Vaccinium myrtillus L., commonly known as bilberry, whortleberry, or European blueberry, is a low-growing, deciduousperennialshrub in the family Ericaceae, characterized by slender branches arising from extensive rhizomes, reaching heights of 10–50 cm, with bright green ovate leaves, urn-shaped pinkish-white to reddish flowers, and globular dark blue to black berries covered in a waxy bloom.[1][2][3]Native to circumboreal regions, V. myrtillus thrives in acidic, moist soils of coniferous forests, heaths, moors, bogs, and montane to alpine zones, often forming dense colonies that contribute to soil stabilization and understory vegetation in boreal ecosystems.[1][3] Its distribution spans Europe, Asia, and North America, from Greenland and Iceland across temperate Eurasia to western Canada and the northwestern United States (including Alaska, British Columbia, and states like Idaho, Montana, Oregon, and Washington), with disjunct populations in the Rocky Mountains extending south to New Mexico and Arizona.[1][4][3]Ecologically, V. myrtillus plays a vital role as a food source for wildlife, including birds such as grouse and thrushes, and mammals like black bears, deer, and small rodents, while its berries and foliage support pollinators and serve as browse for larger herbivores.[1] The plant's berries are rich in anthocyanins, flavonoids, and vitamins, contributing to its traditional and modern uses in food products like jams, juices, and wines, as well as in herbal medicine for purported benefits in eye health, circulation, and antioxidant support, though clinical evidence varies.[2]Taxonomically, V. myrtillus belongs to section Myrtillus within the genus Vaccinium, which comprises about 500 species of shrubs and small trees, and is classified as follows: Kingdom Plantae, Phylum Tracheophyta, Class Magnoliopsida, Order Ericales, Family Ericaceae, Genus Vaccinium.[5][3] It is distinguished from similar North American species like V. membranaceum by its more prostrate growth and bluish berries with a distinct flavor profile.[1]
Taxonomy and Nomenclature
Taxonomic Classification
Vaccinium myrtillus belongs to the kingdom Plantae, phylum Tracheophyta, class Magnoliopsida, order Ericales, family Ericaceae, genus Vaccinium, and species V. myrtillus.[5] This classification places it within the diverse Ericaceae family, which includes other ericoid shrubs adapted to acidic soils. The species authority is attributed to Carl Linnaeus, who formally described it in his 1753 work Species Plantarum.[3]Within the genus Vaccinium, V. myrtillus is classified in section Myrtillus, a group characterized by low-growing, deciduous shrubs with urceolate corollas and berries borne singly.[6] This section distinguishes V. myrtillus from other subgenera, such as subgenus Cyanococcus, which encompasses taller, more upright species like the highbush blueberry (V. corymbosum) used in commercial cultivation and featuring racemose inflorescences.[7] Historical synonyms include the homotypic name Vitis-idaea myrtillus (L.) Moench from 1794, reflecting early taxonomic reassignments before stabilization under Vaccinium.[3]Close relatives include V. uliginosum (bog bilberry), which shares section Myrtillus and similar deciduous habits, and V. vitis-idaea (lingonberry) from section Vitis-idaea, noted for evergreen leaves. Genetic studies highlight similarities among these species within the broader Vaccinium clade, particularly in population structure and diversity across shared northern habitats.[6][8]
Etymology and Common Names
The scientific name Vaccinium myrtillus derives from classical Latin roots. The genus name Vaccinium refers to a berry-bearing plant, possibly the bilberry itself or a related species like hyacinth, and is likely derived from bacca, meaning "berry," with historical usage in classical Latin for similar plants.[9] The specific epithetmyrtillus is a diminutive form of myrtus (myrtle), highlighting the plant's small leaves and berries that resemble those of the myrtle shrub; this binomial nomenclature was formalized by Carl Linnaeus in his Species Plantarum in 1753.[3]Common names for Vaccinium myrtillus vary regionally, reflecting linguistic and cultural differences across its native range. In English-speaking areas, it is primarily known as bilberry, a term originating from the Danish bollebar or bølle, denoting a "dark berry" or "whortleberry."[10] Regional British variants include blaeberry in Scotland and northern England, derived from Old Englishblā (blue or blue-black) combined with berry, emphasizing the fruit's dark hue; whortleberry, wimberry, and whinberry in southwestern England and Wales, with whortleberry tracing to Middle Englishhurtilbery via Danish bølle for the berry type.[11] In continental Europe, it is called myrtille in French, Heidelbeere in German (sometimes distinguished from the larger Vaccinium corymbosum but often used interchangeably for wild blueberries), and borówka czarna (black bilberry) in Polish.[12][13][14]
Description
Morphology
Vaccinium myrtillus is a low-growing, deciduousshrub typically reaching heights of 10–60 cm, characterized by slender, branching stems that arise from extensive underground rhizomes, forming dense mats or open colonies. The plant exhibits a spreading habit, with stems that are green to reddish-brown, sharply angled, and either glabrous or slightly puberulent, often tinged orange-red in sunlight. These stems support the overall structure, with maximum aerial shoot ages reaching 15–18 years.[1][15][16]The leaves are small, simple, and alternate, oval to elliptical in shape, measuring 1–3 cm in length and 0.5–2 cm in width, with finely serrated margins, an acute to obtuse apex, and a rounded to cuneate base. They are bright green during the growing season and turn shades of red, yellow, or brown in autumn. Flowers emerge singly or in pairs from the axils of new leaves on short pedicels; they are small, urn- or bell-shaped (urceolate to campanulate), pinkish-white to reddish, 4–6 mm long, and pendulous, blooming from April to July depending on location. Pollination is primarily achieved by insects, such as bumblebees, which are effective in transferring pollen for fruit set.[1][15][16][17]The fruit is a spherical berry, 5–10 mm in diameter, dark blue to black with a waxy, glaucous bloom coating the surface; the flesh is purple-pigmented and contains numerous small nutlet-like seeds (up to about 70 per berry). Berries ripen 50–70 days after flowering, typically from July to September. As a perennial chamaephyte, V. myrtillus exhibits rhizomatous growth, with new shoots emerging in spring (March–April) and active growth lasting about 5 months before winter dormancy. Reproduction occurs vegetatively through rhizome extension and budding, forming clonal networks that can persist for decades (rhizomes up to 34 years old), alongside sexual reproduction via insect-pollinated seeds that germinate under suitable conditions.[1][15][16][18][19][20]
Chemical Composition
The fruits of Vaccinium myrtillus are particularly rich in anthocyanins, which serve as the primary pigments responsible for their characteristic blue color and contribute to their antioxidant properties. These compounds typically constitute 300–700 mg per 100 g of fresh fruit, with the main types including glycosides of delphinidin, cyanidin, and malvidin, such as delphinidin-3-O-galactoside, delphinidin-3-O-arabinoside, cyanidin-3-O-galactoside, and malvidin-3-O-galactoside. On a dry weight basis, total anthocyanin content can range from 1,971 to 3,803 mg per 100 g, varying by population and environmental factors. Anthocyanins represent the largest fraction of the phenolic content in the fruits.Other phenolic compounds in V. myrtillus fruits include flavonoids such as quercetin (approximately 3 mg per 100 g fresh weight) and catechins (around 20 mg per 100 g fresh weight), as well as phenolic acids like chlorogenic acid. The leaves, in contrast, contain higher levels of tannins, ranging from 0.8% to 6.7% of dry weight, and arbutin (hydroquinone β-D-glucopyranoside), with contents varying from 1,000 to 2,400 mg per 100 g dry weight. These differences highlight the fruit's emphasis on anthocyanin-rich pigmentation and the leaves' prominence in astringenttannins and related phenolics.In addition to phenolics, V. myrtillus fruits provide notable vitamins and minerals. Vitamin C content is approximately 12–15 mg per 100 g fresh weight, while vitamin E is present in smaller amounts. The fruits are also a source of manganese, with levels up to approximately 30 mg per kg fresh weight—which is substantially higher than in cultivated blueberries (around 3.4 mg per kg)—and dietary fiber, at about 3.0–3.5 g per 100 g fresh weight.[21]Volatile compounds contribute to the aroma of V. myrtillus fruits, with key examples including the terpenes linalool, α-pinene, and 1,8-cineole, alongside other monoterpenes that form a complex profile detected via headspace analysis.
Distribution and Ecology
Geographic Distribution
Vaccinium myrtillus exhibits a Holarctic native distribution, spanning the northern regions of Europe, Asia, and North America. In Europe, it is widespread from the United Kingdom and Iceland westward through Scandinavia, where it extends north to approximately 70°N, to Russia and the Baltic states, as well as central and southern mountainous areas including the Alps, Pyrenees, and Carpathians.[3][1] In Asia, the species ranges from Siberia and the Russian Far East, including regions like Krasnoyarsk and Yakutiya, eastward to Japan, Mongolia, Kazakhstan, and parts of northern China such as Xinjiang.[3][22] In North America, it occurs natively from British Columbia and Alberta southward through the Rocky Mountains to central Oregon, northern New Mexico, and Arizona.[1][23] Populations in southwestern Greenland are also native, likely originating from European stock via historical dispersal.[1]The species' historical spread is tied to post-glacial migration following the Last Glacial Maximum, with rapid colonization of deglaciated landscapes through seed dispersal by birds and mammals, establishing dominance in boreal forest understories across its range.[24] This circumboreal pattern reflects adaptation to temperate and subarctic climates, with fossil evidence indicating expansion into northern latitudes as ice sheets retreated around 10,000–15,000 years ago.[25]Introduced ranges are limited, primarily involving cultivation for berry production in temperate parts of North America and Europe outside its native core, such as experimental plantings in the Pacific Northwest and central European lowlands; it has naturalized sporadically in some disturbed temperate habitats but does not form extensive invasive populations.[26] Currently, V. myrtillus covers vast areas within its native range, estimated at several million square kilometers in European boreal zones alone, with populations generally stable yet occurring in patchy distributions influenced by forest cover and soil conditions.[27]
Habitat and Growth Requirements
Vaccinium myrtillus thrives in a variety of acidic, nutrient-poor environments across temperate and boreal regions, including moors, heaths, coniferous forests, bogs, and alpine meadows. These habitats typically feature moderate shade levels, with optimal growth occurring under 30-70% canopy cover, where partial shading from overstory trees supports its understory dominance without excessive competition for light.[28][29][30]The plant requires well-drained, humus-rich soils that are sandy or loamy in texture, with a pH range of 4.0 to 5.5 to prevent chlorosis and promote root health. It favors cool, humid climates characteristic of boreal zones, with annual rainfall typically between 500 and 1000 mm to maintain soil moisture without waterlogging. Vaccinium myrtillus exhibits strong frost tolerance, surviving temperatures as low as -30°C, which enables its persistence in regions with harsh winters.[15][31][1]Ecologically, Vaccinium myrtillus plays a key role as a dominant understory species in boreal forests, contributing to ground cover and biodiversity. It forms symbiotic ericoid mycorrhizal associations with fungi such as Oidiodendron maius and Rhizoscyphus ericae, which enhance nutrient uptake, particularly nitrogen and phosphorus, in nutrient-poor soils. The plant interacts with herbivores like deer, rodents, and bears, which consume its foliage and berries, while pollinators including bees facilitate its reproduction through cross-pollination of its flowers.[28][32][33]Growth in Vaccinium myrtillus is characteristically slow, with annual height increments of 5-10 cm, reflecting its adaptation to stable, low-resource environments. In optimal light conditions within its preferred shade range, it achieves high berry yields, up to 1 kg per square meter in productive years, supporting its ecological and economic value.[1][15][34]
Cultivation and Conservation
Cultivation Methods
Vaccinium myrtillus is propagated primarily through seeds, semi-hardwood cuttings, or micropropagation techniques for enhanced conservation efforts. Seed propagation involves extracting seeds from ripe berries collected in late summer, followed by cold stratification at 4°C for approximately 90 days to break dormancy, after which they are surface-sown in a moist, acidic medium under light conditions for germination.[35] Semi-hardwood cuttings, taken in summer from current-season growth (8-12 cm long), are treated with 2000 ppm indole-3-butyric acid (IBA) and rooted in a 50:50 perlite-peat mix under mist with bottom heat at 21°C, achieving about 48% rooting success in 14 weeks.[35]Micropropagation via tissue culture uses nodal explants on Murashige and Skoog medium supplemented with cytokinins like N6-isopentenyladenine, enabling rapid multiplication and rooting rates up to 71.9% with potassium indolebutyrate, particularly useful for preserving genetic diversity.[36]Site preparation for cultivation requires acidic, well-drained loamy or sandy soils with a pH of 4.0-5.5, achieved by amending neutral or alkaline soils with peat moss, pine needles, or elemental sulfur several months in advance.[31]Plants thrive in partial shade mimicking woodland conditions, with spacing of 30-50 cm between shrubs to allow for their spreading habit up to 1 m wide. Raised beds or containers filled with ericaceous compost are recommended for poor drainage sites to prevent waterlogging.[31]Ongoing care includes mulching with pine bark or leaf mold in spring to retain moisture and suppress weeds, while maintaining consistent soil moisture using rainwater to avoid tap water's higher pH. Fertilization should be minimal and low in nitrogen, such as ammonium sulfate or ericaceous formulations applied sparingly in spring (e.g., 13-13-13 Osmocote at 1 g per container), to prevent excessive vegetative growth. Pruning in late winter involves removing dead wood and thinning crowded stems to promote airflow and fruiting on 2-3-year-old wood. Harvest occurs from July to August when berries turn deep blue and detach easily, ideally using hand-picking or combs for larger plantings.[31][35]Challenges in cultivation include high susceptibility to root rot from Phytophthora species in alkaline or waterlogged soils, necessitating strict pH and drainage management. Commercial production remains limited, with most berries wild-harvested, though potential expansion exists in Nordic countries through managed forest clearings and fertilization to boost yields. Mature plants (3-5 years old) typically yield 0.5-2 kg of fruit per shrub under optimal conditions, far below highbush blueberries but valuable for niche markets.[20][15]
Conservation Status
Vaccinium myrtillus is assessed as Least Concern on the IUCN Red List (as of 2016), indicating a low risk of extinction globally due to its widespread distribution across the Holarctic region and large population sizes. Similarly, NatureServe ranks it as Secure (G5), reflecting its abundance in native ranges, particularly in boreal forests of Europe and North America. Despite this overall stability, local populations face threats that could impact regional viability if not managed.[37][38]Regional threats include habitat loss from intensive forestry practices, such as clear-cutting, which negatively affects understory cover and berry production in boreal ecosystems. In northern Europe, particularly Finland and Sweden, commercial and recreational harvesting exerts pressure on populations, with annual wild bilberry yields estimated at 200-300 million kg across the region, though actual harvested amounts are lower at around 20-30 million kg combined, representing a sustainable fraction but potentially unsustainable locally with unregulated picking. Fires and overcollection have led to drastic reductions in abundance in specific areas, such as the Tropoja region in Albania, where habitat degradation compounds these issues. Climate change poses an additional risk, with warming temperatures causing upward shifts in alpine ranges and potential declines in subalpine populations due to altered phenology, increased mortality, and niche contraction in some habitats; however, recent studies indicate expansion in certain subalpine belts due to reduced land management and warming.[39][40][41][42][43]Conservation measures focus on protecting wild populations through integration into broader habitat directives, including the EU Habitats Directive, which safeguards associated boreal woodlands and heaths where V. myrtillus is a key species. Sustainable harvesting is promoted via guidelines and quotas in Nordic countries to prevent overexploitation, alongside research into in vitro propagation for germplasm banks to preserve genetic diversity. Reintroduction efforts in degraded sites, such as those in Tropoja, Albania, aim to restore local populations through propagation and habitat rehabilitation. Overall population trends remain stable, with high genetic diversity observed in northern European populations supporting resilience, though declines are noted in certain subalpine areas vulnerable to warming.[44][45][46][41][47]
Uses
Culinary Applications
The fruits of Vaccinium myrtillus, commonly known as bilberries, are widely used in culinary preparations due to their tart-sweet flavor and vibrant color. They are consumed fresh, often in Nordic countries where they are paired with cream or incorporated into porridges for a simple dessert. Processed forms include jams, pies, juices, and liqueurs; for instance, traditional Swedish liqueurs draw from wild bilberry harvests to create flavored spirits enjoyed as after-dinner drinks. Dried bilberries are steeped to make herbal teas, providing a mild, fruity infusion suitable for everyday consumption.[16][48][49]Bilberries offer a low-calorie profile, with approximately 57 kcal per 100 g, and are notably high in dietary fiber, contributing to their appeal in health-conscious diets. Their rich anthocyanin content imparts antioxidant properties, enhancing the nutritional value of dishes while adding a deep purple hue to baked goods like muffins and sauces. In Finnish cuisine, bilberries feature prominently in porridges and baked treats, where their natural pigments intensify flavors without added sugars.[50][51][52]Freezing bilberries effectively preserves their color and texture, allowing year-round use in recipes by preventing anthocyanin degradation during storage. Commercially, products like bilberry syrup are popular in Finland, often diluted for beverages or drizzled over desserts. Historically, indigenous groups such as the Sami foraged bilberries for winter storage, drying or cooking them into stews and breads to sustain communities through harsh seasons. Today, Europe's wild bilberry market yields around 5,000 tons annually, primarily through foraging in Nordic and Alpine regions, supporting both local and export demands.[48][53][54][55]
Medicinal Applications
Vaccinium myrtillus, commonly known as bilberry, has been employed in traditional Europeanfolkmedicine since medieval times, primarily using its fruit to address gastrointestinal and urinary ailments. The dried fruit was traditionally prepared as a tea to treat nonspecific diarrhea and dysentery, leveraging its astringent properties for symptomatic relief. Additionally, bilberryfruit has a long history of use for urinary tract issues, such as infections and inflammation, often consumed as an infusion or juice. In the context of vision health, a notable piece of folklore emerged during World War II, when British Royal Air Force pilots reportedly consumed bilberry jam before night flights, believing it enhanced their night vision capabilities, a tradition rooted in earlier European herbal practices for eye disorders like cataracts and retinopathy. Teas made from the fruit were also used to soothe inflammation in the mouth and throat.The leaves of Vaccinium myrtillus have been utilized in folk medicine, particularly for managing diabetes and related conditions. Decoctions of the leaves were traditionally administered to help control blood sugar levels, a practice dating back to the late 19th century and predating the discovery of insulin, with widespread use in European herbalism for ameliorating diabetic symptoms. These preparations were also applied topically as rinses or gargles for mouth and throat infections, as well as for wound healing, including ulcers. In Northern European traditions, such as those in Scandinavia, bilberry leaves served as a digestive aid, often in the form of infusions to alleviate stomach upset and vomiting.In cultural contexts, bilberry holds significance in Sami and Finnish folk medicine as a remedy for digestive disorders, reflecting its role in indigenous practices for gastrointestinal support. Today, these traditional uses have influenced modern herbal supplements, which often feature standardized extracts of the fruit rich in anthocyanins, the plant's key bioactive compounds.Common dosage forms in traditional applications include infusions of 1-2 g of dried leaves per day, steeped in 150 mL of boiling water for 5-10 minutes, and fruit extracts standardized to provide 100-200 mg of anthocyanins daily. Dried fruit was typically used at 20-60 g per day in teas or preserves.
Pharmacology and Safety
Pharmacological Effects
Vaccinium myrtillus, commonly known as bilberry, contains anthocyanins that contribute to its pharmacological effects, primarily through antioxidant and anti-inflammatory mechanisms observed in preclinical and clinical studies. These compounds scavenge free radicals, inhibit lipid peroxidation, and modulate oxidative stress pathways, as demonstrated in rat liver microsomes where bilberry extract reduced superoxide and hydroxyl radical activity.[56] Clinical trials have shown reductions in inflammatory markers such as high-sensitivity C-reactive protein (hsCRP) and interleukin-6 (IL-6) following supplementation with 330 mL bilberry juice daily for 4 weeks in adults at cardiovascular risk.[10] Similarly, 400 g of fresh bilberries daily for 8 weeks lowered hsCRP and IL-6 levels in individuals with metabolic syndrome, supporting improved endothelial function via nitric oxide pathways.[57]In ocular health, bilberry extracts improve microcirculation and show mild benefits for conditions like retinopathy and dry eyes, based on limited clinical evidence. A double-blind trial with 160 mg bilberry extract twice daily for 1 month reported 77-90% improvement in symptoms of diabetic and hypertensive retinopathy, attributed to enhanced retinal blood flow.[56] For dry eyes, supplementation with 160 mg bilberry extract daily for 30 days improved the Ocular Surface Disease Index score in patients compared to placebo.[16] However, meta-analyses and reviews indicate only small improvements overall, with no robust evidence for preventing or treating retinopathy progression beyond short-term symptom relief. Regarding vision enhancement, randomized placebo-controlled trials, such as one involving 160 mg extract (25% anthocyanosides) three times daily for 21 days, found no improvement in night visual acuity or contrast sensitivity in healthy young adults.[58]Metabolic effects include potential blood glucose lowering through alpha-glucosidase inhibition and lipid modulation in preclinical models, with mixed clinical outcomes. A 2025 meta-analysis of randomized controlled trials reported a marginal reduction in HbA1c (weighted mean difference: -1.63%, p=0.06) but no significant change in fasting blood glucose following bilberry supplementation.[59] The same meta-analysis, encompassing 8 RCTs with 409 participants, found no significant overall effects on lipids, including a non-significant decrease in total cholesterol (WMD: -0.11 mmol/L, p=0.27) and triglycerides (WMD: -0.07 mmol/L, p=0.62), no change in HDL (WMD: -0.02 mmol/L, p=0.70), but a small significant increase in LDL (WMD: 0.07 mmol/L, p=0.01); earlier individual studies reported decreases in LDL and increases in HDL in specific populations such as those with hypercholesterolemia.[59][57] Preclinical studies further support hypoglycemic action, with bilberryleaf extract reducing plasma glucose by 26% in diabetic rats.[56]Additional effects encompass anticarcinogenic potential and cardiovascular protection, primarily from in vitro and animal data. Bilberry powder at 10-25 mg/mL inhibited oral carcinoma cell proliferation, migration, and invasion in HSC-3 cells, with reduced tumor area in zebrafish models.[60] For cardiovascular health, extracts at 100 mg/kg reduced ischemia-reperfusion injury and leukocyte adhesion in hamsters, while human trials with 320 mg anthocyanins daily improved flow-mediated dilation.[56][57] The European Medicines Agency's 2015 assessment recognizes bilberry for traditional use in minor venous circulatory disturbances at doses of 80-180 mg extract (providing 60-160 mg anthocyanins) daily, though large-scale clinical trials are needed to confirm broader efficacy.[56] A 2025 systematic review and meta-analysis confirmed reductions in some inflammatory markers but emphasized the need for more high-quality studies on cardiometabolic indices.[59]
Toxicity Profile
Vaccinium myrtillus fruit is generally recognized as safe (GRAS) for consumption in amounts typically found in foods, with no significant toxicity reported at these levels.[61] In contrast, the leaves contain high concentrations of tannins, which can lead to adverse effects such as constipation or nausea when consumed in high doses exceeding 10 g per day or over extended periods.[62]Bilberry leaf extracts are considered possibly unsafe in such scenarios due to these tannin-related risks.[63]Acute toxicity studies indicate low risk, with no lethal effects observed in rodents at oral doses exceeding 2000 mg/kg body weight, suggesting an LD50 value well above this threshold.[27]Chronic toxicity assessments in animal models, including rats and dogs over periods up to 6 months, showed no significant adverse outcomes at doses up to 500 mg/kg, though minor changes like darkened urine or stool were noted.[27] There is no evidence of genotoxicity for the fruit, while leaves exhibit weak mutagenic potential in vitro; no carcinogenicity data are available, but no such risks have been identified in available studies.[27] Rare allergic reactions, including hives, swelling, or difficulty breathing, may occur, particularly in individuals sensitive to tannins or related berries.[64]Bilberry may interact with antidiabetic medications by enhancing their blood sugar-lowering effects, potentially leading to hypoglycemia, so blood glucose monitoring is advised in diabetic patients.[63] It has also been associated with increased bleeding risk when combined with anticoagulants like warfarin, based on case reports of elevated INR levels, though this interaction is not consistently confirmed at standard doses.[65]The European Medicines Agency (EMA) reports no major safety concerns for fruit preparations used traditionally, approving them for short-term oral or oromucosal use, while recommending leaves be limited to short-term application due to insufficient long-term data.[27] For vulnerable groups, fruit is likely safe during pregnancy at food levels, but supplements are not recommended due to limited data; leaves should be avoided in pregnancy and lactation.[63] Use in children under 2 years is contraindicated, with caution advised for those under 12, particularly for leaf preparations.[27] Diabetics should monitor blood sugar closely when using any bilberry products.[61]