Mortiño (Vaccinium floribundum Kunth)

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This entry is adapted from the peer-reviewed paper 10.3390/horticulturae8050358

Mortiño is a member of the Ericaceae family native to the Andes that has been used by local communities for centuries. This species has shown potential in the areas of medicine, agronomy, and green technology. A multidisciplinary approach was used to review aspects related to the ecology, horticulture, composition and potential biotechnological applications of mortiño.

ecology genetic diversity domestication attempts polyphenols

1. Origin, History, and Botany of Mortiño

Native to the tropical Andes of Colombia, Ecuador, and Perú ^[1]^[2], mortiño is commonly found at high altitudes on the edges of cold and humid paramos ^[3]. Mortiño commonly grows at temperatures between 7 and 18 °C ^[1], on rocky surfaces as a shrub reaching 2.5 m height, often prostrate or scandent ^[4]. It is one of the first species to grow back from root sprouts after fires in the paramos, playing a key ecological role in the regeneration of the ecosystem ^[5]. Mortiño leaves are small, coriaceous, elliptic, and ovate to ovate–lanceolate (Figure 1). The flowers are white to pink or red developed on racemes of 6–10 flowers, while the fruits are small (5–8 mm diameter) blue-black glabrous spherical berries at maturity ^[6] (Figure 2).

Figure 1. Description of Vaccinium floribundum Kunth: Semi-woody stem that can measure up to 2.5 m in height (a), Roots with their root hairs that, when they come to the surface, give rise to a seedling (b), White to lilac flowers in racemes of 6 to 10 (c), Elliptical, oval, or oval–lanceolate leathery leaves, with crenate-serrated edge, cuneate or round base, and slightly rounded acuminate apex (d), Corolla and Calyx (e), Vertical view of stamens (f), Gynoecium (g), Nectaries (h), Floral diagram of the Ericaceae (i), Round bittersweet berries of bluish to black color (j), Fruit in longitudinal section (k), Recalcitrant seeds of approximately 1 mm in size (l). Source: Herbarium of the Interpretation Center of the Protective Forest “La Prosperina” (BPP—Ar001E), of the Escuela Superior Politécnica del Litoral (ESPOL), Guayaquil—Ecuador, by Botanist Jaime Naranjo (Co-author).

Figure 2. Mortiño plant (Vaccinium floribundum Kunth) with its floral primordia (A), A small bunch of flowers with some pigmentation (B), Serrated leaves and its ripe berries (C). Source: This entry.

During its development, the fruit color transitions from green to white-pink, to pink, and finally to blue-black ^[7]. Fruit development after anthesis takes roughly between 60 and 100 days under natural conditions ^[8]. Taxonomically, V. floribundum has traditionally been classified within the Pyxothamnus section in the genus Vaccinium along with V. consanguineum and V. ovatum. However, phylogenetic analysis of tribe Vaccineae suggested Vaccinium is not monophyletic with V. floribundum, forming a clade along with V. consanguineum and V. meridionale, separate from other Vaccinium spp. ^[8].

Since it was first described in 1825 by Kunth as Vaccinium floribundum from the collections of Bonpland, the taxonomy of mortiño has varied little over the years. However, some common synonyms for the species have been reported, including: V. crenulatum, V. marginatum, V. ramosissimun, V. polystachium, V. mortinia, V. moritzianum, V. dasygynun, Metagonia crenulate, and M. marginata ^[3]. Common names for V. floribundum are many and often used ambiguously to identify other related species such as V. meridionale, Thibaudia floribunda, or Macleania rupestris. For instance, in Ecuador, mortiño is the most used vernacular name for the species; however, it is also known as uva de los Andes (grape from the Andes), manzanilla del cerro (chamomile from the hill), raspadura quemada (burnt panela), and uva de monte (mountain grape). In Peru, it is known as pushgay, uvitas, congama, and macha macha, whereas in Colombia it is also known as mortiño, agraz, agracejo, and chivaco; nonetheless, here, the name mortiño is more often associated with V. meridionale ^[1].

The consumption of this fruit was common in the Andean region before the arrival of the Europeans; later, it was assimilated into criollo traditions associated with the All Souls’ day, which continues to this day ^[1]. Currently, the species is mainly found in the wild, but it is also often present in smallholder farms ^[9].

2. Ecology and Genetic Diversity

Like many other neotropical Ericaceaes, V. floribundum predominates in belts of moist and cool montane forest preceding the transition towards the colder Paramo (cold and moist ecosystem typical of the high mountain in the Andes between the treeline and the snowline) between 3000 and 4500 m.a.s.l. ^[10]^[11]. On the slopes of the Rumiñahui volcano, near Quito (Ecuador), V. floribundum has commonly been observed in landscapes dominated by Calamagrostis intermedia and Carex jamesonii, although many other species, mostly dicots including other Ericaceae such as Pernettya prostrata, have been found growing in close spatial association to V. floribundum ^[12].

V. floribundum typically thrives in cold, nutrient-poor, moist though well-drained, shallow, and acidic soils ^[5]. The shrubs show remarkable adaptations to these conditions, such as shallow and almost horizontal root systems as well as profuse sprouting from roots and other vegetative tissues. These characteristics make V. floribundum one of the first species to regenerate damaged paramo ecosystems ^[5], drawing attention to ecosystem restoration programs in the Andes. Furthermore, V. floribundum has been reported as one of the species more often visited by a variety of bird and insect pollinators ^[13]^[14]. Adaptation to these environmental conditions is probably aided by association with specific soil microorganisms, as evidence of interactions between the V. floribundum rhizospheric microbiome and soil chemical properties has been reported ^[15]. Similarly, ericoid mycorrhiza forming fungi have been reported in V. floribundum roots, potentially contributing to the development of the species in nutrient-limited soils ^[12].

Currently, the conservation status of V. floribundum populations remains unknown. However, in Ecuador, mortiño is considered highly diverse—a typical characteristic of wild populations—and has shown both geographical and altitudinal patterns of diversification ^[16]. Still, concerns about the conservation of the species in scenarios of increasing demand of the berry and unsuccessful domestication efforts could pose risks, both to the species and to the paramos where it grows. For these reasons, interest in developing propagation techniques amenable for commercial cultivation has been recently growing in Colombia, Ecuador, and Peru ^[17].

3. Chemical Composition of Mortiño

The berries from Vaccinium spp. are known for their substantial amounts of sugars, polyphenols, vitamins B and C, minerals, and anthocyanins ^[18]. Table 1 shows the bromatological parameters and Table 2 describes works on mortiño phytochemistry.

Table 1. Physical and bromatological parameters of the Vaccinium floribundum berry.

Parameter/Units	Value	Reference
Soluble solids g/100 g	10.9	^[19]
Ash g/100 g	0.4	^[19]
Ash g/100 g	0.4	^[20]
Protein g/100 g	0.6	^[19]
Protein g/100 g	0.7	^[20]
Carbohydrates g/100 g	14.5	^[19]
	16.9	^[20]
	18.1	^[21]
Fat g/100 g	0.6	^[19]
Fat g/100 g	1	^[20]
Calories kcal/100 g FW *	84
Calories kcal/100 g FW *	75	^[21]
Water %	80	^[20]
Fiber %	7.6	^[20]
Fiber %	2.9	^[21]
pH	3.8	^[21]
Fe (mg/100 g FW)	0.64 ± 0.2	^[20]
K (mg/100 g FW)	607 ± 73	^[20]
Ca (mg/100 g FW)	17.0 ± 2.3	^[20]
Mg (mg/100 g FW)	10.2 ± 1.1	^[20]
Cu (mg/100 g FW)	0.12 ± 0.02	^[20]
Zn (mg/100 g FW)	0.13 ± 0.02	^[20]

* FW = Fresh Weight.

Table 2. Bioactive compounds from Vaccinium floribundum.

Compounds/Analysis Technique/Units *			Reference
Total polyphenols By the Folin-Ciocalteu method (mg GAE/100 g)	Fresh:	Powder:	^[22]
	524.4 ± 4.5	495.6 ± 9.1	^[22]
	204.01 ± 12.50		^[23]
	882 ± 38		^[19]
	2167 ± 835		^[24]
	7254.62 ± 1209.17		^[25]
	Chimborazo Province:	Pichincha Province:	^[23]
	107.4 ± 6.7	146.1 ± 13.4	^[23]
Total phenols By spectrophotometry:
Total phenols (mg GAE/100 mg)	608.05		^[20]
Total phenols (mgPgEq/g)	1.9 ± 0.7		^[26]
Total phenols (mg GAE/g)	9.3 ± 1.4		^[26]
Anthocyanins	Chimborazo Province:	Pichincha Province:
Total anthocyanin (pH differential) mg/100 g	89.9 ± 0.6	1095.4 ± 19.2	^[23]
Anthocyanin mg/g; (C3G equivalent)	10.6		^[27]
Total anthocyanin (%)	Fresh:	Powder:	^[24]
Total anthocyanin (%)	11.1 ± 0.5	2.3 ± 0.6	^[24]
Anthocyanin (mg cyanidin/100 g)	345		^[19]
Anthocyanin mg/100 g	376.2 ± 49.9		^[28]
Proanthocyanidin mg/g dry weight (epicatechin equivalent)	5.2		^[29]
Proanthocyanidin (%)	Fresh:	Powder:	^[22]
Proanthocyanidin (%)	5.3 ± 0.5	4.6 ± 0.3	^[22]
Flavonoids (Spectrophotometry):			^[26]
Total flavonoids (mg EC/g)	6.5 ± 0.7		^[26]
Flavonols total content mg/100 g	41.6 ± 10.2		^[28]
Flavonols glycosides	38		^[20]
Other compounds (By spectrophotometry)			^[26]
Total tannins (mg TAEq/g)	4.2 ± 0.8
Vit C mg/100 g	45.9 ± 6.7
Carotenes (μg/100 g):
-Β-carotene	70.6 ± 2.0
-Lutein	866.6 ± 7.5
Neochlorogenic acid	1.5 ± 0.5		^[28]
Chlorogenic acid	9.5 ± 2.9		^[28]
Quercetin and myricetin	-3-O-hexosides, -3-0pentosides and -3-0-deoxyhexosides		^[20]
Hydroxycinnamic acids	Predominance of acids: chlorogenic, neochlorogenic and derivatives of caffeic/ferulic acid		^[20]

* GAE = Gallic acid equivalent; C3G = cyanidin-3-glucoside; TAEq = tannic acid equivalent.

V. floribundum berries have shown high amounts of polyphenols, with levels reaching up to 7254.62 ± 10.86 mg GAE/100 g ^[25]. The most representative phenolic acids and flavonols reported in mortiño were quercetin-3-O-arabinofuranoside, chlorogenic acid, and quercetin-3-O-galactoside ^[28].

In comparative studies, other superfruits such as Prunus serotina showed lower levels of polyphenols than those found in mortiño, with reported concentrations of 1494 ± 385 and 2167 ± 835 mg GAE/100 g for each species, respectively ^[24]. Similarly, a higher content of phenolic acids and flavonols was observed in V. floribundum (41.6 ± 10.2 mg/100 g FW) when compared to V. myrtillus (13.7 ± 0.2 mg/100 g FW) ^[28]. In another study, the content of total soluble phenolic of mortiño (882 ± 38 mg GAE/100 g FW) was almost double that of guava (462 mg GAE/100 g FW) and plum (440 mg GAE/100 g FW); furthermore, it was four times higher than the values observed in strawberry (238 mg GAE/100 g FW) but less than half the content in the Andean blackberry (2167 mg GAE/100 g FW) ^[20].

Anthocyanins are other important components in V. floribundum berries, accounting for up to 67% of their total phenolic compounds ^[20]. Delphinidin-3-arabinose and cyanidin-3-arabinose have been reported as the most abundant anthocyanins in mortiño ^[22]. Values from 376.2 ± 49.9 ^[28] to 1095.39 mg/100 g FW ^[23] of anthocyanin content in blueberry have been reported, exceeding that observed in V. myrtillus (568.8 ± 8.8 mg/100 g FW) ^[28]. However, sample processing can reduce the levels of anthocyanins, as commercial mortiño powder showed lower levels of these bioactive compounds compared to fresh berries ^[22]. On the other hand, the levels of proanthocyanidins observed in V. floribundum (5.2 mg/g dry weight epicatechin equivalent) were higher than those in Aristotelia chilensis berries (4.0 mg/g DW EE) but lower than the values observed in V. myrtillus (13.7 mg/g DW EE) ^[22].

Lastly, it is also very important to know the content of P, Mn, Se, and I of mortiño berries. Unfortunately, to date there are no studies that provide this information. Further research is needed to assess the mineral content of mortiño berries.

4. Biological Activities of Mortiño

Due to the high amounts of polyphenols, anthocyanins, and antioxidants, mortiño has shown various bioactive properties. Table 3 shows some of the antimicrobial and medicinal properties of mortiño. Several studies have shown the antimicrobial capacity of Vaccinium spp. For example, V. corymbosum extracts inhibited the growth of various pathogens such as Salmonella spp. ^[30] and Listeria monocytogenes ^[31], while V. macrocarpon prevented the growth of Bacillus cereus and Micrococcus luteus ^[32]. Similarly, two studies have reported the antimicrobial capacity of mortiño. In one study, mortiño extracts obtained from lyophilized fruits or leaves in 70% ethanol inhibited the growth of 12 species of human pathogens, with inhibition halos ranging from 4.3 ± 0.3 to 39.7 ± 0.2 mm. The reported inhibition halos were greater than those observed when the antibiotic ampicillin was used ^[23]. Similarly, aqueous pulp and peel extracts of mortiño inhibited the growth of Streptococcus mutans ATCC35668 ^[27].

Table 3. Medicinal properties of Vaccinium floribundum.

Biological Activity	Main Findings	References
Modulatory capacity of adipogenesis	- Phenolic extracts inhibited lipid accumulation by 10.8% to 37.9% - Proanthocyanidin increased Pref-1 expression in preadipocytes	^[22]
Anti-inflammatory capacity	- Phenolic extracts decreased NO production by 25.5%; inducible NO synthase expression by 55.5%; prostaglandin E2 (ng/mL) by 20.1%; and chlooxygenase-2 conversion by 62.0%	^[22]
Diabetes therapy potential	- Proanthocyanidins inhibited α-glucosidase with IC₅₀ = 35 μg/mL and α-amilase with IC₅₀ = 25 μg/mL	^[29]
Chemopreventive activity	- No significant antimutagenicity and antigenotoxicity activity was observed	^[28]
Protection against oxidative stress	- Crude extracts attenuated the oxidative damage in HDFa caused by the stressor 2,2′-azobis(2-amidinopropane) dihydrochloride (AAPH) and improved the oxidative damage markers compared to the control.	^[26]
Antimicrobial capacity	- Fruit and leaf extracts significantly inhibited the growth of 8 Gram-negative and 3 Gram-positive bacterial strains	^[23]
Antimicrobial capacity	- The pulp and peel extracts of the mortiño inhibited the growth of S. mutans at the in vitro level	^[27]

Various medicinal properties have been attributed to V. floribundum, including potential applications in managing the symptoms of diabetes ^[29] and protection against oxidative stress ^[26]. Additionally, anti-inflammatory properties of mortiño have been suggested, as phenolic extracts of this species decreased the production of inflammatory mediators such as nitric oxide (NO), prostaglandin E2, and cycloxygenase-2 in lipopolysaccharide-stimulated RAW 264.7 macrophages ^[22]. Similarly, mortiño has been proposed as an inhibitor of adipogenesis, since proanthocyanidin-rich extracts of V. floribundum significantly prevented lipid accumulation in adipocytes and increased the expression of the preadipocyte factor 1 (Pref-1) by 4% in preadipocytes, a value higher than the 2.2% reached by A. chilensis but similar to the 5.9% achieved by epigallocatechin gallate used as a positive control (5.9%) ^[22]. Likewise, proanthocyanidins from mortiño successfully inhibited the enzymes α-glucosidase and α-amylase in vitro, with a 50% inhibition concentrations (IC₅₀) of 35 and 25 μg/mL, respectively, suggesting a potential use of this berry for diabetes therapy ^[29].

The berries of V. floribundum have also shown a high antioxidant capacity ranging from 0.339 ± 0.01 g/mL to 0.69 ± 0.03 g/mL ^[17]^[22], comparable to that of V. myrtillus (0.42 ± 0.01 g/mL) ^[28]. Through the Trolox Equivalent Antioxidant Capacity test, 250.01 ± 2.0 μmol TEq/g FW was reported in mortiño, which was higher than the 1.52 ± 3.1 μmol TEq/g FW observed in the berries of Rubus glaucus ^[26]. Due to its high antioxidant capacity, mortiño has the potential to protect human cells against oxidative stress. Crude extracts of V. floribundum attenuated the damage to Human Dermal Fibroblasts (HDFa) caused by oxidative stress, providing better protection than Rubus glaucus extracts ^[26]. However, no chemopreventive activity of mortiño was observed in mutagenicity and genotoxicity tests against 4-nitroquinoline-1-oxide (4-NQO) using SOS Chromotests ^[28]. Further research is needed to assess the potential antioxidant and radical scavenging applications of V. floribundum. In vivo studies, either in clinics or other controlled environments, are still needed to confirm the health benefits of mortiño.

5. Other Uses of Mortiño

There is a growing interest on plant antioxidants for green technology applications. For this reason, mortiño has been used in the synthesis of nanoparticles and solar cells.

The synthesis of nanoparticles commonly requires the use of toxic chemicals that serve as reducing agents. As a result of its high antioxidant capacity, mortiño has the potential to replace hazardous molecules for green production of nanoparticles. Mortiño extracts with high antioxidant capacity have been used for the green synthesis of graphene and functionalization of this material, with silver nanoparticles yielding a highly efficient photocatalyst ^[33]. Similarly, mortiño extracts were applied as a reducing and stabilizing agent for multicomponent nanoparticles (MCNPs) ^[34] and zero-valent iron nanoparticles (nZVIs) for bioremediation studies ^[19]^[35]. The resulting MCNPs showed >99% removal efficiency of toxic metals in water ^[34], whereas the zVINs removed at least 80% of total petroleum hydrocarbons (TPH) in contaminated water and soil ^[35]. The researchers indicated that the efficient formation and stabilization of the nZVIs was probably related to the -OH and -COOH groups from the berry polyphenols.

Additionally, mortiño extracts have also served as sensitizers for dye-sensitized solar cells, yielding efficiencies between 0.18 and 0.26% ^[36]. Table 4 shows some of the applications of mortiño.

Table 4. Other applications of Vaccinium floribundum.

Applications	Main Findings	Reference
Synthesis of zero-valent iron nanoparticles (nZVIs) for environmental remediation	- V. floribundum was used as a reducing and stabilizing agent yielding nZVIs of 13.2 ± 5.9 nm diameter	^[24]
	- V. floribundum was used as a reducing and stabilizing agent yielding nZVIs of 5 to 10 nm diameter - Removal of 88.3% of TPH from water (94.20 mg/L) - Removal of 81.9% of TPH from the soil (5000 mg/kg)	^[35]
Production of multicomponent nanoparticles (MCNPs) for removal/immobilization of heavy metals from water and in soils	- V. floribundum was used as a reducing and stabilizing agent yielding MCNPs of 12 nm diameter - MCNPs made from mortiño removed or immobilized heavy metals from water and soils with >95% efficiency	^[34]
Synthesis of silver–graphene nanocomposites with photocatalytic activity	- V. floribundum was used as a reducing agent. - The nanocomposites showed high photocatalytic degradation of methylene blue (92%; k = 0.0163283 min⁻¹) and methylene orange (84%; k = 0.0140985 min⁻¹)	^[33]
Biosynthesis of silver nanoparticles with photocatalytic activity	- V. floribundum was used as a reducing agent yielding nanoparticles of 20.5 ± 1.5 nm diameter - The nanoparticles showed high photocatalytic activity against 5 mg/L methylene blue with k = 0.00707788 min⁻¹	^[37]
Elaboration of dye-sensitized solar cells (DSSCs)	- V. floribundum was used as a photosensitizer in DSSCs showing a power conversion efficiency of 0.33%, open circuit voltages of 0.520 V, and short-circuit photocurrent densities of 1.014 mA cm⁻²	^[38]
Elaboration of dye-sensitized solar cells (DSSCs)	- V. floribundum was used as a photosensitizer in DSSCs showing an energy conversion efficiency of 0.18–0.26%	^[36]

Mortiño has also been used in the wine industry ^[39]^[40], baking ^[41], and even in the production of mortiño gummies ^[42]. The bioactive compounds of mortiño have attracted much interest in various sectors, which is why several methods have been proposed for the conservation and long-term storage of this berry, including short exposure to UV-C ^[43] and drying pretreatments ^[44] (Table 5). In a study, it was shown that after a storage period of 21 days, UV-C-treated (12.5 kJ m⁻²) mortiño retained 90% of the original anthocyanin levels compared to 76.85% of the untreated berries. However, the concentration of polyphenols was similar in both UV-C-treated and untreated berries ^[43]. Surprisingly, dry mortiño retained 93% of the anthocyanins and all the polyphenols in a storage period of 8 weeks ^[44]. However, other food processing technologies have been detrimental for the bioactive compounds of mortiño. Compared to lyophilization, heating in a sand bath to obtain a commercial mortiño powder yielded significantly lower levels of bioactive compounds such as anthocyanins (2.3 ± 0.6 vs. 11.1 ± 0.5%), proanthocyanidins (4.6 ± 0.3 vs. 5.3 ± 0.5%), total phenols (495.6 ± 9.1 vs. 524.4 ± 4.5 mg/g), and antioxidant capacity (3.3 ± 0.1 vs. 8.3 ± 0.4 mmol/g Trolox equivalents estimated by Oxygen Radical Absorbance Capacity (ORAC)) ^[22] (Table 5). In general, the processing of berries can degrade the anthocyanins naturally present in the fruits ^[45] (Table 3). Therefore, the pre-treatment of mortiño using UV-C represents a non-chemical approach to complement the treatment for low temperature storage, especially to maintain the anthocyanin concentration. Table 5 shows some of the preservation studies carried out on mortiño.

Table 5. Berry quality preservation studies.

Application	Main Findings	References
UV-C treatments for quality preservation of V. floribundum berries	- The exposure to UV-C prior to storage improved the antioxidant capacity (AC) retention of the berry - Retained 90% of the anthocyanins compared to 76.85% of the control	^[43]
Drying pretreatment for V. floribundum	- Best drying conditions: 40 °C, airflow of 0.24 m³/s, immersion bath concentration of 1.55% K₂CO₃ (w/w)—1.25% of ethyl oleate (v/v), concentration of SO₂ 2000 ppm and sulfidation per 2 h - The dried product retained 98.2% of anthocyanins and 99.7% of polyphenols - The mortiño dried by a solar dryer hybrid retained 88.5% of anthocyanins and 99.7% of polyphenols	^[44]
Winemaking	- Andean berries are a suitable raw material to produce wines with antioxidant capacity comparable to red grape wines	^[46]

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Subjects: Plant Sciences

Contributors MDPI registered users' name will be linked to their SciProfiles pages. To register with us, please refer to https://encyclopedia.pub/register :

Susana Llivisaca

Fabian León-Tamariz

Patricia Manzano-Santtana

JENNY RUALES

Lizette Serrano

Eduardo Chica

Juan Cevallos-Cevallos

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Entry Collection: Environmental Sciences

Update Date: 05 May 2022

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