1 
 
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Taylor & Francis: 
 
This is an Accepted Manuscript version of the following article, accepted for 
publication in: 
 
JOURNAL Critical Reviews in Food Science and Nutrition, 
CITATION 
 
 
Xueying Ma, Wei Yang, Heikki Kallio & Baoru Yang 
(2021) Health promoting properties and sensory characteristics 
of phytochemicals in berries and leaves of sea buckthorn 
(Hippophaë rhamnoides), 
DOI https://doi.org/10.1080/10408398.2020.1869921 
   
  
 2 
Health promoting properties and sensory characteristics of 
phytochemicals in berries and leaves of sea buckthorn 
(Hippophaë rhamnoides) 
Xueying Ma b, Wei Yang b, Heikki Kallio b, Baoru Yang a,b*  
 
a Shanxi Center for Testing of Functional Agro-Products, Shanxi Agricultural University, 030000 
Taiyuan, China 
 
 b Food Chemistry and Food Development, Department of Biochemistry, University of Turku, FI-
20014, Turku, Finland 
 
Xueying Ma   E-mail: xueying.ma@utu.fi 
     ORCID 0000-0002-0887-2693 
Wei Yang E-mail: yanwei@utu.fi 
ORCID: 0000-0002-7889-6172 
Heikki Kallio E-mail: heikki.kallio@utu.fi 
 
* Corresponding author: 
       Baoru Yang   
  E-mail address: baoru.yang@utu.fi 
 Tel: +358 29 450 2917; +358452737988 
 
 
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Abstract  
Sea buckthorn (Hippophaë rhamnoides L., SB), as a multi-functional plant, is widely grown 
in Asia, Europe and Canada. The berries and leaves of SB contain a diverse array of health-
supporting phytochemicals, which are also related to the sensory qualities of berry and berry 
products. This review summarizes the biologically active key-compounds of the berries and 
leaves of SB, their health-promoting effects, as well as the contributions to the sensory quality 
of the berries. The target compounds consist of sugars, sugar derivatives, organic acids, 
phenolic compounds and lipophilic compounds (mainly carotenoids and tocopherols), which 
play an important role in anti-inflammatory and antioxidant functions, as well as in metabolic 
health. In addition, these compounds contribute to the orosensory qualities of SB berries, which 
are closely related to consumer acceptance and preference of the products. Studies regarding 
the bioavailability of the compounds and influence of the processing conditions are also part 
of this review. Finally, the role of the sensory properties is emphasized in the development of 
SB products to increase utilization of the berry as a common meal component and to obtain 
value-added products to support human health. 
 
 
Keywords: Bioactive compounds, bioavailability, health effects, processing conditions, sea 
buckhorn, sensory quality  
 
 
 
Introduction 
Consumption of fruits and vegetables has been recommended in dietary guidelines to reduce 
the risk of various diseases and disorders (Martin et al. 2013). Sea buckthorn (Hippophaë 
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rhamnoides L., SB), as a multi-functional plant, has attracted much attention due to the diverse 
array of nutrients and phytochemicals in its berries and leaves, such as polyphenols, carotenoids, 
and vitamin E (Bal et al. 2011; Ciesarová et al. 2020). These compounds have been claimed to 
have many health-promoting properties, related especially to anti-inflammatory activities 
(Ganju et al. 2005; Yang and Kortesniemi 2015), cellular antioxidant activities and anti-
proliferative properties in vitro (Guo et al. 2017; Kim et al. 2010), as well as positive effects 
on the cardiovascular system (Olas 2016; Suomela et al. 2006; Xu  et al. 2011). The nutritional 
attributes, medicinal and therapeutic potential, as well as health-related application of SB have 
been well reviewed (Olas 2018; Stobdan et al. 2013; Suryakumar and Gupta 2011). 
Despite the indicated health-promoting properties, sea buckthorn berries are less commonly 
consumed as food or food ingredients due to their sensory properties (Laaksonen et al. 2016; 
Ma et al. 2017a). Their sensory characteristics are typically described as strongly sour, 
astringent and bitter, with a very low level of sweetness (Tang, Kälviäinen, and Tuorila 2001; 
Tiitinen, Hakala, and Kallio 2005). From the view of consumers, orosensory properties are the 
most important factors affecting the preferences, choices and acceptance of foods (Geertsen et 
al. 2016; Laaksonen et al. 2016). In order to obtain palatable products, malolactic fermentation 
(MLF) is used to reduce the high acidity of SB juice, and some other fruits are added to balance 
bitterness and astringency of sea buckthorn (Markkinen et al. 2019; Selvamuthukumaran, 
Khanum, and Bawa 2007; Tiitinen et al. 2006a; Tiitinen et al. 2007). Improving the sensory 
qualities of SB berries and related products will add their value, increase consumption, and 
create new markets for these commodities. 
Generally, the content of sugars and acids, especially, the sugar/acid ratio plays an 
important role in determining the flavor and consumer acceptance of berries and berry-based 
products (Laaksonen et al. 2013; Laaksonen et al. 2016). Along with glucose and fructose, SB 
berries contain some less common sugar derivatives, e.g. ethyl β-D-glucopyranoside (β-EG) 
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and sugar alcohols such as L-quebrachitol (1 L-2-O-methyl-chiro-inositol) and other 
derivatives of inositol in varying quantities, which may contribute to the sensory properties 
(Ma et al. 2017a; Yang 2009; Zheng et al. 2012). Flavonol glycosides (FGs) and 
proanthocyanidins (PAs) are the main phenolic compounds present in SB berries, which can 
cause astringency and bitterness of the plant-based foods and beverages (Ma et al. 2016; 
Teleszko et al. 2015; Yang et al. 2017). Comparing to PAs, FGs have lower threshold to induce 
diverse drying or puckering astringent mouthfeels in sensory evaluations (Bajec and Pickering 
2008; Hufnagel and Hofmann 2008; Scharbert et al. 2004).  
In addition to SB berries, the composition and content of bioactive compounds in the leaves 
have been well investigated, among which ellagitannins are the most abundant polyphenols 
(Ciesarová et al. 2020; Ma et al. 2019). The extracts of SB leaves have shown various 
pharmacological activities, such as anti-inflammatory activity and hepatoprotective effect 
(Geetha et al. 2005; Geetha et al. 2008; Marnila et al. 2013). The animal and human studies 
have indicated potential of these extracts in reducing the incidence of CVD, certain cancers, 
and age-related degenerative diseases (Guliyev et al. 2004). Recently, due to the increasing 
interest and consumption of herbal infusions from edible plants, there has been an increasing 
interest in utilization of SB leaves and converting them into tea-type beverage products (Ma et 
al. 2019). Despite these aforementioned health benefits and utilization, the majority of the SB 
leaves still remains under-utilized as agricultural waste after berry harvesting.  
This current review aims to provide an overview of the bioactive compounds 
characteristic in SB berries and leaves, as well as their health-promoting properties and 
contributions to sensory properties of SB. Special emphasis is placed on the importance of SB 
as part of a healthy diet for reduction of risk of various chronic diseases. The first part presents 
the bioactive components, their bioavailability and health effects. The second part reviews 
research on contributions of these compounds to the sensory properties of the berries and berry 
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products. The third part draws attention to the effects of processing conditions on the bioactive 
compounds. Moreover, the challenges of the development and processing of SB products are 
discussed. 
Bioactive compounds in berries and leaves of sea buckthorn 
Several epidemiological studies have linked diet, especially substantial consumption of 
plant-based foods, with decreased risk of chronic diseases. Sea buckthorn berries and leaves 
are characterized by high content of bioactive compounds (Table 1), which have been 
suggested to have various health-beneficial effects (Figure 1).  
Flavonol glycosides 
Flavonol glycosides (FGs) constitute the most abundant group of phenolic compounds in 
sea buckthorn berries. The total content of FGs varied from 23 ~ 400 mg/100 g FW (Ma et al. 
2016; Teleszko et al. 2015; Yang et al. 2009; Zheng et al. 2016), which is up to 150 times 
higher than the levels reported in tomato fruit (2 mg/100 g FW), and 10 times higher than those 
in chokeberry (Aronia melanocarpa, 27 mg/100 g FW) and American cranberry (Vaccinium 
macrocarpon, 21 mg/100 g FW) (Mikulic-Petkovsek et al. 2012; Stewart et al. 2000). Among 
the flavonols, isorhamnetin is the typically most abundant aglycone (mean ~60 % of total 
flavonol aglycones), as isorhamnetin-3-O-rutinoside (I-3-R, mean ~30 % of total FGs) and 
isorhamnetin-3-O-glucoside-7-O-rhamnoside (I-3-G-7-Rh, mean ~10 % of total FGs) are two 
major FGs (Ma et al. 2016; Pop et al. 2013). The content of flavonols in the berries varies 
considerably due to origin, weather conditions, growth location, latitude, and harvest date 
(Kortesniemi et al. 2017; Kortesniemi et al. 2014; Ma et al. 2016; Yang et al. 2017; Zheng et 
al. 2012). In other fruits, isorhamnetin is less common than quercetin and kaempferol, except 
the copao fruit (Eulychnia acida Phil.), which is rich in I-3-R with content up to 54 g/100 g in 
phenolic extracts of epicarp (Belitz et al. 2004; Jiménez-Aspee et al. 2014).  
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On a dry weight basis, the content of flavonol glycosides in sea buckthorn leaves was 
reported to be close to the levels found in the berries (Pop et al. 2013). Leaves of SB contain 
flavonols up to 1400 mg/100 g DW comprising glycosides of isorhamnetin, quercetin and 
kaempferol, among which glycosides of isorhamnetin (~ 50 % of total FGs) represent the 
highest percentage. I-3-G-7-Rh and I-3-R are the two major FGs in leaves, which were also the 
major flavonols reported in the berries (Ma et al. 2019). Other FGs, such as isorhamnetin-3-O-
glucoside (I-3-G) and rutin, were also found in both the leaves and the berries (Ma et al. 2016; 
Ma et al. 2019). 
Flavonol glycosides in diet have gained increasing interest due to their particularly high 
antioxidant capacities and concomitantly strong health-promoting effects, such as anticancer 
activity, immunomodulatory activity and antimicrobial activity (Jacques et al. 2013; Xiao et al. 
2016; Xue et al. 2015; Yang et al. 2018). It has been reported that increasing dietary intake of 
FGs of SB berry may play an important role in reducing the cardiovascular mortality (Cheng 
et al. 2003; Clair et al. 2002).  
Various experiments showed that isorhamnetin and its derivatives have anti-inflammatory 
effects in vivo and in vitro (Yang et al. 2013) as well as anti-proliferative (Guo et al. 2017; Li 
et al. 2014; Teng et al. 2006), anti-viral (Apers et al. 2002), and anti-oxidative stress activities 
in vitro (Yang et al. 2014). Among the flavonoid aglycones of SB berries, isorhamnetin showed 
the highest anti-proliferative activity towards human liver cancer cell HepG2. Weaker anti-
proliferative activity was observed by isorhamnetin-3-O-sophoroside-7-O-rhamnoside (I-3-S-
7-Rh) and I-3-G-7-Rh, which, however, exhibited large amount in SB berries (Guo et al. 2017). 
The earlier results demonstrated that isorhamnetin extracted from the SB berries may reduce 
the lipid peroxidation and protein carbonylation in human plasma and oxidative stress induced 
by H2O2/Fe (Skalski et al. 2019). Isorhamnetin and isorhamnetin-3-O-glucuronide are reported 
 8 
to have a significant cytotoxic effects on human breast cancer MCF-7 cells via a ROS-
dependent apoptosis pathway (Wu et al. 2018). Further, isorhamnetin is reported to inhibit 
atherosclerotic plaque development in apolipoprotein E knockout mouse (apoE −/−) (Luo et al. 
2015). It is also demonstrated that this compound is a potent immunosuppressive agent and 
may be used to prevent and treat inflammatory and autoimmune diseases, as well as 
transplantation rejections (Shi et al. 2018). Moreover, a C-glucoside of isorhamnetin is one of 
the best radical scavengers among polyphenols of SB leaves (Radenkovs et al. 2018). 
In human nutrition, quercetin represents the main flavonol (Ahn et al. 2008). There is 
growing evidence showing that quercetin is a potent natural antioxidant. There are also 
indications that quercetin may have a potential in management of chronic diseases, such as 
cardiovascular diseases and cancer (Lesjak et al. 2018; Nabavi et al. 2012; Russo et al. 2012). 
Besides quercetin, kaempferol glycosides display anti-obesity and anti-diabetic potential by 
reducing fat accumulation in adipose tissues, improving hyperlipidemic and diabetic conditions 
in obese mice (Zang et al. 2015).   
The sugar moieties of the flavonol glycosides strongly influence the absorption efficiency 
of flavonols (Manach et al. 2005; Petersen et al. 2016). Flavonol aglycones and only some 
glucosides are absorbed in the small intestine. E.g. quercetin-3-O-glucoside is transported into 
the enterocytes by the sodium-dependent SGLUT1 transporter (Wolffram et al. 2002). 
Flavonols linked e.g. to a rhamnose may not be absorbed in the small intestine but they are 
fermented and degraded by the colonic microflora (Hollman and Katan 1997; Kahle et al. 2006). 
The bioavailability of quercetin glucoside is about 5-fold compared with that of rutinoside in 
humans (Graefe et al. 2001; Walle et al. 2000). An in vitro study suggested that the metabolites 
of I-3-G yielded by human intestinal flora, such as isorhamnetin, kaempferol, and quercetin, 
may improve the absorption of I-3-G by colonic mucosa (Du et al. 2014). This was supported 
by the findings of another study conducted by in vivo pharmacokinetics model, where quercetin 
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and kaempferol increased the intestinal absorption of isorhamnetin extracted from the SB 
berries by regulating the activity and expression of the multidrug resistance-associated protein 
2 (Xiao et al. 2019). Moreover, the addition of a small amount of SB oil was showed to enhance 
the absorption and bioavailability of the SB flavonols in humans (Suomela et al. 2006). 
Tannins 
Proanthocyanidins 
     Tannins consist of condensed and hydrolyzable tannins. Condensed tannins, known as 
proanthocyanidins (PAs), are oligomers and polymers of flavan-3-ol units produced through 
the biosynthetic flavonoid pathway widely distributed in the plant kingdom. The agricultural 
benefits as well as chemistry and biochemistry of PAs have been well summarized in literature 
(Aerts et al. 1999; Ferreira and Slade 2002; Marles et al. 2003).  
The difference in subunits and bond positions/configurations results in the wide structural 
diversity in PAs, and the number of isomers increases markedly with an increase in the degree 
of polymerization (DP). Procyanidins (PCs) and prodelphinidins (PDs) consist only of 
(epi)catechin or (epi)gallocatechin, respectively. The most common in foods are B-type PAs, 
in which the monomeric units of flavan-3-ols are linked by C–C bonds (via C4–C8 and/or C4–
C6). Again, the A-type PAs are less common and present in a few specific foods only, such as 
cranberries, lingonberries and avocados (Deng et al. 2013).  
B-type PAs with DP from 2 to 11 have been detected by HILIC-ESI-MS in SB berries 
(Kallio et al. 2014). More than 60 combinations of (epi)catechins and (epi)gallocatechins of 
PA dimers and trimers were found in SB berries, in which (epi)gallocatechins were the main 
monomeric units (Kallio et al. 2014). The PA oligomers (DP 2−4, 4−26 mg/100 g DW) formed 
only a small part of the total PA pool (390−1940 mg/100 g DW) in the berries studied (Yang 
et al. 2017). PA oligomers (DP 6−9) of the SB pomace are composed mainly of PDs and smaller 
 10 
amounts of PCs (Rösch et al. 2004a). The PAs of SB seeds consist of catechin, epicatechin, 
gallocatechin and epigallocatechin (Fan et al. 2007). The majority of PAs, especially PAs with 
DP > 4 in SB berries, are reported to be present mainly in the seeds (Ma et al. 2017b). Genetic 
background and environmental factors have a significant influence on the content of PAs in SB 
berries. The content of total PAs is 2−3 fold in the berries of H. rhamnoides ssp. rhamnoides 
in northern Finland compared with the berries from southern Finland, but for oligomeric PAs 
(DP 2−4) the situation is vice versa (Yang et al. 2016). The environmental conditions, including 
the length and radiation sum of the growth season, as well as the temperature sum, are 
positively correlated with PA oligomers (DP 2−4), but negatively with the content of total PAs 
(Yang et al. 2016; Yang et al. 2017). According to Arimboor and Arumughan (2011), SB leaves 
contain a smaller amount of PAs compared to berries, and the average DP is 10.6.  
Diet rich in PAs is associated with reduced risks of cardiovascular disease, 
neurodegenerative conditions, and metabolic syndrome (Aron and Kennedy 2008; Langhans 
2017). PAs of SB pomace possessed higher or comparable antioxidant capacities to that of 
ascorbic acid or Trolox (Rösch et al. 2004b). No significant difference was detected in the 
antioxidant capacity measured by scavenging Fremy’s salt between monomeric flavan-3-ols 
and dimeric PAs in SB (Rösch et al. 2004b). The PAs of SB seeds are reported to have the 
protective effect against light-induced retinal degeneration in vivo via antioxidant, anti-
inflammatory and anti-apoptotic mechanisms (Wang et al. 2016). Moreover, anti-proliferative 
effects of PAs extracted from SB has been shown in human colon and liver cancer cell lines 
(Grey et al. 2010).  
The biological activities of PAs in vivo depend on the DP value, which affects greatly the 
absorption of PAs. Data on absorption of flavan-3-ols has been conflicting. Absorption of 
flavan-3-ol dimers have been detected in humans and rats in vivo (Baba et al. 2002; McKay et 
al. 2015), whereas some studies failed to show absorption of PA dimers (Donovan et al. 2002; 
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Gonthier et al. 2003). Despite the high content PAs in some foods, flavan-3-ols are poorly 
absorbed. The polymers with a DP > 4 are not absorbable because of their large molecular size 
and the gut barrier (Manach et al. 2005). It has been shown that PAs of SB berries and leaves 
have the ability to precipitate proteins and to inhibit digestive enzymes, which might alter the 
digestion and bioavailability of proteins and phenolics (Arimboor and Arumughan 2011). 
Phenylvalerolactones and phenolic acids are the main microbial metabolites of PAs resulted 
from the action of the colon microbiota, which may contribute to the health-promoting 
properties of PAs in vivo (Ou and Gu 2014).  
Ellagitannins 
Ellagitannins (ETs) form a complex class of polyphenols, which are water-soluble and able 
to precipitate proteins and alkaloids. ETs are characterized by one or more 
hexahydroxydiphenoyl (HHDP) moieties esterified to a sugar. They are less frequently 
encountered, but berries such as strawberries and raspberries are major sources of intake of 
ETs in western diets.  
The leaves of Finnish sea buckthorn (Hippophaë rhamnoides ssp. rhamnoides) are rich in 
ellagitannins (mean 76 mg/g DW), of which ten major compounds are identified and quantified 
(Moilanen et al. 2015; Suvanto et al. 2018). Among these compounds, hippophaenin C, 
stachyurin, and casuarinin were on average the most abundant, with their total concentrations 
adding up to 63 % of the total ETs in SB leaves (Suvanto et al. 2018). After dry-processing, 
the SB leaf products (tea-type beverages) are still rich in ETs with the concentrations ranging 
from 10 to 20 mg/100 mL infusions (Ma et al. 2019).   
Literature concerning the absorption and metabolism of ETs in humans is limited. ETs are 
hydrolyzed to ellagic acid (EA) by the action of gastric acid and the intestinal enzymes 
(Mertens-Talcott et al. 2006). Part of EA is absorbed in the small intestine, while the remaining 
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EA and ETs are further metabolized to urolithins and hydroxy-6H-dibenzopyran-6-one 
derivatives by the colon microbiota (Cerdá et al. 2005). The gut microbiota may play an 
important role in the biological effects of ETs (Puupponen-Pimiä et al. 2013). EA is also 
proposed to be considered as a biomarker for human studies related to consumption of dietary 
ETs (Andreux et al. 2019; Ludwig et al. 2015; Seeram et al. 2004). 
Despite the poor absorption, ETs may have important beneficial effects on gut health and 
physiological impacts on the whole body due to the metabolites produced through fermentation 
of ETs by gut microbiota. ETs, EA and their metabolites have a wide range of biological 
activities and health-promoting properties, such as antioxidative, anti-inflammatory and 
prebiotic effects (Heinonen 2007; Landete 2011). The isomers of galloyl-bis-HHDP-glucoses 
and HHDP-galloylglucoside which belong to ETs, had been found to be the most effective 
natural antioxidants among phenolics extracted from SB leaves (Radenkovs et al. 2018). 
Majority of the plant extracts ETs showed good metal ion chelating ability (Moilanen et al. 
2016). Moreover, the antioxidative activities of ETs in SB leaves and leaf products are more 
likely due to the capacity of chelating metal ions than the radical scavenging activities by 2-
deoxyribose assay (Ma et al. 2019). Strictinin, isostrictinin, and casuarictin purified from SB 
leaves, have antiviral and interferon-inducing activities, and these compounds have been used 
to develop therapeutic agents (Korekar et al. 2011).  
Further, dietary ETs and EA have displayed antioxidative and anti-carcinogenic effects in 
the gastrointestinal tract. The colonic microbiota further metabolizes EA to urolithins (urolithin 
A, B, C, and D), which have been shown to have high bioavailability compared to their 
precursors, ETs and EA (Landete 2011; Puupponen-Pimiä et al. 2013). Urolithins, urolithin A 
in particular, were the most active compounds to reduce lipopolysaccaride-induced NF-κB 
activity (Rønning et al. 2020). Moreover, urolithin A stimulated mitophagy and improved 
 13 
muscle health in elderly individuals and in preclinical ageing people (Andreux et al. 2019). 
Although the microbial metabolite urolithin B showed lower antioxidant activity than ETs, 
urolithins may still display health benefits, such as estrogenic and/or anti-estrogenic activities 
(Cerdá et al. 2005). It is, however, worth to note that ETs may inhibit protease activities and 
affect the protein digestion (Mcdougall and Stewart 2005). 
Phenolic acids 
 Both sea buckthorn berries and leaves are rich in phenolic acids, such as gallic, salicylic 
and caffeic acids. Gallic acid is dominant in both free and bound forms, contributing 66 % and 
85 % of the total phenolic acids in SB pulp (soft parts of berry, ~1000 mg/kg DW) and leaves 
(~5000 mg/kg DW), respectively (Arimboor et al., 2008). Zadernowski et al (2005) reported 
that the salicylic acid (21−47 mg/kg DW) was the principal phenolic acid (55−74 % of the total 
phenolic acids) in SB berries of the six cultivars studied. The major fraction (approximately 
70 %) of the phenolic acids in SB berries is found to be concentrated in seeds. Furthermore, 
the total phenolic acid content in the seed kernel (~5700 mg/kg DW) is higher than in the berry 
pulp and the seed coat (Arimboor et al. 2008). Genetic background has a great effect on the 
content of total phenolic acid, which has been reported to be highest in H. rhamnoides ssp. 
yunnanensis, followed by ssp. mongolica, ssp. turkestanica and ssp. sinensis (Guo et al. 2017). 
Dietary phenolic acids can be rapidly absorbed and metabolized in humans (Lafay and Gil-
Izquierdo 2008; Nardini et al. 2009). The influence of phenolic acids on both the expression 
and activity of enzymes has been studied with cell and animal models, involved in production 
of inflammatory mediators and in response to mediators of growth factors, cytokines and 
endotoxins (Russell, Wendy and Duthie 2011; Russell et al. 2008). Phenolic acids extracted 
from SB berries have been reported to be stronger free radical scavengers than flavonols (Tkacz 
et al., 2020). Among the phenolic acids of SB berries, gallic acid was observed to have stronger 
 14 
anti-proliferative activity towards human liver cancer cell HepG2 compared to protocatechuic 
acid and ferulic acid (Guo et al. 2017). Obesity leads in multiple inner organs to DNA damage 
which can be prevented by low amounts of gallic acid (Setayesh et al. 2019). Salicylic acid has 
an anti-inflammatory activity, which has been reported to reduce the risk of colorectal cancer 
(Paterson and Lawrence 2001). Caffeic acid displays the major effective anti-proliferative 
action when T47D human breast cancer cells were tested, even at low concentrations (Kampa 
et al. 2004). 
Non-phenolic bioactive compounds   
Lipophilic compounds 
Sea buckthorn pulp oil is abundant in unsaturated fatty acids, such as palmitoleic, oleic, 
linoleic and linolenic acids. The high content of palmitoleic acid (16:1n-7) is typical for SB 
pulp/peel oil, but the content varies widely (20−45 %) depending on the origin of the plant 
(Yang and Kallio 2001). SB pulp/peel oil has been used for development of cosmetic products 
and for healing purposes due to palmitoleic acid forms a significant part of the epidermal lipids 
of human skin. Besides, palmitoleic acid has beneficial effects on epithelial tissues including 
the digestive, respiratory and urological organs, the vagina among postmenopausal women and 
the inner eyelid (dry eye syndrome) (Larmo et al. 2019; Larmo et al. 2014; Olas 2018). 
However, palmitoleic acid is absent in the seed oil, but linoleic (18:2n-6) and α-linolenic acids 
(18:3n-3) (close to 1∶1) comprises about 70 % of seed oil fatty acids (Yang and Kallio, 2001). 
Linolenic acid is important for the water permeability barrier of skin as a structural component 
of ceramides (Larmo et al. 2014). The composition of polyunsaturated fatty acids (PUFA) 
differs from those of most vegetable oils, e.g. the PUFAs of canola oil contain linoleic and α-
linolenic acids at a ratio of close to 2:1 (Ursin 2003).  
 15 
The color of sea buckthorn berries is mainly affected by the content and composition of 
carotenoids, the total content of which varies from 1 to 120 mg/100 g FW of berries (Yang 
2001). Forty-one different carotenoids have been reported in various cultivars of SB berries 
(Bekker and Glushenkova 2001), the major compound being β-carotene (15–55 % of the total) 
(Yang 2001). The content of β-carotene varies from 0.2 to 17 mg/100 g FW in SB berries, and 
the concentrations of α-, γ-, and dihydroxy-β-carotene, zeaxanthin, and canthaxanthin exist at 
lower levels (Yang 2001; Yang and Kallio 2002). Lycopene is an important antioxidant present 
at high levels (8 mg/100 g FW) in SB berries, typically even higher in tomatoes (0.9–11 mg) 
(Malinowska and Olas 2016). Compared to the content in the berries, the level of total 
carotenoids is lower in sea buckthorn leaves (∼4 mg/100 g DW) (Pop et al. 2014). 
Tocopherols and tocotrienols (also referred to as vitamin E) are important bioactive 
components in SB berries. They are in human plasma the major lipid-soluble antioxidants, 
which play an important role in protection against oxidative stress (Ranard et al. 2019). Their 
total concentrations ranged from 8 to 32 mg/100 g FW in seeds and from 6 to14 mg/100 g FW 
in the whole berries, respectively (Kallio, Yang, and Peippo 2002). The large variation in 
amounts and composition of tocopherols and tocotrienols were found to be due to the genetic 
background, harvest year and date. The berries of ssp. sinensis contain more total tocopherols 
and tocotrienols than ssp. mongolica, among which α-tocopherol is the major compound (~80 % 
of total), the content varying from 43 to 116 mg/kg FW in berries (Kallio, Yang, Peippo, 
Tahvonen, et al. 2002). SB berries have higher levels of α-tocopherol in the early ripening 
period, while at later dates, δ-tocopherol levels increase (Andersson et al. 2008). Concentration 
of γ-tocopherol is higher in green berries, but the content rapidly declines to traces when berries 
turned from green to olive-yellow (Zadernowski et al. 2003). Compared to berries, SB leaves 
contain higher level of tocopherols (72–154 g/kg DW) (Sytařová et al., 2020). 
 16 
Oils from SB seeds and pulp are rich in phytosterols at concentration varying from 1.2 to 
1.8 mg/g in the seed oil, and from 12 to 23 mg/g in the pulp oil of berries (Li et al. 2007; 
Sajfrtová et al. 2010). The total quantity of phytosterols in SB oil exceeds that of soybean oil 
by 4-20 times (Stobdan et al. 2013). Twenty-two different phytosterols have been reported in 
SB seed oil, among which β-sitosterol is the most important constituting 57−76 % of the seed 
sterols and 61−83 % of the pulp/peel sterols, respectively (Li et al. 2007; Yang et al. 2001). 
The concentration shows only little variation between subspecies and collection sites of SB, 
but depends on the method of extraction (Li et al. 2007; Yang et al. 2001). Phytosterols of SB 
have been reported to lower serum cholesterol concentrations and decrease the risk of 
hypercholesterolemia-induced cardiovascular disorders (Olas 2018). 
SB oil displays anti-atherogenic, cardio-protective, anti-platelet aggregation, antiulcer and 
anti-depressive properties, which have been attributed to carotenoids, tocopherols, tocotrienols 
and sitosterol, and demonstrated using cell cultures, animal models, and clinical trials in 
humans (Basu et al. 2007; Johansson et al. 2000; Olas 2018; Xing et al. 2002). An eyelid spray 
emulsion-enriched SB oil has a significant beneficial effect on eye dryness, burning and other 
symptoms of human (Larmo et al. 2019).  
Sugars, sugar derivatives and sugar alcohols  
The major monosaccharides in sea buckthorn berries are glucose and fructose, and trace 
amounts of sucrose has also been determined (Tiitinen, Hakala, and Kallio 2005; Yang 2009; 
Zheng et al. 2012). β-EG is a compound characteristic for SB and rarely found in other edible 
berries or fruits. The presence of β-EG was reported in SB for the first time in 2006 (Tiitinen 
et al. 2006b). Also, presence of the corresponding methyl derivative in SB was reported later 
(Lindstedt et al. 2014). In some berries of H. rhamnoides ssp. rhamnoides, β-EG is the most 
abundant compound (1.9 g/100 ml juice) in the sugar fraction (Yang 2009). The content of β-
 17 
EG varies significantly with genetic background, harvesting time, and origin (Yang 2009; 
Zheng et al. 2011; Zheng et al. 2012). During ripening and over-ripening, the content of β-EG 
in the berries of ssp. rhamnoides increased, whereas the content of glucose decreased (Yang 
2009).  
Ethyl and methyl glucosides have been reported to have some relevance to physiological 
effects (Higgins et al. 1996; Storlien et al. 1986). However, literature is hardly available on 
their absorption and metabolism in humans or animals. Based on a NMR metabolomics study, 
both ethyl and methyl glucosides have been detected in human plasma and urine after a single 
meal of SB berries (Lindstedt et al. 2014) Moreover, β-EG was shown to decrease blood 
pressure in SHR rats and stroke-prone spontaneously hypertensive rats after intravenous 
injection of the compound (Matsubara et al. 1989; Sawabe and Matsubara 1999). β-EG was 
rapidly absorbed in the blood stream and easily excreted in urine after oral administration, but 
traces of β-EG were recognized in the rat even 24 h after oral administration (Mishima et al. 
2008).  
The presence of inositols, methyl inositols and L-quebrachitol in SB was reported for the 
first time in 2009 (Kallio et al. 2009). In the berries of ssp. sinensis, L-quebrachitol was the 
most abundant sugar alcohol (0.2–1.1 g/100 mL juice), followed by methyl-myo-inositol, and 
myo-inositol (Zheng et al. 2011). The genetic background of SB berries affects the content of 
methyl inositols, which is higher in ssp. sinensis (means 0.8 g/100 mL juice) than in ssp. 
rhamnoides (means 0.3 g/100 mL juice) and ssp. mongolica (means 0.2 g/100 mL juice) (Yang 
2009). Based on the NMR fingerprints, L-quebrachitol is important for the identification of ssp. 
sinensis as the most distinctive marker in SB berries (Su et al. 2014). L-quebrachitol, inositols 
and methyl inositols are significant regulators of physiological processes of plants and humans. 
L-quebrachitol may increase the fasting insulin level in plasma in db/db mice (Xue et al. 2015). 
The earlier studies found that orally administered inositols can decrease hyperglycemia and 
 18 
hyperlipidemia, and have an influence on insulin secretion and generation of mediators in 
diabetic rats (Fonteles et al. 2000; Nascimento et al. 2006). Further, insulin resistance-related 
diseases, such as polycystic ovary syndrome and gestational diabetes are related to 
derangements in inositol metabolism (Muscogiuri et al. 2016).   
Organic acids 
      Malic acid is the most abundant organic acid in SB berries, followed by quinic and ascorbic 
acids (vitamin C) (Tiitinen, Hakala, and Kallio 2005; Yang 2009; Zheng et al. 2012). Ascorbic 
acid and even citric acid have antioxidant properties (Hraš et al. 2000). Ascorbic acid is among 
the major antioxidants in SB berries, and its content correlates positively with the free-radical 
scavenging capacity of the corresponding fraction (Gao et al. 2000). Besides this, ascorbic acid 
cures the variety of clinical symptoms known as scurvy, repairs the mucosa injuries, as well as 
acts as a co-substrate of many important dioxygenases, and has an indirect effect on mRNA 
transcription (Arrigoni and De Tullio 2002; Gupta et al. 2006; Levavasseur et al. 2015). In 
some studies, ascorbic acid content varied from 50 to 4 000 mg/100 g in fresh berries, and the 
mean value was around 1 000 mg/100 g FW, which is higher than the levels found in many 
other berries, such as strawberries (65 mg/100 g FW), raspberries (29 mg/100 g FW), and 
blackberry (21 mg/100 g FW) (Korekar et al. 2014; Lee and Kader 2000; Tang 2002). Genotype, 
time of harvest, processing techniques, and storage conditions all affect the content of ascorbic 
acid in SB berries and berry products (Gutzeit et al. 2008; Kallio, Yang, and Peippo 2002; Xu 
et al. 2015). Moreover, ascorbic acid content has shown a positive correlation with the total 
phenolics and carotenoids in SB berries (Korekar et al. 2014). The SB leaves were reported to 
contain ascorbic acid on average 2100 mg/kg DW (Jaroszewska and Biel 2017).  
Bioactive compounds eliciting taste of sea buckthorn 
 19 
Consumers preferred a red color of SB berry, however, color and intensity of aroma effect 
only less on the pleasantness of berry (Tang, Kälviäinen, and Tuorila 2001). A balanced sweet 
taste may be the major characteristics contributing to the sensory appeal of berries and berry 
products, and it also affects the preferences and choices of consumers (Geertsen et al. 2016; 
Laaksonen et al. 2016). The sugar/acid ratio, a crucial factor for sweetness, is much lower in 
SB than in most of the common berries. Astringency and bitterness of SB are typically 
perceived as negative factors limiting its consumption (Ma et al. 2017a). These sensory 
properties also present challenges for exploitation of SB berries and berry-based products as 
ingredients in food industry. Hence, it is crucial to study the role of various metabolites in the 
sensory profile of SB. More knowledge is needed for improving the sensory quality and 
consumer acceptance of SB products. Figure 2 illustrates sensory profile and how bioactive 
compounds contribute to the taste properties of sea buckthorn berries.   
Sweetness and sourness 
Sweetness is a key factor influencing the overall liking of a berry, thus affecting also the 
pleasantness of sea buckthorn berries and berry products (Laaksonen et al. 2016; Tang, 
Kälviäinen, and Tuorila 2001). SB is perceived with low intensity of sweetness. The two major 
sugars, glucose and fructose, as well as the sugar/acid ratio, are responsible for the sweetness 
(Tiitinen, Hakala, and Kallio 2005). The sugar derivatives or alcohols, e.g. L-quebrachitol 
showed only little correlation with the sensory attributes (Ma et al. 2017b).  
Sea buckthorn berries are characterized with intense sourness due to the high content of 
malic acid (Tiitinen, Hakala, and Kallio 2005; Tiitinen et al. 2006b). Ascorbic and quinic acids 
also contribute to sourness. Moreover, quinic acids has a close association with astringency in 
SB juices (Ma et al. 2017a). The intensity of sourness is negatively related to sweetness but 
positively with astringency of the juices (Tiitinen, Hakala, and Kallio 2005; Tiitinen et al. 
 20 
2006b). Citric acid is related to the liking of SB juices, but it has less effect on the sourness 
(Ma et al. 2017a)  
Astringency 
     Astringency is an important sensory characteristic of SB berries, of which mouth-drying 
and puckering astringency are the main sub-qualities (Ma et al. 2017b; Tiitinen, Hakala, and 
Kallio 2005). Phenolic compounds are considered as the major contributors to astringency of 
foods and beverages (Ma et al. 2014; Soares et al. 2017). Notable abundance of FGs and PAs 
have been reported in SB berries (Ma et al. 2016; Yang et al. 2017). Compared to the PAs, FGs 
played a more important role in astringency of SB purees (Ma et al. 2017b). FGs have been 
found to induce a multi-astringent sensation even at low concentration. Among them, rutin has 
been reported to elicit astringent perceptions already at very low concentrations (0.61 mg/L) in 
bottled water (Scharbert, Holzmann, and Hofmann 2004; Schwarz and Hofmann 2007). SB is 
rich in rutin, the average level being 10 mg/100 g FW in the berries of ssp. sinensis (Ma et al. 
2016). However, quercetin glycosides are reported to have little impact on the astringency in 
SB purees of six cultivars studied (Ma et al. 2017b). Isorhamnetin glycosides dominate in the 
FGs of SB berries, of which isorhamnetin-3-O-sophoroside-7-O-rhamnoside (Is-3-S-7-Rh) has 
close association with the astringency of SB purees, although the sensory thresholds of 
individual isorhamnetin glycosides for astringency have not been established (Ma et al. 2016; 
Ma et al. 2017b).  
Proanthocyanidins are key compounds for astringency of wine, the composition of the 
monomeric units playing a more important role than other PA variables such as the total 
concentration and degree of polymerization (Quijada-Morín et al. 2012). Moreover, the type 
of specific linkage between the monomeric units also affects the astringent intensity, e.g. 
catechin‐(4→8)‐catechin, being less astringent than either catechin‐(4→6)‐catechin or 
catechin‐(4→8)‐epicatechin (Peleg et al. 1999). Composition and contents of the PA oligomers, 
 21 
such as dimers and trimers, have been investigated in the SB berries (Kallio et al. 2014). 
Monomers are significantly less astringent than the dimers or trimers, whereas the latter two 
do not differ significantly from each other. Monomers and dimers are crucial astringent 
compounds in red wine (Hufnagel and Hofmann 2008). However, these well-known astringent 
components have been reported to show less astringent impact than FGs in the SB purees 
studied (Ma et al. 2017b). The content of PAs and the PC/PD ratio affect the astringent 
properties of black currant juices. Furthermore, higher PC/PD ratio and lower content of PAs 
associate with decreased astringency (Laaksonen et al. 2015). 
The astringent sensory thresholds of the monomer (-)-epicatechin (270 mg/L) and the 
trimeric procyanidin C1 (260 mg/L) were about 2.5 times higher than all the dimeric 
procyanidins B1, B2, and B3 (110-120 mg/L), and the monomer (+)-catechin had similar 
threshold as the dimers (Table 2). Procyanidins are more astringent than ETs, but the latter 
have lower thresholds for astringent detection, ranging from 0.2 to 6.3 μmol/L as shown by 
studies carried out with tannins extracted from oak wood chips (Quercus alba L. and Quercus 
robur L.) (Glabasnia and Hofmann 2006). The astringent response is a combined function of 
the ability of a given tannin to bind soluble proteins and its tendency to interact with proteins 
via hydrophobic binding. The ability of PCs to saturate soluble proteins and then to bind 
membrane-bound proteins was more efficient than that of the ETs (Hofmann et al. 2006). SB 
leaves are rich in ETs. However, their contributions to the sensory quality of products derived 
from SB leaves has not been investigated, although the sensory thresholds of two individual 
ETs for astringency and bitterness have been reported (Table 2).  
In addition to the phenolic compounds described above, it is well documented that various 
phenolic acid derivatives and organic acids contribute to the astringent oral sensations of food 
products (Challacombe et al. 2012; Hufnagel and Hofmann 2008; Lawless et al. 1996). 
Puckering astringency has been reported to relate positively to sourness elicited by various 
 22 
organic acids (malic, quinic and ascorbic acids) and the total acid content in SB juices (Ma et 
al. 2017b). An increase in astringency of phenolic compounds is often associated with a 
decrease in pH (Peleg and Noble 1999). Although it has been stated that astringency of bread 
and crackers was linked with p-hydroxybenzoic and vanillic acids (Challacombe et al. 2012), 
their roles in the sensory quality of SB berries or berry product have not been investigated. Due 
to the effects of food matrices on the astringency perceived, some astringent substances, tannic 
acid and alum will taste more astringent in water solution than in more complex matrices, such 
as wine or orange juice (Valentová et al. 2002).  
Bitterness 
The sensation of bitterness is considered to be the most complex taste quality in humans, 
due to the wide variety of compounds that elicit bitterness and to the apparently large number 
of genes encoding receptors for this taste modality. Flavonol aglycones (FAs) may activate the 
taste receptors of bitterness and are perceived as bitter (Roland et al. 2013). Various monomeric 
flavonoids, such as (+)-catechin, (−)-epicatechin and (−)-epigallocatechin, have been reported 
to activate the receptors TAS2R14 and TAS2R39 of human bitter taste (Roland et al. 2013; 
Soares et al. 2013). Among the PAs, the maximum intensity of bitterness follows the order:  
monomers > dimers > trimers (Peleg et al. 1999). Moreover, the specific linkage between 
monomeric units has an influence on bitterness, e.g. the catechin-catechin dimer linked by a 
4→6 bond is more bitter than the catechin-(4→8)-catechin dimer (Peleg et al. 1999). 
In general, most of the phenolic compounds may be perceived as astringent at notably low 
concentrations, but as bitter only at high concentrations. For instance, the taste threshold of 
(+)-catechin for bitterness was 290 mg/L in water in a triangle test, which is much higher than 
the astringent threshold reported (119 mg/L, Table 2) (Hufnagel and Hofmann 2008). 
Castalagin and vescalagin were identified in SB leaves and their thresholds for bitterness were 
 23 
both around 1600 mg/L, which was 1500-fold higher than the astringency thresholds (Table 2) 
(Glabasnia and Hofmann 2006; Hofmann et al. 2006; Moilanen et al. 2015). Additionally, some 
non-bitter, astringent phenolic compounds such as rutin, may enhance intensity of the 
perceived bitterness of some bitter compounds, such as caffeine (Scharbert and Hofmann 2005).  
Interestingly, the well-known bitter PA oligomers (dimers, trimers and tetramers) did not 
correlate closely to the bitter taste in SB purees (Ma et al. 2017b). However, the ratios between 
acids and phenolic compounds, such as acids/PA dimers and acids/FGs, correlated strongly 
and positively to bitterness (Ma et al. 2017b). Beta-D-glucopyranosides are perceived as bitter 
through the binding of TAS2R16 receptor, which is one of 25 TAS2R receptors involved in 
perception of bitterness (Bufe et al. 2002; Meyerhof et al. 2010; Sakurai et al. 2010). β-EG is 
positively related to the bitterness of SB juices, so are the ratios β-EG/acids and β-EG/sugars 
(Ma et al. 2017a). 
Based on the relationship between chemical compounds and sensory quality in a previously 
established PLS model, a model was investigated to predict the sensory properties of SB berries 
(Ma et al. 2017b; Ma et al. 2020). In this model, the mouth-drying astringency of SB berries 
can be predicted the most reliably, having the highest regression coefficients with quinic acid 
(0.12), total PAs (0.12) and I-3-S-7-Rh (0.11). I-3-S-7-Rh is also a strong factor for predicted 
sweetness (-0.07) and sourness (0.08)(Ma et al. 2020).  
Effect of processing on bioactive compounds 
The processing methods and parameters, such as temperature, pressure, pH and time, can 
markedly affect the bio-accessibility and nutritional quality of berry products (Amarowicz et 
al. 2009; Bakkalbaşi, Menteş and Artik 2008; Rodríguez-Roque et al. 2015). Processing of SB 
juice is a complex operation, in which many factors affect the quality of the final products. 
Conventional thermal processing, such as high-temperature-short-time (HTST) treatment, is 
 24 
used for assuring safety and to extend the shelf life of foods. It often causes undesirable changes 
in quality and sensory characteristics of the products, resulting in loss of flavor and 
development of off-flavor. Good stability of ascorbic acid is recognized in the SB juice during 
short-time heating as well as high retention during juicing and consequent technological 
processing (Seglina et al. 2006). Content of the total ascorbic acid in SB juice reduced by 5−11 % 
during the HTST processing which is negligible in comparison to the production of juice 
concentrate. During this process, the loss of ascorbic acid can reach the level of 50 % due to 
the multiple steps such as clarification, filtration and thermo-vacuum evaporation (Gutzeit et 
al. 2008). Non-thermal, high pressure (200−600 MPa) pasteurization process is also applied 
for commercial production of SB juices of superior quality, retaining high antioxidant activity 
to meet demands of the consumers (Alexandrakis et al. 2014). Additionally, high pressure 
homogenization can be used to alter the physical properties and thus to enhance the color by 
reducing the size of the particles in the puree, leading to lighter and more yellow SB products 
(Aaby et al. 2020).  
Sea buckthorn oil is the most recognized product, which can be pressed or extracted from 
the berry pulp or seeds. Supercritical carbon dioxide (SC-CO2) has been widely applied to 
extract oils from both seeds and pulp of SB berries (Yang 2001; Yang and Kallio 2002; Yang 
et al. 2011). The SC-CO2 extraction increases the yield and reduces the thermal and oxidative 
damages of the bioactive components in the extract compared to solvent extraction. For 
example, a higher yield of phytosterols (β-sitosterol) from SB seeds is achieved compared to 
extraction with hexane (Sajfrtová et al. 2010; Yang et al. 2011).             
Although the processing methods of SB have been widely investigated in many studies, 
limited information is available concerning behavior of the bioactive compounds in different 
processes. Heating (50–100 °C) affects significantly the contents of flavonoids in SB extracts, 
but less the contents of other polyphenols and carotenoids (Ursache et al. 2017). However, 
 25 
Aaby et al. (2020) reported that pre-heating treatment (80 °C, 10 min) of the SB berries prior 
to sieving increased the yield of the purees (80 % to 84 %) and concentration of the total 
carotenoids (27 % to 65 %), but had an effect neither on ascorbic acid nor the total phenolics. 
Zvaigzne (2018) showed that ultra-high temperature treatment decreased ~10 % the content of 
total carotenoids compared to its content in fresh SB juices, but had less effect on the highest 
acidity and the contents of ascorbic acid, total phenolics and vitamin E, as well as antioxidant 
capacities (DPPH, FRAP and ABTS). Polyphenols of SB berries, when added to sausages, have 
remarkable thermal stability (remain 70 % on average) (Püssa et al. 2015). Contents of various 
linoleic acid oxylipins and of isorhamnetin glycosides decreased during cooking of the 
enriched sausages, whereas the content of quercetin glycosides remained constant (yield 99 %) 
(Püssa et al. 2015). Along with changes in bioactive compounds, processing conditions have a 
crucial impact on antioxidant activity of SB products (Bakkalbaşi, Menteş and Artik 2008; 
Kyriakopoulou et al. 2013; Ma et al. 2019). The radical scavenging ability of phenolics of SB 
berries extracted by methanol after freeze-drying, was higher than that of the phenolics 
obtained with microwave extraction of fresh berries (Kyriakopoulou et al. 2013).  
Numerous tea-type beverages and powdered products can be manufactured of SB leaves 
through drying, which cause changes in the phytochemicals, texture, shape, and color of the 
leaves (Kyriakopoulou et al. 2013; Ma et al. 2019). After drying of the leaves at four different  
temperatures (50, 60, 80 and 100 °C), contents of the total phenolics, carotenoids and 
chlorophyll reduced by 28–40 %, 42–62 % and 45–71 %, respectively (Guan, Cenkowski, and 
Hydamaka 2005). The SB leaves processed by steam at high temperature (80–90 °C) resulted 
in the highest content of total ETs and phenolics in the tea-type infusions of the leaves. The 
non-thermal drying methods (freeze-drying and air-drying) led to higher antioxidant activities 
in the infusions than the thermal drying (Ma et al. 2019). The change in phytochemicals 
depends on the drying method, time, temperature, and type of compounds (Aaby et al. 2020; 
 26 
Chan et al. 2009; Guan, Cenkowski, and Hydamaka 2005); Ma et al. 2019). The highly 
astringent ETs, such as vescalagin and castalagin, may be converted into less astringent 
degradation products during thermal process (Glabasnia and Hofmann 2007). Thus, the unit 
processes should be designed and optimized to improve sensory quality and maximize the 
protection of targeted bioactive components in SB products.  
Development and challenge of sea buckthorn products in the future 
    Sea buckthorn berries are rich in health-supporting phytochemicals. Therefore, it is 
worthwhile to develop more food and health-care products of different parts of the plant and to 
promote its large-scale exploitation. Selection of the raw materials and technologies influence 
the quality of the products. Furthermore, special challenges in development are due to the 
sensory qualities of SB berries, especially astringency and bitterness elicited by the wide set of 
the phenolic compounds. Astringency and bitterness are generally considered as negative 
factors reducing the consumer acceptance and liking of berry products (Lesschaeve and Noble 
2005; Verbeke 2006). However, astringency and bitterness can provide complex and balanced 
tastes in some specific types of beverages, such as coffee, tea, beer and wine (Geertsen et al. 
2016). Certain degree of complexity in taste may enhance consumer acceptance of fruit juices 
as well.    
The oral sensations of pure SB juice (average pH 3.1) characterized by high acidity need to 
be balanced to obtain palatable products (Beveridge et al. 1999; Laaksonen et al. 2016; Tiitinen, 
Hakala, and Kallio 2005). Thus, to reduce the sourness and astringency, some fruits, such as 
papaya and grapes, are often added to SB juice at varying proportions to produce mixed 
beverages (Selvamuthukumaran, Khanum, and Bawa 2007). Malolactic fermentation (MLF) 
may also be used to reduce the content of malic acid and thus the sourness of SB juice, 
maintaining simultaneously the health-promoting phenolic compounds (Markkinen et al. 2019; 
 27 
Tiitinen et al. 2006a; Tiitinen et al. 2007). Additionally, the perception of bitterness can be 
reduced by blocking the bitter taste receptor hTAS2R39. E.g. three 6-methoxyflavanones were 
shown to reduce hTAS2R39 activation by epicatechin gallate (Roland et al. 2014). SB berries 
have been used as a novel probiotic cell immobilization carrier, which improved the 
physicochemical characteristics and the aroma of frozen yogurts (Terpou et al. 2019). Although 
the sensory properties limit the consumption of SB berries as fresh fruit and as raw material for 
food processing to some extent, SB products from berries and leaves have a great potential as 
healthy food as well as raw materials and ingredients for health care products and animal 
nutrition. 
Recently, tea-type beverages processed from sea buckthorn leaves have been developed, 
which are sensory acceptable to consumers and contain high content of antioxidants (Ma et al. 
2019). Besides the SB leaves, the current side-streams such as branches and bark, as well as 
residues from juice and oil extraction can be processed into value-added products, since they 
still retain abundant valuable nutrients and non-nutritive bioactive compounds (Puganen et al. 
2018; Sharma et al. 2008). 
Conclusions  
Sea buckthorn is rich in nutritive and non-nutritive bioactive components, which are 
associated with various health-promoting effects. Extensive research efforts and results have 
shown the potential health effects of different fractions and products of sea buckthorn berries 
and leaves in management of chronic health problems, such as metabolic disorder and 
inflammation. However, the majority of the research has been carried out in vitro and in animal 
models. More human intervention studies are needed to validate the health benefits of sea 
buckthorn berries and products. Further, there is an insufficient understanding of the 
contribution of individual bioactive compounds or groups of bioactive compounds to 
 28 
physiological effects observed of sea buckthorn berries, leaves, as well as fractions and 
products derived from the berries and leaves.  
Consumption of sea buckthorn is often limited by the unique sensory properties 
characterized by high intensities of acidity, astringency and bitterness. Only a very limited 
number of studies have been performed to determine the components contributing to sensory 
attributes of sea buckthorn berries and products. Bioactive constituents, such as phenolic 
compounds, vitamin C, organic acids, and sugar derivatives, likely have a strong influence on 
the sensory properties, especially astringency and bitterness of the sea buckthorn berries and 
berry-based products. More research is needed to understand the contribution of different 
compounds and their interaction with food matrix on the perception of the sensory qualities. It 
is also important to explore the role of lipophilic components as well as interactions between 
different components of different groups of compounds (lipophilic vs. hydrophilic; volatile vs. 
non-volatile) in the determining the taste and flavor profile of sea buckthorn. Further studies 
are also needed to improve the understanding on the impact of processing on bioactive and 
flavor compounds in sea buckthorn and to develop innovative processing technologies for 
targeted modification of the composition and sensory properties of sea buckthorn products. 
Such knowledge is needed to guide plant breeding and product development to improve 
orosensory qualities of sea buckthorn berries and products, which is essential for promoting 
the consumer acceptance and consumption of sea buckthorn. 
 
 
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Figure captions 
Figure 1. The classification, example structures and potential health benefits of bioactive 
compounds in the sea buckthorn berries and leaves.  
 46 
Figure 2. Sensory profile and compounds contributing to taste properties of sea buckthorn 
berries. The data of sensory profile is based on the mean of 4 European cultivars (Tytti, Terhi, 
Hergo and Leikora)(Ma et al. 2017b). 
 
Table 1. Concentrations of main bioactive components in sea buckthorn berries and leaves. 
Components1  Berries2 References Leaves2 References 
Flavonol glycosides  
Q-3-O-rutinoside 
3.8–10 mg/100 g FW (Ma et al. 2016) 0.5–0.8 mg/g DW (Ma et al. 2019) 
1.3-7.4 mg/100 g FW (Yang et al. 2009) 
0–76.1 mg/100 g 
DW (Pop et al. 2013)       
Q-3-O-glucoside 3.2–8.8 mg/100 g FW (Ma et al. 2016)   
Q-3-O-sophoroside-7-
O-rhamnoside 
1-120 mg/100 g DW (Chen et al. 2007) ~20 mg/100 g DW (Pop et al. 2013) 
I-3-O-glucoside 
5.3–11 mg/100 g FW (Ma et al. 2016) 0.3–0.4 mg/g DW (Ma et al. 2019) 
1-134 mg/100g DW (Chen et al. 2007)   
I-3-O-rutinoside 15–38 mg/100 g FW (Ma et al. 2016) 0.7–1.8 mg/g DW (Ma et al. 2019) 
 11-292 mg/100g DW (Chen et al. 2007) 
0–49.6 mg/100 g 
DW 
(Pop et al. 2013) 
I-3-O-glucoside-7-O-
rhamnoside 
8.5–22 mg/100 g FW (Ma et al. 2016) 2.0 mg/g DW (Ma et al. 2019) 
 15-44.7 mg/100 g FW (Yang et al. 2009) 
69.1–111 mg/100 g 
DW 
(Pop et al. 2013) 
I-3-O-sophoroside-7-
O-rhamnoside 
5-176 mg/100 g DW (Chen et al. 2007) 
25-44.6 mg/100 g 
DW 
(Pop et al. 2013) 
K-7-O-rhamnoside 4-48 mg/100 g DW (Chen et al. 2007)   
Acylated FGs 
157–260 mg/100 g 
DW 
(Chen et al., 2013) 
24–115 mg/100 g 
DW 
(Chen et al. 2013) 
Total FGs 
23–407 mg/100 g FW 
 
~300 mg/100 g FW 
 
27-130 mg/100 g FW 
(Ma et al. 2016) 
 
(Teleszko et al. 2015) 
 
(Yang et al. 2009) 
14 mg/g DW 
 
1118 mg/100 g DW 
(Ma et al. 2019) 
 
(Pop et al. 2013) 
  
Proanthocyanidins (mg/100g DW)  
Dimers 1.0–8.9   
(Yang et al. 2017; Yang 
et al. 2016; Kallio et al. 
2014) 
 
 
Trimers 1.3–9.5   
tetramers 0.8–7.1   
PA oligomers 3.7−25.5   
Total PAs 167–1941  
    
Ellagitannins (mg/g DW)    
Stachyurin   7.8–8.5 (Ma et al. 2019) 
Casuarinin   11.5-13.0 (Ma et al. 2019) 
mean    76 
(Moilanen et al. 
2015; Suvanto et al. 
2018). 
Total ETs   29.2–109.1  
(Ma et al. 2019; 
Suvanto et al. 2018) 
Phenolic acids (mg/kg DW)    
 47 
Gallic acid 7.9–15.9 (Bittová et al. 2014) 
19.1–79.1 
 
(Bittová et al. 2014) 
 0.2–7  
(Zadernowski et al. 
2005) 
28.4–102.4 
(Sytařová et al. 
2020) 
 1–2.6 mg/ml juice (Rösch et al. 2003)   
Salicylic acid 21–47.5 
(Zadernowski et al. 
2005) 
  
p-coumaric acid 1.4–22.3 
(Zadernowski et al. 
2005) 
8.4–13.4 (Bittová et al. 2014) 
Total phenolic acids 3570–4439  
(Zadernowski et al. 
2005) 
804-4988 
(Arimboor et al. 
2008) 
Total  Phenolics     
 
260-490 
mg/100 g FW 
(Olas 2016) 98 mg/g DW (Ma et al. 2019) 
     
Sugars and orgainic 
acids 
    
β-EG 
0.1–2.0  g/100 mL 
juice 
(Ma et al. 2017; Yang 
2009) 
  
Methyl inosotol 
0.1–1.4 g/100 mL 
juice 
(Yang 2009)   
L-quebrachitol 
0.2–1.1 g/100 mL 
juice 
(Kortesniemi et al. 
2014; Zheng et al. 
2012) 
  
Ascorbic acid 
56–3909 mg/100 g 
FW 
360-1676 mg/100 g 
FW 
52- 130 mg/100 g FW 
(Korekar et al. 2014) 
 
(Tiitinen et al. 2006b) 
 
(Teleszko et al. 2015) 
2139.6 mg/kg DW 
(Korekar et al. 
2014)  
 0.98–3.65 g/kg DW (Sytařová et al. 2020) 
22.81–46.32 g/kg 
DW 
Sytařová et al. 
2020) 
Lipids     
triacylglycerols  15–85 g/kg FW 
(Kallio, Yang, Peippo, 
Tahvonen, et al. 2002) 
   
glycerophospholipids  1.8–2.5 g/kg FW 
(Kallio, Yang, Peippo, 
Tahvonen, et al. 2002) 
  
Total tocopherols and 
tocotrienols 
6–14 mg/100 g FW 
(Kallio, Yang, Peippo, 
Tahvonen, et al. 2002; 
Yang 2001) 
 
 
 
 6.98–29.91 g/kg DW Sytařová et al. 2020) 
71.54–153.99 
g/kgDW 
Sytařová et al. 
2020) 
α-tocopherol 
4.3–11.6 mg/100 g 
DW 
(Kallio, Yang, Peippo, 
Tahvonen, et al. 2002) 
  
Total carotenoids 
1–120 mg/100 g FW (Yang 2001) 
3.5–4.2 mg/100 g 
DW 
(Pop et al. 2014) 53–97 mg/100 g DW (Pop et al. 2014) 
6–24 mg/100 g FW (Teleszko et al. 2015) 
β-carotene 0.2–17 mg/100 g FW (Yang 2001)   
Lycopene 8 mg/100 g FW 
(Malinowska and Olas 
2016) 
  
Total sterols 34–52mg/100g FW (Yang et al. 2001)   
Phytosterols 13–20 g/kg FW (Bal et al. 2011)   
 
6–13 mg/100 mL lipid 
fraction 
(Teleszko et al. 2015)   
1 I, isorhamnetin; Q, quercetin; K, kaempherol; FGs, flavonol glycosides; PAs, proanthocyanins; ETs, ellagitannins; β-EG, 
Ethyl β-D-glucopyranoside 
 2 FW = fresh weight, DW = dry weight  
 48 
 
Table 2. Human taste recognition thresholds for astringency, bitterness and sourness of main 
bioactive components in sea buckthorn. 
           Taste threshold (mg/L) 
Component Astringency Bitterness Sourness 
Sensory 
test 
Medium References 
Flavonols        
Q-3-O-rutinoside 0.61     
Triangle 
test 
Water 
(Scharbert, Holzmann, and 
Hofmann 2004) 
I-3-O-glucoside 1.1   nd  
Triangle 
test 
Water 
(Hufnagel and Hofmann 
2008) 
Q-3-O-glucoside 4.6    
Triangle 
test 
Water 
Hufnagel and Hofmann 
2008) 
Q-3-O-
rhamnoside 
8.9 8.9  Pairs Water (Delcour et al. 1984) 
I   0.04  ns Water (Roland et al. 2013) 
Q 10 10  Pairs 
5% 
ethanol 
(Roland et al. 2013) 
K 20 20  Pairs 
5% 
ethanol 
(Roland et al. 2013) 
Flavanols         
(+)-Catechin 119 290   
Half-
tongue 
test 
Water 
(Hufnagel and Hofmann 
2008) 
(-)-Epicatechin 270  270   
Half-
tongue 
test 
Water 
(Hufnagel and Hofmann 
2008) 
Theaflavins 9    
Half-
tongue 
test 
Water 
(Scharbert, Holzmann, and 
Hofmann 2004) 
Procyanidin B1 139  231   
Half-
tongue 
test 
Water 
(Hufnagel and Hofmann 
2008) 
Procyanidin B2 110  280   
Half-
tongue 
test 
Water 
(Hufnagel and Hofmann 
2008) 
Procyanidin B3 116  289   
Half-
tongue 
test 
Water 
Hufnagel and Hofmann 
2008) 
Procyanidin C1 260  347   
Half-
tongue 
test 
Water 
(Hufnagel and Hofmann 
2008) 
Trimers and 
tetramers 
4.1 4.1  
Triangle 
test 
Water (Delcour et al. 1984) 
Oligomers 0.2    
Half-
tongue 
test 
Water 
(Rossi and Singleton 
1966) 
Condensed 
tannins 
0.035 0.12  Pairs Water 
(Rossi and Singleton 
1966) 
Ellagitannins         
Castalagin 1.03  1578  
Half-
tongue 
test 
Water 
(Glabasnia and Hofmann 
2006) 
Vescalagin 1.03  1578   
Half-
tongue 
test 
Water 
(Glabasnia and Hofmann 
2006) 
 Phenolic acids       
Gallic acid 45   nd  
Triangle 
test 
Water 
(Hufnagel and Hofmann 
2008) 
 49 
Caffeic acid 13 nd  
Triangle 
test 
Water 
(Hufnagel and Hofmann 
2008) 
Ellagic acid 1.99     
Triangle 
test 
Water (Hofmann et al. 2006) 
Chlorogenic acid   10 10 
Triangle 
test 
5% 
ethanol 
(Maga and Lorenz 1973) 
 Sugar derivative       
β-EG   1100  
Duo-tri 
test 
Water (Ma et al. 2017a) 
 Non-phenolic organic acids       
Malic acid      494 
Triangle 
test 
Water 
(Rotzoll et al. 2006; 
Scharbert and Hofmann 
2005) 
Citric acid     499 
Triangle 
test 
Water 
(Rotzoll et al. 2006; 
Scharbert and Hofmann 
2005) 
Quinic acid  10   ns Water (Frank et al. 2006) 
I, isorhamnetin; Q, quercetin; K, kaempherol; β-EG, Ethyl β-D-glucopyranoside. 
ns, not specified, meaning that methodology for evaluation of recognition sensory threshold is not specified in reference.  
nd, not detected for bitter thresholds.
 50 
Figure 1. 
 51 
Figure 2.