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Apple

This article is about the fruit. For the technology company, see Apple Inc.. The apple is the edible fruit produced by the apple tree (Malus domestica), a species of the rose family (Rosaceae). It is one of the most widely cultivated tree fruits, with over 7,500 known cultivars varying in size, shape, color (typically shades of red, yellow, or green), flavor, and uses. The fruit is botanically a pome, featuring a fleshy outer layer surrounding a core with seeds. Apples are propagated primarily by grafting to maintain desirable traits, as seedlings often differ from the parent tree.^[1]^[2] Native to Central Asia, the cultivated apple likely originated from the wild ancestor Malus sieversii in the Tien Shan mountains, where it was domesticated around 4,000 years ago. From there, it spread via ancient trade routes, including the Silk Road, to Europe, the Middle East, and eventually the Americas with European colonists. Apple trees thrive in temperate climates and require cross-pollination for optimal fruit set. Globally, they are grown commercially in over 100 countries, with major producers including China, the United States, Turkey, Poland, and India. The 2024/25 global harvest is forecasted at 84 million metric tons, down slightly from the previous year due to weather impacts in key regions.^[1]^[3] Beyond nutrition and cuisine—where apples are eaten fresh, cooked, or processed into cider, juice, and pies—the fruit holds deep cultural and symbolic significance. In mythology and religion, it represents knowledge (as in the biblical Tree of Knowledge), love (Greek goddess Aphrodite), immortality (Norse Idunn's apples), and health (the proverb "an apple a day keeps the doctor away"). Apples feature prominently in folklore, art, and modern idioms across cultures.^[1]^[4]

Botanical Characteristics

Etymology

The English word "apple" derives from Old English æppel, referring to the fruit or any fruit in general, which traces back to Proto-Germanic aplaz and ultimately to the Proto-Indo-European root h₂ébōl, denoting an apple or similar fruit.^[5]^[6] This root is reflected in other Indo-European languages, such as Lithuanian obuolys and Old Irish ubull, highlighting the shared linguistic heritage across branches of the family.^[7] In classical languages, the term evolved through borrowing and adaptation; Latin mālum specifically meant "apple" or a tree fruit, adopted from Ancient Greek mēlon (Doric malon), which broadly signified any apple-like or foreign fruit and likely originated from a pre-Greek Mediterranean substrate language.^[8] These Greek and Latin words influenced broader Indo-European terminology for fruits, though mālum and mēlon also carried connotations of exotic or mast-like items in some contexts.^[8] Regional variations demonstrate parallel developments within Indo-European branches: French pomme stems from Latin pōmum (fruit or apple), reanalyzed from the plural pōma and derived from Proto-Italic poomos, possibly meaning "picked off" or "taken," emphasizing the act of harvesting.^[9]^[10] Similarly, German Apfel comes from Middle High German apfel and Old High German apful, directly cognate with the English form through the shared Proto-Germanic aplaz.^[11]^[6] Historically, "apple" and its cognates often denoted any fruit rather than the specific Malus domestica; this broader sense persisted into Middle English and appears in biblical translations, where the Hebrew pərî (generic fruit) was rendered in the Latin Vulgate as malum, fostering the later cultural link to apples despite no explicit identification in the original text.^[6]^[12]

Physical Description

The apple tree (Malus domestica) is a deciduous species typically reaching heights of 5 to 10 meters, with a central trunk that branches into a rounded or spreading canopy.^[13] Its leaves are alternate, simple, and oval to lanceolate in shape, measuring 5 to 12 centimeters in length, with serrated margins, dark green upper surfaces, and paler undersides often covered in fine hairs.^[14] Flowers emerge in spring as clusters of 4 to 6 blooms in flat-topped corymbs, featuring five white to pale pink petals, and they measure about 2.5 to 3.5 centimeters across.^[15] These hermaphroditic flowers give rise to the pome fruit following pollination and fertilization.^[14] The apple fruit is a pome, an accessory fruit derived from the hypanthium—a fused structure of the floral tube incorporating the receptacle, calyx, corolla, and stamen bases—that enlarges around the true fruit.^[16] The outer skin, or epicarp, is thin and waxy, providing protection and varying in color from green and yellow to red across cultivars, while the underlying flesh, or mesocarp, is crisp and juicy.^[17] At the center lies the core, formed by the endocarp enclosing five syncarpous carpels that contain 2 to 5 seeds each, with the persistent calyx often visible at the fruit's blossom end.^[18] Fruits typically measure 5 to 8 centimeters in diameter, though sizes can range from 3 to 10 centimeters depending on variety and growing conditions.^[15] Apple fruit development begins with flowering in spring, when buds open to reveal the clustered blossoms that last about 7 to 10 days before petal fall.^[19] Successful pollination triggers fruit set, where the fertilized ovary and surrounding hypanthium initiate rapid cell division, forming a small green fruitlet attached to the branch by a short peduncle.^[20] Over the growing season, which spans 120 to 200 days to maturity, the fruit undergoes expansion through cell enlargement, with the peduncle strengthening to support increasing weight.^[21] Toward maturity, an abscission layer develops at the peduncle-fruit junction, facilitating natural drop or mechanical harvest by weakening the connection.^[20]

Taxonomy and Classification

The apple, scientifically known as Malus × domestica Borkh., belongs to the genus Malus within the family Rosaceae, subfamily Amygdaloideae, and order Rosales in the kingdom Plantae.^[22] The binomial name incorporates the multiplication symbol (×) to denote its hybrid origin, reflecting ancient interspecific crosses primarily between wild Central Asian and European Malus species.^[23] This classification places the cultivated apple among approximately 30–55 species in the Malus genus, which are characterized by small deciduous trees or shrubs producing pome fruits.^[24] The evolutionary history of M. × domestica traces back to domestication from the wild progenitor Malus sieversii (Ledeb.) M. Roem., native to the Tian Shan mountains of Central Asia, particularly in modern-day Kazakhstan, Kyrgyzstan, and Tajikistan.^[25] This process began around 4,000 years ago, with genetic evidence indicating that cultivated apples arose through selective breeding and natural hybridization along ancient trade routes like the Silk Road.^[26] Subsequent introgression from the European wild apple Malus sylvestris (L.) Mill. contributed additional genetic diversity, particularly traits for adaptation to temperate climates in Europe and beyond. A 2025 genomic study using 218 whole genomes further confirms significant gene flow from M. sylvestris and selection pressures that shaped the modern cultivated gene pool.^[25]^[27] While M. × domestica is distinguished as the primary cultivated species, it differs from wild relatives such as M. sylvestris, the European crab apple, which features smaller, more bitter fruits and greater thorniness, serving as a secondary progenitor through hybridization.^[26] Crabapples, encompassing various small-fruited Malus species like M. sylvestris and M. baccata (L.) Borkh., represent the genus's wilder forms and are often interfertile with cultivated apples, leading to hybrid swarms in regions of overlap.^[28] No formal subspecies are recognized within M. × domestica, but its hybrid nature results in extensive variability among cultivars, maintained through vegetative propagation to preserve desirable traits.^[23] Phylogenetically, Malus is closely related to the pear genus Pyrus L. and other pome-bearing genera (e.g., Sorbus L.) within the tribe Maleae of Rosaceae, sharing an ancient whole-genome duplication event that facilitated the evolution of fleshy pomes from dry-fruited ancestors.^[29] This relationship is evidenced by molecular analyses showing Malus and Pyrus clustering together in the Maleae clade, with hybrid zones in Eurasia where intergeneric crosses occasionally occur, though rare due to ploidy differences.^[30] The pome structure, an accessory fruit derived from a hypanthium enclosing the true fruit, underscores these ties among Maleae genera.^[29]

Genetic Makeup

The genome of the domesticated apple (Malus × domestica) comprises 17 chromosomes arranged in a diploid set (2n=34), with an assembled size of approximately 750 Mb based on the reference sequence of the 'Golden Delicious' cultivar.^[31] More recent haplotype-phased assemblies, such as for 'Red Fuji' (2024, contig N50 of 25.6 Mb, assembled size ~669 Mb) and 'WA 38' (Cosmic Crisp, 2024), provide enhanced resolution of the highly heterozygous genome.^[32]^[33] This genome exhibits high heterozygosity, typically ranging from 0.88% to 1.67% across cultivars, reflecting the species' outcrossing nature and complicating efforts to resolve haplotype structures during sequencing.^[34] The high repeat content, exceeding 50% of the genome, further contributes to assembly challenges but also underscores the evolutionary history of whole-genome duplication events in the Rosaceae family.^[31] Apples display varying ploidy levels, with the majority of cultivars being diploid (2n=34), though triploid (2n=51) and occasional tetraploid (2n=68) forms occur naturally or through breeding.^[35] Triploid apples arise from crosses between diploid and tetraploid parents or via unreduced gametes, leading to distinct inheritance patterns where triploids often show sterility and reliance on diploid pollinators for fruit set.^[36] This ploidy variation influences trait expression, such as larger fruit size in triploids due to heterosis, and necessitates specific strategies in genetic mapping and selection.^[36] Several key genes underpin important phenotypic traits in apples. For fruit color, R2R3-MYB transcription factors like MdMYB10 activate the anthocyanin biosynthetic pathway, promoting red pigmentation in skin and flesh; allelic variations in this gene explain color differences across cultivars.^[37] Regarding flavor and quality, polyphenol oxidase genes such as MdPPO7 catalyze the oxidation of phenols into quinones, influencing enzymatic browning that impacts texture and taste post-harvest.^[38] Disease resistance is mediated by nucleotide-binding leucine-rich repeat (NLR) genes, including Rvi6 (formerly Vf), which triggers hypersensitive responses against the apple scab pathogen Venturia inaequalis through recognition of fungal effectors.^[39] Genetic diversity in domesticated apples is constrained by historical domestication bottlenecks, resulting in reduced nucleotide variation compared to wild progenitors, with effective population sizes estimated to have declined significantly along the Silk Road migration routes.^[25] This narrow base, exacerbated by clonal propagation, heightens vulnerability to pests and environmental stresses, but has been mitigated through deliberate introgression of alleles from wild relatives like Malus sieversii and Malus sylvestris, introducing novel resistance and adaptive traits.^[40] Such hybridization efforts restore heterozygosity and expand the gene pool, supporting sustainable breeding without evidence of severe founder effects in core genomic regions.^[26]

Cultivation Practices

Historical Development

The domesticated apple (Malus domestica) originated from the wild ancestor Malus sieversii in the Tian Shan mountains of Central Asia, particularly in modern-day Kazakhstan, where genetic evidence indicates domestication began approximately 4,000 to 10,000 years ago.^[25] This wild species contributed the majority of the genetic makeup to modern cultivars, with genomic studies showing that about 46% of the domestic apple genome derives from Kazakh populations of M. sieversii.^[41] From there, apple cultivation spread westward along ancient trade routes, including proto-Silk Road paths, reaching regions like Mesopotamia by around 1500 BCE, as evidenced by cuneiform records from Nuzi documenting the sale of an apple orchard.^[42] Early written references to apples appear in Assyrian and Near Eastern texts from the late second millennium BCE, highlighting their role in royal gardens and agriculture.^[43] By the classical period, Greek scholars like Theophrastus (c. 371–287 BCE) described apple cultivation in detail in his Historia Plantarum, noting over 40 varieties, their flowering cycles, and propagation methods, while Romans expanded orchards across their empire, integrating apples into viticulture and pomology.^[44] These advancements laid the foundation for systematic fruit growing in the Mediterranean. In medieval Europe, monastic communities preserved and expanded apple cultivation, maintaining orchards within cloistered gardens for sustenance, cider production, and medicinal use, with records from the 10th to 15th centuries documenting hundreds of varieties tended by orders like the Cistercians.^[45] During the Renaissance, grafting techniques advanced significantly, particularly in 15th-century France, enabling clonal propagation and the preservation of specific traits, which increased orchard efficiency and variety diversity.^[46] Colonial expansion in the 17th century introduced apples to the Americas, where European settlers planted seeds and grafts starting with French Jesuits in the late 1500s and English colonists in Jamestown by 1607, leading to hybridized cultivars adapted to New World climates.^[47] In 1788, Captain William Bligh planted the first apple trees in Australia at Adventure Bay, Tasmania, aboard HMS Bounty, initiating cultivation on the continent and facilitating global dissemination through imperial trade networks.^[48] The 20th and 21st centuries marked the industrialization of apple production, with mechanized harvesting, chemical pest management, and high-density planting systems emerging post-World War II to boost yields. In the United States, innovations in storage and transportation post-World War II supported growth in the fresh fruit market, with production expanding from around 5 billion pounds in the mid-20th century to 11.7 billion pounds as of the 2025 forecast.

Breeding Techniques

Apple breeding traditionally relies on controlled cross-pollination between parent varieties selected for complementary traits such as flavor, yield, and disease resistance, followed by the evaluation and selection of seedlings over multiple generations. This labor-intensive process begins with hand-pollination of flowers to create hybrid seeds, which are then planted and observed for fruit quality, tree vigor, and other characteristics through successive years of growth and harvesting. On average, developing a new cultivar from the initial cross to commercial release takes 20-30 years, due to the time required for trees to mature, produce sufficient fruit for assessment, and demonstrate consistent performance across environments.^[49]^[50]^[51] To accelerate selection for specific traits, marker-assisted selection (MAS) employs DNA markers linked to genes of interest, reducing the need for extensive field testing. For instance, MAS targets the Vf gene, derived from Malus floribunda, which confers resistance to apple scab (Venturia inaequalis); markers such as RAPD-derived sequences tightly linked to Vf enable early identification of resistant seedlings in breeding populations. This approach has been integrated into programs to pyramid multiple resistance genes, enhancing durability against pathogens while preserving desirable horticultural qualities.^[52]^[53] Grafting remains essential for propagating elite apple varieties, with the scion (desired cultivar) budded onto rootstocks that influence tree size, productivity, and adaptability. Dwarfing rootstocks like M.9, originating from the East Malling series, limit mature tree height to 25-30% of standard rootstocks, facilitating high-density plantings of up to 2,000 trees per hectare and improving light interception for higher yields per area. These rootstocks also promote early bearing and easier management, though they often require support systems to prevent toppling.^[54]^[55] Post-2010 developments have introduced CRISPR/Cas9 genome editing as a precise tool for apple breeding, enabling targeted modifications to enhance traits like disease resistance and fruit quality. Demonstrated in 2016, CRISPR/Cas9 successfully edited apple genes such as those controlling chlorophyll production, proving its efficiency in stable transformation without off-target effects in key cultivars. More recent applications target genes influencing texture and shelf-life, including MdPG1 (polygalacturonase 1), which regulates cell wall degradation and fruit softening; editing this gene holds potential to maintain firmness during storage, addressing postharvest losses. Ongoing research in pome fruits emphasizes CRISPR for accelerating breeding cycles beyond traditional methods.^[56]^[57]^[58]

Pollination Methods

Apple flowers are perfect, containing both male and female reproductive organs, and typically feature five white petals, five sepals, approximately 20 stamens arranged in a ring around the central pistil, and an inferior ovary composed of five fused carpels, each with two ovules.^[59] The pistil consists of five styles that may unite at the base, facilitating pollen tube growth toward the ovules.^[60] Apple pollination is primarily entomophilous, relying on insects to transfer pollen, with honey bees (Apis mellifera) serving as the dominant pollinators due to their efficiency in visiting multiple flowers within an orchard.^[60] Most commercial apple cultivars (Malus domestica) exhibit gametophytic self-incompatibility, a genetic mechanism that prevents self-pollination and requires cross-pollination from a compatible variety to achieve adequate fruit set.^[61] This incompatibility arises from interactions at the S-locus, where pollen sharing an S-allele with the style halts tube growth, ensuring genetic diversity but necessitating careful selection of pollinizer trees, often planted at a ratio of 1:8 with the main cultivar.^[61] Apple bloom typically occurs in mid-spring, with timing varying by cultivar (early, mid-season, or late) and region, generally spanning late April in temperate zones like the Mid-Atlantic U.S., during which flowers open sequentially in clusters, starting with the central "king" bloom.^[62] Successful cross-pollination demands overlapping bloom periods between compatible varieties to maximize pollen transfer.^[60] Orchard management for pollination includes introducing managed honey bee hives at rates of 1-2 per acre, distributed evenly to ensure uniform coverage, supplemented by wild pollinators like mason bees (Osmia spp.) for enhanced efficiency.^[63] Key factors influencing pollination success encompass weather conditions, such as cool temperatures below 15°C (59°F) or rain that reduce bee foraging activity and pollen viability; pesticide applications, particularly insecticides during bloom, which can cause pollinator mortality or repellency; and genetic factors like S-allele mismatches leading to incompatible pollen.^[64] Mitigating these involves timing sprays to avoid bloom, providing bee-friendly habitats, and selecting compatible pollinizers based on S-genotyping.^[60]

Growth and Harvesting

Apple trees thrive in temperate climates, requiring 500 to 1,000 chill hours—defined as hours between 0°C and 7.2°C during winter—to break dormancy and ensure proper flowering and fruit set.^[65] These conditions are typical in USDA hardiness zones 4 through 8, where cold winters prevent excessive vegetative growth and promote balanced fruit production. Well-drained soils with a pH of 6.0 to 7.0 are essential to avoid root rot, with loamy or sandy loam textures providing optimal aeration and nutrient availability.^[66] Tree spacing varies by rootstock vigor: dwarf varieties are planted 3 to 4 meters apart, semi-dwarf at 4.5 to 6 meters, allowing for adequate light penetration, air circulation, and mechanical access in commercial orchards.^[67] The maturation timeline of apple trees follows an annual cycle beginning with winter dormancy, during which growth halts and the tree conserves energy after leaf drop in late fall.^[68] Planting typically occurs in early spring using grafted nursery stock, with roots establishing quickly in the first year while the canopy develops. Dwarf trees on rootstocks like M.9 may bear their first fruit in 2 to 3 years, semi-dwarf in 3 to 5 years, and standard trees in 5 to 8 years, depending on variety, site conditions, and pruning practices.^[69] Once mature, trees enter a productive phase lasting 20 to 50 years, with vegetative growth resuming in spring, followed by flowering, fruit set, and enlargement through summer, culminating in harvest before returning to dormancy. Harvest timing occurs in late summer to fall, typically from August to November in the Northern Hemisphere, guided by physiological indicators of maturity to ensure optimal flavor and storability. Soluble solids content, measured as Brix levels of 12% to 15%, signals sufficient sugar accumulation for sweetness.^[21] Background skin color shifts from green to yellow or light tan, while increased ethylene production triggers climacteric ripening, softening the fruit and enhancing aroma compounds. Yields from mature trees range from 20 to 50 kg per tree annually, varying by cultivar and management, with dwarf trees producing around 22 to 45 kg and standards up to 90 kg in favorable conditions.^[67] Hand harvesting remains predominant for fresh-market apples, involving selective picking into padded bins to minimize bruising, especially for premium varieties where laborers climb ladders or use platforms. Mechanical harvesting, suitable for processing or cider apples, employs trunk shakers or over-row machines to dislodge fruit onto conveyors, reducing labor by up to 70% but risking higher damage rates of 10-20%. Post-harvest handling includes rapid cooling to 0-4°C and gentle transport to packing facilities, with brief reference to controlled atmosphere storage for extended shelf life as detailed elsewhere.^[21]

Storage and Preservation

After harvest, apples are highly perishable due to ongoing respiration and ethylene production, which accelerate ripening and senescence. Controlled atmosphere (CA) storage is a primary commercial technique to extend shelf life by altering the gaseous environment in sealed storage rooms. This method reduces oxygen levels to 1-3% and elevates carbon dioxide to 3-5%, while maintaining temperatures between 0-4°C and relative humidity at 90-95%.^[70]^[71] These conditions slow metabolic processes, inhibit ethylene synthesis, and minimize fungal growth, allowing many varieties to remain marketable for 6-12 months.^[70] CA storage was first developed in the 1940s and has since become standard in major apple-producing regions, significantly reducing spoilage compared to traditional cold storage.^[72] Several factors influence the effectiveness of apple storage and overall shelf life. Ethylene, a plant hormone produced by apples, promotes ripening and must be managed, as even low levels (below 1 ppm) can trigger rapid deterioration; CA storage helps by limiting its accumulation through low oxygen.^[70] High humidity (90-95%) prevents water loss and shriveling, which can reduce weight by up to 5-10% in dry conditions, while temperatures above 4°C accelerate enzymatic breakdown.^[73] Varietal differences play a key role; for instance, Granny Smith apples, with their firm texture and low ethylene production, can store for 8-10 months under optimal CA conditions, outperforming softer varieties like McIntosh, which may last only 2-4 months.^[74] These biochemical changes during storage, such as softening and acid loss, are influenced by initial fruit maturity at harvest.^[75] For home preservation, refrigeration at 0-4°C in the crisper drawer with high humidity extends usability to 1-2 months for most varieties, by slowing respiration similar to commercial methods.^[76] Waxing, typically a commercial practice using food-grade coatings to seal the skin and reduce transpiration, can be mimicked at home with natural alternatives like beeswax dips, though it adds minimal extension beyond proper cooling.^[77] Processing into cider involves grinding and pressing fresh or stored apples, followed by pasteurization or fermentation for longer-term storage up to several months in sealed containers.^[78] Globally, post-harvest losses for apples range from 20-30%, primarily from improper storage leading to decay, physical damage, and physiological disorders during transport and holding.^[79]^[80] The adoption of CA technology since the 1940s has mitigated these losses by 50% or more in equipped facilities, enabling year-round availability and reducing waste in supply chains.^[72]

Pests, Diseases, and Management

Apple production faces significant threats from various pests and diseases, which can result in substantial economic losses if not managed effectively. Without proper intervention, these biotic factors collectively cause primary yield losses of approximately 25% in apple crops, with secondary losses potentially reaching 38% due to reduced fruit quality and marketability. Integrated Pest Management (IPM) approaches, combining monitoring, cultural practices, biological controls, and targeted chemical applications, are widely recommended to minimize these impacts while promoting sustainable production. Among the key insect pests, the codling moth (Cydia pomonella) is a primary concern, as its larvae bore into developing fruit, leading to internal damage and premature drop. Effective management includes monitoring with pheromone traps to time interventions, mating disruption using synthetic pheromones to confuse male moths, and biological controls such as releases of parasitoid wasps like Trichogramma species. Similarly, the apple maggot (Rhagoletis pomonella) infests fruit through oviposition by adult flies, resulting in maggot tunneling that renders apples unmarketable. Control strategies emphasize sanitation by removing and destroying fallen or infested fruit to break the life cycle, supplemented by yellow sticky traps for monitoring and insecticides applied during peak adult emergence. Major diseases include apple scab, caused by the fungus Venturia inaequalis, which produces olive-green to black lesions on leaves, fruit, and twigs, severely reducing photosynthesis and fruit quality. Fungicide programs, typically involving protectant applications like sulfur or captan starting at green tip stage and continued through petal fall, are essential, with sanitation practices such as raking and composting infected debris to reduce inoculum. Fire blight, a devastating bacterial disease incited by Erwinia amylovora, affects blossoms, shoots, and roots, causing wilting and cankers that can girdle branches. Management relies on copper-based bactericides applied during dormancy and bloom, antibiotics like streptomycin during flowering, and the use of resistant rootstocks from the Geneva series (e.g., G.935, G.41) to limit spread and improve tree vigor. Emerging challenges, exacerbated by climate change, include the northward expansion of the invasive brown marmorated stink bug (Halyomorpha halys), which pierces fruit skin to feed on developing apples, causing corky dimples and deformities that lower cosmetic value. Warmer temperatures have enabled more generations per season and broader range shifts into previously unaffected northern regions. For organic systems, alternatives like low-metallic copper sprays (e.g., during dormancy for fire blight suppression) and kaolin clay barriers offer viable options, though they require integration with IPM to achieve comparable efficacy to conventional methods.

Major Cultivars

Apple cultivars, or varieties, exhibit diverse traits in color, flavor, texture, and adaptability, influencing their popularity in different regions. Among red-skinned varieties, Red Delicious stands out as a longtime favorite in the United States, prized for its striking deep crimson hue and mild, sweet flavor, though modern iterations often feature a mealy texture due to selective breeding for appearance over taste.^[81] Originating as a chance seedling in the 1870s on Jesse Hiatt's farm in Peru, Iowa, it became commercially available in the early 20th century and remains one of the most widely grown apples globally for its reliable productivity.^[82] Another prominent red cultivar, Fuji, developed in Japan during the 1930s as a cross between Red Delicious and Ralls Janet, is renowned for its exceptional sweetness, crisp texture, and dense, juicy flesh that resists browning.^[83] This variety's long storage life, often extending several months under proper conditions, has fueled its rise as a global export favorite originating from Aomori Prefecture.^[84] Green and yellow cultivars offer contrasting tartness and versatility, with Granny Smith emerging as a benchmark for acidity. Discovered in 1868 near Sydney, Australia, as a chance seedling by Maria Ann Smith from discarded crab apples, Granny Smith features bright green skin, firm white flesh, and a sharp, tangy flavor that maintains integrity in cooking.^[85] Its late-season harvest and resistance to bruising have made it a staple in Australian and international markets since the late 19th century.^[86] Complementing this is Golden Delicious, a yellow-green apple found as a chance seedling in the early 1900s on Anderson Mullins' farm in Clay County, West Virginia, valued for its mellow sweetness, thin skin, and adaptability to both fresh eating and processing.^[87] Released commercially in 1914, its high yields and ease of growth have established it as a foundational parent in many hybrid varieties worldwide.^[88] Heirloom varieties like Cox's Orange Pippin represent traditional excellence in flavor complexity, originating in 1825 from a seedling cultivated by Richard Cox in Colnbrook, Buckinghamshire, England. This aromatic apple boasts orange-red streaked skin, fine-grained yellow flesh, and a rich bouquet of citrus, berry, and tropical notes, earning acclaim as one of the finest dessert apples despite its susceptibility to diseases and modest yields.^[89] In contrast, modern breeding efforts have produced cultivars like Honeycrisp, developed at the University of Minnesota in the 1990s from a 1960 cross of Keepsake and Northern Spy, celebrated for its explosive crispness, high juice content, and balanced sweet-tart profile. Selected for superior texture in controlled evaluations, Honeycrisp's patented status and rapid adoption highlight advancements in prioritizing sensory qualities over historical constraints.^[90] Regional leaders further diversify the landscape, with Gala from New Zealand exemplifying mild sweetness and extended shelf life. Bred in the 1930s as a cross between Kidd's Orange Red and Golden Delicious and commercialized in the 1970s by grower J.H. Kidd, Gala features small to medium size, mottled red-orange skin, and firm flesh that stores well for up to two months under refrigeration, supporting its dominance in Southern Hemisphere exports.^[91] Similarly, Pink Lady (Cripps Pink), developed in Australia in the 1970s at the CSIRO by crossing Golden Delicious and Lady Williams, offers a crisp, balanced flavor with pink-blushed skin and notable disease resistance, including moderate tolerance to scab and mildew, enabling robust production in warmer climates.^[92] These traits, combined with its late ripening, have propelled Pink Lady to international acclaim since its 1990s trademarking.^[93]

Global Production Trends

In 2024/2025, global apple production reached approximately 84 million metric tons, with China leading as the top producer at 48 million metric tons, followed by the United States at 4.9 million metric tons, Turkey at 4.8 million metric tons, and Poland at 4.3 million metric tons.^[94]^[95] These four countries accounted for over 60% of the world's output, highlighting the concentration of production in a few key regions.^[96] A notable trend in recent years has been the increasing dominance of Asia in global apple production, which held about 60% of the total share in 2024/2025, driven primarily by expansions in China and India.^[97] Climate variability has influenced yields, such as the severe droughts across Europe in 2022, which contributed to a 5-10% decline in regional production due to water stress and heat damage.^[98] In 2025, forecasts indicate continued growth in U.S. production to 11.7 billion pounds, supported by favorable weather in key states like Washington. International trade in fresh apples generated export values exceeding $7 billion annually, supporting economic ties between major producers and importing markets like the European Union and Russia.^[99] Sustainability efforts in apple cultivation focus on resource efficiency, with the water footprint averaging around 822 liters per kilogram of apples produced, primarily from irrigation and rainfall in growing regions.^[100] Advances in mechanization, including automated harvesting and precision pruning, have boosted average yields to 20-40 tons per hectare in modern orchards, enhancing productivity while reducing labor costs.^[101] Recent developments as of 2025 emphasize climate-resilient breeding and reduced pesticide use to address shifting weather patterns and regulatory pressures. Looking ahead to 2030, production is projected to grow modestly to about 90 million metric tons globally, with significant expansions anticipated in emerging markets like India and Brazil due to improved cultivars and expanded acreage.^[102] However, climate change poses challenges, including earlier blooming periods—advanced by 1-2 weeks in many temperate zones—which increase vulnerability to late spring frosts and disrupt pollination timing.^[103]

Chemical and Nutritional Profile

Biochemical Composition

Apples are composed primarily of water, which constitutes about 86% of their fresh weight, providing hydration and structural integrity to the fruit. The remaining composition includes carbohydrates, predominantly sugars at approximately 10-13% of the total weight, with fructose as the dominant sugar (around 6 g per 100 g), followed by glucose and sucrose; these sugars contribute to the fruit's sweetness and energy content. Dietary fiber accounts for roughly 2% (about 2.4 g per 100 g), primarily in the form of pectin concentrated in the cell walls, which aids in maintaining tissue firmness and texture.^[104]^[105] Polyphenols represent a key class of secondary metabolites in apples, serving as antioxidants that protect against oxidative stress. Notable compounds include quercetin, primarily located in the peel at levels of 4-50 mg per 100 g, and chlorogenic acid, more prevalent in the flesh at 10-100 mg per 100 g, with total polyphenol content varying from 100 to 500 mg per 100 g depending on cultivar and growing conditions.^[106]^[107] The characteristic aroma of apples arises from volatile compounds, particularly esters produced during ripening, such as hexyl acetate, which imparts fruity notes and is especially prominent in mature fruit. These volatiles are synthesized in the fruit's tissues and contribute to sensory appeal. Additionally, enzymes like polyphenol oxidase play a critical role in post-harvest biochemistry, catalyzing the oxidation of polyphenols to quinones upon tissue disruption, resulting in the enzymatic browning of cut surfaces.^[108]^[109]

Nutritional Value

Apples are a nutrient-dense fruit, providing a modest calorie content primarily from carbohydrates while being virtually fat-free. Per 100 grams of raw apple with skin, the macronutrient profile includes approximately 52 calories, 13.8 grams of carbohydrates (of which about 10.4 grams are sugars), 0.26 grams of protein, and 0.17 grams of total fat, with no cholesterol or sodium.^[104] The dietary fiber content stands at 2.4 grams per 100 grams, which supports digestive health by promoting regular bowel movements and potentially reducing the risk of constipation.^[104] In terms of micronutrients, apples offer notable amounts of vitamin C and potassium. A 100-gram serving contains 4.6 milligrams of vitamin C, fulfilling about 5% of the daily value (DV), which contributes to immune function and antioxidant protection. Potassium is present at 107 milligrams per 100 grams (2% DV), aiding in blood pressure regulation and muscle function. Other micronutrients, such as vitamin K (2.2 micrograms, 2% DV) and smaller traces of vitamin A and B vitamins, add to the fruit's overall nutritional profile.^[104] The consumption of apples has been linked to several health benefits, particularly in cardiovascular health. The soluble fiber, such as pectin, and polyphenols in apples help lower cholesterol levels and reduce inflammation, contributing to a decreased risk of heart disease; a systematic review and meta-analysis found that apple or apple polyphenol intake is associated with reduced cardiovascular disease risk factors.^[110] Additionally, apples have a low glycemic index of approximately 36-39, meaning they cause a gradual rise in blood sugar levels, making them suitable for blood glucose management.^[111] A medium-sized apple, weighing about 182 grams, provides around 95 calories and 4.4 grams of fiber, which can fulfill roughly 15-18% of the recommended daily fiber intake of 25-30 grams for adults, supporting satiety and gut health when incorporated into a balanced diet.^[112]

Toxicity Concerns

Apple seeds contain amygdalin, a cyanogenic glycoside that hydrolyzes to release hydrogen cyanide (HCN), a toxic compound, when the seeds are chewed or crushed.^[113] The amygdalin concentration in seeds from various apple cultivars typically ranges from 1 to 4 mg per gram.^[114] Upon breakdown, this yields approximately 0.06 to 0.2 mg of cyanide equivalents per gram of seeds.^[115] The lethal oral dose of cyanide for humans is estimated at 1 to 2 mg per kg of body weight, equivalent to roughly 70 to 140 mg for a 70 kg adult.^[116] However, intact seeds are largely indigestible and pass through the gastrointestinal tract without significant HCN release, rendering accidental ingestion of small quantities harmless.^[113] Pesticide residues, often concentrated on the apple's skin, pose another potential toxicity concern, particularly from organophosphates and other chemicals used in conventional cultivation.^[117] Regulatory bodies enforce strict limits to ensure safety; for instance, the European Union sets default Maximum Residue Levels (MRLs) at 0.01 mg/kg for pesticides not explicitly listed, with specific higher thresholds for approved substances on apples.^[118] Washing apples with water or baking soda solutions can remove a portion of surface residues, while peeling eliminates most, though systemic pesticides absorbed into the flesh may persist at low levels.^[119] In rare instances, apples cultivated in soils contaminated by industrial pollution may accumulate heavy metals such as lead, copper, and arsenic, with concentrations in fruit tissues increasing in line with soil levels in affected areas.^[120] Such cases are uncommon and typically confined to specific polluted regions, where monitoring programs assess risks to consumers.^[121] To minimize toxicity risks, guidelines advise against consuming apple seeds or the core, emphasizing the edible flesh as the safe portion; thorough washing and optional peeling further reduce potential exposures from residues.^[113]

Allergic Reactions

Allergic reactions to apples primarily manifest as immunoglobulin E (IgE)-mediated hypersensitivities, with the most common form being oral allergy syndrome (OAS), which arises from cross-reactivity between apple proteins and birch pollen allergens.^[122] In OAS, the major allergen Mal d 1, a pathogenesis-related protein homologous to Bet v 1 in birch pollen, triggers mild symptoms such as oral itching, tingling, or swelling of the lips, mouth, and throat shortly after consuming raw apples.^[122] This cross-reactivity affects approximately 50-70% of individuals allergic to birch pollen, particularly in temperate regions like Central and Northern Europe where birch sensitization is prevalent.^[123] True IgE-mediated apple allergy, independent of pollen cross-reactivity, is rarer and typically involves sensitization to lipid transfer proteins (LTPs) such as Mal d 3, leading to more severe systemic reactions including urticaria, gastrointestinal distress, or anaphylaxis.^[124] This form is more common in Mediterranean countries, where primary food allergies to LTPs in Rosaceae fruits like apples are frequent, often resulting in life-threatening episodes upon ingestion.^[125] Unlike OAS allergens, Mal d 3 is heat-stable, persisting in cooked or processed apples and contributing to broader reactivity with other LTP-containing foods.^[126] The overall prevalence of apple allergy in the general population is estimated at 1-2%, though it rises significantly among atopic individuals with pollen allergies, reaching up to 3% in birch-endemic areas.^[127] For OAS specifically, symptoms are often seasonal and confined to raw fruit consumption, as cooking or peeling denatures heat-labile proteins like Mal d 1, reducing allergenicity.^[122] In contrast, LTP-mediated reactions show no such mitigation from processing, highlighting regional differences in allergy profiles.^[125] Management of apple allergies centers on avoidance of raw apples for OAS sufferers and comprehensive dietary exclusion for those with LTP sensitization, supplemented by emergency measures like epinephrine for severe cases.^[128] Diagnosis typically involves skin prick testing with fresh apple (prick-prick method) or specific IgE blood tests to confirm sensitization to Mal d 1 or Mal d 3.^[129] For birch pollen-related OAS, sublingual immunotherapy targeting birch allergens has shown efficacy in increasing tolerance to apples and reducing skin reactivity, though apple-specific immunotherapy remains experimental.^[130]

Practical Applications

Culinary Applications

Apples are widely utilized in culinary contexts due to their versatility, crisp texture, and balance of sweetness and tartness, making them suitable for both raw and cooked preparations across various global cuisines. In fresh applications, apples serve as a popular ingredient in salads and snacks, where their juicy flesh adds crunch and flavor. For instance, varieties like Granny Smith, known for their tart profile, are often sliced into green salads with greens, nuts, and vinaigrette, while sweeter types such as Fuji provide a refreshing contrast in fruit platters or as standalone snacks. Tart apples like Braeburn are particularly favored for baking due to their firm texture that holds up during cooking without becoming mushy. Cooked preparations highlight apples' ability to soften and release natural sugars, forming the basis of iconic desserts and condiments. In the United States, apple pie—typically filled with a spiced mixture of sliced apples, cinnamon, and sugar encased in pastry—is considered a national symbol of comfort food, with recipes dating back to colonial times. Applesauce, made by simmering peeled and cored apples with water or cider until they break down into a smooth puree, is a simple side dish or baby food staple. Fermented apple cider, produced by yeast conversion of apple sugars, yields a beverage with 4-8% alcohol by volume, commonly enjoyed during autumn festivities. Internationally, apples feature prominently in diverse dishes that showcase regional techniques. Austrian Apfelstrudel consists of thinly stretched dough wrapped around a filling of grated apples, raisins, breadcrumbs, and cinnamon, then baked to a flaky golden crisp. The French tarte tatin inverts caramelized apples baked under pastry, creating a rustic upside-down tart that emphasizes the fruit's caramelized sweetness. In the United Kingdom, toffee apples—whole fruits coated in a hard sugar syrup—are a traditional treat at fairs and Halloween celebrations. Apples pair harmoniously with savory elements, enhancing dishes through their acidity and subtle sweetness; for example, they are commonly roasted alongside pork to cut through the meat's richness, as in German-style schnitzel accompaniments, or matched with cheeses like sharp cheddar or brie for balanced charcuterie boards. Their seasonal availability peaks in fall harvests, prompting culinary traditions like harvest festivals where fresh-picked apples are incorporated into immediate preparations such as mulled ciders or baked goods to celebrate the autumn yield.

Industrial and Other Products

Apples are commercially processed into a variety of beverages, with apple juice and cider being the most prominent. Apple juice is produced by crushing and pressing apples, followed by filtration and pasteurization, accounting for a substantial share of U.S. apple utilization; approximately 52% of apples available for domestic consumption are used to produce juice or cider.^[131] In the United States, juice and cider represent about 13% of total apple production, making it a key industrial output.^[132] Cider, an alcoholic beverage, is obtained through the fermentation of apple juice using yeast, converting sugars into ethanol, and is a traditional product in regions like North America and Europe.^[133] Apple cider vinegar is derived from cider via a secondary acetic acid fermentation process, where acetic acid bacteria (such as Acetobacter species) oxidize the ethanol into acetic acid, typically achieving 4-5% acidity for commercial use.^[134] Beyond beverages, apples are transformed into various food derivatives on an industrial scale. Dried apple slices are prepared by washing, peeling, coring, slicing, and dehydrating fresh apples using methods like hot air drying or freeze-drying to extend shelf life and concentrate flavors, meeting standards set by the USDA for moisture content and texture.^[135] Apple puree, a smooth blend of cooked and mashed apples, serves as a base for baby food, providing essential nutrients like vitamins and fiber in commercial formulations that undergo heat processing for safety and consistency.^[136] Pectin, a soluble fiber extracted primarily from apple pomace through hot acidified water extraction and precipitation, is widely used as a gelling agent in jams, jellies, and confectionery, with apple-derived pectin comprising about 14% of global production due to its effective gel-forming properties in low-sugar environments.^[137] In non-food applications, apples contribute to cosmetics, biofuels, and animal feed. Malic acid, naturally abundant in apples, is isolated and incorporated into skincare products as a gentle alpha-hydroxy acid (AHA) for exfoliation, promoting smoother skin by removing dead cells without irritation, often sourced from apple extracts in organic formulations.^[138] Apple waste, including pomace, is fermented to produce bioethanol as a biofuel; industrial processes hydrolyze sugars and ferment them with yeast like Saccharomyces cerevisiae, yielding 0.134 g ethanol per gram of dry pomace, offering a sustainable alternative to fossil fuels.^[139] Dried or ensiled apple pomace is utilized as a nutritious animal feed ingredient, providing digestible fiber (8-14% sugars) and energy equivalent to corn silage, safely included at up to 20-30% in diets for pigs, cattle, and poultry to enhance palatability and growth without compromising performance.^[140] Apple processing generates significant byproducts, notably pomace, which constitutes 25-30% of the fruit weight after juice extraction and represents about 25% of the processed harvest volume globally.^[141] This fibrous residue, rich in organic matter, is composted or applied directly as a biofertilizer to improve soil microbial activity, nitrogen transformations, and crop yields, such as enhancing spring wheat productivity by supplying nutrients and reducing nitrate leaching.^[142]

Organic Farming Methods

Organic apple farming emphasizes chemical-free cultivation practices that prioritize soil health, biodiversity, and ecological balance to produce apples without synthetic inputs. These methods align with certification standards established by programs like the USDA National Organic Program (NOP), which prohibit the use of synthetic pesticides, herbicides, and fertilizers in crop production. Instead, organic apple growers rely on natural amendments and cultural practices to maintain fertility and suppress weeds, ensuring that land used for organic production has had no prohibited substances applied for at least 36 months prior to harvest.^[143] Key soil health practices include the integration of cover crops, such as clover or vetch, between orchard rows to prevent erosion, enhance nutrient cycling, and provide habitat for beneficial insects. Compost, prepared according to NOP guidelines (e.g., reaching temperatures of 131–170°F for 3–15 days), serves as a primary fertilizer to build organic matter and improve soil structure, reducing the need for external inputs. These approaches not only sustain long-term orchard productivity but also mitigate environmental impacts associated with conventional farming.^[144]^[143] Among the techniques employed, integrated pest management (IPM) plays a central role, focusing on monitoring pest populations and deploying natural predators to control outbreaks. For instance, conserving predatory mites (e.g., Typhlodromus pyri) and parasitic wasps through selective spraying and habitat enhancement helps suppress common pests like codling moth and plum curculio without broad-spectrum interventions. Biodynamic farming represents an extension of these organic principles, incorporating specific preparations—such as fermented herbal mixtures applied to soil and trees—to boost vitality and resilience in apple orchards, as demonstrated in comparative studies of biodynamic and conventional systems.^[145]^[146] Organic apple production faces notable challenges, including elevated pest and disease pressure due to the absence of synthetic controls. Apple scab (Venturia inaequalis), a primary concern, is managed through applications of sulfur-based fungicides, which require precise timing—often before or during rainy periods—to be effective, yet they can leave residues that affect fruit quality. Yields in organic systems are typically 20–40% lower than in conventional orchards, averaging 10–20 tons per acre compared to higher outputs in non-organic settings, largely owing to these management constraints and weather variability.^[144] In terms of market dynamics, organic apples constitute approximately 5–10% of total production in major regions like the United States and Europe, with certified organic acreage accounting for about 7% of U.S. apple land as of 2024.^[147] This segment has experienced steady growth since the 2000s, driven by rising consumer demand for sustainable produce— with 16% of Washington's 2024 crop being organic—leading to premium pricing often 40–70% higher than conventional apples, which supports economic viability despite lower yields. Expansion has been particularly pronounced in Europe and the U.S., where retail distribution and policy incentives have bolstered adoption.^[148]^[149]^[150]^[132]

Genetic Modifications for Quality

Genetic modifications in apples primarily target quality enhancements such as reduced enzymatic browning, improved nutritional profiles, and increased resistance to diseases, aiming to minimize post-harvest losses and boost consumer appeal. These transgenic approaches involve inserting or silencing specific genes to alter biochemical pathways, distinct from conventional breeding methods. One prominent example is the development of non-browning varieties, which address the oxidation reaction triggered by polyphenol oxidase (PPO) enzymes when apple flesh is exposed to air.^[151] The Arctic Apple, developed by Okanagan Specialty Fruits, exemplifies RNAi-based genetic engineering to silence the PPO gene. This technique uses RNA interference to suppress expression of the apple's own PPO genes, preventing the enzymatic browning that occurs upon cutting or bruising. Approved for deregulation by the U.S. Department of Agriculture in 2015 for Golden Delicious and Granny Smith varieties, the Arctic Apple maintains its fresh appearance for up to 28 days after slicing, compared to rapid discoloration in conventional apples within hours. Canada also approved these varieties in 2015, allowing commercial planting and sale. The modification does not alter taste, texture, or nutritional content but extends shelf life for pre-sliced products, potentially reducing food waste in retail and food service sectors.^[152]^[153]^[154]^[155] Beyond browning resistance, genetic modifications have targeted nutritional enhancements, particularly elevating antioxidant levels. Researchers have engineered apples to overexpress genes for flavonoids and anthocyanins, key polyphenols that combat oxidative stress. For instance, transgenic apples with increased anthocyanin production in the flesh achieve up to a 120-fold higher content—around 11,000 micrograms per apple—compared to standard varieties, potentially amplifying health benefits like reduced inflammation. These modifications draw from natural genetic variation but use transgenic insertion for stable, high-level expression. Similarly, efforts to boost overall antioxidant capacity involve altering pathways for phenolic compounds, as seen in laboratory-modified lines with elevated quercetin and catechin levels.^[156]^[157] Disease resistance represents another quality-focused application, paralleling successes like the virus-resistant papaya. In apples, genetic engineering has introduced genes for resistance to fire blight, a bacterial disease caused by Erwinia amylovora that devastates orchards. Transgenic Gala apples incorporating the fb_E2 attacin gene from wild relatives exhibit reduced susceptibility, with field tests showing minimal lesion development post-inoculation. Cisgenic approaches, using only apple-derived genes, have similarly conferred fire blight tolerance by overexpressing resistance loci like Mr5 from Malus wild species. For fungal threats like apple scab (Venturia inaequalis), the HcrVf2 gene from wild apple Malus floribunda has been transgenically inserted into susceptible cultivars such as Gala, resulting in immune responses in transformed lines. These modifications aim to lower pesticide needs while preserving fruit quality.^[158]^[159]^[160] Regulatory approval for GM apples varies globally, with the U.S. and Canada permitting commercialization of the Arctic varieties since 2015, based on assessments confirming no increased risks to health or the environment. In contrast, the European Union maintains strict oversight under Directive 2001/18/EC, requiring case-by-case authorizations; while field trials for GM apples have occurred, no transgenic varieties like Arctic have been approved for cultivation or sale due to precautionary principles and traceability demands. Public concerns center on mandatory labeling to inform consumer choice, with U.S. bioengineered food disclosure rules implemented in 2022 addressing transparency, though advocacy groups highlight potential allergenicity and long-term ecological impacts. In Canada, voluntary labeling persists amid calls for reform.^[161]^[162]^[163]^[164] Adoption of GM apples remains limited, comprising less than 0.1% of the global apple market as of 2024, constrained by consumer skepticism and supply chain preferences for non-GM produce. The focus is on waste reduction in processing, where non-browning traits cut trim losses by up to 20% in slicing operations, supporting sustainability in an industry where post-harvest discards account for significant economic losses. Commercial planting of Arctic varieties is confined to select U.S. and Canadian regions, with gradual expansion into retail channels.^[165]^[166]^[167]

Cultural and Symbolic Roles

Mythological References

In Greek mythology, the golden apples of the Hesperides were sacred fruits grown in a remote garden at the western edge of the world, guarded by the nymphs known as the Hesperides and a watchful dragon named Ladon.^[168] These apples, a wedding gift from Gaia to Hera, were believed to confer immortality upon those who consumed them, symbolizing eternal life and divine favor.^[169] As one of his Twelve Labors, the hero Heracles was tasked with retrieving these apples, navigating perilous obstacles to obtain three of them before returning them to their sacred grove.^[168] The golden apple also features prominently in the myth of Atalanta, a swift-footed huntress who challenged suitors to a footrace, with the winner claiming her hand in marriage or facing death upon defeat.^[170] Her suitor Hippomenes (or Melanion in some variants), aided by Aphrodite, received three golden apples from the Hesperides; during the race, he dropped them one by one, distracting Atalanta as she paused to retrieve their gleaming allure, allowing him to win.^[170] Another pivotal role for the apple in Greek lore is the Apple of Discord, thrown by the goddess Eris at the wedding feast of Peleus and Thetis to sow strife among the gods.^[171] Inscribed with the words "for the fairest," this golden apple ignited a contest between Hera, Athena, and Aphrodite, leading Zeus to assign the mortal Paris of Troy as judge; his award to Aphrodite in exchange for the love of Helen sparked the Trojan War.^[171] In Norse mythology, the goddess Iðunn safeguarded a basket of golden apples that granted eternal youth and vitality to the Aesir gods, preventing them from aging as long as they partook of the fruit regularly. Detailed in Snorri Sturluson's Prose Edda (Gylfaginning), the myth recounts how Loki's deception led to Iðunn's abduction by the giant Thjazi, causing the gods to wither until Loki retrieved her and the apples, restoring their vigor. The Biblical account in the Book of Genesis describes the forbidden fruit from the Tree of the Knowledge of Good and Evil in the Garden of Eden as simply "fruit," with no specific identification provided in the Hebrew text.^[172] However, by the 12th century in European Christian tradition, particularly influenced by Latin Vulgate translations and artistic depictions, this fruit became widely interpreted as an apple, symbolizing temptation and the fall of humanity through Adam and Eve's disobedience.^[173] This association, though not scriptural, permeated medieval literature and iconography, linking the apple to themes of original sin and divine prohibition.^[172] In Arthurian legend, Avalon—known as the Isle of Apples (Ynys Afallon in Welsh)—served as a mystical otherworldly realm where King Arthur was taken to heal from his fatal wounds after the Battle of Camlann.^[174] Described in Geoffrey of Monmouth's Historia Regum Britanniae (c. 1136), the island's abundant apple orchards evoked Celtic motifs of abundance and rejuvenation, with the fruits implying restorative powers tied to immortality and the sidhe.^[174] Slavic folklore features apples of youth in tales like Alexander Afanasyev's "The Bold Knight, the Apples of Youth, and the Water of Life," where golden apples from a distant garden restore youth and cure ailments for those who obtain them.^[175] In this Russian narrative, a tsar's sons embark on quests to retrieve these magical apples from a guarded tree, encountering dragons and trials that underscore the fruits' role as elixirs of rejuvenation and symbols of heroic triumph over mortality.^[175] Similar motifs appear in Romanian variants, such as "Prâslea the Brave and the Golden Apples," reinforcing the apple's enduring association with vitality across Eastern European oral traditions.

Religious and Folklore Significance

In Christianity, the apple became symbolically linked to the forbidden fruit in the Garden of Eden story through post-medieval artistic traditions, where it was frequently depicted in European religious art as the fruit tempting Eve, despite the Bible not specifying its type. This association arose from the Latin word malum, which means both "apple" and "evil," influencing medieval and Renaissance interpretations by scholars and artists who portrayed the apple as a symbol of original sin and temptation.^[173]^[172] Additionally, apple bobbing emerged as a custom tied to All Saints' Day celebrations, originating from harvest rituals that honored saints and the dead, where participants would attempt to catch floating apples with their teeth as a playful divination for future spouses or prosperity.^[176] In Celtic traditions, apples held significance during Samhain, the ancient festival marking the end of harvest and the thinning of the veil between worlds, where they were offered as food for wandering spirits or buried in graves to sustain souls awaiting rebirth, symbolizing fertility and the cycle of life.^[177]^[178] Irish folklore further associates apples with fairies, as seen in tales like Echtra Condla, where a fairy maiden offers a magical apple to the hero Connla, enchanting him and drawing him to the Otherworld, representing immortality and otherworldly allure.^[179]^[180] In Chinese folklore, the apple, known as ping guo, symbolizes peace and safety due to its phonetic similarity to ping an, meaning "peace" or "safety," a homophone that imbues the fruit with auspicious connotations in traditional beliefs.^[181]^[182] During Lunar New Year customs, red apples are commonly gifted or displayed in homes to invoke harmony and protection for the coming year, often arranged in platters alongside other fruits to attract good fortune and familial well-being.^[183]^[184] In Germanic paganism, particularly within Norse mythology, apples are tied to fertility rites through the goddess Iðunn, who safeguards golden apples that grant eternal youth to the gods, linking the fruit to renewal, vitality, and the Vanir deities associated with fertility and prosperity.^[185]^[186] These apples underscore themes of life-giving abundance in pre-Christian rituals, where such symbols reinforced communal hopes for bountiful harvests and generational continuity.^[187]

Proverbial and Idiomatic Uses

The idiom "apple of one's eye" refers to someone or something cherished above all others, originating from the literal description of the pupil of the eye, which was metaphorically valued for its preciousness due to the importance of sight.^[188] This phrase first appeared in Old English in the late 9th century during the reign of King Alfred, who used it figuratively to denote endearment.^[188] It gained prominence through biblical references in the King James Version, such as Deuteronomy 32:10, "He kept him as the apple of his eye," emphasizing protection and value.^[188] Another common English expression is "one bad apple spoils the bunch," which warns that a single corrupt or problematic individual can negatively influence an entire group.^[189] The proverb traces back to at least 1340 in English, with roots in earlier Latin forms like "A rotten apple quickly infects its neighbor," highlighting the literal spoilage of fruit through ethylene gas emission.^[189] Benjamin Franklin adapted it in 1736 as "The rotten apple spoils his companion" in Poor Richard's Almanack, evolving into the modern variant by the 19th century to underscore moral contagion.^[189] The proverb "An apple a day keeps the doctor away" promotes daily apple consumption for health benefits, serving as a longstanding slogan for preventive wellness.^[190] Its earliest recorded form appeared in the February 1866 issue of Notes and Queries magazine as a Pembrokeshire proverb: "Eat an apple on going to bed, And you’ll keep the doctor from earning his bread."^[190] By 1913, it had standardized in Elizabeth Wright's Rustic Speech and Folk-lore as "An apple a day keeps the doctor away," reflecting growing awareness of the fruit's nutritional value in promoting health.^[190] Cross-culturally, similar idioms exist, such as the French "Il suffit d'une pomme pourrie pour gâter tout le tas" (One rotten apple is enough to spoil the whole pile), which mirrors the English warning about the spread of negativity.^[191] In Japanese, the proverb "Kinomi wa moto e ochiru" (The fruit falls close to the root) equates to "The apple doesn't fall far from the tree," symbolizing how traits are inherited or behaviors persist within families or groups.^[192] Over time, apples in proverbial and idiomatic uses have evolved from literal references to the fruit's physical properties—such as spoilage or visual appeal—to broader metaphors representing health through nutrition, temptation via cultural associations, corruption in social dynamics, and perfection in valued possessions.^[189]^[190]^[188] This shift underscores the apple's symbolic versatility across languages, drawing on its ubiquity in agriculture and daily life to convey enduring human experiences.

Contemporary Cultural Impact

In contemporary literature and film, the apple retains powerful symbolic resonance, often representing temptation and transformation. The poisoned apple in Disney's 1937 adaptation of Snow White and the Seven Dwarfs has become an iconic emblem of deceit and the loss of innocence, influencing popular culture through countless retellings and parodies in media up to the 2020s.^[4]^[193] In modern novels, such as Margaret Atwood's The Handmaid's Tale (1985), apples appear in scenes representing class hierarchies and domestic control within the dystopian framework, a motif that persists in adaptations like the Hulu series.^[194] In visual art, Paul Cézanne's late-19th-century still lifes featuring apples, such as The Basket of Apples (c. 1893), continue to exert influence on modern aesthetics by elevating everyday objects to explore form, space, and perception, paving the way for Cubism and abstract art.^[195]^[196] This enduring legacy manifests in contemporary installations, where apples address ecological concerns; for instance, in 2024, artists Antje Majewski and Paweł Freisler organized the "Superhost" project at M HKA in Antwerp, displaying a year's worth of community-grown apples to promote urban greening and climate resilience through participatory workshops.^[197] Similarly, a 2025 initiative by UK artists planted apple trees inspired by Newton's legend at the Royal Academy of Arts, using the fruit as a "visual marker" of climate change impacts on agriculture.^[198]^[199] The apple's role in branding and public health campaigns further amplifies its cultural presence. Apple Inc.'s 1977 logo, featuring a bitten apple, draws on biblical allusions to knowledge and temptation from the Garden of Eden, evolving into a global symbol of innovation that permeates digital culture.^[200]^[201] Health initiatives like the UK's NHS "5 A Day" program, launched in 2003 and ongoing, prominently feature apples to encourage daily fruit consumption for disease prevention, contributing to widespread awareness of nutritional benefits.^[202] Recent trends highlight the apple's integration into sustainability and digital realms. In vegan culture, apples have surged in popularity as versatile, ethical staples, with 2024-2025 surveys indicating nearly half of U.S. consumers planning increased plant-based intake, often centering affordable fruits like apples in snack trends.^[203]^[204] Sustainability efforts by U.S. apple growers, including conservation and pollinator management practices, support climate-conscious consumerism.^[205]

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Jul 2, 2023 · There are several tests that are used to determine apple fruit maturity, and these will be described throughout this article.Apple Fruit Maturity Indices · Fruit Firmness Changes · Starch Content Changes
[28]
Taxon: Malus domestica - USDA
Malus domestica. Details ... Kingdom: Phylum: Class: Order: Plantae; Tracheophyta; Magnoliopsida; Rosales. Family: Genus: Species: Sub-Species: Variety: Rosaceae ...
[29]
Malus domestica (apple) | CABI Compendium
The domesticated apple has long been referred to as M. domestica Borkh (or M. x domestica to indicate its hybrid origin) (Korban and Skirvin, 1984), although ...
[30]
Malus x domestica | GDR
The domesticated apple (Malus x domestica) is the most economically important species in the Malus genus. Apples are one of the 20 most productive crops in the ...
[31]
Genome re-sequencing reveals the history of apple and supports a ...
Aug 15, 2017 · Cultivated apples likely originate from Malus sieversii in Kazakhstan, followed by intensive introgressions from M. sylvestris. M. sieversii in ...
[32]
New Insight into the History of Domesticated Apple - PubMed Central
May 10, 2012 · Here, we investigated the origins of the domesticated apple Malus domestica Borkh., one of the most emblematic and widespread fruit crops in ...
[33]
Crop-to-wild introgression in the European wild apple Malus ...
Sep 28, 2018 · Malus sylvestris (L.) Mill, the European wild apple, is a small tree native to Europe which reaches its north-western limit in Northern Britain.Introduction · Methods · Results · DiscussionMissing: subspecies | Show results with:subspecies
[34]
Evolution of Rosaceae Fruit Types Based on Nuclear Phylogeny in ...
... Pyrus, and Malus all have inferior ovaries, suggesting ... Phylogenetic relationships in tribe Spiraeeae (Rosaceae) inferred from nucleotide sequence data.
[35]
Phylogenomics insights into gene evolution, rapid species ...
Aug 3, 2023 · Phylogenomics insights into gene evolution, rapid species diversification, and morphological innovation of the apple tribe (Maleae, Rosaceae)
[36]
The genome of the domesticated apple (Malus × domestica Borkh.)
Aug 29, 2010 · Riccardo Velasco and colleagues report the genome sequence of the ... The genome of the domesticated apple (Malus × domestica Borkh.).
[37]
Pan-genome analysis of 13 Malus accessions reveals structural and ...
Nov 15, 2023 · The genome assembly sizes were estimated to be between 668.25–679.75 Mb, and to have heterozygosity rates of 0.88–1.67% and repeat rates of ...
[38]
Population Structure and Association Mapping for Agronomical and ...
Mar 9, 2023 · Most domesticated apple cultivars are diploid (2n = 2x = 34), even though triploid (2n = 3x = 51) and tetraploid (2n = 4x = 68) cultivars can ...
[39]
Population structure, relatedness and ploidy levels in an apple gene ...
Aug 15, 2018 · Using GBS to identify ploidy levels Triploid apples are relatively common among cultivars: they have a large fruit size and vigorous growth. We ...
[40]
Expression of the polyphenol oxidase gene MdPPO7 is modulated ...
Dec 23, 2024 · MdPPO7 contributes to apple browning after cutting. MdWRKY3 binds to MdPPO7's promoter, positively regulating its expression to promote ...
[41]
Rvi6-mediated defense mechanisms against apple scab - PMC
Aug 26, 2025 · Although the key resistance gene Rvi6 (resistance to Venturia inaequalis 6) has been widely deployed in apple scab-resistant breeding programs, ...
[42]
Genomic consequences of apple improvement | Horticulture Research
Jan 1, 2021 · Velasco, R. et al. The genome of the domesticated apple (Malus x domestica Borkh.). Nat. Genet. 42, 833–9 (2010). Article CAS PubMed ...
[43]
How the Silk Road Created the Modern Apple - Smithsonian Magazine
Aug 21, 2017 · The analysis reveals that about 46 percent of the domestic apple genome comes from the Kazakhstan population of the Malus sieversii, while 21 ...
[44]
Fruit: A History of Apples - Hobby Farms
A tablet found in Mesopotamia dating from 1500 B.C. records the sale of an apple orchard by an Assyrian named Tupkitilla. The price: the princely sum of three ...Missing: Near | Show results with:Near
[45]
Apple Farming in Washington - HistoryLink.org
Sep 6, 2021 · The industry thrived during the World War II years because of a strong apple market. ... The History of the Apple Industry in the Wenatchee ...
[46]
Theophrastus, Enquiry Into Plants - ToposText
The fig grows in all the ways mentioned, except from root-stock and cleft wood; apple and pear grow also from branches, but rarely. ... apple with the first cold ...
[47]
Orchard History: Fruit Introduction and Colonization, 1600-1800
Oct 20, 2022 · The 1600 to 1800 period of fruit introduction and colonization in America centered on the 13 original colonies and the Spanish missions of the Southwest.
[48]
HWWG Natural & Human History of Grafting and Budding
Clonal selection of apples, at least partially by grafting, was underway in France during the 15th century. The Frenchman, Caylus, wrote a Historie du ...
[49]
Origins of Apples in America | NC Historic Sites
The first apple trees in North America grew from seeds brought by French Jesuits in the late sixteenth century. It is not known if the settlers of Jamestown ...
[50]
1788 First apple trees in Tasmania - Australian Food Timeline
In 1788, William Bligh planted seven apple trees at Adventure Bay when he anchored there in the Bounty, the first apple trees in Tasmania.
[51]
Orchard History: Fruit Monoculture and Orchard Intensification, 1946 ...
Apr 5, 2023 · The trend towards higher density, dwarf fruit tree orchards in the period from World War II to the present was fueled by the need to lower costs of production.Missing: peak | Show results with:peak
[52]
[PDF] Globalization of the Washington Apple Industry
States, where production averaged between 4.5 and 5.5 billion pounds. Technological developments did continue after the war, with one of the more ...
[53]
Apple breeding at the University of Minnesota
Apple breeding takes time and patience. From the initial cross to release date, it takes on average 20-30 years for a new apple variety to hit the market. Our ...Missing: methods | Show results with:methods
[54]
Why It Takes Decades To Breed New Apple Varieties
Nov 18, 2022 · The growing process is time-consuming, and it can take six to seven years before a young tree produces the number of apples needed for proper ...
[55]
Crossing Apples - Rittman Orchards
May 23, 2019 · Crossing apples involves using pollen from one variety to pollinate another, resulting in a new variety with half the genetics of each parent.
[56]
Molecular selection in apple for resistance to scab caused ... - PubMed
We present here the transformation of two RAPD markers, which we found previously to be linked to the major scab resistance gene Vf, into more reliable and ...
[57]
MARKER ASSISTED SELECTION FOR SCAB RESISTANT APPLE
A previous molecular analysis, conducted randomly on a part of the population, showed that the Vf resistance gene and the scab resistance gene present in the ...
[58]
Rootstocks - WSU Tree Fruit - Washington State University
In order to get trees that are more suited to high-density plantings, rootstocks with dwarfing traits and precocity need to be used. In many parts of the ...
[59]
Influence of nine dwarfing apple rootstocks on vigour and ...
Jul 27, 2011 · Rootstock M.9 has become most dominant for apples because of its suitability for high-density plantings (Wertheim, 1997). At present, more ...
[60]
Efficient Genome Editing in Apple Using a CRISPR/Cas9 system
Aug 17, 2016 · Four guide RNAs (gRNAs) were designed and stably transformed with Cas9 separately in apple. Clear and partial albino phenotypes were observed in ...
[61]
Targeted Gene Editing in Pome Fruit Genetics and Breeding
Dec 15, 2023 · In particular the emergence of genome-editing tools such as the CRISPR/Cas9 technology can significantly improve the speed and accuracy of pome ...
[62]
Promoter Variation of Key Apple Gene MdPG1 & Upstream Regulation
Mar 26, 2023 · MdPG1 encoding polygalacturonase in apple (Malus × domestica) is a key gene associated with fruit firmness and texture variations among apple cultivars.
[63]
[PDF] Pollination of apples by honey bees
FLOWER MORPHOLOGY. Apple blossoms have five petals and numerous stamens, but the ovary consists of five carpels, each of which contain two ovules. For ...
[64]
https://extension.psu.edu/orchard-pollination-pollinizers-pollinators-and-weather/
[65]
Cross-compatibility of apple cultivars and pollinizers - WSU Tree Fruit
Many members of the family Rosaceae, including cultivated apples (Malus ×domestica) and their relatives (Malus spp.), exhibit gametophytic self-incompatibility.
[66]
Home Orchard Calendar - Penn State Extension
Jun 21, 2023 · Calendar applies to Zone 6, in the Mid-Atlantic Region. Apples bloom generally the last week of April. Adjust timings for other regions.
[67]
Honey Bee Management in Tree Fruit Orchards
Apr 1, 2022 · Depending on the crop, using one to two hives per acre is adequate for most orchards. Distribute hives throughout the orchard bearing in mind ...
[68]
Orchard Pollination: Pollinizers, Pollinators, and Weather
Apr 13, 2023 · Pollination involves the integration of several biological and physical factors, including cultivar compatibility, synchronous blooming, insects, and proper ...
[69]
Apple | UC Marin Master Gardeners
Scientific Name Malus domestica. General Information Apple trees are deciduous, attractive trees native to central Asia and cultivated worldwide.<|separator|>
[70]
[PDF] apples_2015.pdf - Aggie Horticulture - Texas A&M University
Consecutive low-chilling years can actually kill unadapted apple trees. Apple trees prefer deep, well-drained soils with a pH of 6.5 to 7.0. Production is ...
[71]
Growing Apples in Virginia | VCE Publications
Not recommended except on very well-drained soils. Dwarf. General: Mature dwarf trees may produce 50 to 100 lbs. of fruit. Trees on dwarfing rootstocks and on M ...
[72]
[PDF] Apple Tree Growth & Development
Their major growth phases are propagation and juvenility, vegetative de- velopment, maturity, and senescence. This life phase lasts two years.
[73]
Growing apples in the home garden | UMN Extension
Dwarf apple trees will start bearing fruit 2 to 3 years after planting. Standard size trees can take up to 8 years to bear fruit. Some varieties are more ...Selecting Plants · Apple Varieties For Home... · How To Keep Your Apple Trees...
[74]
Controlled Atmosphere Storage of Apples
Sep 10, 2021 · Controlled atmosphere (CA) storage manipulates CO2 and O2 to maintain apple quality and shelf life by slowing respiration, increasing shelf ...
[75]
Controlled Atmosphere Storage - Cooperative Extension: Tree Fruits
Controlled atmosphere (CA) storage lengthens fruit life by reducing oxygen and increasing carbon dioxide, slowing ripening and maintaining firmness. Oxygen is ...
[76]
Thanks To Science, You Can Eat An Apple Every Day - NPR
Nov 26, 2018 · After harvest, apples can be stored for months in controlled atmosphere storage rooms where the temperature, oxygen, carbon dioxide, ...
[77]
Controlled Atmosphere Storage - Northwest Horticultural Council
CA storage controls temperature, humidity, oxygen, and carbon dioxide to extend shelf life of fresh products by reducing oxygen levels.
[78]
Postharvest Quality of Granny Smith Apples: Interplay of Harvest ...
Depending on the variety, apples can be successfully stored for 4 to 10 months under optimal conditions; for example, chilling-insensitive Granny Smith can be ...
[79]
A Review of Storage Temperature Recommendations for Apples ...
The storage temperature can affect the humidity and vapour pressure deficit (driving force) in the storage room.
[80]
Storing Apples | Home & Garden Information Center - Clemson HGIC
Nov 8, 2021 · Thicker skinned and harder apples tend to keep longer; 'Arkansas Black', 'Stayman Winesap', 'Rome', 'Fuji' or 'Pink Lady' can last five months ...
[81]
What are the ways to extend the shelf life of apples? - Citrus Freight
Sep 10, 2025 · Extend apple shelf life with enhanced packaging, ethylene control, cold chain management, humidity monitoring, pre-cooling, and controlled ...
[82]
Let's Preserve: Apples - Penn State Extension
Jul 10, 2025 · A quick-reference guide on how to properly preserve apples, including tips on preparation, freezing, canning, and more.
[83]
Characterization and Quantification of Postharvest Losses of Apple ...
The apple losses ranged from 18.3% to 26.6%, considering the 4-year mean for all physiological disorders and fungal decay during CA storage, plus the fungal ...Missing: global | Show results with:global
[84]
Review on postharvest quality and handling of apple
Annual losses of many fruits, including apple, vary from 5 to 35% and for developing countries, it ranges between 20%-50%. The most common causes are poor ...Missing: percentage | Show results with:percentage<|control11|><|separator|>
[85]
The Red Delicious is an apple atrocity. Why are we growing billions ...
Mar 13, 2018 · To understand the modern Red Delicious, it helps to know its roots. In the 1870s, a farmer named Jesse Hiatt discovered a rogue apple tree ...
[86]
Malus domestica 'Red Delicious'
This tree grows well in slightly acidic, moist, and well-drained soil. This apple tree does not tolerate a long drought period.
[87]
Malus domestica 'Fuji' (Fuji Apple)
The Fuji apple tree is a dwarf cultivar of M. domestica created in Japan in the 1930's with two American apple varieties, Red Delicious and old Virginia Ralls ...
[88]
Apple - Fuji - tasting notes, identification, reviews - Orange Pippin
Fuji is is by far the most successful Japanese apple variety, known for its crisp sweet-flavored flesh and long storage potential.
[89]
History of the Granny Smith Apple - HGTV
It all began in 1868 with a chance seedling discovered growing from a pile of discarded crab apples somewhere near Sydney, Australia.Missing: traits | Show results with:traits
[90]
Apple - Granny Smith - tasting notes, identification, reviews
Like all the best old varieties it has a bizarre history, being discovered in Austrialia in the 1860s as a seedling growing in the remains of a rubbish tip. The ...
[91]
Golden Delicious Apples Information and Facts - Specialty Produce
Geography/History Golden Delicious apples arose as a chance seedling on a family farm in the hamlet of Odessa in Clay County, West Virginia. In 1891, J.M. ...
[92]
Apple - Golden Delicious - tasting notes, identification, reviews
Golden Delicious is a popular, easy-to-grow apple, with a sweet, rich flavor when ripe, and is versatile for dessert and cooking.
[93]
Apple - Cox's Orange Pippin - tasting notes, identification, reviews
This is the classic English apple, often regarded as the finest of all dessert apples, and the inspiration for this website.
[94]
An apple a day … the scientific way | University of Minnesota
Oct 6, 2022 · The U of M apple breeding program has been sustaining Minnesota's small but thriving commercial apple industry since the early 1900s.Missing: 1990s crispness
[95]
Apple - Gala - tasting notes, identification, reviews - Orange Pippin
Because Gala is so widely grown, and keeps well in storage, it is not unusual around April/May and September/October to see Gala apples for sale from both ...United States · United Kingdom
[96]
Apple - Pink Lady - tasting notes, identification, reviews
'Pink Lady' is a registered trade mark of Apple and Pear Australia Limited ... Disease resistance: Average; Scab (Apple and Pear): Some susceptibility ...
[97]
Apple scab | Pome fruits | Plant diseases | Biosecurity
Mar 6, 2024 · The most highly susceptible cultivars are: Cripps Pink (including Pink Lady™); Lady William. Some cultivars show some resistance, including:.
[98]
[PDF] Fresh Apples, Grapes, and Pears: World Markets and Trade
Dec 2, 2024 · Output in Washington, normally the top pear-producing state, is projected to drop more than 30 percent and be a smaller harvest than Oregon.
[99]
World Apple Production by Country - AtlasBig.com
China is the largest apple producer in the world with 45,984,858 tonnes production per year. · Turkey comes second with 4,493,264 tonnes yearly production. · With ...
[100]
[PDF] MARKET INTELLIGENCE REPORT APPLES
minor apple-producing countries, collectively produced 6,354,450,000 tons of apples (USDA, 2024). Figure 2: Top global apple-producing countries in 2023.Missing: tonnes | Show results with:tonnes
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https://link.springer.com/article/10.1007/s13593-017-0422-1
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Apples, fresh | Imports and Exports | 2023 - TrendEconomy.com
Nov 7, 2024 · In 2023, the world exports of "Apples, fresh" exceeded $6.69 billion (according to external trade statistics of 95 countries). It was $6.63 ...
[103]
Water Footprint of Food Guide
The Water Footprint of Food Guide gives you water footprint information for more than 100 common foods and beverages.
[104]
Ecosystem services in orchards. A review
Apr 4, 2017 · Fruit yield, which could reach up to 140 t/ha in apple orchards, is increased by light interception, carbon allocation, and nitrogen and water ...
[105]
Apple Market - Size, Size, Growth & Analysis - Mordor Intelligence
Sep 29, 2025 · The apple market size reached USD 107.2 billion in 2025 and is projected to grow to USD 130.1 billion by 2030, registering a CAGR of 3.9%. The ...
[106]
Climate Change Threatens Apple Production
In 2012, an exceptionally warm January and February caused the apple trees to break bud and bloom three to four weeks earlier than expected. Soon after, the ...
[107]
https://pmc.ncbi.nlm.nih.gov/articles/PMC6770235/
[108]
Apples and Cardiovascular Health—Is the Gut Microbiota a Core ...
Apples are a rich source of polyphenols and fiber. The main polyphenol classes in increasing order are: Dihydrochalcones, flavonols, hydroxycinnamates and ...
[109]
Malus domestica: A Review on Nutritional Features, Chemical ...
Oct 22, 2020 · The main structural classes of apple constituents include polyphenols, polysaccharides (pectin), phytosterols, and pentacyclic triterpenes.Missing: biochemical | Show results with:biochemical
[110]
Chemical Composition and Antioxidant Activity of the Main Fruits ...
Sep 10, 2019 · [10]: low (<100 mg of GAE/100 g FW), medium (100–500 mg of GAE/100 g FW), and high contents (>500 mg of GAE/100 g FW). According to this ...2. Material And Methods · 3.1. Total Phenolic... · 3.3. Organic Acids Contents<|separator|>
[111]
Identification, Comparison and Classification of Volatile Compounds ...
May 11, 2021 · Hexyl acetate, hexyl hexanoate, and hexyl 2-methylbutyrate, which are the most important esters, have a great influence on apple aroma because ...
[112]
Determination of Post-Harvest Biochemical Composition, Enzymatic ...
Jan 18, 2021 · Polyphenol oxidase (PPO; EC 1.10.3.1) is the most prevalent enzyme involved in the apple flesh browning process [14]. Generally, in plant cells, ...
[113]
a systematic review and meta-analysis - PubMed
Sep 24, 2021 · Apple or apple polyphenol intake is associated with a reduced risk of cardiovascular diseases. These results are consistent with the old saying that eating an ...
[114]
Best Fruits for Diabetes: Glycemic Index, List of Healthy Fruits
Jul 31, 2024 · The glycemic index (GI) measures how a food affects your blood sugar. ... Apples (GI of 39); Bananas (GI of 55); Pears (GI of 30). Blueberries ...
[115]
Apples - SNAP-Ed Connection - USDA
Apples are in season in spring, summer, fall, and winter. They come in over 2,500 varieties, and a medium apple has 95 calories.
[116]
Apple seeds: Are they poisonous? - MedicalNewsToday
Feb 16, 2024 · When a person chews an apple seed, a compound within it releases cyanide, which is toxic ... A lethal dose of cyanide is around 50–300 milligrams ...
[117]
Determination of amygdalin in apple seeds, fresh apples and ...
Amygdalin contents of seeds from fifteen varieties of apples ranged from 1 mg g−1 to 4 mg g−1. The amygdalin content of commercially-available apple juice was ...
[118]
Determination of amygdalin in apple seeds, fresh apples and ...
Depending on the cultivar, amygdalin is present in seeds around 1À4 mg/g, and it can release 0.06À0. 2 mg/g of cyanide equivalents per gram of apple seeds ( ...<|control11|><|separator|>
[119]
Are Apple Seeds Poisonous? - Healthline
May 29, 2018 · According to the Centers for Disease Control and Prevention (CDC)Trusted Source , 1–2 mg/kg is a fatal oral dose of cyanide for a 154 lbs. (70 ...Missing: cyanogenic HCN
[120]
EWG's 2025 Shopper's Guide to Pesticides in Produce | Apples
Most conventionally grown apples are drenched in diphenylamine, a pesticide banned in Europe, to prevent discoloration while in cold storage.
[121]
EU legislation on MRLs - Food Safety - European Commission
EU legislation harmonizes pesticide MRLs, applies to 315 fresh products, covers around 1100 pesticides, and has a default MRL of 0.01 mg/kg.
[122]
Effectiveness of Commercial and Homemade Washing Agents in ...
Nov 8, 2017 · In practical application, washing apples with NaHCO3 solution can reduce pesticides mostly from the surface. Peeling is more effective to remove ...
[123]
Distribution of Heavy Metals in Apple Tissues Grown in the Soils of ...
The concentration of Ni, Pb, Cr, Fe etc. in the apple tissue increased along with the heavy metal content in soils in the polluted area. The high accumulation ...
[124]
Human Health Risk Assessment on the Consumption of Apples ...
Feb 5, 2021 · The results of the study showed that the maximum levels of heavy metals were exceeded in apple samples from all sampling sites: Pb—from 11.47 to ...
[125]
Apple allergy: Causes and factors influencing fruits allergenic ...
Jun 2, 2021 · These symptoms known as an Oral Allergy Syndrome (OAS) are usually mild and occur directly after exposure to the allergens. The apple allergens ...
[126]
Revisiting Fruit Allergy: Prevalence across the Globe, Diagnosis ...
The study showed 70% of birch-sensitized patients had allergic symptoms to fruits, especially Rosaceae fruits (apple, pear, cherry, peach, plum, apricot, etc.), ...
[127]
f435 Mal d 3 - Thermo Fisher Scientific
Mal d 3 sensitization can serve as a marker for primary sensitization to apple allergy and can differentiate true apple allergy from the birch-apple syndrome.Missing: mediated | Show results with:mediated
[128]
Apple allergy: Causes and factors influencing fruits allergenic ...
Apr 14, 2021 · In the Mediterranean, apple allergies are less frequent but severe and associated with sensitisation to LTPs. (Mal d 3). Variation in the Mal d ...
[129]
Comprehensive review of pollen-food allergy syndrome
For example, Mal d 1 in apple is PR-10, which rarely causes systemic symptoms, whereas Mal d 3 is LTP, which maintains IgE reactivity despite heat treatment ...Review Article · Epidemiological Studies · Prevention & TreatmentMissing: true | Show results with:true
[130]
Assessment of allelic diversity in intron-containing Mal d 1 genes ...
The prevalence of apple allergic individuals mounts up to ~3% in Central and Northern Europe. This type of apple allergy is caused by cross reactivity of IgE ...
[131]
Apple Allergy Symptoms & Treatment - New York ASC
During a skin prick test, a small amount of apple extract is placed on the skin, and a small prick is made through the extract. An allergist may also test if ...
[132]
Allergy information for: Apple (Malus domestica)
Often however, apple allergy in the Mediterranean area is seen independently from pollen allergies. In these cases, another allergen is responsible for the ...Missing: true | Show results with:true
[133]
Oral birch pollen immunotherapy with apples: Results of a phase II ...
Feb 23, 2021 · After 8 months of therapy, patients showed increased tolerance to apples (p < .001) and a decreased skin reactivity to apples.Missing: management | Show results with:management
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https://chem.libretexts.org/Bookshelves/Biological_Chemistry/Fermentation_in_Food_Chemistry_%28Graham%29/01%253A_Modules/1.06%253A_Acetic_Acid_Fermentation
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[PDF] Industry Outlook 2022 - U.S. Apple Association
Aug 12, 2022 · Based on the most recent estimate from the U.S. Department of Agriculture (USDA), U.S. apple production for the 2022/23 crop year (CY) will be ...
[136]
Apple Cider - an overview | ScienceDirect Topics
Apple cider can be defined as a fermented alcoholic beverage made from apple juice. ... Apple beverages including cider can be sources of probiotics, due to which ...
[137]
1.6: Vinegar and Acetic Acid Fermentation - Chemistry LibreTexts
Jul 6, 2022 · Several species of acetic acid bacteria are used in industry for production of certain foods and chemicals. Commonly used feeds include apple ...
[138]
Dried Apple Grades and Standards - Agricultural Marketing Service
The prepared apple segments are properly dried to remove the greater portion of moisture to produce a semi-dry texture. The product may be sulfured sufficiently ...
[139]
Preparation of Semi-Solid Apple-Based Baby Food - ResearchGate
Aug 6, 2025 · To develop commercial semi-solid apple baby food, the physicochemical characteristics of apple puree in relation to different preparing ...
[140]
Apple pomace as food fortification ingredient: A systematic review ...
Sep 23, 2020 · The systematic review confirmed that the apple pomace can be successfully used as a fortification ingredient in plant food products, in meat products, and ...Missing: fertilizer | Show results with:fertilizer<|control11|><|separator|>
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https://pmc.ncbi.nlm.nih.gov/articles/PMC3551016/
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Conversion of apple pomace waste to ethanol at industrial relevant ...
Jun 30, 2016 · These results show that apple pomace is an excellent feedstock for producing ethanol that could be either used as biofuel or as beverage.
[143]
Apple pomace and culled apples - Feedipedia
As apple pomace represents about 25% of apples processed for juice or cider, it can be estimated that 1.4 million t of fresh apple pomace were produced in 2016.
[144]
Utilization of pomace from apple processing industries: a review - PMC
In large scale apple juice industry, about 75% of apple is utilized for juice and the remaining 25% is the by-product, apple pomace.
[145]
The Use of Apple Pomace as a Soil Amendment Enhances the ...
Apr 27, 2021 · The Use of Apple Pomace as a Soil Amendment Enhances the Activity of Soil Microorganisms and Nitrogen Transformations and Affects Crop Growth.
[146]
[PDF] Guide for Organic Crop Producers - Agricultural Marketing Service
If manures are composted according to USDA organic regulations, then they are considered compost, not manure, and may be applied without restrictions. If ...
[147]
Apples: Organic Production Guide - ATTRA – Sustainable Agriculture
This publication provides information on organic apple production from recent research and producer experience.
[148]
[PDF] A Grower's Guide to Organic Apples - Cornell eCommons
The ideal cover crop should be low growing, non-competitive ... It is a potent carcinogen and under NOP regulations, may not be used in organic crop production.
[149]
Soil microbial dynamics in organic (biodynamic) and integrated ...
Aug 9, 2025 · Soil samples were collected from three different apple orchards (each with biodynamic and integrated plots) during three dif-ferent seasons.Missing: credible | Show results with:credible
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https://www.freshfruitportal.com/news/2024/10/08/its-bio-notes-opportunities-for-organic-apples-in-europe/
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[PDF] U.S. Organic Production, Markets, Consumers, and Policy, 2000-2021
Mar 15, 2023 · Consumer demand for organically produced products has driven an expansion in U.S. organic produc- tion since 2000. The premiums paid by ...
[152]
IT'S BIO notes opportunities for organic apples in Europe
Oct 8, 2024 · The organic apple market in Italy currently accounts for 5-10% of total apple production, a relatively stable share over time, ...Missing: 2000s | Show results with:2000s
[153]
USDA Approves Genetically Engineered Apple Despite Health ...
Feb 13, 2015 · The Arctic Apple has been engineered to reduce polyphenol oxidase (PPO) enzymes responsible for browning in apple flesh after bruising.
[154]
USDA Has Approved GE Non-Browning Apples. Is the Public Ready ...
Mar 11, 2015 · Arctic apples have been engineered using a gene silencing technique called ribonucleic acid interference (RNAi), which is based on ...
[155]
USDA approves world's first GMO apples - The Ecologist
Feb 16, 2015 · The US Department of Agriculture has 'deregulated' two varieties of Arctic® apple, the world's first genetically engineered apple produced by Canadian ...
[156]
Okanagan Specialty Fruits & the Arctic® Apple
... United States of America (U.S.A.), and both were approved in 2015 (“Nonbrowning GM apple cleared,” n.d.). The Arctic® Fuji apple was approved for U.S.A. ...
[157]
Fourth non-browning apple, made in Summerland, gets FDA approval
Oct 8, 2024 · That means apples that are sliced into remain looking fresh, extending the shelf life of sliced Arctic apple products to as long as 28 days.Missing: 30 | Show results with:30
[158]
The truth behind genetically modified produce - Food News - Particle
Feb 1, 2018 · “Our engineered apples contain 11,000 micrograms [of anthocyanin] per apple. That's a 120-fold increase!” says Andrew, comparing the red-fleshed ...
[159]
Dietary Flavonoids from Modified Apple Reduce Inflammation ...
Studies in humans have shown that apple can increase antioxidant concentrations in plasma (37) and serum (38) and reduce DNA damage and oxidative stress levels ...
[160]
[PDF] Genetic Engineering of Apple for Resistance to Fire Blight
The ulti- mate solution to fire blight, other diseases, and insect pests, would be resistant vari- eties and rootstocks. However, conven- tional breeding of ...
[161]
Fire-blight resistant apples | ETH Zurich
Mar 13, 2014 · With the aid of so-called cis-genetic engineering, they transfered a resistance gene from a wild apple into the genome of a Gala apple. Tests in ...
[162]
The HcrVf2 gene from a wild apple confers scab resistance ... - PNAS
The reported data show that the HcrVf2 gene can confer resistance to apple scab when transformed in the scabsusceptible cv. Gala. Of the five transgenic lines, ...
[163]
[PDF] Apple (Malus x domestica) Events GD743 and GS784 - usda aphis
• Malus x soulardii (Soulard crab), a hybrid of ioensis x domestica is found from southern ... taxonomically with coronaria; the common name of the hybrid is big ...
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Arctic Apple Events GD743 and GS784 - Canada.ca
May 10, 2024 · Two varieties of Arctic apple were approved for growth and sale in Canada, the Arctic Golden Delicious and the Arctic Granny Smith.
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Recent Developments and Strategies for the Application of ... - NIH
Jun 30, 2022 · The last three field trials with genetically modified apples in the EU ended between 2019 and 2021. One field trial, which was carried out ...
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[PDF] The Regulation of GMOs in Europe and the United States.pdf
This paper seeks to place the divergent approaches of the European Union and United States toward the introduction and marketing of genetically modified ...
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https://foodsafety.institute/food-biotechnology/role-of-gm-crops-reducing-food-waste/
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Apples - MarketResearch.com
Jun 1, 2025 · The global market for Apples estimated at US$84.4 Billion in the year 2024, is expected to reach US$93.8 Billion by 2030, growing at a CAGR of ...
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Role of GM Crops in Reducing Food Waste - Food Safety Institute
Mar 14, 2024 · GM crops can address food waste through extended shelf life, enhanced disease and pest resistance, improved post-harvest characteristics, ...
[170]
HESPERIDES - Greek Goddess-Nymphs of the Evening & Sunsets
The three nymphs and their glowing, golden apples were regarded as the source of the golden light of sunset--a phenomena which celebrated the bridal of Zeus and ...
[171]
Hercules and the Apples of the Hesperides, 11th Labor
Jan 10, 2025 · These were no ordinary fruits; they were golden apples, a wedding gift from Gaia to Hera, said to grant immortality to those who consumed them.Missing: primary | Show results with:primary
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ATALANTA (Atalante) - Arcadian Heroine of Greek Mythology
When many had already perished, Melanion came to run for love of her, bringing golden apples from Aphrodite, and being pursued he threw them down, and she, ...
[173]
ERIS - Greek Goddess of Strife & Discord (Roman Discordia)
Eris is the Greek goddess of strife, discord, and rivalry, associated with war and the desire for bloodshed. She is also the sister of Ares.
[174]
How the Forbidden Fruit Became an Apple - TheTorah.com
Oct 20, 2022 · Many prominent Christian authors either implied or outright asserted that the forbidden fruit was a fig, including Augustine, Theodore and ...
[175]
How the Forbidden Fruit Became an Apple | Rutgers University
Feb 16, 2023 · Latin authors most commonly refer to the forbidden fruit as a pomum, a Latin word meaning "fruit" or "tree fruit." Not surprisingly, Old ...
[176]
Avalon in Literature and History - Vault
This is Geoffrey's Latin interpretation for the Welsh name Ynys Avallach, and he dubs Avalon the "isle of the apples," an association that will remain with ...Missing: primary | Show results with:primary
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The Bold Knight, the Apples of Youth, and the Water of Life
Once there was a Tsar who had three sons. In his garden grew an apple tree that bore special golden apples said to restore youth.
[178]
All Saints Day - Catholic Doors Ministry
That is why it is customary to celebrate with apple-bobbing and pumpkin carving and other harvest games. And that is why All Saints's Day is begun with a night ...<|separator|>
[179]
Why Do We Give Out Candy on Halloween? - Irish Myths
Oct 19, 2022 · We give out candy on Halloween because the ancient Celts made offerings of fruits and nuts on Samhain to appease the gods.
[180]
Samhain ~ Traditions with Apples - Sacred Mists Shoppe
Oct 19, 2017 · Apples and pomegranates serve as a gracious food offering and will aid in your spirits safe travel as they pass through the veil between the worldly realms.
[181]
Apple Tree | Tree Lore | Druidy - OBOD
In Celtic tradition, the Otherwordly Avalon was also known as the Avallach, the Isle of Apples, ruled by Fairy Queen, Morgan le Fay (Freeman, page 196). This is ...
[182]
Celtic Fairy Tales: Notes and References | Sacred Texts Archive
The fairy apple of gold occurs in Cormac Mac Art's visit to the Brug of ... Yeats' Irish Folk and Fairy Tales, 52-9. In the Hibernian Ta/es, "Mann o ...Iv. Horned Women · Xiii. The Wooing Of Olwen · Xxi. Beth Gellert
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Fruity Symbolism in China
Apples. Apples symbolize safety and peace in China because the Chinese word for apple, ping guo, shares the same sound as the word for safety, ping an. · Pears.Missing: folklore | Show results with:folklore
[184]
7 Lucky Fruits for Chinese New Year - Top China Travel
Chinese people believe that one can live a peaceful and harmonious life by eating apples. They also like to buy a box of apples as Chinese New Year gift for ...7 Lucky Fruits For Chinese... · 3. Apples (苹果 Píngguǒ)... · Cultural Tour During Chinese...
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Good Fortune Fruit: Lucky Picks for Lunar New Year - The FruitGuys
Jan 23, 2025 · An apple a day keeps misfortune away! To invite peace, safety, and calm, lay out a platter of apples during the holiday. Many people attribute ...
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https://historycooperative.org/idunn/
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Idunn - World History Encyclopedia
Sep 10, 2021 · Idunn (pronounced Ih-dune) is a fertility goddess in Norse mythology who holds the apples of eternal youth the gods rely on to remain young and healthy.
[188]
Idunn: The Norse Goddess of Youth, Rejuvenation, and… Apples
Idunn is a significant deity in Norse mythology. She is known as the goddess of youth, fertility, and the guardian of the golden apples of immortality.
[189]
Iðunn – Norse Goddess of the Apples - Wyrd Designs
Sep 25, 2022 · In Snorri's Prose Edda we find mentions to her in both Gylfaginning and Skáldskaparmál. In Gylfaginning we are briefly introduced to her, the ...
[190]
Apple of one’s eye
### Origin, Meaning, and Historical Usage of "Apple of One’s Eye"
[191]
'A few bad apples': Phrase describing rotten police officers used to ...
Jun 14, 2020 · "The rotten apple spoils his companion" appears in published work by Benjamin Franklin in 1736. That one eventually morphed to, "One bad apple ...
[192]
An Apple A Day Keeps The Doctor Away - Meaning & Origin Of The ...
What's the meaning and origin of the phrase 'An apple a day keeps the doctor away ... The February 1866 edition of Notes and Queries magazine includes this: “A ...
[193]
Idiom: One bad apple spoils the bunch (English) - Lyrics Translations
Sep 28, 2017 · Idiom: One bad apple spoils the bunch; Language: English; Idiomatic translations / equivalents: French, Spanish; Explained meaning: English, ...
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7 Japanese Idioms & Proverbs to Impress - Listen & Learn USA
Apr 17, 2014 · The Japanese equivalent of 'the apple doesn't fall far from the tree' is 'the fruit of a tree falls to its root'. It means that children often ...Missing: harmony | Show results with:harmony
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How the apple became an important symbol in culture, science and ...
Nov 22, 2023 · The idea that apples can represent some kind of a fall persists in the fairy tale of Snow White. When the queen hears the mirror tell her that ...
[196]
History & Symbolism of Snow White's Poison Apple
Through the power of Disney's storytelling, the most ordinary object—an apple—has been transformed into a potent emblem of temptation, jealousy, and deceit. As ...
[197]
The Handmaid's Tale: Symbols | SparkNotes
The red color of the costumes worn by the Handmaids symbolizes fertility, which is the caste's primary function.
[198]
The Handmaid's Tale – Stewed Apple and Macarons as a symbol of ...
Aug 10, 2019 · The foods the handmaids are given are symbols of fertility (apples, pears, oranges, eggs, chickens, bread – baking in the oven).
[199]
Paul Cézanne, The Basket of Apples - Smarthistory
It was the very fact that still life was so neglected that seems to have attracted Cézanne to it. So outmoded was the iconography (symbolic forms and references) ...
[200]
Paul Cézanne's Painted Apples
Jan 19, 2021 · It was with the humble apple that artist Paul Cézanne chose to challenge both his own method of painting but also the art world of Paris.
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This is Superhost 2024 – a year's worth of apples on display | M HKA
Sep 11, 2024 · The artists propose a new tradition of community apple tree planting in urban spaces, in order to foster collective engagement and the greening ...
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https://vegoutmag.com/news/nat-vegan-eating-habits-in-2025-heres-what-people-are-actually-putting-on-their-plates/
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Apple's Logo and Eden's Forbidden Fruit - The Gospel Coalition
Mar 4, 2021 · Apple's “fruit with a bite mark” iconography alludes to Eden and temptation, whether it was intended to or not, Brett McCracken writes.
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Unraveling the tale behind the Apple logo - CNN
Oct 7, 2011 · The logo on the back of your iPhone or Mac is a tribute to Alan Turing, the man who laid the foundations for the modern-day computer.Missing: cultural | Show results with:cultural
[206]
Why 5 A Day? - NHS
Evidence shows there are significant health benefits to getting at least 5 portions of a variety of fruit and vegetables every day.
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Vegan eating habits in 2025: Here's what people are actually putting ...
Jun 18, 2025 · Nearly half of Americans (47%) surveyed in late 2024 plan to eat more plant-based foods in 2025, and 34% are actively looking to reduce their ...
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Apple Trends: Data And Analytics - Tastewise
The fastest-growing consumer need associated with apples is for snacks, and vegan diets continue to dominate apple consumption. ... Food trends 2025 · Press ...<|separator|>
[209]
[PDF] Health Marketing: “An Apple a Day” Takes on New Meaning
To encourage consumers to eat more apples for their better health, USApple is sponsoring new apple nutrition research and promoting apple health benefits ...
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AI-generated green apple image - Facebook
Jul 19, 2025 · Red & green apple beautiful photo ✨⚡ #facebookpost #artlife #design #artist #artwork #AI #digitalart #aiart #DigitalDiviner #aiArtofIndia # ...<|control11|><|separator|>