Botany

Nutritional composition of saskatoonberries - A review

Journal: Botany

Manuscript ID cjb-2019-0191.R2

Manuscript Type: Review

Date Submitted by the 20-Oct-2020 Author:

Complete List of Authors: Fang, Jim; University of Saskatchewan,

Keyword: Saskatoon berry, nutrition, , , fiber

Is the invited manuscript for consideration in a Special Not applicable (regular submission) Issue? : Draft

© The Author(s) or their Institution(s) Page 1 of 37 Botany

Nutritional composition of saskatoonberries - A review

Jim Fang*

College of Pharmacy and Nutrition

University of Saskatchewan

Saskatoon, Saskatchewan S7N 5E5

Canada

Draft Running title: Saskatoonberries

*Correspondence

College of Pharmacy and Nutrition

107 Wiggins road

University of Saskatchewan

Saskatoon, Saskatchewan S7N 5E5

Canada

E-mail: [email protected]

© The Author(s) or their Institution(s) Botany Page 2 of 37

Abstract

Saskatoonberry ( alnifolia Nutt., ), also known as serviceberry, is a native to the northern prairies and plains of North

America. Saskatoonberries are an excellent source of typical health-promoting nutrients such as fibers, minerals (manganese, , , , iron) and vitamins (tocopherol, , , ascorbic acid, riboflavin, thiamin, ). Saskatoonberries are rich in health-promoting phytochemical compounds, mainly (-3-galactoside, cyanidin-3-glucoside, cyanidin-3-arabinoside, and cyanidin-3-xyloside), flavonols (-3-galactoside, quercetin-3-glucoside, quercetin-3-arabinoglucoside, quercetin-3-arabinoside, quercetin-3-rutinoside, and quercetin-3-xyloside),Draft proanthocyanidins (A- and B-type procyanidins with different degrees of polymerization), phenolic acids, and volatile components. Saskatoonberry seed oil contains 99 g/100g of total fat and out of which

9.8 g/100g is present as saturated fat, 31 g/100g as monounsaturated fat and 58 g/100g as cis-cis polyunsaturated fat. Saskatoonberry seed oil contains approximately 10 g/kg of phytosterols in total (β-sitosterol, Δ5-avenasterol, campesterol and stigmasterol). In mature (purple) saskatoonberries, the content of cyanogenic compounds amygdalin and prunasin were found to be 42-118 mg/kg and 5-17 mg/kg of fresh weight, respectively.

These compounds are confined to seeds which greatly slows down their absorption when saskatoon are consumed. In summary, as an emerging functional food, saskatoonberries are an excellent source of minerals, fiber and phenolic compounds, such as anthocyanins and proanthocyanidins.

Keywords: Saskatoon berry, Amelanchier alnifolia Nutt., Rosaceae, nutrition, composition, polyphenol, mineral, fiber

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Introduction

The name “saskatoon” is derived from the Cree language word misâskwatômina

(Mis-sack-qua-too-mina), which means “the of the tree of many branches”.

Saskatoonberry (Amelanchier alnifolia Nutt., Rosaceae) is a deciduous shrub native to

the northern prairies and plains of North America. The USDA recognizes

saskatoonberry as serviceberry for trade purposes. While the saskatoon fruit is often

called a berry, it is actually a pome fruit, reflecting the ’s placement within the

Rosaceae. The city of Saskatoon in Saskatchewan, Canada is named after this local

fruit.

Saskatoonberries are gaining popularity in Canada and internationally partly due to a resemblance to the fruit of the blueberries.Draft Saskatoonberry and blueberry share many similar characteristics that are important to consumers, including fruit color,

shape, texture and nutrition. However, the blueberries are a member of the Ericaceae

family, and related to cranberry (V. macrocarpon Ait.) and the lingonberry (V. vitisidaea

L.).

Several review papers have been published on saskatoonberries (Mazza 2005;

Jurikova et al. 2013). The current article systematically summarizes the latest findings

on the composition of this fruit variety.

Characteristics and Commercialization

In 1918, W. D. Albright started the first saskatoon orchard in Peace River, Alberta

and the fruit has been grown on the Canadian prairies commercially since the mid-

1960s. The current annual production is about 654 metric tons (AAFC 2017). Recently,

saskatoonberries have been introduced and cultivated in regions with a cold climate,

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such as northern Europe (Jurikova et al. 2012; Lavola et al. 2012; Zurawicz et al. 2014).

Saskatoonberries adapted to local climate so well that new varieties of saskatoonberries are being developed in Poland (Butorova et al. 2016; Lachowicz et al. 2017). Some of the new have significantly higher sugar content (Lachowicz et al. 2017). The cultivation of saskatoonberries is much easier than high-bush blueberries, because it is less demanding in terms of soil quality and weather conditions.

Currently, all Canadian commercial cultivars of Amelanchier alnifolia have been selected from wild populations for their resistance to diseases, high yields, as well as fruit size, flavor, color and texture. Common cultivars of saskatoonberries grown in Canada include Honeywood, Martin,Draft Northline, Pembina, Smoky and Thiessen (Table 1) (Zatylny et al. 2005), each with subtly different flavor profiles (Lutz 1994; Kidd 2006).

Other Canadian varieties include: Altaglow, Bluff, Buffalo, Elizabeth, Forestburg,

Killarney, Lee #3, Moonlake, Nelson, Paleface, Parkhill, Pearson II, Regent, Sturgeon, and Success. Some of the new cultivars from the Polish breeding program are

Lamarckii Balerina, Ostravsky, Tisnovsky skolsky, Tisnovsky velkoplody (Butorova et al.

2016). The mature fruit is a purple blueberry-like pome 1–1.7 cm in diameter.

Saskatoonberries are good to eat raw, tasting somewhat like blueberries with a chewier texture, accented by the almond-like flavor of the seeds.

[Table 1 near here]

The indigenous peoples of the North America have used saskatoonberries in their diet for centuries. Berries provided them with the much-needed minerals and vitamins along with the flavor and sweetness. Native communities consumed

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saskatoonberries in many different ways. Fruits and juice are added to lichens, meat

soups and puddings, or dried fruits are mixed with meat (FAO 1991). Juice was used to

heal stomach ailments, and used as eye and eardrops.

The early settlers in the Canadian prairies described the dark blue sweet berry as

a staple in the local diet providing nourishment that likely prevented malnutrition-related

diseases such as scurvy (FAO 1991). Saskatoonberries were first picked from the wild,

then from home gardens and now from commercial plantings. The fruit was eaten fresh,

dried, or preserved to add both interest and nutrition to winter menus. Preserves

included canned saskatoonberries with added sugar or honey, jams, conserves, and

jellies. Saskatoonberries can be dried whole or as a fruit leather or pemmican type product. The berries also produce juice,Draft which can be added to warm and cold beverages.

Modern methods of processing, freezing and packaging have greatly increased

the year-round consumption of saskatoonberries. Table 2 lists the saskatoonberry

products currently being sold in Canada.

[Table 2 near here]

Essential nutrients

The macronutrient composition of saskatoonberries is summarized in Table 3.

Mazza (2005) reported that saskatoon fruits are slightly lower in water content than

blueberries (79.6% vs. 84.2%) (Mazza 2005). Accordingly, fresh saskatoonberries

contain more fiber and protein than blueberries and can better retain their form when

used in baked goods. Each 100g of fresh saskatoonberries contains 0.63 g of ash,

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which is more than twice that of blueberries. Ash is an indication of the presence of minerals in the fruit. Essential nutrient composition data for saskatoonberries are presented in Tables 4-7. Fatty acids are confined to the seeds (see Table 13).

[Table 3 near here]

Carbohydrate

Fresh saskatoonberries contain 18.5% carbohydrates (Table 3) similar to that of blueberries. The composition and amount of sugars in saskatoonberries are also similar to those in blueberries (Mazza 2005). Saskatoonberries contain around 2.4 to 3.9 g sugar per 100g (Table 4). These sugarsDraft are mainly glucose and fructose, which are known to taste sweeter than sucrose.

[Table 4 near here]

Protein

Most amino acids are present in higher concentrations in saskatoonberries than in blueberries, possibly due to the higher protein content of saskatoonberries (Mazza

2005). Saskatoonberries contain more number of seeds, which are rich in protein. The berries carry all nine indispensable amino acids for humans, though not in significant quantities (Table 5).

[Table 5 near here]

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Vitamins

The vitamin composition of saskatoonberries is given in Table 6. Vitamins found

in saskatoonberries include vitamins A, B9, C, B2, and E. Small amounts of B1, B6, B5,

B6, and H are also found (Table 6). Although saskatoonberries are not one of the best

sources of vitamins, their vitamin content is similar to that of blueberries (Mazza 2005).

[Table 6 near here]

Minerals Mineral composition of the fruitDraft depends heavily on growing conditions. The average mineral content of saskatoonberries is listed in Table 7. On average,

saskatoonberries have significantly higher iron, magnesium, potassium, calcium and

phosphorus levels than blueberries. Saskatoonberries can contribute significantly to the

daily required intake of iron, magnesium, potassium and calcium.

[Table 7 near here]

Phytochemicals

The antioxidant properties of fruits are attributed to . In addition to

being antioxidants, some of these polyphenols are shown to have more specific

biological activities that contribute to overall health benefits (Del Bo et al. 2019; Potì et

al. 2019). It is thus important to study the composition of polyphenols.

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Anthocyanins

The blue and purple colors of saskatoonberries and blueberries originate mainly from anthocyanins. Anthocyanins are water-soluble pigments responsible for the blue, purple, and red color of many fruits, , and . Types and concentrations of anthocyanins vary widely in food (Bhagwat and Haytowitz 2015; Fang 2015). Similar to blueberries, the pigments of saskatoonberries are located primarily in the skin of the fruit.

Anthocyanin contents in saskatoonberries increase as the fruits mature (Green and Mazza 1986; Rogiers and Knowles 1997; Ozga et al. 2006). Saskatoonberries contain a distinct spectrum of anthocyaninsDraft that is different from blueberries, the major components being cyanidin-3-galactoside, cyanidin-3-glucoside, cyanidin-3-arabinoside, and cyanidin-3-xyloside (Ozga et al. 2007; Bakowska-Barczak and Kolodziejczyk 2008;

Lavola et al. 2012). All major saskatoonberry anthocyanins are cyanidin glycosides.

Cyanidin glycosides can be metabolized to protocatechuic acid and vanillic acid (Fang

2014). Phenolic acid metabolites may contribute to the health benefits of anthocyanins because some of these metabolites exhibit antioxidant, anti-inflammatory, and other beneficial properties (Masella et al. 2012). Fruits with higher cyanidin content, such as chokeberries, black raspberries, and bilberries, are more likely to produce anti- inflammatory effects (Joseph et al. 2014). On the other hand, blueberries contain mainly , , and glycosides, which produce unstable phenolic acid metabolites (Fang 2015).

[Table 8 near here]

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Flavonols

Quercetin glycosides are commonly found in fruits of the Roseaceae family such

as apple, peach, plum, and chokeberry (aronia). Quercetin glycosides were also

identified in saskatoonberries. Quercetin-3-galactoside is the most abundant quercetin

glycosides (Table 8). Minor quercetin glycosides include quercetin-3-glucoside,

quercetin-3-arabinoglucoside, quercetin-3-arabinoside, quercetin-3-rutinoside, and

quercetin-3-xyloside.

Proanthocyanidins

Proanthocyanidins are condensed tannins widespread in the plant kingdom where they provide flavor and astringencyDraft to the fruit when consumed. Procyanidins are members of the proanthocyanidin class of flavonoids. They are oligomers with (+)-

catechin or (-)-epicatechin as constituent units. Procyanidins in food can be classified

into A-type and B-type. Proanthocyanidin-B molecules are ubiquitous in nature (e.g.,

strawberry, blueberry, raspberry, blackberry, grape, apple, cherry, mango, banana,

peach, apricot and so on) and may be important as antioxidants. In B-type procyanidins,

the (epi)catechin units are linked through the C4→C8 or C4→C6 interflavan bonds. The

(epi)catechin units in A-type procyanidins are linked by an additional ether bond

between C2 and O7. Cranberry, plums and avocados contain proanthocyanidin-A.

Saskatoon proanthocyanidins are essentially of the procyanidin type (Hellstrom

et al. 2007; Dudonne et al. 2015; Lachowicz et al. 2017). The saskatoonberry

proanthocyanidins range from dimers through heptamers and higher polymers. Thus,

saskatoonberry proanthocyanidins are very similar to those in apple (Hellstrom et al.

2007).

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The reported contents of extractable procyanidin in saskatoonberries varies significantly in the literature (Hellstrom et al. 2007; Hosseinian et al. 2007; Dudonne et al. 2015; Lachowicz et al. 2017). The reason for these differences is not clear and may be due to differences in cultivars, growing conditions, stabilities during storage and analytical methods used. Proanthocyanidins often occur as complex mixtures, which makes their separation difficult. Proanthocyanidin oligomers can be separated by normal phase HPLC according to their degrees of polymerization up to decamers.

However, the isomers often results in overlapping retention times between different procyanidins when UV (Hosseinian et al. 2007) and fluorescence detectors (Hellstrom and Mattila 2008; Dudonne et al. 2015) were used for their quantification. In a recent study using HPLC-MS/MSDraft techniques, saskatoonberries were found to contain both type A and B procyanidins (Lachowicz et al. 2017). Contents of A- and

B-type procyanidins with different degrees of polymerization are presented in Table 9.

The procyanidin content of saskatoonberries need to be reevaluated using normal phase HPLAC-MS/MS methods as more reference standards become available.

[Table 9 near here]

Non-extractable proanthocyanidins (NEPA) are proanthocyanidins which remain in the residues after extraction with aqueous and organic extraction. This is due to their ability to form macromolecules (tannins) or to bind to polysaccharides, protein, and cell walls by covalent bonding (esters and ether), hydrogen bonding, and hydrophobic and hydrophilic interactions. Significant amounts of polyphenolic compounds can be released from the extraction residues when they are treated with acids or enzymes

(cellulases, proteases, digestive enzymes, bacterial enzymes). The content of NEPA

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was shown to range from 11 mg/100 g for pear to 6.7 g/100 g dry weight for red grape

pomace (Perez-Jimenez et al. 2009).

Unextractable proanthocyanidins were measured in saskatoonberries (Hellstrom

and Mattila 2008). Freeze-dried and grounded saskatoonberries were first extracted

three times with a mixture of acetone/methanol/water (2:2:1) (which yield the extractable

proanthocyanidins fraction). The extraction residue was freeze-dried and acid-

hydrolyzed with benzyl mercaptan. The resultant terminal units were quantified using an

HPLC-UV method. Total unextractable procyanidins was determined to be 101 mg/100

g of fresh weight which accounts for over one-third of total proanthocyanidins (Hellstrom

and Mattila 2008). Interestingly, the proanthocyanidinDraft contents in leaves and stems were found to be much higher than in saskatoonberries accounting for 10%–14% of dry biomass

(Lavola et al. 2012). The leaves and stems were suggested to be a potential source for

bioactive procyanidins and condensed tannins.

The analysis of proanthocyanidins poses a challenge and there is currently no

standardized analytical method available (Domínguez-Rodríguez et al. 2017). A more

detailed study is required to clarify the extractable and unextractable proanthocyanidin

profile of saskatoonberries.

Phenolic acids

Phenolic acids are present in berries in free and bound forms. Bound phenolics

may be linked to various plant components through ester, ether, or acetal bonds

(Robbins 2003). Bound phenolic acids were found to be the predominant phenolic acids

in saskatoonberries. 5-Caffeoylquinic acid (neochlorogenic acid), an ester of caffeic acid

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and quinic acid, was found as the major phenolic acid (Table 10) (Dudonne et al. 2015).

This is consistent with the findings of Ozga and coworkers (Ozga et al. 2007). However, a third report found chlorogenic acid as the major phenolic acid in saskatoonberries with neochlorogenic acid as a remote second (Lavola et al. 2012). 3-Caffeoylquinic acid

(chlorogenic acid), 4-caffeoylquinic acid, and ellagic acid, were found to be the minor phenolic acids (Table 10). Protocatechuic acid contents were found to be similar to those reported by other researchers (Lavola et al. 2012).

A number of unknown cinnamic acid derivatives were also detected in saskatoonberries (Ozga et al. 2007; Lavola et al. 2012). In European Juneberries (Amelanchier ovalis), a native EuropeanDraft berry similar to saskatoonberries, hydroxycinnamic acids liberated from soluble esters and glucosides account for 53% and 45% of total hydroxycinnamic acids (Zadernowski et al. 2005). Hydroxybenzoic acids liberated from soluble esters, and glucosides account for 70% and 27% of total hydroxybenzoic acids.

[Table 10 near here]

Organic acids

Organic acids account for 0.5% of the fresh weight of ripe saskatoonberries

(Rogiers and Knowles 1997). Malate was the most concentrated organic acid in mature fruit accounting for about 50% of the total organic acids. Succinate was the predominant organic acid in immature fruit; however, levels declined 14- to 18-folds with advancing

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maturity. The minor organic acids identified in saskatoonberries included quinate,

galacturonate, citrate, pyruvate, cis-aconitate, fumarate, and oxalate.

Each organic acid has its own distinctive taste and contributes to the flavour

differences among different fruits. Acid profile differences may also contribute to other

important characteristics such as fruit color development, decay susceptibility, and

insect and bird predation. Major organic acids in lowbush blueberries are citric, malic,

and quinic acids accounting for 36%, 31% and 20% to total acidity. Citric (75%) and

succinic (17%) acids are the most abundant acids in highbush blueberries (Ehlenfeldt et

al. 1994).

Abscisic acid has recently been shown to improve glucose tolerance and insulin sensitivity and to reduce circulating triglycerideDraft levels and obesity-related inflammation (Zocchi et al. 2017). Saskatoonberries contain a total of 45 μg/100g fresh fruit weight of

abscisic acids (Table 11). Dihydrophaseic acid, phaseic acid and cis-abscisic acid are

the major abscisic acid derivatives found in saskatoonberries (Dudonne et al. 2015).

[Table 11 near here]

Volatile constituents

Earlier studies revealed that benzaldehyde is the major aroma component of

saskatoonberries and amounts to 76-96% of the essence (Mazza and Hodgins 1985). In

a recent study, the typical fruity, sweet, grassy and/or floral aroma of saskatoonberries

was attributed to twenty four compounds (Butorova et al. 2016). The overall profile of

volatiles in saskatoonberries was similar to other berry fruits, and characterized by the

dominant presence of aldehydes (benzaldehyde, hexanal, etc.), alcohols (ethanol,

hexan-1-ol, etc.), esters (ethyl-ethanoate, methyl-ethanoate, etc.) and acids (Butorova

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et al. 2016) (Table 12). The volatile profiles of saskatoonberry are strongly influenced by environmental factors (year to year changes) and cultivars. Significant differences among different production seasons were found in benzaldehyde, hexanal, ethylethanoate and methyl-ethanoate. The blueberry aroma profile is more complex than saskatoonberries with 43 aroma active volatiles identified (Du and Rouseff 2014).

[Table 12 near here]

Composition of saskatoonberry seeds

The composition of saskatoonberry seed oil consists predominantly of fatty acids.

Phytosterols, tocopherols and cyanogenic compounds were identified in saskatoonberry seeds. Other constituents of the seeds Draftare poorly understood and need more characterization.

Composition of seed oil

Fatty acids

The fats and oils are products of fatty acids and glycerol. Each glycerol molecule has three potential binding sites to hold a maximum of three molecules of fatty acids. In the intestine, lipases hydrolyze glycerides to release fatty acids from glycerol. The fatty acids are responsible for the nutritional and health effects of fats. The body uses glycerol to synthesize fat or to produce energy through the glycolysis pathway.

Plant cell membranes and seeds contain fatty acids. Studying the fatty acid composition of saskatoons is important since seeds and pomace are by-products of the saskatoonberry industry. As shown in the Table 13, each 100 g of saskatoonberry seed oil has 99 g of total fat, out of which 9.8 g present as saturated fat, 31 g as

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monounsaturated fat and 58 g is cis-cis polyunsaturated fat. Saskatoonberries do not

contain natural trans-fat.

[Table 13 near here]

Saskatoon oil contains lower concentrations of saturated fatty acids than

unsaturated fatty acids (Table 13). Palmitic acid (16:0) is the dominant saturated fatty

acid of saskatoons, which is 6.6% of the weight of the oil. Stearic acid (18:0), arachidic

acid (20:0) and behenic acid (22:0) are the other significant saturated fatty acids of

saskatoon oil, however, each contributes to less than 1.5 percentage points of the total fat. Draft The compositions of mono- and polyunsaturated fatty acids of saskatoons are

given in the Table 13. Oleic acid (18:1) is the major monounsaturated fatty acid of

saskatoon seed oil, which contributes 30% of the weight of total fat. Trace quantities

(less than 1.5% of total fatty acids) of gadoleic acid (20:1) and palmitoleic acid (16:1)

are present in saskatoon seed oil.

About 58% of fatty acids of saskatoonberries are polyunsaturated fatty acids

(Table 13). The cis-cis linoleic acid (18:2) is the dominant polyunsaturated fatty acids of

saskatoonberries and contributes to 55-59% of the weight of the oil. The fatty acid

profile of saskatoonberries is similar to that of grape seed oil (Prado et al. 2012) and

canola oil (Ackman and Sebedio 1981).

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Phytosterols

As important constituents of plant cell membrane, phytosterols (i.e., sterols and stanols) play an important role in seed germination and development. Humans and animals have an analogous sterol, cholesterol, the structure of which differs from phytosterols only slightly. The phytosterol absorption in the intestine is several times lower than cholesterol absorption. Phytosterols inhibit absorption of cholesterol in the intestine (Fatahi et al. 2019).

Saskatoonberry oil contain approximately 10 mg/kg of phytosterols in total. The phytosterol profile of saskatoons is given in the Table 14. Accordingly, beta-sitosterol is the major phytosterol of saskatoons and each kg of oil contains 7890 mg of the compound. Beta-sitosterol is consideredDraft as the plant sterol with the highest potential to produce steroidal drugs and cholesterol lowering products. Δ5-avenasterol, campesterol and stigmasterol are the other important phytosterols of saskatoons and present in concentrations of 750, 633 and 219 mg/kg oil, respectively.

[Table 14 near here]

Tocopherols

Saskatoonberry seed oils contain high levels of tocopherols (Table 14). Total tocopherols are higher than in virgin olive oil, soybean, peanut, corn, sunflower, and canola oils (260-1000 mg/kg of oil) (Gunstone 2002; Ahmed et al. 2005; Cunha et al.

2006). α-Tocopherol was the predominant derivative and accounted for 87% of the total tocopherols, followed by γ-tocopherol (12%) and δ-tocopherol (0.6%). α-

Tocopherol is the most active tocopherol and one of the most potent phenolic

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antioxidants (Thompson and Cooney 2020). The high levels of α-tocopherol in

saskatoonberry seed oils would also protect the oil from oxidative degradation during

processing and storage.

Cyanogenic glucosides in seeds

Cyanogenic glycosides are a group of natural substances found in that

release cyanide when degraded by enzymes or organic acids. Table 15 lists the berry

seed content of amygdalin, prunasin, and calculated potential content of HCN from the

berries. In mature (purple) berries, the content of amygdalin and prunasin were found to

be 42-118 mg/kg and 5.0-16.5 mg/kg of fresh weight, respectively. The content of cyanogenic compounds is dependentDraft on cultivars. Smoky and Thiessen berries contain higher levels of amygdalin and prunasin than other cultivars. These values are lower

than those reported for some other fruit. For example, the prunasin content of passion

fruit is 285 mg/kg (Chassagne et al. 1996).

No references could be found in the literature concerning any acute toxicity to

humans from saskatoonberries. Saskatoonberries are a traditional food of Native

Americans and is a commercial crop since the1960s. U-pick is one of the favorite

summer activities on the Canadian prairies when consumers can eat as much

saskatoonberries as they can. The cyanogenic compounds in saskatoonberries are

located in the seeds which slows down their absorption (Mandalari et al. 2008).

Furthermore, thousands of plants produce cyanogenic compounds, including common

crops such as mango, cassava, lima beans, bamboo shoots, sorghum, and flax.

Traditional and current consumption trends indicate that levels of cyanogenic

compounds in saskatoonberries is safe. Nevertheless, it is advisable to remove seeds

when processing saskatoonberry powders, because the cyanogenic compounds could

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be more readily absorbed when the seeds are granulated. It has also been suggested that it is possible to breed new saskatoonberry cultivars with larger fruit size and fewer seeds (McGarry et al. 2001).

[Table 15 near here]

Conclusion

Saskatoons have high concentrations of several nutritionally significant minerals.

Each 100 g portion of fresh fruits contain 203 mg potassium, 65 mg calcium, 32 mg magnesium, 26 mg phosphorus andDraft 1.2 mg iron. Saskatoons carry 5.9 g of total in each 100 g of fresh berries. As such, saskatoons are a good source of minerals and fiber from which to potentially develop nutritional supplements.

Phenolic compounds, particularly the anthocyanins and proanthocyanins appear to be the major functional components of saskatoonberries. Major anthocyanins in ripe saskatoonberries are all cyanidin glycosides such as cyanidin-3-galactoside and cyanidin-3-glucoside, cyanidin 3-arabinoside, and cyanidin-3-xyloside.

Saskatoonberries are rich in health-promoting proanthocyanidins, which consist of A- and B-type procyanidins with different degrees of polymerization.

Research carried out during the last thirty years indicates that saskatoonberries are a functional food rich in antioxidants. Functional foods and natural health products are considered as alternative approaches to improve health. Functional foods, carrying physiologically active compounds, are believed to improve the biological defense mechanisms, prevent and reduce the risk of certain diseases, improve cognition and slow down the ageing process.

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Acknowledgment

The authors wish to thank Anula Perera for expert comments on the manuscript

and Gen Clark for assisting with the preparation of the manuscript.

Draft

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References

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Mazza, G., and Hodgins, M.W. 1985. Benzaldehyde, a Major Aroma Component of Saskatoon Berries. Hortscience 20(4): 742-744. Available from ://WOS:A1985ANR0700168 [accessed. Mazza, G., and Cottrell, T. 2008. Carotenoids and cyanogenic glucosides in Saskatoon berries (Amelanchier alnifolia Nutt.). J. Food Compost. Anal. 21(3): 249-254. doi:DOI 10.1016/jjfca.2007.11.003. McGarry, R., Ozga, J.A., and Reinecke, D.M. 2001. Differences in fruit development among large-and small-fruited cultivars of saskatoon (Amelanchier alnifolia). J. Am. Soc. Hortic. Sci. 126(4): 381- 385. Ozga, J.A., Saeed, A., and Reinecke, D.M. 2006. Anthocyanins and nutrient components of saskatoon fruits (Amelanchier alnifolia Nutt.). Can. J. Plant Sci. 86(1): 193-197. doi:10.4141/P05-139. Ozga, J.A., Saeed, A., Wismer, W., and Reinecke, D.M. 2007. Characterization of cyanidin- and quercetin- derived flavonolds and other phenolics in mature saskatoon fruits (Amelanchier alnifolia Nutt.). J. Agric. Food Chem. 55(25): 10414-10424. doi:Doi 10.1021/Jf072949b. Perez-Jimenez, J., Arranz, S., and Saura-Calixto, F. 2009. Proanthocyanidin content in foods is largely underestimated in the literature data: An approach to quantification of the missing proanthocyanidins. Food Res. Int. 42(10): 1381-1388. doi:DOI 10.1016/j.foodres.2009.07.002. Potì, F., Santi, D., Spaggiari, G., Zimetti, F., and Zanotti, I. 2019. Polyphenol health effects on cardiovascular and neurodegenerative disorders: A review and meta-analysis. Int. J. Mol. Sci. 20(2): 351. Prado, J.M., Dalmolin, I., Carareto, N.D., Basso, R.C., Meirelles, A.J., Oliveira, J.V., et al. 2012. Supercritical fluid extraction of grape seed: Process scale-up, extract chemical composition and economic evaluation. J. Food Eng. 109(2): 249-257. Robbins, R.J. 2003. Phenolic acids in foods:Draft an overview of analytical methodology. J. Agric. Food Chem. 51(10): 2866-2887. Rogiers, S.Y., and Knowles, N.R. 1997. Physical and chemical changes during growth, maturation, and ripening of saskatoon (Amelanchier alnifolia) fruit. Can. J. Bot. 75(8): 1215-1225. Available from ://WOS:A1997YA70900002 [accessed. Thompson, M.D., and Cooney, R.V. 2020. The Potential Physiological Role of γ-Tocopherol in Human Health: A Qualitative Review. Nutr. Cancer 72(5): 808-825. Zadernowski, R., Naczk, M., and Nesterowicz, J. 2005. Phenolic acid profiles in some small berries. J. Agric. Food Chem. 53(6): 2118-2124. doi:Doi 10.1021/Jf040411p. Zatylny, A.M., Ziehl, W.D., and St-Pierre, R.G. 2005. Physicochemical properties of fruit of 16 saskatoon (Amelanchier alnifolia Nutt.) cultivars. Can. J. Plant Sci. 85(4): 933-938. Available from ://WOS:000233890600026 [accessed. Zocchi, E., Hontecillas, R., Leber, A., Einerhand, A., Carbo, A., Bruzzone, S., et al. 2017. Abscisic acid: a novel nutraceutical for glycemic control. Front. Nutr. 4: 24. Zurawicz, E., Pluta, S., and Kucharska, D. 2014. Amelanchier - a New Berry Crop in Poland with Good Potential for Commercial Cultivation. X International Symposium on Vaccinium and Other Superfruits 1017: 251-255. Available from ://WOS:000333701000032 [accessed.

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Table 1. Characteristics of different varieties of saskatoonberry Varieties Weight a Total Soluble pH a Titratable Sugar c Seeds d (g/berry) solids a solids a acidity (g/100g (g/100g (%) (%) (% malic acid) FW) FW) Honeywood 1.07 21.7 15.8 4.14 0.31 a 3.75 3.0 Martin 1.66 20.8 15.3 3.71 0.52 a 2.17 2.3 Thiessen 1.62 20.4 14.8 3.76 0.46 a DNA 2.6 Smoky 1.09 21.5 16.0 4.18 0.31 a 2.56 4.4 Northline 1.06 23.0 16.5 3.78 0.53 a DNA 2.9 Pembina 0.85 27.9 20.1 3.95 0.45 b 4.05 3.8 JB30 1.35 19.9 14.0 3.78 0.50 a DNA DNA a Source: (Zatylny et al. 2005) b Source: (Lachowicz et al. 2017) c Sum of fructose, sorbitol, glucose, sucrose (see Table 4). Source: (Lachowicz et al. 2017) d Source: (Bakowska-Barczak et al. 2009) DNA: Data not available. Notes: Characteristics of fruit can vary from orchard to orchard depending on growing conditions.

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Table 2. Processed saskatoonberry products Product group Products Baked goods Pies, muffins, biscuits, breads, cakes Beverages Juice, concentrate, tea, , liqueur Preserved products Jam, jelly, sauce, pie filling, syrups, salad dressings Dried products Whole berry, powder, infused berries, fruit leather Sweets Chocolate coated, maraschino type, gelatin type As ingredients in other food products Meat stuffing and sauces for meat dishes, soups, ingredients in salads, etc.

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Table 3. Summary of macronutrient composition of saskatoonberries and blueberries Composition Saskatoonberrya Blueberrya (FW) (FW) Water (g/100g) 79.6 84.2 Energy (Calories/100 g) 84.9 57 Carbohydrates (g/100 g) 18.5 14.5 Total dietary fiber (g/100 g) 5.93 2.4 Protein (g/100 g) 1.33 0.74 Total fat (g/100 g) 0.49 0.33 Ash (g/100 g) 0.63 0.24 a Source: (Mazza 2005)

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Table 4. Sugar composition of saskatoonberriesa Sugars Honeywood Martin Pembina Smoky Average (g/100 g FW) Sucrose 0.11 0.13 0.14 0.12 0.13 Glucose (dextrose) 1.71 0.58 1.43 1.66 1.35 Fructose 1.58 1.15 1.18 1.15 1.26 Sorbitol 0.52 0.56 0.43 0.78 0.57 Total 3.93 2.42 3.17 3.71 3.31 a Source: (Lachowicz et al. 2017). Water content of the berries was 76.82%.

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Table 5. Amino acid composition of saskatoonberries Amino acid Saskatoonberrya (g/100 g FW) Glutamic acid 0.212 Arginine 0.099 Leucine 0.076 Glycine 0.061 Valine 0.053 Lysine 0.038 Alanine 0.053 Phenylalanine 0.046 Serine 0.053 Proline 0.053 Isoleucine 0.046 Threonine 0.038 Tyrosine 0.030 Histidine 0.023 Aspartic acid 0.189 Tryptophan 0.023 Methionine 0.023 Cystine 0.008 Crude protein Draft a Source: (Mazza 2005)

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Table 6. Vitamin composition Vitamin Saskatoonberrya (IU/100 g FW) 10.68 Vitamin A-RAE (RAE/100 g FW) 10.91

Vitamin B1 (thiamine) (mg/100 g FW) 0.04

Vitamin B2 (riboflavin) (mg/100 g FW) 3.54

Vitamin B3 (niacin) (mg/100 g FW) 0

Vitamin B5 (panthothenic acid) (mg/100 g FW) 0.31

Vitamin B6 (pyridoxine) (µg/100 g FW) 0.03

Vitamin B9 (, total) (µg/100 g FW) 4.55 (ascorbic acid) (mg/100 g FW) 3.55 Vitamin E (α-tocopherol) (mg/100 g FW) 1.12 Vitamin H () (mg/100 g FW) 0.02 a Source: (Mazza 2005) FW – fresh weight.

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Table 7. Mineral composition of saskatoonberriesa Mineral Content Range (mg/100g FW) (mg/100g FW) Potassium 203.1 162 – 244 Calcium 65.0 42 – 88 Magnesium 32.2 24 – 40 Phosphorous 26.1 20 – 32 Manganese 1.39 1.35 – 1.42 Iron 1.16 0.96 – 1.35 0.56 0.48 – 0.64 Zinc 0.25 0.17 – 0.33 a Source: (Mazza 1982) (Mazza 2005)

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Table 8. Concentrations of Individual flavonoid in saskatoonberries Flavonoids Content Range (mg/100g FW) (mg/100g FW) Anthocyanins Cyanidin-3-galactoside 155.4 57.5 - 331.4 Cyanidin-3-glucoside 48.1 10.1 - 115.9 Cyanidin-3-arabinoside 22.4 4.6 - 51.5 Cyanidin-3-xyloside 14.7 3.5 - 32.8 anthocyanins total 240.7

Flavonols Quercetin-3-galactoside 23.0 1.8 - 51.9 Quercetin-3-glucoside 6.4 2.3 - 11.6 Quercetin-3-arabinoglucoside 4.5 1.8 - 14.9 Quercetin-3-rutinoside 2.9 0.69 - 8.7 Quercetin-3-xyloside 1.4 0.66 - 2.4 Quercetin-3-arabinoside 1.2 0.16 - 3.3 Flavonols total 39.5 Source: Average values from (Lavola et al. 2012), (Dudonne et al. 2015), (Lachowicz et al. 2017), (Ozga et al. 2007). Draft

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Table 9. Extractable proanthocyanidin contents in saskatoonberries Proanthocyanidin Proanthocyanidin Saskatoonberry a polymer type (mg/100g FW) P1 (monomers) (+)-Catechin 7.2 (−)-Epicatechin 31.8 P2 (dimers) A-type 8.8 B-type 1.1 P3 (trimers) A-type 14.1 B-type 7.6 P4 (Tetramers) B-type 14.0

>P4 445.7 Total extractable 530.3 proanthocyanidins a Source: (Lachowicz et al. 2017). Values are averages of four cultivars: honeywood, martin, Pembina and smoky.

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Table 10. Concentrations of individual phenolic acids saskatoonberriesa Phenolic acids Content (mg/100 g FW) p-Hydroxybenzoic acid 0.01 Protocatechuic acid 0.59 Gallic acid 0.01 p-Coumaric acid 0.02 Caffeic acid 0.06 Ferulic acid 0.02 Coumaric acid glucoside 0.97 Caffeic acid glucoside 0.35 Shikimic acid 0.04 Ellagic acid 1.03 3-Caffeoylquinic acid (chlorogenic acid) 3.2 4-Caffeoylquinic acid 2.07 5-Caffeoylquinic acid (neochlorogenic acid) 16.3 a Source: (Dudonne et al. 2015)

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Table 11. Abscisic acids contents in saskatoonberriesa Abscisic acid Content (μg/100g FW) cis-Abscisic acid 9.57 ± 0.30 trans-Abscisic acid 1.04 ± 0.10 7′-OH-Abscisic acid 1.07 ± 0.02 Phaseic acid 10.9 ± 0.54 Dihydrophaseic acid 19.6 ± 0.95 Abscisic acid-glucose ester 2.61 ± 0.08 neo-PA 0.04 ± 0.01

Total 44.8 ± 1.37 a Source: (Dudonne et al. 2015)

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Table 12. Volatile aroma compounds in saskatoonberries a Compound Average Range (μg/kg FW) Benzaldehyde 208 18.8 - 985 Hexanal 45.2 13.8 - 87.5 Trans-2-hexenal 5.69* 0 - 9.5* 2-methylpropan-1-ol 842 0 - 2515 3-methylbutan-1-ol 3.7 0 - 5.6 Butan-1-ol 44.1 0 - 85.1 Cis-3-.hexenol 101 0 - 101 Ethanol 271* 128 - 469* Phenylmethanol 666 0 - 1221 Heptan-2-ol 2.54 0 - 3.5 Hexan-1-ol 66.0 37.7 - 104 Methanol 1.17** 0.8 - 1.4** Pentan-1-ol 12.9 0 - 17.5 Ethyl-decanoate 7.60 4.8 - 16.2 Ethyl-ethanoate 205 0 - 601 Methyl-ethanoate 210 0 - 533 a Source: (Butorova et al. 2016). * Content in milligrams per kilogram. Draft ** Content in grams per kilogram. Values are from cultivars: Lamarckii Balerina, Ostravsky, Tisnovsky skolsky, Thiessen, Tisnovsky velkoplody harvested in 2012-2013.

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Table 13. Fatty acid profile of saskatoonberries Fatty acid averagea Rangea (g/100 g seed oil) (g/100 g seed oil) Saturated fatty acids 16:0 Palmitic 6.62 6.52 - 6.72 18:0 Stearic 1.08 0.87 - 1.29 20:0 Arachidic 1.34 1.19 - 1.49 22:00 Behenic 0.57 0.50 - 0.64 24:00 Lignoceric 0.235 0.20 - 0.27 Total Saturated 9.845

Monounsaturated fatty acids 16:1 Palmitoleic 0.365 0.32 - 0.41 18:1 Oleic 29.72 27.8 - 31.6 20:1 Gadoleic 1.205 1.13 - 1.28 Total Monounsaturated 31.29

Polyunsaturated fatty acids 18:2 Linoleic (cis-cis) (omega-6) 57.185 55.1 - 59.3 18:3 α-Linolenic (cis-cis) (omega-3) 0.88 0.82 - 0.94 20:2 Eicosadienoic (cis-cis) 0.11 0.11 - 0.11 Total Polyunsaturated 58.175Draft

Total unsaturated 89.47 a Source (Bakowska-Barczak et al. 2007) and (Bakowska-Barczak et al. 2009).

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Table 14. Phytosterol and tocopherol profile of saskatoonberry seed oila

Content (mg/kg oil) Phytosterol Campesterol 633 Stigmasterol 219 β-sitosterol 7890 Δ5-avenasterol 750 Δ7-avenasterol 135 α-amyrin 176 citrostadienol 145 Total 9949

Tocopherols α-tocopherol (mg/kg of oil) 1173 δ-tocopherol (mg/kg of oil) 8.7 γ-tocopherol (mg/kg of oil) 163 Total 1344 a Source: (Bakowska-Barczak et al. 2009) Draft

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Table 15. Berry data and content of amygdalin, prunasin and potential HCN for fully ripe saskatoonberries* Content Honeywood Northline Martin Smoky Thiessen Seed content (% w/w) 4.63 6.35 3.90 5.62 4.69 Amygdalin (mg/kg FW) 42.0 44.0 44.4 118.2 118.3 Prunasin (mg/kg FW) 6.50 9.92 5.01 16.5 12.64 HCN potential (mg/kg FW) 3.08 3.51 3.08 8.5 8.15 * Source: (Mazza and Cottrell 2008).

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