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Caper

The caper, derived from the flower buds and young fruits of the perennial shrub Capparis spinosa L. (family Capparaceae), is a tangy, brined condiment widely used in Mediterranean and global cuisines, harvested from a drought-tolerant plant native to arid regions of the Mediterranean Basin, Western Asia, and North Africa.^[1]^[2] Capparis spinosa, commonly known as the caper bush, is a spiny, semi-deciduous shrub typically growing 1–3 meters tall, with trailing branches, leathery leaves, and large white to pinkish flowers that bloom from spring to autumn.^[1]^[2] It thrives in xerophytic environments, tolerating poor, rocky soils and extreme temperatures, which has facilitated its natural distribution across Saharo-Arabian and Mediterranean zones since prehistoric times.^[2]^[3] Archaeological evidence indicates capers were consumed as early as the Bronze Age in northern Syria, highlighting their longstanding role in human diets.^[3] In culinary applications, unopened flower buds—known as capers—are harvested by hand, sorted by size, and preserved in brine, vinegar, or salt to develop their characteristic piquant flavor from glucosinolates like glucocapparin.^[2]^[4] Larger caper berries (immature fruits) are also pickled and used in sauces, salads, fish dishes, and pasta, adding a briny, lemony note; notable examples include Italian caponata, French sauce gribiche, and pairings with smoked salmon.^[1]^[5] Commercial production centers in countries like Italy, Spain, Turkey, and Morocco, where the plant is cultivated on terraced slopes or gathered wild, yielding buds that are graded from non-pareil (smallest, most flavorful) to larger sizes.^[2]^[3] Beyond gastronomy, capers offer nutritional value, providing antioxidants such as quercetin and rutin, vitamins (B1, B3, B6, B9, and E), and minerals, while traditional medicine employs various plant parts for anti-inflammatory, diuretic, and antidiabetic effects, supported by phytochemical studies on its bioactive compounds.^[4]^[2] Cultivation challenges include sensitivity to frost and the labor-intensive harvest, but its resilience makes it valuable for sustainable agriculture in marginal lands.^[1]^[2]

Taxonomy and Morphology

Taxonomy

The caper plant is scientifically classified as Capparis spinosa L., belonging to the genus Capparis in the family Capparaceae, within the order Brassicales of the class Magnoliopsida in the kingdom Plantae.^[6]^[7] The name "caper" derives from the Latin capparis, which in turn comes from the ancient Greek kapparis, the classical term for the plant and its edible buds.^[8]^[9] Historically, Capparis spinosa has been placed in the family Capparaceae (formerly Capparidaceae), but taxonomic debates have centered on its close affinity to Brassicaceae, the mustard family, leading to proposals for merging Capparaceae into Brassicaceae sensu lato in earlier systems like APG II (2003).^[10] However, the APG IV classification (2016) maintains Capparaceae as a distinct family, emphasizing molecular evidence that separates it while confirming shared ancestry within Brassicales.^[11] Synonyms for C. spinosa include Capparis inermis Turra (referring to spineless forms) and Capparis orientalis Veill., reflecting regional morphological variations recognized in older nomenclature.^[12] Infraspecific classification of C. spinosa remains variable across revisions, with up to 11 subspecies historically proposed based on morphological differences such as spine presence, leaf shape, and growth habit, though recent treatments recognize fewer.^[13] Accepted subspecies include C. spinosa subsp. spinosa, characterized by spiny branches and prostrate to erect growth in Mediterranean lowlands; subsp. rupestris (Sm.) Nyman, which is typically spineless with pendulous branches adapted to rocky, rupestral habitats; subsp. cordifolia (Lam.) Jafri, distinguished by heart-shaped leaves and distribution in arid regions; subsp. himalayensis (Jafri) Fici, featuring smaller leaves and adaptation to high-altitude environments in Asia; subsp. parviflora (Boiss.) Fici, with reduced flower size and occurrence in the Middle East; subsp. nummularia (Decne.) Fici, noted for rounded, coin-like leaves in semi-desert areas; subsp. canescens (Coss.) Maire, covered in grayish hairs for xeric tolerance; subsp. aegyptia (Lam.) Maire, adapted to saline soils with elongated fruits; subsp. mariana (Duch.) Fici, limited to island endemism with compact habit; and subsp. sicula Fici, endemic to Sicily with finer spines.^[14]^[12] The taxonomy of the C. spinosa group remains under revision, with some entities formerly treated as subspecies now recognized as separate species based on molecular evidence (e.g., C. cartilaginea).^[15] These distinctions aid in understanding regional adaptations, though hybridization complicates boundaries.^[13] Phylogenetic research since 2010, using chloroplast DNA (cpDNA) and nuclear markers, has reinforced C. spinosa's position within Capparaceae and its sister relationship to Brassicaceae, with studies revealing Tethyan relict patterns and genetic divergence among subspecies driven by geographic isolation.^[16] For instance, genome assembly of C. spinosa var. herbacea (2022) highlighted conserved synteny with brassicalean relatives, supporting evolutionary ties to mustard-like glucosinolate biosynthesis pathways.^[17] Recent analyses (2024–2025), including studies on Mediterranean and Australian populations, have revealed genetic differentiation and supported DNA-based taxonomic revisions over purely morphological ones.^[18]^[19]

Morphology

Capparis spinosa is a thorny, perennial shrub or climber that typically reaches 1–3 meters in height, though its branches can extend up to 4 meters in length, forming a scrambling or sprawling habit with arching, often tortuous or zigzag stems that may be green, red, or yellow and sparsely haired.^[2] The plant is semi-evergreen to deciduous depending on climate, with a multi-branched structure adapted for growth in rocky or arid terrains.^[1] The leaves are simple, alternate, and fleshy, measuring 2–5 cm in length, with shapes ranging from ovate to rounded, elliptic, or obcordate; they feature a glossy, glabrous surface, prominent veins, and petioles 0–2 cm long, while stipules often develop into sharp, curved spines up to 6 mm.^[2]^[1] Flowers are large and showy, solitary, and somewhat zygomorphic, with a diameter of 4–5 cm; they consist of four white to pinkish-white sepals, four oblong to obovate petals, numerous (up to 100) long purple stamens with filaments reaching 5 cm, and a gynophore 3–6 mm long, topped by a single stigma.^[2] These hermaphroditic flowers bloom nocturnally during the hot, dry summer months and are short-lived, lasting 12–24 hours before wilting, primarily attracting insect pollinators.^[2]^[1] The fruit is an oblong to ellipsoidal berry, 2–3 cm long, initially green and turning reddish or purple when ripe, containing numerous small, reddish-brown seeds (approximately 150–160 per gram of fruit).^[2] The seeds feature a hard coat and an oily endosperm, with an oil content of about 27%, rich in unsaturated fatty acids such as oleic acid.^[20] The root system is extensive and deep, forming a robust taproot that can extend 6–10 meters, comprising up to 62.5% of the plant's total biomass and enabling access to deep soil moisture for survival in arid environments.^[4]^[2]

Distribution and Ecology

Geographic Distribution

The caper plant, Capparis spinosa L., is native to the Mediterranean Basin, encompassing southern Europe (including Spain, Italy, Greece, and the Balkans), North Africa (from Morocco to Egypt), and extending eastward to Western Asia (including Iran, the Arabian Peninsula, and parts of the Levant).^[2] Its range also includes isolated Atlantic islands such as the Canary Islands and Cape Verde off the African coast, as well as northeastern and eastern Africa through to Sudan and Somalia.^[21] In addition, the species is native to arid and semi-arid regions of Australia, particularly northern areas like the Kimberley, Pilbara, and Queensland, where the subspecies C. spinosa subsp. nummularia occurs.^[22] Introduced ranges of C. spinosa include the Americas, where it was brought by Spanish colonizers in the post-1500s period for culinary purposes, becoming established in arid zones of California, Mexico, and parts of South America such as Chile and Argentina.^[23]^[9] The plant has naturalized in some of these areas, particularly in dry coastal and inland regions, though evidence for pre-Columbian presence in the Americas remains unsubstantiated and is generally attributed to later human introduction.^[23] It has also been introduced to parts of Asia beyond its native range, such as India and Pakistan, and to other Pacific islands, often escaping cultivation to form feral populations in suitable arid environments.^[24] The current global extent of C. spinosa includes an estimated area of occupancy exceeding 2,000 km² (approximately 200,000 hectares), primarily under wild or semi-wild conditions across its native and introduced ranges.^[25] Commercial production relies heavily on wild harvesting, with an annual global output of around 10,000–20,000 tonnes as of 2018, led by Mediterranean countries: Turkey (3,500–4,500 tonnes), Spain (500–1,000 tonnes), and Italy (approximately 1,500 tonnes).^[25]^[26] Cultivated areas remain limited, totaling about 3,600 hectares in intensive production zones of Spain (2,600 ha) and Italy (1,000 ha).^[27] C. spinosa predominantly occupies Mediterranean shrubland biomes, such as maquis and garrigue in Europe and North Africa, as well as semi-desert and steppe ecoregions in Western Asia and Australia, where it thrives in rocky, drought-prone landscapes.^[24]^[2] These habitats support its xerophytic adaptations, with populations often concentrated along coastal dunes, cliffs, and wadis.^[25]

Habitat and Environmental Requirements

Capparis spinosa, commonly known as the caper bush, thrives in Mediterranean-type climates characterized by hot, dry summers and mild, wet winters, making it well-suited to arid and semi-arid environments across its native range.^[2] It is hardy in USDA zones 8 to 10, tolerating temperatures as low as -10°C and as high as over 40°C, which allows it to persist in regions with significant seasonal temperature fluctuations.^[28]^[1] This thermal resilience contributes to its role in xerophytic plant communities, where it occupies niches on rocky slopes, coastal areas, and degraded soils, helping to stabilize landscapes and reduce erosion. The species is assessed as Least Concern by the IUCN due to its wide distribution and lack of major threats.^[29]^[30] The plant requires well-drained, rocky or sandy soils with low fertility, preferring calcareous substrates and a pH range of 6.5 to 8.5 to support optimal growth.^[28]^[31] It exhibits high tolerance for saline and nutrient-poor conditions, which enables it to colonize harsh, low-organic-matter environments without supplemental fertilization.^[2] Regarding water needs, C. spinosa is highly drought-resistant, relying on an extensive root system for efficient water uptake; it requires an annual rainfall of 300 to 500 mm, primarily during winter and spring, with minimal irrigation once established.^[21]^[12] Full sun exposure is essential, with the plant demanding at least 6-8 hours of direct sunlight daily to avoid shade intolerance and maintain vigorous growth.^[31] It can grow from sea level up to 1,500 meters in elevation, adapting to montane Mediterranean habitats while favoring lower altitudes in coastal and inland arid zones.^[32] Ecologically, C. spinosa integrates into xerophytic assemblages, where it interacts with pollinators such as bees and nocturnal moths that visit its white, fragrant flowers, and seed dispersers including birds and lizards that consume and transport its fleshy fruits.^[33]^[34] These associations enhance its propagation in natural settings and support biodiversity in dryland ecosystems.^[2]

Cultivation

Propagation and Establishment

Caper plants (Capparis spinosa) are primarily propagated by seeds or vegetatively through cuttings and grafting, with establishment involving careful site preparation to ensure long-term survival in arid conditions.^[35] Seed propagation requires overcoming the hard seed coat and physiological dormancy, which limit natural germination rates to 1-10% without treatment. Scarification using sulfuric acid for 40 minutes, followed by soaking in gibberellic acid (GA3) at 400 ppm for 2 hours, can achieve germination rates up to 62%. Germination typically occurs in 2-4 weeks at temperatures of 20-25°C in a well-draining medium.^[36]^[37]^[38] Vegetative propagation is preferred for maintaining desirable traits and is achieved through semi-hardwood cuttings of 10-15 cm length, rooted in a 1:1 sand-perlite mix under high humidity, yielding rooting success rates of 50-70% with auxin treatments. Grafting onto resistant rootstocks, such as using bark grafting techniques, provides disease resistance and has success rates around 60% in Mediterranean cultivation systems.^[39]^[2] For establishment, seedlings or cuttings are planted at densities of 1,000-4,000 plants per hectare, with row spacing of 2-3 meters to accommodate the shrub's spreading habit and allow mechanical access in commercial settings. Initial fertilization uses low-nitrogen NPK formulations (e.g., 16-16-16 at 2-3 applications in the first year) to promote root development without excessive vegetative growth.^[40]^[27]^[41] Challenges in propagation include inherently low germination in wild-collected seeds due to dormancy, addressed by GA3 applications in studies from 2015-2023 showing up to 60% improvement. Recent advances in tissue culture micropropagation enable production of uniform, disease-free cultivars, overcoming genetic variability issues and achieving multiplication rates of 5-10 fold per cycle. Recent research (as of 2025) shows that inoculation with arbuscular mycorrhizal fungi (AMF) enhances plant growth, root development, and nutrient levels, supporting sustainable cultivation.^[38]^[42]^[35]^[43]

Management and Harvest

Orchard management of Capparis spinosa focuses on practices that promote bushy growth and maximize flower bud production while adapting to the plant's drought tolerance in arid or semi-arid environments. Pruning is typically performed in late winter, removing dead wood, water sprouts, and overcrowded branches to stimulate new growth on one-year-old wood where buds form, with heavy pruning essential for maintaining plant vigor and facilitating access for harvest.^[44] Irrigation is minimal after establishment, relying on drip systems to deliver 200-600 mm annually, with young plants requiring more consistent moisture in the first two years before transitioning to drought-resistant conditions.^[44]^[41] Pest management employs integrated pest management (IPM) strategies, targeting key threats like the caper moth (Cydia capparidana), which bores into buds causing deformation, through yellow sticky traps for monitoring and mass trapping, combined with cultural practices such as post-harvest ploughing to destroy infested material.^[45] Aphids, such as Brevicoryne brassicae, are controlled with mineral oil sprays during severe infestations to prevent sap feeding and potential virus transmission, emphasizing monitoring to minimize chemical interventions.^[45] Fertilization requirements are low due to the plant's adaptation to nutrient-poor soils, with minimal applications of balanced NPK (e.g., 10-5-5 or 16-16-16) in spring and summer for young plants to support establishment without promoting excessive vegetative growth that could reduce bud quality.^[41]^[46] Emphasis is placed on potassium-rich amendments to enhance bud firmness and flavor intensity, while sustainable systems incorporate organic compost annually to maintain soil health without over-fertilization.^[44] Harvesting occurs manually by hand-picking unopened flower buds (1-2 cm in diameter, dark green and firm) daily during the peak season from May to July in Mediterranean climates, ensuring buds are collected in the morning to avoid opening and maintain quality.^[1]^[44] Caper berries, the matured fruits, are gathered later in the season (July to September) when firm but unripe. Yields typically reach 1-3 kg of buds per mature plant annually after three years, with higher yields up to 10 kg possible in optimal conditions; full productivity achieved in 3-4 years and plantations productive for 20-30 years.^[44]^[27]^[47] Post-harvest processing begins with immediate sorting by size, where premium grades like non-pareil (under 7 mm) and surfine (7-8 mm) command higher value for their tenderness and intense flavor.^[1] Buds are then preserved through dry salting (up to 20% NaCl) or brining in vinegar-salt solutions, which cures the product over several days to weeks, enhancing flavor by hydrolyzing glucosinolates and reducing inherent bitterness while preventing microbial spoilage.^[44]^[48] Recent advancements include breeding programs in Italy and Spain developing climate-resilient cultivars through transcriptome analysis and marker development to improve drought tolerance and yield stability amid changing environmental conditions.^[49]

Uses

Culinary Applications

Capers, the unopened flower buds of Capparis spinosa, and caper berries, the mature fruits of the same plant, are primarily consumed after pickling to mitigate their inherent bitterness and enhance palatability.^[1] Preservation methods include dry-salting, where buds are layered in sea salt to draw out moisture and develop flavor over weeks, or brining in vinegar solutions, often with added salt, for a quicker process that yields a shelf-stable product.^[50] These techniques trigger the enzymatic breakdown of glucocapparin, a mustard oil glucoside, releasing methyl isothiocyanate and other compounds that impart the characteristic briny, tangy profile.^[27] The flavor of capers is pungent and multifaceted, featuring lemony acidity, peppery sharpness, and a subtle floral undertone, with the intensity varying by bud size due to higher concentrations of volatile oils in smaller specimens.^[27] Capers are graded by diameter, from the tiniest nonpareils (under 7 mm), prized for their concentrated, delicate zest, to larger capucines (8-9 mm), which offer a bolder, more robust taste suitable for heartier applications. Recent food chemistry analyses confirm that isothiocyanates, such as methyl isothiocyanate, dominate the aroma profile, contributing to the buds' sensory appeal through volatile sulfur compounds detected via gas chromatography-mass spectrometry.^[51] In Mediterranean cuisine, capers serve as an essential condiment, adding brightness and salinity to dishes like spaghetti alla puttanesca, where they balance the heat of chili and richness of tomatoes and anchovies, or chicken piccata, enhancing the sauce's lemon-butter base.^[52] They pair seamlessly with seafood such as grilled fish or salmon, lean meats like veal, and fresh salads, where their tang cuts through oils and vinaigrettes; in a pinch, chopped green olives can substitute for a similar briny effect.^[50] Globally, variations highlight regional adaptations: in Italian cooking, capers feature in Sicilian pesto pantesco, blending with tomatoes, almonds, and basil for pasta sauces; Greek preparations incorporate them into yogurt-based dips akin to tzatziki variations, providing a salty counterpoint to cucumber and dill; and Middle Eastern recipes use them in tahini sauces, mingling with sesame paste for vegetable or falafel accompaniments. Caper berries, larger and milder, often garnish charcuterie or cocktails, while creative applications include caper butter—made by mashing rinsed capers into softened butter with lemon zest for spreading on bread or fish—and infusions in olive oil or vinegar for dressings.^[53]

Nutritional and Health Benefits

Capers are low in calories, providing approximately 23 kcal per 100 g serving, primarily due to their minimal macronutrient content, including 2.36 g of protein, 0.86 g of fat, and 4.89 g of carbohydrates, of which 3.2 g is dietary fiber.^[54] The high sodium content, around 2,348 mg per 100 g, primarily results from the brining process used in preservation.^[54] In terms of micronutrients, capers offer notable amounts of vitamin K at 24.6 µg per 100 g (about 21% of the daily value), along with smaller quantities of vitamin A (7 µg RAE) and iron (1.67 mg).^[54] They also contain other minerals such as magnesium (33 mg) and calcium (40 mg) per 100 g, contributing to their role as a nutrient-dense condiment. The complete nutrient profile, as documented by the USDA FoodData Central database through 2024, underscores capers' value in providing antioxidants alongside these vitamins and minerals.^[54] Capers are rich in bioactive compounds, including glucosinolates such as glucocapparin, which are characteristic of the Capparaceae family.^[4] Key flavonoids include kaempferol (approximately 131 mg per 100 g) and quercetin (around 173 mg per 100 g) in canned capers.^[55] These compounds support substantial antioxidant capacity.^[56] The health benefits of capers stem largely from their bioactive compounds, particularly quercetin, which demonstrates anti-inflammatory effects by modulating cytokines such as TNF-α and IL-1β, as evidenced in rodent studies and reviews spanning 2015 to 2023.^[4]^[57] Quercetin and other flavonoids may also offer potential anti-diabetic support by reducing fasting blood glucose levels, with caper extracts showing up to 20% reductions in diabetic models at doses equivalent to 400 mg/kg.^[4] Cardiovascular benefits include improved lipid profiles, potentially lowering cholesterol through polyphenol activity.^[4] Additionally, capers have a low glycemic index, estimated at near 0 due to their low net carbohydrate content (1.69 g per 100 g), making them suitable for blood sugar management.^[58] Despite these advantages, capers pose risks for certain individuals, primarily due to their elevated sodium levels, which may exacerbate hypertension or related conditions in sensitive populations.^[54] Allergenicity is rare but documented, with cases of food allergy and contact dermatitis reported, potentially linked to cross-reactivity with related plant families like Brassicaceae.^[59]

Other Uses

Beyond its culinary roles, the caper plant (Capparis spinosa) finds applications in medicine, cosmetics, industry, and horticulture. Traditionally, decoctions of its root bark or teas from young shoots have been used to alleviate rheumatism and joint pain, while buds or leaves prepared as herbal teas serve as remedies for diabetes management in regions like Morocco and China.^[60]^[4] Modern research highlights the antimicrobial properties of caper extracts, with ethanol extracts from fruits demonstrating moderate activity against methicillin-sensitive and methicillin-resistant Staphylococcus aureus, achieving minimum inhibitory concentrations of 21-32 mg/mL.^[61] In cosmetics and pharmaceuticals, rutin—a flavonoid abundant in caper leaves (up to 62 mg/100 g)—is extracted for use in anti-aging creams due to its antioxidant effects that protect skin from oxidative stress and support collagen production.^[4]^[62] Fruit extracts, including those with characteristic aromatic compounds, contribute to perfumes and skincare formulations for their soothing and fragrant properties.^[63] Studies, primarily in animal models, indicate that leaf extracts accelerate wound healing by reducing inflammation and promoting tissue repair in burn and incisional wounds.^[64] Industrially, caper seeds, containing 27-38% oil rich in unsaturated fatty acids like linoleic acid, show promise as a feedstock for biodiesel production, particularly in arid regions where the plant thrives.^[65] The plant's thorny stems enable its use as a natural boundary marker or living fence for property delineation and livestock containment.^[12] Its extensive root system and biomass contribute to erosion control on slopes and degraded soils, stabilizing terrain in semi-arid environments.^[66] As an ornamental, C. spinosa is cultivated in arid and rocky gardens for its drought tolerance, glossy foliage, and showy white-pink flowers with prominent purple stamens, often as a sprawling shrub or groundcover in Mediterranean-style landscapes.^[28] Emerging research post-2022 emphasizes its biofuel potential, with the 2022 genome sequencing of C. spinosa var. herbacea providing a foundation for genetic engineering to enhance seed oil yield and adaptability for sustainable biodiesel in marginal lands.^[67]^[68] As of 2025, studies continue to explore caper-derived biofuels for arid agriculture integration.^[69]

History and Cultural Significance

Historical Development

Archaeological evidence indicates remains of Capparis spinosa from Mesolithic sites in the Mediterranean region, dating to approximately 9500–9000 years before present, suggesting possible early foraging in arid environments, though confirmed consumption is attested from the Bronze Age onward.^[70] Pollen records from the Neolithic era (around 8000 BCE) further attest to its presence in local flora across southern Europe and the Near East.^[70] In ancient civilizations, capers appear in Sumerian texts from around 2700 BCE, where they are referenced in the Epic of Gilgamesh as a gathered wild plant, likely valued for its pungent buds in early culinary or preservative practices.^[71] The Hebrew Bible alludes to the caper berry in Ecclesiastes 12:5, symbolizing the waning vitality of old age, though interpretations debate its exact identification and debated references in other passages like Job highlight its cultural significance in ancient Near Eastern lore.^[72] By the 1st century CE, Greek physician Dioscorides documented capers extensively in De Materia Medica for their medicinal properties, including as a diuretic, anti-inflammatory, and remedy for digestive ailments, establishing their role in classical pharmacology.^[73] During the Roman era, Pliny the Elder described caper cultivation in Natural History, noting its propagation in rocky soils and trade value, with evidence of export along routes extending to the Silk Road, as seed remains from 2800-year-old tombs in northwestern China indicate transcontinental exchange by the 8th century BCE.^[27]^[70] In the medieval period, Islamic scholar Avicenna (Ibn Sina) expanded on these uses in his Canon of Medicine (11th century), detailing capers' pharmacological benefits for treating hypertension, liver disorders, and as an expectorant, influencing Eurasian medical traditions.^[74] Post-1500s European herbals, such as those by John Gerard in 1633, reinforced capers' medicinal and culinary roles, recommending them for rheumatism and as a condiment, while wild collection predominated until the 19th century.^[75] Organized domestication emerged in the 20th century, with specialized farming beginning around 1970 in Italy and Spain, marking the shift from foraging to commercial production in Mediterranean agriculture.^[2]

Cultural and Economic Importance

Capers hold symbolic importance in Mediterranean folklore, where the caper bush (Capparis spinosa) is associated with endurance and resilience due to its ability to thrive in harsh, arid environments.^[76] This symbolism extends to themes of fertility and protection in some traditional narratives, reflecting the plant's persistent growth in rocky terrains as a metaphor for life's tenacity.^[76] The plant features prominently in cultural festivals, notably the Sagra del Cappero on the island of Pantelleria, Italy, an annual event celebrating the harvest with local cuisine, music, and traditions that highlight capers' role in Sicilian heritage.^[77] Economically, the global caper market is valued at approximately $50-70 million annually as of 2023, driven by demand for the flower buds in culinary applications.^[78] Major exporters include Turkey, accounting for about 40% of production, and Spain with around 30%, alongside emerging players like China and Morocco.^[79] Sustainable practices are emphasized through certifications such as the EU Protected Designation of Origin (PDO) for Sicilian capers from Pantelleria, which ensures traditional wild harvesting methods and quality standards to support local economies.^[77] Conservation efforts underscore capers' vulnerability despite their widespread distribution. Capparis spinosa is classified as Least Concern by the IUCN, but faces local threats from overharvesting for commercial use and climate change impacts like increased drought and temperature shifts in Mediterranean habitats.^[32] In Spain, protected areas such as national parks in Andalusia safeguard caper populations, integrating the plant into biodiversity conservation strategies to mitigate overgrazing and habitat loss.^[25] Socially, caper production provides essential rural employment in North Africa, particularly in Morocco and Tunisia, where seasonal harvesting supports thousands of workers in arid regions with limited agricultural options.^[80] In Italy, caper cultivation fosters agritourism, with farms on Pantelleria offering visitor experiences like guided harvests and cooking classes, boosting local incomes and preserving cultural practices.^[81]

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Capparis spinosa (Capers) - Gardenia.net
A sun-loving, drought-tolerant, evergreen shrub that also adds beauty to gardens with its glossy green leaves and dramatic flowers.Missing: morphology | Show results with:morphology
[27]
Impacts of climate change on Capparis spinosa L. based on ...
Oct 16, 2018 · The species grows mainly in arid and semi-arid environments in tropical regions, and thus can be used to combat desertification and soil erosion ...
[28]
Capparis spinosa - Plant Finder - Missouri Botanical Garden
Family: Capparaceae. Native Range: Southern Eurasia ... The genus name Capparis comes from kapparis which is the ancient Greek name for the caper bush.
[29]
(PDF) Capparis spinosa - ResearchGate
Apr 29, 2020 · It is a species of Saharo-Arabian and Mediterranean origins, which is characterized by food, ecological, socio-economic and ornamental ...Missing: hectares | Show results with:hectares
[30]
Intrafloral patterns of color and scent in Capparis spinosa L. and the ...
Nov 13, 2022 · Ecological research has indicated a mixed pollination system including predominantly bees and, less often, nocturnal hawkmoths (Dafni et al ...Missing: dispersers | Show results with:dispersers
[31]
The effect of lizards on the dispersal and germination of Capparis ...
Feb 26, 2021 · spinosa seeds enhanced the permeability of the seed coats, which promoted fast water uptake, broke seed dormancy and increased the seed ...
[32]
(PDF) In Vitro Propagation of Caper (Capparis spinosa L.): A Review
Oct 13, 2025 · Conventional propagation has been successfully achieved by seeds, stem cuttings, and by grafting. In vitro propagation has been achieved through ...
[33]
(PDF) Germination studies of Capparis spinosa L. - ResearchGate
Aug 6, 2025 · Highest germination of 62% was obtained when seeds were pretreated with H2SO4 (40 min.), followed by 400 ppm of gibberellic acid soaking (2 hr).
[34]
https://pmc.ncbi.nlm.nih.gov/articles/PMC7909692/
[35]
Water Uptake and Germination of Caper (Capparis spinosa L.) Seeds
Caper seeds show physiological dormancy, requiring gibberellic acid for high germination. Water uptake follows a triphasic model, and they lack a water- ...
[36]
[PDF] type of cuttings and rooting medium of (capparis spinosa l) cutting
The treatments included medium culture at two levels (Sand or sand and mixed 1/1 sand and Perlite), cutting at two levels (semi-hardwood cuttings, softwood.
[37]
Caper Bush Technical Information - Richters Herbs
Jan 26, 2025 · Plants are transplanted in winter and spring, 2-2.5 metres apart, about 2,000 plants per hectare. Yields reach maximum in 3-4 years, about 1-3 ...
[38]
Grow and Use Capers: Learn About the Caper Bush | Gardener's Path
Jul 5, 2019 · Native to the Mediterranean, C. spinosa plants require dry heat and lots of sun to grow. They will not survive temperatures below 18°F. In their ...Missing: elevation rainfall
[39]
Can the Caper (Capparis spinosa L.) Still Be Considered a Difficult ...
Sep 16, 2021 · As a perennial xerophytic shrub, characterized by plesiomorphic features, the caper (Capparis spinosa L.) is naturally spread throughout the ...1. Introduction · 2. Seed Propagation · 4. In Vitro PropagationMissing: Americas | Show results with:Americas
[40]
[PDF] Capparis spinosa L, An important medicinal plant from Sistan ... - SID
Planting, Irrigation, Pruning and Fertilization. For commercial ... Clonal propagation and medium-term conservation of Capparis spinosa: A medicinal plant.
[41]
https://gardenerspath.com/plants/herbs/how-to-grow-capers/
[42]
A Complete Caper Berry Farming Guide - Organic Farm
The branches develop spines on some varieties, so wear gloves when working around plants. Spineless varieties like 'Eureka' or Capparis spinosa var. inermis are ...
[43]
The Current State of Knowledge about the Biological Activity of ... - NIH
All Capparis species are sources of sulfur-rich metabolites called glucosinolates, which are derived from amino acids. They can be divided into three groups ...
[44]
Transcriptome analysis and codominant markers development in ...
Jul 18, 2019 · Caper (Capparis spinosa L.) is a xerophilous crop showing a remarkable adaptability to harsh environments, and with promising potentialities for ...<|control11|><|separator|>
[45]
Some physical and mechanical properties of caper - ResearchGate
Dec 13, 2020 · ... Capparis spinosa, the caper bush, is a perennial winter-deciduous ... The results obtained through this study can be used in designing harvest ...
[46]
All About Capers, the Powerhouse Pantry Staple - Serious Eats
Like olives, both capers and caper berries are most palatable when cured—curing releases mustard oil from the bud, which gives it its characteristic zing. Here ...<|separator|>
[47]
(PDF) Volatile Profile in Different Aerial Parts of Two Caper Cultivars ...
Feb 2, 2021 · Capparis spinosa L. is cultivated in the Mediterranean basin. In Spain, flower buds, unripe fruits, and tender shoots, either pickled or ...
[48]
15 Great Capers Recipes - A Couple Cooks
30 minFeb 10, 2021 · Capers are used in classic recipes like French-style salmon, Italian pasta puttanesca, and modern dishes like vegan Caesar salad. Capers add a ...<|separator|>
[49]
Buttery Lemon Caper Sauce Recipe - Allrecipes
Rating 4.6 (57) · 15 minMar 25, 2025 · This simple lemon caper sauce is made with butter, capers, fresh savory, and lots of lemon juice for the perfect accompaniment to sautéed ...
[50]
https://www.seriouseats.com/what-to-do-with-capers-caper-berries
[51]
[PDF] USDA Database for the Flavonoid Content of Selected Foods ...
... Capers, canned (Capparis spinosa). Flavonols. Kaempferol. 131.34. 20. 12.13. 59.49. 247.97. B. 93, 120. Quercetin. 172.55. 20. 26.49. 45.05. 519.85. B. 93, 120.
[52]
Nutraceutical Value of Pantelleria Capers (Capparis spinosa L.)
Jul 11, 2019 · Person's coefficient indicated a high correlation between total phenolic content and anti-DPPH radical capacity (R2 = 0.665), as well as between ...
[53]
Capers Nutrition facts and Health benefits
In fact, they are the largest sources of rutin; 100 grams contain 332 mg of this compound. Also, Caper is also very rich in quercetin (180 mg/100 g), second ...
[54]
Recent Advances in Potential Health Benefits of Quercetin - PMC
Anti-Inflammatory. It has been demonstrated that quercetin has strong anti-inflammatory effects and a lengthy half-life. Quercetin and galangin, two flavanols, ...
[55]
Capers, canned: Glycemic index, Glycemic Load and Nutrition facts
May 17, 2023 · Glycemic Index. Capers, canned is likely to have a low glycemic index, as estimated by the AI Glycemic Index Estimator.
[56]
Food allergy to caper (Capparis spinosa) - PubMed
Food allergy to caper (Capparis spinosa)
[57]
Capparis spinosa L. as a potential source of nutrition and its health ...
May 30, 2023 · This plant has also been used traditionally to treat various diseases, such as rheumatism, gout, hypertension, diabetes, malaria, typhoid ...
[58]
Caper (Capparis spinosa L.): An Updated Review on Its ... - Frontiers
Cluster 2 is colored in green and represents the antioxidant activity of Caper and its correlation with compounds, especially quercetin and kaempferol.<|control11|><|separator|>
[59]
Investigation of Rutin Content in Different Plant Parts of Wild Caper ...
Aug 6, 2025 · Rutin was found in large amounts in leaves (62 mg/100 g) followed by flowers (44 mg/ 100 g) and fruits (6 mg/100 g). ... Caper (Capparis spinosa ...
[60]
capparis spinosa fruit extract, 89958-23-6
Cosmetic Uses: skin conditioning ; Odor · characteristic ; Use · Genuine gemmo-cosmetic, oily active extracted from caper buds, Gatuline ; Odor · characteristic ; Use ...
[61]
The Topical Effect of Cappariss spinosa Extract on Burn Wound ...
This study indicated that Capparis spinosa leaves extract has potential effect to accelerate wound healing and decrease inflammation in the treated rats.
[62]
Glucosinolates and fatty acid, sterol, and tocopherol composition of ...
The oil content of the seeds ranged from 27.3 to 37.6 g/100 g (C. spinosa) and from 14.6 to 38.0 g/100 g (C. ovata). The dominating fatty acid of both species ...Missing: percentage | Show results with:percentage
[63]
(PDF) Eco-physiology of capparis spinosa l. : A plant suitable for ...
Aug 10, 2025 · The long roots and wide ecological amplitude allow it to withstand harsh environments.
[64]
The Capparis spinosa var. herbacea genome provides ... - PubMed
The Capparis spinosa var. herbacea genome provides the first genomic instrument for a diversity and evolution study of the Capparaceae family.
[65]
https://pubmed.ncbi.nlm.nih.gov/16131121/
[66]
The discovery of Capparis spinosa L. (Capparidaceae) in the ...
Remains of Capparis spinosa were unearthed in archaeological sites as early as the lower Mesolithic (9500–9000 b. p.)Missing: prehistoric | Show results with:prehistoric
[67]
Capparis species: A potential source of bioactives and high-value ...
These plants were used first time about 2000 years BC by Sumerians (Aghel et al., 2010, Romeo et al., 2007). The roots, flowers, and fruits of these plants ...
[68]
A Biblical Spice Rack
Capers (Capparis spinosa L.) “The almond tree blossoms, the grasshopper drags itself along, and the caper berry shall fail; because man goes to his eternal ...<|separator|>
[69]
Capparis species: A potential source of bioactives and high-value ...
Capparis species, also known as Caper plants, are recognized as a potential source of valuable nutrients and biochemical compounds with physiological ...
[70]
Phenolic Compounds and Vitamin Antioxidants of Caper (Capparis ...
Aug 6, 2025 · In (1020) IbnSina (Avicenna) in canon medicine have been used plant Capparis Spinosa for treatment of hypertension, fruits and puds of the ...
[71]
What could be a more useful garden addition than a caper bush ...
Apr 18, 1991 · But of course, the caper's contemporary value lies in its use as a condiment. Pickled capers have been known in Western Europe since AD 1500.
[72]
The Intriguing World of Caper Bush: Symbolism and Significance
May 31, 2024 · Cultural Symbolism In Mediterranean cultures, the caper bush carries deep symbolic meaning. It is often associated with endurance, an attribute ...Missing: fertility protection festivals Pantelleria Jewish Christian
[73]
Capers - SmellyBlog
Jun 15, 2017 · Legend has it that the caper plants originate in the Western Wall, where the little folded letters containing the prayers, dreams and promises ...Missing: cultural significance protection festivals Pantelleria religious Christian
[74]
Pantelleria capers: tradition, history and tasty recipes - Sicilian Food
May 22, 2025 · The Pantelleria caper is the flower bud of a particular plant cultivated on the island of Pantelleria, in the province of Trapani, and since 1996 recognized as ...Missing: fertility festivals religious Jewish Christian
[75]
2024 Caper global market overview today - Tridge
Discover price data of wholesale and farmgate prices of Caper. See the price changes and trends of your interested countries or varieties.Missing: hectares FAO Turkey
[76]
Caper global exports and top exporters 2024 - Tridge
In 2023, China was the top exporting country of Caper, followed by Belgium, Spain, Mexico, Netherlands, Poland, Ecuador, Egypt, France, Turkiye. This ranking is ...Missing: Turkey PDO Sicilian sustainable
[77]
[PDF] DIVERSIFYING RURAL ACTIVITY - CIHEAM Montpellier
Designations of quality and origin affect a territory on several levels: the effects may be economic (essentially direct or indirect employment), social ( ...<|separator|>
[78]
Val Talks Capers (pt.1) - Zingerman's Community of Businesses
Jun 20, 2013 · This essay recounts her visit to the island of Pantelleria, source of some of the best capers in the world.Missing: cultural significance folklore fertility festivals religious uses Jewish Christian
[79]
Exploratory study into climate change adaptation plans and ...
Nov 7, 2024 · This exploratory study highlights the different climate change adaptations that Fairtrade producer organisations are identifying and implementing as part of ...Missing: capers 2020-2025