of plant-based protein, healthy foods and gluten-free products, and ethanol and biofuel production has emerged ^[33]. The digestibility of sorghum starch has been shown to vary dependent upon variety and may therefore be a useful flour-based ingredient for the optimisation of the glycaemic index of starch-based foods ^[34].

2.6. Millet

Millets are small-seeded species of cereal crops belonging to the family Poaceae, which originated in the arid and semi-arid regions of Asia and Africa. It has a short growing season and is resistant to pests and diseases. Millet has five genera: Panicum, Setaria, Echinochloa, Pennisetum and Paspalum ^[16]. The most important cultivated varieties of millets are foxtail millet (Setaria italica), pearl millet (Pennisetum glaucum), proso millet (Panicum miliaceum), barnyard millet (Echinochola crusgalli), finger millet (Eleusine coracana), brown top millet (Panicum ramosum), kodo millet (Paspalum scrobiculatum) and teff millet (Eragrostis tef). Millet is the sixth most high-yielding grain in the world, with a total annual production of 30.4 million tons, but is still considered an underutilised grain ^[16]^[23]. Millet seeds have small and round shapes with different colours. The seed size varies between 3 and 4 mm, the 1000-kernel weight of millet varieties is about 2.5–3.0 g and the bulk density and true density are about 0.67–0.55 gmL⁻¹ and 1.36–1.79 gmL⁻¹, respectively ^[16]^[35]. The main constituent of millet is its starch (62 to 70%), and some reports indicated that millet contains about 4–5% resistant starch, 6–7% slowly digestible starch and ~10–11% rapidly digestible starch (RDS) ^[36]. The second major component of millet is protein. The amino acid profile of pearl millet is better than that of sorghum and maize, and is comparable to that of wheat, barley and rice, and lysine is the first limited amino acid in millet cultivars ^[37]. Among millets, finger millet is relatively better balanced in essential amino acids because it contains more lysine, threonine and valine. The crude fat content in finger millet has been reported in the range of 1.54 to 3.77%. Linolenic acid and oleic acid are the two dominant fatty acids in the millet varieties ^[38].

Among millets, finger millet is the richest source of calcium and iron, with levels higher than those of sorghum, barley, maize and wheat. Millet grains are rich in several phytochemicals, particularly phenolic compounds. Finger millet has been shown to have the highest phenolic content and antioxidant activities compared to proso and foxtail millets ^[38]. Millets also have antinutrients, such as phytic acid (296–620 mg/100 g), tannins (31–343 mg/100 g) and trypsin inhibitors, which may reduce the bioavailability of minerals ^[39]. Millets are often subjected to different processing methods such as dehulling, decortication, soaking, germination, malting, milling, cooking, roasting, popping, radiation and fermentation to improve the nutritional and sensory properties of millets for developing new food (Xiu et al., 2022). Millet has some food applications, including the production of gluten-free foods, bakery products and porridge ^[39]^[40].

2.7. Wild Rice

Wild rice, known as a health-promoting grain, is the seed of an aquatic plant belonging to the genus Zizania, family Poaceae ^[41]. Wild rice (Zizania spp.) originated from North America over 10,000 years ago and then dispersed into East Asia and other parts of the world ^[42]. It consists of four species: Zizania palustris L., Zizania aquatica L., Zizania texana H. and Zizania latifolia G ^[43].

The seeds of wild rice have long and narrow cylindrical shapes approximately 4.7 to 9.2 mm long and 1.6–2.8 mm wide. The grain colour of these wild rice varies from light red–brown to dark brown with a 1000-kernel weight of 23–37 g ^[42]. Wild rice is rich in minerals, vitamins, starch, dietary fibre, protein and antioxidant phytochemicals, and is low in fat. Wild rice contains about 56–79% starch as the main constituent. Wild rice starch has shorter chains of amylose and longer chains of amylopectin, which causes a slower in vitro digestion rate compared to that of domesticated rice. It contains about 60% rapidly digestible starch, ~4% slowly digestible starch and ~5% resistant starch ^[41]^[43]. The resistant starch content of the wild rice is about 10.8%, which is significantly higher than white rice (~1.4%) and red rice (~0.95%). It also contains about 6.8% dietary fibre content, which is considerably higher than that of red rice (~2.6%) and white rice (~0.42%) ^[44].

Protein (10–15.5%) is the second main constituent of wild rice, which is much higher in content and efficiency ratio than that in white rice (~10%) and red rice (~11%). The essential amino-acid profile of wild rice is generally better and more balanced than that of other grains. Threonine and lysine are the limiting amino acids in all varieties of wild rice ^[42]^[43].

As a whole grain, wild rice is a rich source of phenolic compounds and flavonoids, and this level of antioxidant phenolic compounds is 10–15 times higher than that of white rice. Ferulic acid is the predominant phenolic acid, followed by sinapic acid and p-coumaric acid. Other phytochemical constituents of wild rice are flavonoid glycosides and flavan-3-ols. In addition to phenolic compounds, anthocyanins and carotenoids such as lutein were found in wild rice, thus providing a more complete profile of the antioxidants in wild rice ^[42]^[43]^[44]. Traditionally, wild rice has been exploited to treat a variety of ailments in Chinese medicinal practice ^[41].

2.8. Amaranth

Amaranth (Amaranthus spp.), a pseudocereal and a member of the Amaranthaceae family, is a less explored species with an excellent nutritional profile for human consumption. Amaranth has a diverse range of 60 species but has three common species (Amaranth hypochodriacus, Amaranth cruentus and Amaranth caudatus) domesticated for their seeds ^[45]. China is the largest producer of amaranth in the world, followed by the United States, Canada and Argentina. Owing to its high nutritional quality, such as balanced content of essential amino acids and unsaturated fatty acids, as well as being gluten-free, amaranth is gaining importance among consumers, food producers and the scientific community ^[46]. Amaranth protein contains a high amount of lysine, which is a limited amino acid in almost all cereals and other pseudocereal grains ^[47]. Its protein is also abundant in cysteine and methionine, two essential amino acids that contain sulphur. The Amaranthus species is recognised as a source of important vitamins, such as vitamin C, carotene, folate and B₆, among cereals and vegetables. Aside from its nutritional value, amaranth grain includes several bioactive compounds with potential health benefits. The total phenolic content in amaranth grains ranges from 21.2 to 57.0 mg gallic acid/100 g dry weight, mainly containing ferulic acid followed by quercetin and isorhamnetin. Phytate (0.09%) and saponins (4.96 mg/100 g) are the main antinutrients in amaranth ^[48]^[49].

2.9. Quinoa

Quinoa (Chenopodium quinoa Willd.) is a pseudocereal commonly known as the “golden grain” and has long been considered a source of nourishment and sustenance for Andean indigenous societies. Quinoa grain is mainly cultivated in the South American Andes region; however, over the past decades, it has been introduced in North America, Europe, Africa and Australia. Quinoa production has continuously expanded over the last few decades, and by 2013, the international year of quinoa, quinoa production and consumption had increased dramatically ^[48]^[49].

Quinoa flour is used to make a variety of toasted and baked goods, including bread, cookies, biscuits, noodles, pasta and pancakes. In addition, quinoa grains can be fermented to produce alcoholic beverages such as beer owing to its high starch level. Owing to its high nutritional quality and adaptability, quinoa is traditionally used in livestock feeding. Quinoa grains contain no gluten. Additionally, it has a high amount of nutrient ingredients such as proteins, dietary fibres, vitamins, fatty acids and minerals. The protein content of quinoa grains varied from 12.8 to 16.7%, which is higher than those of corn, rice and barley. The two main storage proteins in quinoa grain are albumins (35%) and globulins (37%). Quinoa proteins are recognised as high-quality proteins due to their great amount and well-balanced composition of essential amino acids. Quinoa protein contains a high concentration of lysine (2.4–7.8 g/100 g protein), methionine (0.3–9.1 g/100 g protein) and threonine (2.1–8.9 g/100 g protein), which are the limiting amino acids in ancient cereals such as maize and wheat ^[48]^[49]^[50].

Similar to other grains, starch is the most important carbohydrate component (32–69% of total carbohydrates). Its total dietary fibre content (7.0–16.5%) is comparable to modern cereals such as wheat. In addition to having a high protein content and good bioavailability, quinoa also has an intriguing lipid content (3.9–7.4%) that is higher than that of wheat and rice, making it a viable oil seed alternative source. The vitamin content, such as for vitamin C, E and folic acid, are greater than those of most other grains, and there is great potential to use quinoa as a functional food ingredient in mainstay food-processing applications. Quinoa has several health benefits in high-risk groups such as children and the elderly, as well as having prebiotic and probiotic effects ^[51]. However, it also contains phytate, saponin, tannins and protease inhibitor as the main antinutrients ^[52].

2.10. Teff

Teff (Eragrostis tef) is a nutritious, gluten-free pseudocereal grain that is native to Ethiopia and Eritrea. It is a staple food in these countries and is often used to make traditional dishes such as injera (a sour fermented pancake-like flat bread). Teff is a rich source of protein (12–15%) and fibre (6–8%) ^[53]. Teff contains a high level of lysine, which is an essential amino acid that is important for growth and tissue repair. Teff is a good source of minerals, including iron and calcium, which are beneficial for individuals with anaemia or osteoporosis. Teff is also a good source of resistant starch, which can help improve digestion and blood sugar control ^[52]^[53].

Teff also contains a variety of phytochemicals and antioxidants, including phenolic acids and flavonoids, which have been shown to have anti-inflammatory and anti-cancer properties and can reduce the risk of chronic diseases ^[54].

Phytic acid, tannins and protease inhibitors are the main antinutritional factors in teff ^[53]. To minimise the negative effects of these compounds, traditional methods of processing, such as fermentation, soaking and germination, can be used to reduce the levels of anti-nutritional compounds in teff. Teff can be ground into flour and used to make a variety of baked goods, including bread, pancakes and cakes. It can also be cooked and eaten as a porridge or added to salads and stews. Phytate, tannins, oxalates and saponins are the main antinutrients in teff ^[53]^[55].

2.11. Chia

Chia (Salvia hispanica) is a pseudocereal native to Mexico and Central America. It is a member of the mint family (Lamiaceae) and is closely related to other species such as sage and oregano. The chia seeds are small and oval in shape, measuring about 1–2 mm in diameter. They are black, brown or white in colour and have a glossy surface. The chia plant is drought-tolerant, making it suitable for dryland farming ^[56]^[57]^[58]. Chia seeds are an excellent source of dietary fibre (~34%), lipids (~33%) and protein (~18%). The protein content in chia seeds is composed of essential amino acids, such as lysine and arginine, and non-essential amino acids, such as alanine and aspartic acid. The chia seed lipid is rich in polyunsaturated acids with beneficial health impacts and, recently, has been extracted and characterised for food applications. Chia seeds are great sources of bioactive compounds such as omega-3 fatty acids (60–64%) and are a good source of minerals. Phytate and trypsin inhibitors are the major antinutrients in chia seeds ^[56]^[57]^[58]^[59].

2.12. Buckwheat

Buckwheat (Fagopyrum esculentum) is a pseudocereal that belongs to the family Polygonaceae. It is small and dark-coloured, typically brown or black, and is often used as a grain-like food source. Buckwheat is a hardy plant that can grow in a variety of soil types and climates, it is tolerant to frost and can be grown as a cover crop or as a green manure crop ^[49]^[60]. The seed of the buckwheat plant is a good source of carbohydrates (~65%), mainly in the form of complex carbohydrates, such as starch and dietary fibre. Additionally, it has a significant amount of protein (14–16%) of high quality, as it includes all essential amino acids, including lysine and arginine, which are often not present in other plant-based protein sources. Buckwheat is also rich in vitamins, such as B and E, and minerals. Some studies have revealed that the buckwheat seed contains a small amount of phytate, trypsin inhibitors and lectins, which can reduce the digestibility of proteins and cause allergic reactions in some individuals. The high levels of flavonoids present in buckwheat, particularly rutin, have been found to have antioxidant and anti-inflammatory properties, and they also have prebiotic and probiotic benefits ^[49].

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