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Juglans
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Juglans
Juglans major
Morton Arboretum acc. 614-47*1
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Rosids
Order: Fagales
Family: Juglandaceae
Subfamily: Juglandoideae
Tribe: Juglandeae
Subtribe: Juglandinae
Genus: Juglans
L.
Type species
Juglans regia
Species

See text

Native ranges of Juglans spp.
Synonyms[1]

Wallia Alef

Walnut trees are any species of tree in the plant genus Juglans, the type genus of the family Juglandaceae, the seeds of which are referred to as walnuts. All species are deciduous trees, 10–40 metres (33–131 ft) tall, with pinnate leaves 200–900 millimetres (7.9–35.4 in), with 5–25 leaflets; the shoots have chambered pith, a character shared with the wingnuts (Pterocarya), but not the hickories (Carya) in the same family.

The 21 species in the genus range across the north temperate Old World from southeast Europe east to Japan, and more widely in the New World from southeast Canada west to California and south to Argentina.

Edible walnuts, which are consumed worldwide, are usually harvested from cultivated varieties of the species Juglans regia. China produces half of the world total of walnuts.

Etymology

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The common name walnut derives from Old English wealhhnutu, literally 'foreign nut' (from wealh 'foreign' + hnutu 'nut'),[2] because it was introduced from Gaul and Italy.[3] The Latin name for the walnut was nux Gallica, "Gallic nut". The name Juglans was apparently conjured by Linnaeus himself,[4] replacing the prior Nux,[disputeddiscuss] by combining Ju from Jupiter with the Latin glans meaning 'acorn',[5] referring to the association of the plant with Jupiter by the Romans.

Folklore

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Tradition has it that a walnut tree should be beaten. This would have the benefit of removing dead wood and stimulating shoot formation.[6]

Production

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Walnut (in shell) production – 2017
Country (tonnes)
 China 1,925,403
 United States 571,526
 Iran 349,192
 Turkey 210,000
 Mexico 147,198
 Ukraine 108,660
World 3,829,626
Source: FAOSTAT of the United Nations[7]

In 2017, world production of walnuts (in shell) was 3.8 million tonnes, led by China with producing half of the world total (table). Other major producers were the United States (15%) and Iran (9%).

Cultivation and uses

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Walnut is one of the main ingredients of Baklava and Turkish cuisine.

The two most commercially important species are J. regia for timber and nuts, and J. nigra for timber. Both species have similar cultivation requirements and are widely grown in temperate zones.

Walnuts are light-demanding species that benefit from protection from wind. Walnuts are also very hardy against drought.

Interplanting walnut plantations with a nitrogen fixing plant, such as Elaeagnus × ebbingei or Elaeagnus umbellata, and various Alnus species, results in a 30% increase in tree height and girth (Hemery 2001).

When grown for nuts, care must be taken to select cultivars that are compatible for pollination purposes; although some cultivars are marketed as "self fertile", they will generally fruit better with a different pollination partner. Many different cultivars are available for growers, and offer different growth habits, flowering and leafing, kernel flavours and shell thicknesses. A key trait for more northerly latitudes of North America and Europe is phenology, with ‘late flushing’ being particularly important to avoid frost damage in spring. Some cultivars have been developed for novel ‘hedge’ production systems developed in Europe and would not suit more traditional orchard systems.

Flowers

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The leaves and blossoms of the walnut tree normally appear in spring. The male cylindrical catkins are developed from leafless shoots from the past year; they are about 10 cm (3.9 in) in length and have a large number of little flowers. Female flowers appear in a cluster at the peak of the current year’s leafy shoots.[8]

Fruit

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The fruits of the walnut are a type of accessory fruit known as a pseudodrupe (or drupe-like nut), the outer covering of the fruit is an involucre - in a drupe the covering would be derived from the carpel.[9]

Nuts and kernels

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Persian walnut (Juglans regia) seeds

The nut kernels of all the species are edible, but the walnuts most commonly traded are from the J. regia, the only species which has a large nut and thin shell. J. nigra kernels are also produced commercially in the US.

Two-thirds of the world export market[citation needed][how?][10][11] and 99% of US walnuts are grown in California's Central Valley and in Coastal Valleys, from Redding in the north to Bakersfield in the south.[12] Of the more than 30 varieties of J. regia grown there, Chandler and Hartley account for over half of total production.[11] In California commercial production, the Hinds' black walnut (J. hindsii) and the hybrid between J. hindsii and J. regia, Juglans x paradox, are widely used as rootstocks for J. regia cultivars because of their resistance to Phytophthora and to a very limited degree, the oak root fungus. However, trees grafted on these rootstocks often succumb to black line.[13]

In some countries, immature nuts in their husks are preserved in vinegar. In the UK, these are called pickled walnuts and this is one of the major uses for fresh nuts from the small scale plantings. In Armenian cuisine, unripe walnuts, including husks, are preserved in sugar syrup and eaten whole. In Italy, liqueurs called Nocino and Nocello are flavoured with walnuts, while Salsa di Noci (walnut sauce) is a pasta sauce originating from Liguria. In Georgia, walnuts are ground with other ingredients to make walnut sauce.

Green leaves of a walnut tree with budding walnuts, in Kashmir Valley.

Walnuts are heavily used in India. In Jammu, they are used widely as a prasad (offering) to Mother Goddess Vaisnav Devi and, generally, as a dry food in the season of festivals such as Diwali.

The nuts are rich in oil, and are widely eaten both fresh and in cookery. Walnut oil is expensive and consequently is used sparingly; most often in salad dressings. Walnut oil has been used in oil paint, as an effective binding medium, known for its clear, glossy consistency and nontoxicity.

Manos and Stone studied the composition of seed oils from several species of the Rhoipteleaceae and Juglandaceae and found the nut oils were generally more unsaturated from species which grow in the temperate zones and more saturated for species which grow in the tropical zones.[14] In the northerly-growing section Trachycaryon, J. cinerea oil was reported to contain 15% linolenate (the report did not specify whether the linolenate was the alpha (n-3) or gamma (n-6) isomer, or perhaps a mixture), 2% of saturated palmitate, and a maximum concentration of 71% linoleate. In the section Juglans, J. regia nut oil was found to contain from 10% to 11% linolenate, 6% to 7% palmitate, and a maximum concentration of linoleate (62% to 68%). In the section Cardiocaryon, the nut oils of J. ailantifolia and J. mandshurica were reported to contain (respectively) 7% and 5% of linolenate, 2% of palmitate, and maximum concentrations of 74% and 79% linoleate. Within the section Rhysocaryon, the nut oils of the U.S. native black walnuts J. microcarpa and J. nigra were reported to contain (respectively) 7% and 3% linolenate, 4% and 3% palmitate, and 70% and 69% linoleate. The remaining results for black walnuts were: J. australis contained 2% linolenate, 7% palmitate, and 61% linoleate; J. boliviana contained 4% linolenate, 4% palmitate, and 70% linoleate; J. hirsuta contained 2% linolenate, 5% palmitate, and 75% linoleate; J. mollis contained 0% linolenate, 5% palmitate, 46% linoleate, and 49% oleate; J. neotropica contained 3% linolenate, 5% palmitate, and 50% linoleate; and J. olanchana contained only a trace of linolenate, 9% palmitate, and 73% linoleate;

Shells

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The shells of walnuts

The walnut shell has a wide variety of uses. Eastern black walnut (J. nigra) shell is the hardest of the walnut shells, and therefore has the highest resistance to breakdown.

Cleansing and polishing
Walnut shells are mostly used to clean soft metals, fiberglass, plastics, wood and stone. This environmentally friendly and recyclable soft grit abrasive is well suited for air blasting, deburring, descaling, and polishing operations because of its elasticity and resilience. Uses include cleaning automobile and jet engines, electronic circuit boards, and paint and graffiti removal. For example: In the early days of jet transportation, crushed walnut shells were used to scour the compressor airfoils clean, but when engines with air cooled vanes and blades in the turbine started being manufactured, this practice was stopped because the crushed shells tended to plug up the cooling passages to the turbine, resulting in turbine failures due to overheating.
Oil well drilling
The shell is used widely in oil well drilling for lost circulation material in making and maintaining seals in fracture zones and unconsolidated formations.
Flour
Flour from walnut shells can be used in thermoplastic starch composites to substitute oil derivatives.[15]
Paint thickener
Walnut shells are added to paint to give it a thicker consistency for "plaster effect" ranges.
Explosives
Used as a filler in dynamite.
Cosmetic cleaner
Occasionally used in soap and exfoliating cleansers.

Husks

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Staining from handling walnuts with husks

Walnut husks are often used to create a rich yellow-brown to dark brown dye used for dyeing fabric, yarn or wood and for other purposes. The dye does not require a mordant and will readily stain the hand if picked without gloves.

Wood

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Walnut shoot cut longitudinally to show chambered pith, scale in mm

The common walnut, and the black walnut and its allies, are important for their attractive timber, which is hard, dense, tight-grained and polishes to a very smooth finish. The color is dark chocolate or similar in the heartwood changing by a sharp boundary to creamy white in the sapwood. When kiln-dried, walnut wood tends toward a dull brown color, but when air-dried can become a rich purplish-brown. Because of its color, hardness and grain, it is a prized furniture and carving wood.

When walnut vascular cambium is involved in a crotch (a branch fork), it behaves unusually, producing characteristic "crotch figure" in the wood which it makes. The grain figure exposed when a crotch in a walnut log is cut in the plane of its one entering branch and two exiting branches is attractive and sought after.

There are some differences between the wood of the European walnut (Juglans regia) and the wood of the black walnut (Juglans nigra). For example, Juglans regia wood sometimes has patches with a wavy texture.[16] Black walnut wood tends to be darker than European walnut wood, and can suffer from paler sapwood that only really comes to light when the wood has been planed.

Walnut wood has been the timber of choice for gun makers for centuries, including the Gewehr 98 and Lee–Enfield rifles of the First World War. It remains one of the most popular choices for rifle and shotgun stocks, and is generally considered to be the premium – as well as the most traditional – wood for gun stocks, due to its resilience to compression along the grain. Walnut is also used in lutherie and for the body of pipe organs.

Walnut burls (or "burrs" in the rest of the world) are commonly used to create bowls and other turned pieces. Walnut burl veneer is one of the most valuable and highly prized by cabinet makers and prestige car manufacturers.

The wood of the butternut and related Asian species is of much lower value, softer, coarser, less strong and heavy, and paler in colour.

Freshly sawn walnut heartwood may be greenish in color, but with exposure to air this color quickly changes to brown due to oxidation of the pigment.

In North America, forestry research has been undertaken, mostly on J. nigra, aiming to improve the quality of planting stock and markets. In some areas of the US, black walnut is the most valuable commercial timber species.[17] The Walnut Council[18] is the key body linking growers with scientists. In Europe, various EU-led scientific programmes have studied walnut growing for timber.[19]

The Cherokee Indians would produce a black dye from walnut bark, which they used to dye cloth.[20] As early as the 2nd century CE, shells and kernels of the edible walnut were used to make a dye solution in the Levant.[21][22]

Parkland and garden trees

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Walnuts are very attractive trees in parks and large gardens. Walnut trees are easily propagated from the nuts. Seedlings grow rapidly on good soils.[17] The Japanese walnut in particular is known for its huge leaves, which have a tropical appearance.

Walnut tree in a garden

As garden trees, they have some drawbacks, in particular the falling nuts, and the releasing of the allelopathic compound juglone, though a number of gardeners do grow them.[23][24] However, different walnut species vary in the amount of juglone they release from the roots and fallen leaves - J. nigra, in particular, is known for its toxicity, both to plants and horses.[25] Juglone is toxic to plants such as tomato, apple, and birch, and may cause stunting and death of nearby vegetation. Juglone appears to be one of the walnut's primary defence mechanisms against potential competitors for resources (water, nutrients and sunlight), and its effects are felt most strongly inside the tree's "drip line" (the circle around the tree marked by the horizontal distance of its outermost branches). However, even plants at a seemingly great distance outside the drip line can be affected, and juglone can linger in the soil for several years even after a walnut is removed as its roots slowly decompose and release juglone into the soil.

Walnut as wildlife food plants

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Walnut species are used as food plants by the larvae of some Lepidoptera species. These include[citation needed]:

The nuts are consumed by other animals, such as mice and squirrels.

In California (US) and Switzerland, crows have been witnessed taking walnuts into their beaks, flying up to 60 feet or so in the air, and dropping them to the ground to crack the shells and eat the nut inside.[26]

Nutritional information

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The raw edible seed of walnut is composed of 4% water, 14% carbohydrates, 15% protein, and 65% fat.[27] In a 100 gram amount, walnuts provide 654 calories and are a rich source (≥20% of Daily Value) of protein, dietary fiber, the B vitamins, niacin, vitamin B6, and folate, and several dietary minerals, particularly manganese.[27]

Walnut oil is composed mostly of polyunsaturated fatty acids, particularly alpha-linolenic acid and linoleic acid, although it also contains oleic acid, a monounsaturated fat and 31% of total fat is saturated fat.[27]

Systematics

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Taxonomy

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The genus Juglans is divided into four sections.[28]

Sections and species

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Section Description Image Name Common name Subspecies Distribution
Section Cardiocaryon Leaves are very large (40–90 cm), with 11–19 broad leaflets, softly downy, margins serrated. The wood is soft, and the fruits borne in racemes of up to 20. The nuts have thick shells. Native to northeast Asia. J. ailantifolia Carr. (J. cordiformis Maxim., J. sieboldiana Maxim.) Japanese walnut Japan and Sakhalin
J. mandshurica Maxim. (J. cathayensis Dode, J. formosana Hayata, J. hopeiensis Dode, J. stenocarpa Maxim.) Manchurian walnut or Chinese walnut China, Russian Far East, Korea
Section Juglans Leaves are large (20–45 cm), with 5–9 broad leaflets, hairless, margins entire. The wood is hard. Native to southeast Europe to central Asia. J. regia L. (J. duclouxiana Dode, J. fallax Dode, J. orientis Dode) common walnut, Persian, English, or Carpathian walnut Balkans eastward to Himalaya, China
J. sigillata Dode iron walnut (doubtfully distinct from J. regia) China
Section Rhysocaryon (black walnuts) Leaves are large (20–50 cm), with 11–23 slender leaflets, finely pubescent, margins serrated. Native to North America and South America. J. australis Griseb. (J. brasiliensis Dode) Argentine walnut, Brazilian walnut Argentina, Bolivia
J. boliviana (C. DC.) Dode Bolivian walnut, Peruvian walnut Andes of Bolivia and Peru
J. californica S.Wats. California black walnut California
J. hindsii (Jepson) R.E.Smith Hinds' black walnut California
J. hirsuta Manning Nuevo León walnut Mexico
J. jamaicensis C.DC. (J. insularis Griseb.) West Indies walnut Cuba, the Dominican Republic, Haiti, and Puerto Rico
J. major (Torrey) Heller (J. arizonica Dode, J. elaeopyron Dode, J. torreyi Dode) Arizona black walnut
  • J. major var. glabrata Manning
Mexico, United States
J. microcarpa Berlandier (J. rupestris Engelm.) Texas black walnut
  • J. microcarpa var. microcarpa
  • J. microcarpa var. stewartii (Johnston) Manning
United States
J. mollis Engelm. Mexican walnut Mexico
J. neotropica Diels (J. honorei Dode) Andean walnut, cedro negro, cedro nogal, nogal, nogal Bogotano Colombia, Ecuador, and Peru
J. nigra L. Eastern black walnut Canada, United States
J. olanchana Standl. & L.O.Williams cedro negro, nogal, walnut
  • J. olanchana var. olanchana
  • J. olanchana var. standleyi
Central America, Mexico
J. soratensis Manning Bolivia
J. steyermarkii Manning Guatemalan walnut Guatemala
J. venezuelensis Manning Venezuelan walnut Venezuela
Section Trachycaryon Leaves are very large (40–90 cm), with 11–19 broad leaflets, softly downy, margins serrated. The wood is soft. Fruits are borne in clusters of two to three. The nuts have a thick, rough shell bearing distinct, sharp ridges. Native to eastern North America. J. cinerea L. Butternut Canada, United States

The best-known member of the genus is the Persian walnut (J. regia, literally "royal walnut"), native from the Balkans in southeast Europe, southwest and central Asia to the Himalaya and southwest China. Walnuts are a traditional feature of Iranian cuisine; the nation has extensive orchards which are an important feature of regional economies. In Kyrgyzstan alone, there are 230,700 ha of walnut-fruit forest, where J. regia is the dominant overstory tree (Hemery and Popov 1998). In non-European English-speaking nations, the nut of the J. regia is often called the "English walnut"; in Great Britain, the "common walnut."

The eastern black walnut (J. nigra) is a common species in its native eastern North America, and is also widely cultivated elsewhere. The nuts are edible, and though they are often used in expensive baked goods, the Persian walnut is preferred for everyday use because it is easier to extract the nutmeat. The wood is particularly valuable.

The Hinds' black walnut (J. hindsii) is native to northern California, where it has been widely used commercially as a rootstock for J. regia trees. Hinds' black walnut shells do not have the deep grooves characteristic of the eastern black walnut.

Japanese walnut foliage and nuts

The Japanese walnut (J. ailantifolia) is similar to butternut, distinguished by the larger leaves up to 90 cm long, and round (not oval) nuts. The variety cordiformis, often called the heartnut has heart-shaped nuts; the common name of this variety is the source of the sectional name Cardiocaryon.

The butternut (J. cinerea) is also native to eastern North America, where it is currently endangered by an introduced disease, butternut canker, caused by the fungus Sirococcus clavigignenti-juglandacearum. Its leaves are 40–60 cm long, the fruits are oval, the shell has very tall, very slender ridges, and the kernel is especially high in fat.

Hybrids

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  • J. × bixbyi Rehd.—J. ailantifolia x J. cinerea
  • J. × intermedia Carr.—J. nigra x J. regia
  • J. × notha Rehd.—J. ailantifolia x J. regia
  • J. × quadrangulata (Carr.) Rehd.—J. cinerea x J. regia
  • J. × sinensis (D. C.) Rehd.—J. mandschurica x J. regia
  • J. × paradox Burbank—J. hindsii x J. regia
  • J. × royal Burbank—J. hindsii x J. nigra

Phylogeny

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A study[29] of sequenced nuclear DNA from the external transcribed spacer (ETS) of ribosomal DNA (rDNA), the internal transcribed spacer (ITS) of rDNA, and the second intron of the LEAFY gene taken from at least one individual of most of the species of Juglans has supported several conclusions:

  • The genus Juglans is monophyletic;
  • Sect. Cardiocaryon is sister to Sect. Trachycaryon;
  • Sect. Juglans is sister to Sect. Cardiocaryon and Sect. Trachycaryon together;
  • Sect. Rhysocaryon is monophyletic and sister to Sect. Juglans, Sect. Cardiocaryon, and Sect. Trachycaryon together;
  • Sect. Rhysocaryon, the black walnuts, contains two clades:
    • one comprises the more northerly species J. californica, J. hindsii, J. hirsuta, J. major, J. microcarpa, and J. nigra;
    • the other comprises the more southerly species J. australis, J. boliviana, J. jamaicensis, J. molis, J. neotropica, J. olanchana, J. steyermarkii, and J. venezuelensis
  • J. olanchana var. standleyi seems to be more closely related to J. steyermarkii than to J. olanchana var. olanchana, suggesting J. olanchana var. standleyi might be better understood as either a separate species or a variety of J. steyermarkii.

The paper presenting these results did not publish any new names for the subdivisions of sect. Rhysocaryon, for any combinations of the other sections, or for J. olanchana var. standleyi.

Paleontological history

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Fossils of Juglans nuts have been described from the Tertiary period of North America.[30] The paleontological history of Juglans regia in Europe shows signs of a post-Ice-Age re-expansíon from refugia in the southeast, much influenced by people carrying walnut nuts about after the numbers of humans had been much increased by the start of agriculture.[citation needed]

See also

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References

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[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia
from Grokipedia
Juglans, commonly known as the , is a of about 21 of trees and shrubs in the family , native to the and western , including North and , the Mediterranean, and eastern to . These are characterized by pinnate, aromatic compound leaves, twigs with chambered , and monoecious flowers borne in catkins (staminate) or solitary (pistillate), producing distinctive drupaceous fruits that enclose large, seeds within a hard, ridged shell. Species in this genus typically grow to heights of 20–40 meters, thriving in temperate to subtropical climates with well-drained soils, though many exhibit sensitivity to and poor drainage. The genus is best known for its economic and cultural significance, with all species producing nuts that are harvested for food, though some contain bitter or inedible kernels without processing. Prominent species include the Persian or English walnut (J. regia), widely cultivated for its high-quality nuts and timber in regions like California and the Mediterranean, and the black walnut (J. nigra), a North American native prized for its durable wood used in furniture and gunstocks but noted for its allelopathic compound juglone, which inhibits understory plant growth. Other notable taxa, such as the butternut (J. cinerea), threatened by butternut canker, and the California walnut (J. californica), susceptible to thousand cankers disease, face serious threats impacting conservation efforts. Beyond , Juglans have historical and medicinal value; for instance, bark, leaves, and husks have been used in traditional remedies for their and properties, supported by studies identifying and polyphenols. The genus's reflects ancient Tertiary disjunctions between the and , with ongoing research using to resolve relationships and hybridization events.

Etymology and cultural aspects

Etymology

The genus name Juglans derives from the Latin iūglans, a contraction of Jovis glans, meaning "Jupiter's acorn" or "Jupiter's nut," which reflects the ancient Roman reverence for the walnut tree as a plant sacred to the god Jupiter. This etymology underscores the nut's perceived divine qualities in classical antiquity, where walnuts were associated with strength and fertility akin to the chief deity. The common English name "" evolved from wealhhnutu, literally translating to "foreign nut," combining wealh (meaning "foreign" or "Celtic," referring to non-Germanic peoples) with hnutu (nut); this term arose because walnut trees (Juglans spp.) were introduced to Britain from by the Romans. Over time, the name persisted in as walnot or walnutte, adapting to describe the nut's exotic origin relative to native British flora. Linguistic variations for walnut in Romance languages trace back to the Latin nux (nut), emphasizing the fruit's hard-shelled, seed characteristic. In Spanish, nuez (walnut) directly inherits from Latin nucem, the accusative form of nux, and broadly denotes nuts while specifically applying to walnuts in culinary contexts. Similarly, French noix (walnut) stems from nois, also from Latin nucem, capturing the nut's kernel-enclosing structure and use in both everyday and gourmet traditions.

Folklore and historical significance

In ancient Greek mythology, the is associated with the story of Carya, a princess of Laconia whom the god transformed into a after her death, symbolizing the nut's enduring cultural reverence. Romans further elevated walnuts in rituals, where guests scattered them at ceremonies to invoke and abundance, viewing the nuts as emblems of procreation and prosperity due to their prolific yield. During the in , walnuts were employed in herbalism under the doctrine of signatures, with their brain-like appearance leading to their use as remedies for head ailments, madness, and cognitive disorders. Folk beliefs also attributed protective qualities to walnuts, believing they could ward off and the when carried or used in charms, reflecting their role in countering malevolent forces. In Chinese traditions, walnuts (known as hu tao) symbolize and intellectual vitality, a association rooted in ancient texts like Shennong's Materia Medica Classic (circa 200–250 AD), which classified them as a superior tonic for nourishing the brain, extending life, and promoting overall vitality. This symbolism persists in cultural practices, where walnuts are gifted to convey wishes for and enduring . The historical dissemination of (common walnut) from Persia along ancient trade routes underscores its cultural significance, with humans facilitating its spread via the Persian Royal Road (circa 500–330 BCE) from and the Trans-Caucasus to , and later through the networks starting around 138 BCE, reaching by the AD during Roman expansion. These corridors not only traded the nuts but also embedded walnuts in diverse folk narratives across .

Botanical description

Growth habit and morphology

Juglans species are trees that typically reach heights of 10 to 40 , forming a straight trunk and a broad, rounded crown in mature individuals. The twigs have chambered , a diagnostic feature of the . The leaves are alternate and pinnately compound, often measuring up to 1 meter in length, composed of 5 to 25 leaflets that emit a characteristic aromatic scent due to the presence of resinous peltate glands. The bark on mature trees is rough and deeply furrowed, ranging in color from gray to dark brown, providing a distinctive textured appearance that develops with age. Growth rates for Juglans trees are generally slow to moderate, allowing them to achieve substantial size over time, with some exhibiting of 200 to 300 years under favorable conditions. The root system features a prominent deep supplemented by wide-spreading lateral , which the effectively in well-drained soils and contribute to its to various site conditions. These exhibit allelopathic properties through the production of , a primarily released from , bark, and leaves, which inhibits the growth of nearby plants by disrupting metabolic processes.

Flowers and reproduction

Juglans species are monoecious, producing separate flowers on the same individual , a characteristic trait that facilitates within the genus. flowers, or staminate blooms, develop in pendulous catkins that emerge from scars of the previous year's growth, typically measuring 5–15 cm in length and consisting of numerous small, scalelike structures lacking petals. Female flowers, or pistillate blooms, form in short terminal spikes at the tips of current-season shoots, usually comprising 2–5 flowers per spike, each featuring a prominent, feathery stigma adapted for capturing airborne and an inferior . Reproduction in Juglans relies on wind pollination, or anemophily, where lightweight grains are released from the anthers of male catkins and carried by air currents to receptive female stigmas. Flowering typically occurs in spring, from to in temperate regions, with the exact timing varying by and local climate; for instance, in , male bloom may precede or follow female receptivity depending on the . Many exhibit dichogamy, where the maturation of male and female flowers on the same tree is temporally offset, reducing ; this is often heterodichogamous, with populations containing both protandrous (male-first) and protogynous (female-first) individuals at roughly equal ratios to promote cross-pollination between trees. Following successful , seeds develop within the drupelike fruits and are primarily dispersed by animals such as squirrels and that cache the nuts after they fall by beneath the parent . Germination rates are generally low due to , necessitating a period of cold stratification—typically 90–120 days at 0–4°C under moist conditions—to break physiological barriers and initiate growth in spring. This requirement ensures that seedlings emerge under favorable post-winter conditions, though viability can be further challenged by husk and exposure if not properly managed.

Fruit, nuts, and husks

The fruits of Juglans species are pseudodrupes, characterized by a fleshy, green outer that encloses a hard-shelled nut containing an edible kernel. The , which develops from the inferior and associated floral structures, is indehiscent—meaning it does not split open naturally—and typically remains firmly attached to the nut until manually removed. This structure protects the nut during development and dispersal, with the overall ranging from 3 to 5 cm in diameter depending on the . In Juglans nigra (black ), the is particularly thick and leathery, providing robust protection, whereas in species like Juglans regia (English ), it tends to be thinner and more easily separated post-maturity. The nut itself features a lignified endocarp, or shell, composed primarily of sclerified cells with high lignin content for rigidity and durability. This shell often exhibits prominent ridges and furrows on its surface, which vary in depth and pattern across species—for instance, deeply ridged in J. nigra and smoother in some Asian walnuts like J. mandshurica. Inside the shell lies the kernel, a cotyledon-rich seed with an oil content typically ranging from 50% to 70% of its dry weight, predominantly composed of unsaturated fatty acids such as linoleic and α-linolenic acids (over 70% of total lipids). These oils contribute to the kernel's nutritional value but also make it susceptible to oxidation if not stored properly. The is chemically rich, containing (a ) and high levels of , which impart allelopathic properties and contribute to the plant's ecological interactions. , synthesized in the and other tissues, can inhibit the growth of nearby plants and certain microorganisms, while cause the characteristic dark staining on , fabrics, and tools upon contact due to their polyphenolic . These compounds also render the toxic to some animals and plants, limiting husk consumption but aiding in pest deterrence. Fruits typically mature in autumn, approximately 5 to 7 months after flowering, with the turning from green to yellowish-brown as the kernel reaches full size and the shell hardens.

Systematics and evolution

Taxonomy and classification

The genus Juglans is placed within the family , which belongs to the order in the class Magnoliopsida. The genus was formally established by in the first edition of published in 1753, where he described several species based on morphological characteristics such as pinnate leaves and drupaceous fruits. Linnaeus designated L., the common or English walnut, as the type species for the genus. Throughout its nomenclatural history, Juglans has faced challenges with synonymy, particularly among Asian species. For instance, East Asian taxa in section Cardiocaryon, such as those formerly confused under names like Juglans ailantifolia and Juglans mandshurica, have been subject to historical misclassifications due to overlapping morphological traits and incomplete records, leading to inconsistent synonymy across floras. These issues stem from early botanical explorations that often relied on limited specimens from regions like , , and Korea, resulting in provisional names that were later revised. As of recent assessments, the comprises approximately 21 accepted , though this number reflects ongoing taxonomic revisions driven by molecular phylogenetic studies. Analyses using nuclear and chloroplast DNA sequences have clarified relationships and prompted re-evaluations of species boundaries, especially in resolving ambiguities among Asian and American taxa. These molecular approaches have highlighted the need for further integration of genomic data to stabilize the classification.

Sections and species

The genus Juglans is divided into four main infrageneric sections based on morphological and phylogenetic analyses: Cardiocaryon, Dioscaryon, Rhysocaryon, and Trachycaryon. These sections reflect distinct evolutionary lineages supported by molecular data from matK and ITS sequences, with differences primarily in leaf morphology, fruit husk dehiscence, and nut shell structure. All species exhibit chambered pith in twigs, a key diagnostic trait distinguishing Juglans from related genera like Carya, though pith color varies subtly between sections—darker brown in Trachycaryon compared to lighter tones in others. Section Cardiocaryon, comprising Asiatic walnuts or butternuts, includes three : J. ailantifolia, J. mandshurica, and J. cathayensis. These are characterized by large compound leaves with 11–19 serrate leaflets, elongated fruits with sticky, indehiscent husks that produce oil-rich nuts, and a growth habit favoring temperate East Asian forests. The section is phylogenetically close to Trachycaryon, sharing butternut-like traits such as furrowed nut shells and pubescent twigs. Section Dioscaryon (synonymous with section Juglans in some classifications) contains primarily J. regia and J. sigillata. It features fewer leaflets (5–9, often entire-margined), dehiscent fruit husks that split at maturity, and smooth, thin-shelled nuts with shallow, winged sutures. These traits adapt the section to drier, mountainous habitats in . Section Rhysocaryon, the black walnuts, is the most species-rich, encompassing about 12 taxa primarily from the , including J. nigra, J. major, J. microcarpa, J. californica, J. hindsii, J. neotropica, and J. olanchana. Morphological hallmarks include 15–23 serrate leaflets per leaf, thick indehiscent husks, and deeply ridged, hard-shelled nuts; the chambered pith is typically pale to tan. This section shows greater diversity in tropical to temperate adaptations. Section Trachycaryon is monotypic, represented solely by J. cinerea. It shares butternut features with Cardiocaryon, such as sticky husks and elongated, easily cracked nuts, but differs in having 11–17 leaflets and darker chambered ; leaves are often more pubescent, suiting mesic North American woodlands. Among key species, (English or Persian walnut) is a reaching 15–25 m in height, with a broad crown and pinnate leaves bearing 5–9 ovate leaflets up to 15 cm long. Native to the mountainous regions of from the to southwest , it is the primary cultivated worldwide for its edible, thin-shelled nuts enclosed in a green husk. (black ), from section Rhysocaryon, grows as a large to 30–40 m tall with dark, furrowed bark and compound leaves of 15–23 lanceolate leaflets. Endemic to eastern , from to northern and west to , it yields high-value timber and flavorful nuts, though husks stain and inhibit growth via . Juglans cinerea (butternut), the sole Trachycaryon species, attains 20–30 m with lighter gray bark and leaves of 11–17 leaflets; its oily nuts are elongated and ridged. Native across eastern from to , it prefers rich, moist soils but faces decline from fungal cankers. Notable endemics include Juglans olanchana in section Rhysocaryon, a tree up to 40 m tall with large leaves (17–21 leaflets) and round-husked fruits containing edible nuts. Restricted to mid-elevation moist forests in , from southern through to , it provides valuable similar to J. nigra but with deeper chocolate tones. Hybrids, such as those between Rhysocaryon and Trachycaryon species like J. nigra × J. cinerea, exhibit intermediate traits and are documented in cultivation.

Hybrids and genetic diversity

Juglans species readily form natural interspecific hybrids due to overlapping flowering phenologies and sharing, with notable examples including Juglans × intermedia (a cross between J. nigra and J. regia). This hybrid demonstrates enhanced growth rates and improved resistance to diseases compared to its parental species, making it suitable for timber production and as a in systems across and . Another common natural hybrid occurs between butternut (J. cinerea) and Japanese walnut (J. ailantifolia), which exhibits resistance to the butternut (Sirococcus clavigignenti-juglandacearum) and has been identified using markers in natural populations. Artificial hybridization programs for Juglans began in the late 19th and early 20th centuries, pioneered by figures like Luther Burbank, who conducted controlled crosses to combine desirable traits from different species. Throughout the 20th century, breeding efforts expanded globally, focusing on enhancing disease resistance—such as to anthracnose and Phytophthora root rot—and improving nut quality, yield, and adaptability, with programs in the United States, Europe, and Asia developing hybrid rootstocks like the Paradox hybrid (J. hindsii × J. regia) for superior vigor and tolerance. These initiatives have produced fertile hybrids that support sustainable cultivation, though challenges like post-zygotic barriers can reduce hybrid viability. Genetic diversity in Juglans is typically high, reflecting the outcrossing nature of most species, as assessed through simple sequence repeat (SSR) markers that reveal substantial heterozygosity. For instance, studies on J. regia populations report observed heterozygosity (H_O) averaging 0.56 across loci, with expected heterozygosity (H_E) similarly elevated at around 0.44–0.59, indicating robust allelic variation that supports adaptive potential in breeding. SSR analyses also highlight population-level differentiation, with gene flow facilitating diversity maintenance in natural stands. In conservation contexts, some endangered Juglans species exhibit reduced due to and , posing risks for long-term viability. For example, J. neotropica (Andean walnut) populations in the Amazonas region show low plastid diversity, linked to deforestation-induced , which underscores the need for targeted ex situ preservation to retain adaptive alleles.

Phylogeny

The genus Juglans diverged from other lineages within the family approximately 40–50 million years ago during the Eocene, based on estimates from chloroplast DNA sequence data. This divergence reflects the broader radiation of the walnut family in the following the Cretaceous-Paleogene boundary. Molecular phylogenetic studies, primarily using noncoding spacers and nuclear ribosomal ITS sequences, resolve Juglans with section Rhysocaryon (American black walnuts) as the basal , sister to a derived comprising section Dioscaryon (Eurasian walnuts) and sections Cardiocaryon + Trachycaryon (Asian butternuts and American butternut), the latter two being closely related. The American species form the to the lineages, indicating an early divergence in , with section Trachycaryon phylogenetically closer to Cardiocaryon than to Rhysocaryon. Key divergence events between these s occurred during the (around 23–5 million years ago), driven by tectonic uplift and climate shifts that promoted vicariance and dispersal across . Morphological analyses reinforce these molecular findings, with traits such as in twigs identified as a synapomorphy defining the Rhysocaryon section, distinguishing it from other sections and supporting its basal position among living species. Post-2010 studies employing next-generation sequencing technologies, including complete genome assemblies and restriction-site associated DNA (RAD-Seq), have robustly confirmed the of Juglans sections and refined interclade relationships with higher resolution than earlier marker-based approaches. These genomic datasets also highlight low sequence divergence within sections, consistent with relatively recent radiations.

Fossil record

The fossil record of Juglans (walnuts) extends back to the Eocene epoch, with the earliest known specimens consisting of fruits and leaves from North American sites, such as Juglans clarnensis dated to approximately 44 million years ago. These early fossils, including well-preserved nuts from the Green River Formation in the , indicate that the genus was already established in subtropical to temperate forests during a period of global warming known as the Eocene climatic optimum. Additional Eocene records from high-latitude locations, like the mummified forests of in the Canadian , reveal species such as Juglans eoarctica, J. nathorstii, and J. cordata, highlighting the genus's adaptation to diverse paleoenvironments across northern continents. During the and epochs, Juglans underwent significant diversification, with fossils documenting its spread across , the northern supercontinent comprising and . This expansion is evidenced by leaf, fruit, and pollen remains in sedimentary deposits from , , and , reflecting the genus's migration via land bridges such as and the North Atlantic connections. The Oligocene-Miocene warming and subsequent cooling phases facilitated the evolution of deciduous forms adapted to seasonal climates, leading to a broader Holarctic distribution that included both black walnut (sect. Rhysocaryon) and butternut (sects. Cardiocaryon and Trachycaryon) lineages. In , Juglans fossils become prominent from the onward but show a marked decline after the , coinciding with the onset of Pleistocene glaciations around 2.6 million years ago. This retreat is attributed to climatic cooling and expansion, which fragmented habitats and led to the local of many European lineages, as seen in reduced records from Alpine and Mediterranean sites. The ancient Holarctic range preserved in the fossil record provides key insights into biogeographic patterns, explaining the current disjunct distribution of Juglans between the and through vicariance and long-distance dispersal events.

Distribution and ecology

Native and introduced ranges

The genus Juglans comprises approximately 21 species of trees, with 15 native to the and 6 to the . Native distributions span temperate and subtropical regions across both hemispheres, reflecting the genus's evolutionary divergence. In the , Juglans species are concentrated in North, Central, and , extending from southeastern and the southward to northwestern and . For example, (black walnut) is native to the , ranging from southeastern through the central and eastern U.S., typically in mixed forests. Other species, such as (butternut), occupy similar eastern North American ranges, while South American taxa like Juglans neotropica occur in Andean cloud forests. In , the six native species are primarily distributed from the Mediterranean Basin eastward to the and Japan, with (Persian or English walnut) originating in , specifically from northeastern through the . This species thrives in riparian and foothill habitats in its native range, often forming pure stands. Other species, such as Juglans mandshurica, extend into temperate eastern . Several Juglans species have been widely introduced beyond their native ranges for timber, nuts, and ornamental purposes. J. regia, in particular, was introduced to by Spanish missionaries in the early and has since become a major crop there. It reached during the via European colonization and is now cultivated extensively in the country's central valleys. In , J. regia was first planted in around 1820, with widespread establishment across southern regions during the . Some introduced Juglans species exhibit and potential invasiveness outside their native ranges. For instance, J. nigra has become naturalized in parts of since its introduction in the 17th century and shows invasive tendencies in certain habitats due to its competitive growth and allelopathic effects. Similarly, J. regia displays high invasiveness in Central European farmlands, altering local plant communities. Juglans predominantly occupy zones, with some extending into subtropical montane areas; they occur from to elevations of up to 3,000 m, as seen in Andean and Himalayan populations.

Habitat and environmental preferences

of the genus Juglans thrive in deep, fertile, well-drained loamy that support their extensive root systems. These are typically neutral to slightly alkaline, with optimal pH levels ranging from 6.0 to 7.5, allowing for efficient nutrient uptake. Juglans exhibit low tolerance to waterlogging, as prolonged saturation can lead to and reduced vigor. In terms of climate, Juglans species are adapted to temperate regions characterized by cold winters and moderate summers, corresponding to USDA hardiness zones 4 through 9. They require annual precipitation between approximately 750 and 1500 mm to sustain growth, with distributions often aligned to areas receiving this range in their native habitats. Juglans trees demand full sun exposure for robust development and nut production, showing intolerance to shaded conditions. Due to their broad crowns and the allelopathic effects of juglone—a compound exuded from roots and hulls that inhibits nearby plant growth—wide spacing is essential in natural stands, often exceeding 10 meters between mature individuals to minimize competition and toxicity. Certain species, such as J. major in the arid Southwest United States, demonstrate notable drought tolerance through deep root penetration to access groundwater, enabling persistence in riparian zones with limited surface moisture.

Ecological interactions

Juglans species exhibit significant allelopathic effects through the production of juglone, a naphthoquinone compound synthesized in roots, leaves, and husks, which inhibits the growth of competing vegetation. This chemical is exuded from roots into the soil and released upon leaf decomposition, creating zones of suppression around the trees that can extend up to 50-80 feet from the trunk, particularly affecting sensitive species such as apple trees (Malus spp.) and pines (Pinus spp.). Allelopathy provides Juglans with a competitive advantage in nutrient-limited environments by reducing understory competition for resources like light and water. Juglans trees form symbiotic associations with mycorrhizal fungi, including both arbuscular mycorrhizal fungi (AMF) and, in some species or conditions, ectomycorrhizal fungi, which enhance nutrient uptake in nutrient-poor soils. These associations involve the exchange of plant-derived carbohydrates for fungal-mediated acquisition of , , and , improving tree vigor and tolerance to environmental stresses such as . For instance, AMF colonization in roots has been shown to increase absorption by up to 40% in low-fertility soils, supporting growth in marginal habitats. The nuts of Juglans serve as a key food source for wildlife, particularly rodents like eastern fox squirrels (Sciurus niger), which incorporate them into about 10% of their diet, aiding in seed dispersal through caching behavior. Birds such as blue jays (Cyanocitta cristata) and wild turkeys (Meleagris gallopavo), along with deer (Odocoileus virginianus), also consume the nuts, with uneaten caches contributing to natural regeneration. Husks are occasionally gnawed and consumed by rodents including wood mice (Apodemus speciosus), though their high tannin content limits intake. Leaves are browsed sparingly by white-tailed deer, which find them palatable but not preferred due to juglone content. In natural ecosystems, the canopy of Juglans trees provides shaded and nesting sites for birds and small mammals, fostering localized by supporting over 100 of larvae as a host . However, in stands, the combined effects of and dense shading reduce understory diversity, often leading to sparse herbaceous layers dominated by tolerant and diminished overall phytocoenosis richness compared to mixed forests. This impact is particularly pronounced in introduced ranges, where lack of co-evolved competitors exacerbates suppression of native .

Cultivation and production

Commercial cultivation

Commercial cultivation of Juglans species, primarily J. regia (English ), relies on vegetative to maintain desirable cultivars while leveraging rootstocks for improved performance in environments. Trees are typically propagated by or scion wood onto rootstocks, with common options including J. regia itself for standard adaptability or the hybrid (J. hindsii × J. regia) for enhanced vigor, deeper rooting, and resistance to soilborne pathogens like . This method ensures uniformity in nut quality and yield potential, as seed alone would not preserve varietal traits. Orchard establishment begins with planting grafted nursery stock at densities of 56 to 75 trees per acre, corresponding to spacings of approximately 6 to 8 (20 to 26 feet) between trees and rows in square or offset configurations. This spacing accommodates mature canopy development while allowing machinery access and optimizing light exposure for nut production. Trees generally require 4 to 6 years after planting to reach bearing age, during which initial growth focuses on structural development rather than set. Pruning and training are essential from the first few years to shape trees into productive forms, with the modified central leader system being widely adopted to create a balanced that promotes even light penetration throughout the canopy. In this approach, a dominant central trunk is maintained, with 4 to 6 primary scaffolds selected and headed at wide angles (45–60 degrees) during in years 1 through 6, removing competing shoots and watersprouts to direct energy toward fruiting wood. Such practices reduce in the lower canopy, enhancing nut fill and overall orchard productivity. In regions with limited rainfall, systems are standard for delivering water directly to the root zone, minimizing evaporation and growth while supporting consistent levels critical for nut development. Fertilization emphasizes to sustain vigorous growth and high yields, with applications typically ranging from 100 to 200 pounds per acre annually for mature orchards, often via fertigation through drip lines to improve uptake efficiency on fertile soils. These inputs are timed with to avoid leaching, particularly in sandy or low-fertility sites.

Major production regions

China is the world's largest producer of walnuts (Juglans spp.), accounting for approximately 56% of global output with an estimated 1.5 million metric tons in the 2024/2025 marketing year. The United States ranks second, primarily through production in California, yielding 547,032 metric tons (20% of the total) in the 2024 crop year. Chile contributed approximately 152,500 metric tons (6%) in 2024/2025. Other significant producers include Iran with an estimated 180,000 metric tons and Turkey with 67,000 metric tons in 2024/2025. Global walnut production for 2024/2025 is estimated at approximately 2.6 million metric tons, reflecting adjustments from initial forecasts due to lower yields in key regions. Yields in commercial J. regia orchards typically average 2-4 tons per for mature trees under standard management, though optimized conditions can exceed 5 tons per . Production has shown an average annual growth rate of about 3-5% from 2015 to 2024, fueled by rising global demand for nuts in and products, though 2024 saw a dip in some regions. For the 2025 crop, production is forecasted at 644,000 metric tons, up 18% from 2024. The United States is a major exporter, shipping walnuts valued at over $300 million in inshell form in 2023, with primary markets in the European Union and Asia. However, exports faced challenges from retaliatory tariffs imposed during the U.S.-China trade wars starting in 2018, which reduced shipments to China and pressured prices. Post-2020 trade agreements have partially mitigated these issues, but ongoing tariffs continue to influence market dynamics. Historical production shifts include significant expansion in the since the 1990s, particularly in , where planted area grew from modest levels to over 46,000 hectares by 2023, enabling counter-seasonal harvests for northern markets. has similarly increased its walnut acreage from around 500 hectares in the early 1990s to over 3,000 hectares by the 2010s, supporting year-round global supply chains.

Harvesting and processing

Harvesting of Juglans nuts, primarily from species like (English walnut), occurs in the fall when the outer green husks begin to split, typically from mid-September to early in major production regions such as . This timing ensures nut maturity, with mechanical shakers used to dislodge the majority of nuts from trees once approximately 95% of husks have split, minimizing damage and facilitating efficient collection. Prompt harvest prevents over-maturity, which can lead to nut drop and potential quality loss. Following collection, dehulling removes the green husks mechanically using hullers equipped with wet scrubbers or rollers to separate the hull from the hard shell without excessive force. This step is critical to prevent of the shell by compounds in the husk and to avoid mold development from trapped . Dehulled nuts are then dried using forced hot air systems, such as column or tray dryers, at temperatures around 110°F (43°C) until reaching 8% content on a wet basis, which typically takes 24 hours or less. This process halts enzymatic activity and microbial growth, preserving kernel integrity. Post-drying, nuts undergo sorting to ensure quality, often employing optical sorters that use near-infrared or to detect defects like discoloration, damage, or foreign materials based on size, color, and density. High-quality in-shell nuts are then cracked using precision shelling machines that apply controlled centrifugal or impact forces to the shell while minimizing kernel damage, achieving shelling rates above 90% with kernel recovery often exceeding 50% intact. These machines grade nuts by size beforehand to optimize cracking efficiency and reduce breakage. For storage, in-shell Juglans nuts are kept in controlled environments at 0-5°C (32-41°F) with relative humidity below 65% to maintain quality and extend up to one year, preventing rancidity from oxidation. Shelled kernels require similar cool temperatures but benefit from nitrogen-flushed, moisture-barrier packaging to further inhibit oxidative degradation.

Uses and economic importance

Nutritional profile

Juglans kernels, especially from (English walnut), exhibit a high-calorie of 654 kcal per 100 g, primarily due to their macronutrient composition of approximately 65 g , 15 g protein, and 14 g total carbohydrates (including 7 g ). The content is dominated by polyunsaturated fatty acids, with alpha-linolenic acid (ALA), an essential omega-3 fatty acid, comprising 9.08 g per 100 g, making walnuts one of the richest plant-based sources of this nutrient. This profile positions Juglans kernels as a concentrated energy source suitable for moderate dietary inclusion. Micronutrients in kernels are particularly abundant in minerals and . levels reach 3.41 mg per 100 g (about 148% of the daily value), supporting enzymatic functions in and , while provides 1.59 mg per 100 g (176% daily value), aiding in iron absorption and formation. , primarily in the gamma-tocopherol form at around 21 mg per 100 g, contributes protection against , alongside derived from ellagitannins, which exhibits potent free radical-scavenging properties. Clinical evidence links regular Juglans kernel consumption to cardiovascular benefits, with meta-analyses indicating that a daily intake of approximately 30 g reduces (LDL) cholesterol by 2.8–4.3% and total by 3.5–5.1% compared to control diets, potentially lowering heart disease risk through improved lipid profiles and effects. These outcomes are attributed to the synergistic action of ALA, polyphenols, and . Nutritional variations exist across ; for instance, J. regia kernels contain significantly higher ALA (9.08 g per 100 g) than J. nigra (black walnut) kernels (2.68 g per 100 g), influencing their relative omega-3 contributions.

Food and culinary applications

Walnuts from species such as Juglans regia and Juglans nigra are consumed raw for their fresh, mildly bitter flavor or processed through roasting to enhance nuttiness and texture. Roasting at temperatures between 120°C and 150°C for 15-20 minutes yields optimal sensory properties, including improved aroma and reduced astringency, without excessive hardening. This process breaks down cell walls, releasing oils that intensify the earthy taste while preserving nutritional integrity. In global cuisines, walnuts feature prominently in both sweet and savory dishes, reflecting regional traditions. In Middle Eastern and Mediterranean cooking, they form a key filling in , a layered phyllo pastry soaked in honey syrup, where chopped walnuts provide crunch and richness alongside spices like . Italian from employs walnuts in salsa di noci, a creamy sauce blended with bread, milk, garlic, and cheese, typically tossed with pasta or for a nutty, garlicky contrast. In American confections, walnuts appear in baked goods such as brownies, , and cookies, adding texture and a subtle bitterness that balances sweetness. Value-added products expand walnuts' culinary versatility. , pressed from kernels, has a of 160°C in its unrefined form, making it suitable for low-heat or dressings where its delicate, fruity notes shine without bitterness. butter, ground from roasted nuts, serves as a spread on toast or an ingredient in smoothies and , offering a creamy alternative to with added omega-3s. Ground into , walnuts enable gluten-free , such as in quick breads or cakes, where their high oil content contributes moisture and a tender crumb without wheat. Despite these applications, walnuts pose risks for those with tree nut allergies, which affect approximately 1% of the U.S. . Clinical is high between walnuts and pecans due to shared protein structures, with up to two-thirds of walnut-allergic individuals also reacting to pecans. [](https://www.nist.gov/news-events/news/2022/06/nist-tree-nut-reference-materials-support-food-allergen-testing

Industrial and material uses

The wood of , commonly known as black walnut, is prized for its dark heartwood, which ranges from chocolate brown to deep purplish black, often with straight grain and a fine, even texture that enhances its workability. This species exhibits a Janka hardness of 1,010 lbf, indicating moderate resistance to wear while remaining machinable for precision applications. Due to these properties, black walnut is extensively used in high-end furniture, , and gunstocks, where its stability, shock resistance, and aesthetic appeal provide durability and visual elegance. Globally, Juglans species, particularly J. regia, contribute to a walnut nut market valued at over $4 billion USD as of 2023, with production totaling around 3.5 million metric tons, led by (1.8 million tons). Walnut shells, a of nut processing, are ground into grit for use as an blasting media, valued for its non-toxic, biodegradable nature and low dust generation compared to silica-based alternatives. With a Mohs hardness of approximately 3.5, it effectively cleans soft surfaces like wood, aluminum, and without causing pitting or scratching, and is also employed in filtration systems for due to its porous structure. The husks, rich in and , serve as a source for textiles, yielding brown to black hues on protein and fibers when mordanted or used directly, historically applied in fabric coloration for earthy tones. Juglone, extracted from walnut husks, leaves, and bark, demonstrates antifungal properties comparable to certain commercial agents, inhibiting fungal growth in vitro and showing potential in pharmaceutical applications for treating infections. Walnut byproducts, including leaves and husks, can be composted after detoxification, as juglone breaks down rapidly—within weeks—through microbial activity and exposure to air and moisture, rendering the material safe for soil amendment without allelopathic risks. The U.S. black walnut lumber market supports significant economic activity; as of 2020, black walnut sawtimber volume stands at 13.1 billion board feet, with annual net growth of approximately 122 million cubic feet for growing stock. Stumpage prices for high-quality logs typically range from $1,000 to $5,000 or more per thousand board feet, depending on grade and region, as of 2024-2025, underscoring the premium placed on this resource.

Ornamental and environmental uses

Species of Juglans, particularly J. nigra (black walnut) and J. regia (English walnut), are employed in ornamental for their stately growth and aesthetic qualities. Black walnut trees are cultivated as shade trees in residential and park settings, valued for their broad, rounded crowns reaching up to 100 feet in height and their dark green, pinnate foliage that provides dense summer shade. Their open winter structure and rich bark texture add year-round visual interest, though maintenance is required to manage and nut drop. English walnut, with its spreading, dome-shaped canopy, serves as a specimen in expansive landscapes, offering seasonal appeal through catkins in spring, glossy summer leaves, golden fall color, and pendulous fruits. However, the allelopathic compound produced by Juglans roots, leaves, and husks inhibits growth of many nearby plants, necessitating careful site planning to avoid incompatibility with gardens or turf. Raised beds or distance from sensitive species are recommended for successful integration. J. cinerea (butternut) is less commonly used ornamentally due to its smaller stature and susceptibility to disease, though it can enhance naturalized areas with its upright form. Environmentally, Juglans species support by providing and for . The nuts of J. nigra constitute a significant portion of diets for eastern fox squirrels (up to 10%) and various birds, while its leaves serve as browse for , and the tree offers roosting sites for species like the eastern screech-owl. J. regia acts as a host plant for moth larvae, including the hickory horned devil () and , and its nuts feed small mammals such as squirrels and chipmunks. These contributions enhance local ecosystems, promoting and vertebrate populations in wooded or settings. In restoration ecology, Juglans s play a role in rehabilitation. J. nigra demonstrates tolerance to harsh conditions, achieving 30-50% higher rates on surface-mined lands when paired with , aiding and efforts. Ongoing conservation programs for like J. cinerea involve planting resistant strains to restore native riparian and upland forests, preserving genetic diversity amid pathogens like butternut canker. Additionally, J. regia improves through nutrient cycling and supports in mixed agroecosystems, contributing to overall .

References

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