The increasing market demand for functional foods and value-added products is pushing producers to think about incorporating novel additives that can improve their goods and fight lifestyle diseases. Sea buckthorn's health benefits have been known for centuries. This plant is rich in bioactive substances, such as vitamins C, K, and E, amino acids, antioxidants, phytosterols, essential fatty acids, and comprises a potent natural preservative. Its distinct scent is also enhanced by a low sugar content and a broad range of volatiles. Sea buckthorn has antiviral and antimicrobial qualities, making it a possible cosmeceutical or nutraceutical. It has been demonstrated to assist in the treatment of gastrointestinal and skin issues, diabetes, tumors, and cardiovascular disease. Sea buckthorn is a great option for new food products because of its many health advantages.
China is the original home of sea buckthorn, which is now found in all of the world's major temperate zones, including France, Russia, Mongolia, India, Great Britain, Denmark, the Netherlands, Germany, Poland, Finland, and Norway [1]. Air pollution, frost, and drought are just a few of the extreme environments in which it can grow [2]. The generic name Hippophae comes from the fact that sea buckthorn was once fed to horses in ancient Greece in order to give them glossy coats (Greek: hippos, horse; phaos, shiny) [3,4]. Reaching up to 7 meters in height, this prickly, dioecious, deciduous shrub with yellow or orange berries is a member of the oleaster family (Elaeagnaceae). Based on genetic variations, Hippophae rhamnoides is classified into nine subspecies, two of which are unique to China: H. rhamnoides ssp. sinensis Rousi and H. rhamnoides ssp. yunnanensis Rousi [7].
The underside of the long, lanceolate leaves of sea buckthorn is adorned with distinctive silver hairs. It produces a large number of tiny green and brown flowers that grow together to form clusters when it blooms in late April or early May. Berries eventually form, typically yellow or orange in color. September is the season of ripening. A tiny, elongated, grooved stone covers an oily seed in each fruit [2,6,8, 9]. The berries taste and smell like pineapple, with a distinct bitter and sour flavor [2,3,4]. Because of this, sea buckthorn is sometimes referred to as Siberian pineapple [2]. Harvesting is quite challenging due to the berries' dense arrangement and the abundance of thorns (Figure 1). Because of this, complete bunches are occasionally cut off from the shrub; however, doing so stunts the growth of subsequent crops [5,6]. Berries can therefore only be harvested once every two years [6,10]. Sea buckthorn fruit is still harvested by hand in developing nations, particularly China. Additional harvesting techniques include direct harvesting, which uses a vacuum suction harvester, and indirect harvesting, which involves shaking a section of the plant using either cutting or vibration harvesting. Fruit separates from the stem when forces are applied to the trunk or branches [6].
Numerous bioactive substances are found in sea buckthorn, not only in the berries but also in the seeds and leaves [12,13,14,15,16]. The antioxidant content varies among the plant's parts, indicating that sea buckthorn has beneficial biological, physiological, and therapeutic effects. Ascorbic acid, which is present in berries, juice, and leaves [17], phytosterols like cycloartenol and campesterol [18], citrostadienol and sitosterol [19], carotenoids, including lutein, zeaxanthin, α-carotene, β-carotene, and γ-carotene [20,21], tocopherols—α-tocopherol, β-tocopherol, and γ-tocopherol [3]; the most common flavonoids, isorhamnetin and quercetin [22,23], or polyphenolic compounds, like gallic acid in leaves and berries, and lesser levels of caffeic acid, p-coumaric acid, and ferulic acid [24]. In both Europe and Asia, traditional medicine has utilized the plant since ancient times [25]. In the production of food and feed, it is widely used (Figure 2). The entire fruit is pressed to produce a liquid that, when centrifuged, can be used as juice. Oil or oil products are made from the remaining portion [26]. Tea is the most widely consumed dish made with sea buckthorn leaves. Berries are also used in the brewing of this hot beverage [27]. Sea buckthorn fruit can be used to make antioxidant-rich jam, or it can be combined with other fruits like papaya, grapes, or watermelon [28, 29]. India has the potential to use sea buckthorn leaves, seeds, and fruit leftovers as animal and poultry feed [30].
When ripe, the berries on female sea buckthorn plants can be yellow, orange, or red in color. The spherical fruits are 3–8 mm in size [33]. Wax is applied in a thin layer to the skin. Every berry has a single sheathed seed inside, encased in a cellular structure filled with juice [34]. A sea buckthorn berry's general composition is 23% seed, 7.75% skin, and 68% pulp [35,36,37]. Sea buckthorn berries are highly acidic and astringent, making them unpleasant to eat raw. Nonetheless, a number of techniques can be employed to lessen astringency, including frosting before additional processing and/or blending with juices that are sweeter, like apple or grape.
The nutritional value and chemical makeup of sea buckthorn berries can differ depending on the subspecies, place of origin, climate, harvesting period, and processing techniques [38]. Sea buckthorn berries and seed oil contain a variety of bioactive compounds [39]. Li, Schroeder, and Yao [10] claim that sea buckthorn fruits are among the healthiest and highest in vitamins that a plant can yield. The nutritional value of sea buckthorn berries is higher than that of other berries due to their abundance in flavonoids and other antioxidants, fat- or water-soluble vitamins (like lycopene, C and E, and β-carotene), phytosterols, polyunsaturated fatty acids (like omega-7 palmitoleic acid), amino acids, and minerals (like calcium, iron, etc.) in addition to carbohydrates and proteins [26, 40]. When compared to other fruits and vegetables that are high in carotenoids, sea buckthorn berries have remarkably high levels of lipids. Carotenoids' bioavailability is increased by lipids, which makes it easier for people to absorb them [13]. Juice without filtering can have a freezing point as low as −18.5 °C due to its high lipid content. Juice can stay liquid even at below-freezing temperatures, which is an important benefit for storage [41].
Vitamins
Sea buckthorn berries are most known for their exceptionally high vitamin C content. Berries from the European-growing subspecies Rhamnoides contain 360 mg/100 g of vitamin C, while berries from the Chinese subspecies Sinensis contain 2500 mg/100 g [45,46]. Fruit cultivated in the United States has an average of 695 mg of vitamin C per 100 g, but can contain anywhere between 114 and 1550 mg. Compared to oranges, this has roughly a 12-fold higher vitamin C content. Compared to sea buckthorn berries, other fruits that are often thought to be rich sources of vitamin C—like strawberries, kiwis, tomatoes, carrots, and hawthorns—also have substantially lower concentrations [47, 48]. The local environmental factors that exist during the brief reproductive period are linked to the variations in vitamin C levels found in sea buckthorn berries from various regions [45]. The pulp of sea buckthorn berries has 223.2 mg of vitamin C per 100 g, according to Arimboor [49]. After processing, the juice retains about 75% of the vitamin C found in the pulp, with the final juice having 168.3–184.0 mg/100 g. The fruit of sea buckthorn trees also contains vitamin E. Vitamin E content has been measured as follows: 160 mg/100 g for berries [50,51], 162–255 mg/100 g for juice [51], 481 mg/100 g for pulp [52], and 40.1–103.0 mg/100 g for seeds [53]. Vitamin K is found in berries at levels of 110–230 mg/100 g and in seeds at 109.8–230.0 mg/100 g. It is essential for the post-synthesis modification of proteins involved in blood coagulation as well as for regulating calcium binding in bones and other tissues [54, 55].
Phytosterols
The total sterol content in the seeds, fresh pulp/peel, and whole berries of two subspecies of sea buckthorn (sinensis and rhamnoides) ranges from 1200–1800 mg/kg, 240–400 mg/kg, and 340–520 mg/kg, according to Yang and colleagues [38]. The highest concentrations were found in sitosterol derivatives. The other main sterols identified were R-amyrin, stigmastanol, and cholesterol. Depending on the subspecies and collection sites, these compounds' contents and composition varied slightly. Human plasma cholesterol levels can be lowered through plant sterol ingestion. Treatment for coronary heart disease involves lowering cholesterol [33].
Amino Acids
Of the 22 known amino acids, sea buckthorn berries contain eighteen (Table 1). It's common knowledge that amino acids are the building blocks of proteins. About half of them are involved in vital bodily functions like energy production, muscle and cell growth, fat loss, and vital brain and mood processes. Threonine, valine, methionine, leucine, lysine, tryptophan, isoleucine, and phenylalanine are among the essential amino acids [51].
Amino Acid Content (mg/100 g)
Aspartic acid 426.6
Serine 28.1
Glutamine 19.4
Glycine 16.7
Alanine 21.2
Cysteine 3.3
Valine 21.8
Ammonia 41.8
Tyrosine 13.4
Isoleucine 17.4
Methionine 2.3
Proline 45.2
Phenylalanine 20.0
Histidine 13.7
Lysine 27.2
Threonine 36.8
Arginine 11.3
Organic Acids
One important organic acid found in many species is sorbic acid, which acts as a preservative. The antimicrobial property of sea buckthorn fruits is linked to the presence of sorbic acid, a common acid found in many Rosaceae fruits. The actual amount varies greatly, with some species comprising amounts below detectability. Malic and quinic acids are two other primary organic acids present in sea buckthorn fruits. Though the amounts also differ amongst species, these three acids collectively make up about 90% of all fruit acids in sea buckthorn plants. Russian berries have comparatively lower total acidity concentrations (2.1–3.2 g/100 mL). Finnish genotypes range in total acidity from 4.2 to 6.5 g/100 mL. The highest organic acid concentrations, ranging from 3.5 to 9.1 g/100 mL, are found in Chinese genotypes [51, 53, 56].
Mineral Elements
Sea buckthorn fruit is rich in iron, magnesium, copper, zinc, sodium, potassium, phosphorus, calcium, nitrogen, and manganese, among other mineral elements [57,58]. Potassium is the most prevalent mineral element in berries and juice [56], with a concentration of 9.33–13.42 ppm in the seed and 10.12–14.84 ppm in the pulp [55]. Eight components in berries from various nations were compared in a study conducted by Kallio and associates. Compared to Chinese fruit, Finnish species had higher levels of cadmium but lower levels of iron, calcium, and lead. The amount of N, Ca, K, Na, Mg, Cu, Fe, Zn, and Mn is influenced by the plant's maturity [56,59].
Sugars
Sea buckthorn berries contain three main types of sugars: glucose, fructose, and xylose. Between 5.6% and 22.7% of raw juice in the species cultivated in China is made up of total soluble sugars [51,53,56, 60]. China-sourced berries had the highest sugar content, with Russian and Finnish berries coming in lower order [56]. Three distinct subspecies (Hippophae rhamnoides ssp. sinensis, rhamnoides, and mongolica) harvested over a four-year period in China, Finland, and Russia were the subjects of Yang's [61] study on sugar levels. The juice from Finnish fruit (ssp. rhamnoides) had combined levels of glucose and fructose ranging from 0.6 g/100 mL to 24.2 g/100 mL in the juice pressed from wild Chinese berries (ssp. sinensis). Various batches contained varying amounts of sugar. The slightly different harvest dates and weather in each of the years the crops were grown can account for these variations. Glucose is the primary sugar found in sea buckthorn berries of all varieties. In berries harvested in Russia and China, glucose and fructose make up about 90% of the total sugar content. Lower concentrations of these sugars (60%) are a characteristic of Finnish species [53,56,62].
Volatile Compounds
Because sea buckthorn berries contain volatile compounds, they have a distinct scent. The primary components of these volatile substances are alcohols, branched or n-fatty acids, and short chain esters. The volatiles' composition is determined by when they are harvested [38]. Finnish berries are rich in ethyl 2-methylbutanoate, ethyl 3-methylbutanoate, and ethyl hexanoate; in contrast, Chinese berries have higher concentrations of butyl pentanoate, 2-methylpropyl 3-methylbutanoate, pentyl 3-methylbutanoate, and ethyl 3-methylbutanoate [56,63]. Using combined gas chromatography-mass spectrometry, 60 volatile compounds were identified in a study by Hirvi and Honkanen [64]. Ethyl hexanoate, 3-methylbutyl 3-methylbutanoate, 3-methylbutanoic acid, 3-methylbutyl hexanoate, 3-methylbutyl benzoate, and 3-methylbutyl octanoate were the compounds that were found in the highest concentrations. Traces of aromatic compounds and terpenes were discovered. Cakir found 30 compounds in the volatile oil of sea buckthorn by employing combined gas chromatography-mass spectrometry as well. Decanol (5.6%), ethyl decanoate (5.5%), ethyl dodecanoate (3.7%), and ethyl octanoate (9.9%) were the main ingredients [65].
Structures of the most relevant phytochemicals with present in sea buckthorn with medicinal value
(A)—quercetin, (B)—isorhamnetin, (C)—catechin,
(D)—gallocatechin, (E)—epigallocatechin,
(F)—ursolic acid [72,73,74,75,76,77]
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